Bothalia

‘N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Vol. 25,2

Oct./Okt. 1995

TECHNICAL PUBLICATIONS OL THE NATIONAL BOTANICAL INSTITUTE, PRETORIA

TEGNIESE PUBLIKASIES VAN DIE NASIONALE BOTANIESE INSTITUUT, PRETORIA

Obtainable from the National Botanical Institute, Private Bag X 1 0 1 ,

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 1-20 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-

bosch Botanic Gardens. Published: 1 & 2.

Verkrygbaar van die Nasionale Botaniese Instituut, Privaatsak X 1 0 1 ,

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

bare publikasies kan aangevra word.

Bothalia is vemoem 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.

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

The memoirs are individual treatises usually of an ecological nature, but ‘n Reeks van losstaande omvattende verhandelings oor vernaamlik eko-

sometimes dealing with taxonomy or economic botany. Published: Nos logiese, maar soms ook taksonomiese of plantekonomiese onderwerpe.

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

for illustration.

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

An index to Vols 1^-9 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 limited 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 gepub-

liseer 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 ekonomi-

ese waarde vir afbeelding.

Vanaf Vol. 55 sal 20 plate op ‘n keer met ongereelde tussenposes gepub-

liseer word..

‘n Indeks tot Vols 1 — 49 is beskikbaar.

‘n Taksonomiese verhandeling oor the flora van die Republiek van Suid-Af-

rika, Lesotho, Swaziland, Namibie en Botswana. Die FSA bevat beskry-

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

genusse en spesies, sinonimie, literatuur, verwysings na enkele eksemplare,

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:

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

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

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

able from/Beskikbaar van: A. A. Balkema Marketing, Box/Posbus 317, Claremont 7735, RSA.

BOTHALIA

'N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Volume 25,2

Scientific EditorAVetenskaplike Redakteur: O.A. Leistner

Technical Editor/Tegniese Redakteur: B.A. Momberg

NATIONAL

^Botanical

INSTITUTE

NASIONALE BOTANIE*

INSTITUUT S

| Private b>ag X101 PRETORiAOOC

1995 -12- 0 1

I X tgr h-r?~TORtA

I national F o-TA^c^-iL

(MSTiTunrE

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag XI 01, Pretoria 0001

ISSN 0006 8241

Oct./Okt.l995

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 25,2

1. Three subspecies of Pelargonium laevigatum (Geraniaceae). C.M. VAN WYK (nee SCHONKEN) and J.J.A.

VAN DER WALT 133

2. A systematic study of the genus Pseudopentameris (Arundinoideae: Poaceae). N.P. BARKER 141

3. A note on Combretum subgenus Combretum section Macrostigmatea (Combretaceae). E.F. HENNESSY and

S. RODMAN 149

4. New records and new species of Asclepiadaceae from Namibia. P.V. BRUYNS 155

5. An annotated list of Urediniomycetes (rust fungi) from South Africa 1 : Melampsoraceae and Pucciniaceae,

excluding Puccinia and Uromyces. M. VAN REENEN 173

6. Studies in the Marchantiales (Hepaticae) from southern Africa. 9. The genus Marchantia and its five local

species. S.M. PEROLD 183

7. New combinations and a complete list of Asparagus species in southern Africa (Asparagaceae). A.C.

FELLINGHAM and N.L. MEYER 205

8. The taxonomic history of the Ricciaceae (1937-1995) and a classification of sub-Saharan Ricciae. S.M.

PEROLD 211

9. Notes on African plants:

Author citation ‘ex Krauss’ in nomenclature. B.A. MOMBERG 245

Asteraceae. A note on Ursinia brachyloba. P.P.J. HERMAN 244

Asteraceae. A new combination in Cotula. P.P.J. HERMAN 244

Boraginaceae. Typification of the sections in Lobostemon. M.H. BUYS and J.J.A. VAN DER WALT 233

Cyperaceae. New records for southern Africa and KwaZulu-Natal. J. BROWNING, K.D. GORDON-

GRAY and C.J. WARD 238

Ericaceae. A new species of Erica from the Western Cape. E.G.H. OLIVER 242

Lunulariaceae. Fruiting Lunularia cruciata , now also reported from southern Africa. S.M. PEROLD 239

Podocarpaceae. Notes on Podocarpus in southern Africa and Madagascar. O.A. LEISTNER, G.F.

SMITH and H.F. GLEN 233

Pteridophyta. New distribution records of South African pteridophytes. J.E. BURROWS and N.R.

CROUCH 236

10. The psocid Liposcelis bostrychophilus Badonnel (Psocoptera: Liposcelidae): an occasional herbarium pest.

E. RETIEF, A. NICHOLAS and H. BAIJNATH 247

1 1 . Obituaries: Winsome (Buddy) Barker (1907-1994). J.P. ROURKE 255

Alan Percy-Lancaster (1944-1995). G.F. SMITH and H.F. GLEN 259

Friedrich von Breitenbach OMS (1916-1995). H.F. GLEN and W.G. WELMAN 260

12. National Botanical Institute: list of staff and publications, 31st March 1995. Compiler: B.A. MOMBERG 265

13. Guide for authors to Bothalia 277

Digitized by the Internet Archive

in 2016

https://archive.org/details/bothaliavolume2525unse_0

Bothalia 25,2: 133-139(1995)

Three subspecies of Pelargonium laevigatum (Geraniaceae)

C.M. VAN WYK* (nee SCHONKEN) and J.J.A. VAN DER WALT**

Keywords: Geraniaceae, Pelargonium laevigatum , subspecies, taxonomy

ABSTRACT

Pelargonium laevigatum (L. f.) Willd. is a variable species. The macromorphology, anatomy of the leaves, chromosome

numbers and geographical distribution of the species were studied, and it is concluded that three subspecies should be

distinguished. The subsp. diversifolium (Wendl.) Schonken stat. nov. and subsp. oxyphyllum (DC.) Schonken stat. nov. were

originally described as species. A key to identify the three subspecies, a distribution map, and illustrations of the macromor-

phological characters as well as anatomical features of the leaves, are provided.

UITTREKSEL

Pelargonium laevigatum (L. f.) Willd. is 'n varierende spesie. Die makromorfologie, anatomie van die blare, chromosoom-

getalle en geografiese verspreiding van die spesie is bestudeer en daar is tot die gevolgtrekking gekom dat drie subspesies

onderskei moet word. Die subsp. diversifolium (Wendl.) Schonken stat. nov. en subsp. oxyphyllum (DC.) Schonken stat. nov.

is oorspronklik as spesies beskryf. ‘n Sleutel om die drie subspesies te identifiseer, 'n verspreidingskaart, en illustrasies van die

makromorfologiese kenmerke sowel as anatomiese kenmerke van die blare, word voorsien.

INTRODUCTION

Harvey (1860) described the section Glaucophyllum

and recognized Pelargonium laevigatum (L. f.) Willd. as

one of the species in the section. Van der Walt et al. (1990)

altered the composition of the section Glaucophyllum but

concluded that P. laevigatum should remain in the section.

P. laevigatum is a variable species and Schonken

(1980) suggested that three subspecies should be distin-

guished. For this study the macromorphology, leaf anat-

omy, chromosome numbers and geographical distribution

of the species have been studied to determine whether it

is justified to recognize infraspecific taxa.

MATERIAL AND METHODS

For the morphological and geographical distribution

studies, specimens of altogether 17 herbaria were studied.

This was followed by extensive field work on different

populations in the entire distribution area, mainly to de-

termine the variation in morphological features.

Transverse sections, 12-15 pm thick, of wax-embed-

ded terminal leaflets and petioles were cut with a rotary

microtome and stained with Alcian Green-Safranin (Joel

1983). The sections were made through the middle part

of the terminal leaflets and petioles.

Specimens examined for leaf anatomy and chromosome

numbers

1. Subsp. laevigatum : Maggs 16, Willowmore 3502+; Van der Walt

734 , 2n = 22, Swartberg Pass, Oudtshoom 1444+; Van der Walt 1594,

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

**Department of Botany, University of Stellenbosch Stellenbosch 7600.

MS. received: 1994-05-1 1.

+ project locality no.

Rooiberg Pass, Calitzdorp 4093+; 2n = 22, Schonken 150, Swartberg

Pass 2014+.

2. Subsp. diversifolium : Van der Walt 1592, Tweedside, Matjiesfon-

tein 405 1+; Van der Walt 1063, 2n = 22; Gydouw, Ceres 4132+.

3. Subsp. oxyphyllum : Van der Walt 1604, 2n = 22, Cedarberg 4133+;

Stirton 6401, Wupperthal 1085+.

For mitosis, root tips were treated with 0.002 mol 8-

hydroxiquinoline for 24 h at 20°C, fixed in 3:1 absolute

ethanol and glacial acid, and stained with aceto-cannine.

RESULTS

Pelargonium laevigatum (L. f) Willd., Species

plantarum 3,1: 685 (1800); Knuth: 436 (1912). Type:

‘Cap. Bonae Spei’, Thunberg s.n. ( UPS 15620, lecto.!,

here designated).

Erect and compact or prostrate and lax subshrub, much

branched, evergreen, 0.3-0.5 (-lm) high and 0. 3-0.5 m

in diameter. Stems smooth, herbaceous, glabrous or

glabrescent, grey or glaucous. Leaves unifoliolate to tri-

foliolate, articulated to petiole, subsucculent, glabrous or

pilose to hirsute, with sparse glandular hairs, glaucous;

pinnae sessile, lanceolate to terete, bases cuneate, apices

acute, margins entire or irregularly dentate to incised with

linear segments, pinnae (10-)20-40(-70) x 1-10 mm;

petiole (4—) 15-20(^40) mm long; stipules free, subulate,

3-11 x 0. 2-1.0 mm, somewhat succulent, glabrous to pi-

lose. Inflorescence : flowers borne singly or in reduced

pseudo-umbels of 2-5 flowers each, peduncle (6-)30-50

(-100) mm long, glabrous to pilose. Pedicel 0.5-13 mm

long, glabrous or with glandular hairs. Hypanthium 4-38

mm long, glabrous or with glandular hairs. Sepals 5, nar-

rowly ovate, adaxially concave, apices acute, glabrous or

pilose to hirsute, green to reddish brown with hyaline mar-

gins, 6-12 x 1.5-3. 5 mm. Petals 5, white or cream-col-

oured or pink; posterior 2 spathulate with apices obtuse

134

Bothalia 25,2 (1995)

or rounded or retuse, 1 1-30 x 4—9 mm, reflexed at ± 90°

from the middle, with reddish purple markings at base; an-

terior 3 spathulate to narrowly obovate with apices obtuse

or rounded or retuse, somewhat smaller (11-20 x 3-6 mm)

and somewhat lighter in colour than posterior petals, gradu-

ally recurved so that apices are at an angle of ± 90° with

bases, with fine purplish lines near bases. Stamens : fertile 7

(4 long, 1 medium, 2 short), staminodes 3, filaments basally

fused forming a staminal column 2-3 mm long; pollen yel-

low to orange. Ovary ovate, 5-lobed, pilose; style glabrous

or sparsely pilose basally, 1^4 mm long; stigmas 5, purplish,

1-2 mm long. Mericarps: 20-30 mm long; bases 3.5-6.0

mm long, tails 16-24 mm long. 2n = 22.

Key to subspecies

la Leaves trifoliolate, pinnae incised, southern Cape to E Cape

1. subsp. laevigatum

lb Leaves unifoliolate or trifoliolate, pinnae not incised, SW Cape

to southern Cape:

2a Leaves, glabrous, trifoliolate, flowers usually bome singly, re-

stricted to Cedarberg Mountains 3. subsp. oxyphyllum

2b Leaves hirsute, unifoliolate to trifoliolate, usually more than 1

flower per pseudo-umbel, Western Cape

2. subsp. diversifolium

1. subsp. laevigatum

Pelargonium laevigatum (L. f.) Willd., Species plantarum 3,1: 685

(1800); Pers.: 233 (1806); DC.: 667 (1824); Drege: 122 (1843); Harv.:

297 (1860); Compton: 295 (1931); Clifford: 218 (1970); Van der Walt

& Vorster: 87 (1981). Type: as for P. laevigatum (L. f.) Willd.

Geranium laevigatum L. f.: 306 (1781); L.: 619 (1784); Cav.: 255

(1787); Thunb.: 115 (1794); Thunb.: 522 (1823). Eumorpha laevigata

(L. f.) Eckl. & Zeyh.: 78 (1834-1837).

G. acuminatum Thunb.: 526 (1823). Pelargonium acuminatum

(Thunb.) DC.: 680 (1824); R. Knuth: 542 (1912). Type: locality and

collector unknown, specimen UPS 15555 with Thunberg’s handwriting

(lecto.!, here designated).

P. laevigatum (L. f.) Willd. var. compositum L'Her. ex DC.: 667

(1824). Type: locality and collector unknown, specimen G-DC 667-190

with De Candolle’s handwriting (holo.!).

P. macowanii Bolus: 157 (1890). Type: Cape Province, Boschberg

near Somerset East, MacOwan 1647 (BOL, holo.!; GRA, K!, SAM!, Z!).

Diagnostic morphological features

Leaves trifoliolate, pinnae variously incised, glabrous

to sparsely hirsute, flattened, 1-5 flowers per pseudo-um-

bel (Figure 1).

I IGURE 1 . — P. laevigatum subsp. lae-

vigatum, Ward-Hilhorst 185. A,

flowering branch, x 0.7; B, pet-

als, x 1 ; C, androecium, x 2; D,

gynoecium, x 2.

Bothalia 25,2(1995)

135

FIGURE 2. — Transverse section of petiole of P. laevigatum. A, subsp.

laevigatum, Maggs 16: B, subsp. diversifolium, Van der Walt

1603: C, subsp. oxyphyllum. Van der Walt 1604.

Diagnostic anatomical features of leaves

Petiole : cordiform in transverse section, glandular hairs

multicellular, non-glandular hairs unicellular, hypodermis

not clearly differentiated, parenchymatous cortex 4-6 lay-

ers, extraxylary sclerenchyma cylinder continuous and

2-4 layers wide, main vascular bundles 4, smaller bundles

2-6, medullary bundle usually lacking (Figure 2A). Lam-

ina: V-shaped to more or less flattened in transverse sec-

tion, trichomes as on petiole, stomata evenly distributed

in ad- and abaxial epidermis, mesophyll not clearly dif-

ferentiated in palisade and spongy parenchyma, 2 adaxial

palisade layers larger than 1 abaxial layer (Figure 3A).

Specimens examined

EASTERN CAPE. — 3225 (Somerset East): Boschberg, (-DC), Mac-

Owan 1479 (SAM), 1647 (BOL, GRA. K, MEL, SAM. Z). 3323 (Willow-

more): Slypsteenberg, (-AC), Esterhuysen 6278 (BOL); near Willowmore,

(-AD), Maggs 16 (STEU); Buispoort, (-AD), Theron 1011 (PRE);

Baviaanskloof, (-BC), Van Wyk 384 (STE); 37 km E of Willowmore, (-DA),

Olivier 1643 (STEU); between Miskraal and Smitskraal, (-DA). Hugo 1451

(STE); Studtis, (-DB), Martson 53, 67 (STE); 15 km W of Joubertina,

(-DC), Van der Walt 856 (STEU); Joubertina, (-DD), Esterhuysen s.n.

(BOL), Fourcade 3217 (BOL), Horn s.n. (PRE); near Joubertina, (-DD),

Esterhuysen 7040, 22797 (BOL). 3324 (Steytlerville): Langkloof, (-AB),

Ecldon & Zeyher609 (SAM); Kouga Hills, (-CA), Esterhuysen 6661 (BOL,

PRE); Kouga Mountains, (-CA), Bayliss 6382 (MO. NBG. WAG); Kouga,

(-CA), Compton 10525 (BOL, NBG); between Cambria and Smitskraal,

(-CB), Oliver 4551 (STE); Kareedouw, (-CD), Compton 4516 (BOL); Mor-

denaarskloof, (-CD), Stayner s.n. (NBG); Baviaanskloof, (-DA), Bayliss

4423 (MO); NW of Cambria, Baviaanskloof, (-DA), Oliver 4535 (PRE.

STE). 3325 (Port Elizabeth): Otterford Forest Reserve, (-CC). Oliver 4443

(PRE). Rodin 1109 (BOL, MO, PRE). 3424 (Humansdorp): Krommerivier

Heights, (-BA), Fourcade 2704 (BOL, PRE, STE).

WESTERN CAPE. — 3321 (Ladismith): Rooiberg, (-DA), Compton

3917 (BOL, NBG), Esterhuysen 17144 (BOL), Oliver 5330 (PRE, STE),

Schonken 150 (STEU), Taylor 9796 (STE). Van der Walt 1594 (STEU);

Gamka Mountain Reserve, (-DB), Boshojf 360 (STE). 3322 (Oudts-

hoom): Baviaanskloof. (-AC), Bayliss 4423 (NBG), Boucher 38 (STEU),

Gill 19 (BOL, PRE); Swartberg Pass, (-AC), Bolus 11739 (BOL, PRE).

Compton 10419 (NBG), Hafstrom & Acocks 753 (BOL, PRE), Moffett

696 (STEU), Pocock 120 (PRE), Schonken 150. 189. 190 (STEU), Stokoe

s.n. (SAM), Thompson 2195 (PRE, STE) Van der Walt 734, 1148, 1589

(STEU); Kamanassie Mountains, (-DA/DB), Zinn s.n. (SAM). 3323

(Willowmore): 13 km W of Uniondale, (-CA), Wells 2835 (GRA. PRE);

Bo-Koega, (-CB), Bayliss 7081 (MO); Kouga Mountains, (-CB). Wisura

2173 (NBG); Prince Alfred Pass, (-CC), Fourcade 2091 (BOL).

Subsp. laevigatum occurs in the southern and eastern

Cape, and it is known from Rooiberg near Calitzdorp, east-

wards to Otterford Forest Reserve near Port Elizabeth (Figure

4). It has been collected from 180 m above sea level near

Humansdorp to 1 600 m in the Swartberg Range. The rainfall

in its distribution area varies from 200-600 mm per annum.

Subsp. laevigatum is morphologically very variable but

a continuous variation pattern makes it impossible to di-

vide it into different taxa. Varying characters are the

number of flowers per pseudo-umbel and the size of the

petals, as well as the indumentum and degree of incision

of the leaf margins. To complicate matters, there seem to

be natural hybrids between subsp. laevigatum and Pelar-

gonium fruticosum (Cav.) Willd. P. laevigatum depicted

in Van der Walt & Vorster (1981) with greenish leaves is

most probably an example of such a hybrid. These sus-

pected hybrids occur in the Swartberg Pass where subsp.

laevigatum and P. fruticosum are common.

The plants occurring at Boschberg near Somerset East

have finely divided leaves superficially resembling those

of P. fruticosum. This form was described by Bolus (1890)

as P. macowanii and quoted by Knuth (1912) as P. divari-

cation (Thunb.) DC. var. scabrum Harv. However, the

glaucous leaves and floral characters of these plants are

typical of P. laevigatum and we concluded that they rep-

resent a form of subsp. laevigatum. The atypical leaves

are probably the result of the local environmental condi-

tions and geographical isolation.

Some plants growing in Long Kloof (Fourcade 2704)

and in the Otterford Forest Reserve ( Wilman 1109 ) have

up to five flowers per pseudo-umbel and the petals are

exceptionally large. The pinnae of these plants are also

relatively wide.

136

Bothalia 25,2(1995)

FIGURE 3. — Transverse sections of terminal leaflets of P. laevigatum. A, subsp. laevigatum , Maggs 16; B, subsp. diversifolium. Van der Walt 1603.

C, D, subsp. oxyphyllum : C, flattened leaf. Van der Walt 1501; D, centric leaf. Van der Walt 1604.

FIGURE 4. — Known geographical distribution of/5, laevigatum: subsp. laevigatum, •; subsp. diversifolium, ■; subsp. oxyphyllum, ▲.

Bothalia 25,2 (1995)

137

2. subsp. diversifolium (Wendl.) Schonken stat. nov.

Pelargonium diversifolium Wendl., Botanische Beobachtungen: 52

(1798); Willd.: 664 (1800); Pers.: 230 (1806); Wendl.: 53 (1809). Icono-

type: Wendland, Collectio plantarum 2,4: t. 58 (1809).

Geranium trifoliatum Andr.: (1805). Pelargonium trifoliatum (Andr.)

Sweet: 294 (1826). Iconotype: Andrews, Geraniums c. ic. (1805).

Diagnostic morphological features

Leaves unifoliolate to trifoliolate, pinnae not incised,

hirsute to densely hirsute, flattened, 1-2 flowers per

pseudo-umbel (Figure 5).

Diagnostic anatomical features of leaves

Petiole : adaxially flattened, glandular hairs multicellular,

non-glandular hairs unicellular, hypodermis clearly differen-

tiated, parenchymatous cortex 5-6 layers, extraxylary

sclerenchyma cylinder continuous and 2-6 layers wide,

main vascular bundles 4, smaller bundles 4—8, medullary

bundle 1 or 0 (Figure 2B ). Lamina : flattened, trichomes as

on petiole, more stomata abaxially, mesophyll differentiated

in palisade and spongy parenchyma, 2 adaxial palisade layers

larger than 1 abaxial layer (Figure 3B).

Specimens examined

WESTERN CAPE. — 3319 (Worcester): Agter-Witzenberg, (-AB),

Marais 47 (STEU); Elandsfontein, Skurweberge, (-AB), Schlechter 1750

(PRE), 10020 (BOL, E, MO, P, PRE, W, Z); 11 km N of Gydouw Pass,

(-AB), Hutchinson 1040 (BOL), Van der Walt 1603 (STEU); Farm Me-

rino, Theronsberg, (-AD), Cillie 686 (STEU); Baviaansberg, Ceres,

(-BA), Compton 12850 (NBG), Stokoe s.n. (SAM); Ertjieslandkloof, Ce-

res, (-BA), Wilman 2266 (BOL, MO); Hottentotskloof, (-BC), Pearson

4945 (BOL); Bokkerivier farms, Ceres, (-BD), Barker 10144 (NBG),

Booysen 60 (NBG); between Concordia and Triangle, (-BD). Barnard

754 (SAM); Bonteberg, Worcester, (-BD), Compton 9936 (NBG),

Esterhuysen 3712 (BOL); Eendracht, (-DB), Lewis 1777 (SAM). 3320

FIGURE 5. — P. laevigatum subsp. di-

versifolium, Ward-Hilhorst

111. A, flowering branch, x 0.8;

B, petals, x 1.6; C, androecium,

x 3.2; D, gynoecium x 3.2.

Bothalia 25,2 (1995)

138

FIGURE 6. — P. laevigatum

subsp. oxyphyllum. Ward-

Hilhorst 210. A, flower-

ing branch, x 0.7; B, pet-

als, x 1 ; C, androecium, x

3; D, gynoecium, x 3.

(Montagu): Tweedside, (-AB), Barker 20602 (BOL), Marloth 10848,

12096 (PRE, STE), Van der Walt 1592 (STEU); Cabidu, (-AB). Hall

171 (NBG), Oliver s.n. (NBG); Pieter Meintjies, (-AD), Rogers 17887

(PRE); Witteberg, (-AD/BC), Adamson s.n. (BOL), Compton 2680

(BOL), 12213 (NBG); Matjiesfontein, (-BA), Marloth 2961 (BOL,

PRE); Whitehall, (-BA), Compton 7957 (NBG), Fisantekraal, (-BC),

Compton 21 1 10 (NBG). 3321 (Ladisniith): Elandskloof between Vleiland

and Seweweekspoort, (-AD), Maggs 28 (STEU); Ladismith, (-BD),

Geldenhuys 454 (STE); Rietkloof, (-BD), Lewis 1168 (SAM). 3322

(Oudtshoom): Robinson Pass, (-CC), Taylor s.n. (BOL).

Subsp. diversifolium occurs from Gydouw near Ceres

eastwards to the Robinson Pass between Oudtshoom and

Mossel Bay (Figure 4). It is often found in mountainous

habitats in a variety of vegetation types. Its distribution

area falls entirely in the winter rainfall region, but it usu-

ally occurs on the lower slopes of mountains or on hills

where the rainfall is relatively low in comparison to habi-

tats higher up on the mountains.

3. subsp. oxyphyllum (DC.) Schonken stat. nov.

Pelargonium oxyphyllum DC., Prodromus systematis naturalis regni

vegetabilis I: 667 (1824); Drege: 209 (1843). Type: locality and collector

unknown, specimen C-DC 667-190 with De Candolle’s handwriting

(holo. !).

P. laevigatum (L. f.) Willd. var. oxyphyllum (DC.) Harv.: 297 (1860).

Diagnostic morphological features

Leaves trifoliolate, pinnae not incised, glabrous, sub-

terete to flattened, usually 1 flower per pseudo-umbel

(Figure 6).

Diagnostic anatomical features of leaves

Petiole : adaxially flattened to cordiform, without

trichomes, hypodermis not clearly differentiated, paren-

chyma cortex 5-6 cell layers, extraxylary sclerenchyma

cylinder continuous and 2-5 layers wide, main vascular

bundles 4, smaller bundles 6-8, medullary bundle 1 or 0

(Figure 2C). Lamina : more or less flattened to centric,

more stomata abaxially, mesophyll differentiated in pali-

sade and spongy parenchyma, 1 adaxial palisade layer

Bothalia 25,2 (1995)

139

larger than 2 abaxial layers, mesophyll in centric leaves

not clearly differentiated (Figure 3C, D).

Specimens examined

WESTERN CAPE. — 3219 (Wupperthal): Heuningvlei, Wupperthal

road, (-AA), Schonken 86 (STEU); Middelberg, (-AA/AC), Compton

7503 (NBG); Kerfoot 6150 (NBG); Sneeukop road. (-AC), Strauss 63

(NBG); between Algeria and Dwarsrivier, (-AC), Schonken 81 (STEU);

Hoogvertoon, (-AC), Haynes 1205 (PRE, STE); Cedarberg, (-AC), Van

der Walt 1604 (STEU); Matjesrivier, (-AC), Wagener 281 (NBG);

Dwarsrivier, (-AC), Rycroft 2641 (MO, NBG, STE); Tafelberg, (-AC),

Barnard s.n. (SAM), Lamb 1 (STE); between Eselbank and Wupperthal,

(-AC), Van der Walt 1501. 1502 (STEU); Eselbank, (-AC), Drege s.n.

(MEL, PRE); Wupperthal, (-AC), Stirton 6401 (PRE, STEU), Thode

A1965 (NH, PRE); Gideonskop, (-CB), Stokoe s.n. (NBG); Juriesberg,

(-CB), Compton 12742 (NBG); Zoo Ridge, Suurvlakte, (-CD), Taylor

6120 (PRE, STE); Skurweberge E of Citrusdal, (-CD), Prismos s.n.

(SAM).

Subsp. oxyphyllum is confined to the Cedarberg Range,

occurring from Heuningvlei in the vicinity of Wupperthal

southwards to the district of Citrusdal (Figure 4). It grows

in flat areas on deep sandy soil derived from sandstone.

The annual rainfall in its distribution area varies from

200-500 mm.

DISCUSSION

P laevigatum is taxonomically a complicated species

because it displays a high degree of phenoplasticity. Al-

most all the populations studied have unique combinations

of characters. The floral structure and many leaf characters

of the plants investigated are, however, basically identical

and therefore it was decided to consider them as conspe-

cific. The three subspecies which are distinguished, have

the same somatic chromosome number of 2n = 22. The

rank of subspecies rather than variety was chosen for the

three infraspecific taxa because they are allopatric.

The phenoplasticity of the species is also reflected in the

leaf anatomy. The leaflets of subsp. oxyphyllum for example

vary from flattened to centric. A medullary vascular bundle

can be present or absent in the petioles of all three subspecies,

even in different leaves of the same plant. The occurrence

of hairs on the leaves of subsp. laevigatum is also extremely

variable. Anatomical features of the leaves of P. laevigatum

are therefore of limited taxonomic importance.

ACKNOWLEDGEMENTS

We are indebted to Prof. Dr Focke Albers for assisting

us to determine the chromosome numbers and the Bren-

thurst Library (copyright) for permission to reproduce the

water-colour paintings by Ellaphie Ward-Hilhorst.

REFERENCES

ANDREWS, H.C. 1805. Geraniums: ora monograph of the genus Gera-

nium, Vol. 2. Taylor, London.

BOLUS, H. 1890. Contributions to South African botany. Journal of the

Linnean Society, Botany 25: 157.

CAVANILLES, A.J. 1787. Monadelphiae classis dissertationes decern.

Quarta dissertatio botanica, de Geranio, F.A. Didot, Paris.

CLIFFORD, D. 1970. Pelargoniums including the popular Geranium.

edn 2. Blandford Press, London.

COMPTON, R.H. 1931. The flora of the Whitehill District. Transactions

of the Royal Society of South Africa 19: 295.

DE CANDOLLE, A.L.L.P. 1824. Prodromus systematis naturalis regni

vegetabilis. Treutel & Wiirtz, Paris.

DREGE, J.F. 1843. Zwei pflanzengeographische Documente nebst einer

Einleitung von E. Meyer. Jena & Leipzig.

ECKLON, C.F. & ZEYHER, K.L.P. 1834—1837. Enumeratio plantarum

africae australis extratropicae. Perthes, Flamburgii.

HARVEY, W.H. 1860. Geraniaceae. Flora capensis 1: 254-308.

JOEL, D.M. 1983. A.G.S. (Alcian Green-Safranin) — a simple differential

staining of plant material for light microscopy. Proceedings of the

Royal Microscopy Society 18/3: 1 49-15 1 .

KNUTH, R.G.P. 1912. Geraniaceae. Das Pflanzenreich 4. 129 (Heft 53).

Wilhelm Engelmann. Berlin.

LINNAEUS, C. (fil.) 1781. Supplementum plantarum. Impensis or-

phanotrophei, Braunschweig.

LINNAEUS, C. 1784. Systema vegetabilium , 4th edn. Gottingen.

PERSOON, C.H. 1806. Synopsis plantarum 2, Cotta, Tubingen.

SCHONKEN, C.M. 1980. 'n Morfologiese en taksonomiese studie van

die seksie Glaucophyllum van Pelargonium. M.Sc. thesis. Univer-

sity of Stellenbosch.

SWEET. R. 1826. Geraniaceae, Vol. 3: 294. James Ridgway, London.

THUNBERG, C.P. 1794. Prodromus plantarum capensium. Part 1. Ed-

man, Uppsala.

THUNBERG, C.P. 1823. Flora capensis, edn 3. Stuttgart.

VAN DER WALT, J.J.A., ALBERS, F. & GIBBY, M. 1990. Delimitation

of Pelargonium sect. Glaucophyllum (Geraniaceae). Plant Sys-

tematics and Evolution 171: 15-26.

VAN DER WALT, J.J.A. & VORSTER, P.J. 1981. Pelargoniums of

southern Africa, Vol. 2. Juta, Cape Town.

WENDLAND, J.C. 1798. Botanische Beobachtungen. Hannover.

WENDLAND, J.C. 1809. Collectio plantarum. Hannover.

WILLDENOW, C.L. 1800. Caroli a Linne species, 4th edn, Vol. 3. Nauk,

Berlin.

Bothalia 25,2: 141-148 (1995)

A systematic study of the genus Pseudopentameris (Arundinoideae: Poaceae)

N.P. BARKER*

Keywords: Arundineae, Arundinoideae, conservation, leaf anatomy, phenetics, phylogenetics, Poaceae, Pseudopentameris, systematics

ABSTRACT

The genus Pseudopentameris Conert is examined morphologically and anatomically. A phenetic study of the morphologi-

cally variable species P. macrantha indicates that two taxa should be recognised. One of these, P. caespitosa N.P. Barker, is

described as new. In addition, the study supports the inclusion of Pentameris obtusifolia in Pseudopentameris. The genus

Pseudopentameris is re-delimited to accommodate the new taxa, and a key to species is provided. Details of the cytology,

phylogeny and conservation status of taxa in the genus are also discussed.

UITTREKSEL

Die genus Pseudopentameris Conert word morfologies en anatomies ondersoek. ’n Fenetiese ondersoek van die varierende

spesie P. macrantha toon dat twee taksons erken moet word. Een van hulle, P. caespitosa N.P. Barker, word as nuut beskryf.

Verder ondersteun die ondersoek die insluiting van Pentameris obtusifolia in Pseudopentameris. Die genus Pseudopentameris

word heromgrens om die nuwe taksons te huisves, en ’n sleutel tot die spesies word verskaf. Besonderhede van die sitologie,

filogenie en bewaringstatus van taksons in die genus word ook bespreek.

INTRODUCTION

The genus Pseudopentameris Conert is endemic to the

lowland fynbos of the southwestern Cape Province, South

Africa. This genus was created by Conert (1971), who

thought that two species of the genus Danthonia DC., D.

macrantha Schrad. and D. brachyphylla Stapf, merited

recognition as a separate genus. This new genus was char-

acterised by unusually large spikelets which, while similar

to both those of Pentameris P Beauv. and Danthonia , dif-

fered in having many-nerved glumes. The nature of the

fruit of these two species also set them apart.

Studies of the leaf anatomy (discussed below) provided

further evidence of their singularity (De Wet 1956, 1960;

Ellis 1985a). Ellis (1985b) observed that the leaf anatomy

of the taxon he called Pentameris sp. nov., bore a strong

resemblance to the anatomy of the two known species of

Pseudopentameris, and postulated a hybrid origin for this

species. Additional studies on the fruit morphology

(Barker 1986, 1989, 1993) and leaf anatomy (presented

below) also suggested that this unusual species has close

affinities with the other species of Pseudopentameris. Be-

cause of the nomenclatural confusion surrounding this

taxon (see 1. Pseudopentameris obtusifolia below), past

studies have referred to it by a number of names. Barker

(1989) and Ellis (1985b) referred to it as Pentameris sp.

nov. whereas Linder & Ellis (1990) refer to it as Pen-

tameris squarrosa Stapf. Some herbaria (in particular PRE

and STE) misapplied this name to the taxon now known

as Pentameris oerophila N.P. Barker (1993), whereas BOL

has used the synonym Pentameris squarrosa Stapf.

* Department of Botany, University of Cape Town, Private Bag, Ronde-

bosch 7700. Present address: Department of Botany, University of the

Western Cape, Private Bag XI 7, Bellville 7535.

Email: nigel@botany.uwc.ac.za

MS. received: 1994-09-27.

This study encapsulates all these observations, and

documents the anatomical and morphological variation

within the genus, and presents the relevant nomenclatural

modifications that are required as a result of these obser-

vations.

MORPHOLOGY

Pseudopentameris is unusual in having two species ( P.

macrantha and P. obtusifolia) with aerial stems that pos-

sess the anatomy of a rhizome (H.P. Linder unpublished).

Until the anatomy and homology of these structures is

better understood, the term ‘stem’ is used, as it serves to

differentiate this structure from below-ground rhizomes

and the annually produced flowering culm.

These stems are often branched, and the annual flow-

ering culms are produced from their apical buds. Sub-

sequent shoot growth takes place from what Linder &

Ellis (1990) term caulescent innovation buds on the stems,

which results in branched stems. This branching can lead

to a shrub-like growth form in older plants. This type of

growth form has also been described in some species of

Pentameris (Barker 1993). In Pseudopentameris obtusifo-

lia these branched aerial stems are scandent, spreading

out from the base of the plant, over and between the other

vegetation. Linder & Ellis (1990) illustrate this species

(named in their work as P. squarrosa) and consider its

unusual growth form to be an adaptation enabling the

plant to survive long interfire periods that are charac-

teristic of the fynbos biome. The two species which do

not have these stems (P. brachyphylla and P. caespitosa

sp. nov. described below) produce the annual flowering

culm from an underground rhizome that can become fi-

brous or woody in older plants.

In P. obtusifolia the stems are covered by the persistent

remains of the leaf sheaths, the dead leaf blades being

142

Bothalia 25,2 (1995)

FIGURE 1. — First two axes of Princi-

pal Components Analysis

(PCA) of 59 specimens match-

ing the previous description of

Pseudopentameris macrantha.

Specimens measured for six

floral characters. First axis:

81.7% of variation present in

data. Second axis: 6.8%. Lines

drawn around two groups were

placed by eye. Each symbol

represents an individual speci-

men. Specimens with rhi-

zomatous growth form, P.

macrantha, O', specimens with

caespitose growth form, P.

caespitosa. A; specimens with

no recorded growth form, ★.

deciduous. In the other species in the genus, the dead leaf

blades are persistent, and in P brachyphylla (Stapf) Conert

they curl in a characteristic manner at the base of the

plant. The living leaf blades are short, open to rolled and

rigid in P. obtusifolia, but long, somewhat flexuous and

open to rolled in the other species.

The inflorescence is a contracted panicle, which be-

comes somewhat lax during anthesis and comprises 8-80

spikelets. Spikelet size is variable between the species.

The glumes are 3-nerved in P. obtusifolia (the lateral veins

are visible only at the base of the glumes) and 5-9-nerved

in the other species. The lemmas are variously pubescent,

with a geniculate central awn. The lemma lobes are acu-

minate and variously adnate to the lateral bristle. The

palea is bicarinate, bifid at the apex and longer than the

lemma body. The lodicules are many-nerved (G.A. Ver-

boom unpublished data), cuneate, glabrous or ciliolate.

The latter condition is developed to the extreme in P. ob-

tusifolia, which has densely ciliolate lodicules, the cilia

being as long as the lodicule bodies.

Each floret has three large anthers (up to 9 mm). The

ovaries of all taxa possess what Barker (1990, 1994)

termed pseudostigmata. These structures have the appear-

ance of stigmatic hairs, and arise from the region between

the points of origin of the style branches, although the

style branches themselves appear to be absent (or alter-

natively have stigmata along their entire length, from the

apex of the ovary upwards). As the pseudostigmata are

easily deciduous, they are seldom observed on mature

fruit. Their presence may, however, explain the references

by De Wet (1954b) and Tomlinson (1985) to hairs which

are decurrent on the inner side of the caryopsis and join

over the top of the ovary. A similar description of these

structures is given by Chippindall (1955). These features,

although not labelled as pseudostigmata, appear in the fig-

ures published by Barker (1986, 1990). The fruit is a cary-

opsis (Clayton & Renvoize 1986), up to 6 mm long, with

a long, canaliculate hilum extending almost the full length

of the fruit. The embryo is a quarter to a fifth the length

of the fruit (Conert 1971; Barker 1986, 1989, 1990). The

surface of the caryopsis is reticulate (Barker 1986, 1989).

Barker (1990) noted substantial variation in the size of

the floral parts in specimens identified as P. macrantha.

This prompted an additional study in which measurements

of six floral characters from 59 specimens were taken,

including those examined in the earlier study. The

phenetic package NTSYS-pc version 1.4 (Numerical Tax-

onomy and SYStematics package; Rohlf 1988) was used

to perform a Principal Components Analysis (PCA).

The first three principal components (axes) account for

92.9% of the variation in the data. When the specimens

are plotted out on the first two axes (accounting for 88.5%

of the variation), two clusters are apparent (Figure 1).

Each cluster contains specimens from a wide geographic

area, and the geographic areas of the two clusters overlap

extensively. Furthermore, specimens from each cluster

have been observed to co-occur. Despite this sympatric

distribution, a one-sided T-test showed that the differences

in the means of each of the six characters from the two

clusters were statistically significant at a 99.9% confi-

dence interval. Table 1 presents the sample means and

standard deviations of these characters in each cluster.

TABLE 1. — Sample means and standard deviations (mm) for six quan-

titative floral characters of two clusters of specimens correspond-

ing to P. macrantha and P. caespitosa. A one-sided T-test shows

that differences between means for all characters are significant

at 99.9% confidence interval

P. macrantha P. caespitosa

Mean Std. dev. Mean Std. dev.

Bothalia 25,2 (1995)

143

A character not included in the PCA, but overlaid onto

the PCA plot, is that of growth form, as related to the

presence of the branching stems. This feature was ob-

served in the field for some of the specimens included in

the PCA, and the position in the PCA plot of these speci-

mens is shown in Figure I by means of open symbols.

As can be seen from Figure 1, each cluster has a specific

growth form. The cluster with the smaller floral parts in-

cludes specimens with branching stems, whereas the

specimens with larger floral parts do not possess stems.

These two clusters are considered here to be two species:

P. macrantha and P. caespitosa N.P. Barker sp. nov. De-

scriptions and details on nomenclature of these two spe-

cies appear below.

LEAF ANATOMY

Pseudopentameris has been well examined anatomi-

cally by De Wet (1956, 1960), Renvoize (1981), Ellis

(1985a), Tomlinson (1985) and Barker (1990). The genus

as a whole possesses a sclerophyllous type of arundinoid

leaf anatomy (Ellis & Linder 1992). Ellis (1985a) found

minor anatomical differences between P. macrantha and

P. brachyphylla. The anatomy of P obtusifolia is only

slightly different from the former two species (Ellis

1985b). All the taxa share characters such as the presence

of large, inflated abaxial epidermal cells, dumbbell-shaped

silica bodies and the absence of abaxial micro- and macro-

hairs. This knowledge was supplemented in this study by

a Scanning Electron Microscope (SEM) survey of the leaf

epidermides of all the species in the genus.

The SEM study of both adaxial and abaxial leaf anat-

omy was carried out on leaf blade material of nine her-

barium specimens of P. macrantha , nine specimens of P.

caespitosa, five specimens of P. brachyphylla and 1 1

specimens of P. obtusifolia. Leaf material was mounted

on SEM stubs and then coated in gold-palladium. The

specimens were examined using an ISI-SX-25 Scanning

Electron Microscope.

The abaxial epidermis of all specimens examined

lacked both microhairs and macrohairs and, as noted by

Ellis (1985a), the abaxial epidermides of all the species

are almost indistinguishable. However, occasional needle-

like prickles were observed. Figure 2A illustrates the

abaxial epidermis of P. obtusifolia.

The adaxial epidermides of P. macrantha, P. caespitosa

and P. brachyphylla were almost identical (Figure 2B, C),

but the epidermis of P. obtusifolia differed, in that it had

a denser distribution of prickles (Figure 2D). In all the

FIGURE 2. — Leaf blade SEM of Pseudopentameris. A, D. P. obtusifolia, Ellis 2342: A, abaxial epidermis showing large inflated epidermal

cells and small prickles; D, deeply ribbed adaxial epidermis heavily adorned with prickles. B, P. brachyphylla, Boucher 357a,

adaxial epidermis showing open furrows and ribs with prickles. C, E, P. macrantha, Ellis 2515: C, adaxial prickles, note also

dumbbell-shaped silica bodies; E, microhair from adaxial epidermis, note minute apical cell. Scale bars: A, 225 gm; B. 240 pm;

C, 1 15 gm; D, 325 gm; E, 17 gm.

144

Bothalia 25,2 (1995)

species, the prickles are needle-like and angled almost par-

allel to the epidermis. These prickles appear to be modi-

fications of the epidermal cell surface (shown magnified

in Figure 2C). These prickles are unlike those found in

Pentameris (Barker 1993), Pentaschistis (Ellis & Linder

1992) and at least one species of Merxmuellera (Barker

& Ellis 1991). The prickles of these latter taxa are slightly

inflated basally, variable in length but longer than in

Pseudopentameris , and protrude away from the epidermis.

Microhairs with minute, deflated apical cells were ob-

served at the bottom of the adaxial furrows (Figure 2E).

These bicellular microhairs are similar, except in size, to

those reported in Pentameris (Barker 1993). Microhair

size ranged from 55-67 pm, with a mean of 64.8 pm (five

samples), and (unlike in Pentameris) no species level size

differences were apparent. No microhairs were observed

on the abaxial leaf surface.

The presence, distribution and nature of microhairs in

this genus has been a contentious issue. De Wet (1956,

1960) implies that, as Pseudopentameris has a panicoid

epidermis, bicellular hairs (microhairs) would be present,

presumably on the abaxial surface. In contrast, Renvoize

(1981) considered microhairs to be absent, but later he

described ‘long slender papillae, which in a few instances

appear to bear the remains of a thin-walled apical cell’

which were associated with the adaxial surface (Renvoize

1986). This description matches the structure shown in

Figure 2E, and it appears that these papillae are in fact

microhairs.

CYTOLOGY

The cytology of the genus Danthonia was examined

by De Wet (1953, 1954a, b), who was of the opinion that

the base chromosome number of this group is x = 6. The

subsequent division of the genus Danthonia resulted in

limited cytological knowledge for each of the segregate

genera. Additional research by Spies & Du Plessis (1988)

and Du Plessis & Spies (1988) has increased cytological

knowledge about the Danthonia segregates but no chro-

mosome counts of the species of Pseudopentameris have

been published. From analyses of the relationship between

stomatal size and chromosome counts, De Wet (1954a)

predicted a count of 2n = 24. This figure is credible, as

many of the African arundinoid genera display intraspe-

cific polyploidy (see the study on Cliaetobromus by Spies

et al. 1990).

CONSERVATION STATUS

Two species in the genus are mentioned in local con-

servation texts. Pseudopentameris obtusifolia (under the

name of Pentameris obtusifolia) is listed by Hall et al.

(1980) as a threatened species, the conservation status of

which is given as ‘uncertain’. However, owing to the no-

menclatural difficulties experienced with this taxon (dis-

cussed below and in Barker 1993) and the lack of voucher

citations, the exact identity of this taxon is not certain.

Pseudopentameris brachyphylla is considered by Bond &

Goldblatt (1984) to be rare, as it is restricted to the Cale-

don area. Both these species are restricted to relatively

small areas at a low to mid altitude. Therefore, although

not presently under threat, their narrow habitat range

places them in a category that may best be described as

‘potentially threatened’.

PHYLOGENETIC RELATIONSHIPS

The genus Pseudopentameris now contains four spe-

cies. These species are united by an unusual fruit mor-

phology and the presence of pseudostigmata (described

above). This fruit morphology is rare in the southern Af-

rican Arundineae. The only other arundinoid taxon with

a similar fruit type is Cliaetobromus Nees, the fruit of

which are somewhat smaller, but similar in all other re-

spects except the possession of pseudostigmata (Barker

1994). The close relationship between these two genera

is further supported by DNA-based molecular studies

(Barker et al. in prep.). As Pseudopentameris has both

anatomical and morphological autapomorphies (the pres-

ence of inflated abaxial epidermal cells and the presence

of the pseudostigmata on the apex of the ovary) the genus

is distinct from Chaetobromus, and the two genera ought

not to be combined. Autapomorphies for Chaetobromus

include spikelets which disarticulate below the glumes and

dimorphic florets. In Pseudopentameris , and many other

arundinoid genera, the spikelets disarticulate above the

glumes. Chaetobromus is the only arundinoid genus which

has dimorphic florets: the basal ones are glabrous, the up-

per ones variously hirsute.

Affinities between Pseudopentameris and other 2-flow-

ered arundinoids, Pentameris and Pentaschistis, have also

been postulated (e.g. Ellis 1985b; Ellis & Linder 1992). Pen-

tameris is presently defined by fruit characters (an achene

with an apical crown of hairs; Barker 1993), whereas Pen-

taschistis, as presently defined, has no obvious autapomor-

phy, and may include both Pentameris and Prionanthium

Desv. (H.R Linder pers. comm.). While the presence of 2-

flowered spikelets may indicate some degree of relationship

of Pseudopentameris to these genera, other features such as

the sclerophyllous type of arundinoid leaf anatomy suggest

that these relationships extend beyond these taxa to genera

such as Merxmuellera. The assessment of phylogenetic re-

lationships of Pseudopentameris and other arundinoid taxa

therefore needs to be done in the broadest possible context,

and should include all taxa and not be restricted to the south-

ern African genera. Such a study is presently under way

(H.R Linder et al. in prep.).

TAXONOMY

Pseudopentameris Conert in Mitteilungen der Bo-

tanischen Staatssammlung, Miinchen 10: 304 (1971). Type

species: Pseudopentameris macrantha (Schrad.) Conert.

Plants perennial; caespitose, scandent or sometimes

branched. Stems 0.4—1 ,4(-2) m long, woody and persistent

or herbaceous from woody base; flowering culms up to

0.8 m long. Leaves', sheaths appressed to culm, persistent;

sheath mouth sometimes auriculate; ligule a fringe of

hairs; leaf blades linear, 25-500 mm long, rigid or soft,

open and flat to rolled and rigid. Inflorescence paniculate,

lanceolate and somewhat contracted. Spikelets 6-80, lat-

erally compressed, 2-fiowered, highly reduced third floret

Bothaiia 25,2 (1995)

145

FIGURE 3. — The distribution of

Pseudopentameris spp. in the

southwestern Cape, South Af-

rica. A, P. obtusifolia; B, P.

brachyphylla’, C, P. macran-

tha; D, P. caespitosa.

rarely present; glumes two, more or less equal, 1 8-60 mm

long, 3-7(-9)-nerved, chartaceous, minutely scabrid or

glaucous; lemma body chartaceous to cartilaginous, vari-

ously pubescent. 4.5-11.0 mm long, 7-9-nerved, nerves

anastomosing into awn base and 8-27 mm long lateral

bristle; lemma lobes 1 .5^1.5 mm long, acute to acuminate,

partly to fully adnate to bristles; central awn geniculate,

scabrid, contorted basally, 5-17 mm long from base to

knee. 12-34 mm long from knee to tip; palea longer than

lemma body, bifid at apex. Lodicules two, glabrous,

sparsely to densely ciliolate. Stamens three, 4.5-9.0 mm

long. Ovary glabrous, with apical pseudostigmata, styles

short to absent. Fruit a caryopsis, 4—6 mm long, hilum

% as long as fruit, embryo % as long as fruit, surface

reticulately sculptured.

Key to species

la Lower glumes 18-25 mm long, 3-nerved basally, central nerve

extending the length of glume; lodicules densely ciliolate

1. P. obtusifolia

lb Lower glumes (27— )35 — 60 mm long and (3-)5-9-nerved;

lodicules sparsely ciliolate or glabrous:

2a Lemmas pubescent over entire abaxial surface . .2. P. brachyphylla

2b Lemmas basally glabrous, apically pubescent, including

lemma lobes:

3a Lemma body 5-8 mm long; awn (knee to tip) 1 1-18 mm

long; lemma bristle 9-15 mm long; plants with

branched above-ground stems, especially in older

plants 3. P macrantha

3b Lemma body 8-10 mm long; awn (knee to tip) 17-27

mm long; bristle 15-30 mm long; plants with woody

or fibrous underground base, branched above-ground

stems absent 4. P. caespitosa

1 . Pseudopentameris obtusifolia (Hochst.) N.P. Barker

Danthonia obtusifolia Hochst. in Flora 29: 120 (1846); Stapf: 536

(1899). Pentameris obtusifolia (Hochst.) Schweick.: 91 (1938); N.R

Barker in Gibbs Russell et ah: 252 (1990). Type: Pr. b. spei, Ludwig s.n.

(OXF, iso.(?)!).

Avena rigida Steud.: 482 (1829), non A. rigida M. Bieb. (1808). Dan-

thonia rigida Steud.: 243 (1855). Type: Pr. b. spei, Ludwig s.n. (OXF, holo.!

or iso.!).

Pentameris squarrosa Stapf: 515 (1899); Linder & Ellis; 91-103

(1990). Type: Caledon Div., Nieuw Kloof in Houw Hoek Mountains,

Burchell 8076 (K, holo.; PRE, iso.!).

Pentameris sp. nov., Ellis: 567 (1985b); Barker: 136 (1989).

Plants perennial; scandent, sometimes branched. Stems

branching, 0.4-1. 5(— 3.0) m long, woody and persistent,

flowering culms produced annually from apical growth

points. Leaves : sheaths appressed to culm, persistent;

sheath mouth slightly bearded; leaf blades linear, 25-120

mm long, somewhat rolled and rigid. Inflorescence pani-

culate, lanceolate and contracted. Spikelets 8-45, laterally

compressed, 2-flowered; glumes two, more or less equal,

18-25 mm long, chartaceous, minutely scabrid, 3-nerved

basally, central nerve extending to glume apex; lemma

body uniformly pubescent, 4. 5-6.0 mm long, 9-nerved,

nerves anastomosing into awn base and a 8-12 mm long

lateral bristle; lemma lobes 1. 5-3.0 mm long, acute to

acuminate, partly to fully adnate to bristles; central awn

geniculate, scabrid. contorted basally, 7.5-11.0 mm long

from base to knee, 12-16 mm long from knee to tip; palea

longer than lemma body, bifid at apex. Lodicules two,

densely ciliolate, cilia as long as lodicule body. Stamens

three, 4. 5-7. 5 mm long. Ovary glabrous, with apical

pseudostigmata, styles short to absent. Fruit 4 mm long.

Flowering time November to January. The known distri-

bution of this species is indicated in Figure 3A.

Schweickerdt (1938) points out that Steudel (1829) and

Hochstetter ( 1 846) probably described new taxa from dupli-

cate material collected by Baron von Ludwig. Hochstetter

called his species Danthonia obtusifolia Hochst., whereas

Steudel named the material Avena rigida Steud. Schweick-

erdt showed that the name Avena rigida had been used earlier

by Marschall von Bieberstein (1808) and therefore Steudefs

later homonym is rejected. The Ludwig specimen at OXF

bears the name Avena rigida , and is considered to be the

146

Bothalia 25,2 (1995)

holotype or perhaps an isotype of this name. The specimen

seen by Hochstetter was not traced, so the same specimen

(Ludwig specimen at OXF) is considered to be probably

the isotype of the name Danthonia obtusifolia. Hochstetter’s

(1846) D. obtusifolia has priority over Stapf’s (1897) I).

squarrosa. Schweickerdt notes that had Stapf seen the type

material on which Steudel had based his description, he

would have been able to identify his species as Danthonia

obtusifolia Hochst.

The type of Stapf’s Pentameris squarrosa is a Burchell

specimen, Burchell 8076. There is a fragment of this

specimen in PRE, which originates from Kew. The enve-

lope is annotated by Schweickerdt, who wrote 'a fair

match of the type of Avena rigida Steud. in the Herb.

Fielding, Oxford". This fragment, consisting of two leaves

and two spikelets, comprises sufficient material to allow

comparison on the basis of a number of characters. It is

a good match of the type specimen of Avena rigida. It

can therefore be confirmed that Pentameris squarrosa is

a synonym of Pseudopentameris obtusifolia.

This nomenclatural confusion is reflected in past stud-

ies and curatorial practices (cited above). The use of these

confusing names without voucher citations in red data

books (e.g. Hall et al. 1980) results in further confusion

and renders the information in such texts meaningless. In

order to assist in the identification and curation of speci-

mens of this taxon, an extended list of voucher specimens

is provided. The herbarium code JF is used for the her-

barium at Jonkerhoek Forestry Station.

Vouchers: N. Barker 300. 331. 332. 333 (PRE); Burchell 8076 (PRE);

Ellis 2342 (PRE); Esterhuysen 29029. 35396 (BOL, PRE); Esterhuysen

35416 (BOL); Haynes 770 (JF. PRE, STE); Hilton-Taylor 350. 351. 352.

353 (PRE); Kruger KR565 (JF); Levyns 10460 (BOL); Macdonald 634

(PRE, STE); Stokoe 7197 (BOL); Taylor 3023 (PRE, STE).

2. Pseudopentameris brachyphylla (Stapf) Conert

in Mitteilungen der Botanischen Staatssammlung,

Miinchen 10: 304 (1971); N.P. Barker in Gibbs Russell

et al. 278 (1990). Type: Locality unknown, Zeyher 1826a

(K, holo. !).

Danthonia brachyphylla Stapf: 520 (1899).

Plants perennial, caespitose, bases woody and fibrous.

Stems absent. Culms 400-750 mm long. Leaves : sheaths

appressed to culm, persistent; sheath mouth slightly

auriculate; leaf blades 45-150 mm long, open and fiat or

folded. Inflorescence paniculate, lanceolate and con-

tracted. Spikelets 8-20, laterally compressed, 2-flowered,

highly reduced third floret rarely present; glumes two,

more or less equal, 27-37 mm long, 5-7-nerved, char-

taceous, minutely scabrid; lemma body cartilaginous, uni-

formly pubescent, 5. 5-6.5 mm long, 7-9-nerved, nerves

anastomosing into awn base and 12-20 mm long lateral

bristle; lemma lobes 2.0-3. 5 mm long, acute to acuminate,

partly to completely adnate to bristles; central awn geni-

culate, scabrid, contorted basally, 7.5-11.0 mm long from

base to knee, 12-20 mm long from knee to tip; palea

longer than lemma body, bifid at apex. Lodicules two,

glabrous or sparsely ciliolate. Stamens three, 5-8 mm

long. Ovary glabrous, with apical pseudostigmata, styles

short to absent. Fruit 4. 5-6.0 mm long. Flowering time

September-October. The distribution of this species is

given in Figure 3B.

The holotype, Zeyher 1826a housed at Kew is accom-

panied by drawings and annotations in Stapf’s hand. Con-

ert (1971) also cites this specimen, and it carries a ‘det.

slip' bearing his signature. Although the type locality is

not known, PRE has a Zeyher collection of the same

taxon, Zeyher 7825(3, collected at Hemel en Aarde, Swel-

lendam. Although speculative, it is possible that these con-

secutively numbered Zeyher specimens were both

collected from this locality.

Vouchers: N. Barker 731 (PRE); Burman 1180 (BOL); De Vos 1401

(STE); Esterhuysen 33258 (BOL, PRE); Williams 2542 (NBG, PRE).

3. Pseudopentameris maerantha (Schrad). Conert

in Mitteilungen der Botanischen Staatssammlung,

Miinchen 10: 304 (1971); N.P. Barker in Gibbs Russell

et al. 279 (1990). Type: Table Mountain above Kirsten-

bosch. White 5518 (PRE, neo.! here designated).

Danthonia maerantha Schrad. in J.A. Schultes: 385 (1824); Trinius:

t. 63 (1827); Pentameris maerantha (Schrad.) Nees: 312 (1832). Type:

P.B.S., Hesse, s.n. (not traced).

Plants perennial, basally woody, caespitose, becoming

shrub-like with age. Stems branched, 0.5-1. 4 m long, cov-

ered in persistent leaf sheaths. Culms 0. 5-0.8 m long.

Leaves : sheaths appressed to culm, persistent; sheath

mouth somewhat auriculate; leaf blades 45-350 mm long,

open and flat or rolled and filiform. Inflorescence pani-

culate, lanceolate and somewhat contracted. Spikelets

1 0— 25(— 50), laterally compressed, 2-flowered, highly re-

duced third floret rarely present; glumes two, more or less

equal, chartaceous, minutely scabrid or glaucous, 22-50

mm long, 5(-7)-nerved; lemma body cartilaginous, api-

cally pubescent, basally glabrous, 5-8 mm long, 9—1 1 -

nerved, nerves anastomosing into awn base and 9-15 mm

long lateral bristle; lemma lobes 1.5-3. 5 mm long, acute

to acuminate, partly to fully adnate to bristles; central awn

geniculate, scabrid, contorted basally, 5.5-1 1 .0 mm long

from base to knee, 11-18 mm long from knee to tip; palea

longer than lemma body, bifid at apex. Lodicules two,

glabrous or sparsely ciliolate. Stamens three, 5.0-7. 5 mm

long. Ovaty glabrous, with apical pseudostigmata, styles

short to absent. Fruit 4. 5-6.0 mm long. Flowering time

August to September. The distribution of this species is

shown in Figure 3C.

Conert (1971) combined Pentameris maerantha Nees

and Danthonia maerantha Schrad. The Hesse s.n. speci-

men on which Schrader (1824) based his original descrip-

tion of Danthonia maerantha has not been traced, and it

is not known if this specimen was seen by Conert ( 1971 ).

A neotype is designated here in its place.

The absence of a type specimen, and thus accurate flo-

ral measurement data, caused some problems in the allo-

cation of the epithet maerantha to the relevant cluster in

the PCA study. Despite the lack of the type specimen,

Schrader's ( 1824) description includes the spikelel length,

given as 1.5 inches (37.5 mm). As the glumes extend well

beyond any floret parts except the awns, this spikelet

measurement is interpreted to equate to that of the glumes.

As this figure (37.5 mm) is close to the mean glume size

Bothalia 25,2 (1995)

147

(37.14 mm) for one of the two clusters distinguished by

the PCA, this cluster is allocated the epithet macrantha

(Figure 1 ).

In contrast to the paucity of information provided by

Schrader. Trinius (1827) provides an elegant description

of this species, referring to an elongate woody rhizome

and a branched culm. This description is accompanied by

a detailed illustration showing a well-developed rhizome.

These characters clearly match the morphology of the

specimens in the rhizomatous cluster of the PCA, which

Schrader’s description implies represents P. macrantha.

Unfortunately, although citing Schrader's description,

Trinius does not provide a collector and number for the

specimen on which his description or illustration is based,

and mentions no type specimen. Therefore the link be-

tween his clear description and the Hesse specimen on

which Schrader’s description was based is tenuous at best.

The Trinius herbarium is presently housed at St Peters-

burg (Leningrad). The grass collection, including Trinius’s

types and the material on which his illustrations are based,

have recently been databased (R. Soreng, Cornell Univer-

sity pers. comm). Computer records indicate that these

include several specimens of Danthonia macrantha. al-

though the type status of these is not certain. All attempts

to communicate with this herbarium have failed, and thus

the unsatisfactory solution of neotypification has been im-

plemented. The second cluster in the PCA is considered

to be a new species and is described below.

Vouchers: Adamson 1251 (BOL): Boucher 2661 (STE); Ellis 2308

(PRE); Morely 176 (STE); White 5518 (PRE).

4. Pseudopentameris caespitosa N.R Barker sp.

nov. a P. macrantha habitu caespitoso, culmis simplicibus,

partibus floralibus grandioribus, glumis 33-60 mm longis,

arista centrali lemmatis 27-M-5 mm longa differt.

Plants perennial with woody or fibrous basal under-

ground parts, caespitose. Stems absent. Culms annual,

0.7- 1.0 m long, sometimes decumbent. Leaves', sheaths

basal, persistent, often purple, appressed to culm; sheath

mouth somewhat auriculate; leaf blades 45-500 mm long,

open and Hat or rolled. Inflorescence paniculate, lanceo-

late and contracted, upper culm and panicle branches pur-

ple. Spikelets 10-20. laterally compressed, 2-flowered,

highly reduced third floret rarely present; glumes two,

more or less equal, 33-60 mm long, 5-9-nerved, char-

taceous, minutely scabrid or glaucous, dark purple when

young; lemma body cartilaginous, apically pubescent,

basally glabrous, 8-10 mm long, 9- 11 -nerved, nerves

anastomosing into awn base and 15-30 mm long lateral

bristle; lemma lobes 2-A mm long, acute to acuminate,

partly to fully adnate to bristles; central awn geniculate,

scabrid, contorted basally, 10-18 mm long from base to

knee. 17-27 mm long from knee to tip; palea longer than

lemma body, bifid at apex. Lodicules two, glabrous or

sparsely ciliolate. Stamens three, up to 9 mm long. Ovary

glabrous, with apical pseudostigmata, styles short to ab-

sent. Fnut 5-6 mm long. Flowering time October-Novem-

ber. The known distribution of this species is shown in

Figure 3D.

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

(-DC), eastern comer of Farm Buffeljagt, 19-10-1987,

Linder 4362 (BOL, holo.!).

This species is characterised by both its larger floral

structures and its caespitose, unbranched growth form,

hence the specific epithet. The glumes, culms and panicle

branches are often dark purple when young, turning a dark

brown biscuit colour with age, in comparison to the pale

yellow of the glumes of P. macrantha. Although occurring

with P. macrantha in some localities (e.g. Cape Point Na-

ture Reserve, where they have been observed growing

within a metre of each other), P. caespitosa flowers sub-

stantially later than P. macrantha , thus precluding any pos-

sible hybridisation.

In addition to the morphological differences described

above, certain differences in the leaf anatomy have also

been observed. In a phenetic study on a limited number

of anatomical samples. Barker (1990) noted that speci-

mens of what was then named P. macrantha were sepa-

rable into two groups on the basis of three anatomical

characters. One of these groups corresponds to the taxon

here named P. macrantha and has a leaf which is V-shaped

in cross section, with narrow furrows between the ribs

and prickles distributed over the exposed adaxial rib sur-

face. The other group corresponds to P. caespitosa (de-

scribed above) in which the leaf is U-shaped, the furrows

are open and the prickles are restricted to the edges of

the adaxial ribs.

Vouchers: N. Barker 24 (PRE); Boucher & Shepard 4821 (STE);

Ellis 1171. 2515 (PRE); Esterhuysen 31747 (BOL).

ACKNOWLEDGEMENTS

I would like to extend my thanks and appreciation to

Mrs A. Romanowski (National Botanical Institute, Preto-

ria) and R. Carelse (University of Cape Town) for the

photographic work. Dr J. Rourke (NBI. Kirstenbosch) for

the Latin translation and nomenclatural advice, Drs E.R.

Robinson (University of the Witwatersrand). R.P Ellis

(Roodeplaat Grasslands Institute), and Prof. H.P. Linder

(University of Cape Town) for their comments on various

aspects of this work. The curators of B, BOL. G. GRA,

K, NBG, OXF; PH, PRE, S, SAM and STE are thanked

for assisting in the tracing and loan of type material. Most

of this work was undertaken for an M.Sc. study at the

University of the Witwatersrand, and was carried out

while the author was in the employ of the National Bo-

tanical Institute, Pretoria.

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A note on Combretum subgenus Combretum section Macrostigmatea (Com-

bretaceae)

E.F. HENNESSY* and S. RODMAN**

Keywords: Combretum, section Macrostigmatea, southern Africa, taxonomy

ABSTRACT

The history of Combretum section Macrostigmatea, its circumscription and its representation in the Flora of southern Africa

region are provided. A specimen from northern Botswana, Miller B/II99, initially misidentified as C. engleri. is shown to be C.

kirkii, the first record of this taxon in the FSA region. Combretum mkuzense is placed in synonymy in C. zeyheri section

Spathulipetala.

UITTREKSEL

Die geskiedenis van Combretum seksie Macrostigmatea en die omskrywing en verteenwoordiging daarvan in die gebied van

die Flora van suidelike Afrika word voorsien. Daar word aangetoon dat ’n eksemplaar uit noordelike Botswana, Miller B/1199,

wat aanvanklik verkeerd as C. engleri gei'dentifiseer is, C. kirkii is, die eerste rekord van hierdie takson in die FSA-gebied.

Combretum mkuzense word in sinonimie in C. zeyheri seksie Spathulipetala geplaas.

INTRODUCTION

Section Macrostigmatea of Combretum was erected by

Engler & Diels in 1899 to accommodate two taxa, C.

macrostigmateum Engl. & Diels (1899) and C. kirkii M.A.

Lawson (1871). Both taxa have the distal part of the style

expanded, markedly so in C. schumannii , less obviously

so in C. kirkii. In habit and distribution these taxa differ,

C. schumannii being a tree or shrub known from Kenya,

Tanzania, Malawi, Zimbabwe and Mozambique, and C.

kirkii a scandent shrub or liana known to occur only in

the drainage basin of the Zambezi River system and hith-

erto recorded from Zambia, Zimbabwe, Malawi and

Mozambique.

A third taxon, C. engleri Schinz (1901), a multistem-

med virgate shrub, was described from northern Namibia.

This entity has since been recorded in Angola, southwest-

ern Zambia and northern Botswana, growing in substrata

derived from Kalahari sands. Liben (1965) described a

fourth member of the section, C. gillettianum from mate-

rial collected by Glover in northern Zambia. Plants as-

signed to this taxon are small trees or shrubs which to

date have been recorded from Zaire, northern Zambia and

Tanzania.

Of these four taxa, two, C. schumannii and C. engleri ,

(Figure 1 A, B) have Bowers with a glabrous nectariferous

disc, whereas the flowers of C. kirkii and C. gillettianum

(Figure 1C, D) have a pilose disc margin.

Exell (1970) recognized the four taxa enumerated

above as constituting section Macrostigmatea , at the same

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

X54001 , Durban 4000. Present address: Department of Botany, Univer-

sity of Natal, Private Bag X01, Scottsville 3209.

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

X54001. Durban 4000. Present address: 14138 S.E. 238th Lane, Kent,

WA 98042, USA.

MS. received: 1992-11-20.

time expressing the opinion that C. schumannii and C.

engleri might prove to be conspecific, a view shared by

Wickens (1973). Subsequently Exell (1978) formally re-

duced C. engleri to conspecificity in C. schumannii on

the basis that the flowers of both entities have a glabrous

nectariferous disc with a very short (less than 1 mm) free

margin, the fruit of both is glabrous except for peltate

secretory scales and of similar dimensions, and the shrub

habit is common to both.

DISCUSSION

Section Macrostigmatea in southern Africa

Since, of the four entities recognized as species in

1970, only one, C. engleri , has been recorded from the

Flora of southern Africa (FSA) region, it was necessary

to decide whether to follow Exell’s (1978) treatment or

to maintain C. engleri as a species. It was therefore nec-

essary to examine flowering, fruiting and vegetative ma-

terial of C. engleri sensu lato and of C. schumannii sensu

stricto. Of the material labelled C. engleri borrowed from

or seen in situ in southern African herbaria, only a single

specimen. Miller B/1199 from Serondela, Chobe District

in northern Botswana, bore flowers. This specimen, de-

scribed as a straggling shrub, was cited by Exell (1970)

as C. engleri and by Exell (1978) as C. schumannii. When

we dissected flowers we found them to have a pilose disc

margin (Figure 1C), a character which immediately ex-

cluded the specimen from C. engleri and C. schumannii.

Examination of peltate secretory scales of the specimen

(Figure 2A-C) showed that these were structurally similar

to those described (Stace 1969) for C. kirkii and different

from those of a specimen of C. engleri , Maguire 1597 ,

collected in the Caprivi Strip (Figure 3A). Subsequent ex-

amination of peltate secretory scales from type material

of C. kirkii, Kirk s.n. (Figure 2D) and Menyharth s.n.,

1 50

Bothalia 25.2(1995)

FIGURE 1. — Half-flower drawings, x 10: A, Combretum schuman-

nii, Burtt 3826: B, C. engleri, Gossweiler 3241 ; C, C. kirkii ,

Miller B1 199\ D, C. gillettianum, Brenan 8172.

confirmed that, despite its smaller than average flowers,

the depauperate specimen Miller B/II99 should be as-

signed to C. kirkii M.A. Lawson. The locality from which

it was collected, which lies within the drainage basin of

the Zambezi River system, and the straggling shrubby

habit are appropriate for this species. This taxon has not

previously been recorded from the FSA region. It is im-

perative to note that without examination of llowers and

fruit and, in particular, without microscopic examination

of peltate secretory scales at magnification of at least

400 x, it is all too easy to fail to recognize the shrubby

form of C. kirkii. The peltate secretory scales of C. gil-

letlianum (Figure 2E-G) are less divided and mostly

smaller than those of C. kirkii.

All other specimens labelled C. engleri from the FSA

region, all described as shrubs, lacked llowers. Some bore

fruit, some were sterile, but the peltate secretory scales of

all which were examined microscopically were structur-

ally alike and it was therefore concluded that all were

representatives of a single taxon. Specimens of C. engleri ,

among them Gossweiler 3241 (holotype of C. chlorocar-

pum Exell) and Exell & Mendonca 1325 were examined

and found to have scales of the same kind (Figure 2J-M)

as those of specimens labelled C. engleri from the FSA

region (Figure 3A).

Exsiccata of C. schumannii collected in Tanzania and

Malawi, including an isotype, Holst 2375 , were examined

at K and BM. The material of C. schumannii and C.

engleri was compared (Table 1).

Cair (1988) made the significant point that whereas C.

schumannii has been successfully propagated from seed

in cultivation, all attempts to germinate seed of C. engleri

in cultivation to date have failed. All these observations,

together with the fact that as far as one can tell from

existing records there is no overlap in the distribution

ranges of C. schumannii and C. engleri , lead to the con-

clusion that these taxa are not conspecific and therefore

the name C. engleri should be retained, at least until such

time as germination tests have been carried out on material

from Mozambique which is reported to be intermediate

between C. schumannii and C. engleri.

The status of C. mkuzense J.D. Carr & Retief

The reasons for the establishment of C. mkuzense as a

discrete taxon and for its placement in section Macrostig-

matea (Carr & Retief 1989) are unclear since the type

material, Carr 187 (PRE), is patently a specimen of C.

zeyheri Sond. (section Spatlmlipetala). Carr & Retief

(1989) state that the peltate epidermal scales of C.

mkuzense agree well with those of other representatives

of section Macrostigmatea. They fail to mention that the

scales of C. mkuzense and those of C. zeyheri are struc-

turally identical (Figure 3B, C; Table 2), yet they note

that C. zeyheri is present in the area where the specimen

Carr 187 was collected and that the fruit of C. zeyheri

and that of C. mkuzense are alike. They do, however, point

out that C. zeyheri is a small to medium-sized tree,

whereas C. mkuzense is a scrambling shrub. Carr & Retief

in referring to analyses by Rogers of triterpenoid and fla-

vonoid-type compounds of local Combretum spp. (Can- &

Rogers 1987) state that 'examination of the profiles of C.

mkuzense and C. kirkii shows similarities but no signifi-

cant difference’, yet there is no mention in Carr & Ro-

gers’s ( 1987) paper of C. kirkii l They fail to point out that

in the same paper Rogers records that the leaves of C.

mkuzense and C. zeyheri contain exactly the same triter-

penoids and flavonoid-type compounds in the same rela-

tive concentrations. Based on these comments and on

analysis of selected, taxonomically important characters

performed by the second author (Table 2), we conclude

that C. mkuzense cannot be maintained as a discrete taxon,

but must be reduced to synonymy in C. zeyheri Sond.

Combretum Loefl. subgenus Combretum section Macro-

stigmatea Engl. & Diels in Engl., Monographieen afrikan-

ischer Pflanzenfamilien und Gattungen 3: 24 (1899); Stace:

158 (1969); Exell: 176 (1970); Wickens: 21 (1973); Exell:

115 (1978); Stace: 336 (1981). Lectotype: Combretum

FIGURE 2. — Peltate secretory scale heads. A-D, Combretum kirkii : A-C, Miller B/1199: D, Kirk s.n.. E-G, C. gillettianum, Hutchinson & Gillett

3956: H. I. C. schumannii , Burn 3940. J-M, C. engleri : J, K, Gossweiler 3241: L. M, Exell & Mendonga 1325.

macrostigmateum Engl. & Diels = C. schumannii Engl., des-

ignated by Stace (1981).

Small tree, shrub, scandent shrub or liana. Inflorescence

an elongate, subcapitate or capitate spike. Flowers tetramer-

ous; hypanthial tube cupulifonn or broadly infundibuliform,

broader than long; petals ovate to subcircular with entire or

entarginate apex, glabrous; stamens 8, insertion uniseriate

near rim of nectariferous disc; disc concealed within hypan-

thial tube, the narrow free rim glabrous or pilose; style ex-

panded (markedly or slightly) at apex. Fruit sanraroid,

medium-sized to large, 22-55 x 21-50 mm with 4 thinly

chartaceous wings and slender stipe 7-30 mm long; germi-

nation epigeal with cotyledons arising above soil level

(where known); mature peltate secretory scale-heads (35-)

45-85 pm in diameter divided by 8 primary radial walls,

1-several secondary and partial radial walls and several

tangential walls to form 1— 2(— 3) concentric zones of cells.

CONCLUSIONS

Two species of Combretum subgenus Combretum section

Macrostigmatea are currently recorded in tire FSA region, C.

engleri Schinz (which is maintained as distinct from C. schu-

mannii Engl.) and C. kirkii M.A. Lawson which is recorded

in this region for the first time. The entity described by Carr

& Retief ( 1987), C. mkuzense , is shown to be neither a discrete

species nor a member of section Macrostigmatea. but a misi-

dentified specimen of C. zeyheri Sond., section Spathuli-

petala.

TABLE 1. — A comparison of taxonomically important characters of Combretum schumannii and C. engleri

152

Bothalia 25,2 (1995)

FIGURE 3. — Peltate secretory scales on abaxial surface of lamina. A,

Combretum engleri, Maguire 1597: B, C. mkuzense (synonym of C.

zeyheri), Carr 187: C, C. zeyheri , Ward 3165. Scale bars: 10 pm.

SPECIMENS EXAMINED

Combretum schumannii

TANZANIA. — 0438: Lushoto Dist., Simbili, Holst 2375 (K, iso.!).

0636: Mpwapwa (flowering material), Hornby 311 (BM). Grid ref. un-

known, flowering material: Burn 3826, 3940 (BM).

MALAWI. — 1435: Fort Johnston, Burtl 5986 (BM, K).

C. gillettianum

ZAMBIA. — 0831 (Abercorn Dist.): 1.6 k E of Mpulungu, Angus

773A (BM); flowering material, Brenan & Greenway sub Brenan 8172

(K); Mpulungu, Hutchinson & Gillett 3956 (K); flowering material,

Richards 18416 (K).

C. kirkii

MOZAMBIQUE. — Grid ref. unknown: 14°-19°S, Tete, Kirk s.n. (K.

holo.!); Sambesi, Boroma (Chnore), Menyharth s.n. (K, fragment of

holo.! of C. menyhartii Engl. & Diels).

ZIMBABWE.— 1628: Kariba, Goldsmith 23/59 (BM, K).

BOTSWANA. — 1824 (Kachikau): Serondela, Chobe Dist., flowering

material. Miller B/1199 (PRE).

C. engleri

ANGOLA. — Northeastern region, Lunda Province: Biula, Chicoso

R., Exell & Mendonga 1325 (BM); R. Coxi, Exell & Mendonga 1362

(BM). Southern region, Huila Province: Baixo Cunene. Rocadas, ao km

200 da estrada Sa da Bandeira-Rocadas, Borges 287 (BM, K); Distrito

da Huila Rocadas, Centro de Estudos do Cunene, Da Silva 2943 (K);

Gambos, Chibemba no Tchimbolelo, Menezes 569 (K); Mucope, desta

para Bicuar, flowering material, Menezes 3472 (K). Cuando Cubango

Province: between Kuiriri and Kuito, flowering material, Gossweiler

3241 (BM, holo.! of C. chlorocarpum Exell).

NAMIBIA. — 1714 (Ruacana Falls): 8 Meilen siidlich Otjehua, Kaoko-

veld. (-CA), Giess & Leippert 7581a (NBG). 1715 (Ondangwa): bor-

dering Angola near Oshikango, Ovamboiand, about 32 miles (51.2 km)

east at Eenhana, (-BD), Rodin 2669 (K). 1716 (Eenhana): south of air-

strip at Eenhana, (-AD), Roux 151 (NBG); Amboland, Ombalambuenge,

Ondonga, (-CA), Rautanen 236 (BM, photo.! of holo. ex Z). 1719

(Rundu): Runtu, near Okavango River. Okavango Native Reserve, (-DD),

Maguire 1597 (NBG). 1721 (Mbambi): Kavango, 2 km W of Mbambi,

(-CC), Mueller & Giess 585 (K). 1723 (Singalamwe): Wes Caprivi, Oli-

fantkamp, (-CD), Pienaar & Vahrmeijer 483 (K). 1820 (Tarikora):

Okavango Native Territory, 15 m (24 km) W of Nyangana on road to

Runtu, (-BA), De Winter & Marais 4589 (K). 1821 (Andara): Okavango

Native Territory, 14.8 m (23.7 km) E of Nyangana Mission Station,

(-AA), De Winter & Marais 4778 (K); Okavango Native Territory, An-

dara Mission Station, (-AB). De Winter 4242 (K); Grootfontein-Nord/Ca-

privizipfel, an der Pad von Andara nach Bagani, (-AB/BA), Merxmueller

& Giess 2002 (BM). 1918 (Grootfontein): Neitsas, Diinen, (-BC), Dinter

668 (BM, fragment of syn.! of C. myrtillifolium Engl.); Neitsas, Diinen,

Dinter 7278 (BM, isosyn.!); Hereroland, Otjituuo Reserve 236, 4 km S

of Otjituuo depot in Omatako Omuramba, (-DA), D. Edwards 04398

(K). 2120 (Rietfontein): Hereroland, Epukiro Go 329 Reserve, 10 km S

of Rooiboklaagte Omuramba along red line fence, (-AA), D. Edwards

04416 (K).

BOTSWANA. — 1821 (Andara): Ngamiland, 25 km NE of Tsodilo Hill,

(-DA/DB), Mueller & Biegel 2299 (K); northern base of second highest of

Tsodilo Hills, Mueller & Biegel 2322 (K); slopes of Mount Female, Tsodilo

Hills, Ngamiland, (-DB), Banks 31 (PRE). 1822 (Kangara): near Samoqoma

lediba, flowering material, (-CD), P.A. Smith 2864 (K). 1823 (Siambisso):

‘rhino thicket’ near Xauna Pan, (-AA), P.A. Smith 322 (K). 1920

(Tsumkwe): 38 km north of Aha Hills, (-DB), Wild & Drummond 6986

(K). 2022 (Lake Ngami): 28 m (44.8 km) W of Mabele Pudi Hills, (-CD),

Blair Rains & Yalala 10 (K); Mabelepudi Hills, (-CD), Story 5093 (K).

C. zeyheri

NORTH-WEST.— 2527 (Rustenburg): ‘Macalisberg’, (-DA/DB),

Zeyher 552 (SAM, iso.!).

KWAZULU-NATAL.— 2632 (Bela Vista): Ndumu Hill, Ndumu

Game Reserve, (-CD), Pooley 479 (NU); Mkonyane, Ndumu Game Re-

serve, Pooley 676 (NH). 2732 (Ubombo): Ingwavuma Hills, (-AA), Strey

8170 (NH); 4 m (6.4 km) south of Ingwavuma, Ward 2048 (NH); Ing-

wavuma Dist., Ward 3165 (NH); Ingwavuma, (-AC), Ward 5661 (NH,

UDW); Mkuze Game Reserve, African staff quarters area, (-CA), Can-

187 (PRE, holo.! of C. mkuzense Carr & Retief); Mkuze Game Reserve,

Msinga Sand Forest, White 10388 (PRE); Mkuze Game Reserve,

(-CA/CB), Ward 3568 (PRE); east side of farm, ‘Shotton 13810', (-CD),

Ward 8793 (PRE).

ACKNOWLEDGEMENTS

The authors thank the keepers and curators of the her-

baria cited for the loan of specimens and for permitting

the use of facilities. Dr J.R. Lawton and Mrs Y. Naidoo

Bothalia 25,2 (1995)

153

TABLE 2. — A comparison of taxonomically important characters of Combretum zeyheri and C. mk.uz.ense. Only personally observed characters

have been used in table construction

of the Electron Microscope Unit at the University of Durban-

Westville for help in producing micrographs, Mrs J.B.M.

Browning of the Botany Department, University of Natal

for valuable assistance with the illustrations, Mrs E.L. van

Hooff and Mr A. Rajh for help in the printing of photo-

graphs, and the Foundation for Research Development

(FRD) for financial support.

REFERENCES

CARR, J.D. 1988. Combretaceae in southern Africa. Tree Society of

Southern Africa, Johannesburg.

CARR, J.D. & RETIEF. E. 1989. A new species of Combretum from

Natal. Bothalia 19: 38-40.

CARR, J.D. & ROGERS, B.C. 1987. Chemosystematic studies of the

genus Combretum (Combretaceae), part 1. South African Journal

of Botany 53: 173-176.

ENGLER, H.G.A. & DIELS, F.L.E. 1899. Combretaceae. In

H.G.A.Engler, Monographieen afrikanischer Pflanzenfamilien

und Gattungen 3: 12-189. Engelmann, Leipzig.

EXELL, A.W. 1970. Summary of the Combretaceae of Flora zambesiaca.

Kirkia 7: 159-252.

EXELL, A.W. 1978. Combretaceae. Flora zambesiaca 4: 100-183. Flora

Zambesiaca Managing Committee, London.

LAWSON, M.A. 1871. Combretaceae. Flora of tropica! Africa 2:

413M-36. Reeve, London.

LIBEN, L. 1965. Note sur quelques Combretaceae ( Combretum Loeffl.,

Terminalia L. et Strephonema Hook. f. ) du Congo, du Rwanda et

du Burundi. Bulletin du Jardin Botanique de VEtat, Bruxelles 35:

167-184.

SCH1NZ, H. 1901. Combretaceae. In H. Schinz, Beitrage zur Kenntnis

der afrikanischen Flora (Neue Folge). Bulletin de FHerbier Bois-

sier , Ser. 2,1: 877-879.

STACE, C.A. 1969. The significance of the leaf epidermis in the taxon-

omy of the Combretaceae U. The genus Combretum subgenus

Combretum in Africa. Botanical Journal of the Linnean Society

62: 131-168.

STACE. C.A. 1981. The significance of the leaf epidermis in the taxon-

omy of the Combretaceae: conclusions. Botanical Journal of the

Linnean Society 8 1 : 327-339.

WICKENS, G.E. 1973. Combretaceae. In R. M. Polhill. Flora of tropical

East Africa: 1-99. Crown agents for Overseas Governments and

Administrations, London.

Bothalia 25,2: 155-172(1995)

New records and new species of Asclepiadaceae from Namibia

P.V. BRUYNS*

Keywords: Asclepiadaceae, Namibia, new records, new species, taxonomy

ABSTRACT

Eleven species of Asclepiadaceae not previously recorded from Namibia are discussed. Of these, Brachystelma codonan-

thum Bruyns, B. recurvatum Bruyns and Schizoglossum saccatum Bruyns are new species. The new combinations Brachy-

stelma gymnopodum (Schltr.) Bruyns (for Ceropegia pygmaea Schinz) and B. schultzei (Schltr.) Bruyns [for Tenaris schultzei

(Schltr.) E. Phillips] are proposed and the delimitation of Brachystelma R. Br. against Ceropegia L. and Tenaris E. Mey. is

discussed. Notes are also included on little-known species, and the species of Gomphocarpus R. Br. in Namibia are enumerated.

UITTREKSEL

Elf spesies van die Asclepiadaceae wat nog nie voorheen in Namibia aangeteken is nie, word bespreek. Hiervan is

Brachystelma codonanthum Bruyns, B. recurvatum Bruyns en Schizoglossum saccatum Bruyns nuwe spesies. Die nuwe

kombinasies Brachystelma gymnopodum (Schltr.) Bruyns (vir Ceropegia pygmaea Schinz) en B. schultzei (Schltr.) Bruyns [vir

Tenaris schultzei (Schltr.) E. Phillips] word voorgestel en die omgrensing van Brachystelma R. Br. teenoor Ceropegia L. en

Tenaris E. Mey. word bespreek. Notas oor min bekende spesies word ook ingesluit en die spesies van Gomphocarpus R. Br. in

Namibia word opgenoem.

INTRODUCTION

Several collecting trips to Namibia since 1987 have

brought to light species of Asclepiadaceae not previously

recorded from Namibia. Searches through the material

preserved in the National Herbarium, Windhoek (WIND)

have also revealed a few already recorded but not dealt

with in Huber (1967).

The recently published list of Namibian plants (Kol-

berg et al. 1992) has various errors in the account of the

Asclepiadaceae. Some of these are rectified here. Keys

are provided to the taxa where possible. The very con-

fused state of the taxonomy of the complex of genera

around Asclepias does not allow for an exact statement

as to the generic placing of all the species. This problem

is being addressed by Nicholas (pers. comm. 1994) and

once this has been sorted out it should be possible to con-

struct keys to these genera. The genera are arranged al-

phabetically except for those in the Schizoglossum

complex which are all discussed together.

1. ASCLEPIAS L.

Apart from the larger, shrubby species of Asclepias ,

which are here placed under Gomphocarpus , no others

are listed by Huber (1967). However, at least two collec-

tions of small, herbaceous species are known from Na-

mibia. Their identification poses somewhat of a problem

as generally species of the narrow-leaved, herbaceous type

in Asclepias , as with Pachycarpus E. Mey., are known

from the eastern side of southern Africa only (Nicholas

* Bolus Herbarium, University of Cape Town, Private Bag, Rondebosch

7700.

MS. received: 1994-12-02.

1982: 450). Since the genus Asclepias is now intended to

be restricted to the American species (see discussion under

Gomphocarpus ), these will all be moved to other genera

in due course.

Key to the herbaceous species of Asclepias for Namibia

la Leaves at least 3 mm broad at middle, plant branching pro-

fusely from base with spreading stems A. randii

lb Leaves not more than 1.5 mm broad at middle, plant usually

single-stemmed, erect A. aurea

1.1 Asclepias aurea ( Schltr ) Schltr.

A. aurea is widespread along the eastern side of south-

ern Africa and also occurs in Zimbabwe (Brown 1 902—

1904). According to Nicholas ( 1982: 452), it does not oc-

cur in the Cape Province but I have seen plants undoubt-

edly belonging to this species south of Kuruman in the

northern Cape (Bruyns 5836, BOL).

There is a small specimen in WIND which is probably

also referable to A. aurea. However, whereas material

from Kuruman fits the description of A. aurea exactly, in

this Namibian collection there are some differences. It is

slightly more hairy than A. aurea (where the leaves are

glabrous except sometimes for a few hairs on the midrib

beneath) and the peduncles are also much shorter. A brief

description of this material follows.

Plant 100-250 mm tall, stems just over 1 mm thick,

erect, with fine hairs in a single line, intemodes 15-20

mm long. Leaves up to 70.0 x 1.1 mm, scantily hairy on

both sides, ± sessile. Peduncle up to 20 mm long, finely

hairy in a single line, bearing 4 flowers. Pedicel 10-12

mm long, very scantily hairy. Corolla about 8 mm diam.,

lobes with few hairs on dorsal surface. Corona lobes about

156

Bothalia 25,2 (1995)

FIGURE 1 . — Asclepias randii, Bruyns 5487. A, portion of plant; B, flower; C, gynostegium with one corona lobe removed; D, half flower, papillae

on inner face of corona lobes clearly visible; E, pollinarium; F, cross-section of leaf, taken near middle. Scale bars: A, 10 mm; B, 2 mm; C,

F, 1 mm; D, 1 mm; E, 0.25 mm.

2.5 mm long, with many papillae inside hood. Otherwise

as in A. aurea.

Specimen examined

NAMIBIA.— 2217 (Windhoek): near Windhoek, (-CA), Seydel4109 ,

coll. Jan. 1965 (WIND).

1 .2 Asclepias randii S. Moore. Figure 1 .

Small herbs belonging probably to this species were

observed among short grasses on a gentle slope in the

Otavi Mountains. According to Brown (1902-1904: 351),

A. randii is known only from central Zimbabwe. However,

the species is very similar to A. modesta N.E. Br. (Nicho-

las pers. comm. 1994) which is known to be widespread

in south tropical Africa with known localities in Tanzania,

Zambia, Malawi and Angola (Brown 1902-1904). The

two seem to be separated mainly by the rather long, erect

apices of the corona lobes in A. randii , which project

somewhat above the gynostegium.

A. modesta was made a synonym of Trachycalymma

pulchellum (Decne.) Bullock (1953: 351), a genus which

Bullock defined mainly by the papillae or ‘papilliform

hairs' on the inner face of the hood formed by the corona.

These papillae are present in both the material illustrated

here of A. randii (Figure ID) and A. aurea but neither

appears to have been referred to Trachycalymma Bullock.

Specimen examined

NAMIBIA. — 1917 (Tsumeb): Otavi Mountains, (-DB), Bruyns 5487

(BOL; K, alcohol only).

2. BRACHYSTELMA Sims

Two new species of Brachystelma are described. The

generic boundaries of Brachystelma against Ceropegia

and Tenaris E. Mey. are discussed and this results in two

new combinations in Brachystelma. A key to these genera

and Macropetalum Burch, is given and also one to all the

species of Brachystelma in Namibia.

2.1 Brachystelma recurvatum Bruyns sp. nov., a

ceteris speciebus lobis corollae pedicellisque longissimis

angustissimis differt. Typus: Namibia, 1917 (Tsumeb):

Otavi Mountains, (-DB), 5-2-1993, Bruyns 5486 (BOL,

holo.; WIND, K, iso.). Figures 2, 3.

Plant with flattened subglobose tuber up to 80 mm

diam., with 1 -several erect simple to branched stems

reaching 60-300 mm tall. Stems 1-2 mm thick, pubescent.

FIGURE 2. — Brachystelma recurvatum , Bruyns 5486 , showing very

slender pedicels and recurved petals.

Bothalia 25,2(1995)

157

FIGURE 3. — Brachystelma recurvatum , Bruyns5486. A, plant, much reduced in length (up to 300 mm tall); B, flower and bud; C, dissected flower;

D, gynostegium; E, pollinarium. Scale bars: A, 10 mm; B, 2 mm; C, 1 mm; D, 0.5 mm; E, 0.25 mm (at B).

green to purplish. Leaves 30-90 x 3-13 mm, narrowly

lanceolate, margins recurved, pubescent on both surfaces,

green, petiole 1.0-1. 5 mm long. Inflorescence extra-axil-

lary, bearing 1-4 successively developing (lowers, pubes-

cent; peduncle 0-2 x 0.5 mm; pedicels 10-16 x 0.25 mm,

purplish. Corolla 7-8 mm diarn.. outside green finely pu-

bescent, inside with fine crinkled white hairs, with very

shallow tube (± 0.5 mm deep) enclosing base of

gynostegium; lobes within greenish black blotched with

very pale yellow (nearly colourless), with ± deltoid basal

portion about 1 .5 mm broad at base then abruptly narrow-

ing into recurved tail ± 6 x 0.5 mm (seen in bud as long

beak), margins recurved. Gynostegium ± 1 .5 mm diani.,

glabrous; outer corona lobes truncate -bifid, pale yellow

with large greenish black patch on inner surface; inner

lobes with dorsal projection confluent with outer lobes

and narrow limb incumbent on anthers nearly equalling

them, with droplets of colourless fluid secreted on backs,

practically colourless (pale yellow).

Brachystelma recurvatum is unlike any other species

known in Namibia, with exceptionally slender pedicels

and corolla lobes. The corolla lobes are strongly recurved

behind the corolla with the narrow tail then sometimes

bent back and partly obscuring the gynostegium, hence

the epithet recurvatum. The corona structure is typical of

the genus with short, truncate-bifid outer corona lobes

and very small inner lobes. Plants were found among

short grasses on a gentle slope between dolomite outcrops

growing in reddish loam. This is the ‘ Tenaris spec, indet

mentioned and illustrated rather poorly in Bruyns (1984)

from a specimen collected west of Outjo (no material

preserved). Since there seems to be some vagueness

about the distinctions between Tenaris and Brachystelma ,

this is discussed separately under 2.4 and 2.5.

2.2 Brachystelma codonanthum Bruyns sp. nov„

a B. dinteri Schltr. cognatum sed (lore campanulato coro-

naque exteriore cupulata, a B. cupulato R.A. Dyer lobis

corollae deltoideis et corona exteriore quinqueloba dis-

cedit. Typus: Namibia, 1918 (Grootfontein): northeast of

Grootfontein, (-BB), Bruyns 5518 (BOL). Figure 4.

Plant arising from a depressed-globose tuber up to 20

x 30 mm. Stem usually solitary, simple, ± erect, 20-150

x ± 1.5 mm. Leaves 11-22 x 9-15 mm, elliptic, tapering

into petiole 1-2 mm long. Flowers in small sessile groups

extra-axillary at nodes, opening successively. Pedicel

spreading, 2-3 x 0.5 mm. Sepals lanceolate, acute, 2 x

0.5 mm. Corolla 5 x 9-10 mm, outside pale green to

brown, inside glabrous obscurely transversely rugose

green to brown with greenish tips, campanulate; tube ±

2.5 x 4.0 mm (across mouth), broadly conical; lobes 3^1

x 3 mm, deltoid, spreading. Gynostegium 1.5 x nearly 3.0

mm, dark brown, glabrous, cupular; outer corona lobes

rising above rim of cup, bifid, narrowing towards apex;

inner lobes cushion-like, arising lower down in cup and

adpressed to backs of anthers near their base, very short

and usually leaving most of anthers uncovered.

Specimens examined

NAMIBIA.— 1819 (Karakuwisa): (-CC), Bruyns 5990 (BOL).1918

(Grootfontein): (-BB), Bruyns 5518 (BOL).

With its solitary, erect stem and elliptical leaves, this

species resembles B. dinteri vegetatively and, when col-

lected, was assumed to be that species. On flowering, it

turned out to have a completely different, campanulate

flower with somewhat cupular corona. In B. dinteri the

outer corona lobes arise between the broad dorsal parts

of the inner lobes which determine the pentagonal shape

of the corona. In B. codonanthum the inner corona lobes

arise on the inside of the cupular outer corona as in B.

cupulatum. From B. cupulatum it is distinguished by dif-

ferently-shaped corolla lobes with rugose surface (oblong

in B. cupulatum , deltoid in B. codonanthum) and the outer

corona cup which is deeply 5-lobed in B. codonanthum

and notched around the apex in B. cupulatum. B. codonan-

158

Bothalia 25,2 ( 1995)

HUURL 4. — Brat hysiclina codonanthum, Bruyns 55/<V. A. portion ol plant; B, flower; C. dissected flower; D, gynoslegium; E, pollinarium. Scale-

bars: A, 10 mm; B, 2 mm; C, 1 mm; D, 0.5 mm; E, 0.25 mm.

thum was found in two places northeast of Grootfontein

growing in deep white sand in open forest.

2.3 Brachystelma blepharanthera (Schltr. ) H.E. Huber

This small species of Brachystelma was not dealt with

in Bruyns (1984) but, alongside that account, Muller

(1984) recorded its ‘rediscovery’ in 1983 after 70 years

since Dinter’s original collections in 1909 and 1910. Ac-

tually it had been collected long before this in 1968 by

C. Grabow near Gobabis, but this and another collection

by Woodman from north of Okahandja remained uniden-

tified at WIND until recently.

More intensive collecting over southern Namibia has

revealed that this species is widely distributed in the

southern half of the country from Gobabis and Okahandja

southwards to Helmeringhausen. In the southern parts ii

is the only known species of Brachystelma (Figure 5).

Unlike other species, B. blepharanthera grows in arid

places on pebbly or stony ground usually in the open or

among short bushes (sometimes even among Rhigozum

trichotomum). From Gobabis to Windhoek and south

wards to Nauchas it was mainly seen in spots with a

scanty cover of small bushes with the ground densely

strewn wilh quartz and other small pebbles. Several times

it was observed growing with Lithops pseudotruncatella.

Around Helmeringhausen il was usually found on stony,

north-facing mountain slopes.

The tuber of B. blepharanthera is usually cylindrical

to irregularly top-shaped (Figure 6A) or very occasionally

flattened discoid. The largest seen was ± 80 x 50 mm and

several up to 80 mm long and only 20-30 mm thick were

found. This shape is unique in Brachystelma. It is often

tightly wedged among stones and can be difficult to ex-

tract. In ihe northern part of the distribution, plants seen

were rarely more than 50 mm tall above the soil with the

branches spreading horizontally up to about 100 mm. Fur-

ther south, the plant may be erect wilh stems up to 300

mm tall. The cylindrical stems are almost always finely

mottled with purple and the small leaves are usually crin-

kled (as in the largest plant in Figure 6A).

Generally in this species the sepals are green to

brownish, the corolla tube is white (sometimes lightly suf-

fused with pink), the lobes are green (sometimes with pur-

plish margin), the inner corona lobes are green whereas

the rest of the gynostegium is white. In the northern part

of the range the corolla lobes vary from 2^4 mm long.

1'IGUKE 5. Dlsirihuiion ol Brachystelma hlcpluiranthera,9 ; and

Pachycctrpus schweinfurthii, A, in Namibia.

Bothalia 25,2(1995)

159

FIGURE 6. — A, B, Brachystelma blepharanthera, Bruyns 5818\ A, three

plants excavated to show peculiar elongated tubers, usually crin-

kled leaves clearly visible in plant with largest tuber: B, plant with

the flowers are nodding (Figure 6B) and the club-shaped

inner corona lobes are widely divergent to almost hori-

zontally spreading. Towards the base the inner surface of

the corolla lobe is usually covered with papillae each

topped by a fine hair. Around Maltahohe plants were col-

lected with flowers held erect and with the base of the

corolla lobes entirely smooth within. Near Helmering-

hausen the flower was nodding once more (Figure 7K),

the corolla lobes were much longer (5 mm) and narrower,

the papillae with hairs were again present but the corona

lobes, though still club-shaped, were much less thickened

towards their tips and were connivent rather than widely

diverging (Figure 7L).

Specimens examined

NAMIBIA.— 2116 (Okahandja): (-BA), Woortman 221 (WIND).

2118 (Steinhausen): (-DC), Grabow 19 (WIND). 2216 (Otjimbingwe):

(-DB), Van Vuuren s.n. (WIND); (-DD), Bnivns 5626 (K). 2217 (Wind-

hoek): (-CA), Van Vuuren s.n. (WIND). 2218 (Gobabis): (-BA), Bruyns

5818 (WIND). 2316 (Nauchas): (-AD), Pehlemann 1334 (WIND);

(-CB), Bruyns 5639 (BOL). 2416 (Maltahohe): (-DD), Bruyns 5658

(BOL). 2516 (Helmeringhausen): (-BC), Bruyns 5681 (BOL); (-CD),

Bruyns 5703 (BOL); (-DC), Bruyns 5734 (BOL).

Brachystelma gymnopodum , a new name for

Ceropegia pygmaea Schinz

The genera Brachystelma and Ceropegia have similar

distributions around the perimeter of the Indian Ocean

from southern Africa to Australia. The only recent regional

treatments of both genera have covered southern Africa

(Dyer 1980) and Australasia (Forster 1988; Bruyns & For-

ster 1989). None of these accounts deals with the problem

of the boundaries between the two genera which is not

as insignificant as this neglect may imply. Dyer (1980)

followed Brown ( 1907-1909) in separating the two genera

as follows:

la Corolla tubular with the tube usually 2 to several times as

long as its diameter at middle Ceropegia

lb Corolla opening nearly flat or tube campanulate, when tubular

then less than twice as long as its diameter at middle . .

Brachystelma

Many genera within the Stapelieae exhibit variation in

the corolla from nearly rotate through campanulate to tu-

bular (cf. Stapelia leendertziae N.E. Br., Tridentea cho-

anantha (Lavranos & Hall) L.C. Leach, Echidnopsis

mijerteina Lavranos as extreme examples within their re-

spective genera) and it is therefore not surprising to find

this variation in Brachystelma. However, the tubular-flow-

ered species of Brachystelma (such as B. oianthum Schltr.)

may begin to resemble Ceropegia as it is defined above.

Similarly some species may have been placed in Ceropegia

because of this single character while their closest relatives

are in Brachystelma.

The generic position of Ceropegia pygmaea is of con-

cern here. It is clear from Brown (1902-1904), Huber

(1957) and Dyer (1980) that C. pygmaea occupies an iso-

lated position in Ceropegia , where its very long corolla

tube places it according to the above definition.

In general the gynostegium of Ceropegia has a shal-

lowly cupular outer corona with the usually bifid lobes

only slightly overtopping the bases of the inner lobes. The

dorsiventrally flattened inner lobes are incumbent on the

backs of the anthers and then rise up, connivent, in a col-

umn above the centre of the style head. Whereas exactly

this inner corona structure is found randomly in several

genera, but is useful there only to separate species [cf.

Lavrania cactiformis (Hook.) Bruyns and L. picta (N.E.

Br.) Bruyns, (1993)], it is remarkably reliable in separating

Ceropegia from its allies. As far as I am aware there are

only three species out of about 160 where this is not

found: C. pygmaea (southern Africa), C. loranthiflora K.

Schum. (Ethiopia: own unpublished drawings of material

at K) and C. dorjei C.E.C. Fisch. [Bhutan: illustrated in

Huber (1957) but gynostegium possibly based on a juve-

nile flower].

The gynostegium in Brachystelma on the other hand,

has an extremely variable outer corona: from slender bifid

lobes to a tube containing the whole gynostegium. The

inner corona lobes are invariably small and often only just

touch the backs of the anthers near their base. In a few

cases they exceed them, rising up connivent in a column

above the style head as in Ceropegia: B. blepharanthera ,

B. dyeri K. Balkwill, B. gracile E.A. Bruce, B. pachypo-

diutn R.A. Dyer and B. schizoglossoides (Schltr.) N.E. Br.

are the only examples known to me. In all these the co-

rolla tube is shorter than it is broad and so these species

separate out (according to Dyer 1980) clearly as Brachy-

stelma, where their closest relatives are as well.

160

Bothalia 25,2 (1995)

In Ceropegia pygmaea the outer corona forms a deep

cup hiding the rest of the gynostegium (Figure 8G, H).

The inner corona lobes are short, touching the backs of

the anthers and do not exceed them (Figure 8K). As can

be seen, the corolla is very variable in length, ranging

from twice its maximum breadth with the mouth up-

turned (Figure 8B) to about five times its maximum

breadth (Figures 8C; 9) with the mouth turned down to-

wards the soil and usually touching it or even becoming

slightly buried among leaf litter and loose sand [exactly

as observed in Echidnopsis ballyi (Mam. -Lap.) P.R.O.

Bally, (Bruyns 1988)].

In C. pvgmaea the leaves vary from broadly ovate

to linear. Narrow-leaved forms are difficult to distin-

guish when sterile from Brachystelma stenophyllum

(Schltr.) R.A. Dyer and, in fact, vegetatively these two

are very similar: both are small plants arising from a

flattened-globose tuber with short stems branched just

above ground level, producing a cluster of leaves usually

close to the ground with flowers developing on short, hori-

zontal peduncles near the base of the stems. Their

gynostegia are almost identical. A similar corona structure

is also found in B. cupulatum and B. brevipedicellatum

Turrill. These four species possibly form a group with

corolla increasing in tubularity from the nearly rotate

state in B. brevipedicellatum via B. stenophyllum (illus-

trated in Bruyns 1984) to the deep tube of C. pygmaea.

Whereas C. pygmaea occupies an isolated position in

Ceropegia, its close affinities to B. stenophyllum suggest

that it would be much better accommodated in Brachy-

stelma and it is therefore transferred to that genus. How-

ever, it cannot be called Brachystelma pygmaeum because

of the earlier and validly published Brachystelma pyg-

maeum (Schltr.) N.E. Br. Therefore a new combination is

required:

FIGURE 1 .—Brachystelma blepharanthera. A, D, H, K, flowers; B, E, I, L, dissected flower showing gynostegium from side; F, face view of

gynostegium; J, half flower; C, G, M, pollinaria. Scale bars: A, D, H, K, I mm (at A); B, E, F, I, J, L, 0.5 mm (at B); C, G, M, 0.25 mm (at

C). A-C, Bruyns 5626 ; D-G, Bruyns 5659; H-J, Bruyns 565<S; K-M, Bruyns 5703.

Bothalia 25,2 (1995)

161

FIGURE 8. — Brachystelma gymnopodum ( Ceropegia pygmaea). A-C, flowers; D-F, corolla dissected; G, H, side view of corona; I, J, face view of

gynostegium; K, dissected gynostegium; L, M, pollinaria; N, O, hairs in corolla tube. Scale bars: A, B, D, E, F, 3 mm (at A); C, 5 mm: G. H,

I, J, 1 mm (at G); K, 1 mm; L, M, 0.25 mm (at L); N, O, 0.5 mm (at N). Drawn from: A, D, G, I, K, L, N, Bricyns 2078 , Gravelotte, eastern

2.4 Brachystelma gymnopodum (Schltr.) Bniyns ,

comb. nov. Ceropegia gymnopoda Schltr. in Bulletin de

l'Herbier Boissier 4: 450 (1896). Type; Namibia, Rait-

tanen 82 (Z).

Ceropegia pygmaea Schinz (1888). Type: Namibia, Schinz 147 (Z).

Figures 8, 9.

Tenaris E. Mey. in Namibia? A new combination

in Brachystelma for Tenaris schultzei (Schltr.)

E. Phillips

Brown (1902-1904) listed three species of Tenaris

from tropical Africa (two from Somalia and one from Tan-

zania) and commented that There is little beyond habit to

distinguish this genus from Brachystelma.' . In the key he

used the Terminal raceme or lax panicle’ of flowers to

distinguish these species from Brachystelma. Brown

(1907-1909) listed four species of Tenaris for South Af-

rica. In this account he used characters of the flower ‘co-

rolla tube very short; lobes free, not reflexed’ etc. to

distinguish Tenaris from Brachystelma but it is not clear

how this could succeed in separating these genera, for

there are many species of Brachystelma to which this

would apply equally well. It appears that T. chlorantha

Schltr., where there is no Terminal raceme or lax panicle',

may be the reason for Brown’s abandoning his earlier dis-

tinctions for Tenaris.

The seven species listed in Brown (1902-1909) appear

to share the following characteristics;

Flowering stems slender, erect, unbranched, glabrous;

leaves linear 1-2 mm broad or filiform, blade narrower

than midrib on flowering stems, glabrous; peduncle and

FIGURE 9. — Brachystelma gymnopodum. Bniyns 5512. Plant with

broad leaves and very long corolla touching ground at anthesis.

162

Bothalia 25,2 (1995)

pedicel glabrous; exterior of corolla glabrous; follicles

glabrous. Some of these characters are found in species

of Brachystelma but no Brachystelma has all of them.

Another species, Tenaris schultzei (Schltr.) E. Phillips was

added in 1941. Here, although the plant often has only a

single stem and linear leaves 1-2 mm broad, the stem may

branch above and the flowers are not borne on long, slender,

unbranched stems. The entire plant is pilose. Schlechter

(1914) intended describing this species as a Brachystelma

but decided, after a more thorough examination of the

gynostegium, that the long, connivent inner corona lobes

were more similar to those in Ceropegia than Brachystelma.

However, the corolla was entirely different from Ceropegia

and hence he placed it in a separate genus Kinepetalum

Schltr. Phillips’ (1941) placing of this species in Tenaris was

not justified by any discussion and, indeed, if it is to be

accepted, then it is unclear how Tenaris should be defined.

Dyer (1975), partly following Brown (1907-1909), charac-

terised Tenaris by ’Corolla tube almost obsolete, with linear,

linear-spathulate or sometimes filiform lobes, slender erect

herbs with tubers’. However, ‘corolla tube almost obsolete’

and ‘slender erect herbs with tubers’ would fit many species

of Brachystelma and T. schultzei [as illustrated in Bruyns

(1984)] would not fit comfortably into ‘slender erect herbs’.

Also ‘linear’ or ‘filiform lobes’ would fit some species of

Brachystelma as well (e.g. B. recurvation , B. gracile).

Schlechter was in favour of segregating many species

from Brachystelma but most of his segregate genera were

abandoned by Brown (1902-1904). Bullock (1953)

seems to have intended reviving some of these segregates

and started moving species to Dichaelia, but this was

abandoned by Dyer (1980). If one is to make the treat-

ment of this group consistent with that being applied in

the Asclepiadeae, and elsewhere in the Stapelieae, then

one will have to examine Bullock’s statement (1953: 358)

that Brachystelma is unsatisfactorily ‘heterogeneous’ in

some detail. In my opinion Bullock’s statement is not

true except perhaps in respect of the inclusion within the

genus of those species with fusiform roots. These species

differ in their vegetative parts and florally quite substan-

tially from the remainder of the genus. Also, Dichaelia ,

as Bullock envisaged it, is unsatisfactory and it would

be unwise to resurrect the genera Siphonostelma Schltr.

and Kinepetalum which seem to be superfluous.

Nevertheless there appears to be a case for maintaining

Tenaris as distinct from Brachystelma provided it is con-

fined to the seven species of Brown (1902-1904; 1907-1909).

Brachystelma christianeae Peckover (1992) also belongs to

this genus. Furthermore, it needs to be considered carefully

whether Tenaris should be united with the monotypic

Macropetalum which also shares most of the above-mentioned

characters. There seems to be no justification for placing T.

schultzei in Tenaris and it should be moved to Brachystelma.

This change is effected below.

2.5 Brachystelma schultzei (Schltr.) Bruyns comb. nov.

Kinepetalum schultzei Schltr. in Botanische Jahrbiicher 51: 150

(1914). Tenaris schultzei (Schltr.) E. Phillips (1941). Lectotype: Namibia,

Okakuja, Dinter 2528 (SAM).

To summarise the information above, one may separate

the four genera Brachystelma , Ceropegia , Macropetalum

and Tenaris as follows:

la Corolla rotate to campanulate, tube broader than long, if

longer, then inner corona lobes not exceeding anthers:

2a Flowering stems slender, unbranched glabrous; leaves linear

or filiform, 1-2 mm broad, glabrous; exterior of corolla

glabrous:

3a Pedicel glabrous, anthers horizontal, pollinia inserted at

level of top of style head Tenaris

3b Pedicel pilose, anthers erect, pollinia inserted about half-

way up style head Macropetalum

2b Flowers amongst leaves on short often spreading, branched

stems; stems pubescent; leaves pilose, usually with distinct

blade narrowing into petiole and to apex; exterior of co-

rolla pilose [at least one of these must apply, many species

will not satisfy all of them] Brachystelma

1 b Corolla with tube longer than max. breadth; inner corona lobes

exceeding anthers, connivent into column above style

head Ceropegia

The key to Brachystelma in Namibia in Bruyns (1984)

is incomplete, dealing only with the five species discussed

there. There are now 1 1 species recognised from Namibia

and it is necessary to construct a key to them. This key

is somewhat complicated, using characters of the corona

as well as more obvious ones.

Key to Brachystelma spp. in Namibia

1 a Outer corona lobes united above anthers into an entire to 5-lobed cup with inner lobes cushion-like fused to inside of cup ± half-way up:

2a Corolla tube more than twice as long as lobes B. gymnopodum

2b Corolla tube shorter than lobes:

3a Corolla lobes ovate-lanceolate becoming linear above, fused at apices B. stenophyllum

3b Corolla lobes deltoid to oblong, not becoming linear above, free at apices:

4a Corolla lobes oblong 4.5-6.0 mm long, cup formed by outer corona with 5 small notches in margin B. cupulatum

4b Corolla lobes deltoid 3-4 mm long, outer corona cup deeply 5-lobed with each lobe bifid and narrowing towards apex . ... B. codonanthum

lb Outer and inner corona lobes not as above:

5a inner corona lobes much longer than anthers, rising up and connivent above style head or widely separating:

6a Corolla lobes lanceolate-oblong to narrowly deltoid, 2^4 x as long as broad at base, inner corona lobes clavate B. blepharanthera

6b Corolla lobes ± filiform, about 10 x as long as broad at base, inner corona lobes uniformly wide to tips B. schultzei

5b Inner corona lobes shorter than anthers, not rising up above style head:

7a Pedicel 10-35 mm long:

8a Corolla rotate, lobes not recurved, with clavate cilia along margins B. schinzii

8b Corolla with recurved lobes without cilia along margins:

9a Corolla glabrous within, lobes 2-3 x 1 mm B. amotii

9b Corolla pubescent within, lobes ± 7.5 x 1.5 mm B. recurvatum

7b Pedicel 1-5 mm long:

10a Corolla faintly rugose within, lobes ± deltoid (1.0-1. 5 x as long as broad at base) B. dinteri

10b Corolla smooth within, lobes linear to filiform (at least 4 x as long as broad at base) B. circinatum

Bothalia 25,2(1995)

163

3. CEROPEGIA L.

3.1 Ceropegia filiformis (Burch.) Schltr.

The collections cited below are the first known from

Namibia and extend the known distribution of this species

about 450 km to the west. There are no known localities

from the intervening territory between these Namibian

gatherings and that of William Burchell, who originally

discovered it in 1812 between Kloof and Witwater, south-

west of Griquastad in the northern Cape. The distribution

map (Figure 10) has been compiled from material seen

in BOL, NBG, from PRECIS records, collections (2922

BB, 2824 CD) cited by Wilman (1946) and two collec-

tions in cultivation in Cape Town. From the map the ex-

tremely wide and scattered range is clear with the records

from Namibia providing a remarkable case of vicariance.

Most of the other known localities lie on the Great Karoo

in the Cape from Richmond eastwards to Colesburg and

south to Pearston and Somerset East with further unusual

records in the southwest between Sutherland and Fraser-

burg. There is a single collection from the Orange Free

State near Liickhoff. This distribution is unique in the ge-

nus (Bruyns 1985).

In January 1989, after good rains had fallen, several

colonies of plants were found on the eastern side of the

Great Karas Mountains. Plants were scrambling in the

open among stones or climbing on Rhigozum trichotomum

bushes on flat areas on the summit of mountains. On the

western side (seen February 1993) they are also quite

plentiful at higher levels and usually found climbing on

Rhigozum trichotomum.

Specimens of Ceropegia filiformis are extremely in-

conspicuous. The plant usually consists of an unbranched,

thin (about 1 mm thick) cylindrical stem (up to 0.8 m but

usually 150-300 mm long) arising from a dense cluster

of thick, spreading, whitish, fusiform roots. The leaves are

narrow, linear-lanceolate. On the other hand the flowers,

which are 50-60 mm long, are quite striking. Most of the

exterior is greyish mottled with purple. The contrast on

the lobes of white on the medial keel against the dull

purple remainder and striking, narrow, replicate, purple-

black margin makes them easily visible. A peculiar feature

of this deeply coloured margin, which has particularly

large epidermal cells, is the secretion along its length of

regularly spaced droplets of a clear liquid. The function

of this liquid remains unknown. The inside of the basal

inflation of the corolla is white with longitudinal green

stripes on which rows of scattered green papillae are lo-

cated. These stripes continue up into the constricted part

of the tube without the papillae. There are no hairs at all

inside the corolla tube (Figure 11).

The outer corona has unusual, thickened, brownish or

purple-red apices. The rest of the corona is white to yel-

low-white, flecked with purple-red spots, quite densely on

the inner corona lobes, sparsely on the outer lobes. The

raised style-head projecting above the anthers (Figure

11F) is most unusual. This is known in a few Brachy-

stelma species but not in any other Ceropegia.

C. filiformis is unusual among the species with fusi-

form roots, slender non-succulent stems and leaves in that

the stems are cylindrical and not angled. Its affinities

among the other southern African species remain unclear.

Specimens examined

NAMIBIA. — 2718 (Grtinau): Great Karas Mountains, (-BC), ± 1 650

m, Bruyns 5767 (WIND): (-BD), 1 600-1 700 m, Bruyns 3523 (BOL,

WIND); (-DA), ± 1 600 m, Bruyns 5806 (WIND).

NORTHERN CAPE.— 3123 (Victoria West): (-DD), Tyson 410

(BOL); (-CA), Bruyns 4230 (no specimen). 3124 (Hanover): (-AD),

Bruyns 3275 (BOL); (-CC), Meve 347 (BOL). 3221 (Merweville):

(-AA), Bruyns 4840 (BOL); (-BC), Bruyns 5969 (BOL).

EASTERN CAPE. — 3125 (Steynsburg): (-CC), Bruyns 5060 (no

specimen). 3224 (Graaff-Reinet): (-BD), Bruyns I960 (NBG). 3225

(Somerset East): (-AB), Bruyns 1767 (NBG); (-BA), J.R. James (BOL).

3.2 Ceropegia dinteri Schltr.

Dinter first collected this species around Rehoboth,

where it has not been seen again. Several gatherings have

been made from west of Outjo to Grootfontein (Bruyns

1984). The four collections listed below extend the known

distribution over 400 km to the north and make it possible

that this rare and inconspicuous species occurs in Angola.

Plants were found on dolomite and granite hillsides

growing between rocks or in stony bare patches under

trees. Tubers were observed from 25-60 mm in diameter.

The stem (1.0- 1.5 mm thick) is frail and spindly but

reaches a height of over 500 mm with leaves up to 100

mm long. If the plant is growing in the open among short

grasses, the stem is short — the stem in Figure 12 was

about 150 mm long. Flowers are exceptionally variable

with no two the same in size, shape or colouring. The

differences between flowers illustrated in Dyer (1983: t.

17) and the present Figures 12 & 13C give some idea of

this variability.

Specimens examined

NAMIBIA. — 1713 (Swartbooisdrif): (-AC), 1 200-1 500 m, Bruyns

4092 (BOL); (-DA), 1 170 m, Bruyns 4095. 1813 (Opuwo): (-DD), Bruyns

5548 (BOL); south of Kaoko Otavi, 18-02-1993, Bruyns 5557 (K).

164

Bothalia 25,2(1995)

FIGURE 1 1 . — Ceropegiafiliformis. A, plan!; B, bud; C-E, flower; F, l.s. of inflated base of flower: longitudinal ridges of papillae; G, papillae from within basal

inflation of flower. H-L, gynostegium: J, one corona lobe removed; L, top, outer corona and one inner corona lobe removed, anthers and ± elongated

style-head. M, N, poll inaria. Scale bars: A, 10 mm; B-F, 3 mm (at C); G-K, 1 mm (at H); L, 0.5 mm; M, N, 0.25 mm (at H). A, B, C, 1, L, Bmyns 3275;

D, K, N, Bmyns 3523 ; E, J. M, Bmyns I960', F, G, H, Meve 347.

Bothaliu 25,2 (1995)

165

FIGURE 12. — Ceropegia dinteri , Bruyns 5557. Flower: corolla tube 41

mm long.

4. GOMPHOCARPUS R. Br.

According to Bullock (1952: 406) and Nicholas &

Goyder (1992), the genus Asclepias should be confined

to species from the New World since the type species, A.

syriaca L., is an American species. Most of the larger,

shrubby species from Africa placed in Asclepias by Brown

should be referred to Gomphocarpus R. Br. The majority

of the names have already been transferred to Gompho-

carpus and it is only necessary now to decide which spe-

cies occur in Namibia.

In Kolberg et al. (1992), Asclepias buchenaviana and

A. fruticosa appear under Asclepias whereas Gomphocar-

pus tomentosus is listed under Gomphocarpus. If Bul-

lock’s system is to be used then clearly this is inconsistent

and all three should appear under Gomphocarpus as in

Bullock (1952). Three other species are also known from

Namibia and are listed neither in Huber (1967) nor Kol-

berg et al. (1992).

A key is provided for all the species now known for

Namibia and, since the synonymy of some is rather com-

plicated, this is listed and discussed briefly.

Key to Gomphocarpus in Namibia

la Leaves not more than 2 mm broad, follicles 4—6 mm broad

when mature G. filifonnis

lb Leaves broader than 4 mm, follicles usually much more than

15 mm broad when mature:

2a Plant glabrous in all parts, leaves at least 25 mm broad with

rounded lobes clasping stem at base G. glaucophyllus

2b Plant pubescent or puberulous at least on young shoots and

leaves, leaves less than 20 mm broad at max., subsessile

to shortly petiolate without stem-clasping basal lobes:

3a Leaves oblong-lanceolate to elliptic, usually rounded at

base, coriaceous and somewhat rigid G. cancellatus

3b Leaves linear or linear-lanceolate, acute, tapering gradually

into petiole, thin:

4a Follicle without bristles on exterior, produced into apical

beak at least 25 mm long G. rosiratus

4b Follicle covered with soft bristles, beak not more than

15 mm long:

5a Young parts of stem, leaves and inflorescences densely

white-tomentose (plant grey-green)

G. tomentosus

5b Young parts of stem, leaves and inflorescence not

densely white-tomentose (plant yellow-green) . .

G. fruticosus

4.1 Gomphocarpus cancellatus (Bunn, f) Bruyns ,

comb. nov. Asclepias ccincellata Burnt, f., Flora Indica:

cui accedit series zoophytorum indicorum nec non pro-

dromus florae capensis: 7 (betw. 1 March and 6 Apr.

1768).

A. rotandifolia Miller (1768).

A. arborescens L.: 216 (1771). Gomphocarpus arborescens (L.) W.T.

Aiton: 79 (1811).

A. pubescens L.: 215 (1771).

Brown (1907-1909) was incorrect in using the name

A. rotundifolia Miller as this was published on 16 April

1768, slightly after A. cancellata. Several other names ex-

ist for this plant, of which only Asclepias arborescens L.

has been transferred to Gomphocarpus.

G. cancellatus is common in the southwestern Cape

and even occurs in places as a roadside weed. In

Namaqualand and the Richtersveld it is rare and there

appears to be only a single record of its occurrence in

Namibia, some distance north of Rosh Pinah but well

within the winter rainfall zone.

Specimen examined

NAMIBIA. — 2716 (Witpiitz): (-DA), Bruyns 3927 (BOL).

4.2 Gomphocarpus filiformis ( E . Mey.) Dietr..

Synopsis plantarum 2: 900 (1839-1852). Lagarinthus fili-

formis E. Mey.: 203 (1836-1838). Asclepias filifonnis (E.

Mey.) O. Kunze: 268 (1886) non Jacq. (1760).

Asclepias buchenaviana Schinz: 261 (1888). A. filifonnis var.

buchenaviana (Schinz) N.E. Br.: 336 (1902-1904).

The name Asclepias buchenaviana has become famil-

iar. The species was first described as Lagarinthus filifor-

mis by Meyer in 1837 but this name cannot be used in

Asclepias because of Asclepias filifonnis Jacq. (1760), a

species from the Caribbean. However, there is no reason

for it not to be used in Gomphocarpus.

4.3 Gomphocarpus fruticosus (L.) W.T. Aiton. Hoitus

kewensis (1810-1813).

Asclepias fruticosa L.: 216 (1753).

166

Bolhalia 25,2 (1995)

4.4 Gomphocarpus glaucophyllus Schltr. 19(1 894).

Asclepias glaucophylla (Schltr.) Schltr.: 455 (1896).

Rare, only found in the northeastern portion of Namibia.

Specimen examined

NAMIBIA. — 1718 (Nkurenkuru): (-CD), De Winter 3881 (WIND).

4.5 Gomphocarpus rostratus {N.E. Br.) Bullock in

Kew Bulletin: 410 (1952).

Asdepias rostrata N.E. Br.: 331 (1902-1904).

A widespread species in tropical Africa known from

Kenya, Tanzania (Bullock l.c.), Angola and Botswana

(Brown l.c.) and from two collections in Namibia.

FIGURE 13. — Ceropegia dinteri. A, plant; B, cross-section of leaf taken more or less in middle; C, flower; D, base of corolla dissected; E, F,

gynostegium; G, pollinarium. Scale bars: A, 10 mm; B, I mm; C, 3 mm; D, 2 mm; E, F, 0.5 mm; G, 0.25 mm. F drawn from Bruyns 2062 ,

Outjo, remainder from Bruyns 4095.

Bothalia 25,2(1995)

167

Specimens examined

NAMIBIA. — 1719 (Rundu): (-DC), De Winter & Marais 5037

(WIND). 1724 (Katima Mulilo): (-DD), Killick c£ Leistner 3153

(WIND).

4.6 Gomphocarpus tomentosus Burch., Travels 1 :

453 (1822). Asclepias burchellii Schltr. : 336 (1895).

Bullock (1952: 406) regarded this taxon as a synonym

of G. fruticosus but, as Huber (1967) pointed out, two

distinct species appear to be involved.

5. PACHYCARPUS E. Mey.

Smith (1988) revised the southern African species of

Pachycarpus, recognising 24 species for the region. All

of them seem to occur only on the eastern side of southern

Africa and neither there nor in Huber (1967) are any spe-

cies mentioned for Namibia. Nevertheless, as early as

1955, B. de Winter gathered a plant clearly identifiable

as a Pachycarpus in northern Namibia. Recent collections

by B. Strohbach and myself have again brought to light

the same taxon.

5.1 Pachycarpus schweinfurthii (N.E. Br.) Bullock

According to Bullock (1953), P. schweinfurthii is very

widely distributed in tropical Africa from Nigeria and Su-

dan southwards to Angola and Zimbabwe. In Namibia it

only occurs in the moister northeastern region (Figure 5)

and its occurrence in Botswana is also likely.

I have attempted to insert P. schweinfurthii into Smith’s

( 1 988) key to the southern African species of Pachycarpus

(see below) by adding an extra couplet, and a description

is also provided. Unfortunately the drawings in Smith’s

paper are not clear and it is hard to see what the characters

used in the key really refer to, especially as there is no

explanation of the terminology adopted for the different

floral parts. In the specimen here the guide rails (anther

wings of Smith) are a little over 3 mm long. The distance

from the edge of the rail to the centre of the flower in-

creases as one goes from the corpusculum towards the

base of the flower (clearly seen in Figure 14D) and I

assume that this is what Smith means by ‘outer margin

concave’. I have changed this to ‘with concave margin’

as there is no ‘inner’ margin which the use of 'outer' im-

plies.

It may be of use to observe here that the ‘corona lobes’

referred to above and throughout Smith (1988) are the

‘staminal corona’ lobes of Kunze (1982) corresponding to

the ‘inner corona lobes’ of the Stapelieae. It seems to be

assumed generally that in the species in the Asclepias -

Gowp/iocwpits-Pachycarpus-Schizoglossum complex the

corona only consists of these five lobes of staminal origin.

However, in most species 1 have looked at, there are five

more lobes which are of interstaminal origin (Kunze

1982). Unlike in the Stapelieae (where these ‘outer corona

lobes’ are generally well developed), here they are minute

and mostly hidden behind and below the large mouth at

FIGURE 14. — Pachycarpus schweinfurthii, Bruyns 5484. A, flower; B, gynostegium; C, gynostegium with one corona lobe removed; D, half flower;

E, pollinarium. Scale bars; A. 6 mm; B. 3 mm; C, D. 2 mm (at C); E, 0.5 mm.

168

Bothalia 25,2 (1995)

FIGURE 15. — Pachycarpus schweinfurthii , Bruyns 5484 , in habitat.

the base of the guide rail so they have probably been

missed. They are clearly visible in Figure 14C, each lobe

divided into two narrow, widely diverging teeth encircling

the base of the guide rail.

Part of key taken from Smith ( 1988)

19a Guide rail (anther wings) 1. 3-2.4 mm long with ± straight

margin P. natalensis

19b Guide rail 3. 0-7. 2 mm long with concave margin:

20a Flower rotate-campanulate, corolla lobes spreading but

curved inwards in upper half P. schweinfurthii

20b Corolla lobes reflexed:

20ba Keel of corona lobe taller than broad P. reflectens

20bb Keel of corona lobe not taller than broad . . P appendiculatus

Plant to 450 mm tall, consisting of 1 -several ascending

stems, all parts stiffly pubescent except where indicated oth-

erwise (Figure 15). Stems 3-5 mm thick. Leaf 30-10 x 20-50

mm, ovate to ovate-lanceolate, apex rounded, base truncate

to cordate, green with many prominent parallel veins, petiole

4—10 mm long. Peduncle 20-55 mm long, ± 2 mm thick,

bearing 4—10 ± simultaneously opening flowers, bracts nar-

rowly lanceolate, 6-14 mm long, pedicels 15-18 mm x

1.0-1. 5 mm thick. Sepals 10-12 x 2-3 mm (at base), wid-

ening very slightly above base then lanceolate, pubescent

below, glabrous above. Corolla 30-35 mm diam., rotate-

campanulate; lobes 15x9 mm, ovate, finely pubescent out-

side and on one margin, glabrous within, white suffused with

reddish outside, inside white with conspicuous greyish red

veining. Gynostegium 14-15 mm across; outer corona lobes

minute, bifid and spreading adpressed to base of guide rails;

inner corona lobes ascending, ± 7 mm long, exceeding

height of style-head, with keel 2 mm tall, just equalling style-

head, keel and pail below it white, part above keel pinkish

red.

Specimens examined

NAMIBIA. — 1917 (Tsumeb), Otavi Mountains, (-DB), Bruyns 5484

(BOL, K). 1920 (Tsumkwe): (-BC), Strohbach 1268 (WIND).'

6. SCHIZOGLOSSUM E. Mey. sensu lato in Namibia

During recent expeditions to the northern parts of Na-

mibia several plants belonging to the former genus Schi-

zoglossum have been collected. These have proved very

difficult to identify to species level and information on

these collections is presented here in the hope that they

may add to the knowledge of the group. This group was

worked on extensively by Kupicha (1984) who, following

Bullock (1952), resurrected the genus Aspidoglossum E.

Mey. and created a new genus Miraglossum Kupicha as

well as concurring with Bullock’s resurrection of

Stenostelma Schltr. and Glossostelma Schltr. However, she

provided no keys to all these genera and various lengthy

diagnoses have to be perused to sift out the salient details

for identifications.

In Kolberg et al. (1992) various species were left er-

roneously in Schizoglossum with Kupicha cited as the

authority for this. In fact she placed S. baumii Schltr.

ex N.E. Br. under Aspidoglossum masaicum (N.E. Br.)

Kupicha (not S. bidens subsp. atrorubens (Schltr.)

Kupicha) and S. pentheri Schltr. tentatively under Aspi-

doglossum erubescens (Schltr.) Bullock so that Schizo-

glossum sensu stricto is not known in Namibia. Never-

theless, I have included it in the key which follows.

The differentia that Kupicha (1984) gives for Steno-

stelma seem to me to have so much overlap with the

other genera that I have reverted to the shape of the

style-head to separate this genus from the others as

Schlechter (1894) originally did: this works for S. cap-

ense but may cause problems with the other species

tentatively assigned to this genus by Kupicha.

Key to genera of the Schizoglossum complex in Namibia

la Style-head conical, anther appendages much exceeding style-

head connivent in centre and hiding it entirely, caudicles

conspicuously angled, broadly attached to corpusculum .

Stenostelma

lb Style-head flat with slight depression in centre, anther append-

ages adherent to sides of apex and not covering it, caudi-

cles not conspicuously angled with narrow attachment to

corpusculum:

2a Inflorescences pedunculate, pollinia medially attached to

caudicle with germinating mouth on outer side

Schizoglossum

2b Inflorescences sessile, pollinia subapically attached usually

without germinating mouth, if this is present then lo-

cated at point of attachment of pollinium to caudicle:

3a Corona lobes fleshy, pollinia without germinating mouth

Miraglossum

3b Corona lobes thin, pollinia often with germinating mouth

Aspidoglossum

6.1 Aspidoglossum carinatum ( Schltr ) Kupicha

The taxon illustrated in Figure 16 keys out in Huber

(1967) as Schizoglossum pentheri. In Kupicha (1984) one

arrives at Aspidoglossum carinatum (Schltr.) Kupicha rather

than A. erubescens which is where she tentatively placed S.

pentheri in synonymy. The pollinium lacks the conspicuous

germinating zone supposedly typical of A. erubescens and

so we are unlikely to be dealing with that species.

According to Kupicha (1984: 640), A. carinatum oc-

curs in the eastern Cape Province and Natal. The present,

rather exceptional extension of its distribution into north-

Bothalia 25,2(1995)

169

em Namibia stretches credibility somewhat. In addition,

the corona does not correspond very exactly with that of

A. carinatum : the ventral appendage of the corona is slen-

der and much exceeds the apical process (not broadly tri-

angular). However, there appears to be nowhere else to

place this collection and since the taxonomy of this whole

group remains such a mess, I would prefer to avoid de-

scribing new taxa which might further confound the situa-

tion.

It should be observed that the stem here is uniformly

finely pubescent all round and the leaves are pubescent

on the ventral surface and glabrous on the dorsal except

for the pubescent midrib (Figure 16F). Unless damaged,

the plant always consists of a simple, slender, erect stem

up to 300 mm tall with narrow leaves and many small

clusters of nearly sessile flowers. There is a small, neat,

carrot-like tuber just beneath the surface of the soil (one

excavated was ± 60 mm long and 10 mm broad at apex).

Specimen examined

NAMIBIA. — 1917 (Tsumeb): Otavi Mountains, (-DB), Bruyns 5488

(BOL).

6.2 Schizoglossum saccatum Bruyns , sp. nov., a

ceteris speciebus pilis longis densis in lobis corollae et

lobis coronae crassioribus processa apicale horizontale

breve discedit. Typus: Namibia, 1813 (Opuwo), south of

Kaoko Otavi, (-DD), 1 400 m, Bruyns 5558 (BOL, holo.;

K, WIND, iso.).

Plant much branched at ground level from an under-

ground rootstock, with milky latex. Stems 70-200 x

1. 5-2.0 mm, ascending, hairy all round, intemodes 7-25

mm long. Leaves 20-70 x 2.5-4.0 mm, narrowly linear,

sessile, ventral surface pilose, dorsal glabrous except on

midrib, margins recurved, acute at apex and at base. Flow-

ers in fascicles mostly of 4 arising alongside all except

the basal nodes; peduncle 7-20 x ± 1 mm, pilose along

one side only, with several filiform bracts up to 3 mm

long at apex; pedicel 7-9 mm long, about 0.5 mm thick,

pilose. Sepals dorsally pilose, ventral surface glabrous,

4-5 x 1 mm (at base), lanceolate, usually recurved to-

wards apex, green. Corolla divided right to base; lobes

6.0 x 2.5 mm (at base), ovate-lanceolate, ascending-

spreading with recurved margins and apex, dorsal sur-

face glabrous, ventral surface convex and densely

white-pilose especially towards apex with hairs up to

1 .5 mm long, green with faint longitudinal purplish

stripes. Gynostegium 2.0 x 5.5 mm, glabrous, white ex-

cept for purplish pink dorsal apex; outer corona lobes

minute, truncate-emarginate, beneath guide rails; inner

lobes massive, inflated, ± ovoid when viewed from above

with small apical process projecting horizontally onto an-

ther appendage and broad rounded dorsal process. Fruit

and seed not seen. Figure 17.

FIGURE 16. — Aspidoglossum carinatum , Bruyns 5488. A, plant, much reduced in length (up to 450 mm tall); B, flower; C, gynostegium; D. half

flower; E, pollinarium; F, cross-section of leaf taken near middle showing distribution of thickened hairs. Scale bars; A, 10 mm; B. C, F, 1

mm (at B); D, 0.5 mm; E, 0.25 mm.

170

Bothalia 25,2 ( 1995)

FIGURE 1 7. — Schizoglossum saccatum, Bruyns 5558. A, portion of plant; B, flower; C, face view of part of corona; D, gynostegium with one corona

lobe removed; E, half flower; F, pollinarium. Scale bars: A, 10 mm; B, 2 mm; C, D, 1 mm (at D); E, 1 mm; F, 0.25 mm.

This species is only provisionally placed under Schi-

zoglossum with which it shares the pedunculate inflores-

cence but, if Kupicha’s (1984) sense of the genus is

applied, little else. In Miraglossum , where the fleshy co-

rona lobes and lack of germinating mouth on the pollinia

suggest it might belong, it would be a very odd bedfellow

with the other species of this small, compactly distributed

genus: as Kupicha (in litt.) points out, most of them have

whorled leaves in the flowering part of the stem, sessile

inflorescences and pollinia attached to the caudicle some-

what on their inner side rather than apically as in this new

species. It also seems that the fleshiness of the corona

lobes exceeds anything known in Miraglossum. D. Goyder

at Kew has tried to match it with existing material or

existing genera without success.

Plants were common among limestone chunks in a flat

Sesamothamnus- dominated area, growing with a small

spiny Euphorbia and many other succulents. They are

generally small and inconspicuous and, once the grass be-

gins to grow, are probably difficult to find.

6.3 Stenostelma capense Schltr.

This species seems to be widespread but relatively un-

common in northern Namibia. Schlechter (1894) based

Stenostelma on the conical shape of the style-head. From

the half flowers shown here (compare Figure 18H with

Figure ID) it can be seen that the style-head is indeed

differently shaped to that typical amongst the genera As-

clepias , Gomphocarpus, P achy carpus and Schizoglossum.

The anther appendages are also much more conspicuous.

Instead of being adpressed to the edge of the pentagonal

style-head they here only touch the edge, rise up and are

connivent in the centre, in so doing hiding the entire style-

head from view.

In the material illustrated here the stems are pilose in

a longitudinal band on one side (the side changing after

each node) and are not pilose all round as given by Huber

(1967). On the other hand the corolla lobes are papillate

all over the inner surface and not glabrous as Schlechter

described them. The very tiny outer corona lobes consist

of a single, erect tooth beneath the guide rail (Figure 18G).

Specimens examined

NAMIBIA. — 1918 (Grootfontein): Gobis Water, (-AC), Bruyns 5499

(BOL). 1920 (Tsumkwe): 15 km southeast of Tsumkwe, (-DA), Bruyns

5834 (BOL).

7. Stigmatorhynchus hereroensis Schltr.

The genera of the Marsdenieae are so poorly known

that every attempt should be made to obtain and publish

information on them so that an eventual re-assessment of

the tribe is possible.

S. hereroensis is common and fairly widespread in the

northern portion of Namibia usually growing in areas with

much surface limestone. The distribution (Figure 19,

Bothalia 25,2 (1995) 171

FIGURE 18. — Stenostelma capense, Bruyns 5834. A, part of plant: stem virtually glabrous in patch below insertion of each petiole, peduncle only

hairy on upper side. Flowers produced in dense, simultaneously opening clusters on several such peduncles towards apex of stem; B. C, flower;

D, flower with sepals removed; E, flower with two corolla lobes excised; F, gynostegium; G, gynostegium with two corona lobes removed;

H, half flower (small oval area demarked inside cut corona lobe is hollow); I, pollinarium; J, cross-section of leaf, taken near middle. Scale

bars: A, 2 mm; B-E, J, 1 mm (at B); F-H, 0.5 mm (at G); I, 0.25 mm.

drawn up from material in WIND and BOL) seems to be

bounded to the north and east by regions covered with

deep Kalahari sand but it remains possible that it extends

into Angola and Botswana. Plants often grow in the open

but may also be found growing inside other bushes. They

are usually heavily grazed.

S. hereroensis is a shrub-forming species. Plants usu-

ally form a rigid, round bushlet up to about 300 mm tall

but, if sheltered, may reach 1 m in height, although they

never show any tendency to climb. The grey, woody stems

contain a milky latex.

Plants bear small white flowers in nearly sessile, si-

multaneously opening clusters along the stems. Within

each flower is a dense beard of whitish hairs, beginning

on the lobes, extending into the tube and blocking the

throat of the tube. Peculiar to this species (to the genus?)

is the very long, cylindrical style-head which protrudes

slightly from the tube, and the pair of ovules in each carpel.

One of these ovules seems to abort as each pod contains

only a single, quite large seed. The pods may be paired or

solitary and when paired are divergent from one another at

about 120°. The pollinia are very small and attached by long,

slender caudicles to the small corpusculum (Figure 20).

Schlechter (1914) placed three species in Stigma-

torhynchus , of which the other two, S. umbelliferus (K.

Schum.) Schltr. and S. stelostigma (K. Schum.) Schltr.

(from Tanzania and Somalia respectively) are unknown

FIGURE 19. — Distribution of Stigmatorhynchus hereroensis in Na-

mibia (from records in WIND, BOL).

172

Bothalia 25,2 (1995)

FIGURE 20. — Stigmatorhynchus hereroensis, Bruyns 5833. A, portion of plant with paired follicles; B, flower; C, dissected flower; D, gynostegium;

E, half flower, each carpel has two ovules; F, pollinarium with caudicles stretched out. Scale bars: A, 10 mm; B. 2 mm; C, 1 mm; D, 0.5

mm; E, 0.5 mm; F, 0.25 mm.

to me. Bullock (1954) transferred S. stelostigma back to

Marsdenia R. Br. so that, at present, Stigmatorhynchus

contains only two species. It should be noted that

Schlechter’s illustration (1914) is not very accurate: the

pollinaria are very different to what he has shown and the

style head maintains a roughly equal diameter right to the

apex, where it may sometimes be bifid.

ACKNOWLEDGEMENTS

I wish to thank A. Nicholas and D.J. Goyder for useful

information on members of the Asclepiadeae, where my

knowledge is patchy. The Curators of WIND and PRE

are thanked for the loan of material.

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SCHLECHTER, R. 1914. Asclepiadaceae africanae. Botanische

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Bothalia 25,2: 173-181 (1995)

An annotated list of Urediniomycetes (rust fungi) from South Africa 1:

Melampsoraceae and Pucciniaceae, excluding Puccinia and Uromyces

M. VAN REENEN*

Keywords: checklist, new records, taxonomy, Urediniomycetes

ABSTRACT

Up-to-date information is given on the identity, occurrence, nomenclature, host ranges, geographical distribution and

literature references of the Urediniomycetes in South Africa. Species of the family Melampsoraceae and all species of the family

Pucciniaceae other than those in the genera Puccinia Pers. and Uromyces (Link) Unger are listed. New species records since

1945 include the following: Melampsora allii-populina Kleb.. Melampsora coleosporioides Dietel, Melampsora larici-

populina Kleb., Melampsora medusae Thiim., Frommea obtusa-duchesneae (Arthur) Arthur, Hemileia gardeniae-thunbergiae

Maubl. & Roger, Phakopsora gossypii (Arthur) Hirats. f., Uromycladium tepperianum (Sacc.) McAlpine and Tranzschelia

discolor (Fuckel) Tranzschel & M.A. Litv. Although many species of rusts have already been recorded, these are thought to

represent only a small percentage of the rust population in southern Africa. Furthermore, only a small portion of rust specimens

at present housed at PREM have so far been studied. Continued research on this economically important group of fungi is

therefore essential.

UITTREKSEL

Die jongste inligting oor die identiteit. voorkoms, nomenklatuur, gasheerreekse, geografiese verspreiding en literatuur-

verwysings van die Urediniomycetes in Suid-Afrika word gegee, Spesies van die familie Melampsoraceae asook alle spesies

van die familie Pucciniaceae behalwe die van die genusse Puccinia Pers. en Uromyces (Link) Unger word gelys. Nuwe

spesierekords sedert 1945 sluit die volgende in: Melampsora allii-populina Kleb., Melampsora coleosporioides Dietel,

Melampsora larici-populina Kleb., Melampsora medusae Thiim., Frommea obtusa-duchesneae (Arthur) Arthur, Hemilea

gardeniae-thunbergiae Maubl. & Roger, Phakopsora gossypii (Arthur) Hirats. f„ Uromycladium tepperianum (Sacc.) McAlpine

en Tranzschelia discolor (Fuckel) Tranzschel & M.A. Litv. Alhoewel talle spesies reeds aangeteken is, verteenwoordig hulle

vermoedelik slegs ‘n klein persentasie van die roespopulasie van suidelike Afrika; boonop is slegs ‘n klein gedeelte van die

roes-eksemplare wat in PREM gehuisves word, reeds bestudeer. Daarom bly navorsing op hierdie ekonomies belangrike groep

swamme steeds noodsaaklik.

INTRODUCTION

The rust fungi (Urediniomycetes). being obligate plant

parasites, constitute one of the most economically impor-

tant groups of fungi. The rusts in South Africa have been

neglected for some time and our knowledge of the occur-

rence of rust fungi in this country is based largely on the

publication by Doidge (1950). A few subsequent lists of

plant diseases in South Africa (Doidge et al. 1953; Gorter

1977, 1979, 1981. 1982) have provided additional infor-

mation on certain species.

As the records in the National Collection of Fungi rep-

resent mainly early collections up to 1945, catalogued by

Doidge ( 1950), a significant number of new records of gen-

era and species are to be expected. Furthermore, many of

the recorded rusts and their hosts have undergone a change

of name, and several new host records have been reported.

The need for an up-to-date list is therefore evident.

This publication is the first in a series of lists on the

rust fungi recorded for South Africa. It deals with the fam-

ily Melampsoraceae and all genera of the family Puc-

* Former address, to which enquiries should be sent: Mycology Unit,

Biosystematics Division, Plant Protection Research Institute, Private Bag

X 1 34, Pretoria 000 1 .

MS. received: 1993-03-09.

ciniaceae other than Puccinia Pers. and Uromyces (Link)

Unger. These will be dealt with in separate papers, as will

the form-genera Aecidium Pers. and Uredo Pers. The aim

of the present publication is to combine all previous in-

formation in one list. Doidge (1950) included 17 species

under the family Melampsoraceae, 264 species under the

Pucciniaceae and 143 species under the Imperfect rusts.

In this paper nine new species records are added, seven

species previously recorded from South Africa are ex-

cluded as they were based on specimens from outside the

region covered by the present list, five species are placed

in synonymy and one previously incorrectly identified

species is correctly named.

The list covers two families: Melampsoraceae and Puc-

ciniaceae. Species are listed alphabetically. Each name (in

bold type) is given with the relevant literature references,

synonyms (in italics, preceded by =), correct host names

(in italics) and reference to specimens in the National Col-

lection of Fungi (PREM) together with their localities.

Information on the geographical distribution of all species

in South Africa is summarised in Table 1. The geographi-

cal area referred to as South Africa in this publication

includes the Republic of South Africa with the former

Republics of Bophuthatswana, Ciskei, Transkei and

Venda. An alphabetical list with hosts is included in Table

2. An alphabetical list of hosts (Table 3) is also added:

the scientific names are the names listed by the National

174

Bothalia 25,2 (1995)

TABLE 1. — Summary of rust distribution in South Africa (1993)

TABLE 2. — Alphabetical list of rusts with hosts

Bothalia 25,2(1995)

175

TABLE 2. — Alphabetical list of rusts with hosts (continued)

TABLE 3. — Alphabetical list of hosts

Acacia

ataxacantha DC.

burkei Benth.

caffra (Thunb.) Willd.

gerrardii Benth. var. gerrardii

hebeclada DC. subsp. hebeclada

karroo Hayne

luederitzii Engl. var. retinens (Sim) J.H. Ross & Brenan

mearnsii De Wild.

mellifera (Vahl) Benth. subsp. detinens (Burch.) Brenan

nigrescens Oliv.

rehmanniana Schinz

robusta Burch, subsp. robusta

saligna (Labill.) Wendl.

sieberiana DC. var. woodii (Burtt Davy) Keay & Brenan

spp.

Agrimonia

bracteata E. Mey. ex C.A. Mey.

procera Wallr.

Albizia

gummifera (J.F. Gmel.) C.A. Sm.

versicolor Welw. ex Oliv.

Ancylanthos fulgides Welw.

Annonaceae undet.

Artabotrys monteiroae Oliv.

Calpumia aurea (Aiton) Benth. subsp. sylvatica (Burch.) Brummitt

Cant hi um

ciliatum (Klotzsch) Kuntze

mundianum Cham. & Schltdl.

orbiculare Good

spp.

Cassia sp.

Cheilanthes viridis (Forssk.) Sw. var. viridis

Clematis

brachiata Thunb.

oweniae Harv.

Clerodendrum

glabrum E. Mey. var. glabrum

sp.

Coffea arabica L.

Croton

gratissimus Burch, var. gratissimus

sylvaticus Hochst.

Dalbergia

armata E. Mey.

sp.

Dichrostachys cinerea (L.) Wight & Arn. subsp. nyassana (Taub.)

Brenan

Dissotis

incana (Naudin) Triana

longicaudata Cogn.

princeps (Kunth) Triana var. princeps

Duchesnea indica (Andrews) Focke

Elephantorrhiza elephantina (Burch.) Skeels

Euclea

lanceolata E. Mey.

natalensis A. DC. subsp. natalensis

schimperi (A. DC.) Dandy subsp. schimperi

Euphorbia

helioscopia L.

heterophylla L.

kraussiana Bemh. var. kraussiana

striata Thunb. var. striata

spp.

Fabaceae undet.

Fadogia

cienkowskii Schweinf.

tetraquetra Krause var. tetraquetra

sp.

Faidherbia albida (Delile) A. Chev.

Ficus

carica L.

cordata Thunb. subsp. salicifolia (Vahl) C.C. Berg.

Gardenia

volkensii K. Schum. subsp. volkensii var. volkensii

thunbergia Thunb.

Gossypium sp.

Grewia

bicolor Juss.

caffra Meisn.

monticola Sond.

occidentalis L.

spp.

Helichrysum sp.

Hewittea sublobata (L. f.) Kuntze

Hypericum

aethiopicum Thunb. subsp. aethiopicum

aethiopicum Thunb. subsp. sonderi (Bredell) N. Robson

lalandii Choisy

Indigofera sp.

Kohautia amatymbica Eckl. & Zeyh.

Linum

africanum L.

usitatissimum L.

Melinis repens (Willd.) Zizka subsp. repens

Millettia grandis (E. Mey) Skeels

Mundulea sericea (Willd.) A. Chev.

Oldenlandia sp.

Ormocarpum trichocarpum (Taub.) Engl.

Pachystigma

latifolium Sond.

pygmaeum (Schltr.) Robyns

Polypodium lycopodioides L.

Populus

alba L.

deltoides Marshall

nigra L. var. italica

wislizenii Sarg.

spp.

Prunus

armeniaca L.

persica (L.) Batsch

salicina Lindl.

Psydrax livida (Hiem) Bridson

Pteridium aquilinum (L.) Kuhn

Pygmaeothamnus

pygmaeum (Schltr.) Robyns

zeyheri (Sond.) Robyns var. zeyheri

Rhamnus prinoides L’Her.

176

Bothalia 25,2 (1995)

TABLE 3. — Alphabetical list of hosts (continued)

Rhoicissus rhomboidea (E. Mey. & Harv.) Planch.

Ricinus communis L.

Rosa spp.

Rubus

apetalus Poir.

ludwigii Eckl. & Zeyh. subsp. ludwigii

pinnatus Willd.

rigidus Sm.

transvaaliensis Gust

ursinus Cham. & Schltdl.

Rumohra adiantiforme (G. Forst.) Ching

Salix

babylonica L.

mucronata Thunb. subsp. capensis (Thunb.) Immelman

spp.

Securinega virosa (Roxb. ex Willd.) Pax & K. Hoffm.

Senna

petersiana (Bolle) Lock

sinqueana (Delile) Lock

Tephrosia

heckmanniana Harms

macropoda (E. Mey.) Harv. var. macropoda

Tricalysia

capensis (Meisn. ex Hochst.) Sim var. capensis

sonde riana Hiem

Vangueria infausta Burch, subsp. infausta

Vangueriopsis lanciflora (Hiem) Robyns ex Good

Vernonia colorata (Willd.) Drake subsp. colorata

Ziziphus

mucronata Willd. subsp. mucronata

spp.

Botanical Institute (Gibbs Russell et al. 1985, 1987, 1988;

De Wet et al. 1990; Arnold & De Wet 1993).

All publications are listed in which references to rusts

collected in South Africa have been traced. However, ref-

erences to the original descriptions of species first de-

scribed from other countries have been omitted. No

attempt has been made to give a complete bibliography

of literature on the South African rusts.

ACKNOWLEDGEMENTS

The assistance of Alice Baxter and Louise Schutte as

well as the encouragement of the staff of the National

Collection of Fungi, Plant Protection Research Institute

are acknowledged. The National Botanical Institute is

thanked for verifying many of the scientific host names.

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CHECKLIST OF RUST FUNGI RECORDED IN SOUTH AFRICA

Host species are listed in italics.

Names of authors of plant names follow Brummitt & Powell ( 1992). All collection numbers given (followed by locality) are PREM numbers (PREM:

National Collection of Fungi), unless preceded by one of the following codes:

S.A.M.: fungus specimens from the South African Museum Herbarium, now incorporated in PREM;

Stell.: specimens from the Mycological Herbarium of the former Stellenbosch-Elsenburg College of Agriculture, now incorporated in PREM;

Van der Byl: specimens from the P.A. van der Bijl Herbarium, now housed at PREM.

Numbers given in square brackets are duplicate numbers for the preceding specimen.

Other codes used:

* specimens cited in Doidge (1950) which could not be located;

** specimens previously recorded for South Africa which fall outside the geographical area of this publication;

• new species record;

+ type specimen;

= synonym.

MELAMPSORACEAE

GENUS COLEOSPORIUM

clematidis Barclay, P. Syd. & Syd. (1915); Doidge (1927, 1950); Dietel

(1928); Doidge et al. (1953); G.J.M. Gorter (1981)

= Caeoma clematidis Thiim.

= Uredo clematidis Berk.

Clematis brachiata Thunb.: 1861 Barberton; 2359 Kentani; 4383,

10160, 20788, 20795 Somerset East; 10580 Natal

Clematis oweniae Harv.: 801, 10581, 11223, 31906* Inanda

hedvotidis Kalchbr. & Cooke (1880); Sacc. (1888); P. Syd. & Syd.

(1915); Doidge (1927, 1950); G.J.M. Gorter (1981)

Kohautia amatymbica Eckl. & Zeyh.: 36, 1256, 1419, 1456, 2194,

6600, 8908. 9692, 23369. 30944 Pretoria; 8910 Bethlehem;

10067 Mooi River; 10156+ Inanda; 14171 without locality;

40997 East Griqualand

Oldenlandia sp.: 32706 Natal

ipomoeae (Schwein.) Burrill. Doidge (1927, 1950); G.J.M. Gorter (1981)

Hewittea sublobata (L. f.) Kuntze: 1598, 2501, 9100 Winkle Spruit;

6633* Isipingo; 9479 Inanda; 12334 Durban

GENUS CRONARTIUM

gilgianum Henn.. Sacc. & P. Syd. (1890); P. Syd. & Syd. (1915); Syd.

& P. Syd. (1918); Doidge (1927, 1950); Nel (1942)

= Cronartium bresadoleanum Henn.

= Crossopsora gilgiana (Henn.) Syd.

= Cronartium bresadoleanum Henn. var. eucleae

Euclea lanceolata E. Mey.: 1737, 1738, 1739 Soutpansberg

Euclea natalensis A. DC. subsp. natalensis: 1606, 1617, 2415. 9496,

32442 Durban; 2509 Winkle Spruit; 6646 Isipingo; 9089, 9115

Kentani; 10630 Natal; 12420 East London; 26006. 32399,

33601 Nelspruit; 22417 Alexandria; 32273 Kruger National

Park; 32410 Sabie; 33487, 35159, 43119 Grahamstown; 33527

Mtunzini; 33597 Entabeni; 41032 Umtentweni

Euclea schimperi (A. DC.) Dandy subsp. schimperi: 12170** [Van der

Byl 1803]; 729 Nelspruit; 1151. 1852, 5620, 18097 Barberton;

1399 Winkle Spruit; 30959, 30960 Kruger National Park

zizyphi Syd. & E.J. Butler. Doidge (1927, 1950)

Ziziphus mucronata Willd. subsp. mucronata: 32724 between Acom-

hoek and Tzaneen; 41983 Lydenburg

Ziziphus spp.: 50573 Ellisras; 50576 Nelspruit; 50696 Kruger National

Park

GENUS MELAMPSORA

aecidioides (see M. populnea)

allii-populina* Kleb.

Populus deltoides Bart, ex Marshall: 43192 Pretoria

coleosporioides* Dietel

Salix babylonica L.: 50566 Grabouw; 50568, 50569 Stellenbosch

helioscopiae G. Winter, Doidge (1927, 1950); G.J.M. Gorter (1981)

Euphorbia helioscopia L.: 41453 Pretoria

Euphorbia heterophylla L.: 29876, 30979, 32278 Pretoria; 32206 Dur-

ban; 33603 Natal; 40167 Tzaneen

Euphorbia kraussiana Bemh. var. kraussiana: 9087 Claridge

178

Bothalia 25,2 (1995)

Euphorbia striata Thunb. var. striata: 17035 Mooi River; 29720 Pre-

toria

Euphorbia spp.: 1065 Bethlehem; 1851 Barberton

Euphorbia heterophylla L.: 420711, 44620 Pretoria

hvpericorum G. Winter var. australis Doidge (1927, 1950); G.J.M.

Goiter (1981)

= Melampsora hyperici J. Schrot.

Hypericum aethiopicwn Thunb. subsp. aethiopicum : 30079* Hogsback

Hypericum aethiopicum Thunb. subsp. sonderi (Bredell) N. Robson:

1074 Lydenburg, 1302, 1946, 1988, 9314, 9480*, 9691, 32090*

Pretoria; 10361 Inanda; 12241 Mooi River

Hypericum lalandii Choisy: 10362 Natal

junodii** Doidge (1927, 1950)

Vemonia colorata (Willd.) Drake subsp. colorata: 11723+

larici-populina* Kleb., G.J.M. Gorter (1979, 1982)

= Melampsora populina (Jacq.) Lev.

Populus deltoides Marshall: 43691 Pietermaritzburg; 43694 Pretoria

Populus nigra L. var. italica: 43730 George; 43732 Nelspruit; 43833

Pretoria; 48510 Harding

Populus wislizenii Sarg.: 43731 George

Populus spp.: 43692, 47995 Pietermaritzburg; 43911, 50565 Pretoria;

47559 Stellenbosch; 48496 Witbank; 48497 Bethal; 48511

Greytown

lini (Ehrenb.) Lev., Verwoerd (1929); Nel (1942); Doidge (1950); Doidge

et al. (1953); G.J.M. Gorter (1977, 1981)

Linum africanum L.: 46119 (Stell. 150) Stellenbosch

Lirtum usitatissimum* L.

medusae* Thiim., T.N. Trench, A.P. Baxter & H. Churchill (1988)

= Melampsora albertensis Arthur

= Uredo medusae (Thiim.) Arthur

Populus deltoides Marshall: 48253, 50554, 50555, 50556, 50557 Grey-

town; 48329 Ermelo; 48330 Howick; 48331 Pietermaritzburg;

48332 Piet Retief

Populus sp.: 48499 Vryheid

populnea (Pers.) P. Karst., G.J.M. Gorter (1977)

= Melampsora aecidioides J. Schrot.

Populus alba L.: 537 Cape Town; 7102 Soutpansberg; 23723, 44353

Pretoria

Populus spp.: 639 Cape Town; 71 1 Groot Drakenstein; 724 Cape; 733

Inchanga; 1514 Wellington; 21936 Bloemfontein; 29859 Sout-

pansberg; 30780 Rustenburg; 32141**, 34319, 34419, 36791

Pretoria; 45985 (Stell. 24), 46795 (Stell. 879) Stellenbosch

puccinioides* G. Winter (1885); Sacc. (1888); P. Syd. & Syd. (1915);

Doidge (1950)

Helichrysum sp.

ricini Pass., P. Syd. & Syd. (1915); Van der Byl (1926); Doidge (1927,

1950); Doidge & Bottomley (1931); Nel (1942), Doidge et al.

(1953); G.J.M. Gorter (1977)

= Caeoma ricini Schltdl.

= Uredo ricini Biv.

= Melampsorella ricini (Biv.) De Toni

= Uromyces ricini Biv.

Ricinus communis L.: 55, 7402, 14203, 30968*, 30969 Pretoria; 179

Soutpansberg; 524**; 738 Kenilworth; 1013, 20778 Graham-

stown; 10161 Inanda; 10163, 20783, 20789 Somerset East;

11505 Buffelspoort; 23363 Boulders; 32300 Durban; 32309

Eshowe; 45988 Stellenbosch

vitellinae Thiim., Doidge (1927, 1950); G.J.M. Gorter (1981, 1982),

= Uredo mixta Duby forma salicis-capensis Thiim.

= Melampsora mixta (Schltdl.) J. Schrot.

Salix mucronata Thunb. subsp. capensis (Thunb.) Immelman: 923

Boshoff; 20766* (S.A.M. 33935) Somerset East; 50505 Betty’s

Bay

Salix spp.: 45875 Grahamstown; 50492, 50753 Royal Natal National

Park; 50656 Pretoria

GENUS MILESINA

dieteliana (Syd.) Magnus, P. Syd. & Syd. (1915); Doidge (1927, 1950)

= Milesia polypodii F.B. White

= Melampsorella dieteliana Syd.

Polypodium lycopodioides L.: 1634 Amanzimtoti

nervisequa (Syd.) Faull, P. Syd. & Syd. (1915), Doidge (1927, 1950)

= Milesia nervisequa Syd.

= Caeoma nervisequum Thiim.

sub Uredo filicum Klotzsch

sub Uredo polypodii (Pers.) DC.

Cheilanthes viridis (Forssk.) Sw. var. viridis: 999 Pretoria; 1370 Dur-

ban; 1836 Soutpansberg; 10162 Natal; 20781 Somerset East

Rumohra adiantiforme** (G. Forst.) Ching: S.A.M. 28729

GENUS PUCCINIASTRUM

agrimoniae (Dietel) Tranzschel, P. Syd. & Syd. (1915); Doidge (1927,

1950); Dietel (1928)

= Coleosporium ochraceum Fuckel

sub Phragmidium obtusum C. Tul.

Agrimonia bracteata E. Mey. ex C.A. Mey.: 303 Fort Beaufort; 332

Inanda; 1150, 1202, 1261, 1407, 2147, 6655, 29771, 29814,

29847 Pretoria; 2313 Harrismith; 2353 Kentani; 32284 without

locality; 11638 Lidgetton; 11649 Tweedie; 24351, 20323 Sout-

pansberg; 10157*, 10158. 20777, 20797 Somerset East; 25977

Barberton; 32311 Natal

Agrimonia procera Wallr.: 50762 Cathkin Park, 50731 Uitsoek State

Forest

GENUS UREDINOPSIS

macrosperma (see U. pteridis)

pteridis Dietel & Holw.

= Uredinopsis macrosperma (Cooke) Magnus

Pteridium aquilinum (L.) Kuhn; 10590, 11224 Inanda

PUCCINI ACEAE

GENUS CEROTELIUM

fici (Castagne) Arthur, Verwoerd (1929); Doidge (1950); Doidge et al.

(1953)

= Kuehneola fici E.J. Butler

= Uredo fici Castagne

= Physopella fici Arthur

Ficus carica L.: 289 Grahamstown; 615 Constantia; 852 Pieter-

maritzburg; 1207, 25159 Cape; 20726* Cape Town; 1315

Renosterfontein; 46129 [Stell. 161], 46799 [Stell. 882] Stellen-

bosch

Ficus cordata Thunb. subsp. salicifolia (Vahl) C.C. Berg: 176, 1298

Pretoria

gossypii (see Phakopsora desmium)

GENUS DIORCH1D1UM

tricholaenae Syd., Syd. & P. Syd. (1912); Sacc. & Trotter (1925);

Doidge (1927, 1950); Doidge & Bottomley (1931)

Melinis repens (Willd.) Zizka subsp. repens: 269*+, 586 Barberton;

1449 Harden Heights, 1605, 2393, 2512 Winkle Spruit; 1607,

1608 Amanzimtoti; 20631 Natal; 30138 Nelspruit; 30292

Ndwedwe; 30974 Estcourt

woodii Kalchbr. & Cooke in Kalchbr. (1882); Sacc. (1888); Magnus

(1891 ); Doidge ( 1927, 1950); Dietel (1928); Doidge & Bottomley

(1931)

= Puccinia woodii (Kalchbr. & Cooke) Syd.

Millettia grandis (E. Mey.) Skeels: 335+, 10448+, 10449+, 22090 Ton-

gaat; 1593, 1978, 2515 Winkle Spruit; 1601 Amanzimtoti, 5606

Port Shepstone; 11893 Umkomaas

GENUS ENDOPHYLLUM

macowanii Pole Evans (1907, 1908); P. Syd. & Syd. (1924); Van der

Byl (1925, 1926); Doidge (1927, 1950); Nel (1942)

= Aecidium elegans Dietel

= Aecidium rhamni Pers. forma rhamni-prinoides

sub Puccinia coronata Corda

Rhamnus prinoides L’Her.: 379, 171 14 George; 1772, 17738, Sout-

pansberg; 2097 Kentani; 2418, 17215 Knysna; 4275, 20735,

20761 Somerset East; 33152 Donny brook

GENUS FROMMEA*

obtusa-duchesneae* (Arthur) Arthur

= Frommea obtusa (F. Strauss) Arthur

= Frommea obtusa (F. Strauss) Arthur var. duchesneae Arthur

Duchesnea indica (Andrews) Focke: 48512 Greytown; 48609, 48610

Pretoria; 50482 Cathkin Park

Bothalia 25,2(1995)

179

obtusa var. duchesneae (see F. obtusa-duchesneae)

GENUS HAMASPORA

longissima (Thiim.) Korn. (1877); Massee (1892); P. Syd. & Syd. (1915);

Doidge (1927, 1950); Dietel (1928); Doidge & Bottomley (1931 );

Doidge et al. (1953)

= Uredo lucida Thiim.

= Phragmidium longissimum Thiim.

Rubus ludwigii Eckl. & Zeyh. subsp. ludwigii : 9776 Spitskop; 27562

between Rested and Clarens; 32303 Durban

Rubus pinnatus Willd.: 1620 Winter’s Kloof; 2350 Kentani; 2404 Cra-

mond; 30375 Donnybrook

Rubus rigidus Sm.: 144, 1825 Soutpansberg; 2351 Kentani; 5161

Komgha; 8387 Pretoria; 10589, 11055, 11205 Inanda; 20721,

20723*, 20742*, 20746* [S.A.M. 28746] Somerset East; 22413

East London; 26051 Barberton; 32741 Rustenburg

Rubus transvaaliensis Gust: 1267, 1379*, 1380, 1513, 1535*, 1891*.

1914, 1949, 2184, 6656, 6657, 6898, Pretoria; 20333 Sout-

pansberg; 26016 Barberton

Rubus spp.: 29753 Pretoria; 33385 Grahamstown; 46490 [Stell. 561] Cape

Town; 46955 [Stell. 1033] Stellenbosch; 50489 Cathkin Park

GENUS HEMILEIA

aneylanthi** Syd.. P. Syd. & Syd. (1915); Doidge (1927, 1950)

= Uredo aneylanthi Henn.

Ancylanthos fulgides Welw.

canthii Berk. & Broome, P. Syd. & Syd. (1915); Doidge (1927, 1950);

G.J.M. Gorter (1981)

= Coleosporium detergibile Thiim.

= Uredo detergibilis Henn.

Canthium ciliatum (Klotzsch) Kuntze: 20779, 33918 [S.A.M. 33918]

Somerset East

Canthium mundianum Cham. & Schltdl.: 50861 Sandton

Canthium orbiculare** Good

Canthium sp.: 9084 Wyebank

Psydrax livida** (Hiem) Bridson

evansii Syd., Syd. & P. Syd. (1912), P. Syd. & Syd. (1915), Sacc. &

Trotter (1925), Doidge (1927, 1950)

Tricalysia sonderiana Hiem: 6635 Umbogintwini

Tricalysia capensis (Meisn. ex Hochst.) Sim. var. capensis : 590**

fadogiae Syd., Syd. & P. Syd. (1912); P. Syd. & Syd. (1915); Sacc. &

Trotter (1925); Doidge (1927, 1950); G.J.M. Gorter (1981)

Fadogia cienkowskii Schweinf.: 757**, 7391 Nelspruit

Fadogia tetraquetra Krause var. tetraquetra : 34482 Pietersburg

Fadogia** sp.

Pygmaeothamnus pygmaeum (Schltr.) Robyns: 48608 Nelshoogte State

Forest

Pygmaeothamnus zeyheri (Sond.) Robyns var. zeyheri: 281+ Heidel-

berg; 2209, 12229, 26376, 34218 Pretoria; 21232 Newcastle;

28811* Waterberg; 42421 Welverdiend

gardeniae-thunbergiae* Maubl. & Roger

Gardenia thunbergia** Thunb.

Gardenia volkensii K. Schunt. subsp. volkensii var. volkensii: 50708

Kruger National Park

Gardenia* sp.

scholzii Syd., Syd. & P. Syd. (1910); P. Syd. & Syd. (1915); Doidge

(1927, 1950)

= Uredo scholzii Henn.

Clerodendrum glabrum E. Mey. var. glabrum: 1569, 1570 Amanzimtoti

Clerodendrum** sp.

vastatrix Berk. & Broome. J.M. Wood (1883, 1906); P. Syd. (1899);

Burtt Davy (1904); Pole Evans (1907); P. Syd. & Syd. (1915);

Van der Byl (1926); Doidge (1927, 1950); Doidge & Bottomley

(1931); G.J.M. Gorter (1977, 1981)

= Uredo gardeniae-thunbergiae Henn.

Coffea arabica L.: 5, 42, 59, 172, 20322 Tzaneen; 323, 349 Natal; 372

Durban; 409 Pretoria; 991 Lemana; 1403 Winkle Spruit; 6824,

7788 Verulam; 20857 Rietvlei; 21933 Lydenburg; 23142 Louis

Trichardt; 23498 Potchefstroom; 24344* Duiwelskloof; 24941

Pinetown; 25469, 30936* Umbogintwini

woodii Kalchbr. & Cooke (1880); Sacc. (1888); Burtt Davy (1904); J.M.

Wood (1906): Pole Evans (1907), P. Syd. & Syd. (1915); Van der

Byl (1926); Doidge (1927, 1950); Nel (1942); G.J.M. Gorter

(1981)

= Uredo vangueriae Cooke

Pachystigma latifolium Sond.: 173 without locality; 340, 10600, 11185

Inanda, 814, 10549, 10599*, 32021* Natal; 1404, 5640 Winkle

Spruit; 6835 Scottburgh; 8250 Krantzkloof; 9762 New Hano-

ver; 14177 Entumeni; 24940 Brighton Beach

Pachystigma pygmaeum (Schltr.) Robyns: 30909 White River

Vangueria infausta Burch, subsp. infausta: 324, 352, 10410 Natal; 1154,

1277, 1278 Barberton; 1679 Table Mountain; 1815 Sout-

pansberg; 2446 Cramond; 2497 Winkle Spruit; 11845 Louis

Trichardt; 14104*; 30941 Inanda; 32403, 50733 Nelspruit

Vangueriopsis lanciflora** (Hiem) Robyns ex Good

GENUS KUEHNEOLA

albida (see K. uredinis)

uredinis (Link) Arthur, G.J.M. Gorter (1977, 1981)

= Kuehneola albida (J.G. Kiihn) Magnus

Rubus apetalus Poir.: 1532 Lydenburg

Rubus rigidus Sm.: 1863 Barberton; 32287 without locality

Rubus transvaaliensis Gust: 642 Barberton; 992 Wellington; 2417

Knysna; 33173 Pietermaritzburg

Rubus ursinus Cham. & Schltdl.: 46001 Stellenbosch [Stell. 39; Van

der Byl 1101], This specimen was previously wrongly identified

as Phragmidium violaceum (C.F. Schultz) G. Winter.

Rubus spp.: 681 Cape Town; 6931 Stellenbosch; 18087 Pretoria; 32125

Knysna; 50483 Cathkin Park

GENUS MASSEEELLA

flueggeae Syd., Doidge (1939, 1950); G.J.M. Gorter (1981)

= Uredo brideliae (Henn. & Pole Evans) Doidge

= Aecidium brideliae Henn. & Pole Evans

Securinega virosa (Roxb. ex Willd.) Pax & K. Hoffm.: 77 Nelspruit;

25968, 26359 Barberton

GENUS PHAKOPSORA*

desmium (see P. gossypii)

gossypii (Arthur) Hirats. f., Cummins (1962)

= Phakopsora desmium (Berk. & Broome) Cummins

Gossypium sp.: 23190 Natal

GENUS PHRAGMIDIUM

disciflorum (see P. mucronatum)

mucronatum Fr.

= Phragmidium disciflorum (Tode) J. James

Rosa spp.: 139 Kimberley; 141, 1171, 1212, 1892, 32830 Pretoria; 421

Bloemfontein; 484**; 787* Durban; 1866 Wellington; 2430

Uniondale; 9310 Kentani; 15041* Port Elizabeth; 28521 Fort

Beaufort; 34257**; 46022 Stellenbosch

violaceum (R. Schultz) G. Winter (This specimen was wrongly identified

and is now filed under Kuehneola uredinis. Phragmidium vio-

laceum has not been collected in South Africa.)

GENUS PUCCINIOSIRA

dissotidis* Wakef., P. Syd. & Syd. (1924); Doidge (1927, 1950)

= Aecidium dissotidis Cooke

= Uredo dissotidis Cooke, l.c.

= Uredo dissotidisdongicaudae Henn.

Dissotis incana (Naudin) Triana: 10288, 11194 Natal

Dissotis longicaudata** Cogn.

Dissotis princeps (Kunth) Triana var. princeps: 10289 Natal

GENUS RAVENELIA

atrides Syd., Syd. & P. Syd. (1912); P. Syd. & Syd. (1915); Dietel

(1928); Sacc. & Trotter (1925); Doidge (1927, 1948. 1950)

= Dasturella atrides Thirum.

Grewia bicolor Juss.: 32731 Kruger National Park

Grewia caffra Meisn.: 1630, 1670+, 2534 Durban

Grewia monticola Sond.: 32401 Nelspmit

Grewia occidentalis L.: 1629 Amanzimtoti

Grewia spp.: 5075, 5076 Kruger National Park

baumiana Henn. ( 1903); Dietel ( 1906); Sacc. & D. Sacc. (1905); P. Syd.

& Syd. (1915); Doidge (1927, 1948, 1950)

= Haploravenelia baumiana (Henn.) Syd.

Senna petersiana (Bolle) Lock: 29870 Rooiwal; 30955 Schagen; 32404

Nelspmit; 50553 Hanglip State Forest

Senna singueana (Delile) Lock: 6886**+

180

Bothalia 25,2 (1995)

bottomleyae** Doidge (1927, 1950)

Albizia versicolor Welw. ex Oliv.: 14166

deformans** (Maubl.) Dietel (1906); P. Syd. & Syd. (1915); Sacc. &

Trotter (1912); Doidge (1927, 1950)

= Pleoravenelia deformans Maubl.

= Cystingophora deformans (Maubl.) Syd.

Acacia sp.

dichrostachydis Doidge (1927, 1950)

Dichrostachys cinerea (L.) Wight. & Am. subsp. nyassana (Taub.)

Brenan: 6806, 7106 Verulam

elephantorrhizae Doidge (1927, 1950)

Elephantorrhiza elephantina (Burch.) Skeels: 8955+, 12225, 12841,

23453 Pretoria; 23359 Enselberg; 29732* Donkerpoort

escharoides Syd., Syd. & P. Syd. (1912); P. Syd. & Syd. (1915); Sacc.

& Trotter (1925); Doidge (1927, 1950); G.J.M. Gorter (1981)

Faidherbia albida ** (Delile) A. Chev.

Acacia burkei Benth.: 534+ Potgietersrus; 10086 Springbok Flats;

20654 Naboomspruit; 27586 Hartebeespoort; 29918 Pienaar’s

River; 50690 Kruger National Park

Acacia nigrescens Oliv.; 50691, 50695 Kruger National Park

evansii Syd., Syd. & P. Syd. (1912); P. Syd. & Syd. (1915); Sacc. &

Trotter (1925); Doidge (1927, 1939, 1950); G.J.M. Gorier (1981)

= Dendroecia evansii Syd.

Acacia gerrardii Benth. var. gerrardii: 30127 Verulam

Acacia rehmanniana Schinz: 5625 Pretoria

Acacia robusta Burch, subsp. robusta: 123, 149, 2 1 8 1 +, 2212, 5175,

6588, 6698, 14144 Pretoria; 1316 Magalies River; 6807, 7105

Verulam; 18085 Loskop; 28954 Magaliesberg; 29933 Wolma-

ransstad; 30124 Umzinto; 50693 Kruger National Park

Acacia sieberiana DC. var. woodii (Burn Davy) Keay & Brenan: 2539+

Estcourt

Acacia spp.: 2403 Cramond; 11670 Pretoria; 33171 Johannesburg

glabra Kalchbr. & Cooke (1880), Cooke (1880); Dietel (1894, 1906); P.

Syd. & Syd. (1915); Doidge (1927, 1950)

sub Ravenelia slictica Berk. & Broome

Calpumia aurea (Aiton) Benth. subsp. sylvatica (Burch.) Brummitt:

2320, 2330, 5165 Kentani; 2375+, 10696+, I0698+, 20728+,

20732+, 20727, 20728*+, 20851 [S.A.M. 28754] Somerset East

halsei Doidge (1939, 1950)

Acacia ataxacantha DC.: 30117+ Ndwedwe; 50571, 50748 Nelspruit

indigoferae** Tranzschel, Dietel (1894); P. Syd. & Syd. (1915); Doidge

(1927, 1950)

sub Ravenelia laevis Dietel & Holw.

Indigofera sp. 15513 Harare, Zimbabwe

inornata Dietel (1894, 1906); Sacc. (1895); P. Syd. & Syd. (1915);

Doidge (1927, 1950); G.J.M. Gorter (1981)

= Aecidium inomatum Kalchbr.

= Cystotelium inomatum (Dietel) Syd.

= Haploravenelia inornata Dietel

Acacia karroo Hayne: 2368 Kentani; 2377+, 4262*+, 20734+ Somerset

East; 2368*, 2541 Komgha; 22375 Kirkwood; 26396 Graham-

stown

le-testui** Maubl., Dietel (1906); Sacc. & Trotter ( 1912); P. Syd. & Syd.

(1915); Doidge (1927, 1950)

Cassia sp.

macowaniana Pazschke, Dietel (1894, 1928); Sacc. (1895); P. Syd. &

Syd. (1915); Doidge (1927, 1948, 1950); G.J.M. Gorter (1981)'

= Aecidium omamentale Kalchbr.

= Ravenelia ornamentalis Dietel

= Cephatelium macowanianum (Pazschke) Syd.

Acacia karroo Hayne: 329, 834*, 10299, 13074 Ladysmith; 929

Leeuwdoorns; 1417 Kroonstad; 1524, 2123, 21224, 21225,

23449, 23452, 1985, 2095, 2170, 11857, 2255, 2447, 6665,

23203, 23452*, 24913, 28603, 29817, 29798*, 29916, 30912,

36935, 47473, 50686, 50687, 50688, 50692, 50697, 50701,

50699, 50700, 50711 Pretoria; 1915, 8757 Kentani; 2077

Indwe; 2135 Plat River; 2355 Zwartfontein; 2357, 5154 Vry-

heid; 2376+, 4067+, 20730+, 2073 1+, 20733+ Somerset East;

2423 Trapp’s Valley; 2552 Bathurst; 7367* Butterworth, 7404,

28597, 507(H) Onderstepoort; 8245 Muden; 8857 Queenstown;

10017 Riverton; 1 1002 Springbok Flats; 11300 Bloemfontein;

15453 Klipplaat; 21052* Asbury; 27585 Hartebeespoort; 28505

Friedenheim; 28606 Prieska; 30327 Wolmaransstad; 30971 Est-

court; 31021 Ventersburg; 34583 Witbank; 40653 Soutpansberg;

44332 Mafikeng; 46145 Knysna; 46208 Oudtshoorn; 47474

Schweizer Reinecke; 50757, 50758 50759, 50760 Boeken-

houtskloof; 50762 Ashton Bay

minima Cooke (1882); Sacc. (1888); Dietel (1894, 1906); P. Syd. &

Syd. (1915); Doidge (1927, 1950); Doidge & Bottomley (1931);

G.J.M. Gorter (1981)

Albizia gummifera (Gmel.) C.A. Sm.: 351, 10697+ Inanda; 1835*,

1400, 1582, 2384 Winkle Spruit; 6827 Scottburgh; 30779 Dur-

ban

modesta Doidge (1939, 1948, 1950); G.J.M. Gorter (1981)

Acacia hebeclada DC. subsp. Iiebeclada : 30110+ Pretoria

Acacia luederitzii Engl. var. retinens (Sim) J.H. Ross & Brenan: 34572

between Hammanskraal and Pienaar's River

natalensis Syd. & Pole Evans, Syd. & P. Syd. (1912); P. Syd. & Syd.

(1915); Sacc. & Trotter (1925); Doidge (1927, 1950)

= Longia natalensis (Syd. & Pole Evans) Syd.

= Haploravenelia natalensis (Syd. & Pole Evans) Dietel

Acacia karroo Hayne: 1584+, 1935+, 2514 Winkle Spruit

peglerae Pole Evans (1918); Doidge (1927, 1950); G.J.M. Gorter (1981)

Acacia caffra (Thunb.) Willd.: 2331+, 2367, 5179, 5626 Kentani;

2544* Butterworth; 5150 Weenen

pienaarii Doidge (1927, 1950); G.J.M. Gorter (1981)

Acacia caffra (Thunb.) Willd.: 2449. 5627+, 6658*+, 11508, 23450*,

23448, 23451, 26395, 49454, 51013 Pretoria; 5159 Weenen

pretoriensis Syd., Syd. & P. Syd. (1912); P. Syd. & Syd. (1915); Sacc.

& Trotter (1925); Doidge (1927, 1939, 1950)

Acacia sp.: 1376+ Pretoria

stictica Berk. & Broome, Sacc. & P. Syd. (1890); P. Syd. & Syd. (1915);

Doidge (1927, 1950); G.J.M. Gorter (1981)

= Ravenelia munduleae Henn.

Mundulea sericea (Willd.) A. Chev.: 124, 1522, 13089, 23447, 2210,

2256, 2227, 19842, 29728, 30716, 32667, 41959, 43144, 49376,

50558 Pretoria; 146 Hartebeespoort; 28255 Potgietersrus

tephrosiae Kalchbr., Sacc. (1888); Dietel (1894, 1906); P. Syd. & Syd.

(1915); Doidge (1927, 1950); G.J.M. Gorter (1981)

Tephrosia heckmanniana** Harms

Tephrosia macropoda (E. Mey.) Harv. var. macropoda: 10700 Inanda;

14190 Entumeni

transvaaliensis Doidge (1939, 1950)

Acacia mellifera (Vahl) Benth. subsp. detinens (Burch.) Brenan:

37382+ Pretoria

woodii* Pazschke, Dietel (1894); P. Syd. & Syd. (1915); Sacc. (1895);

Doidge ( 1927, 1950)

Fabaceae undet.

GENUS SCHROETER1ASTER

doidgeae Syd. (1926); Doidge (1927, 1950)

Croton gratissimus Burch, var. gratissimus: 14173 Waterberg

stratosus (Cooke) Syd., P. Syd. & Syd.; Doidge (1927, 1950); Doidge

& Bottomley (1931)

= Melampsora stratosa Cooke

Croton sylvaticus Hochst.: 363 Natal; 2020, 2383, 2500, 5639 Winkle

Spruit; 10363 Inanda; 33093 Eshowe; 35148 Louis Trichardt

GENUS SKIERKA

robusta Doidge (1927, 1950); Mains (1939)

Rhoicissus rhomboidea (E. Mey. ex Harv.) Planch.: 1635+ Durban;

1824 Lemana; 2398 Winkle Spruit, 2416 Umgeni Beach; 6628

Isipingo; 32781 Umhlanga Rocks; 39125 Southport

GENUS SPHAEROPHRAGMIUM

artabotrydis Doidge (1941, 1950); Monoson (1974)

Artabotrys monteiroae Oliv.: 30761+ Westville

dalbergiae Dietel; Kalchbr. ( 1882); Sacc. ( 1895); P. Syd. & Syd. ( 1915);

Doidge (1927, 1950); Monoson (1974)

Dalbergia armata E. Mey: 881*, 2524 Umgeni Lagoon; 1371. 5144

Durban; 6825 Verulam; 30954 Nelspruit; 35147 Empangeni

Dalbergia sp.: 51015 Nelspruit

GENUS UROMYCLADIUM*

tepperianum* (Sacc.) McAlpine. M.J. Morris, M.J. Wingf. & J. Walker

(1988)

Acacia mearnsii De Wild.: 48897 Bainskloof

Acacia saligna (Labill.) J.C. Wendl.: 50659 Stellenbosch

Bothaiia 25,2 (1995)

181

GENUS UROPYXIS

(?)gerstneri Doidge (1927, 1950); J.W. Baxter (1959)

Annonaceae undet.: 34564+ Zululand

This is a doubtful species. Examination revealed characters that place

this species in the genus Diorchidium.

steudneri Magnus var. rhodesica** Doidge (1948, 1950); J.W. Baxter

(1959)

Ormocarpum trichocarpum (Taub.) Engl.: 34032 Inyati, Zimbabwe

GENUS TRANZSCHELIA*

discolor* (Fuckel) Tranzschel & M.A. Litv.; Nel (1942)

= Tranzschelia punctata (Pers.) Arthur

= Tranzschelia pruni-spinosae var. discolor (Fuckel) Dunegan

= Puccinia discolor Fuckel

Primus armeniaca L.: 5593 Graaff-Reinet; 30966 Pretoria

Primus persica (L.) Batsch: 1 Elsenburg; 125 without locality; 290,

321 Fort Beaufort; 7800 Pretoria; 45982, 46791 Stellen-

bosch

Primus salicina Lindi.: 1250 Lake Chrissie; 1290 Bethal; 2378 Gra-

hamstown; 48612 Pretoria

Bothalia 25,2: 183-204(1995)

Studies in the Marchantiales (Hepaticae) from southern Africa. 9. The genus

Marchantia and its five local species

S.M. PEROLD*

Keywords: distribution, ecology, Hepaticae, Marchantia, M. berteroana, M. debilis, M. paleacea subsp. paleacea, M. pappeana subsp. pappeana, M.

polymorpha subsp. ruderalis, southern Africa, taxonomy

ABSTRACT

In this, the ninth and final article in the series on southern African Marchantiales, a taxonomic account is given of the genus

Marchantia which is locally represented by five species. Two of the three subgenera that are recognized, namely subgenus

Marchantia (with two species, M. polymorpha subsp. ruderalis and M. berteroana ), as well as subgenus Chlamidium (Corda)

Bischl. which is divided into three sections, namely Paleaceae (with M. paleacea subsp. paleacea ), Chlamidium (with M.

pappeana subsp. pappeana ) and Papillatae (with M. debilis ) are treated. The third subgenus, Protomarchantia Schust., is absent

from the region. Corrections need to be made to the Marchantia species given in Magill & Schelpe's (1979) checklist and in

Arnold & De Wet (1993). Descriptions and illustrations of the taxa together with distribution maps, their ecology and a key to

the subgenera and species are provided.

Only traditional taxonomic methods were employed and this treatment closely follows that of Bischler-Causse (1993a), the

recognized world authority on the group. All southern African specimens held at BOL and PRE, as well as a few from other

herbaria have been studied and some new collections have been added. It is shown that M. polymorpha subsp. ruderalis has been

introduced, as it is only known from local nurseries. The presence here of M. paleacea subsp. paleacea is confirmed, although

it is rare. Otherwise little that is new could be added to the exhaustive studies by Bischler-Causse (1993a).

UITTREKSEL

In hierdie, die negende en finale artikel in die reeks oor die Marchantiales van suidelike Afrika, word 'n taksonomiese

verslag gegee oor die genus Marchantia wat plaaslik deur vyf spesies verteenwoordig word. Twee van die drie erkende

subgenusse, nl. subgenus Marchantia (met twee spesies, M. polymorpha subsp. ruderalis en M. berteroana ) sowel as subgenus

Chlamidium (Corda) Bischl. wat verdeel word in drie seksies, nl. Paleaceae (met M. paleacea subsp. paleacea), Chlamidium

(met M. pappeana subsp. pappeana) en Papillatae (met M. debilis), word behandel. Die derde subgenus, Protomarchantia

Schust.. kom nie in die gebied voor nie. Regstellings moet aangebring word aan die Marchantia- spesies soos aangegee in Magill

& Schelpe (1979) se lys en in Arnold & De Wet (1993). Beskrywings en illustrasies van die taksons sowel as verspreidings-

kaarte, die ekologie en ’n sleutel tot die subgenusse en spesies word verskaf.

Slegs tradisionele taksonomiese metodes is toegepas, en word die werk van Bischler-Causse (1993a), die erkende wereld-

gesaghebbende op die groep, word ten nouste nagevolg in hierdie behandeling. Alle eksemplare uit suidelike Afrika in BOLen

PRE asook ’n paar van ander herbariums is bestudeer en nuwe versamelings is bygevoeg. Daar word aangetoon dat M.

polvmorpha subsp. ruderalis uitheems is, aangesien dit slegs in plaaslike kwekerye voorkont. M. paleacea subsp. paleacea se

teenwoordigheid hier word bevestig, maar dit is skaars. Origens kon min tot die uiters volledige studies deur Bischler-Causse

( 1 993a) bygevoeg word.

CONTENTS

Genus Marchantia L 183

Key to local subgenera, sections and species 184

Subgenus Marchantia L 185

1 . M. polymorpha L 185

2. M. berteroana Lehm. & Lindenb 189

Subgenus Chlamidium (Corda) Bischl 192

Section Paleaceae Bischl 192

3. M. paleacea Bertol 193

Section Chlamidium 195

4. M. pappeana Lehm 197

Section Papillatae Bischl 199

5. M. debilis Goebel 201

Acknowledgements 202

References 202

Specimens examined 203

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

MS. received: 1995-03-27.

Marchantia L., Species plantarum edn 1: 1137

(1753) emend. Raddi, Opuscoli Scientifica di Bologna:

358 (1818); Sim: 27 (1926); S.W. Amell: 53 (1963);

Bischl.: 6(1984); Bischl.: 13 (1989a); Bischl.: 13 (1993a);

R.M. Schust.: 305 (1992). Type species: Marchantia po-

lymorpha L. [lecto. fide Leman: 115 (1823)].

Marchantia M. Marchant f.: 229 (1739).

Marchantiopsis Douin & R.C.V. Douin: 135 (1918).

Chlamidium Corda: 647 (1829).

Thalloid, smallish to medium-sized to large, fiat and

ungrooved, relatively delicate or firm and occasionally

rather leathery, green to dark green or greyish green, along

margins sometimes purplish, rarely all over dorsally and

occasionally also ventrally; dorsal face with distinct po-

lygonal areolae, each with a pore; in crowded mats, some-

times in partial rosettes, hygrophytes, requiring high

humidity and mostly unable to sustain life dry; growing

184

Bothalia 25,2(1995)

on damp soil of vertical stream banks, at waterfalls, dams,

weirs and canals, sometimes on wet rocks, rarely on rotting

wood.

Branches broadly band- or narrowly ribbon-shaped,

pseudodichotomously furcate, without lateral or apical inno-

vations from keel, ventrally thickened medianly and tapered

toward margins, which are lobulate, crenulate, undulate or

entire, apex notched, with median scale appendages recurved

over edge. Dorsal epidermal cells generally unistratose, often

without chloroplasts, walls thin or slightly thickened but lack-

ing trigones; air pores compound, encircled by several su-

perimposed concentric rings of cells, some above epidermis,

others projecting into air chambers below, the latter in a shal-

low, single layer, densely packed with 2^1(5 )-celled assimi-

latory filaments; storage tissue compact, sometimes with a

few sclerotic cells and/or mucilage cavities, also with scat-

tered oil cells each containing a single, large oil body, these

present as well elsewhere in thallus and scales; numerous

rhizoids ventrally, some smooth, others pegged, rarely with

inner thickenings spirally connected. Ventral scales in 4 or

more rows, covering part or all of ventral face and rarely

projecting beyond margins, median scales bluntly triangular,

constricted where joined to appendage, laminal scales

smaller, sometimes in double rows, lacking an appendage,

marginal scales rarely present. Cupules borne dorsally on

thalli, hollowed, contracted below, flaring above, margin cili-

ate, lobed-ciliate, dentate or nearly entire, containing discoid

gemmae that reproduce vegetatively.

Dioicous. Gametangiophores stalked. Stalks arising at

apex of main and/or lateral branches, basally sometimes

surrounded by large scales, generally without an append-

age, along their length bearing filiform scales, 2-4 rhizoid

furrows and 0-2 bands of air chambers with compound

air pores. Antheridiophores with receptacular discs rather

flat, lobed or divided into rays, antheridia and compound

air pores on dorsal side, several rows of scales per lobe

beneath. Arche goniophores with receptacles convex

above, lobed or divided into ± flat rays, dorsally with

compound air pores, below with scales and archegonia,

each one surrounded by calyptra and pseudoperianth, in

groups that are enclosed by bivalved involucres, occurring

alternate with the lobes or rays (in African species).

Sporophytes consisting of foot, short seta (elongating

later) and subglobose capsule, its unistratose wall with

annular thickenings, dehiscing irregularly. Spores small,

thin-walled, ± triangular-globular, ornamentation with nu-

merous thin, irregularly convoluted ridges or with few

wide ridges separated by granules, the 2 faces similar or

dissimilar. Elaters tapering at ends, bi- or trispiral. Chro-

mosome number: n = 9 (basically).

On the basis of differences in spore morphology, ger-

mination patterns and flavonoid patterns, the genus is sub-

divided into 3 subgenera: Marchantia , Chlamidium and

Protomarc hand a R.M. Schust. The latter is absent from

the region.

Key to local subgenera, sections and species

la Thalli with margins plicate-crisped, crenulate or lobulate, dorsally with or without dark median band; epidermal

pores with walls of cells bordering inner opening slightly or strongly projecting and pores then cruciate; scales

in 4-6 rows, covering % or all of ventral surface and then also extending beyond thallus margins; appendages

of median scales orbicular or reniform or ovate, marginal cells slightly or strikingly smaller than inner cells;

laminal scales much wider than long; cupules with ciliate lobes at margins, externally with papillae; male

receptacles peltate, shallowly dissected into 6-10 broad lobes; membranous margins of rays crenulate or entire

and then with very small cells; female receptacles deeply dissected into 9-11 terete rays, with or without

papillae; stalks with single broad band of air chambers; margins of involucres with ciliate lobes; spores 8-16

pm in diameter, ornamentation on 2 faces similar, with numerous narrow convoluted ridges

subgenus Marchantia (p. 185):

2a Thalli with margins entire or crenulate, crisped-plicate; dorsally often with dark median band; cells bordering

inner opening of epidermal pores with slightly convex inner walls, pores not cruciate; ventral scales in 6

rows and extending beyond thallus margins; appendage of median scales small, margins bordered with some-

what smaller cells; membranous margin of male receptacle crenulate, composed of slightly smaller cells;

rays of female receptacle with numerous papillae 1. M. polymorpha

2b Thalli with margins lobulate; dorsally without dark median band; cells bordering inner opening of epidermal

pores with strongly convex inner walls and pores cruciate; ventral scales in 4(-6) rows and not extending

beyond thallus margins, marginal scales sometimes partly or altogether absent; appendage of median scales

large, margins entire or slightly crenulate, bordered with strikingly small marginal cells; membranous margin

of male receptacle not distinctly crenulate, composed of very small cells; rays of female receptacle without

papillae 2. M. berteroana

lb Thalli with margins entire, nearly flat or sometimes slightly undulate, dorsally with or without dark median band;

cells bordering inner opening of epidermal pores with straight or convex walls, rarely with strongly projecting

processes and then pores cruciate; scales in 4 rows, covering V4 to almost 2/3 of ventral surface; appendage

of median scales orbicular, ovate or ± triangular, apically acuminate, acute or apiculate, marginal cells smaller

than inner ones; laminal scales as long as or longer than wide; cupules with margins ciliate, almost entire or

rarely with ciliate lobes, externally rarely with papillae; male receptacles palmate or rarely peltate, shallowly

or deeply dissected into rays; membranous margin of lobes entire or slightly crenulate; female receptacles

dissected into 5-9 apically ± flat lobes; margins of involucres ciliate or entire, rarely with ciliate lobes; stalks

with air chambers in 1 or 2 bands; spores 20-35 pm in diameter, ornamentation on 2 faces dissimilar, distally

mostly with wide ridges subgenus Chlamidium (p. 1 92):

3a Thalli medium-sized to large, 6-8(11) mm wide; epidermal pores with inner walls of cells bordering inner

opening strongly projecting and pores cruciate; appendages of median scales with margins entire or slightly

denticulate; cupules with triangular, ciliate lobes and externally with numerous papillae; female receptacle

deeply dissected into lobes, apically truncate or emarginate; margins of involucres with ciliate lobes ....

section Paleaceae (p. 192), 3. M. paleacea

3b Thalli smallish to large, 3.5-10.0 mm wide; epidermal pores with inner walls of cells bordering inner opening

straight or convex; appendages of median scales with margins often distinctly toothed; cupules bearing

long or short cilia or almost entire at margins and externally without or with few papillae; female receptacle

shallowly or deeply dissected into lobes; margins of involucres ciliate or ± entire:

Bothalia 25,2(1995)

185

4a Thalli 6.0-8.5 (—10.0) mm wide; dorsally without dark median band; appendages of median scales large,

520-550 x 375-530 pm, inner cells also comparatively large, 75.0-87.5 x 32.5-42.5 pm, i.e. cell structure

loose; margins of cupules with cilia up to 6 cells long; female receptacle large, shallowly dissected into

broad lobes, sometimes bearing a male ray; margins of involucres ciliate

section Chlamidium (p. 195), 4. M. pappeana

4b Thalli 3.0-7. 5 mm wide; mostly with distinct dark median band dorsally; appendages of median scales small,

350—375 x 275—340 pm, inner cells not markedly ‘loose’, cells 67.5—75.0 x 30.0—37.5 pm; margins of

cupules crenulate or with short cilia, up to 3 cells long; female receptacle rather small, deeply dissected

into narrow lobes; margins of involucres entire or crenulate section Papillatae (p. 199),. 5. M. debilis

Subgenus Marchantia

Marchantia F. sect. Astromarchantia Nees: 60, 61 (1838)

(nom. illegit.)

Marchantia L. (without rank) Stellatae Gottsche, Fin-

denb. & Nees; 522 (1844—1847; published 1846)

Marchantia L. (without rank) Marchantiotypus Dumort.:

150(1874).

Thallus branches wide, from nearly 7 mm to almost

twice as wide; margins crenulate, lobulate or plicate-

crisped. Dorsal epidermis with or without papillae; air

pores surrounded by 4-5(6) concentric rings of cells, at

inner openings sometimes with rounded walls, otherwise

with pronounced processes and pores then cruciate; stor-

age tissue lacking sclerotic cells. Scales in 4-8 rows, cov-

ering 3A to all of ventral face, containing scattered oil cells;

median scales with appendages, ovate, orbicular or reni-

form, not acuminate at apex; margins crenulate, toothed

or entire, sometimes with a row of cells very much smaller

than those in interior; laminal scales wider than long, on

either side of median scales, apically rounded, lacking pa-

pillae, upper cell walls with trigones. Cupules with acutely

triangular, toothed lobes, externally papillate.

Dioicous. Antheridiophores with receptacles ± symmet-

rical. peltate, rays very short and broad; stalks with 2 rhi-

zoid furrows, but lacking air chambers. Archegonio-

phores with receptacle deeply dissected into terete lobes;

stalks with 2 rhizoid furrows and a single band of air

chambers; involucres with toothed or ciliate margins.

Spores small, 8-16 pm in diameter; 2 faces similarly or-

namented, with numerous thin, highly convoluted ridges,

triradiate mark distinct to absent and wing absent.

Marchantia polymorpha , M. berteroanci and M. plicata

(from central and South America) are classified in subge-

nus Marchantia.

1 . Marchantia polymorpha L. in Species plan-

tarum edn 1: 1137 (1753); Bischl.: 34 (1993a). Type:

Europe, Dillenius in Historia Muscorum. t. 76, fig. 6E, F

(1742) [OXF, lecto., typo., fide Bischl. & Boisselier-

Dubayle: 363 (1991) (photo, of typo, in PRE!)].

Synonymy after Bischler-Causse (1993a).

Thallus large, mostly prostrate, sometimes growing

somewhat erect, apical segments broadly obovate, bright

or dark or yellowish green, medianly with or without dark,

continuous or interrupted band dorsally (Figure 1A), mar-

gins reddish brown, crisped, entire, sometimes crenulate;

pores quite small to large, subdorsal air chamber walls

visible from above; margins not raised and incurved when

dry; in crowded, overlying patches, repeatedly pseudodi-

chotomously furcate. Branches with total length up to 35

mm, terminally 2-15 mm long and successive branches

5— 1 0(— 14) mm apart, narrowly divergent to overlapping,

7-10 mm wide, 325-375 pm thick over midrib, laterally

thinning out into wings, apex notched, with appendages

of several purple-red median scales recurved over edge;

margins acute, thin, becoming recurved, flanks sloping

obliquely, ventral face medianly keeled (Figure 1C),

densely covered with rhizoids and on either side with 3

rows of purple or brownish scales, the outermost extend-

ing beyond thallus margins (Figure IB).

Dorsal epidermal cells unistratose, mostly hyaline,

4-6-sided, 35-74 x 18-30 pm, thin-walled, lacking

trigones, in transverse section 18-20 pm thick, marginal

cells in 3(4) rows (Figure 1G), those in outer 1 or 2 rows

smaller, generally rectangular, 15.0-27.5 x 17.5-20.0 pm,

inner cells larger, 27.5-37.5 x 22.5-27.5 pm; air pores

quite numerous, 175-350 pm distant from each other,

raised, compound, round or oval, 30.0-62.5 x 40.0-60.0

pm, suiTounded by 4 or 5 concentric rings of cells, 2 or

3 above epidermis and 2 or 3 projecting into air chambers

(Figure IF), innermost ring of upper cells collapsed, ± 5

pm wide, next ring of cells ± 40 x 10 pm, outer ring of

cells ± 50 x 12 pm, partly overlying epidermal cells (Fig-

ure ID), inner opening mostly surrounded by 4 cells (Fig-

ure IE), ± 27.5 pm long, and ± 12.5 pm wide across

widest parts which protrude into cavity and are covered

with black granular deposit. Assimilation tissue 50-100

pm thick, Vg— V3 as thick as thallus, air chambers in a

single layer, 150-250 pm wide, height of bounding walls

2 or 3(4) cells, round or oval, 17.5-27.5 x 15.0-17.5 pm,

chambers internally crowded with densely chlorophyllose,

mostly 2- or 3-celled filaments (Figure IF), cells round,

oval, elongated or irregular, 20.0-32.5 x 15.0 pm; storage

tissue occupying ± 1 1 rows of cells in ventral 2/3— 5/g of

thickness of thallus medianly, decreasing laterally, cells

angular, 45.0-75.0 x 60.0-65.0 pm. smaller just below

air cavities and where adjoining ventral epidermis; scle-

rotic cells and mucilage cavities absent; rhizoids some

smooth. 1 5-50 pm wide, faintly brown, others tuberculate,

7.5-12.5 pm wide, and still others with internal ‘spirals',

± 12.5 pm wide.

Median scales (Figure IK) pale mauve, in one row

on either side of midrib, ± obliquely triangular, body up

to 750 pm long, width 1575-2250 pm across nearly

straight base, continuing into narrow ‘tail', cells in body

of scale elongated, 4-6-sided, 85-160 x (25-)40-62 pm,

rhizoids arising from a few smaller rhizoid initial cells

and with 5 or 6 scattered oil cells present, ± 25.0 x 22.5

pm, toward margins cells changing orientation and some

in semiradial groups, from centre of which 1 or 2 papillae

project. 35-50 x 10 pm; scale narrowing upwards, deeply

constricted where joined with appendage, which is broadly

186

Bothalia 25,2(1995)

FIGURE I . — Marchantia polymorpha L. subsp. ruderalis. A-G, thallus: A, dorsal lace of apical branch of <? with disc and cupule; B, ventral face of apical

branch; C, t.s. of branch; D, air pore and surrounding cells from above; E, air pore from below; F, t.s. of air pore, dorsal epidermal cells and air

chamber; G, margin, H, cupule margin; I, marginal scale; J, laminal scale; K, median scale; L, appendage of median scale; M, a" receptacle; N, ¥

receptacle from side; O, t.s. of cf stalk; P, t.s. of ¥ stalk; Q, median scale of o' receptacle. R-V, scale: R, along o" stalk; S, ¥ receptacle; T, along ¥

stalk; Ui, U2, loot of d" stalk; Vi, V2, fool of ¥ stalk. W, t.s. of ¥ ray; X, detail of part of section of ¥ ray, showing papillae on epidermal cells and

air chambers; Y, margin of 0" ray; Z, margin of involucre. A, M, O, Q, R, Ui, U2, Y, Perold & Koekemoer 3248\ B-L, N, P, S, T, Vi, V2, Wi, X,

Z, Glen 3728. Scale bars: A, B, M, N, 2 mm; C, 1 mm; D-F, 50 pm; G, X, Y, 100 pm; H-L, O-W, 250 pm.

Bothalia 25,2 (1995)

187

rounded, brown or tinged with purple, (350-) 400-640 x

500-700 pm, central cells 4-6-sided, 42.5-62.5 x

32.5- 50.0 pm, with 1-3 oil cells, up to 30.0 x 22.5 pm,

margin sharply toothed, crenulate or almost entire, cells

small, some conical, ± 22.5 x 15.0 pm, projecting slightly,

others rectangular, with long axis parallel to margins, ±

35.0 x 7.5 pm, or perpendicular, with 10 pm high protu-

berance, sometimes separated as distinct cell, submarginal

cells larger, 25.0-27.5 x 20.0-37.5 pm. Laminal scales

hyaline or pale mauve, in one row on either side of and

lateral to median scales, almost wedge-shaped, but with apex

rounded, base flatly arched (Figure 1 J), 600-825 x

1450-1750 pm, upper cells rather irregularly arranged, some

with slightly thickened walls, 4—6-sided, 37.5-42.5 x

17.5- 27.5 pm, larger lower down, 70-90 x 30-35 pm, oil

cells rare, 5 or 6 rhizoids originating from smaller internal

initial cells, at upper margin, several papillae, ± 47.5 x 12.5

pm. Marginal scales (Figure II) usually projecting beyond

thallus margins, hyaline or brownish, oblong or ovate, apex

rounded, 700-750 x 550-650 pm, upper cells with walls

thickened and brown, ± 47.5 x 32.5 pm, lower down walls

not thickened, cells 70.0-120.0 x 25.0-37.5 pm, up to 5 or

6 scattered oil cells and some rhizoid initials present. Cupules

with ciliate, ± triangular lobes (Figure 1H), ± 240 pm long

and 250-275 pm wide at base, apically with 5 or 6 cells in

a uniseriate, vertical row, top cell conical, ± 20 x 10 pm,

lower cells gradually enlarging, bottom cell trapezoid, ± 27.5

x 32.5 pm, at base of lobes on either side 3 or 4 trans-

versely projecting cellular filaments or cilia, 1-6 cells long,

cells ± 35 x 10 pm, lower down cupule wall several cell

layers thick, exteriorly with numerous 1-3-celled papillae

and internally with many oil cells.

Dioicous. Antheiidiophore arising from apex of termi-

nal segment (Figure 1A) of main or short lateral branch,

raised on stalk, ± 4.5-15.0 mm long, in transverse section

rounded, at widest part along its length, ± 975 x 1000 pm

(Figure 10), cortical cells small, 10.0-17.5 x 7.5-20.0

pm, outer wall slightly thickened and bulging, medullary

cells larger, angular, 22.545.0 x 22.5-35.0 pm, air cham-

bers absent, but with 2 rhizoid furrows, 237.5-250.0 x

87.5- 137.5 pm, lined with purple pigmented cells; scales

at base of stalk (Figure 1 U i , U2), large, with purple bases,

round or oblong, lacking an appendage, 1 250-2000 x 800

pm, cells in body of scale, 50.0-57.5 x 20.0-30.0 pm,

smaller at apex and margin, 22.5-37.5 x 12.5 pm, some-

times with protruding papillae, up to 8 scattered oil cells

present; scales along length of stalk, hyaline, filiform (Fig-

ure 1R) (1)2-3 cells wide; receptacle ± 8 mm in diameter,

shallowly dissected into 8(— 1 0) lobes, ± 375 pm long,

symmetric (Figure 1M), basal sinus up to 30° wide; mar-

gins of lobes (Figure 1Y) membranous, hyaline, crenulate,

cells in outer 1 or 2 rows small, 10.0-17.5 x 15.0-22.5

pm, some with a bulging protrusion, inner cells larger,

30.0-72.5 x 32.5-37.5 pm; median scales on ventral side

of lobes (Figure IQ) hyaline, oblong with rounded apex

and lacking appendage, 1375-1575 x 600-850 pm, cells

in body of scale 4-6-sided, 50.0-75.0 x 27.5-30.0 pm,

4—10 oil cells present, ± 25 x 25 pm, upper marginal cells

thin-walled, small, 7.5-15.0 x 12.5-15.0 pm, sometimes

toward base of lateral margins, with small papillae.

Arche goniophore arising from apex of terminal segment

of main or short lateral branch, raised on stalk, 17-40 mm

long, in transverse section 1000 x 1250 pm, narrower across

single band of air chambers, ± 625 pm (Figure IP), cor-

tical cells small, 10.0-20.0 x 7.5-17.5 pm, outer wall

slightly thickened, medullary cells angular, 37.5—42.5 x

25.0-32.5 pm, with 2 rhizoid furrows, ± 350 x 250 pm;

scales at base of stalk, (Figure IV], V2), large, puiple or

brown or hyaline, rarely with slightly constricted appendage,

mostly without, but some appear to be composite scales,

shape irregular, up to 750 x 1000(4 750) pm, inner cells ±

50 x 45 pm, small at crenulate upper margin, 17.5-25.0 x

12.5-27.5 pm, at lateral margins toward base, sometimes

with papillae, ± 35 x 15 pm; scales along length of stalk

(Figure IT), hyaline or brown, filiform, up to 600 pm long,

apical cell conical, ± 55.0 x 12.5 pm, lower 2(3) cells ±

62.5 x 20.0 pm, serially arranged and then 2 adjacent, some-

times with base of upper cell wedged between them, and

sometimes with a short lateral branch; receptacle 9.5-10.0

mm in diameter, nearly symmetric, with small round projec-

tion dorsally, deeply divided into 9-1 1 rays of slightly un-

equal length (Figure IN), 2500-3200 pm, basal sinus ± 40°,

margins distally decurved and terete in transverse section

(Figure 1W), toward apices with numerous conical papillae

(Figure IX), ± 12.5 x 17.5 pm; involucres with margins

hyaline or occasionally purple-tinged, with tapering ciliate

lobes (Figure 1Z), 300-350 pm long, 90-150 pm wide at

base, inner cells 5- or 6-sided, 52.5-75.0 x 22.5-32.5 pm,

with scattered, smaller oil cells in between, cilia ( 1— )2— 5

cells or up to 230 pm long, top cell conical, 37.547.5 x

10.0 pm, basally with single cell, 72.5 x 25.0 pm, but some-

times with 2; scales of receptacle (Figure IS) hyaline, up to

2000 pm long and 80 pm or 6 cells wide at base, apical

cell conical, ± 62.5 x 12.5 pm, following 3 cells in a vertical

row, ± 57.5 x 15 pm, basal cells 62.5-87.5 x 12.5-20.0 pm,

sometimes branched along the length. Spores 10.0-12.5 pm

in diameter, triangular-globular, yellow; distal face (Figure

2 A, B) with numerous highly convoluted and branched,

smooth ridges; proximal face (Figure 2C, D ) with faint, col-

lapsed triradiate mark, facets finely and densely granular,

narrowly winged. Elaters yellow-brown, 350435 x 5 pm,

gradually tapering to narrow tips, bispiral. Chromosome

number, n = 9 (Bischler-Causse 1993a).

Marchantia polymorpha, which has been known since

classical times, is widespread. It has frequently been de-

scribed but until recently its taxonomy has been unre-

solved because proper lectotypification had not been done.

Amell (1963) reported it in southern Africa from Golden

Gate, near Clarens in the eastern [Orange | Free State and

from Zimbabwe [Southern Rhodesia], near Odnazi River

Bridge, Umtali. These specimens are not held at PRE or

at BOL and up to now, the presence here of M. polymor-

pha under natural conditions has not been confirmed.

Bischler-Causse (1993a) stressed the need for such con-

firmation. It is therefore considered worthy of note that

female specimens of M. polymorpha with cupules have

recently been collected at Johannesburg Botanical Garden,

Glen 3468 & 3728, in a shade house and also at Sterlig

Nursery, Krugersdorp, Perold & Koekemoer CH 13640

(Figure 3). The plants are clearly introduced, but a de-

scription is nevertheless given. To complete the descrip-

tion and illustrations fresh male plants collected on a

pavement in Ledbury, England, Perold & Koekemoer

3248, had to be used.

The specimens above have been referred to M. poly-

morpha subsp. ruderalis Bischl. & Boisselier-Dubayle,

188

Bothalia 25,2(1995)

FIGURE 2. — SEM micrographs of spores. A-D, Marchantia polymorpha , S.M. Perold <£ M. Koekemoer 3248: A, distal face; B, part of distal face

much enlarged; C, proximal face; D, part of proximal face much enlarged. E-I, M. berteroana , Geldenhuys 1332: E, distal face; F, part of

distal face much enlarged; G, proximal face; H, part of proximal face much enlarged; I, side view. A, C, x 3006; B, D, x 5780; E, x 3044; F,

H, x 5596; G, x 2875; I, x 3006.

since they grow as weeds in man-made habitats and agree

in other respects with this subspecies, as distinguished by

Bischler-Causse & Boisselier-Dubayle (1991). They are

characterized by the fresh thalli being prostrate and bright

green, with a dark, rather indistinct and discontinuous me-

dian band with elongated air chambers, by the dorsal air

pores being 50.0-62.5 pm wide, the dorsal epidermal cells

35-55 pm long and the median scale appendages having

toothed margins and being 350-500 x 510-760 pm in size.

The other 2 subspecies of M. polymorpha, polymorpha and

montivagans, have so far not been found in southern Af-

rica. They are distinguished by occurring in natural habi-

tats, with the former growing more or less erect, being dark

green, with a conspicuous median dark band lacking air

chambers, having small air pores, short dorsal epidermal

cells and the median scale appendages having entire mar-

gins; the latter subspecies is prostrate, yellow-green, lacks

a median band, the air pores are larger, the dorsal cells are

of intermediate length and the median scale appendages

are of average size and the margins toothed. The distin-

guishing characters of the subspecies are given, in case M.

polymorpha subsp. polymorpha and M. polymorpha subsp.

montivagans are found in southern Africa in the future.

Lectotypification of M. polymorpha was recently done

by Bischler-Causse & Boisselier-Dubayle (1991) by

means of a Dillenian illustration and by a specimen held

in OXF (i.e. a typotype). This corresponds to the first of

three Linnean varieties (Linnaeus 1753), namely var. [°c],

which was later called 'aquatica' . This first variety [®=]

forms the basis for the application of the specific name

(Isoviita 1970). However, since the lectotype of M. poly-

morpha var. polymorpha of Linnaeus corresponds to the

taxon ‘ aquatica ’ (at any rank), it renders that name ille-

gitimate, because the epithet ‘ polymorpha ’ would be used

for a taxon not including its type. The best procedure

seemed, therefore, for Bischler-Causse & Boisselier-

Dubayle to describe their three newly separated electro-

phoretic groups (in three enzyme systems: esterases,

peroxidases and acid phosphatases) as subspecies, give

two of them new names and select good, recent type

specimens. M. polymorpha L. subsp. polymorpha thus cor-

responds to the taxon formerly called ‘ aquatica M. po\y-

morpha L. subsp. ruderalis Bischl. & Boisselier-Dubayle

corresponds roughly to the former polymorpha ‘ sensu

stricto' and M. polymorpha L. subsp. montivagans Bischl.

& Boisselier-Dubayle roughly to the former ‘ alpestris ' .

Bothalia 25,2 (1995)

189

On chemical evidence Markham et al. (1977) had pre-

viously concluded that the three taxa of M. polymorpha

should be treated as varieties rather than as separate spe-

cies, since they all possess identical flavone glucuronides.

The time of going to press of Schuster’s (1992) monu-

mental work pre-dated the publication by Bischler-Causse

& Boisselier-Dubayle (1991) on the lectotypification of

M. polymorpha , but judging by some of the remarks con-

tained in it, it is by no means certain that he would have

accepted it. He still subscribes to the genetic analyses of

Burgeff (1943), although they have been shown to be out

of date, and he therefore treats the segregates of M. po-

lymorpha as distinct species.

As mentioned above, in southern Africa M. polymor-

pha subsp. ruderalis is only known from man-made sites,

i.e. in nurseries. Its presence in natural habitats has not

been confirmed. Asakawa et al. (1988) refer to the distri-

bution of sesquiterpenoids and cyclic bis-bibenzyls in

southern African collections of M. polymorpha. but it is

not known where the material was obtained from or who

identified it. Magill & Schelpe (1979) and Arnold & De

Wet (1993) list M. polymorpha as occurring in southern

Africa, but these lists are based on erroneous information

in the literature.

Marchantia polymorpha is very similar to M. ber-

teroana. but can be distinguished from it by the marginal

scales which are always present and which mostly extend

beyond the crisped, sometimes crenulate thallus margins;

by the air pores not being cruciate; by the presence of

numerous papillae on the rays of the female receptacle

and by the appendages of the median scales bordered by

larger cells.

2 . Marchantia berteroana Lehm. & Lindenb. i n

Lehm., Novarum et minus cognitarum stirpium, pugillus 6: 2 1

(1834); Gottsche et al.: 481 (1846); Steph.: 393 (1898-1900);

Schift'n.: 41—44 (1896); A. Evans: 246 (1917); Hassel de

Menendez: 160 (1963); S.W. Amell: 55 (1963); E.O. Campb.:

122 (1965); Bischl.: 44 (1984); Bischl.: 81 (1989a); Bischl.:

56 (1993a). Type: ‘In insula Juan Fernandez, legit cl. Bertero

(Herb. Hooked), ( W8294 , holo.; FH, G, NY, PC, STR, W, iso.).

FIGURE 3. — Distribution of M. polymorpha var. ruderalis , •; M.

berteroana. ▲; and M. paleacea, ■ in southern Africa.

M. tabularis Nees: 71 (1838); Sim: 28 (1926). Type: South Africa,

Tafelberg, leg. Ecklon (W8289) (female) W!, syn.; leg. Bergius (male)

STR, syn.

M. contractu Bisch.: 135 (1846) (fide Bischler-Causse (1993a)). Type:

South Africa, Cap, Duyvelsberg, Krauss s.n. (BM. FH, G!, PC. W).

For the rest of the synonymy consult Bischler-Causse (1993a).

Thallus robust, rather flat, almost leathery, apical seg-

ments broadly oblong (Figure 4A), green, yellowish green

or bluish green and occasionally with some purple pig-

mentation, without distinct median band dorsally, margins

hyaline, puiplish or brownish, irregularly lobulate, more

or less plicate but not crisped, entire or minutely crenulate,

pores numerous, small, almost closed, subdorsal air cham-

ber walls indistinct from above, when wet; thallus margins

not raised or incurved, when dry; in crowded, overlying

mats, repeatedly furcate but rather irregularly so. Branches

with total length up to 80 mm, terminally up to 20 mm

long and successive branches 6-10 mm apart, narrowly

to moderately divergent, 9.5— 12.0(— 15.0) mm wide,

600-900 pm thick over midrib, laterally thinning out into

wings (Figure 4C); apex notched with brownish, hyaline

or partly puiplish red appendages of median scales recur-

ved over edge; margins acute, thin, slightly recurved,

flanks sloping obliquely; ventral face (Figure 4B) brownish

or purplish, medianly keeled, covered with rhizoids, and with

2 rows of scales on either side (sometimes with 3), extending

over 2 3 * * * * */4-9/io of thallus width, but never right to the edge.

Dorsal epidermal cells unistratose, seldom bistratose

in patches, hyaline, polygonal, 4-7-sided, (50-)57. 5-77.5

x 20.0-30.0 pm, walls slightly thickened, in transverse

section 15-20 pm thick, occasionally containing an oil

body; along margins (Figure 4G) 3 or 4 rows of cells,

smallest in the outermost row, rectangular or polygonal,

17.5-22.5 x 10.0-15.0 pm, enlarging inwardly, 27.5-37.5 x

25.0- 37.5 pm, air pores numerous, ( 1 07.5—) 1 37.5— 1 85.0

(-225.0) pm distant fom each other, raised, compound, oval

(37. 5-)57. 5-75.0 x 27.5-47.5(-55.0) pm, encircled by

(4)5-7 concentric rings of cells, 3(4) above epidermis and 3

or 4 projecting into air chambers (Figure 4F), innermost ring

with collapsed cells ± 5 pm wide, uppermost ring with 4 or

5 sausage-shaped cells, 50-55 x 10 pm, also 4 or 5 cells in

next ring, 45.0-47.5 x 27.5-32.5 pm, those in outermost

ring irregular in shape as they adjoin or partly overlie dorsal

epidermal cells (Figure 4D), 55.0-62.5 x 12.5-20.0 pm. in-

ner opening with 4 or 5 cells, their inside walls covered with

a granular deposit and strongly protuberant, leaving only a

small, usually cruciate opening (Figure 4Ej, E?). Assimilation

tissue 45— 50(— 60) pm thick. ’/jO-'Ao as thick as thallus, air

chambers in a single layer, 77.5-120.0 pm wide, bounding

walls 3 or 4 cells high, rounded or angular, 7.5-20.0 x

10.0- 12.5 pm, chambers crowded with densely chlorophyl-

lose filaments, cells oval, round or irregularly shaped.

15.0- 20.0 x 12.5-15.0 pm; storage tissue occupying ventral

9/jq— 19A>o of thickness of thallus medianly, decreasing lat-

erally, cells angular, up to 75 pm wide, becoming smaller

lower down, containing scattered starch grains and some

with an oil body, sclerotic cells and mucilage cavities ab-

sent; rhizoids some smooth, ± 27.5 pm wide, others

pegged, 27.5^15.0 pm wide.

Median scales (Figure 41) hyaline or puiplish. in one

row on either side of midrib, body ± obliquely triangular,

up to 1450 pm long, across base 2875-3500 pm, hardly

190

Bothalia 25,2(1995)

FIGURE 4. — Marchanlia berteroana Lehm. & Lindenb, A-G, lhallus: A, dorsal face of apical branch with cupule; B, ventral face of apical branch;

C, t.s. of branch; D, air pore and surrounding cells from above; Ei, E2, air pores from below; F, t.s. of air pore, dorsal epidermal cells and air

chamber; G, margin. H, laminal scale; I, median scale; J 1 , J2, appendages of median scales; K, cupule margin; L, ¥ receptacle; M, o" receptacle;

N, t.s. of <f stalk; O, t.s. of ¥ stalk. P-R. scale: P, from foot of a" stalk; Q, from along <? stalk; R, median scale of ef receptacle. S, margin of

<f ray; T, scale from foot of ¥ stalk; U. scale from ¥ receptacle; V, margin of involucre. A, F, G, L, N, P, R, S, A.E. van Wyk 2066: B-Et,

E2, H— J 1 , h, M, O, T-V, Geldenhuys 1332: K, Q, Pillans 4048. Scale bars: A, B, L, M, 2 mm; C, 1 mm; D-G, 50 pm; H-K, N-R, T, 250

pm; S, U, 100 pm.

Bothalia 25,2 (1995)

191

arched, central cells elongated and narrow, 4 or 5— (6)-

sided, 125.0-212.5 x 17.5-30.0 pm, toward margins outer

7 or 8 rows of cells thinner-walled, orientation irregular

and much smaller in size, 27.5-50.0 x 20.0-22.5 |am, with

as many as 30 scattered oil cells, ± 87.5 x 37.5 pm, dark

brown, scale gradually narrowing upwards and then ±

1875 pm wide, abruptly and deeply constricted at join

with appendage, hyaline or purplish or both, orbicular to

broadly ovate (Figure 4J j, J2), apex rounded to somewhat

obtusely narrowed, basally cordate, 675-950 x 650-800

pm, median cells 4-6-sided, 80.0-95.0 x 45.0-52.5 pm,

with 4 or 5 oil cells, margins entire or minutely crenulate,

bordered by 1 or 2(3) rows of very small cells, 30.0^10.0

x 10.0-20.0 pm, mostly narrowly rectangular to subquad-

rate and orientated with long axis perpendicular to margin,

sometimes alternating with 2 stacked cells parallel to mar-

gin, and even smaller, 12.5 x 12.5-17.5 pm. Lamina!

scales mostly hyaline, yellowish brown or reddish, lateral

to, but also alternating irregularly with median scales,

broadly rounded, lacking apical appendage, base flatly

arched (Figure 4H), 850-1125 x 1875-3375 pm, cells in

body of scale 4-6-sided, walls straight or sinuous,

92.5-125.0 x 22.5-32.5 pm, sometimes in upper part with

thickened walls, but in 6 or 7 rows toward inner margins

thinner-walled, smaller, 32.5-50.0 x 12.5-25.0 pm, and

very variable in orientation, with ± 12 oil cells, margins

entire or slightly crenulate. Marginal scales rarely to

sometimes present, when more or less scattered and never

extending beyond thallus margins, hyaline, crescent-

shaped with rounded apex, 675-1250 x 775 pm, upper

cells small, with thickened comers, 15.0-25.0 x 12.5 pm,

inner cells with slightly sinuous walls, 35.0-58.5 x

15.0- 25.0 pm. toward base cells 5- or 6-sided, up to 112.5

x 22.5 pm. Cupule margins with ciliate, ± triangular lobes

(Figure 4K), up to 675 pm long and 250 pm wide at base,

at apex with 10 or 11 cells in a tapering, uniseriate, vertical

row, top cell conical, ± 32.5 x 7.5 pm, lower cells larger,

up to 37.5 x 30.0 pm, at base of lobes on either side a

few transversely projecting (2— )3 or 4-(-6)-celled cilia,

lower down cupule wall several cell layers thick, exteri-

orly with numerous 1- or 2(-3)-celled papillae, 32.5-50.0

x 20.0 pm and internally with many oil cells.

Dioicous. Antheridiophore arising from apex of termi-

nal segment of main or short lateral branch, raised on stalk

(Figure 4N), 23-30 mm long, in transverse section

rounded, diameter ± 1000 pm, cortical cells small,

10.0- 15.0 x 12.5-17.5 pm, with thickened outer wall,

medullary cells larger, mostly angular, 22.5-45.0 x

15.0- 45.0 pm, band of air chambers absent but with 2

rhizoid furrows, ± 162.5 x 100.0-125.0 um; scales at base

of stalk large (Figure 4P), hyaline or purplish, roughly

triangular or oblong, sometimes slightly bulging, lacking

an appendage, up to 1200 x 1400 pm, cells in body of

scale 4—6-sided, often with sinuous walls, 47.5-87.5 x

22.4-25.0 pm, along upper margins 1 or 2 rows of much

smaller cells, 12.5-25.0 x 12.5-30.0 pm, with up to 10

scattered oil cells throughout; scales along length of stalk

(Figure 4Q) hyaline, filiform, sometimes only 3 cells wide

but sometimes wider and up to 9 cells wide; receptacle

(Figure 4M) up to 10 mm in diameter, shallowly dissected

into 8(9) ± symmetric lobes, basal sinus ± 80° wide; mar-

gins of lobes membranous, brownish or hyaline, minutely

crenulate, with 1 or 2 rows of small cells (Figure 4S),

mostly rectangular across, 15.0-20.0 x 10.0-22.5 pm. in-

ner cells larger, 52.5-87.5 x 37.5-40.0 pm; median scales

on ventral side of lobes (Figure 4R) hyaline, 750-1375 x

1000-1875 pm, cells in body of scale 4-6-sided, up to

125 x 50 pm, thin-walled, at margins smaller, ± 30.0 x

17.5 pm, appendage absent or present, ± 267.5 x 200,0

pm, oblong, rounded apically, not or hardly constricted at

join with scale, inner cells 40.0-42.5 x 15.0-22.5 pm,

smaller at margins.

Arche goniophore arising from apex of terminal seg-

ment of main or short lateral branch; raised on stalk 48-65

mm long, in transverse section (Figure 40) 1000 x 925

pm, constricted on inner side of single band of air cham-

bers, cortical cells small, 12.5-20.0 x 12.5-25.0 pm, outer

wall thicker, medullary cells angular, up to 60 x 45 pm,

in between and at centre of stalk with smaller cells ± 25

x 20 pm, containing 2 rhizoid furrows, ± 240 x 105 pm;

scales at base of stalk (Figure 4T), brownish or hyaline,

without appendage, oblong or ovate, with apex rounded,

large, 1575-2000 x 800-1250 pm, inner cells 4—6-sided,

walls mostly straight, sometimes sinuous, 75.0-125.0 x

27.5—42.5 pm. margins apically occasionally slightly cre-

nulate, with rectangular cells arranged at right angles to

margin, small, 10.0-25.0 x 12.5-25.0 pm; scales along

length of stalk (Figure 4U) hyaline, filiform, up to 6500

pm long, 2 or 3(4) cells wide, their average size ± 1 12.5

x 25.0 pm; receptacle (Figure 4L) up to 10 mm in di-

ameter, nearly symmetric, dorsally with small round me-

dian projection, deeply divided into 9 linear rays, up to

2625 x 550 pm, basal sinus ± 45° wide, margins distally

slightly decurved and terete in transverse section, lacking

papillae toward apex; involucres with margins hyaline or

purplish, with tapering ciliate lobes (Figure 4V). base

100.0- 175.0 x 125.0-182.5 pm, inner cells 5- or 6-sided.

35.0- 37.5 x 22.5-25.0 pm. lower down with numerous

oil bodies, up to 60.0 x 37.5 pm, each one almost filling

a cell, lobes apically and at sides with numerous, tapering,

branched and intertwined cilia, length up to 220 pm or 7

cells in series, 22.5-50.0 x 20.0 pm, with granules on

walls of apical cells; scales of receptacle (Figure 4U) hya-

line, filiform, up to 2150 pm long and 35.0-87.5 pm or

2-5 cells wide at base, often splitting further along and

strands mostly only 2 cells wide, 80.0-87.5 x 20.0-22.5

pm, apical cell ± 75.0 x 17.5 pm. Spores 7.5-12.5 pm in

diameter, ± globular, brownish, distal face (Figure 2E, F,

I) with numerous, much convoluted and branched ridges;

proximal face (Figure 2G, H, I) with faint triradiate mark

and winged, densely covered with numerous fine granules

and tiny irregular ridges. Elaters yellow-brown. (550-)

600-640(-770) x 5 pm, gradually tapering at both ends

for 150-200 pm, bispiral. Chromosome number : n = 9

(Bischler-Causse 1993a).

Marchantia tabularis was described from specimens

collected on Table Mountain and was placed in synonymy

under M. berteroana by Schiffner (1896). Although

Stephani (1898-1900) accepted Schiffner’s decision, he

still applied the epithet ‘ tabularis ' and was followed in

this by Sim (1926). Amell (1963) gives only brief notes

on M. berteroana. Bischler-Causse (1993a) recognized

that M. contracta, collected by Krauss at Devil's Peak,

also belongs here. M. berteroana is quite variable in size,

but generally the plants are large. Sometimes six rows of

ventral scales are present and some authors, e.g. Hassel

de Menendez (1963) and Engel ( 1990) recognize these as

192

Bothalia 25,2(1995)

belonging to a separate variety, namely var. polylepida.

However, several authors (Campbell 1965; Bischler 1984)

have pointed out that the number of rows of scales is too

unstable to warrant the designation of a variety. Schuster

( 1992) has assigned M. berteroana to his new monotypic

section, Berteroanae, on the absence of marginal scales,

on the cruciate inner openings of the epidermal pores with

six rings of cells and on the tiny marginal cells of the

median scale appendages. Bischler-Causse (1993c) is,

however, not convinced that this splitting is necessary,

since marginal scales are fairly frequently present and the

pores are often similar to those in other species of subge-

nus Marchantia.

Marchantia berteroana generally grows on damp soil,

and sometimes on wet rocks, at stream banks, near wa-

terfalls, along paths, in kloofs, passes, ravines, gorges, in

forests under trees or in burnt-over areas.

It is widely distributed in the southern hemisphere,

from South America, south to the Antarctic Peninsula and

north to Costa Rica, islands of the Atlantic and south In-

dian Ocean, southern Africa, Australia, Tasmania, New

Zealand, New Guinea, New Caledonia, Java and Sulawesi

(Bischler-Causse 1993a). In southern Africa (Figure 3) it

is known from the Western Cape, which has winter rain,

and from Kwazulu-Natal as well as the Northern Province

(Northern Transvaal | which have summer rain. Exact lo-

calities of two of these collections by Wilms and by

MacLea are not known. This distribution indicates that

the species seems to be indifferent to the seasonality of

the rainfall, in contrast to several other members of the

Marchantiales which are ± restricted to either winter or

summer rainfall areas.

Of the 50 specimens examined 36% had cupules, 34%

had antheridiophores and 46% archegoniophores; only 6%

had both.

Specimens of M. berteroana are easily distinguished

by the tiny marginal cells of the median scale appendages,

by the cruciate dorsal pores and by the lobulate thallus

margins. The membranous margins of the rays of the male

receptacle also have very small cells. Otherwise it is quite

similar to M. polymorpha, except for the latter having pa-

pillae on the rays of the carpocephalum. The ornamenta-

tion of their spores is also quite similar.

Subgenus Chlamidium (Corda) Bischl., Cryp-

togamie, Bryologie et Lichenologie 3: 362 (1982); Bischl.:

89 (1989a); Bischl.: 65 (1993a).

Marchantia L. sect. Chlamidium (Corda) Nees: 60, 101

(1838). Type: Chlamidium indicum Corda (Sieber flora

mart, exsicc. No. 375) = Marchantia chenopoda L. Type:

Sieber 378 p.p. (W, neo.; STR, iso., fide Bischl. 1984).

Thallus branches rather narrow, from 2.4 to ± 10.0 mm

wide, rarely more, margins nearly Oat, entire, sometimes

slightly undulate, rarely crisped. Dorsal epidermis with-

out papillae; air pores surrounded by (4— )5— 7 concentric

rings of cells, at inner openings with straight or convex

walls, rarely with pronounced, rounded processes, and

pores then cruciate; storage tissue often with scattered

sclerotic cells. Scales in 4 rows, covering V4 to 2Aj of ven-

tral face, with oil cells present or absent, median scales

with appendages variously shaped, orbicular, ovate or tri-

angular, apically often acuminate, acute or apiculate, sel-

dom rounded; margins entire, crenulate-serrate, coarsely

toothed or lobed; laminal scales as long as, or longer than

wide, sometimes in 2 incomplete rows on either side of

median scales, apically acute or obtuse, with papillae, up-

per cell walls lacking trigones. Cupules with margins cili-

ate, almost entire, or with ciliate lobes, externally without,

or rarely with papillae.

Dioicous. Antheridiophores with receptacle symmetric

or asymmetric, palmate or rarely peltate, rays shallowly

or deeply dissected; stalks with 2(-4) rhizoid furrows, air

chambers in a single band or absent. Archegoniophores

with receptacle symmetric, or sometimes asymmetric; dis-

sected into 5-9 rays, flat or convex but never terete; scales

of receptacle in African species apically with marginal

cells rectangular and long axis parallel to margins; involu-

cre margins ciliate or crenulate to entire, rarely with ciliate

lobes; stalks with 2 -4 rhizoid furrows, air chambers in 1

or 2 bands. Spores larger than in subgenus Marchantia ,

20-35 pm in diameter; ornamentation on distal face gen-

erally with thick ridges separated by dense granules; on

proximal face with coarse granules only, triradiate mark

and thick wing usually present.

Subgenus Chlamidium contains three sections, Paleaceae,

Chlamidium and Papillatae. Section Paleaceae contains a

single species. Sections Chlamidium and Papillatae are each

represented in southern Africa by a single species.

Marchantia section Paleaceae Bischl. in Bryophy-

torum Bibliotheca 38: 90 (1989a); Bischl.: 67 (1993a).

Type species: M. paleacea Bertol. (lecto. fide Grolle: 210

(1976)].

Thallus with branches (3.5— )6.0— 8.0(— 1 1 .0) mm wide,

irregularly spaced, narrowly divergent. Dorsal epidermis

without papillae; air pores with inner opening cruciate,

inside walls of bordering cells strongly protuberant. Me-

dian scales with appendage oblong, ovate or suborbicular,

apically rounded, acute or shortly apiculate, basally cor-

date, width across broadest part 650-750 pm, margins en-

tire, crenulate or slightly denticulate; with 1, 2 or more

oil cells. Cupules with margins triangularly lobed and cili-

ate, externally with I - or 2-celled papillae.

Dioicous. Antheridiophore on stalk lacking band of air

chambers and basally surrounded by large scales without

an appendage; receptacle peltate, shallowly dissected into

6-10 broad, rounded lobes, dorsal surface without papil-

lae. Arche goniophore on stalk having a single band of

small air chambers, basally surrounded by large scales,

their apices rounded, rarely with a short appendage; re-

ceptacle bearing prominent median projection dorsally,

deeply divided into 8 or more convex lobes, basally

costate, apically truncate or hardly broadened, emarginate;

involucre margin with ciliate lobes. Spores 19-24 pm in

diameter, ornamentation on distal face lacking areolae,

mostly covered with a rather featureless, granular layer or

with very irregular ridges, broken up or folded in or con-

voluted; proximal face different, thickly winged, triradiate

Bothalia 25,2(1995)

193

mark faint, covered with dense granules or centrally with

narrow, irregular granular ridges. Only M. paleacea be-

longs to this section. The ornamentation of its spores is

distinctive. It also differs from the other two sections in

subgenus Chlamidium , by the shape of the female recep-

tacle and by the ciliate, lobed margins of the involucres

and the cupules.

3. Marchantia paleacea Bertol., Opuscoli scienti-

fic! di Bologna 1: 242 (1817); Bischl.: 55 (1984); Bischl

91 (1989a); Bischl.: 68 (1993a) subsp. paleacea. Type;

Italy, Borgonuovo secus valles in Liguria orientali, D.

Turio , 1810 |BOLO, lecto. fide Grolle (1976)].

M. papillata Raddi var. italica Raddi: 20 (1822). Syntypes: Italy,

Contomi di Firenze, Raddi s.n. (BOLO, FH. FI, G, PC, STR).

For the rest of the synonymy see Bischler-Causse (1993a).

Thallus medium-sized to large, firm and occasionally

somewhat leathery, apical segments oblong (Figure 5A),

bright green to slightly bluish green, sometimes blotched

with dark red pigmentation, generally lacking but now and

then with faint, short stretches of dark median band dor-

sally, margins mostly deep red or pink, entire, proximally,

however, undulate and scalloped, pores numerous, small,

generally closed, subdorsal air chamber walls visible from

above and these flecked with numerous white oil bodies,

when wet; thallus margins ± crinkled, not raised or incur-

ved, when dry; in crowded, overlying mats, repeatedly

furcate. Branches with total length up to 45 mm, termi-

nally up to 15 mm long and successive branches rarely

more than 10 mm, but mostly less apart, narrowly diver-

gent, (3.5— )6.0— 8.0(— 1 1.0) mm wide, 750-800 pm thick

over midrib, laterally thinning out into wings (Figure 5C),

apex notched with dark red to marginally orange append-

ages of median scales recurved over edge; margins acute,

thin, flanks sloping obliquely; ventral face (Figure 5B)

dark red entirely or only medianly, and the remainder

green, with 2 rows of scales on either side of ventral keel,

extending over V3— V2 of thallus width.

Dorsal epidennal cells mostly unistratose, here and

there bistratose, hyaline, long-rectangular or polygonal,

(45.0-)60.0-80.0 x 27.5-37.5 pm, walls thin or slightly

thickened, in transverse section 20.0-27.5 pm thick, oil

bodies usually subdermal, almost filling cell, ± 32.5 x

30.0-40.0 pm, dark brown, globular to subglobular; along

margins (Figure 5G) mostly with 3 rows of cells, narrow-

est in outermost row, long rectangular, 17.5-37.5 x ± 5.0

pm, in next row 4- to 6-sided, 30.0-35.0 x ± 12.5 pm,

and in innermost row 30.0-52.5 x 17.5-27.5 pm; air pores

numerous, 192.5-262.5 pm distant from each other,

raised, compound, oval, 40.0-50.0 x 35.0^4-2.5 pm, en-

circled by 4-6(-7) concentric rings of cells, 2 or 3 above

epidermis and up to 4 projecting into air chambers (Figure

5F), innermost ring with collapsed cells, ± 5 pm wide,

uppermost ring with (3)4 or 5 cells, 37.5-62.5 x 7.5 pm,

cells in next ring 37.5^42.5 x 7.5-12.5 pm and in outer

ring, sometimes up to six, 42.5-62.5 x 15.0-22.5 pm (Fig-

ure 5D), inner opening with 4 or 5 cells, inside walls

mostly with granular deposit and strongly protuberant,

leaving only a small cruciate opening (Figure 5Ep Eo).

Assimilation tissue 75.0-100.0 pm thick, '/iQ-'/g as thick

as thallus, air chambers in a single layer, 112.5-187.5 pm

wide, bounding walls 3 or 4(5) cells high, 22.5-35.0 x

20.0- 27.5 pm, crowded with chlorophyllose filaments,

cells mostly irregularly shaped, 17.5-27.5 x 12.5-20.0

pm; storage tissue occupying ventral 7/b-9/|o of thickness

of thallus medianly, decreasing laterally, cells angular,

crowded together, 52.5-67.5 pm wide, walls pitted, central

area sometimes stained purple, sclerotic cells and mucilage

cavities absent in specimens seen, but reportedly sometimes

present; rhizoids mostly smooth, 17.5-27.5 pm wide, occa-

sionally pegged, 10.0-17.5 pm wide.

Median scales (Figure 51) mauve, in one row on either

side of midrib, body ± obliquely triangular, up to 1050

pm long, base arched and continuing into long ‘tail’, when

up to 3250 pm wide, inner cells elongated, 4-6-sided,

52.5- 87.5 x 20.0-25.0 pm, walls straight or somewhat

sinuous toward margin, cells in outer 8 or 9 rows with

thinner, often sinuous walls, orientation very irregular and

smaller in size, 15.0-55.0 x 12.5-20.0 pm, sometimes 3

or 4 marginal cells grouped together and slightly raised

in a little peak from centre of which 1 or 2 slender papillae

project, scattered throughout up to 25 oil cells, ± 37.5 x

27.5 pm, oil body almost filling cell, scale gradually nar-

rowing upwards and then ± 750 pm wide, abruptly deeply

constricted at join with appendage (Figure 5J j , J2), mar-

ginally orange-brown or purplish, internally pink or occa-

sionally hyaline, ovate, oblong or suborbicular, apically

rounded, acute or shortly apiculate, basally cordate, rarely

with small basal lobe, 810-875 x 650-750 pm, inner cells

5- or 6-sided, 65.0-77.5 x 37.5-52.5 pm, with 1, 2 or

more oil cells or even none; margins entire, crenulate or

slightly denticulate, bordered by 1 row of smaller cells,

quadrate to rectangular, sometimes walls sinuose, 17.5-37.5

x 10.0-27.5 pm, orientated parallel to or perpendicular or

oblique to margin, sometimes partly protruding, submarginal

cells ± 62.5 x 27.5 pm. Laminal scales (Figure 5Hj, fL)

mauve or internally mauve and externally hyaline, in one

row on either side of and lateral to, but alternating irregularly

with median scales, ± oblong, but not quite symmetric, base

slightly arched or oblique, 750-1575 x 500-650 pm. cells

in body of scale elongated, 4—6-sided, walls straight or sinu-

ose, 55.0-100.0 x 17.5-27.5 pm, with up to 10 scattered

oil cells, toward margins cells smaller, 17.5-32.5 x

15.0- 25.0 pm, their orientation very variable, a few along

inner sloping margin grouped together and forming indi-

vidual little peaks from tips of which slender papillae pro-

ject. Cupule margins (Figure 5K) with ciliate, ± triangular

lobes, 380^120 pm long and 200-240 pm wide at base,

apically up to 6 cells in a tapering, uniseriate, vertical

row, top cell conical, 25.0-32.5 x 15.0-20.0 pm, lower

cells, 20.0-40.0 x 20.0-25.0 pm, base of lobes broadened

to 6-9 adjacent cells, 4- or 5-sided, 27.5-35.0 x 22.5-27.5

pm, from margins of lobes on both sides, horizontally

projecting, l-3(4)-celled cilia, lower down cupule wall

several cell layers thick, exteriorly with 1- or 2-celled pa-

pillae, up to 37.5 pm long, and internally with numerous

oil cells.

Dioicous. Antheridiophore not seen as none available

for study. Arche goniophore arising from apex of terminal

segment of main or short lateral branches, raised on stalk.

10-1 1 mm long, partly reddish, in transverse section (Fig-

ure 5N), 750 x 700 pm, cortical cells small, 15.0-25.0 x

12.5- 27.5 pm, outer wall thicker, medullary cells rounded,

42.5^t5.0 x 35XM-2.5 pm, smaller cells in between and

at centre of stalk, 12.5-22.5 x 20.0-25.0 pm, with single

194

Bothalia 25,2 (1995)

FIGURE 5. — Marchantia paleacea Berlol. A-G, ihallus: A, dorsal lace of apical branch; B, ventral face of apical branch; C, l.s. ol branch; D, air

pore and surrounding cells from above; Ei, E2, air pores from below; F, t.s. ol air pore, dorsal epidermal cells and ait chamber, G, margin.

Hi, Hr, laminal scales; I, median scale; Ji, J 2 , appendages of median scales; K, cupule margin; L, ¥ receptacle from side, M, ¥ receptacle

from above; N, t.s. of ¥ stalk. O-Q, scale: Oi, O2, from foot of ¥ stalk; P1-P3, from along ¥ stalk; Qi, Q2, from ¥ receptacle. R, margin of

involucre. A, Ei, E2, G. I, Ji, J2, Rankin 206; B-D, F, Hi, H2, K, S.M. Perold 3264\ L-R, C. Tavares L1SU P 66716. Scale bars: A, B, L, M,

2 mm; C, 1 mm; D-G, 50 pm; H-K, N-R, 250 pm.

Bothalia 25,2(1995)

195

band of 4 small air chambers, ± 62.5 x 30.0 pm, and 2

rhizoid furrows, lined with purple-walled cells, ± 1 1 2.5 x

72.5 pm, scales at base of stalk (Figure 50], O2) con-

spicuous, brown, rounded or partly 2-lobed or irregularly

shaped, often bulging, without or with short apical ap-

pendage, 1025-1150 x 900-1750 pm, inner cells 5- or

6-sided, 52.5-70.0 x 25.0-32.5 pm, with up to ± 10 oil

cells, 30 x 20 pm, margins ± entire, cells quadrate or

rectangular or irregularly shaped, 22.5-47.5 x 15.0-20.0

pm, long axis orientated parallel or perpendicular to mar-

gins; scales along length of stalk (Figure 5P)— P3) hyaline

or purplish, filiform, up to 1750 pm long, toward apex ±

4 single cells in a row, 67.5-75.0 x 10.0-20.0 pm, lower

down 3 or 4 cells wide, 75.0-132.5 x 20.0-22.5 pm; re-

ceptacle (Figure 5L, M) ± 4 mm in diameter, nearly sym-

metric, dorsally with prominent median projection, deeply

divided into 8 lobes, up to 750 pm long, narrow at base

and some of them wider at ± truncate apex, basal sinus

± 100° wide; involucres with margins (Figure 5R) hyaline

or purplish, divided into narrow lobes ± 500 x 90 pm,

fringed with cilia 180-650 pm long, which consist of sev-

eral cells in a uniseriate row, apical cell ± 75 x 25 pm,

lower cells 82.5-92.5 x 20.0-30.0 pm, toward base often

with 1 or 2 serially arranged transverse cells on one or

both sides, originating at join between 2 successive cells;

scales of receptacle (Figure 5Q 1 , Qt) hyaline or purplish,

filiform, up to 2075 pm long, apically cells in a uniseriate

row, 62.5-85.0 x 12.5-22.5 pm, then 2 cells and eventu-

ally 4 cells wide, up to 92.5 x 25.0 pm. Spores and elaters

not available for study. Chromosome number: n = 9

(Bischler 1984, 1988, 1989a).

Marchantia paleacea has been known since the time

of Micheli ( 1729), who described and illustrated it; but it

was not accepted by Linnaeus (1753). After M. polymor-

pha, it was to become only the second species in the genus

to be recognized from Europe, and was described by Ber-

tolini (1817) from material collected in Italy.

It has been placed in subgenus Chlamidium on account

of the four rows of ventral scales which are restricted to

the median part of the thallus. It is assigned to the mono-

typic section Pcileaceae because of the shape of the female

receptacle, the staicture of the margins of the involucres

and the cupules, which have ciliate lobes (Bischler-Causse

1993a) with papillae externally.

The species has a circumtethyan distribution, ranging

from the southern states of the USA. Mexico and Central

America, to the Mediterranean, the Caucasus, the Hima-

layas, and to the Far East (Bischler 1988). In Africa (and

the neighbouring islands), it is known from Algeria, Ethio-

pia, the Azores. Terceira and Reunion. Bischler-Causse

(1993a) states that its presence on the Canary Islands, Ma-

deira and in Morocco needs confirmation. A specimen

from Madeira, Tavares (LISU) was seen by me, and its

archegoniophores were studied and are illustrated in Fig-

ure 5L-R. The presence of M. paleacea on Madeira is

thus confirmed. Bischler-Causse (1993a) thought that the

single specimen, Rankin 206 (BM), from the vicinity of

Pilgrim’s Rest, South Africa, that she had seen, might have

been mislabelled. A computer printout at PRE, however,

revealed that several other bryophytes were collected by

Rankin in the same area, with collecting numbers both

lower and higher than the one referred to, so that it was

unlikely to have been mislabelled. I have recently also

collected it near Pilgrim’s Rest (Figure 3) on a steep earth

bank of the Blyde River, where it grew down to the

water’s edge together with M. debilis. The plants were

sterile unfortunately, as was Rankin’s collection, but both

had cupules. As noted above, archegoniophores from Ma-

deira were used for the description and illustrations, but

no antheridiophores were available for study.

In the Far East a subspecies, M. paleacea subsp. dip-

terci (Nees & Mont.) Hatt., is recognized (Bischler-Causse

1989a). It is distinguished from subsp. paleacea by the

epidermal pores of the thallus usually being surrounded

by 7 or 8 rings of cells, by the frequent presence of non-

functional female receptacles and by the marginal cells of

the median scale appendage having the long axis oblique

to perpendicular to the margins (not parallel to).

Marchantia paleacea subsp. paleacea is regarded as

morphologically stable and can be distinguished from the

other species of subgenus Chlamidium by its cruciate epi-

dermal pores and its cupules which have margins with

ciliate lobes. The appendage of the median scales is ovate

to orbicular. Schuster ( 1992) regards M. berteroana as the

closest ally to M. paleacea , because it also has cruciate

epidermal pores and its cupules have margins with trian-

gular lobes bearing teeth (or cilia) as well. M. berteroana

is however, classified in subgenus Marchantia.

Marchantia section Chlamidium

Thallus with branches (6.0-)7.0-8.5 (-10.0) mm wide,

generally rather remotely spaced and narrowly divergent.

Dorsal epidermis without papillae; air pores with inner

opening bordered by cells, their inside walls convex to

nearly straight or with short, rounded processes. Median

scales with appendage ovate to orbicular or broadly tri-

angular, apically rarely obtuse, mostly acute, sometimes

shortly apiculate, basally rounded or cordate, width across

broadest part 375-530 pm, margins entire or sometimes

bluntly toothed; with 1 or 2 oil cells, rarely more numer-

ous. Cupules with ciliate margins, cilia up to 6 or 7 cells

long and 3 cells wide basally, exteriorly sometimes also

with several cilia.

Dioicous. Antheridiophore on stalk which mostly lacks

bands of air chambers but with 2 or 3 rhizoid furrows;

basally surrounded by quite large scales, often with an

appendage; receptacle palmate, shallowly to deeply dis-

sected into 6-8 (-10) rays, dorsal surface with or without

papillae. Arche goniophore on stalk having 2 bands of air

chambers and 2 rhizoid furrows; basally surrounded by

quite large scales, often with an appendage and rather

similar to median scales of thallus; receptacle with or

without small, median projection dorsally, shortly divided

into 9-1 1 rather flat lobes, sometimes basally narrow, wid-

ening slightly toward truncate apex; involucre margins

shortly to long ciliate. Spores 20-30 pm in diameter, or-

namentation on distal face with wide, irregular, smooth

ridges forming incomplete areolae filled with nodules;

proximal face entirely covered with nodules.

Of the southern African taxa, only M. pappeana be-

longs in this section. The ornamentation of its spores is

196

Bothalia 25,2 (1995)

FIGURE 6. — Marchantia pappeana Lehm. subsp. pappeana. A-G, thallus. A, B, apical branch: A, dorsal face; B, ventral face. C, t.s. ol branch.

D-F, air pore from above; Ei, E2, from below; F, t.s. of dorsal epidermal cells and part of air chamber. G, margin. H-J, scales: Hi . H:, laminal;

1 1 , h, median; J1-J3, appendages of median scales. Ki. K2, margins of cupules; L-Q, receptacles: Li, L2, ¥ from above; M, ¥ from side; N,

cf; O, t.s. of ¥ stalk; P, t.s. of <f stalk; Q, median scale of cf. R-V, scales: R, alongd1 stalk; S, foot of d" stalk; T1-T3, ¥ receptacle; U, toot ot

¥ stalk; V, along ¥ stalk. W, margin of 0" ray; X, margin of involucre. A. G, Hi, I2, J2. Ki, Li, L2, M, T1-T3, X, Koekemoer 1050', B, C, F,

J3, K2, N, Perold & Koekemoer 2918: D, O, V, hurgoyne 2068: Ei, S, H. Anderson 1261 ; E2, 1 1 , J 1 , H2, Perold & Koekemoer 2841 ; P-R, W,

Hilliard & Burn 15460: U, H. Anderson CHI 3278. Scale bars: A, B, L-M, 2 mm; C, I mm; D-G, 50 pm; H-V, X, 250 pm; W, 100 pm.

Bothulia 25,2(1995)

197

referred to as the chenopoda type; there are, however, two

other spore coat ornamentation types in the section (Bischler-

Causse 1989a). The section is distinct in the shape of the

female receptacle and in the ciliate margins of the involucres

and cupules.

4. Marchantia pappeana Lehm. in Novarum et mi-

nus cognitarum stirpium, Pugillus X; 21 (1857); Bischl.:

76 (1993a). Type: South Africa, ‘In Prom. B. S. leg. Pappe’,

ex herb. Lehmann (RO, holo.?; G! ex herb. Univ. di Roma).

M. pappeana subsp. pappeana Bischl.: 82 (1993a).

M. flavescens Steph. in Bonner: 107 (1953). Type:. Fernando Po, 1911

Mildbraed 6275 (G).

M. pan’iloba Steph.: 305 (1895); Vanden Berghen: 46 (1954); S.W.

Amell: 56 (1963). Type: Uganda. Runssoro, um 2 800 m, 10 Juli 1891,

Stuhlmann 2368a (G. lecto. fide Vanden Berghen: 46 (1954); BM,

isolecto. fide Bischl.: 82 (1993a)).

M. planiloba Steph.: 90 (1886); Henriques: '153, 154’; 181, 182

(‘1886’. 1887). Type: Sao Tome, Cachoeira do Rio Manuel Jorge, circa

S. Nicolau, 800 m, 1885. Moller 32 [G, lecto. fide Vanden Berghen: 52

(I960)].

M. planiloba Steph. var. walteri Burgeff: 276 (1943). Type: Tanzania,

‘Nderema in Ost-Usambara, etwa 1 000 m, leg. H. Walter' , syn. fide

Bischl.: 83 (1993a).

M. stephanii Vanden Berghen: 50 (1954) [= M. umbellata Steph.:

305 (1895), nom. illeg.]. Type: Tanzania, Usambara, Holst 692 (FH, G),

syn. fide Bischl.: 83 (1993a).

M. wilmsii Steph: 126 (1892). Type: South Africa, Transvaal. McLea

in Rehmann Hep. austro-afr. exs. I [PC, lecto. fide Vanden Berghen: 44

(1954); BM!, G, NY, S, isolecto. fide Bischl.: 84 (1993a)].

M. winkleri Steph. in Bonner: 112 (1953). Type: Cameroon, Winkler

270 (G).

Thallus medium-sized to large, rather distantly and ir-

regularly branched, not ribbon-like or flat, apical segments

oblong to obovate, light green to yellowish green, often

purplish pink or deeply purple toward margins, or irregu-

larly smudged with purple on dorsal surface, without dark

median band (Figure 6A), margins undulate, scalloped,

usually hyaline, occasionally purple, entire, mostly crisped;

pores quite large and subdorsal air chamber walls visible

from above and flecked with whitish oil cells, when wet;

thallus margins rarely raised, not incurved when dry; in

densely crowded and irregularly overlying mats. Branches

with total length up to 50 mm, terminally 10-15 mm long

and successive branches generally 10-13 mm apart,

mostly narrowly to moderately divergent, (6.0-)7.0-8.5

(-10.0) mm wide, 740-925 pm thick over midrib, laterally

thinning out into wings (Figure 6C); apex notched, with

appendages of purple-brown median scales recurved over

edge (Figure 6A); margins acute, thin; flanks sloping

obliquely; ventral face medianly keeled, with rows of pur-

ple scales on either side, extending over 40-75% of thallus

width (Figure 6B).

Dorsal epidennal cells unistratose, very rarely bi-

stratose in patches, hyaline, mostly 5- or 6-sided (Figure

6D), 42.5-80.0(- 105.0) x 22.5-37.5 pm, thin-walled, not

thickened at comers, rarely containing an oil body, in

transverse section 25.0-32.5 pm thick; along margins

1-2(3) rows of cells, mostly narrowly rectangular (Figure

6G), 22.5-30.0 x 10.0-17.5 pm, sometimes shorter than

broad, ± 17.5 x 35.0 pm, innermost third row of cells

larger, 27.5-32.5 x 27.5 um; air pores fairly numerous, dis-

tance between them (200-)325-575 pm, raised, com-

pound, oval or round, ( 65— )90— 1 05 x 70-100 pm, sur-

rounded by 6 or 7 concentric rings of cells, 3 or 4 above

epidermis and 2 or 3 projecting into air chambers (Figure

6F), innermost ring of upper cells, ± 50.0 x 7.5 pm, next

2 rows of cells ± 40.0 x 7.5-10.0 pm, partly overlying

the outer, larger ring of 7 or 8 polygonal cells, up to 75.0

x 17.5 pm, inner opening with 4 or 5(6) bulging cells

(Figure (6E|, Eo), 30.0-42.5 x 12.5-15.0 pm. inner walls

convex or almost straight, densely covered by dark granules.

Assimilation tissue 60-90 pm thick, V^-’/g as thick as thal-

lus medianly, air chambers in a single layer, with 2 or 3 cells

in vertical bounding walls, 27.5^42.5 x 17.5-30.0 pm, oc-

casionally containing an oil body, ± 35 pm in diameter, air

chambers 275-387 pm wide, crowded with chlorophyllose

filaments (Figure 6F), 1-3-celled, often irregular in shape,

20.0-30.0 x 15.0-17.5 pm; storage tissue occupying ventral

7/8— n/|2 of thickness of thallus medianly, decreasing later-

ally, often with a wide, upper, central, purple-stained band,

cells angular, closely packed, 25-50 pm wide, rather smaller

below, but much larger laterally, sclerotic cells present or

absent, mucilage openings few, up to 200 pm wide, or ab-

sent; rhizoids some smooth, width 1 5 — 45 pm, others pegged,

10-15 pm wide.

Median scales (Figure 61 1, L) purple, in one row on

either side of midrib, body obliquely triangular, 800-1 125

pm high. 1430-1875 pm wide across arched base, mostly

ending below in a long ‘tail’, central cells elongated, 5-

or 6-sided, walls often sinuous, 92.5-107.5 x 30.0-37.5

pm, smaller toward margins and longer and narrower in

‘tail’, oil bodies scattered, margins sometimes with pro-

tniding papillae, ± 50.0 x 12.5 pm, scale narrowing up-

wards, deeply constricted where joined with appendage

(Figure 6J i— J3), purple brown or reddish, ovate to orbicu-

lar or broadly triangular, apically rarely obtuse, mostly

acute, sometimes shortly apiculate, 520-550 x 375-530

pm, apiculus with a vertical row of 2 or 3 cells, 35.0-62.5

x 20.0-27.5 pm, basally rounded or cordate, margins en-

tire or sometimes bluntly toothed, teeth 32.5-50.0 x

20.0-30.0 pm, marginal cells 50.0-62.5 x 17.5-37.5 pm,

only slightly smaller than inner cells, 4-7-sided, 75.0-87.5

x 32.5-42.5 pm, oil cells solitary, rarely more numerous.

Laminal scales (Figure 6Hj, Fh) mauve with hyaline base,

in one row at lateral sides of median scales, obtusely tri-

angular, (750— )980— 1 125 x (360-)630-1075 pm, cells

4-7-sided, 45.0-70.0 x 20.0-27.5 pm, smaller toward

margins, oil cells scattered, appendage lacking, margins

often with protruding papillae. Cupules with margins cili-

ate (Figure 6Kj, Kj), cilia (1 22.5-) 180.0-3 15.0 pm or up

to 6(7) cells long, top cell 45.0-67.5 x 15.0 pm. lower

cells 30.0-75.0 x ( 1 7.5— )22.5— 32.5 pm, basal cells ± 50.0

x 62.5 pm, sometimes 2 adjoining, ± 75.0 x 17.5-30.0

pm, exterior surface occasionally also ciliated.

Dioicous. Antheridiophore arising mostly from apex of

terminal segment of main branch, raised on stalk,

9— 1 8(— 32 ) mm long, diameter 775-925 pm, in transverse

section (Figure 6P) with one row of small cortical cells,

(1 2.5-) 15.0-20.0 x (12.5— )1 5.0—17.0 pm, outer wall

slightly thickened, medullary cells larger, up to 60 pm

wide, but in between with smaller cells, ± 20 pm wide,

air chambers mostly absent, with 2(3) rhizoid furrows, ±

150 pm wide; scales at base of stalk (Figure 6S), 1 or 2,

hyaline, broadly triangular, 1000-1150 x 550-750 pm.

198

Bothalia 25,2 (1995)

FIGURE 7. — SEM micrographs of spores. A-E, Marchantia pappeana, H. Anderson CHI 3278: A, distal face; B, part of distal face much enlarged;

C, side view; D, proximal face. E. part of elater, much enlarged. F-I, M. debilis , Preusss.n.: F. distal face; G. part of distal face much enlarged;

H, proximal face; I, part of proximal face, much enlarged. A, x 1233; B, x 2340; C, D, x 1200; E, x 740; F, x 1412; G, I, x 5148; H, x 1373.

sometimes with appendage, ± 675 x 500 pm, partly or

not constricted at base, cells 75-80 x 30-55 pm, slightly

smaller lower down; scales along length of stalk (Figure

6R). narrowly triangular and tapering toward apex, up to

1300 pm long, base ± 175 pm wide, cells 62.5-100.0 x

27.5- 37.5 pm; receptacle 9-14 mm in diameter, palmate,

divided into 6-8 rays (Figure 6N), 1. 7-3.4 mm long, ±

3.5 mm wide, basal sinus 120°-140°, margins of rays red-

dish, undulating, entire (Figure 6W), outer cells long and

narrow, ± 27.5 x 10.0 pm, here and there alternating with

wider cells, 25.0-37.5 x 25.0 pm; median scales on ven-

tral side of rays brownish, obtusely triangular, body ± 750

x 500 pm. cells 55-75 x 25 pm, mostly with appendage

tapering to acute apex (Figure 6Q), 350-375 x 125-250

pm, cells ± 62.5 x 37.5 pm.

Arche goniophore generally arising from apex of termi-

nal segment of main branch, raised on stalk, 1 6— 32(— 60)

mm long, diameter 850-1050 pm, wider across chambers,

± 1250 pm, in transverse section (Figure 60) with one

row of small, rounded, cortical cells, 15.0-25.0 x

12.5- 25.0 pm, outer wall slightly thickened, medullary

cells angular, ± 50 pm wide, with smaller, ± 20 pm wide

cells in between, air chambers in 2 separate and opposite

bands ± 500 pm wide, containing several small air cham-

bers, ± 62 pm high, also with 2 rhizoid furrows, ± 85 x

195 pm; scales at base of stalk, 1 or 2 present (Figure

6U), larger, ± 1350 x 850 pm, shape irregular, cells

87.5- 107.5 x 25.0 pm, often with ± triangular appendage,

slightly constricted at base or hardly so, 750-900 x

500-550 pm, cells 92.5-137.5 x 25.0-37.5 pm, 4-7-sided,

walls straight, shorter along margin, 37.5-42.5 x

37.5- 50.0 pm; scales along length of stalk (Figure 6V),

hyaline strands, 1300-1950 x 70-200 pm, tapering, api-

cally filamentous; receptacle ( 6.5— )8.0— 1 1.0 mm in di-

ameter, sometimes with small, rounded, median projection

dorsally, shortly and ± symmetrically divided into 9-11

lobes (Figure 6L j , L2, M) (occasionally I or 2 lobes re-

placed by male rays), 1.4-1. 7 mm long, base narrow,

1.0-1.25 mm, widening to 2. 1-3.0 mm, toward truncate

apex, basal sinus ± 30°; margins of involucres hyaline,

ciliate (Figure 6X), cilia delicate, sometimes collapsed,

187.5- 200.0 pm long, consisting of cells, 62.5-100.0 x

17.5- 25.0 pm, occasionally base broader, up to 37.5 pm

wide, inner cells angular, 50.0-62.5 x 32.5 pm, with nu-

merous large oil bodies; scales of receptacle mauve or

hyaline, tapering toward apex, sometimes abruptly so, oc-

casionally forked (Figure 6T1-T3), up to 3250 x 250 pm,

Bothalia 25,2(1995)

199

cells 4-7-sided, 75.0-137.5 x 37.5-40.0 |jm, filamentous

apices ± 450 pm long, with apical cell conical, ± 50 x

15 pm, lower cells ± 65 x 30 pm, several oil cells present,

± 50,0 x 32.5 pm, and marginally with unicellular papil-

lae, ± 37.5 x 15.0 pm. Spores 22.5-30.0 pm in diameter,

triangular-globular, yellow-brown, distal face (Figure 7A,

B) with irregular, ± smooth ridges, forming incomplete

areolae filled with nodules; proximal face (Figure 7C, D)

with faint triradiate ridge, each facet densely covered with

nodules, some discrete, others confluent, narrowly winged,

margin entire. Elaters yellow-brown, up to 825 x 7.5 pm,

tapering toward ends, 5.0 pm wide, bispiral (Figure 7E).

Chromosome number, n = 18 (Bischler-Causse 1993a).

Although M. pappeanci had already been described by

Lehmann in 1857 and was mentioned by Stephani

(1898-1900), the name has been neglected by subsequent

authors and specimens in most herbaria, including PRE,

are labelled as M. parviloba, M. planiloba or M. wilmsii.

The species is listed as M. parx’iloba in Magill & Schelpe

(1979). Several authors had suspected, however, that M.

pan’iloba was synonymous with M. planiloba (Vanden

Berghen 1965; Jones & Harrington 1983). Bischler-

Causse (1993a) found that only seven of the 18 specimens

cited by Stephani, belong to M. pappeana and of the 30

kept in his herbarium, only 1 1 belong here.

Amell (1963) mistakenly considered M. pappeana to

be a synonym of M. berteroana. He may, however, have

been misled by a specimen in S (where Lehmann’s origi-

nal collections are kept) which was labelled M. pappeana ,

but actually contains M. berteroana , as was shown by Bis-

chler-Causse (1993a). Amell (1963) merely surmised that

M. parviloba would be found in the northern parts of

South Africa, but he had probably only seen specimens

from Zaire [Congo], Tanzania [Tanganyika] and Zim-

babwe [Southern Rhodesia],

Marchantia pappeana often grows in the same locali-

ties as M. debilis. on vertical soil banks of streams, at

waterfalls, at sluice canals, very rarely on rotting wood or

on rocks, in open grassland or in forests, sometimes in

deep shade.

It is widely distributed in tropical Africa, Bischler-

Causse (1993a) reporting it from the Cape Verde Islands

to Ethiopia and south to southern Africa, generally at an

altitudinal range of 1 000-2 500 m. In southern Africa

(Figure 8) it is known from the Northern Province [North-

ern Transvaal] and Eastern Transvaal, Gauteng [PWV],

Swaziland, Kwazulu-Natal, eastern [Orange] Free State

and Lesotho. It has also been collected at Kirstenbosch

Botanical Garden a number of times and the type speci-

men is from Promontorium Bonae Spei.

Of the 50 specimens examined, 62% had cupules, 20%

had antheridiophores and 34% had archegoniophores; only

6% had both. Morphologically the species is variable, ex-

hibiting this trait even among specimens from the same

geographical area. Generally, however, it can be distin-

guished by being larger than M. debilis and by lacking a

dark median line on the dorsal surface of the thallus; its

median scale appendages are large and often marginally

toothed, with the inner cells large, presenting a ‘loose’

appearance; its cupules have longer cilia; the female re-

ceptacle is shortly divided into rays and the involucral

margin is ciliate; androgynous branches in the female re-

ceptacle are sometimes present.

Marchantia pappeana subsp. pappeana is distin-

guished from M. pappeana subsp. robusta , a close relative

in South India and Sri Lanka, by the latter having numer-

ous sclerotic cells and mucilage cavities in the thallus; oil

bodies in the median scale appendages are, however, ab-

sent.

Marchantia section Papillatae Bischl. in Cryp-

togamie, Bryologie et Lichenologie 10: 69 ( 1989); Bischl.:

99 (1993a). Type: M. papillata Raddi (PI, lecto., Bischl.:

95(1984).

Thallus with branches rather narrow, (2.1-)4.5-7.3 mm

wide, ribbon-like, often quite regularly spaced, moderately

to widely divergent. Dorsal epidermis without papillae;

air pores with inner opening bordered by cells, their inside

walls convex or straight. Median scales with appendage

orbicular or ovate, apically acute or apiculate, seldom ob-

tuse, basally rounded, width across widest part 275-340

pm, margins toothed; oil cells absent. Cupule margins

with short cilia, 3(4) cells long and I or 2 cells wide

basally, external surface without papillae.

Dioicous. Antheridiophore on stalk without or with 1

or 2 bands of much reduced air chambers and 2 rhizoid

fuiTows, basally surrounded by smaller, narrowly triangu-

lar scales, sometimes divided above into filamentous seg-

ments; receptacle smallish, palmate, asymmetric and

deeply divided into (4 — )5— 7 rays, dorsal surface without

papillae. Arche goniophore on stalk with single band of air

chambers (in African taxon) and 2 rhizoid furrows; basally

surrounded by smaller, ± triangular scales, gradually ta-

pering to filiform apex; receptacle smallish, with a

rounded median projection dorsally, deeply divided into

8-10 lobes, basally convex and costate, apically broad-

ened; involucre margins entire or crenulate. Spores 25-32

pm in diameter, ornamentation on distal face with irregu-

lar coarse ridges, forming incomplete areolae filled with

granules: proximal face densely verrucose.

FIGURE 8. — Distribution of M. pappeana var. pappeana. □: and M.

debilis. •. in southern Africa

200

Bothalia 25,2 (1995)

FIGURE 9. — Marchantia debilis Goebel. A-G, thallus. A, B, apical branch: A, dorsal face, with cupules; B, ventral face. C, t.s. of branch; D-F, air

pore: D, from above; Ei, E2, from below; F, t.s. of dorsal epidermal cells and air chamber. G, margin. Hi, H2, laminal scales; 1 1 , 12, median

scales; J 1 , J2, appendages of median scales; Ki, K2, margins of cupules. L-Q, receptacles: Li, L2, ¥ from above; M, ? from side; N, <f. O,

t.s. of d" stalk; P, t.s. of ¥ stalk; Qi, Q2, median scales of 0" ray. R-V, scale: R, along <f stalk; S, along ¥ stalk; T, base of ¥ stalk; U, base of

stalk; V 1-V3, ¥ receptacle. W, margin of 0" ray; X, margin of involucre. A, H. Anderson CHI223; B, R, U, W, X, Mogg 6172; C-G, H2,

.1 1 , Ki, Condy 90; Hi, li, I2, K2, Bester2544\ J2, Glen 1940; Li, L2, M, P, S, T, Dieterlen 850; N, Sim CH1345; O, S.M. Perold2891; Qi, Q2,

V1-V3, S.M. Perold 3048. Scale bars: A-B, L-N, 2 mm; C, I mm; D-G, 50 pm; H-K2, O-V3, 250 pm; W, X, 100 pm.

Bothalia 25,2(1995)

201

Only M. clebilis of the southern African taxa belongs

to this section. Bischler-Causse (1993a) states that it is

closest to the Asiatic species, M. emarginata subsp.

toscinci. The ornamentation of the spores is of the papillata

type. The section differs from sect. Paleaceae and sect.

Chlamidium in the shape of the female receptacle, with

the lobes costate basally and broadened apically.

5. Marchantia debilis Goebel in Organographie der

Pflanzen. 1. Bryophyten 2, edn 2: 901 (1915); Bischl.:

100 (1993a). Type: Cameroon, Urwaldgebiet von Bipindi,

Zenker, Flora von Kamerun exs. 1339 (BM!, BR, E, F, G,

GOET, M, S, iso.).

M. chevalieri Steph. in Bonner: 103 (1953). Type: Ivory Coast, Haute

Cote d'Ivoire, pays Dijola, environs de Ona, 4-1909, Chevalier s.n. (G,

PC).

Thallus smallish to medium-sized, rather flat and rib-

bon-like, apical segments oblong or broadly lingulate,

green or occasionally purplish all over, when much ex-

posed to bright sun, usually with a narrow, dark, broken,

longitudinal median band dorsally (Figure 9A), margins

narrowly reddish purple or hyaline, entire; pores quite

small, subdorsal air chamber walls visible from above,

when wet; thallus margins not raised or incurved, some-

times with slight longitudinal indentation along midline

and apex occasionally incurved, when dry; in crowded,

overlying patches or occasionally in partial rosettes, up to

75 mm across, repeatedly pseudodichotomously furcate,

often rather regularly. Branches with total length up to ±

38 mm, terminally 5-12 mm long and successive branches

5 — 7( — 9) mm apart, mostly moderately to widely divergent,

4.5-7. 3 mm wide, 675-850 pm thick over midrib, laterally

thinning out into wings (Figure 9C); apex notched, with

appendages of several reddish brown or purplish median

scales recurved over edge; margins acute, thin, occasion-

ally slightly crisped, flanks sloping obliquely; ventral face

medianly keeled and densely covered with long strands

of rhizoids, on either side with rows of purple scales ex-

tending over 70% or more of thallus width (Figure 9B).

Dorsal epidermal cells unistratose, rarely bistratose in

patches, mostly hyaline, polygonal, ( 4— )5— 7-sided,

37.0-62.5 x 22.0-32.0 pm, thin-walled, not thickened at

comers, in transverse section ± 17.5 pm thick, occasion-

ally containing a round oil body, ± 37.5 pm in diameter,

along margin 2 rows of narrow, ± rectangular cells,

(10.0-)20.0-30.0(-37.5) x (7.5— )12.5— 17.5 pm (Figure

9G), innermost third row of cells mostly larger, 30.0-37.5

x 15.0-22.5 pm; air pores quite numerous, (100-)

215-275(^475) pm distant from each other, raised, com-

pound, oval, 37.5-62.5 x 40.0-52.5 pm, surrounded by

(4)5 or 6 concentric rings of cells (Figure 9D), 2 or 3

above epidermis and 2 or 3 projecting into air chambers

(Figure 9F), innermost ring of upper cells ± 7.5 pm wide,

cells of middle ring ± 12.5 pm wide and outermost ling

with 6 or 7 larger cells, 35.0-62.5 x 17.5-22.5 pm, some-

what variable in shape and partly overlying epidermal

cells, inner opening (Figure 9Ep E2) with 4 or 5 rather

swollen cells, 20-25 x 35-42 pm, their inside walls ±

convex or nearly straight. Assimilation tissue up to 150

pm thick, ± '/g as thick as thallus, air chambers in a single

layer (Figure 9F), ( 1 05—) 1 50—300 pm wide, cells in

bounding walls ± 32.5 x 17.5 pm, crowded with densely

chlorophyllose, mostly 3-celled filaments, cells ± 20 x

15-25 pm; storage tissue occupying ventral 5/() of thick-

ness of thallus medianly, decreasing laterally, cells angu-

lar, medianly ± 50 pm wide, compact, much enlarged

laterally, sclerotic cells and mucilage cavities usually ab-

sent; rhizoids some smooth, 12.5-25.0 pm, others pegged,

5.0- 12.5 pm wide.

Median scales (Figure 91 1 , I2) brownish or purplish, in

one row on either side of midrib, body ± obliquely trian-

gular, up to 625 pm long, base slightly arched, its width

up to 3000 pm, central cells 4—6-sided, walls often sinu-

ous, 65.0-87.5 x 17.5-27.5 pm, elongating and narrowing

into long ‘tail’, ( 1 00.0—) 1 37.5—1 82.5 x 12.5-17.5 pm, oil

bodies scattered, upper margins of scale serrate, as it nar-

rows and becomes constricted where joined with append-

age (Figure 9Jp J2), mauve generally, orbicular or ovate,

apically acute or apiculate, rarely rounded, 350-375 x

275-340 pm, median cells 67.5 — 75.0 x 30.0-37.5 pm,

smaller at toothed margin, ± 42.5 x 25.0 pm, teeth conical.

I -celled, ± 37.5 x 20.0, or 2-celled, 45.0-62.5 x 15.0-42.5

pm, oil bodies absent. Laminal scales (Figure 9Hp Fh)

brownish, in one row on lateral sides of median scales, ±

ovate, rounded apically, lacking appendage, 450-560 x

430-670(-900) pm, sometimes with short ‘tail’ (Figure

9Hj), cells (50.0-)58. 5-67.5 x 17.5-25.0 pm, walls often

sinuous, oil bodies rare or absent, sometimes with mar-

ginal papillae, 50.0 x 12.5 pm. Cupules with margins

shortly ciliate (Figure 9K], K2), some cilia unicellular,

others 3 cells and up to 75 pm long, apical cells conical,

20-30 x 15 pm, basally 1 or 2 cells wide.

Dioicous. Antheridiophore arising from apex of termi-

nal segment of main branch or of short lateral branch, on

stalk (2.5— )7.0— 1 6.0 mm long, diameter 625-700 pm, in

transverse section (Figure 90) with 1 row of cortical cells,

17.5-22.5 x 10.0-17.5 pm, outer wall curved, medullary

cells angular, many up to 50 pm wide, but in between

smaller cells, only 17.5 pm wide, air chambers much re-

duced, in single band, rhizoid furrows 2, ± 160 pm wide;

scales at base of stalk (Figure 9LJ ), narrowly triangular,

750-1250 pm long and 150-500 pm wide at base, some-

times ± midway divided into (2)3 segments, only central

part elongated and tapering to filamentous apical string

of ± 5 single cells, top cell ± 30 x 10 pm, basal one ±

45.0 x 27.5 pm, cells in body of scale ± 92.5 x 27.5 pm,

margin with a few papillae, 37.5 x 7.5 pm; scales along

length of stalk filiform (Figure 9R), up to 1 100 x 50 pm,

2 or 3 cells wide, except for apical row of up to 5 single

cells, and sometimes basally with short cellular strands at

sides of long main one; receptacle 7.5-9.0 mm in diame-

ter, asymmetric, above without antheridial papillae, di-

vided into (4)5-7 rays (Figure 9N), 1. 5-3.0 mm long and

± 1.2 mm wide at base, narrowing slightly to tip, basal

sinus ± 130°, margins undulating, entire (Figure 9W), hya-

line, cells in 1 or 2 rows, rectangular, 17.5-32.5 x

10.0- 25.0 pm; median scales on ventral side of rays (Fig-

ure 9Qi, Q2) hyaline or mauve, obtusely triangular, up to

1000 x 670 pm, cells 4-6-sided, 35.0-62.5(- 100.0) x

20.0- 27.5 pm, at margin with some papillae, sometimes

with an apical appendage, constricted at its base or not,

narrowly to broadly triangular, 300-430-x 100-200 pm,

brownish with mauve cell walls, cells 50-60 x 25-35 pm,

smaller toward apex, 20 x 15-22 pm.

202

Bothalia 25,2 (1995)

Arche goniophore at apex of terminal segment of main

branch or of short lateral branch, raised on stalk, 9-16

mm long, diameter 400-550 pm, in transverse section

(Figure 9P) with 1 row of cortical cells, ± 12.5 x 22.5

pm, medullary cells 22-40 x 15-30 pm, with 2 rhizoid

furrows, ± 75 x 130 pm, and mostly a single band of air

chambers, the latter up to 85.0 x 37.5 pm each, rarely 2

bands or a split band present; scale(s) at base of stalk

(Figure 9T), brownish, ± triangular, 900-1000 x 400-450

pm, apex filiform with 3 or 4 single cells in a row, ± 32.5

x 12.5 pm, then widening gradually below, upper cells

thick-walled, 50.0-55.0 x 27.5-37.5 pm, thinner-walled

below, sometimes with short appendage at the side; scales

along length of stalk (Figure 9S) scattered, ± 700 x 112

pm, narrowing above to ± 35 pm wide, cells ± 75 x 15

pm; receptacle (3.5M-.5-7.CK9.0) mm in diameter, dor-

sally with a rounded projection medianly (Figure 9Lj, L2,

M) and deeply divided into 8-10 lobes, 0.8-1.25 mm long,

narrower at base and widening toward apex, basal sinus

40°-60°; margins of involucres hyaline, entire or crenu-

late, cells rectangular across to 5-sided (Figure 9X),

15.0-22.5 x 27.5-37.5 pm; scales of receptacle (Figure

9V1-V3) hyaline or yellow-brown or purple, 800-1000 x

330-370 pm, irregularly shaped, cells 75.0-100.0 x

37.5—42.5 pm, 5- or 6-sided, thick-walled, at basal margin

smaller, ± 32.5 x 15.0 pm, thin-walled and sinuous, at

apex 3 cells in a row, top cell conical, 40-75 x 10-12

pm, gradually widening lower down and sometimes with

filiform appendage at the side. Spores 25.0-32.5 pm in

diameter, faintly triangular-globular, brown, distal face

(Figure 7F, G) with irregular, coarse ridges, forming in-

complete areolae, spaces between filled with granules;

proximal face (Figure 7H, I) with triradiate mark hardly

visible, facets densely verrucose. Elaters brown, up to 295

x ± 7.5 pm, bispiral. Chromosome number: n = 9 (Bischler-

Causse unpublished).

Bischler-Causse (1993a) states that this species, which

is confined to Africa, has not been mentioned since its

description by Goebel (1915) and that it occurs in various

herbaria under 14 different names, but most commonly

under M. wilmsii , M. planiloba and M. chevalieri. She

also remarked that most authors considered M. chevalieri

to be a synonym of M. wilmsii , but the lectotype of M.

wilmsii (McLea!, syntype of Rehmann exsiccate) selected

by Vanden Berghen (1954), belongs to M. pappeana and

is an unfortunate choice.

The syntypes of M. wilmsii cited by Stephani are

McLea in Rehmann Hep. austro-a.fr exs. I and ‘Prope

Lydenburg et Greytown Dr Wilms’. Bischler-Causse

(1993a) states that the McLea specimens are male and

correspond to M. pappeana ; the Wilms specimens from

Lydenburg are female with cupules, or are sterile and also

belong to M. pappeana , whereas the Greytown specimens

are male with cupules and contain M. debilis. Bischler-

Causse (1993a) concludes that Stephani probably had M.

pappeana in mind, and not M. debilis, since he clearly

described M. pappeana under the epithet M. wilmsii. In

Stephani’s herbarium, she found that 13 of the 16 'M.

wilmsii ’ specimens belong to M. pappeana and only three

to M. debilis.

It is indeed fortunate that Bischler, with her long and

intimate experience with Marchantia species, was able to

resolve the taxonomy of this species, since neither Vanden

Berghen ( 1 954, 1 965 ), nor Arnell ( 1 963 ) nor Jones & Har-

rington (1983) managed to do so. Jones & Harrington

( 1 983) regarded 'M. wilmsii ’ as the commonest Marchan-

tia species in tropical Africa and readily recognizable by

the presence of the dark dorsal line. Specimens held at

PRE, that belong here, were mostly identified as M. wilm-

sii. The species is also listed as such in Magill & Schelpe

(1979) and in Arnold & De Wet (1993).

Marchantia debilis generally grows on damp soil on

vertical stream banks or waterfalls, on mud (or occasion-

ally on stones) of stream beds, at weirs, or sluice canals,

on stone dam walls kept wet by spray, and on soil over-

lying sandstone or granite, in open grassland or in forests,

sometimes in partial shade.

It is widely distributed in Africa, Bischler-Causse

(1993b) stating it to occur from Morocco to South Africa,

as well as on Reunion and also in the eastern part of

Madagascar. In southern Africa (Figure 8) it occurs in the

summer rainfall areas of the Northern Province [Northern

Transvaal] and Eastern Transvaal, Gauteng [PWV],

Swaziland, Kwazulu-Natal, eastern [Orange] Free State,

Lesotho and Eastern Cape.

Most of the 98 specimens examined had cupules, but

only 10% had antheridiophores and 16% had archegonio-

phores; 3% had both.

Sterile plants of M. debilis can be distinguished from

M. pappeana , the species it has frequently been confused

with, by the smaller size of the rather ribbon-like thallus,

by the dark median line on the dorsal face of the thallus,

by the smaller appendages of the median scales and by

the shortly ciliate or almost entire margins of the cupules.

Fertile plants should present no problem to identify as the

smallish male and female receptacles, the latter with

deeply divided lobes, are quite distinctive and the margin

of the involucre is entire.

ACKNOWLEDGEMENTS

I wish to sincerely thank Dr H. Bischler-Causse for

refereeing this paper and for her helpful suggestions; the

curators of BM, BOL, BR, G, LISU and W for the loan

of specimens; my colleagues at NBI for collecting speci-

mens; the artist, Mrs M. Steyn; the typist, Mrs J. Mul-

venna, and the photographer, Mrs A. Romanowski, for

their valued contributions.

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LEMAN, D.S. 1823. Cryptogames. In Dictionnaire des sciences

naturelles dans lequel on traite methodiquement des differents

etres de la nature.

LINNAEUS, C. 1753. Species plantarum. Bradbury & Evans, London.

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Africa. Memoirs of the Botanical Survey of South Africa No. 43:

1-39.

MARCHANT, M. (fd.) 1739. Nouvelle decouverte des fleurs et des

graines d’une plante rangee par les botanistes sous le genre Li-

chen. Memoire de I’Histoire de P Academic Royale des Sciences,

Paris 1713: 229-234.

MARKHAM. K.R.. PORTER, L.J. & CAMPBELL, E.O. 1977. The

usefulness of flavonoid characters in studies of the taxonomy and

phylogeny of liverworts. Bryophytorum Bibliotheca 13: 387-398.

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Berteroana Lehm. et Lindenb. und Marchantia tabularis Nees ab

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SPECIMENS EXAMINED

H. Anderson 1223 (5), CH4497 (5), CH4501 (4), CH 13278 (4),

CHI 3279 (5) PRE. S.W Amell 1058 (2) BOL (sub M. tabularis)-, 1075

(2) BOL; 1234 (5) BOL.

Barnard CH 1315 (sub M. tabularis) (2) PRE. Bester 2544 (5) PRE.

Bews CH1321 (sub M. tabularis) (2). CH1348 (5) PRE. Bingham 8181

(5) (Zambia) PRE. Bolus CH1319 (sub M. tabularis) (2) PRE. Bosnian

CH195 (5) PRE. Bottomley CHI 335 (sub M. wilmsii) (4) PRE. Breutel

s.n. (2) W. Burgoyne 1588, 2068 (4) PRE. Bunt Davy 435 (5) PRE.

Child 262 11 (2) BOL. Cholnoky 929 (sub M. wilmsii) (4) PRE. Condx

90 (5), CH 13638 (4) PRE.

204

Bothalia 25,2 ( 1995)

Dieterlen 850 (sub M. polymorpha ) (5) PRE. Doidge CH54 (2) PRE.

Drege s.n., 8286 (2) W. Duthie CH 1320 (sub M. tabularis ) (2) PRE.

Ecklnn 8284, 8289 W; G546, G2072/7G (typus) (2) G. Edwards

CHI 346 (sub M. wilmsii) (4) PRE. Esterhuysen 34979a (2) BOL. Eyles

CH1311 (sub M. tabularis) (2) PRE.

Garside 6186 (2) PRE. Geldenhuys 1061, 1332 (2) PRE. Ger-

mishuizen 4411, 6031 (5) PRE. Gerstner 583 (5) PRE. Glen 1690, 1940,

2189, 2207 pp. (5) PRE; 2207 pp. (4) PRE; 3007 (5) PRE; 3009, 3012,

3125a (4) PRE; 3468, 3728 (1) PRE. Guthrie 52555 (2) BOL.

Hepburn CH1223, CH1343 (5) PRE. Hilliard & Bunt 15460 (sub

M. wilmsii) (4) BOL.

Kluge 2442 (5) PRE. Koekemoer 1049 (5), 1050 (4) PRE. Krauss

8291 (sub M. contracta) (2) G.

Lambert 7 (5) PRE. Lanham 72 (sub M. polymorpha) (4) PRE.

Leendertz 755c (sub M. polymorpha) (5) PRE. Lubenau-Nestle SA269

(5) PRE.

McLea in Rehmann exs. 1 (sub M. wilmsii) (4) BM. Marais ( Duthie

5163) (sub M. wilmsii) (4) BOL. Mogg CH159 (2) PRE; CH1193,

CH4150, 4227, 6172 (5) PRE. Moonsammy 17 (NH 16194) (sub M.

wilmsii ) (4) PRE. Morley 298 (2) PRE.

Oliver 54680 (2) BOL. Onderstall 314 (5) PRE.

Pappe s.n. (typus) (4) G, RO. Pattison 14479 (2) BOL. J.J. Perold,

CH13644 (4) PRE. S.M. Perold, 41, 154, 164, 166, 168, 245, 1073 (5)

PRE; 2500 (4) PRE; 2549 (2) PRE; 2675 (Malawi), 2701, 2891 (5) PRE;

2926 (4) PRE; 2945, 3048, 3239, 3245 (5) PRE; 3246, 3250, 3251 (4)

PRE; 3252 (5) PRE; 3260a (4) PRE; 3261-3263 (5) PRE; 3264, 3265

(3) PRE; 3266 (5) PRE; 3267, 3268 (4) PRE. Perold & Koekemoer 2841,

2848 (4) PRE; 2869, 2875, 2893, 2897 (5) PRE; 2918 (4) PRE; 3181

(5) PRE; 3181a (4) PRE; 3248 (Ledbury, England), 3249 (Wageningen,

Netherlands), CHI 3640 (1) PRE. Pillans 4047, 4048, 4233 (2) BOL.

Playford CH 1324 (sub M. wilmsii) (4) PRE. Pole-Evans 461 (5), CHI 351

(sub M. wilmsii) (4) PRE. Pone r CH3586 (5) PRE. Potts 27, CHI 3 12

(2) PRE. Preiss 8282 (2) PRE. Preuss 1192 (Cameroon) (5) BM.

Rankin 206 (3) BM. C. Reid 816 (5) PRE. Reichenbach 354137 (2)

W. Reinecke CHI 74 (5) PRE. Rennie & Lambert 22 (sub M. polymorpha)

(4) PRE. Rogers TM 2269 (sub M. polymorpha) (4) PRE.

Scheepers 562 (sub M. wilmsii) (4) PRE. Schelpe 2048 (5) BOL;

4226 (sub M. wilmsii) (2) BOL; 3745 (sub M. wilmsii (4) BOL; 5359

(sub M. parviloba) (Zimbabwe) (4) BOL; 25367, 52542 (sub M. wilm-

sii) (2) BOL. Scott CH3693 ( 5) PRE. T.R. Sim CH1313, CH1316-

CH1318, CH1322 (sub M. tabularis) (2) PRE; CH1323 (sub M. wilmsii)

(4) PRE; CHI 325-CH 1327, CH1329, CH1330, CH1333 (5) PRE;

CH1339, CHI 341 (sub M. wilmsii (4) PRE; CH1342, CH1344, CH1345

(5) PRE; CH1347 (sub M. wilmsii) (4) PRE; CH1352, CH1353 (5)

PRE; CHI 356, CHI 358 (sub M. wilmsii) (4) PRE. SL CHI 308 (2) PRE.

Smook 8746 (2) PRE. Stokoe 9468 (sub M. wilmsii) (2) BOL. Strauss

CH13641 (4) PRE. Symons CH1337 (5) PRE.

C. Tavares, P 66716 (3) LISU. Theron 1339 (5) PRE. Thorne

CHI 309, CHI 544, CH3153 (2) PRE.

Univ. Durban-Westville 54 (5) PRE.

Van Breda 1116 (2) PRE. Van der Bijl CH1332, CH1349, CH1354,

NHI6171, NH 16175 (5) PRE. Van Rooy 1012 (5), 1653 (4), 2624 (5)

PRE. A.E. Van Wvk 2066 (2) PRE. Y. Van Wyk 883 (2) PRE. Veltman 3,

5 (4) PRE; 12 (5), 90 (4) PRE. Venter 12200 (5) PRE. Vorster 1785,

1789 (5) PRE.

Wager CH3805 (sub M. wilmsii) (4) PRE. Wagener CH 13341 (2)

PRE. Wawra 10 (2) W.

Zenker 1339 (5) (typus) BM; 1898 (5) BR.

Bothalia 25,2: 205-209 (1995)

New combinations and a complete list of Asparagus species in southern Africa

(Asparagaceae)

A.C. FELLINGHAM* and N.L. MEYER**

Keywords: Asparagaceae, Asparagus , new combinations, southern Africa, taxonomy

ABSTRACT

With only flower sexuality differing, Protasparagus can not be separated from Asparagus , neither at generic nor at

subgeneric level. No significant differences in the degree to which filaments were connivent were observed between species

placed under Myrsiphyllum and those under Protasparagus/ Asparagus. All southern African species of Asparagaceae are

therefore reinstated under Asparagus and listed here without recognition of subgenera. Several new combinations are made.

UITTREKSEL

Met slegs die geslagtelikheid van die blomme wat verskil, kan Protasparagus nie van Asparagus geskei word nie, nog op

genus- nog op subgenusvlak. Geen noemenswaardige verskille in die mate waartoe helmdrade saamneigend is, is waargeneem

tussen spesies wat onder Myrsiphyllum geplaas is en die onder Protasparagus/Asparagus nie. Alle Suider-Afrikaanse spesies

van Asparagaceae word dus heringestel onder Asparagus en hier gelys sonder erkenning van subgenusse. Verskeie nuwe

kombinasies word gemaak.

The family name Asparagaceae was proposed by Jus-

sieu (1789), soon after the genus Asparagus was named

by Linnaeus (1753). The debate on the division of the

genus into separate genera or subgenera, was started by

Willdenow (1808) with his description of the genus Myr-

siphyllum, which was countered by Roemer & Schultes’s

(1829) treatment of Asparagus as a single genus. Kunth

( 1 850) then not only subdivided the genus, but transferred

it back to the Liliaceae. Baker ( 1875) continued the trend

and it remained the standard treatment till Huber (1969)

and Dahlgren & Clifford (1982) once again advanced the

concept of separate genera and the family name, Aspara-

gaceae.

Obermeyer (1983, 1984) adopted Kunth’s (1850) divi-

sion of Asparagus s.l. into three separate genera: Myrsi-

phyllum Willd., Asparagus L. with unisexual flowers and

Asparagopsis Kunth for the bisexual southern African spe-

cies, in preference to Baker’s (1875) single genus with

three subgenera, Euasparagus Baker (= Asparagus),

Asparagopsis (Kunth) Baker and Myrsiphyllum (Willd.)

Baker. As Asparagopsis had been used by Montagne

(1840) for a genus of the Rhodophycaceae and was there-

fore an illegitimate homonym in Kunth’s application,

Protasparagus Oberm. (1983) was published. This stance

was also taken in the treatment of the family for the Flora

of southern Africa by Obermeyer & Immelman (1992).

Malcomber & Sebsebe (1993) came to the conclusion

that, with only flower sexuality differing, Protasparagus

and Asparagus could not be separated at generic level nor

*Stellenbosch Herbarium. National Botanical Institute, RO. Box 471,

Stellenbosch 7599.

**National Herbarium. National Botanical Institute, Private Bag X101,

Pretoria 0001.

MS. received: 1995-02-21.

even kept as separate subgenera, but rather, on the basis

of the shared character of free filaments, Protasparagus

and Asparagus belonged together in the subgenus Aspara-

gus. Neither could Myrsiphyllum be a distinct genus on

the strength of connivent filaments versus free ones in the

rest of the Asparagaceae. This character was, however,

sufficient to distinguish the taxon as a subgenus. They

therefore proposed that Asparagus be the only genus in

the Asparagaceae, with subgenera Myrsiphyllum and As-

paragus, the latter including the species previously placed

in Protasparagus.

We accept the treatment by Malcomber & Sebsebe

(1993) with reservation. The absence of A. juniperoides

Engl, and A. multituberosus R.A. Dyer from the list of

species investigated by these authors, prompted further in-

vestigation. As the close relatives of these two species are

in the subgenus with connivent filaments, it could be ex-

pected to find the same condition here. However, it could

not be demonstrated that filaments were markedly con-

nivent, and no significant differences were observed in

the degree to which filaments were connivent in the two

subgenera. Therefore, we prefer to refrain from dividing

the southern African species of the genus into subgenera.

In addition to the new combination made by Malcom-

ber & Sebsebe (1993), more new combinations as well

as reinstatements are necessary for the southern African

species of the genus Asparagus. The numbers next to the

species names listed alphabetically below (comb. nov.

numbers indented), are the species numbers allocated by

Obermeyer & Immelman (1992); those in Myrsiphyllum

being affixed with an ‘M’.

53. Asparagus acocksii Jessop in Bothalia 9: 74 ( 1966).

Protasparagus acocksii (Jessop) Oberm.: 243 (1983); Oberm. & Im-

melman: 57 (1992).

206

Bothalia 25,2(1995)

46. Asparagus aethiopicus L., Mantissa plantarum: 32

(1767).

Protasparagus aethiopicus (L.) Oberm.: 243 (1983); Oberm. & Im-

melman: 54 (1992).

19. Asparagus africanus Lam. , Encyclopedic methodique 1 :

295(1783).

Protasparagus africanus (Lam.) Oberm.: 243 (1983); Oberm. & Im-

melman: 35 (1992).

48. Asparagus aggregatus (Oberm.) Fellingham &

N.L. Mey. comb. nov.

Protasparagus aggregatus Oberm. in Oberm. & Immelman: 55 (1992).

M8. Asparagus alopecurus (Oberm.) Malcomber &

Sebsebe in Kew Bulletin 48: 73 (1993).

Myrsiphyllum alopecurum Oberm.: 85 (1984); Oberm.: 78 (.1992).

50. Asparagus angusticladus (Jessop) Fellingham

& N.L. Mey. comb. nov.

Asparagus aethiopicus L. var. angusticladus Jessop: 69 (1966). Prot-

asparagus angusticladus (Jessop) Oberm.: 243 (1983); Oberm. & Im-

melman: 56 (1992).

M2. Asparagus asparagoides (L.) W. Wight , Century Dic-

tionary 2: 845 (1909).

Medeola asparagoides L.: 339 (1753). Myrsiphyllum asparagoides

(L.) Willd.: 25 (1808); Oberm.: 78 (1984); Oberm.: 73 (1992).

5 1 . Asparagus aspergillus Jessop in Bothalia 9:71 (1 966).

Protasparagus aspergillus (Jessop) Oberm.: 243 (1983); Oberm. &

Immelman: 56 (1992).

2. Asparagus bayeri (Oberm.) Fellingham & N.L.

Mey. comb. nov.

Protasparagus bayeri Oberm. in Oberm. & Immelman: 21 (1992).

17. Asparagus bechuanicus Baker in W.T. Thiselton-

Dyer, Flora of tropical Africa 7: 429 ( 1 898).

Protasparagus bechuanicus (Baker) Oberm. in Oberm. & Immelman:

33 (1992).

63. Asparagus biflorus (Oberm.) Fellingham &

N.L. Mey. comb. nov.

Protasparagus biflorus Oberm. in Oberm. & Immelman: 63 (1992).

34. Asparagus buchananii Baker in Kew Bulletin 1893:

211 (1893).

Protasparagus buchananii (Baker) Oberm.: 243 (1983); Oberm. &

Immelman: 45 (1992).

8. Asparagus burchellii Baker in Journal of the Linnean

Society of London, Botany 14: 618 (1875).

Protasparagus burchellii (Baker) Oberm. in Oberm. & Immelman:

27 (1992).

4a. Asparagus eapensis L. var. capensis. Species plantarum,

edn 1: 314(1753).

Protasparagus capensis (L.) Oberm.: 243 (1983); Oberm. & Immel-

man: 23 (1992).

4b. Asparagus capensis L. var. litoralis Suess. & J. Karl

in Mitteilungen der Botanischen Staatssammlung, Miinchen

1 : 50 ( 1 950); Jessop: 44 ( 1 966).

Protasparagus capensis L. var. litoralis (Suess. & J. Karl) Oberm.

comb, illeg. in Oberm. & Immelman: 24 (1992).

47. Asparagus clareae (Oberm.) Fellingham & N.L.

Mey. comb. nov.

Protasparagus clareae Oberm. in Oberm. & Immelman: 54 (1992).

45. Asparagus coddii (Oberm.) Fellingham & N.L.

Mey. comb. nov.

Protasparagus coddii Oberm. in Oberm. & Immelman: 53 (1992).

59. Asparagus concinnus (Baker) Kies in Bothalia 6: 1 78

(1951a).

Asparagus africanus Lam. var. concinnus Baker: 619 (1875). Pro-

tasparagus concinnus (Baker) Oberm. & Immelman: 61 (1992).

41. Asparagus confertus K. Krause in Botanische Jahr-

biicher 51 : 449 (1914).

Protasparagus confertus (K. Krause) Oberm. in Oberm. & Immel-

man: 51 (1992).

18. Asparagus cooperi Baker in Gardeners’ Chronicle 1 :

818 (1874a).

Protasparagus cooperi (Baker) Oberm.: 243 (1983); Oberm. & Im-

melman: 33 (1992).

52. Asparagus crassicladus./cs.so/? in Bothalia9: 75 ( 1966).

Protasparagus crassicladus (Jessop) Oberm.: 243 (1983); Oberm. &

Immelman: 57 (1992).

M9. Asparagus declinatus L.. Species plantarum, edn I :

313 (1753).

Myrsiphyllum declination (L.) Oberm.: 86 (1984); Oberm.: 78 (1992).

39. Asparagus densiflorus (Kuntli) Jessop in Bothalia 9:

65 (1966).

Asparagopsis densiflora Kunth: 96 ( 1850). Protasparagus densiflorus

(Kunth) Oberm. in Oberm. & Immelman: 49 (1992).

2 1 . Asparagus denudatus ( Kunth ) Baker in Journal of the

Linnean Society of London, Botany 14: 606 (1875).

Asparagopsis denudatus Kunth: 82 (1850). Protasparagus denudatus

(Kunth) Oberm.: 243 (1983); Oberm. & Immelman: 36 (1992).

30. Asparagus devenishii (Oberm.) Fellingham &

N.L. Mey. comb. nov.

Protasparagus devenishii Oberm. in Oberm. & Immelman: 41 (1992).

69. Asparagus divaricatus (Oberm.) Fellingham &

N.L. Mey. comb. nov.

Protasparagus divaricatus Oberm. in Oberm. & Immelman: 67 (1992).

29. Asparagus edulis (Oberm.) Fellingham & N.L.

Mey. comb. nov.

Protasparagus edulis Oberm. in Oberm. & Immelman: 41 (1992).

10. Asparagus exsertus (Oberm.) Fellingham & N.L.

Mey. comb. nov.

Protasparagus exsertus Oberm. in Oberm. & Immelman: 29 (1992).

Bothalia 25,2 (1995)

207

32. Asparagus exuvialis Burch., Travels in the interior of

southern Africa 1 : 432 ( 1 822).

Protasparagus exuvialis (Burch.) Oberm.: 244 (1983); Oberm. & Im-

melman: 42 (1992).

32a. Asparagus exuvialis Burch, forma ecklonii

(Baker) Fellingham & N.L. Mey. comb. nov.

Asparagus ecklonii Baker: 615 (1875). Protasparagus exuvialis

(Burch.) Oberm. forma ecklonii (Baker) Oberm. in Oberm. & Immelman:

43 (1992).

33. Asparagus falcatus L., Species plantarum, edn 1:313

(1753).

Protasparagus falcatus (L.) Oberm.: 244 (1983); Oberm. & Immel-

man: 43 (1992).

M 1 2. Asparagus faseiculatus Thunb., Flora capensis. edn

2: 329 (1823).

Myrsiphyllum fasciculatum (Thunb.) Oberm.: 87 (1984); Oberm.: 80

(1992).

40. Asparagus filicladus (Oberm.) Fellingham &

N.L. Mey. comb. nov.

Protasparagus filicladus Oberm. in Oberm. & Immelman: 51 (1992).

1 la. Asparagus flavicaulis (Oberm.) Fellingham &

N.L Mey. comb. nov.

Protasparagus flavicaulis Oberm. in Oberm. & Immelman: 29

(1992).

1 lb. Asparagus flavicaulis (Oberm.) Fellingham &

N.L Mey. subsp. setulosus (Oberm.) Fellingham & N.L.

Mey. comb. nov.

Protasparagus flavicaulis Oberm. subsp. setulosus Oberm. in Oberm.

& Immelman: 30 (1992).

61. Asparagus fouriei (Oberm.) Fellingham & N.L.

Mey. comb. nov.

Protasparagus fouriei Oberm. in Oberm. & Immelman: 62 (1992).

28. Asparagus fractiflexus (Oberm.) Fellingham &

N.L. Mey. comb. nov.

Protasparagus fractiflexus Oberm. in Oberm. & Immelman: 40 (1992).

3. Asparagus glaucus Kies in Bothalia 6: 229 (1951b).

Protasparagus glaucus (Kies) Oberm.: 224 (1983); Oberm. & Im-

melman: 23 (1992).

31. Asparagus graniticus (Oberm.) Fellingham &

N.L. Mey. comb. nov.

Protasparagus graniticus Oberm. in Oberm. & Immelman: 42 (1992).

68. Asparagus intricatus (Oberm.) Fellingham &

N.L. Mey. comb. nov.

Protasparagus intricatus Oberm. in Oberm. & Immelman: 65 (1992).

M7. Asparagus juniperoides Engl, in Botanische Jahr-

biicher 10:3(1889).

Myrsiphyllum juniperoides (Engl.) Oberm.: 84 (1984); Oberm.: 78

(1992).

M4. Asparagus kraussianus (Kunth) J.F. Macbr. in Con-

tributions from the Gray Herbarium 56: 17 (1918).

Myrsiphyllum kraussianum Kunth: 107 (1850); Oberm.: 80 (1984);

Oberm.: 76 (1992).

42. Asparagus krebsianus (Kunth) Jessop in Bothalia 9:

74(1966).

Asparagopsis krebsiana Kunth: 93 (1850). Protasparagus krebsianus

(Kunth) Oberm.: 244 (1983); Oberm. & Immelman: 51 (1992).

1 6. Asparagus laricinus Burch., Travels in the interior of

southern Africa 1: 537 (1822).

Protasparagus laricinus (Burch.) Oberm.: 244 (1983); Oberm. & Im-

melman: 33 (1992).

23. Asparagus lignosus Burm.f, Prodromus florae cap-

ensis: 10 (1768).

Protasparagus lignosus (Burm.f.) Oberm. in Oberm. & Immelman:

37 (1992).

20. Asparagus longicladus N.E. Br. in Kew Bulletin

1921: 298 (1921).

Protasparagus longicladus (N.E. Br.) B. Mathew: 181 (1989);

Oberm. & Immelman: 36 (1992).

36. Asparagus lvnetteae (Oberm.) Fellingham &

N.L. Mey. comb. nov.

Protasparagus lynetteae Oberm. in Oberm. & Immelman: 47 (1992).

60. Asparagus maeowanii Baker in Journal of the Lin-

nean Society of London, Botany 14: 609 (1875).

Protasparagus maeowanii (Baker) Oberm.: 244 (1983); Oberm. &

Immelman: 62 (1992).

6. Asparagus mariae (Oberm.) Fellingham & N.L.

Mey. comb. nov.

Protasparagus mariae Oberm. in Oberm. & Immelman: 24 (1992).

57. Asparagus microraphis (Kunth) Baker in Journal of

the Linnean Society of London. Botany 14: 612 (1875).

Asparagopsis microraphis Kunth: 83 (1850). Protasparagus mi-

croraphis (Kunth) Oberm.: 244 (1983): Oberm. & Immelman: 59 (1992).

64. Asparagus minutiflorus (Kunth) Baker in Journal of

the Linnean Society of London. Botany 14: 616 (1875).

Asparagopsis minutiflorus Kunth: 89 (1850). Protasparagus minuti-

florus (Kunth) Oberm.: 244 (1983); Oberm. & Immelman: 64 (1992).

15. Asparagus mollis (Oberm.) Fellingham & N.L.

Mey. comb. nov.

Protasparagus mollis Oberm. in Oberm. & Immelman: 32 (1992).

25. Asparagus mucronatusicsso/? in Bothalia 9: 56 ( 1 966).

Protasparagus mucronatus (Jessop) Oberm. in Oberm. & Immelman:

38 (1992).

27. Asparagus multiflorus Baker in Journal of the Lin-

nean Society of London, Botany 14: 610 (1875).

Protasparagus multiflorus (Baker) Oberm.: 244 (1983); Oberm. &

Immelman: 39 (1992).

M 1 . Asparagus multituberosus R.A. Dyer in Bothalia 6: 442

(1954).

Myrsiphyllum multituberosum (R.A. Dyer) Oberm.: 77 (1984);

Oberm.: 73 (1992).

208

Bothalia 25,2 (1995)

49. Asparagus natalensis (Baker) Fellingham & N.L.

Mey. comb. nov.

Asparagus aethiopicus L. var. natalensis Baker: 272 ( 1 896). Prot-

asparagus natalensis (Baker) Oberm.: 244 (1983); Oberni. & Immelman:

56 (1992).

37. Asparagus nelsii Schinz in Bulletin de 1’Herbier Boissier,

Ser. 1,4, App. 3: 44 (1896).

Protasparagus nelsii (Schinz) Oberm.: 244 (1983); Oberm. & Im-

melman: 48 (1992).

56. Asparagus nodulosus (Oberm.) Fellingham & N.L.

Mey. comb. nov.

Protasparagus nodulosus Oberm. in Oberm. & Immelman: 59 (1992).

12. Asparagus oliveri (Oberm.) Fellingham & N.L.

Mey. comb. nov.

Protasparagus oliveri Oberm. in Oberm. & Immelman: 30 (1992).

M3. Asparagus ovatus T.M. Salter in Journal of South

African Botany 6: 167 (1940).

Myrsiphyllum ovatum (T.M. Salter) Oberm.: 79 (1984); Oberm.: 75

(1992).

38. Asparagus oxyacanthus Baker in Journal of the Lin-

nean Society of London, Botany 14: 625 (1875).

Protasparagus oxyacanthus (Baker) Oberm.: 244 (1983); Oberm. &

Immelman: 48 (1992).

22. Asparagus pearsonii Kies in Bothalia 6: 1 79 ( 1 95 1 a).

Protasparagus pearsonii (Kies) Oberm.: 244 (1983); Oberm. & Im-

melman: 37 (1992).

14. Asparagus pendulus (Oberm.) Fellingham &

N.L. Mey. comb.nov.

Protasparagus pendulus Oberm. in Obeim. & Immelman: 32 (1992).

55. Asparagus plumosus Baker in Journal of the Linnean

Society of London, Botany 14: 613 (1875).

Protasparagus plumosus (Baker) Oberm.: 244 (1983); Oberm. & Im-

melman: 59 (1992).

35. Asparagus racemosus Willd.. Species plantarum edn

4,2:152(1799).

Protasparagus racemosus (Willd.) Oberm.: 244 (1983); Oberm. &

Immelman: 45 (1992).

MIL Asparagus ramosissimus Baker in Gardeners’

Chronicle new series 2: 6 ( 1 874b).

Myrsiphyllum ramosissimum (Baker) Oberm.: 87 (1984); Oberm.: 79

(1992).

1. Asparagus recurvispinus (Oberm.) Fellingham

& N.L. Mey. comb. nov.

Protasparagus recurvispinus Obeim. in Oberm. & Immelman: 19 (1992).

26. Asparagus retrofractus L.. Species plantarum, edn 1:313

(1753).

Protasparagus retrofractus (L.) Oberm.: 244 (1983); Oberm. & Im-

melman: 39 (1992).

65. Asparagus rigidus Jessop in Bothalia 9: 64 ( 1966).

Protasparagus rigidus (Jessop) Oberm.: 244 (1983); Oberm. & Im-

melman: 64 (1992).

24. Asparagus rubicundus P.J. Bergius , Descriptiones

plantarum ex Capite Bonae Spei: 88 (1767).

Protasparagus rubicundus (P.J. Bergius) Oberm.: 244 (1983); Oberm.

& Immelman: 38 (1992).

M 1 0. Asparagus scandens Thunb ., Prodromus plantarum

capensium 1 : 66 (1794).

Myrsiphyllum scandens (Thunb.) Oberm.: 86 (1984); Oberm.: 79

(1992).

44. Asparagus sehroederi Engl, in Botanische Jahr-

biicher 32: 97 (1903).

Protasparagus sehroederi (Engl.) Oberm.: 244 (1983); Oberm. & Im-

melman: 53 (1992).

62. Asparagus sekukuniensis (Oberm.) Felling-

ham & N.L. Mey. comb. nov.

Protasparagus sekukuniensis Oberm. in Oberm. & Immelman: 63

(1992).

54. Asparagus setaceus (Kunth) Jessop in Bothalia 9: 5 1

(1966).

Asparagopsis setacea Kunth: 82 (1850). Protasparagus setaceus

(Kunth) Oberm.: 244 (1983); Oberm. & Immelman: 58 (1992).

9. Asparagus spinescens Steud. ex Roem. & Schult.,

Systema vegetabilium 7, 1 : 334 ( 1 829).

Protasparagus spinescens (Steud. ex Roem. & Schult.) Oberm. in

Oberm. & Immelman: 27 (1992).

58. Asparagus stellatus Baker in Journal of the Linnean

Society of London, Botany 14: 612 (1875).

Protasparagus stellatus (Baker) Obeim. in Oberm. & Immelman: 61

(1992).

5. Asparagus stipulaceus Lam.. Encyclopedic methodique

1:297 (1783).

Protasparagus stipulaceus (Lam.) Obeim.: 244 (1983); Oberm. &

Immelman: 24 (1992).

66. Asparagus striatus (L.f.) Thunb.. Prodromus plan-

tarum capensium 1: 65 (1794).

Dracaena striata L.f.: 204 (1781). Protasparagus striatus (L.f.)

Oberm.: 244 (1983); Oberm. & Immelman: 64 (1992).

7. Asparagus suaveolens Burch.. Travels in the interior

of southern Africa 2: 226 ( 1 824).

Protasparagus suaveolens (Burch.) Oberm.: 244 (1983); Oberm. &

Immelman: 25 (1992).

67. Asparagus subulatus Thunb.. Prodromus plantarum

capensium 1:66(1794).

Protasparagus subulatus (Thunb.) Oberm.: 244 (1983); Oberm. &

Immelman: 65 (1992).

43. Asparagus transvaalensis (Oberm.) Fellingham

& N.L. Mey. comb. nov.

Protasparagus transvaalensis Oberm. in Oberm. & Immelman: 52 ( 1992).

M6. Asparagus undulatus (L.f. ) Thunb., Prodromus plan-

tarum capensium 1 : 66 ( 1 794).

Dracaena undulata L.f.: 203 (1781). Myrsiphyllum undulatum (L.f.)

Kunth: 109 (1850); Oberm.: 83 (1984); Oberm.: 77 (1992).

Bothalia 25,2 (1995)

209

13. Asparagus virgatus Baker in Refugium botanicum 3:

t. 214(1870).

Protasparagus virgatus (Baker) Oberm. in Oberm. & Immelman: 31

(1992).

M5. Asparagus volubilis Thunb., Prodromus plantarum

capensium 1 : 66 (1794).

Myrsiphyllum volubile (Thunb.) Oberm.: 82 (1984); Oberm.: 76

(1992).

ACKNOWLEDGEMENTS

We gratefully acknowledge the help and guidance re-

ceived from Mr E.G.H. Oliver in the preparation of this

article.

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BROWN, N.E. 1921. New plants from tropical and South Africa col-

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BURCHELL, W.J. 1822. Travels in the interior of southern Africa 1.

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comparative study. Academic Press, London.

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ENGLER, H.G.A. 1903. Liliaceae africanae II. Botanische Jahrbiicher

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der Liliifloren. Mitteilungen der Botanischen Staatssammlung,

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KRAUSE, K. 1914. Liliaceae africanae V. Botanische Jahrbiicher 51:

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LAMARCK, J.B.A.P.M. DE 1783. Encyclopedic methodique 1. Mme

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de I'Herbier Boissier Set. 1,4, App. 3: 1-57.

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Africa australi a Dr S. Rehm et aliis collectarum. Mitteilungen der

Botanischen Staatssammlung, Miinchen 1: 50.

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WILLDENOW, C.L. 1808. Nahere Bestimmung einiger Liliengewachse,

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Ordnung des Linneschen Systems gehoren. Gesellschaft naturfor-

schender Freunde zu Berlin Magazin 2: 25, 26.

Bothalia 25,2: 211-231 (1995)

The taxonomic history of the Ricciaceae (1937-1995) and a classification of

sub-Saharan Ricciae

S.M. PEROLD*

Keywords: classification, groups, history, Ricciaceae, sections, species, subgenera, sub-Saharan Africa

ABSTRACT

The taxonomic history of the Ricciaceae (1937-1995) is reviewed. Over the years many attempts have been made to

subdivide and rearrange the taxa in this large and puzzling family, but consensus has still not been reached. Hepaticologists are

therefore urged to undertake more revisions worldwide, using modem methods of investigation to aid them in. hopefully,

coming to an eventual agreement in defining the limits of subgenera, sections, species and subspecies in the Ricciae. In addition,

a classification of sub-Saharan Ricciaceae. partially based on informal groups, is herein proposed.

UITTREKSEL

"n Oorsig van die taksonomiese geskiedenis van die Ricciaceae (1937-1 995 ) word gegee. Oor die jare heen is talle pogings

aangewend om die taksons in hierdie groot en moeilike familie op te deel en te herrangskik, maar eenstemmigheid is nog nie

bereik nie. Hepatikoloe word aldus aangemoedig om verdere hersienings wereldwyd te ondemeem en om modeme ondersoek-

metodes aan te wend, wat hopelik sal help om uiteindelik eenstemmigheid te bereik oor die omskrywing van subgenusse,

seksies, spesies en subspesies in die Ricciae. Verder word 'n klassifikasie van die Ricciaceae van Afrika, suid van die Sahara,

wat gedeeltelik op informele groepe gebaseer is, hier voorgestel.

CONTENTS

Introduction 211

Overview of regional studies 212

1. Duthie & Garside ( 1937, 1939): South Africa. 212

2. Frye & Clark ( 1937-1947): North America .. 212

3. Schuster (1949): central and western New York. 213

4. Schuster (1953): Minnesota and adjacent regions 2 1 3

5. Meijer (1951): The Netherlands 213

6. Muller (1951-1958): Europe 214

7. Amell (1956): Scandinavia 214

8. Jones (1957): tropical Africa 215

9. Pande & Udar ( 1958): India 215

10. Hassel de Menendez (1963): Argentina 215

11. Amell (1963): southern Africa 216

1 2. Vanden Berghen (1972): Zaire. Zambia 217

13. Jovet-Ast ( 1975): diverse areas 217

14. Campbell ( 1 975, 1 977): New Zealand 217

15. Grolle (1976, 1983): Europe and adjacent regions 2 1 7

16. Na-Thalang ( 1980): Australia 217

17. Jovet-Ast (1984): Australia 219

18. Volk (1983): Namibia and southern Africa

(and elsewhere) 219

19. Schuster ( 1984, 1985): diverse areas 219

20. Perold (1986): southern Africa 219

21. Schuster (1985): worldwide 219

22. Scott (1985): southern Australia 220

23. Volk & Perold (1986a): South Africa 220

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

MS received: 1995-04-28.

24. Jovet-Ast ( 1986): Mediterranean countries . . . 220

25. Damsholt & Hallingback (1986): Fennoscandia . 22 1

26. Volk & Perold (1990): South Africa 221

27. Smith (1990): Britain and Ireland 221

28. Van Melick (1991): The Netherlands 222

29. Volk & Perold; Perold & Volk; Perold ( 1984-

1991 ): southern Africa 222

30. Jovet-Ast (1991, 1993): Latin America 223

31. Schuster (1992): North America (and elsewhere) 224

Discussion 226

32. Perold ( 1995): sub-Saharan Africa 227

Acknowledgements 228

References 228

Index to taxa 229

INTRODUCTION

Upon recently re-reading Duthie & Garside’s (1937)

excellent account of the taxonomic history of the Ric-

ciaceae, which begins with John Ray (1696) and spans

the next ± 240 years, I was once more struck by the many

attempts to subdivide and rearrange the taxa in this large

and puzzling family during that time.

This publication aims to cover in chronological order

the most important regional studies of the Ricciaceae, that

span the period between the survey by Duthie & Garside

and the present.

It has now become accepted practice to recognize only

two genera in the family, the monotypic Ricciocarpos and

the species-rich Riccia. Both genera are characterized by

the simplicity of the sporophytes, which are immersed in

the thallus, and by the lack of elaters. The assimilation

212

Bothalia 25,2 ( 1995)

tissue in Ricciocarpos (formerly section Hemiseuma) con-

tains tiered air chambers, whereas in Riccia it has either

narrow air canals or rather wide, uniseriate air chambers.

Other major differences between Ricciocarpos and Riccia

are the following: Ricciocarpos is terrestrial or aquatic; in

the latter the ventral scales are much elongated and pen-

dant, with the margins serrate; single oil bodies are scat-

tered throughout the cells of the scales and thallus; the

antheridia and archegonia are in groups confined to the

median groove of the thallus. Except for the Riccia flui-

tans complex, which is often aquatic and has small ventral

scales that are never pendant, Riccia species are terrestrial

and the scales are sometimes evanescent, but mostly reach

to the thallus margins or extend above them; oil bodies

are absent and the gametangia are not strictly confined to

the median groove of the thallus.

Although there are intermediate forms with regard to

the width of the air spaces in Riccia , the two groups clas-

sified here are generally recognized respectively as sub-

genus (rarely as section) Riccia (formerly Lichenoides or

Euriccia ) and as subgenus Ricciella (formerly, occasion-

ally, as genus or as section). In the latter, two formal sub-

divisions have been recognized for a long time by the

position of the capsule in the thallus and by the direction

of liberation of the spores, i.e. the spores can be liberated

dorsally (section Spongodes Nees) or ventrally (section

Ricciella (A. Br.) Rchb.). Only informal subdivisions of

subgenus Riccia have, however, been recognized until re-

cently. as it was thought that not enough was known about

several aspects of many species in this subgenus, that oc-

curs worldwide, to make formal subdivisions.

OVERVIEW OF REGIONAL STUDIES

In the following overview diagnostic features are not

cited verbatim from each author’s text, but information is

extracted from the description and keys and then com-

bined in an abbreviated form more or less retaining the

author’s choice of words.

1 . DUTHIE & GARSIDE (1937, 1939): South Africa

In 1937 Duthie & Garside published their detailed his-

tory of the group. Although they were aware of at least

1 1 species in the genus Riccia from the Stellenbosch Flats

alone, they dealt with only five species in the section,

then referred to as Ricciella.

1.1 Genus Riccia: upper photosynthetic zone in majority

of species consisting of pillar-like vertical rows of cells

separated by narrow air spaces, terminal cells of vertical

rows usually larger and devoid of chlorophyll.

1.1. 1 Section ‘ Ricciella upper zone consisting of air

chambers, apparently strictly uniseriate and separated by

plates of chlorophyll-containing cells; chambers roofed

over by epidermis perforated by simple air pores which

enlarge and later on cause the dorsal surface to become

pitted; spores liberated from dorsal surface of thallus, but

deeply imbedded and clearly visible from below; com-

mon, annual species.

R. plana Taylor (synonym of R. crystallina L.); R. rau-

tanenii Steph. (synonym of R. cavernosa Hoffm. emend.

Raddi). Spores separate at maturity; monoicous.

R. cupulifera A.V. Duthie. Spores separate at maturity;

dioicous; heterothallic.

R. curtisii (James ex Austin) Austin (corrected author

citation); R. compacta Garside (air chamber layer rather

more compact and firmer). Spores retained in permanent

tetrads; dioicous; heterothallic.

Nowadays species that retain their spores in permanent

tetrads are assigned to subgenus Thallocarpus (Lindb.) Ast

(1975).

Duthie & Garside suggest that the Ricciaceae may be a

polyphyletic group, derived along several lines of descent.

Their work together was discontinued when Duthie retired

in 1939.

2. FRYE & CLARK (1937-1947): North America

Frye & Clark recognized three genera in the Ricciaceae,

namely Riccia, Ricciocarpns forth, var.) and Oxymitra.

2.1 Genus Riccia : thalli growing on damp or wet soil or

suspended in water but not floating; air spaces narrow or

chambered; antheridia and archegonia scattered; sporo-

phyte without involucre; air pores inconspicuous, not stel-

late, the walls radiating from them not greatly thickened;

divided into sections Euriccia and Ricciella.

2.1.1 Section Euriccia: air chambers are spaces between

vertical or subvertical columns of photosynthetic cells;

species assigned here are grouped together by absence or

presence of cilia and by size of scales.

R. dictyospora Howe, R. macallisteri Howe, R. bijurca

Hoffm.; R. glauca L., R. sorocarpa Bisch., R. campbel-

liana Howe, R. nigrella DC., R. albida Sulk, R. austinii

Steph. Margins of thallus without cilia; scales conspicuous

or inconspicuous.

R. californica Austin, R. hirta (Austin) Underw., R.

beyrichiana Hampe ex Lehm., R. tnchocarpa Howe, R.

donnellii Austin. Margins of thallus with cilia.

2.1.2 Section Ricciella: air chambers polyhedral hollows

bounded by unistratose walls; margins of thallus lack cilia;

ventral scales lacking or rudimentary.

R. membranacea Gottsche & Lindenb., R. frostii

Austin, R. crystallina L. (misidentified and is R. caver-

nosa), R. sullivantii Austin. Mature spores not grouped in

tetrads.

R. curtisii. Mature spores remaining attached in tetrads.

R. fluitans L. Thalli free-floating or stranded; segments

narrowly linear.

In all modern treatments, two of the species they in-

cluded in section Ricciella , R. curtisii and R. membrana-

cea, are assigned to different subgenera, namely Thallo-

carpus (as mentioned before) and Leptoriccia (Schuster

1984) respectively.

2.2 Genus Ricciocarpns Corda: R. natans (L.) Corda.

Thalli floating to stranded or rarely growing on very wet

Bothalia 25,2(1995)

213

soil; antheridia and archegonia located in median groove;

dorsal surface distinctly reticulate on account of air cham-

bers visible through it; monotypic.

2.3 Genus Oxymitra Bisch.: O. androgyna Howe. Thalli

growing on soil, air chambers large, subvertical, prismatic

or pyramidal; sporophyte with conspicuous conic or tri-

angular-pyramidal involucre; pores stellate due to greatly

thickened walls radiating from them.

Muller (1940) segregated the family Oxymitraceae

from the family Ricciaceae, to accommodate the genus

Oxymitra in the suborder Ricciineae. Although its sporo-

phytes are similar to those of the Ricciaceae, its thalli and

chromosome number are different, for which reasons it is

best treated in a separate family. Garside (1958), however,

described a new species of Oxymitra, O. cristata, from

South Africa, but referred it to the family Ricciaceae and

not to Muller’s (1940) family Oxymitraceae, although he

mentioned Muller’s publication. This was presumably

done because Garside was unsure whether the name

Oxymitra would be upheld.

3. SCHUSTER ( 1949): central and western New York

In an early study on the Hepaticae of this region,

Schuster treated Ricciocarpus and Riccia (with 6 local

species) in the Ricciaceae.

3.1 Genus Ricciocarpus Corda: R. natans. Aquatic; occur-

ring in circumneutral or somewhat calcareous waters,

shallow and still or occasionally stranded on shore: thallus

broad; ventral scales purple, dense, stiff, sword-like.

3.2 Genus Riccia : no subgenera or sections are mentioned,

but the key distinguishes between the following two

groups:

3.2a Group 1. R. fluitans (a mostly aquatic species), R.

crystallina , R. sullivantii (the latter bracketed, because it

only presumably occurs in the above area), R. frostii.

‘Photosynthetic tissue loose, separated by more or less

widely polyhedral air chambers, walls between chambers

only one cell thick; ventral scales absent or obsolete.'

3.2b Group 2. R. sorocarpa , R. arvensis Austin (a synonym

of R. bifurca Hoffm.). ‘Photosynthetic tissue compact, with

narrow vertical or subvertical, canal-like air chambers (at

times becoming polyhedral near margins); spores areolate to

alveolate, over 65 pm in longer diameter.’

4. SCHUSTER (1953): Minnesota and adjacent regions

Subsequently in reporting on the family Ricciaceae

(Suborder Ricciineae) from Minnesota and adjacent re-

gions, Schuster (1953) followed the conventional treat-

ment of placing in it the genera Ricciocarpus and Riccia.

In the latter he now recognized two subgenera. Ricciella

and Euriccia (with two sections).

4. 1 Genus Ricciocarpus: R. natans. Aquatic or subaquatic,

floating on water surface or stranded; in imperfect, partial

rosettes, with long, linear ventral scales; thallus with nar-

row, sharp, median groove, air pores and scattered oil cells.

4.2 Genus Riccia : thallose forms with sporophyte reduced

and surrounded by thallus tissue; air pores vestigial and

very inconspicuous, surrounded by hardly modified cells;

terrestrial forms in rosettes, but some species normally

aquatic; very variable.

4.2.1 Subgenus Ricciella : internal photosynthetic tissue

loose, forming large, polyhedral chambers (i.e. not com-

pact in structure in cross section); ventral scales rudimen-

tary; marginal cilia of thallus lacking; dorsal surface of

terrestrial forms often lacunose and spongiose with age.

R. fluitans (with related European taxa, R. rhenana Lorb.,

R. duplex Lorb. and R. canaliculata Hoffm. not yet found in

North America). Thallus segments narrowly linear, less than

0.7 mm wide; free-floating under water; sterile.

R. frostii. R. crystallina (misidentified and is R. caver-

nosa), R. sullivantii (the latter with sporangia bulging and

rupturing ventrally). Thallus segments relatively broad, al-

ways terrestrial; spores various.

4.2.2 Subgenus Euriccia: photosynthetic tissue compact;

spores always regularly areolate on external faces, with

or without wing margin.

R. austini (sic ),R. dictyospora, R. macallisteri, R. soro-

carpa, R. ar\>ensis. Thallus margins acute, sharp or rarely

obtuse, without obvious cilia; scales large, hyaline or pur-

ple, protruding beyond thallus margins or not.

R. hirta, R. beyrichicma, R. trichocarpa. Thallus mar-

gins obtuse, never sharp, with distinct cilia.

4.2.2. 1 Section Hirtae: R. hirta, R. beyrichiana, R. arven-

sis, and R. eldeeniae Jacobs (a synonym of R. warnstorfii

Limpr. ex Wamst.). Defined as ‘extreme xeric forms

which may show cilia and development of purplish pig-

mentation of lateral thallus margins under extreme inso-

lation'.

4. 2. 2. 2 Section Papillosae: R. atromarginata Levier. De-

fined as ‘extreme xeric forms abundantly hirsute'. No

mention is made of the author(s) of the two sections, nor

are there Latin descriptions or references that I could trace,

so I must infer that possibly, Schuster himself was the

author of these two sections which were, of course, then

not validly published.

5. MEIJER (1951): The Netherlands

5.1 Genus Riccia is defined by Meijer as including land

and water form species, frequently with ventral scales; the

compact rosette-shaped thalli are less strongly forked in

the land forms than in the water forms, which have thin,

narrow lobes that branch regularly and frequently; chlo-

rophyll-containing cells have smaller or larger air spaces

between; in isolated species reduced air pores are present;

two subgenera are recognized.

5.1.1 Subgenus Euriccia : chlorophyll-containing top portion

of thallus with narrow air canals bounded by 4 cell pillars:

R. glauca. TTiallus 4—5 times wider than thick in trans-

verse section.

R. subbifurca Wamst. ex Crozals (with large scales),

R. bifurca, R. beyrichiana (the latter two species with

small scales), R. sorocarpa (distinguished by the secon-

214

Bothalia 25,2 (1995)

dary epidermis having thickened walls). Thallus l-3‘/2

times wider than thick in tranverse section; thallus margins

with few or no cilia.

5.1.2 Subgenus Ricciella : top portion of thallus contains

large superimposed air chambers separated on all sides by

unicellular walls; (they are said to have the ability to adopt

an amphibian life style).

R. canaliculata (land form). Thallus ± twice wider than

thick, not or seldom branching.

R. crystallina, R. huebeneriana Lindenb. Thallus in a

rosette, spongy in older parts or with pitted appearance

dorsally.

R. rhenana, R. fluitans. Thallus not in a rosette, dor-

sally not pitted.

Meijer remarked on the problems encountered in

studying dried Riccia material and the desirability of ex-

amining living examples.

6. MULLER (1951-1958): Europe

Muller also recognized two genera, Ricciocarpus and

Riccia in the family Ricciaceae.

6. 1 Genus Ricciocarpus : R. natans. Thalli floating on water;

with long, band-shaped ventral scales hanging down; can

also manifest as a land form, when ventral scales are small;

assimilation tissue with large, tiered air chambers, air pores

present; oil bodies scattered in thallus and ventral scales;

gametangia sunken along centre of thallus.

6.2 Genus Riccia'. land forms or submerged in water; ventral

scales small, rarely projecting above thallus margins; assimi-

lation tissue in parallel cell rows or chambered; air pores

sometimes present in subgenus Ricciella ; oil cells lacking

and gametangia scattered over whole surface of thallus.

6.2.1 Subgenus Ricciella'. assimilation tissue consists of

chambers; simple air pores are now and then present in

roof over chambers.

R. frostii, R. crystallina , R. crystallina var. angustior

Nees (= R. cavernosa ), R. huebeneriana. Thalli in small

or large rosettes; spores various.

R. fluitans, R. canaliculata, R. rhenana, R. duplex.

Thalli linear, rarely forked; dorsally often clearly marked

into areas; some mostly aquatic.

Muller attempted (rather like Stephani (1898) and oth-

ers before him) to arrange the large number of species in

the subgenus Euriccia into informal groups, the members

of which are related by one or more shared characters,

e.g. the presence of cilia along the thallus margins or by

papillae on the dorsal surface or by thickened walls in the

(sub)dorsal cells.

6.2.2 Subgenus Euriccia'. assimilation tissue consists of

cell rows, ending in a larger cell on top.

6.2.2a Group Ciliifera

R. ciliifera Link, R. gougetiana Mont., R. gougetiana

var. erinacea Schiffn., R. zachariae Lorb., R. sommieri

Levier, R. melitensis C. Massal., R. macrocarpa Levier.

Some thalli with single cilia, but not a very constant char-

acter.

6.2.2b Group Ciliata

R. ciliata Hoffm., var. epilosa Wamst. var. intumescens

Bisch., var. violacea Kny, R. canescens Steph., R. crozalsii

Levier, R. michelii Raddi, R. bicarinata Lindb. Thallus

margins mostly with numerous long cilia.

6.2.2c Group Bifurca

R. bifurca , R. beyrichiana. Terminal branches with

margins raised and swollen; median groove wide and shal-

low.

6.2. 2d Group Sorocarpa

R. ligula Steph., R. wamstorfii, R. sorocarpa (subdorsal

cells with thickened walls), var. heegii Schiffn., R. breidleri

Jur., R. oelandica C.E.O. Jensen. Small plants, mostly linear

or lingulate, rarely with a few cilia.

6.2.2e Group Papillosa

R. papillosa Moris. Dorsal surface and along margins

with large papillae.

6.2. 2f Group Nigrella

R. subbifurca , R. nigrella, R. atromarginata var. glabra

Levier, R. trabutiana Steph. Thalli mostly in small ro-

settes; sometimes with deep median groove; occasionally

with blunt papillae dorsally.

6.2. 2g Group Lamellosa

R. lamellosa Raddi. Ventral scales large, hyaline and

extending above thallus margins.

6.2. 2h Group Glauca

R. glauca var. subinermis (Lindb.) Wamst. Thalli thin;

at margins occasionally with short hairs; in rosettes; scales

colourless.

7. ARNELL (1956): Scandinavia

In his treatment of the 'Hepaticae of the Moss Flora

of Fennoscandia’, Arnell placed the family Ricciaceae in

the suborder Ricciineae H. Buch and also recognized two

genera, Ricciocarpus and Riccia.

7. 1 Genus Ricciocarpus'. R. natans. Floating form with long

ventral scales hanging down, land fonn with smaller scales;

cordate; leathery; greyish green to dark green; dorsally with

pores surrounded by 5 or 6 thin-walled guard cells; assimi-

lative tissue containing several large air chambers, one lying

above the other; ventral tissue thin; oil cells present;

antheridia in small groups sunken in the middle.

7.2 Genus Riccia'. dorsal surface of thallus not leathery; yel-

lowish to dark green; assimilative tissue consisting of vertical

cell rows with interposed vertical, air-filled canals, or con-

sisting of the walls of the air chambers, the upper of which

each has a little pore in the middle of its roof; antheridia

scattered or in groups, not crowded in special organs.

Bothalia 25,2 (1995)

215

7.2.1 Subgenus ' Euriccia : assimilative tissue consists of

vertical cell rows surrounding narrow vertical air canals.

R. cilifera (sic), R. glauca, R. dalslandica S.W. Amell,

R. beyrichiana, R. bifurca, R. ciliata, R. wamstorfii, R.

subbifurca, R. sorocarpci. Thalli in rosettes or not; small

or large; branches thick or thin; narrow or wide; margins

winged or not, round or acute, naked or with cilia; epi-

dermal cells rarely thick-walled.

7.2.2 Subgenus Ricciella : assimilative tissue consists of

irregular chambers, separated by walls one cell thick.

R. canaliculata, R. crystallina var. angustior (= R. cav-

ernosa), R. rhenana, R. fluitans, R. duplex, R. frostii, R.

huebeneriana. Thalli linear, little-branched or repeatedly

furcate, thick or thin, reticulate, occasionally aquatic,

mostly terrestrial, sometimes in rosettes, large or small,

dorsally pitted.

8. JONES (1957): tropical Africa

The usual two genera were recognized by Jones in The

‘Ricciaceae in Tropical Africa’.

8.1 Genus Ricciocarpus: R. natans. Very widely distributed;

floating in still water; with long pendant ventral scales.

8.2 Genus Riccia : subdivided into the subgenera Ricciella

and Euriccia.

8.2.1 Subgenus Ricciella (A. Braun) Boulay sensu Into : as-

similation tissue with air chambers (aquatic or terrestrial).

R. fluitans. Floating on water or growing on wet soil;

branches elongated, parallel-sided, 0. 5-1.0 mm wide; air

chambers occupying most of thallus; spores rare.

R. membranacea. Branches many times broader than

thick, tapering into thin wings; dorsally areolate, older

parts becoming cavernous; air chambers large, occupying

most of thallus, except for 2 or 3 layers of basal tissue in

median part.

R. moenkemeyeri Steph. Branches tapering, apically

sharply channelled; becoming pitted dorsally; scales dis-

tant, not reaching edge of thallus.

R. rautcmenii (synonym of R. cavernosa). Large, spongy

rosettes; margins of thallus thick.

R. intermedia E.W. Jones. Branches parallel-sided, api-

cally sharply grooved; air chambers medianly narrow and

inconspicuous, like in Euriccia, but wider toward the mar-

gins. Riccia intermedia was regarded by Jones as inter-

mediate between Ricciella and Euriccia with respect to

the width of the air chambers or air canals; it was later

placed in synonymy under R. discolor Lehm. & Lindenb.

by Pande & Udar (1958) and classified in Euriccia.

Jones refers to Duthie & Garside’s publications (1937;

1939) and stresses that Ricciella sensu lato contains sev-

eral well-marked groups of species which are worthy of

sectional, if not of subgeneric rank.

8.2.2 Subgenus Euriccia'. assimilation tissue with closely

packed filaments.

R. trichocarpa. Margins of thallus with long, hyaline

cilia.

R. nigrosquamata E.W. Jones (= R. berriei E.W. Jones),

R. nigerica E.W. Jones, R. lanceolata Steph., R. radicosa

Pearson, R. rhodesiae S.W. Amell, R. runssorensis Steph.,

R. congoana Steph., R. angolensis Steph., R. papillispora

Steph. (R. albomarginata Bisch. and R. limbata Bisch. are

confined to the southwestern Cape). Margins of thallus

without cilia, ventral scales extend to, or greatly exceed

margins of thallus, distant or approximate or imbricate;

hyaline or with hyaline margins or dark violet; thallus

winged or not, when flanks steeply ascending; spores sub-

globose or tetrahedral.

The type specimen of R. rhodesiae, S.W. Amell 1291

(S), is a mixed collection of R. rhodesiae and R. atropur-

purea Sim, and Jones mistakenly identified the R. citropur-

purea portion of the specimen as R. rhodesiae ; R. rhodesiae

and R. nigrosquamata were placed in synonymy under R.

congoana by Perold (1986b).

9. PANDE & UDAR (1958): India

9.1 Genus Riccia'. divided into subgenera Euriccia and

Ricciella.

9.1.1 Subgenus Euriccia'. thalli with compact assimilatory

zone and narrow air spaces.

R. melanospora Kashyap. Thallus bluish green; seg-

ments small; anteriorly sulcate, rest nearly flat; cilia both

on margins and on surface, small and broad.

R. wamstorfii. Thallus yellow-green; segments narrow,

linear, apex rounded; deeply sulcate anteriorly and sulcus

broader behind; cilia large.

R. crozalsii. Thallus bluish green; segments lanceolate;

anteriorly deeply sulcate, rest very broadly channelled.

R. tuberculcita Pande & Udar. Thallus small, about 3

times broader than high; tubercular thickenings on cells

of assimilatory filaments; spores irregularly reticulate or

papillate, winged.

R. discolor. Male plants comparatively smaller than fe-

male; spore reticulate, unwinged.

R. billardieri Mont. & Nees. Thallus large, 4-6 times

broader than high; spores reticulate, with prominent pro-

jections.

R. gangetica S. Ahmad. Thallus small, about three

times broader than high; spores black, with 8-16 small

reticulations across distal face.

9.1.2 Subgenus Ricciella'. thalli with loosely arranged as-

similatory zone and wider air spaces.

R. huebeneriana. Thalli narrow, repeatedly branched;

spores small.

R. plana (synonym of R. crystallina). Thalli narrow,

repeatedly branched; spores much larger.

R. frostii. Thalli forming well-defined rosettes, female

rosette normally larger than male; spores incompletely re-

ticulate.

Riccia cruciata is mentioned in the key, but there is

no description of it.

10. HASSEL DE MENENDEZ (1963): Argentina

Hassel de Menendez treated the Argentinian Ric-

ciaceae.

216

Bothalia 25,2(1995)

10.1 Genus Ricciocarpus: R. natans. Plants floating or

terrestrial; photosynthetic layer formed by several super-

imposed air chambers; ventral layer almost absent; scales

in 3-5 series on both sides of midline.

10.2 Genus Riccia : plants terrestrial; photosynthetic layer

formed by one layer of air spaces surrounded by walls of

adjoining cells or by columns of chlorophyllose cells; ven-

tral layer developed; scales in 1 or 2 series.

10.2.1 Subgenus Ricciella : photosynthetic tissue consist-

ing of air chambers separated by walls one cell thick;

pores surrounded by complete rings or not, cells similar

to the others in dorsal walls of air chambers.

R. paranaensis Hassel, R. stenophylla Spruce. Thalli

linear, capsules ventrally prominent as described by A.

Braun.

R. membranacea, R. crystallina (misidentified and is

R. cavernosa), R. plana, R. frostii, R. paraguayensis

Spruce, R. curtisii. Thalli frequently in rosettes; capsules

slightly or ventrally not prominent; spores separating at

maturity except in R. curtisii, where they are united in

tetrads.

10.2.2 Subgenus Riccia: photosynthetic tissue formed by

chlorophyllose columns separated by narrow canals.

R. lindmanii Steph. Thallus margins without cilia.

R. bialbistrata Hassel. Thallus margins with cilia.

R. iodocheila Howe. Apex of thallus with some purple

papillae.

R. austinii, R. andina Miill.Frib., R. plano-biconvexa

Steph., R. campbelliana, R. squamata Nees, R. fruchartii

Steph. Walls of dorsal cells not thickened.

R. sorocarpa, R. dorsiverrucosa Hassel. Walls of dorsal

cells thickened.

No formal or informal groupings were made in either

subgenus. Riccia crystallina, as treated by Hassel, is ac-

tually R. cavernosa, as the two species were frequently

confused until Jovet-Ast (1964, 1966) cleared up the mat-

ter. Hassel selected R. glauca as the lectotype of the genus

Riccia, because Howe’s lectotype, R. crystallina, belongs

in subgenus Ricciella and it is therefore contrary to pre-

vious practice.

The specimen, leg. Sleumer 1755, identified as R.

campbelliana by Hassel, has large dark red scales, in con-

trast to those from other specimens of this species, in

which they are inconspicuous, fragile and hyaline,

splashed with brown and purple patches (Perold & Volk

1988a). Schuster (1992a) has assigned it to a new sub-

species, R. campbelliana subsp. austrigena. Riccia dorsiver-

rucosa (as well as R. andina Miill.Frib.) were recently placed

in synonymy under R. mauryana Steph. by Jovet-Ast ( 1991 ).

1 I . ARNELL( 1963): southern Africa

In his book Hepaticae of South Africa, Arnell (1963)

recognized Muller's Oxymitraceae and the family Ric-

ciaceae in the suborder Ricciineae. For the rest he fol-

lowed the same classification here as in his earlier work

(Arnell 1956) on Scandinavian Ricciaceae. He made no

attempt to arrange the species in the genus Riccia into

sections or groups.

1 1 . 1 Genus Ricciocarpus: R. natans. Floating on water

(with long, serrate scales) or growing on mud; dorsal sur-

face leathery, reticulate, greyish green to dark green; air

chambers large; pores present, surrounded by 5 or 6 thin-

walled guard cells; ventral tissue thin; ventral face with

several rows of puiplish ventral scales, numerous and long

in aquatic form; oil bodies in scales and thallus; antheridia

in small groups, sunken in the midline of thallus; sporo-

gonia rare, immersed in thallus.

1 1 .2 Genus Riccia: most species only have land forms,

but some species in subgenus Ricciella have both terres-

trial and aquatic forms; dorsal surface of thallus not leath-

ery; pores lacking except in some species of subgenus

Ricciella: ventral scales generally small; green layer of

assimilative tissue consists either of vertical cell pillars

with interposed air-filled spaces or (consists) of walls of

air chambers; oil bodies lacking; antheridia scattered or

in groups, not crowded in special organs.

11.2.1 Subgenus Euriccia: assimilative tissue composed

of cell pillars surrounded by air spaces.

R. canescens (synonym of R. trichocarpa), R. crozalsii.

Margin of thallus with long cilia.

R. villosa Steph., R. albomarginata Bisch., R. concava

Bisch. Margin of thallus usually without cilia; dorsally

velvet-like by the presence of free cell pillars.

R. sorocarpa. Epidermal cells in 2 layers, second layer

with thick walls.

R. coronata Sim, R. natalensis Sim. Epidermal cells

with thin walls, some of them elongated and free.

R. atropurpurea , R. limbata, R. angolensis, R. albolim-

bata S.W. Arnell, R. albosquamata S.W. Arnell, R. cap-

ensis Steph., R. rhodesiae, R. pottsiana Sim, R. warnstorfii

(an introduced species), R. runssorensis, R. okahandjana

S.W. Arnell. Epidermal cells in one layer, not forming free

pillars, but an adherent dorsal layer; scales absent or present,

large or small, hyaline or purple to black.

1 1 .2.2 Subgenus Ricciella: assimilative tissue composed

of large air chambers with walls one cell thick.

R. fluitans, R. stricta A.V. Duthie ined. Thalli with

long, linear segments, usually aquatic plants or rarely to

frequently terrestrial.

R. purpurascens Lehm. Usually terrestrial, large plants;

tuber formation on ventral stolons.

R. perssonii S.A. Kahn, R. curtisii, R. spongosa S.W.

Arnell, R. compacta. Tetrads of spores permanently ad-

herent, male plants much smaller than female ones.

R. cupulifera, R. rautanenii, R. plana, R. crystallina.

Monoicous, except for R. cupulifera which is dioicous;

spores separating when mature; plants in rosettes; male

and female plants the same size or male plants slightly

smaller.

R. bullosa Steph. (Link ex Lindenb. is actually the

author), R. garsidei Sim, R. montaguensis S.W. Arnell, R.

volkii S.W. Arnell, R. rubricollis Garside & A.V. Duthie

ined. Plants not forming complete rosettes, simple or di-

chotomously branched, very thick.

Bothalia 25,2(1995)

217

Arnell (1957) had previously described several new

Riccia species based on collections made by Volk in Na-

mibia, but had failed to recognize that the species with

velvet-like dorsal surfaces (Arnell 1963) form a distinct

group (later named section Pilifer O.H. Volk); R. villosa,

R. albomarginata and R. concava belong together in this

group and are quite distinct from ciliate species; no R.

coronata Sim specimens could be traced; R. capensis, as

interpreted by Arnell, was R. nigrella, whereas R. capensis

Steph. is a synonym of R. limbata Bisch. (Perold & Volk

1988b); R. albosquamata was placed in synonymy under

R. albolimbata (Perold 1989); R. rhodesiae is a synonym

of R. congoana (Perold 1986b); R. pseudolimbata S.W.

Arnell is a synonym of R. angolensis ; R. chrystallina (sic)

is R. cavernosa and R. plana is a synonym of R. crystallina ;

R. montaguensis is a synonym of R. bullosa (Perold 1991c);

R. spongosa is a synonym of R. curtisii (Perold 1991a).

12. VANDEN BERGHEN (1972): Zaire, Zambia

Vanden Berghen reported on the Ricciaceae of Zaire’s

province Shaba and of Zambia. Ricciocarpus natans

specimens were listed, as well as several Riccia species,

but the latter were not assigned to any particular subgenus.

12.1 Genus Ricciocarpus: R. natans. Thallus with wide

air cavities and small pores surrounded by 6-8 cells on

the dorsal face; floating in water, with long violet scales,

or when at water’s edge, without long scales; oil bodies

isolated in specialized cells; antheridia and archegonia sur-

rounded by a rudimentary involucre in the median groove.

12.2 Genus Riccia: plants annual or perennial; terrestrial

or aquatic; thallus dichotomously branched, frequently

forming small rosettes; thalli with air cavities generally

narrow; ventral scales mostly present, sometimes evanes-

cent; without oil cells; antheridia and archegonia not sur-

rounded by an involucre.

R. membrancicea. Thalli very thin, 6— 1 0(— 1 5 ) times

wider than thick; lobes 2. 8^4.0 mm wide.

R. fluitcins. Plants freely Boating in water or stranded

on mud; capsules strongly protruding on ventral face of

thallus; lobes up to 1 mm wide; ventral scales absent or

very reduced and evanescent.

R. cavernosa , R. moenkemeyeri, R. intermedia , R. sy-

moensii Vanden Berghen. Terrestrial plants; capsules not

strongly protruding on ventral face of thallus; lobes gen-

erally 1-2 mm wide; ventral scales absent or very reduced,

or large and violet.

As mentioned before, R. intermedia has been synony-

mised under R. discolor. It also appears, from Vanden

Berghen's description and illustration of the spores of his

so-called R. fluitans , that he was actually dealing with

R. stricta.

13. JOVET-AST ( 1975): diverse areas

Twenty years ago Jovet-Ast studied the species R. pers-

sonii and R. curtisii , which retain their spores in perma-

nent tetrads. Previously, on account of the tetrad spores,

Austin (1869) had placed R. curtisii in his new genus.

Cryptocarpus , but Lindberg ( 1874) called attention to the

fact that the generic name, Cryptocarpus, was already in

use and proposed the name Thallocarpus, instead. Sub-

sequently. however, these plants were treated under sub-

genus Ricciella, until Thallocarpus was reinstated as an

independent subgenus by Jovet-Ast (1975).

14. CAMPBELL (1975, 1977): New Zealand

1 4. 1 Genus Ricciocarpos: R. natans. Plants in spongy, par-

tial rosettes, up to 20 mm in diameter; when floating in

ponds, little-branched and green or yellow-green in col-

our; dorsally firm and convex, with deep median furrow;

upper epidermis interrupted by simple air pores; com-

posed below of several series of large air chambers

bounded by unistratose walls; ventrally with few rhizoids

but with numerous conspicuous and characteristic pendant

scales, toothed, ribbon-shaped and reddish violet or

brownish green in colour; when temporarily stranded on

shore, freely branched; with numerous smooth and tuber-

culate rhizoids; scales only rudimentary; rarely fertile in

either aquatic or land form.

14.2 Genus Riccia: divided into two subgenera.

14.2.1 Subgenus Riccia: with a rather compact photosyn-

thetic layer consisting of more or less vertical filaments

separating narrow air chambers.

R. crozalsii. Margin of thallus with cilia, mainly near

apex, 200-500 pm long.

R. ciliata. Cilia abundant along margin, 600-900 pm

long.

R. sorocarpa. Dorsal groove deep and narrow.

R. bifurca, R. glauca. Dorsal groove shallow and wide.

14.2.2 Subgenus Ricciella: photosynthetic layer spongy

and consisting of unistratose walls enclosing polyhedral

air chambers.

R. fluitans. Thallus free-floating or stranded, not at-

tached: segments narrowly linear.

R. crystallina. In glassy green or grey-green spongy

rosettes; air chambers large and in several layers.

R. vesiculosa (Carrington & Pearson) Steph., formerly

thought to be R. bullosa (pers. comm.), green or olive-

green, strap-shaped and large; median air chambers nar-

row, those of the wings larger and in layers.

15. GROLLE ( 1976, 1983): Europe and adjacent regions

Grolle in his ‘Leberntoose Europas' divided Riccia, as

usual, into the subgenera Riccia and Ricciella (A. Br.)

Boulay. In 1983 he stated the authors of the latter subge-

nus to be (A. Br.) Rchb., with R. fluitans as the newly

selected lectotype. Two sections were recognized, namely

Ricciella and Spongodes (the latter with the newly se-

lected lectotype, R. crystallina). No attempt was made to

arrange the species into groups, nor are diagnostic char-

acters given.

16. NA-THALANG (1980): Australia

In her study of Australian Ricciae, Na-Thalang thought

it incorrect for the two subgenera, Riccia and Ricciella,

218

Bothalia 25,2 (1995)

to be separated on the size of the air chambers, because

the characteristics of air chambers, pores and epidermal

cells ought to be considered more important than the size

of the air chambers.

16.1 Genus Riccia: monoicous or dioicous; complete or

incomplete rosettes or in gregarious patches; mostly land

forms; annual or perennial; upper surface normally sul-

cate; margins sometimes with cilia or scales of various

sizes; dorsal pail with compact tissue in vertical columns

or with loosely arranged tissue forming polyhedral air

chambers; capsules with neither seta nor foot, normally

embedded in thallus.

16.1.1 Subgenus Riccia : upper part of thallus composed

of compact tissue arranged in almost vertical columns; air

chambers narrow, without specialized air pores; epidermal

cells hyaline.

16.1.1.1 Section Vmdisquamata Na-Thalang: R. caroliniana

Na-Thalang. Dorsal part of thallus consists of compact, hya-

line tissue; chlorophyll-bearing layer at ventral side of thallus,

but almost whole thallus pellucid, destitute of chloroplasts

except a few towards apex, margins and ventral layer next

to scales; scales narrowly oblong, green, cells containing

chloroplasts, imbricate and persistent, 1 or 2 layers of cells

in thickness; branches furcate or bifurcate.

16.1.1.2 Section Riccia : chlorophyll-bearing layer at dor-

sal part of thallus; consists of compact vertical columns

of green cells terminated by hyaline epidermal cells, each

air space normally enclosed by 4 vertical columns; in

some species, air chambers close to the margin may be

enlarged, when surrounded by 5 or 6 columns.

Three groups are recognised.

16.1.1.2a Group Ciliatae: thalli bear cilia at margins and

sometimes on dorsal surface.

1 6. 1 . 1 ,2aa Subgroup Longiciliata: plants with long cilia

on thallus and on capsule.

R. longiciliata Na-Thalang, R. crinita Taylor, R. areo-

lata Na-Thalang. Closely related species, but with differ-

ent spore characters.

Seppelt (1983) subsequently reduced Na-Thalang’s R.

longiciliata and R. areolata to synonyms of R. crinita.

Riccia longiciliata , with n = 16, is regarded by him as

only a polyploid of R. crinita, with n = 8, and it was

concluded that the three species cannot be satisfactorily

delimited by spore characteristics, neither can they be dif-

ferentiated by morphological gametophyte characteristics.

16. 1.1. 2ab Subgroup Crozalsii: plants with short cilia on

thallus, none borne on capsule.

R. crozalsii, R. hlackii Na-Thalang, R. asprella Car-

rington & Pearson. Thalli small to larger; margins tumid

and raised; channel broad.

16. 1 . 1 .2b Group Squamatae: thalli with scales reaching to

or extending beyond margins.

16.1.1 .2ba Subgroup Macrospora: thalli with variable pur-

ple scales; spores globose.

R. macrospora Steph., R. gangetica, R. billardieri, R.

discolor. Colour of scales and thalli sensitive to environ-

mental change.

16. 1.1. 2bb Subgroup Limbata: scales dark purple; spores

triangular-globular, proximal face with triradiate mark.

R. limbata, R. nigrella. Colour of scales not easily

changed.

1 6. 1 . 1 ,2bc Subgroup Sorocarpa: scales hyaline; mostly an-

nuals.

R. sorocarpa, R. lamellosa. Introduced species.

R. marginata Carrington & Pearson, R. olgensis Na-

Thalang, R. rorida Na-Thalang. Australian species, prob-

ably endemic.

1 6. 1 . 1 ,2c Group Laevigatae: scales small, only present un-

der apex of thallus.

R. bifurca, R. albida. Introduced species; thallus mar-

gins rounded, the latter species being the only Riccia with

smooth and rounded spores.

16. 1 .2 Subgenus Ricciella: upper part of thallus composed

of rather loosely arranged chlorophyll-bearing tissue form-

ing reticulate one-layered cell plates; air chambers poly-

hedral, normally roofed over by green epidermal cells

with an air pore in the centre.

Two groups are recognized.

16.1.2a Group Terrestriae: land forms, sometimes growing

on mud.

16.1.2aa Subgroup Crystallina: annual plants; in round

and compact rosettes.

R. crystallina, R. cavernosa. Thalli not strap-like,

broader than 1.5 mm; capsules not bulging markedly on

ventral surface.

16. 1.2ab Subgroup Papulosa: large thalli, 3-6 mm wide.

R. papulosa Steph., R. spongiosula Na-Thalang. Plants

not strictly annual; forming rosettes only in young stages.

16.1.2ac Subgroup Vesiculosa: thalli smaller, 2-3(4) mm

wide; hygrophilous plants.

R. vesiculosa (Carrington & Pearson) Steph., R. mus-

cicola Steph., R. deserticola Steph., R. crassa Steph., R.

junghuhniana Nees & Lindb., R. collata Na-Thalang.

Scales mostly small and hyaline.

1 6. 1 ,2b Group Aquaticae-Terrestriae: thalli narrow-linear or

strap-like; multifurcate; capsules bulging on ventral surface.

R. luticola Na-Thalang, R. duplex, R. multifida Steph.

Plants growing in water or on land; upper surface marked

out by air chamber areas or not.

Na-Thalang’s groups were not given formal taxonomic

status, because she considered a worldwide appraisal of

the genus a necessity before formalizing taxa below the

Bothalia 25,2(1995)

219

level of subgenus. Schuster ( 1 992a), however, correctly

points out that these informal groupings are ‘nomenclatu-

rally irrelevant’.

1 7. JOVET-AST (1984): Australia

Jovet-Ast raised Na-Thalang’s (1980) section Viridi-

squamata (containing R. caroliniana) to subgeneric rank,

because of the very unusual structure of its lobes, which

could represent either a primitive stage, i.e. a relictual en-

demic of Australia, or else a parallel lineage to the genus

Riccia. Its spores are, however, comparable to those of

many other species of Riccia. Jovet-Ast, therefore, thought

it preferable to maintain it in the genus Riccia.

17.1 Genus Riccia

17.1.1 Subgenus Viridisquamata (Na-Thalang) Ast: R.

caroliniana. Dorsal part of thallus hyaline and compact;

ventral part of thallus chlorophyll-bearing, i.e. layers in-

verted compared with other species of Riccia ; ventral side

of lobes with rows of chlorophyllose lamellae which are

not scales; distal face of spores reticulate, forming 9 or

10 rounded areolae across diameter; proximal face simi-

larly ornamented.

18. VOLK (1983): Namibia and southern Africa (and

elsewhere)

In 1981 Volk described a new Riccia species, R. al-

bovestita , from Namibia. It, and others like it, are char-

acterized by a dorsal epithelium of loose, erect cell pillars,

which Amell (1963) had referred to as ‘velvet-like’, but

had not segregated into a distinct group. This was sub-

sequently done by Volk (1983), when he created his new

section Pilifer. He also proposed a new classification for

the genus Riccia, in an attempt to avoid the problems

caused by intermediate forms with regard to the width of

the air canals/air chambers. This classification was no

longer to be based on the width of the air chambers, but

rather on the number and size of the air pores in the dorsal

epithelium/epidermis of the thallus.

18.1 Genus: Riccia

18.1.1 Subgenus A. Riccia ( Euriccia auct.): outer dorsal

layer consisting of an epithelium of hyaline cells; without

sharply delimited air pores.

18.1.1.1 Section Viridisquamata: R. caroliniana. Chloro-

phyllose tissue limited to ventral side of thallus and scales.

18.1.1.2 Section Riccia | Lichenoides (Bisch.) Nees]: R.

glauca , R. sorocarpa, R. limbata, R. angolensis etc. Chlo-

rophyll mostly in dorsal tissues; ventral scales not green;

top cells of epithelium close together.

18.1.1.3 Section Pilifer O.H. Volk: R. villosa Steph., R.

albovestita O.H. Volk, R. concava Bisch. etc. Upper cells

of epithelium free, hair-like pillars.

18.1.2 Subgenus B. Spongodes (Nees) O.H. Volk: outer

dorsal layer consisting of an epidermis with chlorophyll

and sharply delimited air pores.

18.1.2.1 Section Thallocarpus: R. perssonii, R. curtisii.

Spores in permanent tetrads.

18.1.2.2 Section Spongodes Nees: R. crystallina, R. cav-

ernosa, R. volkii. Ripe spores free; sporangia opening dor-

sally.

18.1.2.3 Section Ricciella (A. Braun, pro gen.) Bisch.: R.

fluitans, R. canaliculata , R. stricta etc. Sporangia bulging

and opening ventrally.

19. SCHUSTER (1984, 1985): diverse areas

Schuster proposed a new subgenus, Leptoriccia, for R.

membranacea, as well as a third genus in the Ricciaceae,

namely his monotypic Pteroriccia R.M. Schust. to accom-

modate R. villosa Steph.

19.1 Genus Riccia

19.1.1 Subgenus Leptoriccia R.M. Schust.: R. membranacea.

Thallus very thin with vestigial ventral tissue; spores with

numerous papillae, not arising from a reticulum.

19.2 Genus Pteroriccia R.M. Schust.: R. villosa Steph.

Thallus with tall dorsal cell pillars and large erect, imbri-

cate ventral scales, margins serrate.

On account of the above characters Schuster thought

that Pteroriccia was at least as distinct from Riccia as

the water form of the separate, unrelated genus, Riccio-

carpus, which has long, trailing, ventral scales also with

serrate margins.

20. PEROLD (1986): southern Africa

In response to Schuster's publication on Pteroriccia

Perold (1986a) attempted to draw attention to O.H. Volk’s

section Pilifer , which was instituted for the reception of

all the then (and later) known Riccia species with tall,

loose dorsal cell pillars. She also pointed out that scale

margins vary considerably in some species of Riccia, from

crenate to serrate to having long, multicellular appendages

in a then, as yet, undescribed species. The new genus

instituted by Schuster was accordingly regarded as syn-

onymous with Riccia.

21. SCHUSTER (1985): worldwide

Schuster (1985), however, had already reduced the

status of his genus Pteroriccia (here regarded as a syno-

nym of section Pilifer O.H. Volk) to that of subgenus

Pteroriccia (R.M. Schust.) R.M. Schust. after restudying

the Pteroriccia complex and finding that the ventral scale

criterion, which had strikingly isolated his type species,

did not hold for several other taxa in the same taxonomic

unit. He also thought that Pilifer O.H. Volk was illegiti-

mate, as he had assumed (incorrectly, however) that no

type species had been designated, whereas Volk (1983)

had indeed selected R. albomarginata as the type species

of his section Pilifer. Schuster (1985) recognized six sub-

genera in the Ricciaceae.

220

Bothalia 25,2 (1995)

21.1 Genus: Riccia

21.1.1 Subgenus Spongodes (Nees) O.H. Volk: dorsal

epidermis distinct, complete, becoming lacunose, chlo-

rophyllose, with pores; air chamers large, polyhedral;

ventral tissue several cell layers thick.

21.1.2 Subgenus Leptoriccia R.M. Schust.: dorsal epider-

mis distinct, complete, becoming lacunose, chlorophyl-

lose with pores; air chambers large, polyhedral; ventral

tissue vestigial.

21.1.3 Subgenus Thallocarpus [Austin (sic)] Ast: spores

coherent in permanent tetrads; plants unisexual, strikingly

heterothallic with dwarf males.

2 1 . 1 .4 Subgenus Riccia: epithelial tissues in mutually con-

nate cell rows between which vertical air canals are distinct.

21.1.5 Subgenus Pteroriccia (R.M. Schust.) R.M. Schust.:

epithelial tissues free-standing, independent multicellular

uniseriate hairs, formed of elongated cells; ventral scales

typically large, projecting beyond thallus margins.

21.1.6 Subgenus Viridisquamata (Na-Thalang) Ast: chlo-

roplasts limited to ventral strata of thallus; ventral lamellae

bistratose and chlorophyllose.

22. SCOTT (1985): southern Australia

22.1 Genus Riccia : thallus often distinctly coloured;

wedge-shaped, usually wider than thick; bifurcating fre-

quently and forming rosettes; ventral scales conspicuous

or not; antheridia and archegonia sunken in pits along

mid-line of thallus; sporophyte merely a ‘bag of spores’

also sunken in thallus; spores with elaborate sculpturing

on surface which seems to be diagnostic of each species;

divided into two subgenera which are ± distinct.

22.1.1 Subgenus Riccia ( sensu stricto ): upper layer of

thallus firm, consisting of compact vertical pillars of cells,

closely packed together and joined at the top, forming a

firm, close tissue with narrow, vertical air slots and an

even, homogeneous upper surface.

R. asprella, R. crozalsii, R. crinita. Thallus margins

with hairs, fine or coarse, few or abundant, confined to

apex or not.

R. limbata, R. nigrella. Medium-sized or small; bor-

dered by black ventral scales.

R. sorocarpa, R. cartilaginosa Steph. Smallish or large;

ventral scales hyaline and just reaching thallus margin.

R. bifurca. Upper surface broadly channelled.

R. lamellosa , R. rorida. Thai 1 i rimmed by white scales,

extending beyond margins.

R. albida. Upper surface narrowly grooved; chalky

white with calcareous incrustation.

22.1.2 Subgenus Ricciella : upper layer of thallus spongy;

loosely arranged chlorophyllous tissue, forming polyhe-

dral or large and irregular air chambers, usually with defi-

nite air pores; epidermal cells with chloroplasts except

around air pores.

R. crystal lina , R. cavernosa. Plants in circular rosettes;

upper epidermis granular or rough or cavernous even near

apex.

R. vesiculosa , R. spongiosula, R. papulosa. Plants dor-

sally sometimes becoming cavernous in older parts or

honeycombed, except at apex or only at base.

R. multifida, R. duplex. Thalli narrow, strap-like, less

than 1 .5 mm broad; thick or thin; with or without reticu-

lation; grooved along midline or only at apex.

23. VOLK & PEROLD (1986a): South Africa

In 1986 Volk & Perold described a new species, R.

schelpei. which is a very distinctive plant; it was assigned

by them to the new monotypic subgenus Chartacea

Perold, which is restricted to the winter rainfall region of

the Western Cape.

23.1 Genus: Riccia

23.1.1 Subgenus Chartacea: R. schelpei. Thallus when

dry, with papery texture, epidermis parchment-like with

thick-walled hyaline cells; air pores surrounded by a ring

of thin-walled, superimposed cells; spores reticulate-

faveolate on distal face; proximal face with small shallow

areolae, sprinkled with granules.

Later in the same year Volk & Perold (1986b) de-

scribed another new species, R. hirsuta , which was placed

in the new section Micantes O.H. Volk & Perold, subge-

nus Spongodes (Nees) O.H. Volk.

23.1.2 Subgenus: Spongodes

23.1.2.1 Section Micantes : R. hirsuta. Thallus dorsally

densely covered with crowded multicellular, hairlike, hyaline

shiny pillars; assimilation tissue with polyhedral air cham-

bers; ventral scales triangular and apically split into cellular

strands; spores single, ornamentation reticulate; spores with

3-5(6) large central areolae completely or incompletely sub-

divided, proximal face incompletely reticulate.

Unfortunately the description of R. hirsuta was based

on two distinct, but rather similar species; fresh collections

of R. hirsuta showed the assimilation tissue to have air

canals and not polyhedral air chambers and hence to be-

long to section Pilifer, subgenus Riccia ; section Micantes

was accordingly rejected by Volk & Perold (1990).

24. JOVET-AST (1986): Mediterranean countries

Jovet-Ast’s study of Mediterranean Ricciae was the

culmination of the examination of several thousand speci-

mens collected during 22 surveys around the Mediterra-

nean Sea. Although only subgenus Riccia (with section

Riccia ) and subgenus Ricciella (with sections Ricciella

and Spongodes) grow in the area treated, she listed five

subgenera in her classification: Riccia (with sections Ric-

cia and Pilifer ), Ricciella, Thallocarpus. Viridisquamata

and Leptoriccia. In her keys, of which there are several

Bothalia 25,2(1995)

221

to subgenus Ricciella , she referred to Grolle's lectotypifi-

cation of section Spongodes Nees and of section Ricciella.

No formal or informal groups or subgroups were recog-

nized in section Riccia, and varieties based on small dif-

ferences in size and colour were rejected.

24.1 Genus Riccia : thallus in the form of simple, linear

lobes or ramified 1—4 times or in complete or partial ro-

settes; compact or spongiose; dorsal surface flat or with

longitudinal median furrow; margins glabrous or bearing

papillae or cilia; lateral surfaces with scales, small or

large, colourless, or flecked with violet, orange, or dark

violet or entirely dark violet.

24.1.1 Subgenus Riccia : thalli not spongiose, without air

chambers; chlorophyllose tissue consisting of cells ar-

ranged in vertical rows, between them filiform aeration

canals that open dorsally by minute openings; terrestrial

species.

R. crustata Trab., R. lamellosa , R. melitensis, R. soro-

carpa, R. glauca. R. macrocarpa, R. sommieri , R. bifiurca,

R. subbifurca , R. wamstorfii, R. nigrella, R. trabutiana.

Thallus glabrous; upper surface white or not; scales vari-

ous.

R. atromarginata, R. papillosa, R. gougetiana. R. cili-

ifera , R. bicarinata , R. trichocarpa. R. cilicita , R. michelii,

R. beyrichiana, R. crozalsii. R. ligula. Thallus bearing cilia

or papillae.

24.1.2 Subgenus Ricciella'. thalli spongiose and with nu-

merous large lacunae opening to exterior; or thallus not

spongiose, air chambers opening dorsally via pores; sur-

rounding cells arranged stoma-like; air chambers sepa-

rated by mostly unicellular walls.

24.1.2.1 Section Spongodes : thallus spongiose; dorsal sur-

face bearing large lacunae or both lacunae and pores; cap-

sules not very prominent on upper surface of lobes,

enclosed in thallus; terrestrial species.

R. crystallina. R. cavernosa. Monoicous; thallus in ro-

settes.

R. frostii. Dioicous; pronouncedly heterothallic with

male thallus much smaller than female thallus.

24.1.2.2 Section Ricciellcr. thallus mostly not spongiose,

generally with dorsal epidermis bearing pores surrounded

by regularly arranged cells; capsules protuberant on ven-

tral surface of thallus; terrestrial or aquatic species.

R. perennis Steph.. R. huebeneriana , R. Jim tans, R. du-

plex, R. canaliculalu . Thallus with or without clear areo-

lation on dorsal surface; in R. canaliculata ventral apex

of lobes covered by white scale, in other species not.

25. DAMSHOLT & HALLINGBACK (1986): Fenno-

scandia

Also published in 1986 was Damsholt & Hallingback’s

report on the Ricciae of Fennoscandia, which contained

beautiful coloured photoprints to illustrate the different

colours of the thalli as well as outstanding drawings by

the Danish artist, Annette Pagh. Since they were dealing

with local species, only two subgenera were recognized,

neither were any sections separated in either of the sub-

genera.

25.1 Genus: Riccia

25.1.1 Subgenus Ricciella : thallus with air chambers in

assimilation tissue; secondarily adapted to damp or fresh

water environments since air pores are here used as a

floating mechanism.

R. cavernosa, R. huebeneriana. Land plants; in partial

or complete rosettes.

R. canaliculata, R. fluitans. The former species is ter-

restrial, whereas the latter is a land or a water plant; nei-

ther grows in rosettes.

25.1.2 Subgenus Riccia: thallus compact; assimilation tis-

sue with closely spaced vertical cell rows; adapted to dry

areas; upper surface of thallus closes up as the sides meet

and so dehydration is prevented.

R. bifurca, R. beyrichiana, R. sorocarpa, R. wamstorfii,

R. subbifurca, R. gothica Damsh. & Hallingb., R. glauca.

Thallus margins often glabrous.

R. ciliata, R. ciliifera. Thallus margins mostly ciliate.

R. dalslandica is placed in synonymy under R. ciliata

var. epilosa; R. glauca forma colorata S.W. Amell is

treated as a separate species, R. gothica, by the authors.

Jovet-Ast (1994) expresses some doubt that Damsholt &

Hallingback had indeed examined the true type specimen

of R. ciliata and argues that they may have confounded

R. ciliata and R. trichocarpa , since she had examined the

types of both and found them to be different.

26. VOLK & PEROLD (1990): South Africa

Volk & Perold in correcting their initial error (Volk &

Perold 1986b) now referred R. hirsuta to section Pilfer;

R. tomentosa, the other element on which the original de-

scription of R. hirsuta had been partly based, was referred

to subgenus Thallocarpus , section Pannosae (sect, nov.),

since its spores (absent in the original collection) are re-

tained in permanent tetrads. Riccia tomentosa and R. hir-

suta share triangular ventral scales with apical filamentous

strands and elongated dorsal cell pillars.

27. SMITH ( 1990): Britain and Ireland

Smith treated the liverworts of Britain and Ireland. He

commented that there seemed to be no general agreement

about the arrangement of species within the subgenus Riccia,

since Grolle (1983) had arranged them in alphabetical order

and the arrangements of Muller (1951-1958), Jovet-Ast

(1986) and others differed. He chose to follow Jovet-Ast as

hers was, at the time, the most recent authoritative account.

27.1 Genus Ricciocarpus: R. natans. Plants floating in

base-rich or neutral ponds, ditches and canals or on mud

of dried-out ponds; with conspicuous linear-lanceolate

ventral scales (terrestrial forms with small scales); dorsal

epidermis with small distinct pores and scattered oil cells.

222

Bothalia 25,2 ( 1995)

27.2 Genus Riccia : plants terrestrial or, if aquatic, sub-

merged; often winter ephemerals or annuals; ventral scales

small to larger; pores lacking or, if present, vestigial and

not surrounded by specialised cells; oil cells lacking.

27.2.1 Subgenus Ricciella: thallus with chambers; epider-

mis breaking down to form lacunae in older parts of thal-

lus or not; margins without cilia.

27.2.1.1 Section Spongodes: plants terrestrial; dorsal epi-

dermal cells in older parts of thallus breaking down to

form lacunae, rendering thallus spongy in appearance;

capsules embedded in thallus, not protruding strongly on

ventral side.

R. cavernosa, R. crystallina, R. huebeneriana. Plants

terrestrial; not strap-like; older parts spongy in appear-

ance; capsules not protruding strongly on ventral side of

thallus [except in R. huebeneriana (parenthesis mine)].

27.2.1.2 Section Ricciella : plants aquatic or terrestrial; epi-

dermal cells on dorsal side of thallus not breaking down,

older parts of thallus not spongy in appearance; capsules

protruding very strongly on ventral side.

R. canaliculata, R. fluitans, R. rhenana. Thallus not

channelled, except sometimes in older parts; not reticulate

or often reticulate; capsules common or rare.

27.2.2 Subgenus Riccia : thalli never spongy in appear-

ance; margins ciliate or not; dorsal epidermal cells fre-

quently soon lost; chlorophyllose tissue composed of

vertical columns with narrow air spaces, forming a com-

pact layer of green cells.

R. crozalsii. Thallus lobes with ciliate margins, cilia

curved over thallus when dry.

R. subbifurca, R. glaucci, R. bifurca, R. beyrichiana.

Thallus lobes with marginal cilia if growing under moist

conditions; not indexed over thallus when dry.

R. nigrella, R. sorocarpa. Thallus margins naked;

subepidermal cells of thallus lobes thick-walled in R. soro-

carpa.

28. VAN MELICK (1991): The Netherlands

Van Melick undertook a revision of 1 259 Dutch her-

barium collections of Riccia. Earlier bryologists in the

Netherlands evidently did not have a clear concept of

many Riccia species as many specimens had been misi-

dentified. Eleven species were retained by him and his

classification also follows that of Jovet-Ast ( 1986), but he

comments that there was no ‘eenduidige opvatting over

de taxonomische indeling van het genus Riccia ’.

28.1 Genus Riccia: plants once to several times dichoto-

mously furcate or in rosettes; dorsal surface of lobes

smooth or with stomata-like pores or else with raised epi-

dermal cells, otherwise with crater-like openings; ventral

surface of lobes with rhizoids and scales, these laterally

displaced with growth of thallus; on t.s. dorsally with wide

air chambers enclosed by green walls or with compact,

pal lisade-l ike assimilation tissue; ventral tissue paren-

chyma-like, lacking oil cells; monoicous or dioicous; cap-

sules sunken in thallus.

28.1.1 Subgenus Riccia: dorsal side of thallus not spongy,

lacking network of small areas, but epidermis with hya-

line, round or pear-shaped, generally ephemeral cells;

margins without or with cilia; assimilation tissue dorsal,

with compact, vertical cell rows, separated by narrow air

canals; ventral V?-2/ 3 of thallus compact storage tissue.

R. crozalsiii. Thallus lobes at margins with more than

20 cilia that arch over thallus when dry.

R. glauca, R. beyrichiana, R. bifurca, R. warnstorfii.

Thallus lobes without or with fewer than 20 cilia at mar-

gins and not arched over thallus when dry.

R. subbifurca. Thalli scattered, rarely in rosettes.

R. sorocarpa. Thallus lobes with deep, sharp groove;

upper epidermis 2-layered, subepidermis with thick-

walled cells.

28.1.2 Subgenus Ricciella: dorsal side of thallus with ob-

scure or clear network of small areas; epidermis uni-

stratose, sometimes with stoma-like pores, sometimes

lacunose; margins of thallus without cilia; assimilation

layer composed of loose tissue with two or more irregular

layers of large or small air chambers; storage tissue ventral

± V3 of thallus.

28. 1 .2. 1 Section Spongodes: plants growing on soil; dorsal

side of thallus, especially in the older parts, spongy,

caused by openings in epidermis; capsules ventrally bulg-

ing or not.

R. cavernosa. Thalli light green to yellow-green, turn-

ing brown with age; lobes 1. 0-2.5 mm wide; groove ab-

sent; capsules sunken into thallus or dorsally slightly

bulging.

R. huebeneriana. Thalli bright green, mostly with rose

to wine-red flecks; lobes 0.3-1 .0 mm wide; grooved; cap-

sules bulging ventrally.

28.1.2.2 Section Ricciella: plants growing on soil or float-

ing in water; dorsal side of thallus not spongy; epidermis

without openings; capsules bulging ventrally.

R. canaliculata. Thallus lobes narrower at tips; ventral

scales 1. 5-2.0 times wider than high, apical scale reflexed

cap-like over dorsal side.

R. fluitans. Thallus lobes widening at tips; ventral

scales 2-5 times wider than high, apical scale not reflexed

cap-like over dorsal side; land form or water form.

Riccia duplex and R. rhenana were not accepted as

good species, because of the lack of constant diagnostic

characters to differentiate them from R. canaliculata and

R. fluitans respectively.

29. VOLK & PEROLD; PEROLD & VOLK; PEROLD

( 1 984- 1991): southern Africa

The Ricciaceae of southern Africa continued to be

studied by Volk (1979-1984), then by Volk & Perold or

by Perold & Volk (1984-1990) and lately by Perold

(1991). Several new species were described between

them.

Bothalia 25,2 (1995)

223

In her Ph.D. thesis, Perold (1991a) recognized the fol-

lowing taxa in the Ricciae: subgenus Riccia, with two sec-

tions, Riccia (which was divided into the informal groups,

‘Ciliatae’ and ‘Squamatae’) and Pilifer, for which Schuster’s

subgenus Pteroriccia had not been adopted, as the compo-

sition of the air canals is no different from those in Riccia ;

subgenus Ricciella which was also divided into two sections,

Spongodes (with two groups, ‘Crystallina' and ’Vesiculosa’),

and Ricciella ; to subgenus Chartacea , R. schelpei was as-

signed and to subgenus Thallocarpus, R. curtisii, R. corn-

pacta and R. perssonii ; section Pannosae was elevated to

subgenus Pannosae (Perold) Perold. (See Perold’s proposed

classification of sub-Saharan Ricciae on p. 227.)

30. JOVET-AST (1991, 1993): Latin America

Jovet-Ast, in the study of the Ricciae of Latin America,

listed the five subgenera previously recognized by her

(Jovet-Ast 1986): Riccia, Viridisquamata, Ricciella, Thal-

locarpus, Leptoriccia and added a sixth one, Chartacea.

Only four of these are from the area treated in these works,

namely Riccia , Thallocarpus, Ricciella and Leptoriccia.

In the subgenus Riccia, section Riccia, she used Stephani's

(1898) two units, the Ciliatae and the Inermes, but intro-

duced a refinement by distinguishing a third one, the

Papillatae, for species with papillose lobes. Her groups

were not given a specified rank and are informal.

30.1 Genus Riccia : thalli compact or spongiose; dorsal

surface flat or with longitudinal median furrow; margins

glabrous, or bearing papillae or cilia: lateral surfaces with

scales.

30.1.1 Subgenus Riccia : chlorophyllose tissue compact,

consisting of cells arranged in vertical rows with narrow

air spaces between; epidermal pores simple.

30.1.1.1 Papillatae: margins of thallus lobes with papillae.

R. atromarginata var. atrornarginata. Margins of lobes

obtuse, papillae on the margins and lateral surfaces,

60-210 pm long, smooth.

R. violacea Howe var. violacea, R. violacea var. laevis

Ast. Papillae on margins and lateral surfaces of lobes

25-130 pm long, granular or smooth.

R. iodocheila. Margins of lobes acute; papillae rare.

30.1.1.2 Ciliatae: margins of lobes with cilia.

R. horrida Ast, R. trichocarpa, R. subdepilata Ast. R.

enyae Ast, R. cubensis S.W. Amell. Lobes mostly abundantly

and densely ciliate to base.

R. brcisiliensis Schiffn., R. lindmanii. Lobes with few.

sparse cilia.

30.1.1.3 Inermes: thallus margins glabrous, lacking papil-

lae or cilia.

30.1.1.3a Group 1: R. albida. Lobes calcified; spores almost

smooth.

30.1.1.3b Group 2: R. albopunctata Ast, R. campbelliana.

Lobes not calcified; with idioblasts.

30.1.1.3c Group 3: R. nigrella , R. boliviensis Ast, R.

squamata. Lobes not calcified; lacking idioblasts; scales

brilliantly black.

30. 1 . 1 .3d Group 4: R. vitalii Ast, R. ridleyi A.Gepp, R.

wainionis Steph., R. ekmanii S.W. Amell. Lobes not cal-

cified; lacking idioblasts; scales hyaline or violet; cells

with 2 thickened bands.

30.1. 1.3e Group 5: lobes not calcified; lacking idioblasts;

scales hyaline; cells without thickened bands.

30. 1 . 1 ,3ea Subgroup 1 : R. mauryana. Inferior walls of epi-

dermal cells granulose.

30. 1.1. 3eb Subgroup 2: R. sanguineisporis Ast. Inferior

walls of epidermal cells smooth; cells of dorsal tissue all

with thick walls.

30.1.1.3ec Subgroup 3: R. sorocarpa, R. erythrocarpa Ast.

Inferior walls of epidermal cells smooth; cells of dorsal

tissue not all with thick walls.

30.1.1.3ed Subgroup 4: R. lamellosa, R. viannae Ast. In-

ferior walls of epidermal cells smooth; cells of dorsal tis-

sue with thin walls; scales white, extending beyond

margins of lobes.

30. 1.13ee Subgroup 5: R. ianthina Ast, R. subplana

Steph., R. plano-biconvexa, R. grandis Nees, R. australis

Steph., R. taeniaeformis Ast, R. fruchartii , R. elliottii

Steph., R. breutelii Hampe ex Steph.. R. brittonii Howe,

R. howellii Howe. Inferior walls of epidermal cells

smooth; cells of dorsal tissue with thin walls; scales hya-

line or tinted with violet, not extending beyond margins

of lobes.

30.1.2 Subgenus Thallocarpus: R. curtisii. Dorsal tissue

with air chambers; dorsal epidermis with pores enclosed

by cells, sometimes with lacunae; scales absent; spores

fused in tetrad, surrounded by external layer of exine.

30.1.3 Subgenus Leptoriccia: R. membranacea. Thallus

thin, one layer of lacunae and 2 or 3 layers of cells; scales

present or reduced or absent; spores free at maturity,

spherical, 40-67 pm in diameter, external layer of exine

surrounding each spore; sporoderm ornamented with fine

spinules.

30.1.4 Subgenus Ricciella: thallus thick, up to 15 layers

of cells, spongy; spores tetrahedral, diameter greater than

65 pm; sporoderm ornamented with alveoli or tubercles.

30.1.4.1 Section Spongodes: thalli spongy, divided into

wide lobes; capsules hardly projecting on the two surfaces

of lobes; scales small or very reduced or absent.

R. crystallina, R. cavernosa. Thalli in rosettes, spongy.

10-35 mm in diameter.

224

Bothalia 25,2 (1995)

30.1.4.2 Section Ricciella: thalli wide and deeply divided

or in narrow strips; capsules projecting on ventral surface

of lobes; scales small or rarely reaching edge of lobes.

R. frostii , R. paraguayensis Spruce. Dioicous; het-

erothallic; female plants in rosettes.

R. chiapasensis Ast, R. dussiana Steph.; R. geissleriana

Ast, R. hegewcmiana Ast. Monoicous; lobes 1 .5^1.0 mm

wide; grouped together.

R. jovet-astiae Vianna emend. Vianna, R. crassifrons

Spruce, R. bahiensis Steph., R. stenophylla, R. limicola

Ast, R. paranaensis. Lobes 0.3-1. 5 mm wide.

31. SCHUSTER ( 1992): North America (and elsewhere)

Schuster (1992a) maintained that infrageneric classifi-

cations in the genus Riccia were still unresolved. He now

adopted a multiple system with eight recognizable sub-

genera, although he conceded that his attempts at dividing

them up into a series of complexes and sections were

largely provisional, since he collected chiefly in North

America and Europe. He thought that it was up to students

residing in the other areas to test the validity of his pro-

posed new sections, and that other sections from Australia

and Africa would without doubt have to be recognized.

He nevertheless instituted 10 new sections and formally

provided them with Latin descriptions and designated type

species.

31.1 Genus Riccia : plants usually small; closely prostrate,

normally on soil, rarely aquatic; annual to perennial; in

rosettes or hemirosettes, occasionally gregarious; seg-

ments generally with a longitudinal dorsal groove which

disappears proximally; thallus structure either with dis-

tinct, persistent, rarely disintegrating epidermis with pores,

overlying polyhedral air chambers or without a continu-

ous, defined epidermis and aerenchyma with vertical ca-

nal-like air chambers; thallus margins naked or with

papillae, cilia or setae; ventral scales often inconspicuous;

oil bodies absent; antheridia and archegonia sunken into

thallus; sporophyte consisting of spherical capsule con-

taining spores.

31.1.1 Subgenus Riccia : thalli with photosynthetic tissue

compact, arranged in closely contiguous vertical cell rows

separated by slender vertical air canals; dorsal epithelial

cells breaking down, the hypodermal layer persistent,

regularly arranged; scales small to large; cilia or papillae

often present; spore release dorsal; spores single.

31.1.1.1 Section Riccia: thallus margins blunt or rounded,

naked or with short cilia (under 300 pm); lacking special-

ized oil cells; scales relatively small, usually colourless or

vestigial, failing to reach thallus margins; sulcus weakly

impressed, soon widening and disappearing posteriorly.

3 1.1. 1.1 a Subsection Riccia: thalli never calcified and

chalk-white dorsally; nearly always forming rosettes; spo-

rangia tending to bulge more on ventral side of thallus

than on dorsal; spores polar, with clearly defined wing

margins.

R. glauca, R. hirta, R. tenella Jacobs, R. bifurca, R.

warnstorfii, R. beyrichiana, R. califomica, R. setigera

R.M. Schust. Monoicous; thalli ciliate or not.

R. ozarkiana McGregor. Dioicous; dorsal sulcus wide.

3 1.1.1. lb Subsection Albidae R.M. Schust.: dorsal surface

of thallus calcified, chalk-white; pores open into canals

bounded by 5-7 cell rows; deep dorsal groove not wid-

ening posteriorly.

R. albida (= 1R. crustata). Epithelial cells persistent,

tumid or sometimes conoidally inflated, rough with se-

creted calcium carbonate. This subsection was previously

a section in Schuster (1992a).

31.1.1.2 Section Lamellosae R.M. Schust.: plants always

naked on dorsal surface and margins; epidermal cells usu-

ally ephemeral and collapsing to irregular remnants, not

thick-walled; dorsal groove shaip and usually ± persistent;

scales two-ranked, usually well developed.

R. lamellosa Raddi subsp. lamellosa , R. lamellosa

subsp. austini R.M. Schust.. R. andina Mull.Frib. subsp.

andina, R. andina subsp. chionophora R.M. Schust., R.

albolimbata. Ventral scales hyaline, whitish, projecting be-

yond thallus margins.

R. albolimbata is a southern African taxon which

Schuster now also reports from North America.

Jovet-Ast (1994) found spores in the type specimen of

R. andina and has concluded that it is synonymous with

R. mauryana , which Schuster has placed in another sec-

tion, namely Sorocarpae.

R. macallisteri, R. dictyospora, R. nigrella, R. camp-

belliana Howe subsp. campbelliana , R. campbelliana

subsp. austrigena R.M. Schust. Ventral scales not con-

spicuously projecting beyond moist thallus margins, usu-

ally pigmented.

31.1.1.3 Section Sorocarpae R.M. Schust.: monoicous;

thalli light green; less compact than in section Sommieri ;

epithelial cells with thick bases; hypodermal cells firm;

thallus margins lacking cilia; scales two-ranked and small;

dorsal groove deep.

R. sorocarpa subsp. sorocarpa , R. sorocarpa subsp.

arctica R.M. Schust., R. sorocarpa subsp. erythrophora

R.M. Schust., R. dorsiverrucosa, R. mauryana. Ventral

scales small, obscure, not or hardly imbricate, colourless

or castaneous.

In her experience, Jovet-Ast (1994) finds R. sorocarpa

to be a stable species, although there are certain modifi-

cations in the colour of the lobes and scales, in the thick-

ness of the epidermal cell walls and in the diameter of

the spores and size of the wing. She states that she is

inclined to reject Schuster’s subspecies of R. sorocarpa

as they are based on a very small number of specimens

seen by him and that the rank of subspecies is too impor-

tant to designate to such weak variations.

31.1.1.4 Section Sommieri R.M. Schust.: dioicous; thalli

deep green, with vertical purple flanks; very compact and

Bothalia 25,2(1995)

225

thick; epidermal cells delicate and hypodermal ones strik-

ingly thickened.

R. sommieri. Ventral scales large, imbricate, blackish

purple.

31.1.1.5 Section Atromarginatae R.M. Schust.: thalli dull,

bluish green and deep purple or purple-black laterally;

with a tendency to develop both blunt papillae and hyaline

sharp, rigid cilia mostly under 80 pm long; spores opaque,

nonpolar.

R. atromarginata forma atromarginata , R. atromargi-

nata forma glabra Levier ex Mull.Frib., R. violacea var.

violacea Howe, R. violacea var. iodocheila (Howe) R.M.

Schust., R. violacea var. laevis R.M. Schust. Ventral scale

small, often rudimentary, black-purple; margins some-

times with papilliform cells.

This section evidently consists of highly problematical

species, subspecies, varieties and formae that need to be

clarified.

31.1.1.6 Section Ciliiferae R.M. Schust.: thalli robust;

ventrally with large keel and at apices often forming tu-

bers; margins winged, often ciliate; sulcus deep, soon dis-

appearing; scales hyaline, whitish, semicircular.

R. ciliifera, R. gougetiana , R. erinacea Schiffn., R.

melitensis (from Malta). Thallus surfaces and margins

mostly eciliate or with cilia 100-325 pm long.

Jovet-Ast (1994) doubts whether R. melitensis is placed

correctly in this section, since it lacks marginal cilia; she

also prefers to treat R. erinacea as a variety of R.

gougetiana namely var. armatissima Levier ex Mull.Frib. ,

because all its characteristics resemble those of R.

gougetiana , except for the abundance of bristles.

31.1.1.7 Section Ciliatae R.M. Schust.; plants copiously

ciliate with tapered, sharp cilia, 300-1000 pm long; me-

dian sulcus sharp near apices, soon broadened and shal-

low; flanks swollen; scales small, hyaline to purplish.

R. ciliata subsp. ciliata , R. ciliata subsp. canescens

(Steph.) R.M. Schust., R. ciliata subsp. grisea R.M.

Schust., R. ciliata subsp. trichocarpa (Howe) R.M.

Schust., R. crinita. Cilia very thick-walled along at least

one side, up to 1150 pm long; flanks usually purplish to

purplish black.

Jovet-Ast (1983) has shown that R. canescens Steph.

is a synonym of R. trichocarpa and she finds it unaccept-

able (Jovet-Ast 1994) that R. trichocarpa can be consid-

ered a subspecies of R. ciliata.

31.1.1.8 Section Bicarinatae R.M. Schust.: R. bicarinata.

Thalli at margins with groups of 2-3 basally connate,

thick-walled, sharp, roughened cilia; flanks ascending,

blunt at margins; dorsally with sulcus soon widening.

31.1.1.9 Section Albosquamatae R.M. Schust.: R. al-

bosquamata S.W. Amell. Thalli without cilia or papillae;

ventral scales with regularly oriented, almost quadrate,

very firm-walled cells.

Amell (1957, 1963) had based his description of R.

albosquamata on two white-scaled species (Perold 1989),

R. albolimbata and another which was later named R. ar-

genteolimbata, using characters from both. R. albosqua-

mata is regarded as a synonym of R. albolimbata ; Schuster

did not include this new section Albosquamatae in his

later treatment (Schuster 1992b).

3 1 . 1 .2 Subgenus Ricciella : most species terricolous or

sometimes aquatic; dorsal epidermis chlorophyllose,

smooth or with age lacunose; pores present although

sometimes vestigial; thallus margins always lacking cilia;

air chambers large; ventral scales in a single median row,

semilunate, or (if 2-ranked) much reduced; sporophytes

releasing spores ventrally or dorsally.

3 1 . 1 .2. 1 Section Ricciella'. segments narrow and lingulate to

linear, under 0.8 mm wide; dorsal epidermis normally per-

sistent. usually perceptibly areolate; pores not enlarging.

R. fluitans, R. stenophylla, R. rhenana, R. canaliculata,

R. duplex. R. huebeneriana subsp. huebeneriana, R. hue-

beneriana subsp. sullivantii (Austin) R.M. Schust., R.

paranaensis. Scales remaining in a single median row or

soon splitting into 2 rows; capsules protruding ventrally;

some species aquatic or land plants.

Schuster (1992b) places R. perennis in synonymy un-

der R. canaliculata , as Muller (1953-1958) had done be-

fore him, but this is emphatically rejected by Jovet-Ast

(1994) who states that they are so different in appearance

that one can recognize them even with the naked eye.

31.1.2.2 Section Cavemosae R.M. Schust.: thalli in ro-

settes, fleshy and succulent; photosynthetic tissue loose,

air chambers large, polyhedral: dorsal epidermis delicate;

pores enlarging with age and thallus becoming lacunose.

R. cavernosa. R. crystallina. Ventral scales apparently

lacking or small and ephemeral.

Schuster (1992a) remarks that his section Cavemosae

is perhaps identical with Nees’s Spongodes but he finds

the latter ambiguous because it includes both R. bullosa

and R. crystallina. Whether R. bullosa remains in section

Spongodes is not mentioned. Perold (1991a) referred R.

bullosa to section Spongodes , group 'Vesiculosa’.

Schuster (1992b) notes that the prior usage of R. crys-

tallina (for R. cavernosa sensu Jovet-Ast 1964; 1966) and

R. plana (for R. crystallina sensu Jovet-Ast) was adopted

by all workers and should have been preserved by perti-

nent lectotypification. Riccia crystallina has been lecto-

typified by Perold ( 1992) with the typotype held at FI and

shows that Jovet-Ast was correct after all.

31.1.2.3 Section Frostii R.M. Schust.: R. frostii. Growing

in rosettes; heterothallic with small male plants: photo-

synthetic tissue relatively compact with air chambers nar-

row and tall; scales lacking or vestigial; spores dorsally

released.

31.1.3 Subgenus Leptoriccia : R. membranacea. Thalli

delicate, exceedingly flat and thin, chiefly consisting of

226

Bothalia 25,2(1995)

epidermis and air chambers; ventral tissue vestigial; scales

seemingly absent; sporangia ventrally protruding, spores

single, spherical, apolar, covered in spines.

31.1.4 Subgenus Thallocarpus : dorsal epidermis chloro-

phyllose, with pores, becoming lacunose; thalli spongy,

soft-textured; strongly heterothallic with much smaller

male plants; assimilation tissue with large and high air

chambers in 1-3 layers; spore release dorsal; spores in

tetrads; rhizoids mostly smooth or in part conspicuously

tube rcul ate.

3 1 . 1 .4. 1 Section Thallocarpus : thalli with dorsal surface

smooth; ventral scales lacking or vestigial; spores of tetrad

separated by distinct, ± smooth connecting band; spore

surface with spine-like tubercles.

R. curtisii, R. leptothallus R.M. Schust., R. howei R.M.

Schust. Plants forming rosettes or irregularly gregarious,

with or without colour dimorphism in the sexes.

31.1.4.2 Section Chaetoriccia R.M. Schust.: R. tomentosa.

Distinct from sect. Thallocarpus in (a) thalli dorsally

shaggy-haired, with long free-standing filaments formed

of elongated cells; (b) ventral scales conspicuous, hyaline,

triangular, apices bearing filaments formed of several

cells; (c) spore tetrads only obscurely tetrahedral, very

closely armed with papillae.

Riccia tomentosa was placed in subgenus Thallocar-

pus, section Pannosae by Volk & Perold (1990) and Pan-

nosae was later elevated to the rank of subgenus (Perold

1991), but these must have escaped Schuster’s notice,

when he created the new section Chaetoriccia for R.

tomentosa.

31.1.5 Subgenus Chartacea: R. schelpei. Dorsal epidermis

firm, thick-walled, echlorophyllose; pores conspicuous,

surrounded by 5 or 6 small, thin-walled cells elevated

above epidermal cells; thallus grooved, margins acute;

gametangia limited to groove; spores single.

3 1 . 1 .6 Subgenus Micantes : R. hirsuta. Dorsal epidermis

densely invested with tapered ciliform ( 1 )2— 7-celled

‘hairs’; pores obscure; ventral scales large, 2-ranked,

shaggy; spores (95)115-125(130) pm in diameter.

As pointed out above, the original description of R.

hirsuta was based on two distinct, but superficially rather

similar species, R. hirsuta (section Pilifer) and R. tomen-

tosa (subgenus Thallocarpus, section Pannosae, (Volk &

Perold 1990)). It is regrettable that the initial mistake by

Volk & Perold was perpetuated by Schuster (1992a, b).

31.1.7 Subgenus Pteroriccia: epithelial cells in the form

of free-standing, independent multicellular uniseriate and

often tapered hairs, mostly formed of elongated cells; dis-

tal cells of epithelial filaments persistent, typically smaller,

often conoidal and/or conspicuously elongated.

R. villosa, R. parvo-areolata O.H. Volk & Perold.

Mostly with large ventral scales projecting beyond thallus

margins.

31.1.8 Subgenus Viridisquamata : R. caroliniana. Dorsal

tissues devoid of chlorophyll, which is almost wholly lim-

ited to ventral strata of thallus; ventral surface with bi-

stratose chlorophyllose transverse lamellae in V-shaped

configurations, these giving rise to reduced unistratose

ventral scales.

31.2 Genus Ricciocarpus : R. natans. Plants floating, oc-

casionally terrestrial; dorsal surface with distinct pores

surrounded by 5 or 6 cells; air chambers large and con-

spicuous; ventral scales linear, numerous, sword-like; oil

bodies present; antheridia sunken in thallus or along me-

dian groove.

DISCUSSION

The above treatment has resulted in numerous species

being assigned to different sections and groups in the

works by Jovet-Ast (1991, 1993), and Schuster (1992a,

b). One is also faced with the anomalous situation, that

in two articles in the same publication, following directly

on one another (Perold 1993; Fischer 1993), the same

taxon, R. vulcanicola Eb. Fisch., is placed in two different

sections, since Perold followed the time-honoured tradi-

tion of placing R. crystallina (and its allies, in this case

R. vulcanicola ) in section Spongodes Nees, whereas Fis-

cher chose to follow Schuster (1992a, b), and placed it in

section Cavemosae.

Anyone is, of course, free to follow the treatment of

their choice, but for later students, it could be very con-

fusing. Flowever laudable the motive for arranging the

large number of Riccia species into a series of more or

less natural complexes or sections, so that they can be

more easily ‘digested’, six of the 11 new sections pro-

posed by Schuster (1992a) in the genus Riccia have only

one species and four of the subgenera also only have one

representative each. So, are we really making progress or

are we just proliferating the number of higher ranking

taxa, some of which may not stand the test of time? It

calls to mind a remark attributed to the late Prof. Tom

Harris of Reading University, ‘You can always make any

genus (or subgenus for that matter) uniform merely by

making it monospecific, and thereby defeating the object

of classification’.

It is as well to bear in mind Hassel de Menendez’s

(1976) cautionary advice, ‘since the nomenclatural system

does not allow us to abandon completely names which

are no longer required, continuing in this way will only

result in the accumulation of too many names, each of

which will be protected by the Code’.

Until more revisions worldwide have been completed,

using methods such as biochemical and DNA analysis,

enzyme electrophoresis, multivariate analysis, TEM stud-

ies of the ultrastructure of the spore wall, like that of

Thaithong (1982), and cultivation experiments, it may be

wiser to use informal groups. Once sufficient data have

been gathered hepaticologists should come to an agree-

ment in defining the limits of subgenera, sections, species

and subspecies in the Ricciae.

Bothalia 25,2 (1995)

227

I am not well acquainted with the Ricciae of North Amer-

ica, Latin America, the Mediterranean and Australia, but for

sub-Saharan Africa, I propose the following classification,

which is quite similar to that of Perold (1991), which, being