Bothalia

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Vol. 20,2

Oct./Okt. 1990

PUBLICATIONS OF THE NATIONAL BOTANICAL INSTITUTE, PRETORIA

PUBLIKASIES VAN DIE NASIONALE BOTANIESE INSTITUUT, PRETORIA

Obtainable from the National Botanical Institute, Private Bag X101,

Pretoria 0001, Republic of South Africa. A current price list of all avail-

able 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.

Verkrygbaar van die Nasionale Botaniese Instituut, Privaatsak X101,

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

bare publikasies kan aangevra word.

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

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

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

bevordering van die wetenskap van plantkunde. Die hoofgebiede wat gedek

word, is taksonomie, ekologie, anatomie en sitologie. Twee dele van die

tydskrif en ’n indeks van die inhoud, outeurs en onderwerpe verskyn

jaarliks.

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

MEMOIRS VAN DIE BOTANIESE OPNAME VAN SUID-AFRIKA

The memoirs are individual treatises usually of an ecological nature, 'n Reeks van losstaande omvattende verhandelings oar vernaamlik

but sometimes dealing with taxonomy or economic botany. ekologiese, maar soms ook taksonomiese of plantekonomiese onderwerpe.

THE FLOWERING PLANTS OF AFRICA / DIE BLOMPLANTE VAN AFRIKA

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. 50, one part of twenty plates is published annually. A volume

consists of two parts. The publication is available in English and Afrikaans.

Hierdie reeks bied kleurplate van Afrikaanse plante met bygaande teks.

Die skilderye word meestal deur die kunstenaars van die Nationale

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

ekonomiese waarde vir afbeelding.

Vanaf Vol. 50 word een deel, bestaande uit twintig plate, jaarliks

gepubliseer. ’n Volume bestaan uit twee dele. Die publikasie is beskikbaar

in Afrikaans en Engels.

FLORA OF SOUTHERN AFRICA / FLORA VAN SUIDELIKE AFRIKA

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

Ciskei, Transkei, Lesotho, Swaziland, Bophuthatswana, South West

Africa/ Namibia, Botswana and Venda. 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. Also available in the FSA series are the

following:

'n Taksonomiese verhandeling oor die flora van die Republiek van Suid-

Afrika, Ciskei, Transkei, Lesotho, Swaziland, Bophuthatswana, SWA/

Namibie, Botswana en Venda. Die FSA bevat beskrywings van families,

genusse, spesies, infraspesifieke taksons, sleutels tot genusse en spesies,

sinonimie, literatuur, verwysings na enkele eksemplare, asook beknopte

taksonomiese en ekologiese aantekeninge. Ook beskikbaar in die FSA-

reeks is die volgende:

The genera of southern African flowering plants, by/deur R.A. Dyer, Vol. 1 Dicotyledons (1975); Vol. 2

Monocotyledons (1976).

Keys to families and index to the genera of southern African flowering plants, by/deur R.A. Dyer (1977).

Plant exploration of southern Africa by Mary Gunn & L.E. Codd. Obtainable from/Beskikbaar van: A. A. Balkema

Marketing, Box/Posbus 317, Claremont 7735, RSA.

PALAEOFLORA OF SOUTHERN AFRICA / PALAEOFLORA VAN SUIDELIKE AFRIKA

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 Palaeoflora met ’n uitleg vergelykbaar met die van die Flora van suide-

like Afrika. Baie van die inligting word aangebied in die vorm van tabelle

en fotografiese plate waarop fossiele populasies afgebeeld word. Reeds

beskikbaar:

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

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

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

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

BOTH AO A

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Volume 20,2

Editor/Redakteur: O.A. Leistner

Assisted by B.A. Momberg

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, Kirstenbosch, RSA

University of Utrecht, Utrecht, Netherlands

Editorial Committee

Redaksiekotnitee

O.A. Leistner

F. Getliffe Norris

B.A. Momberg

M.C. Rutherford

ISSN 0006 8241

NASIONALE 30TAN1E -

INST1TUUT

'990 -11- 95

K>-« 1 ». • . t

INSTITUTE

Issued by the National Botanical Institute, Private Bag X 10 1 , Pretoria 0001, South Africa

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

1990

Digitized by the Internet Archive

in 2016

https://archive.org/details/bothaliavolume2020unse_0

CONTENTS — INHOUD

Volume 20,2

1. Ascospore development in Ceratocystis sensu lato (Fungi): a review. P.W.J. VAN WYK and M.J.

WINGFIELD 141

2. A preliminary checklist of Xhosa names for trees growing in Transkei. COLIN T. JOHNSON .... 147

3. Nomenclature in Mesembryanthema (Aizoaceae): the generic names by Rappa and Camarrone. H.E.K.

HARTMANN and V. BITTRICH 153

4. A checklist of the plants of the Karkloof Forest, Natal midlands. J.O. WIRMINGHAUS 159

5. Studies in the genus Riccia (Marchantiales) from southern Africa. 17. Three new species in section

Pilifer. R. elongata, R. ampullacea and R. trachyglossum. S.M. PEROLD 167

6. Studies in the genus Riccia (Marchantiales) from southern Africa. 18. New species in section Pilifer

from the NW Cape: R. furfuracea, R. vitrea and R. namaquensis. S.M. PEROLD 175

7. Studies in the genus Riccia (Marchantiales) from southern Africa. 19. Two new species: R. pulveracea ,

section Pilifer and R. bicolorata , section Riccia, group ‘Squamatae’. S.M. PEROLD 185

8. Studies in the genus Riccia (Marchantiales) from southern Africa. 20. R. albovestita and its synonyms,

R. duthieae and R. sarcosa. S.M. PEROLD 191

9. Studies in the genus Riccia (Marchantiales) from southern Africa. 21. R. stricta comb. nov. and R.

purpurascens, subgenus Ricciella. S.M. PEROLD 197

10. Notes on African plants:

Adiantaceae/Pteridaceae. Doryopteris pilosa var. gemmifera, a new fern variety established.

J.E. BURROWS and S.E. STRAUSS 221

Aizoaceae. A new species of Galenia from the Great Karoo. V. BITTRICH 217

Anacardiaceae. A new species of Ozoroa from the Transvaal. E. RETIEF 219

Annonaceae. Uvaria gracilipes, a new southern African record. K. BALKWILL and A.

NICHOLAS 207

Bryophyta. New and interesting records of mosses in the Flora of southern Africa area: 1.

Sphagnaceae— Grimmiaceae. J. VAN ROOY and S.M. PEROLD 211

Convolvulaceae. The correct orthography and author citation of Falckia. D.O. WIJNANDS and

A.D.J. MEEUSE 208

Cucurbitaceae. A new species of Citrullus (Benincaseae) from the Namib Desert, Namibia. B.

DE WINTER 209

Liliaceae/Asphodelaceae. The type of Chortolirion bergerianum (Alooideae). G.F. SMITH .. 213

Marchantiales. Spore germination, early protonema development and vegetative reproduction in

Riccia, section Pilifer. S.M. PEROLD 214

Orchidaceae. Notes on a rare Cape Disa. H.P. LINDER 216

Poaceae. A new species of Eragrostis (Chlorideae) in the E. curvula alliance. B. DE WINTER 208

11. Ordination as a tool for substantiating and interpreting floristic classifications: a case study. G.B. DEALL

and G.K. THERON 223

12. Miscellaneous note:

Descriptive ecological account of intensive spring flowering of ephemeral vegetation in the Boshof

area. Orange Free State, South Africa. P.J. WEISSER and G. GERMISHUIZEN 229

13. The vegetation of the north-western Orange Free State, South Africa. 1. Physical environment. M.S.

KOOIJ, G.J. BREDENKAMP and G.K. THERON 233

14. The vegetation of the north-western Orange Free State, South Africa. 2. The D land type. M.S. KOOIJ,

G.J. BREDENKAMP and G.K. THERON 241

15. New taxa, new records and name changes for southern African plants. B.C. DE WET, G.

GERMISHUIZEN, B.D. SCHRIRE, M. JORDAAN, B.J. PIENAAR, W.G. WELMAN, C. REID,

C.M. VAN WYK, L. FISH, K.L. IMMELMAN, J. VAN ROOY, S. PEROLD, J. TAUSSIG,

N.P. BARKER and H.F. GLEN 249

16. National Botanical Institute, Pretoria: list of staff and publications 267

Bothalia 20,2: 141-145 (1990)

Ascospore development in Ceratocystis sensu lato (Fungi): a review

P.W.J. VAN WYK* and M.J. WINGFIELD**

Keywords: Ascomycetes, ascospore, Ceratocystiopsis, Ceratocystis, cytology, Ophiostoma, perithecium centrum, ultrastructure

ABSTRACT

Ceratocystis, Ceratocystiopsis and Ophiostoma are important pathogens of trees and some agricultural crops and have

recently been found on proteas and forest trees in South Africa. Taxonomic controversy exists regarding these genera and

ultrastructural studies on the development of asci, uniquely shaped ascospores and centrum structure are inadequate. This

review summarises current knowledge of ascospore shape and development of the centrum obtained from light and electron

microscope studies of Ceratocystis sensu lato. Important questions requiring further investigations are outlined. It is furthermore

proposed that additional ultrastructural studies are required to clarify the current taxonomic disagreement in this group.

Such studies could also identify relationships between these fungi and other Ascomycetes.

UITTREKSEL

Ceratocystis, Ceratocystiopsis en Ophiostoma is belangrike patogene van borne en sommige landbougewasse en is ook

onlangs in Suid-Afrika op proteas en in woudbome gevind Daar bestaan nteningsverskille oor die taksonomiese plasing

van hierdie genera en ultrastrukturele ondersoeke van die ontwikkeling van askusse, uitsonderlike askospore en sentrum-

struktuur van die askokarp is tans ontoereikend Hierdie literatuurbespreking bied 'n opsontming van lig- en elektron-

mikroskopiese ondersoeke van askospoorvorm en sentrumontwikkeling by Ceratocystis sensu lato. Belangrike vraagstukke

wat verdere navorsing vereis, word uitgewys. Vervolgens word voorgestel dat bykontstige ultrastrukturele ondersoeke noodsaaklik

is om die huidige taksonomiese meningsverskille op te klaar. Sodanige ondersoeke kan moontlik ook die verwantskappe

tussen hierdie fungi en under Ascomycetes aantoon.

INTRODUCTION

Ceratocystis E1I.& Halst. sensu lato includes the genera

Ophiostoma H .& P. Sydow, Ceratocystis sensu stricto and

Ceratocystiopsis Upadh. & Kendr. (DeHoog & Scheffer

1984; Upadhyay 1981; Upadhyay & Kendrick 1975;

Weijman & DeHoog 1975). These organisms are important

plant pathogens, especially of trees (Boyce 1961; Clark &

Moyer 1988; Marion & French 1967; Smith 1967; Wismer

1961; Wood & French 1963) and have recently been

recorded from South Africa associated with pine bark

beetles (Wingfield & Marasas 1980) and insect-infested

Protea L. inflorescences (Wingfield et al. 1988).

Ceratocystis s.l. are characterized by ostiolate perithecia,

evanescent asci and, in many cases, ascospores with

galeate sheaths. The development of asci and ascospores

in Ophiostoma and Ceratocystis has received little

attention. As far as we are aware, no cytological study

exists on ascospore development in Ceratocystiopsis ,

despite the importance of these organisms as plant

pathogens and the significance that such studies could

have on their taxonomy and determining evolutionary

relationships.

The generic concept in Ceratocystis s.l. has been the

basis of controversy and has changed regularly since the

first cytological studies of these fungi in 1925. In a number

of cases, the same fungi have been studied under different

names which could lead to confusion. In this review we

follow the taxonomic scheme of De Hoog & Scheffer

(1984) where Ceratocystis and Ophiostoma are treated as

distinct genera. Currently used names of fungi treated in

* Department of Botany and Genetics and ** Department of Microbiology

and Biochemistry, University of the Orange Free State, P.O. Box 339,

Bloemfontein 9300.

MS. received: 1989.09.14.

older literature under different taxonomic schemes are

given in Table 1 with the appropriate synonymies.

The first ultrastructural examination of ascospore

development of Ceratocystis s.l. . was that of Stiers (1976)

on C. fimbriata (Ell. & Halst.) Elliot. Subsequent studies

were on O. stenoceras (Robak) Melin & Nannf. (Garrison

et al. 1979) and O. ulmi (Buism.) Nannf. (Jeng & Hubbes

1980a, 1980b). These studies, however, give little insight

into centrum organization. Moreover, ascospores of

Ceratocystis s.l. have unique shapes and previous studies

have provided almost no information on the structure or

development of these spores.

The aim of this review, is to compile data from past

cytological and recent ultrastructural studies, to summarize

current knowledge of centrum structure and organization

as well as ascospore shape, in Ceratocystis s.l. The

possible implications of these observations in the taxonomy

of Ceratocystis s.l. and their relationships with other

Ascomycetes, comparing certain ultrastructural features,

are discussed.

CENTRUM DEVELOPMENT

Controversy exists as to whether speeies of Ceratocystis

* s. I. should be included in the Plectomycetes, characterized

by closed ascocarps (cleistothecia) (Ainsworth et al.

1973; Malloch 1981), or Pyrenomycetes, with ostiolate

ascocarps (perithecia) (Luttrell 1951). Luttrell (1951)

includes members of Ceratocystis s.l., with their distinc-

tive ostiolate ascocarps, in the Pyrenomycetes. In contrast,

Benny et al. (1980) considered the Ophiostomatales as an

order of the Plectomycetes, with the Ophiostomataceae as

the only family.

142

Bothalia 20,2 (1990)

TABLE 1. — Cytological studies of Ceratocystis and Ophiostoma from 1927 to 1981

Genus and species

Author

A Ceratocystis adiposa (Butl.) C. Moreau

= Ceratostomella adiposum (Butl.) Sartoris

= Ophiostoma majus (Van Beyma) Goid.

B Ceratocystis autographa Bakshi

C Ceratocystis fimbriata (Ell. & Ha!st.) Elliot

= Ceratostomella fimbriata Ell. & Halst.

D Ceratocystis moniliformis (Hedge.) C. Moreau

E Ceratocystis paradoxa (Dade) C. Moreau

= Ceratostomella paradoxa Dade

F Ophiostoma ips (Rumb.) Nannf.

= Ceratostomella montium Rumb.

Sartoris (1927)

Hutchinson (1950)

Bakshi (1951)

* Stiers (1976)

Andrus & Harter (1933)

Gwynne-Vaughan & Broadhead (1936)

Moreau & Moreau (1952)

Dade (1928)

Taylor-Vinje (1940)

G Ophiostoma multiannulatum (Hedge. & Davids.) Hendrix

= Ceratostomella multiannulata Hedge. & Davids.

H Ophiostoma piceae (Munch) Bakshi

= Ceratocystis piceae Munch

I Ophiostoma stenoceras (Robak) Melin & Nannf.

= Ceratocystis stenoceras (Robak) C. Moreau

J Ophiostoma ulmi (Buism.) Nannf.

= Ceratocystis ulmi (Buism.) C. Moreau

Andrus (1936)

Bakshi (1951)

* Garrison et al. (1979)

Rosinsky (1961)

* Jeng & Hubbes (1980a)

A— J Current names of genus and species.

* Ultrastructural studies.

An important feature of the Pyrenomycetes is the

organization of the centrum. Luttrell (1951) proposed a

developmental scheme, where centrum development is

initiated by the formation of sterile and fertile cells. The

fertile cells are initiated by curved hyphal branches and

the sterile cells by additional branching hyphae. These

cells, surrounded by other hyphae, give rise to the

maturing perithecium. The central multinucleate fertile cell

or ascogonium lies above the centre of the perithecium,

rather than at its base, the asci therefore develop

progressively and basipetally. The sterile cells are

distributed between the fertile ascogonial cells, and

they have the important function of providing space

for the irregularly developing asci, by disintegrating

during ascus development.

Asci in the Plectomycetes are evanescent and the

ascospores fill the cleistothecial centrum at maturity with

no special discharging mechanism. Luttrell (1951)

emphasized that the irregular distribution of the asci within

the centrum of Ceratocystis s. 1. , is reminiscent of the

Plectomycetes. Redhead & Malloch (1977) concurred with

this opinion and included Ophiostoma and Ceratocystis

in this order. Furthermore, based on the presence of

ascospores with galeate sheaths, these authors included

the genera with numerous yeasts in the Endomycetaceae.

Typically, Plectomycetes have closed ascocarps (Ainsworth

et al. 1973), and thus Ophiostoma and Ceratocystis with

beaked, ostiolate perithecia could equally be excluded

from this group.

Malloch (1981) suggested that the Plectomycete centrum

has evolved towards progressive simplification. Sterile

tissue, the ostiole, as well as forcible discharge of spores

would therefore have been lost. This argument would

justify inclusion of Ceratocystis s.l. in the Plectomycetes,

despite their having ostiolate perithecia. Our knowledge

of ascocarp development in these fungi is based on few

examples with almost no ultrastructural information.

Ultrastructural studies on the development of the centrum

of Ceratocystis , Ophiostoma and related fungi could

provide important clues clarifying their relationships.

The development of the centrum of Thermoascus

aurantiacus and Ceratocystis thermophile is typical of that

of the Plectomycetes, with asci arising from croziers

formed by ascogenous hyphae (Ellis 1981a, 1981b). Ultra-

structural studies of these species, however, provide

insufficient information to make logical comparisons with

Ceratocystis s.l. The single-celled nature of yeasts

apparently related to Ceratocystis s.l. precludes com-

parisons of centrum development.

CYTOLOGY

The earliest cytological study of Ceratocystis was that

of Elliott (1925) on C. fimbriata, associated with black

rot of sweet potato. Subsequent investigations on this spe-

cies were by Andrus & Harter (1933) and Gwynne-Vaughan

& Broadhead (1936). Cytological studies on related

fungi dealt with Ophiostoma multiannulatum (Hedge. &

Davids.) Hendrix (Andrus 1936), C. paradoxa (Dade)

C. Moreau (Dade 1928), C. adiposa (Butl.) C. Moreau

(Sartoris 1927; Hutchinson 1950), and O. ips (Rumb.)

Nannf. (Taylor-Vinje 1940). This was followed by studies

of C. autographa Bakshi (Bakshi 1951), C. moniliformis

(Moreau & Moreau 1952) and O. piceae (Munch) Bakshi

(Bakshi 1951).

Cytological characteristics of each species of Cerato-

cystis s.l. previously studied are given in Table 2. It is

generally accepted in the Pyrenomycetes, that perithecia

Bothalia 20,2 (1990)

143

are derived from coiled hyphae without fertilization

(Luttrell 1951). Coiling of hyphae, was observed by

different authors in all genera that have been studied

(Table 2). Crozier formation has been reported for most

species, except C. adiposa, C. paradoxa and O. stenoceras

(Table 2). In reference to studies of C. fimbriata (Andrus

& Harter 1933; Gwynne-Vaughan & Broadhead 1936) and

C. adiposa (Sartoris 1927), Hutchinson (1950), in a study

of C. adiposa, however, stated that a simple curved hypha

with four terminal nuclei, is unconvincing evidence of

crozier formation. The inference here was that crozier

formation does not occur in Ceratocystis s.l. In contrast,

crozier formation was illustrated in ultrastructural studies

of O. ulmi (Jeng & Hubbes 1980b). Ascogonia were

observed in all species (Table 2), although some authors

described the ascogonium as either a fertile cell in O. multi-

annulatum (Andrus & Harter 1933) or ascogenous hyphae

in O. piceae (Bakshi 1951) and O. ips (Taylor-Vinje 1940).

In developmental studies of Ceratocystis s. 1. , reference

is commonly made to sterile cells, which include both

cushion and pseudoparenchymatous cells (Table 2).

However, their function has seldom been discussed.

Pseudoparenchymatous cells are dispersed amongst the

ascogenous hyphae and their arrangement appears to

characterise centrum organization (Table 2). Although the

ascogenous hyphae generally emerge towards the base of

the perithecium, they may also extend to the base of the

neck, with the asci developing towards the centre (Table

2). The ascogenous hyphae are usually separated from the

perithecial wall by the cushion cells, with asci forming

irregularly throughout the centrum.

According to Luttrell (1951), the ascogonium is

located above the centre of the perithecium (Figure 1A),

developing basipetally (Figure IB). From available

cytological studies we interpret centrum development in

Ceratocystis s.l. as follows: ascus development is preceded

by the development of ascogenous hyphae (= fertile cells)

from the ascogonium. The ascogenous hyphae developing

from the ascogonium, appear to form towards the base of

the perithecium, with cushion cells (= sterile cells)

surrounding them (Figure 1A). The ascogonium therefore

develops basipetally (Figure IB) and asci are formed by

acropetally developing ascogenous hyphae (Figure 1C).

Ascospores then mature basipetally (Figure ID).

Gwynne-Vaughan & Broadhead (1936) stated that there

is inadequate reference in most literature to the charac-

teristic shape of the ascospores in Ceratocystis s.l. The

ascospore shape of each species previously studied

cytologically, reproduced from descriptions or original

illustrations, is compared in Table 2. Hutchinson (1950)

emphasized that the number of perithecial wall layers and

ascus shape are inadequate criteria for determination of

natural relationships. More emphasis should therefore be

placed on detailed cytological and ultrastructural studies

of centrum organization, as well as ascospore shape and

development, as potential criteria in the taxonomy of

Ceratocystis s.l.

ULTRASTRUCTURAL COMPARISON OF CERATOCYSTIS S.L.

WITH OTHER ASCOMYCETES

Ultrastructural studies on ascus and ascospore de-

velopment in the Ascomycetes are numerous (Dyby &

TABLE 2. — Cytological characteristics of perithecium development in Ceratocystis and Ophiostoma

Coil Crozier Ascogonium Cushion parenchym layers Centrum Ascospore

144

Bothalia 20,2 (1990)

FIGURE 1.— Schematic diagrams of perithecium development in

Ceratocystis sensu lato. A, ascogonium (as) formed above the

centre of the perithecium surrounded by wall hyphae (wh);

B, ascogonium developing basipetally forming ascogenous hyphae

(ah) and cushion cells (c) forming surrounding layer; C,

ascogenous hyphae developing acropetally forming asci (ac)

and neck primordial cells (n) developing from perithecial wall

layers (wl); D, evanescent asci maturing basipetally, releasing

mature ascospores (a). Neck (n) elongating and cushion cells

degenerating.

Kimbrough 1987; Furtado 1971; Honegger 1985; Main-

waring 1967; Merkus 1973; Van Brummelen 1987). In

general, previous ultrastructural studies of ascospore

development have investigated wall formation (Lynn &

Magee 1972; Moens 1971), the morphology of nuclear and

membrane structure (Hashimoto et al. 1960; Oso 1969)

and the role of lomasomes and plasmalemmasomes

(Marchant & Moore 1973). Ultrastructural studies of

ascospore delimitation and development have also been

conducted (Beckett 1981; Rosing 1985). A comprehensive

review of this process has been provided elsewhere (Turian

1976). The latter review did not, however, cover Ceratocys-

tis s.l.

Development of ascospores appears to be similar in all

Ascomycetes. The spore walls are formed between two

delimiting membranes (Ellis 1981a; Stiers 1976). In a

preliminary investigation of C. moniliformis (Figure 2A)

and O. minus (Hedge.) H. & R Syd. (Figure 2B) we have

tentatively confirmed the presence of an electron

transparent endospore and an electron dense epispore wall

layer in ascospores of these species. Amongst fungi with

evanescent asci, ultrastructural studies are available for

Thermoascus aurantiacus Miehe (Ellis 1981a) and

Chaetomium thermophile La Touche (Ellis 1981b) as

well as Ceratocystis spp. However, in the former

fungi, ascospores are elliptically shaped with a charac-

teristic germ pore and are therefore incomparable with

Ceratocystis s.l.

Ascospore walls in Ceratocystis sensu stricto (Garrison

et al. 1979; Jeng & Hubbes 1980a; Stiers 1976) are similar

to those of certain yeasts (Bandoni et al. 1967; Beckett

et al. 1973; Black & Gorman 1971; Hashimoto et al. 1960;

Kreger-van Rij & Veenhuis 1975; Kurtzman & Ahearn

1976). Ultrastructural comparisons can therefore be

made between ascospores with hat-shaped sheaths, in

Hansenula anomala (Hansen) H. & R Syd. (Bandoni et

al. 1967), Pichia spartinae Ahearn, Yarrow et Meyers

(Kurtzman & Ahearn 1976) and certain species of

Ceratocystis s. 1. To our knowledge these are the only fungi

with hat-shaped galeate sheaths and they appear to be

ultrastructurally indistinguishable.

CONCLUSION

With the exception of the hat-shaped ascospores in

Ceratocystis fimbriata, the characteristic and unusual

ascospores in other species of Ceratocystis s.l. have not

been illustrated ultrastructurally. It is proposed that

additional ultrastructural studies are required to clarify the

current taxonomic disagreement in this group. Further

ultrastructural studies on centrum organization, ascus

development and ascospore shape may provide new keys

to relationships between genera of these fungi. A better

understanding of Plectomycetes/Pyrenomycetes relation-

ships could result and might aid in interpretation of

taxonomic and evolutionary relationships in the

Ascomycetes as a whole

FIGURE 2. — Electron micrographs

of mature ascospores. A, asco-

spore of Ceratocystis monili-

formis with hat-shaped sheath

(ep = epispore, en = endo-

spore); B, elongated ascospore

of Ophiostoma minus. Scale

bar = 500 pm.

Bothalia 20,2 (1990)

145

ACKNOWLEDGEMENT

We are grateful to Prof. PS. van Wyk for stimulating

discussions and suggestions as well as Dr Z.A. Pretorius

& Dr B.D. Wingfield for constructive criticism of the

manuscript.

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.

Bothalia 20,2: 147-152 (1990)

A preliminary checklist of Xhosa names for trees growing in Transkei

COLIN T. JOHNSON*

Keywords: ethnobotany, Transkei, trees, vernacular names

ABSTRACT

Xhosa names for 284 of the 392 tree taxa recorded for Transkei are listed. The semantics of the common names are discussed.

The main moulding forces for these names were found to be: tree habit, characteristics shared with animals, medicinal uses

and social or historic events.

UITTREKSEL

Xhosa-name vir 284 van die 392 boomtaksons wat in Transkei aangeteken is, word gelys. Die semantiek van die volksname

word bespreek. Daar is bevind dat hierdie name hoofsaaklik op die volgende oorwegings gebaseer is: groeiwyse van boom,

kenmerke in gemeen met diere. geneeskundige gebruike en sosiale of geskiedkundige gebeurtenisse.

INTRODUCTION

Over the years numerous people have collected and

compiled various lists of Xhosa names for plants. Miller

(1923) listed about 321 names of trees and shrubs. Palmer

& Pitman (1972) in their publication give a multilingual

glossary of Xhosa, Zulu and Sotho names. A publication

from the Transkei Forestry Department (Anon. 1975) listed

about 199 Xhosa names for trees in Dwesa Forest. Sim

(1900) commented that ‘with careful handling these native

names are much more intelligible than some of the

European’. While one cannot advocate the use of common

names in official reports, it appears that in Transkei, where

botanical names are generally unknown and where a

well recognised terminology already exists by which the

different species can be distinguished, it is worthwhile

recording these names along with their botanical

equivalents. With the inception of the research project of

compiling a checklist of the flora of Transkei (Johnson

1985) identification of species proved difficult. To alleviate

this problem and to assist future botanists working in

Transkei, a quick reference system was established and

is presented.

After the present paper had been accepted for publica-

tion, F. von Breitenbach (1989) published a list of Xhosa

tree names as part 6 of his series Standard names of trees

in southern Africa. Most of the names chosen by Von

Breitenbach are also given here. The present paper,

however, provides introductory paragraphs on Xhosa

nomenclature and covers a wider range of names and

orthographic variants.

METHODS

The data for this paper were collected over a period of

five years (1981—85) and present a summary of an

unpublished report (Johnson 1985) and the work of

Johnson & Cawe (1987). During the course of this survey

extensive collecting and field studies were undertaken.

* Botany Department, University of the Western Cape, Private Bag X17,

Bellville 7535.

MS. received: 1988.10.24.

Herbarium specimens are housed at KEI. To ensure that

Xhosa names previously recorded did not go unnoticed,

a number of references were consulted (Anon. 1975;

Bigalke 1967; Broster & Brown 1981; De Lange 1963;

Dornan 1932; Ferreira 1949; Friede 1953; Kropf 1915;

Miller 1923; Mzamane 1945; Palmer & Pitman 1972; Rose

1979; Sim 1900, 1907; Shaw 1938 and Von Breitenbach &

Von Breitenbach 1983).

Emphasis was placed on the collecting and recording

of the Xhosa names and to a lesser extent research was

carried out into their semantics, the reason for their

application.

RESULTS AND DISCUSSION

Various moulding forces can be detected when analysing

the Xhosa names. Herd boys learn to recognise trees from

which they use sticks, women must know which trees they

use for firewood and men must know what is best used

for building huts, kraals, sledges etc. Herbalists have to

recognise trees for their medical usage. Of the 392 tree

taxa recorded for Transkei (Johnson & Cawe 1987), 119

taxa were found to lack a Xhosa equivalent. The 284 tree

taxa having Xhosa names, are listed in alphabetical order

in the checklist.

XHOSA NOMENCLATURE

Miller (1923) was the first to point out the presence of

a generic and ‘species’ name in Xhosa botany. For a better

understanding of Xhosa nomenclature, a list of Xhosa plant

names, more complete than is presently available, is

required. However, analysis of the names collected for

trees support the findings of Miller and is to a degree in

accordance with the rules of the Code of Botanical Nomen-

clature regarding rank concepts. Superficial examination

of the Xhosa names shows words that express a generic

concept. This is illustrated by the use of iGalagala or

umGalagala which refers to Buxus macowanii and B.

natalensis while umGxam and umSintsana are applied to

several species of Schotia and Erythrina respectively.

148

Bothalia 20,1 (1990)

The Xhosa name conveys something characteristic about

a particular species and thus helps to distinguish species.

Commiphora harveyi , umHlanguthi, is distinguished from

C. woodii , imiNyele, by an outstanding feature of the

species. The first named species is described as a soft

wooded tree that is used as a hedge round kraals while

the second species is found growing on the edge of forests.

As can be expected, Xhosa nomenclature does not

necessarily correspond to that of botanical nomenclature.

Some plant names are undoubtedly homonyms, that are

given to more than one species. This is best exemplified

by the use of umBomvane which refers to Cassine peraqua ,

C. papillosa, C. crocea, Cassipourea gerrardii, Eugenia

capensis, Memecylon grandiflorum, Olinia radiata and

Pleurostylia capensis. Other names are polynomials,

several names given to one plant species, as is the case

in Baphia racemosa which is recognized as isiFithi or

uTshiphu.

The greater proportion of Xhosa names must have

evolved at a time when these people lived close to nature.

Older names tend to refer to habitat or habit characters,

newer ones to social events and uses. The moulding forces

for Xhosa names can therefore be divided into a number

of categories.

Outstanding features of the tree or its habitat

UmNukane (Ocotea bullata). The wood possesses an

unpleasant smell when freshly cut. It is this, the same

character, that is recorded in the English common name

‘Stinkwood’.

UmSimbithi ( Milletia grandis), the Xhosa name quoted

by Sim (1900), refers to the extremely heavy, hard and

strong character of the wood. The heartwood is dark brown

and the sapwood is yellow. The wood is currently used

and sought after for knobkieries.

The vernacular name umGomgom refers to the beautiful

tree fern ( Cyathea dregei), that grows up to about 3,5 m,

of which the stem with no durable wood is described as

being ‘empty’.

The dead branches of Podocarpus falcatus and P.

latifolius are a common feature in the immediate

surroundings of these trees and are not collected for

firewood, the Xhosa name umKhoba refers to the lightness

of the dead wood. The common name for Acacia cajfra,

umNyamanzi , refers to the fact that this tree is frequently

found along rivers and streams.

Sharing characteristics with an animal

UmNukambiba (Clausena anisata), the strong un-

pleasant smell of the leaves is embodied in the Xhosa name

which means ‘the bad smell of the striped field mouse’.

UbuHlungu benyoka ( Acokanthera oppositifolia) . Except

for the seeds, all parts of this plant contain latex that is

highly toxic. An infusion made from the powdered bark,

wood or leaves was used as a poison (Watt & Breyer-

Brandwijk 1932, 1962). This feature must have been

known to the early Xhosas as the common name compares

the poison of this plant to snakebite.

UmNukambila (Hippobromus pauciflorus) . The Xhosa

name compares the odour of this tree to that of a wild

animal, such as a rock-rabbit or striped field mouse. The

smell is not as pungent as that of umNukambiba (Clausena

anisata).

UmGobandlovu (Pterocelastrus tricuspidatus) . With its

rounded leaf canopy, this plant forms a major constituent

of dune vegetation. The Xhosa name fancies a resemblance

of the growth form of this plant with the back of an

elephant.

Functional characters

IsiThungwa ( Cryptocarya woodii ), this plant is common-

ly used as the plait when constructing the frame of storage

huts or when building a sledge. The Xhosa name refers

to this functional, ‘plaiting’ feature.

Social or historic events

A large number of names relate to rituals or social

functions, thereby implying that the tree has symbolic

values or magical powers (De Jager 1963). Some of the

names have an obscure meaning which might be of con-

siderable historical interest.

UmNgquza (Biaus natalensis) relates to a feast where

the bride prepares for marriage.

Intshilo ( Calpumia sepiaria). The powdered bark is used

by the warriors to wash their bodies, while the green leaves

are burnt, creating smoke. This ritual is believed to pro-

tect the warriors against their enemies.

Imbulanyathi (Osyris lanceolata), refers to a mystical

animal with the power and strength of a buffalo, which

can assume the shape of a human.

UmLungumabele (Zanthoxylum capense). This much

branched tree is characteristically armed with thorny

protuberances. These protuberances, which may be as

much as 60 mm in diameter, are jokingly compared with

the breasts of European women.

Medicinal properties

Plants are commonly used for medicinal purposes

(Mzamane 1945) and of the 392 tree taxa that were

recorded, 212 are believed to have healing power. The com-

mon name seldom refers to the medicinal properties of

the tree but rather describes the condition of the patient.

The rural tribal people are greatly influenced by their

natural surroundings and each family has its own ubuLawu

or iyeza lasekhaya (medicine of the home) which can

include Hippobromus pauciflorus, Acalypha glabra and

Burchellia bubalina.

Nomina nuda names

Finally, some plant names are apparently empty names,

derived from nothing specific, left over from old lore or

obsolete forgotten phrases. This is best exemplified by the

Bothalia 20,2 (1990)

149

use of umCandathambo and umCane which refer to

Allophylus africanus and Sclerocarya birrea subsp. cajfra

respectively. The generic name for Protea, isiQwane and

the common name isiQwane-sehlati for Rapanea

tnelanophloeos are to a degree bare, except that they are

distinguished by the fact that Rapanea tnelanophloeos is

referred to as occurring in a forest.

ACKNOWLEDGEMENTS

The author wishes to express his appreciation to the

inhabitants and informants of the various districts of

Transkei who were willing to share their knowledge. I owe

special thanks to Messrs W. Dutton, T.M. Sokutu and H.

Kalala who provided valuable comments on the

manuscript. A special word of thanks to Sharon Slingers

and Belinda Feris for typing the manuscript.

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Director of Forestry, Transkei. Unpublished.

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districts of Transkei: Bizana, Flagstaff, Libode, Lusikisiki, Qumbu.

Unpublished.

BROSTER, J.A. & BROWN, H.C. 1981. Amagqirha, religion, magic

and medicine in Transkei, 1st edn, 2nd imp. Via Africa Ltd.

DE JAGER, E.J. 1963. Notes on the magical charms of the Cape Nguni

tribes: 293-302. Fort Hare.

DE LANGE, M. 1963. Some traditional cosmetic practices of the Xhosa.

Annals of the Cape Provincial Museum 3: 85—95.

DORNAN, S.S. 1932. Some beliefs and ceremonies connected with the

birth and death of twins among the South African natives. South

African Journal of Science 29: 690-700.

FERREIRA. F.H. 1949. Bantu customs and legends protect trees. African

Wild Life 3: 59.

FRIEDE, H.M. 1953. Sacred trees of the Bantu. Trees of southern Africa

5,2: 4-5.

JOHNSON, C.T. 1985. Identification of pharmaceutical plants and

traditional medicine in Transkei. Department of Botany, University

of Transkei. Unpublished.

JOHNSON, C.T. & CAWE, S. 1987. Analysis of the tree taxa in Transkei.

South African Journal of Botany 53: 387—394.

KROPF, A. 1915. Kaftr-EngUsh Dictionary, 2 edn by R Godrey. S.A

Lovedale Mission Press.

MILLER, O.B 1923. A list of some native names of trees, shrubs etc. .

in use in the Transkeian Territories, Union of South Africa. Forest

Department, Bulletin No. 8, Pretoria.

MZAMANE, G.l.M. 1945. Some medicinal, magical and edible plants

used among some Bantu tribes in South Africa: 29—35. Fort Hare

paper Series No. 1.

PALMER, E. & PITMAN, Z. 1972. Trees of southern Africa, Vols 1—3.

Balkema, Cape Town.

ROSE, B. 1979. Xhosa plant names. Bantu Cancer Research Registry,

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SHAW, E.M. 1938. Native pipes and smoking in South Africa. Annals

of the South African Museum 24: 277—302.

SIM, T.R. 1900. Botanical observations on forests of eastern Pondoland.

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104-114.

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of Good Hope. Taylor & Henderson, Aberdeen.

VON BREITENBACH, F. & VON BREITENBACH, J 1983. Note on

the natural forest of Transkei. Journal of Dendrology 3: 17—53.

VON BREITENBACH, F. 1989. Standard names of trees in southern

Africa. Part 6: Xhosa tree names. Journal of Dendrology 12:

15-22.

WATT, J.M. & BREYER-BRANDWIJK. M.G. 1932. The medicinal and

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Livingstone, Edinburgh and London.

A CHECKLIST OF TREE NAMES AND THEIR XHOSA EQUIVALENT

Acacia

caffra (Thunb.) Willd., umNgatrianzi, umNyamanzi. umTholi, umThole,

umToli, umTolo

karroo Hayne, umNga, umNga-mpunzi

kraussiana Meisn. ex Benth., uBobo

Acalypha glabrata Thunb., iTaboti, uTaboti, umThombothi, isiThombothi,

isiTomboti, isiToboti

Acokanthera

oblongifolia (Hochst.) Codd, iNtlungunyembe

oppositifolia (Lam.) Codd. ubuHlungubenyoka. ubuHlungu, isiHlungu-

sehlathi, iNxinene, inTIungunyembe

Acridocarpus natalitius Juss., umThenenenda, uMabophe

Alberta magna E. Mey., isiQalaba. uMabophe

Albizia adianthifolia (Schumach.) W.F. Wight, isiCangca, umDhlandohi.

umDIandlothi, umHIandlothi, umKhankqa. umNebelela, umNe-

belele

Allophylus

africanus Beauv., iNqala. inQala

decipiens Radik., umCandathambo

dregeanus (Sond.) De Winter, umCandathambo, uThabathani

natalensis (Sond.) De Winter, uZingathi.

Aloe

bainesii T.-Dver, umGxwala, iKalane-enkulu, iMpondondo. iNkala-

neenkulu, uPhondonde

ferox Mill., umHIaba. iKhala

rupestris Bak., uPhondonde

Anastrabe integerrima E. Mey. ex Benth., umNqumaswele, isiPambata,

isiPambato, isiPhambatho, isiPambati

Andrachne ovalis (Sond.) Muell. Arg., umBeza, umBheza. umGqwata,

noGqwata, uNogqwata,

Apodytes dimidiata E. Mey. ex Arn., umDakana, umDakane, umDekana,

umDikane, umKhwenkwe

Arundinaria tesselata (Nees) Munro, uDuli

Avicennia marina (Forssk.) Vierh., isiKhungathi

Azima tetracantha Dim., iGcegceleya, iGcegceya, uGegaya, umGeya

Bachmannia woodii (Oliv.) Gilg, umPhunzisa, umTswantswantsa

Bauhinia bowkeri Harv., umDlandlovu

Baphia racemosa (Hochst.) Bak.. isiFithi, uTshiphu, uTshuphu

Bequaertiodendron natalense (Sond.) Heine & J.H Herns!. . umThong-

wane, umThongwane-wehlathi. umThungwane, umTongwane, unt-

Tongwani, umTungwane

Bcrsama

swinnyi Phill., umDIandla, isiNdiyandiya

tysoniana Oliv. , isiNdiya-ndiya

Boscia albitrunca (Burch.) Gilg & Ben., umGqomo-gqomo, umPunzito,

Bowkeria verticillata (Eckl. & Zeyh.) Schinz. iGqabi-lesiduli,

Brachylaena

discolor DC. subsp. discolor, umPhahla, iPhahla

elliptica (Thunb.) DC., isiDuli, umDuli, isiDulisehlathi. uDuliwehlathi

Bridelia micrantha (Hochst.) Bail/., umHIahla-makwaba. isiHlala-

makwaba, umHIahlangu, uHlahlangulu

Bruguiera gymnorrhiza (L.) Dun., umHIuma, isiKhungathi, isiKhan-

gazi, isiKhungathi, isiQungathi

Buddleja

saligna Willd., umGeba, umNceba, umNquma

salviifolia (L.) Dim., iGcwizigcwizi, iGqange, iGwangi. iHlothiyana.

iLotana, iLothane, iLotyana, iLotyane, iYotwana

Burchellia bubalina (L.f.) Sim. umFinca-thobankomo, umFincamlincane-

wehlathi, umFincane. umFineane-wehlathi, umFince-wehlathi,

ubuHlungu-benyoka. iThobankomo, iThobankone. iThobankone-

ficane wehlathi, iZothwane

Buxus

macowanii Oliv. , untGala-gala, iGala-gala

natalensis (Oliv.) Hutch., iGala-gala, untGala-gala. umNgquzu. isiXeza.

ukuXeza, uXwezo

Calodendrunt capense (L.f. ) Thunb., umBaba, untBhaba. umBhabhu

Calpoon compressum Bcig., umBalunyathi. umBulunyathi. inTekaza. iM-

bulunyathi

150

Bothalia 20,2 (1990)

Calpurnia

aurea (Ail.) Benth., inDlole, iNdloli, umHlahlambedu, umSiphane,

umSitshana

Canthium

inerme (L.f.) Kuntze, ubuChotshana, umNyushulube, iliTyelehlathi,

umVuthwamini

mundianum Cham. & Schlechtd., umNqayimbila. umSantulane, umSan-

dulane, umSuntulane

obovatum Klotzsch, umBombemfene, umBonemfene

spinosum (Klotzsch) Kuntze. isiPhambatha, isiPambato

Capparis

sepiaria L., inTshilo, inTsihlo, uPasimani

tomentosa Lam., umFihlo, imFihlo, umPasimani, inTsihlo

Carissa

bispinosa (L.) Desf. ex Brenan, isiBetha-nkunzi, isaBetha-inkunzi,

isiBetha-umtunzi, isiBetha-nkunzi, umVusankunzi

macrocarpa (Eckl.) A. DC., amaThungula, umThungula, umTungula,

umTungulu

Cassine

aethiopica Thunb., umNqayi, utnGxube, umNguyi, umBomvane

crocea (Thunb.) Kuntze, umMaqundeni, umThunyalele

papillosa (Hochst.) Kuntze, umBomvane, iKhukhuzi, umMakhankatha

peragua L., umBovane, umBomvane, iKhukhuzi. uMaphipha

Cassinopsis

ilicifolia (Hochst.) Kuntze. isiBethankunzi-sehlathi, iCegceya, iGcegceya

tinifolia Harv., umHlaza

Cassipourea

flanaganii (Schinz) Alston, umMemezi

gerrardii (Schinz) Alston, umBomvane. umHlamalala, umKatane,

umKhathane, umMaphiphe, isiThushana

gummiflua TuL, umBiko, umKanga, iQonga

Catha edulis (Vahl) Forssk. ex End/., iGqwaka, umHIwazi

Celtis

africana Bunn, f, umVumvu

gomphophylla Baker, umVumvu

Chaetacme aristata Planch.. umKaboti, umKambothi. umKhovothi. um-

Kombota, umKovoti

Chrysanthemoides monilifera (L. ) Nor!.. iTholonja. ulwAinfiti

Clausena anisata (Willd. ) Hook. f. ex Benth., isiFutu. umFuto, umNukam-

biba, umNukambele, umNukambile, umTuto, isiTutu

Clerodendrum glabrum E. Mey. , isiDwadwa. umKangazani. uNukisi-

qaqa, umQangazani, umQwaqu. Qwaqwana

Cliffortia nitidula (Engl.) R E. & T.C.E. Fries, umBele, uNwele

Clutia pulchella L., uFiyo, umSipane, iQadi

Cola natalensis Oliv, imBalonyathi, umNqayana. umTenenda. um-

Thenenenda

Combretum

bracteosum (Hochst.) Brandis, uQota, uQoto

calf rum (Eckl. & Zeyh.) Kuntze. umDubu, umDubi.

erythrophyllum (Burch.) Sond., umDubu-wehlathi, umDubu

kraussii Hochst., umDubu-wehlathi, umDuba-wehlathi

Commiphora

harveyi (Engl.) Engl., umHIanguthi, umHlunguthi

wtxxlii Engl. , umHlunguthi. umHlunguthi weHlathi. imiNele, imiNyele,

iNyele

Cordia caffra Sond , umHIovu hlovu, umLovu-lovu. umNovu-novu

Croton sylvaticus Hochst., umEgwaqane, untEgwaqanisa, untFeze, um-

Fezu. umGwaqane, umGwaqanisa, uMagwacane, uMagwaqane

Cryptocarya

latilolia Sond., umGxaleba, umGxobothi, umNcatyana, umThungwa,

umThunywa, umTungwa, umXaleba, iXoboti

myrtifolia Stop], umGqebe, umNcatyana, umThungwa, isiThungwa,

umThungwane, umThungwe, umTungwa, umXaleba, iXoboti

woodii Engl., umNcatyana, umThungwa, isiThungwa, isiTungwa, um-

Xaleba, iXoboti

wyliei SiapJ, umNcatyana, iNqanyana, umXaleba, iXobothi

Cunonia capensis L., umQwashube

Curtisia dentata (Burnt J.) C.A. Sm., isiBande, umGuna. umGxima, umG-

zina, umGxina, umHIebe, umLahleni

Cussonia

sphaerocephala Strey, umSenge

spicata Thunb., umSenge

Cyathea

capensis (L.f.) J.E. Sin.. isiHihi, umKhoma-khoma

dregei Kuntze, umGomgom, isiHihi

Dais cotinifolia L., iNtozani, umTozane, inTozani

Dalbergia

armata E. Mey., luBobo, uBobo, uBobu, iTozane, umZungulu

obovata E. Mey., uZungu, noZungu, umZungulu

Deinbollia oblongifolia (E. Mey. ex Arn.) Radik., umAsibe, umAsibele,

umBangabanga, uMasibele, umSibe

Diospyros

lycioides Desf, umBhongisa, umQokolo, umTshiwane, umTshicwane

natalensis (Harv.) Brenan, umQokolo, umSitshana, umSitshane, uTshi-

wane

villosa (L.) De Winter, umBhongisa, umCandane, umTshekisane

simii (Kuntze) De Winter, umBhongisa

whyteana (Hiern) E White, umKhaza, umKhaze, iNtsazimane, um-

Tenatena, umTenatene, inTsazimane, umTshikivane, umTshi-

kivane

Dombeya

cymosa Harv, umSengathi, umZingathi

tiliacea (Endl.) Planch., iTyibe, iTyibo

Dovyalis

caffra (Hook. f. & Harv) Hook, f, umQokolo, iNgali

rhamnoides (Burch, ex DC.) Harv, umNyezana, umNyezane, um-

Qaqoba

Duvernoia adhatodoides E. Mey. ex Nees, isiBiko, iHlwehlwe, isiPheka

Dracaena hookeriana K. Koch, umKhomakhoma, umKobakoba

Dry petes

arguta (Muell. Arg.) Hutch., umGudlamfene, umHlagela, iNyanzane

gerrardii Hutch., umHIakela, umKhiwane

natalensis (Harv) Hutch., umKhiwane

Ehretia rigida (Thunb.) Druce, umBotshani, umHlele, umHIeli, iNqo-

bogobana

Ekebergia capensis Sparrm., umGwenya, umGwenye wezinja, um-

Gwenyezinja, umNyamati, umQwehle, umQwengazinja, um-

Qwenga-izinja

Encephalartos sp., umPanga

Englerodaphne subcordata (Meisn.) Engl., umHIoza, iNtozane, inTonzane

Erythrina

caffra Thunb., umSintsi

humeana Spreng., umSintsana

latissima E. Mey., umGqwane, umKuwane, umNqwane, umSintsana,

umQwane

lysistemon Hutch., umSinsi, umSintsana, umSintsi

Erythroxylum

emarginatum Thonn., uQadansube, uTimani

pictum E. Mey. ex Sond., umDwibi, umGcamashe, iNtozani

Euclea

crispa (Thunb.) Guerke, umGwali, umHlungu, inTshekisane

natalensis A. DC., iDungampunzi, umKhaza

Eugenia capensis (Eckl. & Zeyh.) Harv & Sond., umBelvane, umBom-

vane, isiDuli, isiDuli sehlathi, isiDul-sehlathi, umHela-hela,

uMaphipha, iTolowa, umTsheka, umTsheqa

Euphorbia

grandidens Haw., umHIontlo, umHIonhlo

triangularis Desf, umHIontlo, umHIonhlo

tetragona Haw., umHIontlo

Faurea

macnaughtonii Phill., iCuba lethole, Cuba latole, iSafo, iSefi, isiFo,

isiQwane

saligna Harv, iCuba lethole

Ficus

bizanae Hutch. & Burtt Davy, umKovoti, umThombe

craterostoma Warb. ex Mildbr. & Burr., umThombe

sur Forssk., umKhiwane

natalensis Hochst., uLuzi, umThombe, umTombi, umZombi

Garcinia gerrardii Harv ex Sim, umBande, umBandi, umBinda, um-

Bindi, umNebelele wehlathi

Gardenia

amoena Sims, umThongathi, umThongothi

thunbergii L.f, umKancaza, umKangaza, umKangaze, umKhangazi,

um Khangaza

Grewia

lasiocarpa E. Mey. ex Harv, uHlolo, umHIolo, uLuzi

occidentalis L., umNqabaza, umVilani, umVileni

Greyia

flanaganii H. BoL, uSinya, umVilani-wamaxhegokazi

sutherlandii Hook. & Harv, umBerebere, inDula, umDalu

Halleria lucida L., umBinta, umBinza, umBiza, uBiza

Harpephyllum caffrum Bemh., umGwenye, umGwenyezinja, umGwenye-

wezinja, umGwenye-hangula

Heteromorpha trifoliata (Wendl.) Eckl. & Zeyh., umBangandlatho, um-

Bangandlele, umBanganolala, umBongondlala

Heywoodia lucens Sims, umNebelele, umNebelele wehlathi

Hibiscus tiliaceus L., miLolo, umLolwa

Hippobromus pauciflorus (L.f.) Radik., ulwAtile, ulwAthile, umFazoneng-

xolo, Futa, isiFuta, isiFutha, umHlwathile, uLathile, ubuLawu,

uLwathile, umLungumabele, umNukambila, umNungumabele,

Bothalia 20,2 (1990)

151

umNquma, ulWTile

Homalium dentatum (Harv.) Warb., iGqabela, iGqabile, iKomanci,

iKomanzi

Ilex mitis (L.) Radik., umDuma, isiDumo, isiDuma

Indigofera

cylindrica DC., umSipane

natalensis H. Bol., uSitshana

Jubaeopsis caffra Becc. , iKomba, iNkomba,

Kiggelaria africana/,., umDuma, umHIandlela, umHlandela, umHIin-

zanyathi, umHIinziyati, umKhokonkho, umKokoko, umLuvuluvu,

umVethi, umVeti

Leucosidea sericea Eckl.& Zeyh., isiDwadwa, iNtshitshi , inTshitshi, um-

Tyityi

Linociera

foveolata (E. Mey.) Knobl., umDlebe, umHIebe, umNqumaswele, um-

Qumaswele, umQumaswile

peglerae (C.H. Wright ) Gilg <6 Schellenb., umDlebe

Macaranga capensis (Baill.) Benth., umBengele

Maerua

cafra (DC.) Pax, umQomoqomo

racemulosa (DC.) Gilg & Ben., umBhonziso, umMphunziso, umPhun-

ziso, umPunziso

lanceolata Eorssk., iNtendekiwane, inTendekiwane. umTendekwane,

inTendekiwane, umTentekwane, umTentekwani, isiThende

Maytenus

acuminata (L.f.) Loes., umNama, iNqayi, umNyama, umZungulwa,

umZungulwo

heterophylla (Eckl. & Zeyh.) N.K.B. Robson, umHIongwe, umKoko-

hoha, umQaqoba, umQaqoha, umQuqoba

mossambicensis (Klotzsch) Blakelock, inGqwanganeyehlathi, iNgqwan

ganeyehlathi,

nemorosa (Eckl. & Zeyh.) Marais, umHIangwe,

peduncularis ( Sond .) Loes., umNgqi. umNqayi

procumbens (L.f.) Loes., umPhonophono

undata (Thunb.) Blakelock, umGorane, umGovane, umGurane, um-

Kokane, umKokusa, umNqayimpofu, umQaqoba wehlathi

Memecylon grandiflorum R. & A. Fernandes, umBande, umBomvane,

umBondi

Micrococca capensis (Baill.) Prain, uBubu

Millettia

grandis (E. Mey.) Skeels, umKunye, umSimbithi

sutherlandii Harv., umGunye, umQunye

Mimusops

caffra E. Mey. ex A. DC., umHlungulu, umHlope, umNweba, um-

Thunzi, umTunzi, umYoyiga,

obovata Sond., amaSetole, amaSethole, amaSithole, amaSitole, um-

Thunzi, umTunzi, umTunzi wehlathi

Monanthotaxis caffra (Sond.) Verde., isiDwaba

Noltea africana (L.) Reichb. f, umGlindi, umGlinidi, umKhuthula, um-

Khuthuhla, maKutula, amaLuleka, iPalode, iPhalode, iPhalo

elide, iYeza lesidiya

Nuxia

congesta R. Br. ex Fresen., umKhobeza, umKobess

floribunda Benth., isiKali, isiKhali, inGqota, iNgqotha

Ochna

arborea Burch, ex DC., umTensema, inTensema, umTentsema, um-

Thentsema, umTentsenda, umVithi

natalitia (Meisn.) Walp., umBomvane-ncici, umBomvu, isiBomvu, um-

Thentsama

serrulata (Hochst.) Walp., iliTye

Ocotea

bullata (Burch.) Baill., umHlungulu, umNimbithi, umNimbiti, um-

Nukane

kenyensis (Chiov.) Robyns, umNukani

Olea

africana Mill., umNquma, umQuma

capensis L., iGqwanxe, iGqwanci, iGqwanxe, iGqwangxe, iGxanci, iG-

qwhanxi, iGxwanxi, umHIebe, umNquma-swile, umSinjane

woodiana Knobl., umGqukunqa, umNqurunqa, umNququnya, um-

Qoqunya

Olinia

emarginata Burn Davy, umNgona-lahlo, umNgene-lahle, uQudu

radiata J. Hofmeyer & Phill., umBomvane, umPhanzi

ventosa (L.) Cufod., umPhanzi ongenalahle, umNonono, oNqena-lahle

Osyris lanceolata Hochst. & Steud., umBulunyathi, uMbulunyathi, iN-

tekaza

Oxyanthus speciosus DC., isiBika, umBinda, umBindi, iHlwehlwe, iM-

pekane

Pachystigma macrocalyx (Sond.) Robyns, inTzane, umVilo-wehlathi

Pappea capensis Eckl. & Zeyh., iliLitye, iLitye

Passerina rigida Wikst., iNwele

Pavetta

lanceolata Eckl., umDIesa, umGonogono, umHIeza, iLitoba, um-

Ponyane, iliToba

revoluta Hochst., umCilikishe, isiFufufu, usKolpati, umPonyane,

iNtozane, uSkolpati, iriTozane

Peddiea africana Harv., isiFufufu, Gcamche, uLuzi, iNtozane, um-

Ponyane, inTozane

Phoenix reclinata Jacq., iDama, uSundu

Phyllanthus discoides (Baill.) Mull. Arg., umPanzite, umPhanzithe

Pittosporum virdiflorum Sims, umGqengqwe, umKwenkwe

Pleurostylia capensis (Turcz.) Oliv., umBomvane, umBomvani,

uMaphipha, umMaphipha, umNgqangqa, umNqwanqwa, um-

Thunyelo

Portulacaria afra Jacq. , iGanishe, iGwanishe

Podocarpus

falcatus (Thunb.) R. Br. ex Mirb., umCeya, umKhoba, umKoba, um-

Kolaya, umKoleya, umSonti

henkelii Stapf ex Dallim. & Jacks., umSonti

latifolius (Thunb.) R. Br. ex Mirb., umCeya, umGeya, umKhoba, um-

Koba, umSonti

Premna mooiensis (H.H.W. Pearson) Pieper, umCacabane, umKakabane,

umTyetyembane, umTyetyembani, umTyintyambani

Protea sp., isiAdlunge, isiQwane

Protorhus longifolia (Bemh.) Engl., isiFuce, umHluthi, umHluthi-

wezinja, umKhambathi, iKhubalo, umKomiso, umKumbati, um-

Kumbat, umKupati, umLuthi, uZintlwa

Prunus africana (Hook, f.) Kalkman, umKakase, iTywina-elikhulu

Psychotria capensis (Eckl.) Vatke, umGonogono, unoGqiwathi-wehlathi,

uGqiwathi-wehlathi, umGubhe, iHwehlwe, umSalamiya, iliTye-

lehlathi

Ptaeroxylon obliquum (Thunb.) Radik., umTali, umTati, umThathi, um-

Thati, umThothe

Pterocelastrus

echinatus N.E. Br., iBolo

tricuspidatus (Lam.) Sond., iBholo, iBolo, umGobandlovu, iTywina,

uTwina

Putterlickia pyracantha (L.) SzyszyL, umHIangwe, umKokhoba, iNtlang-

wana, inThlangwana

Quisqualis parviflora Gerr. ex Harv., uQhotho

Rapanea melanophloeos (L.) Mez, isiQalaba-sehlathi, isiQalaba-hlathi,

isiQwane, isiQwane-sehlathi, isiQwane-wehlathi, isiQwan-

dashube, isiQwand-weshube

Rauvolfia caffra Sond., umHlambamase, umHlambamasi, umlela, um-

Jele, umJelo, umTelo, umThondisa, umThundisa, umTundisa

Rawsonia lucida Harv. & Sond., umLongo, umNqagi-maphuthi,

umNqayi-wephuthi, umNqayi-masende, umNqeyi-masende, um-

Pitshi-wehlathi

Rhamnus prinoides L'Herit., umGilindi, umGlindi, umNonye, umNyenye

Rhizophora mucronata Lam., umHluma, umKhangathi, isiKhangathi,

isiKhungathi

Rhoicissus

digita (L.f.) Gilg & Brandt, iSaqoni-esincinci

rhomboidea (E. Mey. ex Harv.) Planch., umThwazi

Rhus

chirindensis Bak., umHlakothi, umHlokothi, umHlakotini, umHla-

kothi-omkhulu, iNcakotsa, iNtlolokotshane, iNtlokotshane-

enkulu, iNyangane, isiNyangane, iTlokotshane-enkulu

fastigiata Eckl. & Zeyh., iNtlokolotshane-encinci, inNtlokotshane-

encinane, iNtlokolotshane-yedobo, inNtolokotshane-enkulu,

inTlolokotshane-yedobo

laevigata L., umHlakoti, iNcakotsa, isiNyangani, umTlakoti

lucida L., iNtlokotshane-ebomvu, inTlokotshane-bomvu , inTlakotshane-

bomvu

macowanii Schonl., umHlakothi

natalensis Bernh., uGana, umGwele, uGwele, unGwele, iNgwele

nebulosa Schonl., umHlokotshane

Rinorea angustifolia (Thouars) Baill., uGudhla-mfeni, uGudla-mfene,

ukuTyakwemfene

Rothmannia

capensis Thunb., iBolo, umGuba, umGubhe, isiGubhu, umGupa, um-

Sugusu, umZukuza

globosa (Hochst.) Keay, umGube, umSugusu

Salix

capensis Thunb. , umBenya, umBhenya, umGculube, umGcunuba, um-

Gcunubi, umNceunch, umNculuba, umNcunube, umNcunubi,

umNgcunube, umSwi, umThentsema

mucronata Thunb., umBenya, umGcwimbe, umNgcwimbe, umNgcu-

nube, umNgcunubi, umYezana

Sapium ellipticum (Hochst.) Pax, umHongolo

152

Bothalia 20,2 (1990)

Schcfflera'umbellifera (Sond.) Bail!., umBengele, umBumba, umGezisa,

umKisiso, umRongo, umSengane, umSenge, umSenge-mbuzi,

umSenge-wempunzi, umSengu-mbute

Sclerochiton harveyanus Nees , umHlalani, isiThibothi

Scholia

afra (L.) Thunb. var. afra, umGxam

brachypetala Sond., umGxam, umQonci

latifolia Jacq., umGxam, umXamo

Sclerocarya birrea (A. Rich.) Hochst. subsp. caffra (Sond.) Kokwaro.,

umCane

Scolopia

mundii (Eckl. & Zeyh.) Warb., umKokoko, isiNqandazembe, isiNqandi-

zembe, umNqanqa, iQumza,

zeyheri (Nees) Harv., iDumza-elinameva, umKhakboda, umKhakhoba,

umQaqoba, iQumza-elinameva

Scutia myrtina (Bunn f.) Kurz, isiBhingo, isiBhingwa, uBobo, umKapula,

umKokwane, isiNcumncum, isiPhingo, isiPinga, isiPingo,

isiPingwa, umQapuma, uQapuma, umQaphula, uQapula,

umQaqoba-omnyama, umQokwane, umSondezo, iTywina

Seemannaralia gerrardii (Seemann) Vig. , uMaweni

Sideroxylon inerme L., umGqwashu, aMasethole, uMasethole, uMaze-

tole, Mazetole uMazetoli, Mazetoli, umQwashu

Smodingium argutum E. Mey. ex Sond., umThomvane, uThovane, um-

Tomvane, umTovane, Tovane, uTovani, Tovani

Solanum giganteum Jacq., iCuba-lasendle

Strelitzia nicolai Regel & Koern., isiGceba, isiGude, iKamanga

Strophanthus speciosus (Ward & Harv.) Reber, umKhukhumeza

Strychnos

decussata (Pappe) Gilg, umHlamalala, umKhangaza, umKhangele, um-

Nama, iNama

henningsii Gilg, umCaloti, umKaloti, umNonono, umNomo

mitis S. Moore, umNdwibi, umNgqongqothi, umNgqungquthi,

uMngqungquthi, umNqunquthi, umNqunquti

spinosa Lam., umHlaba, umHlala, umHleli, iNhalo, inSala

usambarensis Gilg, uNdlunye

Suregada africana (Sond.) Kuntze, isiThubi

Syzygium

cordatum Hochst., umDoni, umJom, umJomi, umSibi, umSwi

gerrardii (Harv. ex Hook, f.) Burn Davy, umBomva, umJome, umJome-

wehlathi, umJomi, uManzani, isiTholothi

Tarchonanthus

camphoratus L., isiDuli, isiDuli-sehlathi, isiDuli-wehlathi, isiDuli-

selinate, isiDuli-sasendle, isiDuli-selindle

trilobus DC., isiDuli sehlathi

Teclea

gerrardii Verdoorn, umBoza

natalensis (Sond.) Engl., umGani, umSane, umSenge-wesane, um-

Senge-mzani, umSingomzane, umSingo, umZane

Tecomaria capensis (Thunb.) Spach, iCakatha, iCakata, iDywadi, umKoto

Trema orientalis (L.) Blume, uBatima, umBengele, Pak, umVangazi,

umVumvu

Tricalysia

capensis (Meisn.) Sim, iNdulwane

lanceolata (Sond.) Burn Davy, umBonyane, Dhlesa, umDlesa, umHleza,

isiNyana, umPonyane, umPonyone

Trichilia dregeana Sond. , umHlakele, umKhuhlu, umKuhlu, umKuhlwa

Trichocladus

crinitus (Thunb.) Pers., iTambo, iThambo

ellipticus Eckl. & Zeyh. ex Walp., uGqonci, umVa wenyathi, umNqundu-

wenyati

Trimeria grandifolia (Hochst.) Warb., iliDhlebe-lendhlovu, iDlebe-

lendlovu, inDlebe-yendlovu, inDlebezendlovu, iNdlebe-yendlovu,

iNdlebezendlovu, iGqabela, uGqabela, iGqabi, uGqabile, Nqa-

bane, umNqabane, iTabatane, iQabela

Turraea floribunda Hochst., umaDlozana, umHIatholana, umLahlana,

umVuma

Umtiza listeriana Sim, umThiza

Urera cameroonensis Wedd., umBabazane

Uvaria caffra E. Mey. ex Sond., iDwaba

Vangueria infausta Burch., umVilo, umViluka

Vepris undulata (Thunb.) Verdoorn & C.A. Sm., umDlebe, umNgama-

zwele, umNqumaswile, umSane, umZani, umZane

Vitellariopsis marginata (N.E. Br.) Aubrev., umTunzi

Voacanga thouarsii Roem. & Schult., umHlambamaas, umHIambamasi,

umThofi, umTomvi, umThondisa, umThundisa, umTundisa

Xymalos monospora (Harv) Baill., umBokoboko, iGxwanci, iKomanci,

iKomanzi, uVete, uVethe, uVeti, umVeti, uVeto

Zanthoxylum

capense (Thunb.) Harv., umLungumabele, umNungumabele. um-

Nungwane

davyi (Verdoorn) Waterm., umLungu mabele, umNungu mabele, um-

Nungwane

Ziziphus mucronata Willd., umPafa, umPhafa

Bothalia 20,2: 153-157 (1990)

Nomenclature in Mesembryanthema (Aizoaceae): the generic names

by Rappa and Camarrone

H.E.K. HARTMANN* and V. BITTRICH*

Keywords: Aizoaceae, Mesembryanthema, nectaries, Pentacoilanthus, Tetracoilanthus

ABSTRACT

A re-examination of the seven generic names used by Rappa and Camarrone reveals that three of them were validly published

(Pentacoilanthus, Tetracoilanthus, Pteropentacoilanthus)', the first one is lectotypified here. The other four names are invalidly

published. Their synonymy is given nevertheless where possible.

UITTREKSEL

'n Herondersoek van die sewe genusname wat Rappa en Camarrone gebruik het, toon dat drie van hulle geldig gepubliseer

is ( Pentacoilanthus , Tetracoilanthus, Pteropentacoilanthus ); die eerste een word hier gelektotipifiseer. Die ander vier name

is ongeldig gepubliseer. Hul sinonimie word nietemin aangegee waar moontlik.

INTRODUCTION

In a survey of all genera of the Aizoaceae (including the

Mesembryanthema) (H.E.K.H.) it was found that the

genera described by Rappa and Camarrone (1953, 1955,

1960) have never been examined critically, and their

validity and synonymy have never been established. Based

on recent investigations in the group involved (V.B.), it

is now possible to settle the nomenclatural uncertainties.

Early investigations of flowers of Mesembryanthema led

Rappa (1912) to the description and distinction of two

different types of nectaries: lophomorphic ones, which are

crest-shapkl (Figure 1A,B), and coilomorphic ones, which

are conchiform or shell-shaped (Figure 2A, B). Nectaries

may also be absent, and consequently Rappa (1912) names

three groups: Lofomorfi, Anettari, and Coilomorfi, which

were later classified as subfamilies (Rappa & Camarrone

1953, 1960). They lack Latin descriptions, however, and

therefore do not conform with article 36 of the Interna-

tional Code of Botanical Nomenclature (Greuter et al.

1988, abbreviated ICBN subsequently). The names are

therefore not validly published.

The fundamental distinction between lophomorphic

and coilomorphic types of nectaries has been confirmed

repeatedly (e.g. Ihlenfeldt 1960). Bittrich (1987) stresses

the fact that the subfamily Mesembryanthemoideae

(— Aptenioideae Schwant. ex Bittrich & H. Hartm.) is

characterized by coilomorphic nectaries, and Bittrich &

Hartmann (1988) mention them as a synapomorphic

character for the subfamily. It is also worth noting that

in Aptenioideae the number of carpels agrees with the

number of perianth lobes as well as with the number of

nectaries. N.E. Brown (1925) already used this character

set in his keys to various genera. It must be remembered,

though, that neither the number of carpels nor that of the

nectaries is always constant within populations (e.g.

Phyllobolus subg. Prenia , Bittrich 1987). Rappa &

Camarrone (1953, 1955, 1960) use the number and shapes

* Institut fur Allgemeine Botanik, Ohnhorststr. 18, D-2000 Hamburg

52, Federal Republic of Germany.

MS. received: 1989.08.24.

of nectaries for further subdivision of the subfamilies,

but only in the Aptenioideae (Coilomorphioideae, Rappa

FIGURE 1 .—Leipoldtia weigangiana (Dinter) Dinter & Schwant. ex

H. Hartm. & Rust. (Ruschioideae) Hartmann 8476 (HBG). A.

view of upper ovary surface, white scale bar = 1 mm; B, enlarged

section of lophomorphic holonectary, the amorphous particles

on the surface are remains of nectar, white scale bar = 0,1 mm.

a, androecium, cut off; c, calyx, cut off; g, gynoecium raised

subapically, with deep fissures between lobes; n, lophomorphic

nectary which forms a complete ring (holonectary); s, styles, cut

off. SEM micrograph: H. Golling.

154

Bothalia 20,2 (1990)

FIGURE 2. — A, Brownanthus kuntzei (Schinz) Ihlenfeldt & Bittrich

(Aptenioideae) Ihlenfeldt & Warda 12158 (HBG), view of

ovary surface with five separate coilomorphic nectaries. B,

Aridaria umbelliflora (Jacq.) Schwant., Jurgens 22261 (HBG),

closer view of a single coilomorphic nectary, a, androecium,

cut off; b, prominent bladder cells; c, calyx, cut off; cn, coilo-

morphic nectary; g, gynoecium; o, ovary; s, styles, cut off. SEM

micrograph: H. Golling. White scale bar = 1 mm.

& Camarrone 1960) do they draw nomenclatural conse-

quences, i.e. name genera.

The guiding principle of Rappa & Camarrone (1953) is

primarily the number of nectaries, complemented later

(1955, 1960) by the absence or presence of valve wings.

In the first step, two genera based on five, respectively

four, nectaries are distinguished: Pentacoilanthus and

Tetracoilanthus (Rappa & Camarrone 1953). Four more

genera are added later after the importance of valve wings

is recognized: Perapentacoilanthus (Rappa & Camarrone

1955), Pteropentacoilanthus, Pterotetracoilanthus , and

Peratetracoilanthus (Rappa & Camarrone 1960). The

names indicate that a certain pattern is followed predicting

particular character combinations for the genera. This

approach can be traced from the recognition of informal

groupings (Rappa & Camarrone 1955) to the formal

descriptions of taxa (Rappa & Camarrone 1960). The

mechanical and even predictive procedure followed is well

illustrated by the description of Pterotetracoilanthus , with

a character combination which is theoretically possible

but which the authors failed to find in any real plant (see

6. below). This unconventional approach is certainly one

reason why the descriptions of Rappa & Camarrone have

been neglected. Another reason is that, in most cases, the

authors cite several species for their genera taken from

various other genera previously described. Also, the new

genera were not typified, making identification extremely

difficult. Nevertheless, none of these shortcomings alone

invalidate the names as such, and a detailed re-investigation

was carried out to establish a reliable taxonomy for future

use. Since each genus poses its own problems, the genera

are discussed separately in chronological order.

DISCUSSION

1. Pentacoilanthus Rappa & Camarrone (1953)

The description conforms with the relevant requirements

(art. 32—36 ICBN), and the name is therefore validly

published. Four species are included, Mesembryanthemum

aitonis, M. granulicaule, M. splendens and M. crystalli-

num. Unfortunately no type species is mentioned and a

lectotype must therefore be chosen.

(1) Mesembryanthemum aitonis Jacq. can be excluded

because, in contrast to the genus description of Rappa &

Camarrone (1953) the nectaries of this species are not

shell-shaped but tubular; these tubes may extend as dee-

ply into the receptacle as the locules (Figure 3). It seems

unlikely that Rappa & Camarrone studied longitudinal sec-

tions and it is therefore assumed that they were unaware

of the different nectary morphology of this species. Sur-

face views do not permit the assessment of the depth of

the nectary (Figure 2A, B). M. aitonis is therefore not

available for selection as a lectotype. At present, the spe-

cies is included in Mesembryanthemum L. subg. Mesem-

bryanthemum (Bittrich 1987).

FIGURE 3. — Mesembryanthemum aitonis Jacq. (Aizoaceae) Ihlenfeldt

& Bittrich 13857 (HBG). Longitudinal section of flower. CA,

prominent dorsal appendage of CL, calyx lobe; N, one of the

five very deep coilomorphic nectaries.

Bothalia 20.2 (1990)

155

The remaining three species do possess shell-shaped

nectaries, and the choice between them is not easy.

(2) Mesembryanthemum granulicaule Haw. (belonging to

the genus Psilocaulon N.E. Br.) is a doubtful species. It

is often synonymized with Psilocaulon articulation

(Thunb.) N.E. Br. (the type species of the genus

Psilocaulon) because of the rough papillose surface of

leaves and stem; the latter species has white to pink

flowers. The drawing of M. granulicaule, however, which

is kept at Kew and is assumed to have been seen by

Haworth (it would have to be chosen as the lectotype of

the species in a revision), shows yellow flowers. The very

few herbarium specimens with yellow flowers were all

collected in a small area in the Orange Free State. They

may well represent the true M. granulicaule as studied

by Haworth. It seems extremely unlikely, though, that

Rappa & Camarrone have investigated living or pickled

material of this very rare species and herbarium material

is useless for the investigation of nectaries. Furthermore,

in species of Psilocaulon, four- and five-locular ovaries

can occur within one species or even on one plant.

Species of Psilocaulon can therefore not be associated

unambiguously with either Pentacoilanthus or Tetracoilan-

thus. Considering the care with which Rappa and

Camarrone led their investigations, it seems highly

unlikely that they saw species, or even specimens, with

both 4- and 5-lobed ovaries.

(3) Mesembryanthemum splendens |now included in

Phyllobolus subg. Aridaria (N.E. Br. ) Bittrich, Bittrich

(1987)| is a doubtful species because its identity is very

difficult to determine. It belongs to a group of about twenty

described species in the subgenus Aridaria most of which

are synonymous. Due to the insufficient knowledge of the

group, however, the exact number and delimitation of the

species is still uncertain. It is scarcely possible to identify

any species of this group with the help of literature only.

The most useful characters are form and sculpture of seeds,

rarely mentioned in descriptions. It is unlikely therefore,

that Rappa & Camarrone investigated the true M.

splendens , particularly since it is very rare in cultivation.

Material kept under this name often belongs to the closely

allied M. umbeUijlorus Jacq. It would seem highly unwise

to choose this species as the lectotype.

(4) Mesembryanthemum erystullinum L. is the only species

figured by Rappa & Camarrone (1953). This species has

not only been well known in Europe for a long time, it

is also naturalized around the Mediterranean and is

sometimes used as a vegetable. The figure mentioned

above shows with great certainty a flower of M. crystal-

linum, because the extremely large bladder cells of the

epidermis of the receptacle are highly characteristic of this

species. Therefore, and in contrast to the two other species

with shell-shaped nectaries discussed above, it is most

probable that M. crystallinum is the plant studied by Rappa

& Camarrone. Consequently, M. crystallinum is chosen

as the lectotype of the genus Pentacoilanthus. At the same

time, M. crystallinum is the type species of the genus

Cryophytum (N.E. Brown 1926). By this lectotypification,

the name Pentacoilanthus becomes superfluous and is

illegitimate (art. 63.1 ICBN). It should be stressed, though,

that the choice was not influenced by this consequence

but is based on agreement of characters with the descrip-

tion and the probable identity of the original material.

It should be mentioned further that the name Pentacoilan-

thus would in any case be a synonym of another generic

name, since all relevant species had been described earlier.

The name Pentacoilanthus crystallinum , is not validly

published because the basionym was not cited clearly and

fully as required (art. 33.2 ICBN). At present, the species

is considered to belong to Mesembryanthemum L. subg.

Cryophytum (N.E. Br.) Bittrich (Bittrich 1987).

2. Tetracoilanthus Rappa & Camarrone (1953)

The description conforms with the relevant requirements

(art. 32—36 ICBN), and the name is therefore validly

published. Since only one species is included, this is

accepted as the indication of the type (art. 37.2 ICBN).

Tetracoilanthus is based on the same type as the genus

Aptenia N.E. Br. (1925), namely A. cordifolia (L.)

Schwant. The name is therefore superfluous and conse-

quently illegitimate (art. 63.1 ICBN).

The combination T. cordifolius (L. f.) Rappa & Camar-

rone is not validly published, because the basionym is not

indicated clearly and fully (art. 33.2 ICBN). At present,

the species is included in the genus Aptenia N.E. Br.

(Bittrich 1987).

3. Perapentacoilanthus Rappa & Camarrone (1955)

The description of this genus is almost identical with

that of Pentacoilanthus Rappa & Camarrone 1953 (see 1.

above) except for the addition of ‘valvis... quae, septis

deficientibus, circa nudum axem consistunt’. This

statement probably refers to the columella, the formation

of which is very variable and changes from low to high

with all intermediates. In Aptenioideae, the septa are

always transformed into expanding keels in their upper

parts, and consequently a remaining collumella will appear

naked in the open capsule. These inconsistencies detract

from the taxonomic value of the character. The two

descriptions are consequently considered to be identical.

Although Rappa and Camarrone did not state it

explicitly, there is no doubt that they coined the name

Perapentacoilanthus to replace Pentacoilanthus Rappa

& Camarrone (1953). They had realised in the mean-

time that the absence or presence of valve wings is an

important character. Consequently they were now planning

to group species and name genera not only according

to the number of nectaries, but also according to the

absence or presence of valve wings, as described in the

introduction above. This is borne out by the following:

a, under the heading ' Pentacoilanthus . Genere privo di

umenoprosteci’ (genus deprived of valve wings), Rappa

& Camarrone (1955) indicate that they intend to alter the

diagnostic characters of Pentacoilanthus, which indeed

they did later (Rappa & Camarrone 1960: 13); b, in the

1960 publication they include all the species originally

mentioned under Pentacoilanthus Rappa & Camarrone

(1953) under Perapentacoilanthus. This action was

obviously initiated with the aim to name the genera

with five, respectively four, coilomorphic nectaries in

parallel fashion ( Pentacoilanthus matching Tetracoilanthus

without valve wings; Pteropentacoilanthus—Pterotetra-

coilanthus with free valve wings; Perapentacoilanthus—

156

Peratetracoilanthus with valve wings forming pockets;

compare list in Rappa & Camarrone 1955: 10).

To declare Perapentacoilanthus a superfluous and

therefore illegitimate name (art. 63.1 ICBN) requires the

definite inclusion of the holotype of the earlier synonym

Pentacoilanthus. This is not possible literally, since Pen-

tacoilanthus 1953 was published without the indication of

a type, but the inclusion of all four species of the original

Pentacoilanthus 1953 under Perapentacoilanthus (Rappa

& Camarrone 1960) makes it clear that the type would

be included, whichever species had been chosen. As a

consequence, Pentacoilanthus 1953 and Perapentacoilan-

thus 1955 must be considered synonymous, being names

for the same genus.

4. Pentacoilanthus Rappa & Camarrone (1960)

As stated above, the characterization of the genus differs

from the Pentacoilanthus of 1953 in an important

diagnostic character, namely the absence of valve wings,

and an entirely different set of species is accordingly

assigned to the genus. This name therefore represents a

later homonym of Pentacoilanthus Rappa & Camarrone

1953 (although no type is mentioned in either genus) and

is consequently illegitimate (art. 64.1 ICBN). Further-

more, the lack of citation of a type at this date makes the

name an invalidly published one (art. 37.1 ICBN ).

All species included in Pentacoilanthus 1960 were taken

from the genus Sceletium N.E. Br., and they are at present

assigned to Phyllobolus N.E. Br. subg. Sceletium (N.E.

Br.) Bittrich (Bittrich 1987).

5. Pteropentacoilanthus Rappa & Camarrone (1960)

Only one species is cited as an example with the

description of the genus and this validates the name (art.

37.2 ICBN). The genus is based on the same species as

Halenbergia Dinter, namely H. hypertrophica (Dinter)

Dinter. The name Pteropentacoilanthus is therefore a

superfluous name and illegitimate (art. 63.1 ICBN).

The combination P hypertrophicum (Dinter) Rappa &

Camarrone is neither validly published, because the

basionym is not cited fully (art. 33.2 ICBN), nor

legitimate, since it includes the type of an earlier named

taxon (art. 63.1 ICBN).

At present, the species is considered to belong to

Mesembryanthemum L. subg. Opophytum (N.E. Br.)

Bittrich (Bittrich 1987).

6. Pterotetracoilanthus Rappa & Camarrone (1960)

No type is named with the description, and the name

is therefore invalid (art. 37.1 ICBN); no species are

mentioned at all.

7. Peratetracoilanthus Rappa & Camarrone (1960)

Eight species are cited with the description, but no type

is chosen, and the name is therefore invalidly published

(art. 37.1 ICBN). The new combinations are invalid as well

because no basionyms are given (art. 33.2 ICBN), and

Bothalia 20,2 (1990)

the name of the genus is invalidly published (art. 43.1

ICBN).

TAXONOMY

1. Pentacoilanthus Rappa & Camarrone in Lavori

dell’Istituto Botanico e Giardino Coloniale Palermo 14:

32 (1953), nom. illeg. Lectotype species: Mesembryanthe-

mum crystallinum L.: 480 (1753) (here designated). Type:

Dillenius: 231, t. 211 (1732).

Cryophytum N.E. Br.: 412 (1925). Mesembryanthemum L. subg.

Cryophytum (N.E. Br.) Bittrich: 72 (1987). Type: C. crystallinum (L.)

Schwant., fide N.E. Brown: 245 (1926).

Perapentacoilanthus Rappa & Camarrone 1960, nom. illeg., nom.

invalid (see 3 below).

2. Tetracoilanthus Rappa & Camarrone in Lavori

dell’Istituto Botanico e Giardino Coloniale Palermo

14: 34 (1953), nom. illeg. Type: Mesembryanthemum

cordifolium L. f. : 260 (1782). Holotype: Thunberg s.n.

1773 in Herb. Montin (S!) [= Aptenia cordifolia (L. f.)

Schwant.: 69 (1928)].

Note on the holotype: there is no doubt that the sheet cited

and seen is the holotype since the description on the back

refers to the original publication. The specimen agrees well

with the description. The holotype was already recognized

by both Norlindh and Glen in 1976, but there seems to be

no published reference to this. In any case, the existence

of the holotype supersedes the selection of a neotype by

Preston & Sell (1988).

Aptenia N.E. Br.: 412 (1925).

Litocarpus L. Bol.: t. 261 (1927), nom. illeg.

3. Perapentacoilanthus Rappa & Camarrone in Lavori

dell’Istituto Botanico e Giardino Coloniale Palermo 15:

6 (1955), nom. illeg., nom. invalid. No type given (see

1 above).

4. Pentacoilanthus Rappa & Camarrone in Lavori

dell’Istituto Botanico e Giardino Coloniale Palermo 18:

13 (1960), nom. illeg., nom. invalid. No type given (see

1 above).

Sceletium N.E. Br. : 412 (1925). Phyllobolus N.E. Br. subg. Sceletium

(N.E. Br.) Bittrich: 75 (1987).

5. Pteropentacoilanthus Rappa & Camarrone in Lavori

dell’Istituto Botanico e Giardino Coloniale Palermo 18:

14 (1960), nom. illeg. Type species: Mesembryanthemum

hypertrophicum Dinter [= Halenbergia hypertrophica

(Dinter) Dinter: 200 (1937)]. Type: Dinter 3875 (B!).

Halenbergia Dinter: 200 (1937).

Opophytum N.E. Br.: 412 (1925). Mesembryanthemum L. subg.

Opophytum (N.E. Br.) Bittrich: 73 (1987).

6. Pterotetracoilanthus Rappa & Camarrone in Lavori

dell’Istituto Botanico e Giardino Coloniale Palermo

18: 14 (1960), nom. invalid. No type given. No species

mentioned.

7. Peratetracoilanthus Rappa & Camarrone in Lavori

dell’Istituto Botanico e Giardino Coloniale Palermo 18:

14—15 (1960), nom. invalid. No type given.

Bothalia 20,2 (1990)

157

ACKNOWLEDGEMENTS

The basic field work on Mesembryanthema was

supported by the German Science foundation and the

National Botanic Gardens of South Africa; to both we are

most grateful. We would like to thank H. Golling for help

with SEM micrographs, and Prof. P. Hiepko, Berlin, for

critically reading the manuscript.

REFERENCES

BITTRICH, V. 1987. Untersuchungen zu Merkmalsbestand, Gliederung

und Abgrenzung der Unterfamilie Mesembryanthemoideae

(Mesembryanthemaceae Fenzl). Mitteilungen des lnstituts fur

Allgemeine Botanik Hamburg 21: 5-116.

BITTRJCH, V, & HARTMANN, H E.K. 1988. The Aizoaceae — a new

approach. Botanical Journal of the Linnean Society 97: 239 - 254.

BOLUS, L. 1927. Litocarpus cordifolius. The Flowering Plants of South

Africa 7: t. 261.

BROWN, N.E. 1925. Aptenia, Cryophxtum. Opophytum, Sceletium.

Gardeners' Chronicle ser. 3,78: 412.

BROWN, N.E. 1926. Cryophxtum. In E.P Phillips, The genera of South

African flowering plants: 245. Cape Town.

DILLENIUS, J.J 1732. Hortus elthamensis: 231, t. 211. London

D1NTER, K, 1923. Succulentenforschung in Sudwcstafrika Feddes

Repertorium, Beiheft 23: 63. Berlin.

DINTER, K. 1937. Halenbergia hypertrophica. Kakteen und andere

Sukkulenten: 200. Berlin.

GREUTER, W, et al. 1988. International Code of Botanical

Nomenclature. Regnum vegetabile 118. Konigstein.

IHLENFELDT, H.-D. 1960. Entwicklungsgeschichtliche, morphologische

und systematische Untersuchungen an Mesembryanthemen

Feddes Repertorium 63: 1 — 104

LINNAEUS, C. 1753. Species plantarum. Stockholm.

LINNAEUS, C. 1782. Supplementum plantarum Brunswick.

PHILLIPS, E P 1926. The genera of South African flowering plants.

1st edn. Pretoria.

PRESTON, C D. & SELL, P.D. 1988. The Aizoaceae naturalized in the

British Isles. Watsonia 17: 221.

RAPPA, F. 1912. Per una classificazione naturaledei Mesembrianthemi.

Bulletin Reale Orto Botanico Giardino Colonial e Palermo II:

21-36.

RAPPA, F. & CAMARRONE, V. 1953. Lavori dell' Istituto Botanico

e Giardino Coloniale Palermo 14: 1 — 39.

RAPPA, F. & CAMARRONE, V. 1955. Lavori dell' Istituto Botanico

e Giardino Coloniale Palermo 15: 1 — 16.

RAPPA, F. & CAMARRONE, V. I960. Lavori dell' Istituto Botanico

e Giardino Coloniale Palermo 18: 1—24.

SCHWANTES, G. 1928. Mesembriaceen unserer Kulturen in neuer

Benennung. Gartenflora 77: 69.

Bothalia 20,2: 159-165 (1990)

A checklist of the plants of the Karkloof Forest, Natal midlands

J.O. WIRMINGHAUS*

Keywords: checklist, Karkloof. forest. Natal

ABSTRACT

A checklist of the plants of the Karkloof Forest is presented. Of the 400 taxa recorded, seven are bryophytes. 69 are

pteridophytes. four are gymnosperms and 320 are angiosperms. At least 17 of the plants recorded are localized or uncommon

in Natal. Two of these are endemic to the area.

UITTREKSEL

'n Kontrolelys van plante van die Karkloofwoud word verskaf. Van die 400 taksons wat aangeteken is. is sewe briofiete,

69 pteridofiete, vier gimnosperme en 320 angiosperme. Minstens 17 van die aangetekende plante is tot sekere lokaliteite

beperk of is seldsaam in Natal. Twee hiervan is endemies in die gebied.

INTRODUCTION

Rycroft (1941) carried out the first detailed study of the

Karkloof Forest’s vegetation and provides a preliminary

checklist of the plants. Taylor (1961) gives a brief account

of the plant communities of the Karkloof Forest although

his paper is directed at its conservation. More recently

Moll (1976) included the Karkloof Forest in his vegetation

survey of the Three Rivers region. Natal. Other than the

above, little has been published on the area, despite the

fact that it is one of the largest remaining patches of

mistbelt forest in Natal.

The absence in regional herbaria of records of species

commonly seen in the area highlighted the need for a

comprehensive ch ecklist of the forest plants.

STUDY AREA AND METHODS

The Karkloof Forest lies in the Natal midlands and

covers an estimated area of 6 000 ha (Cooper 1985). It

extends in length for 40 km, from about 22 km north of

Howick (29°17’S. 30°09’E) to 13 km northeast of New

Hanover (29°14’S, 30°28’E), and comprises a number of

forest patches restricted mainly to the steep eastward side

of the Karkloof Range. The forest ranges in altitude from

800 m near the Karkloof and Umgeni River confluence

up to I 700 m on the upper slopes of Mt Gilboa.

Most of the forests in the midlands region, including

the Karkloof. fall under Acocks’s (1988) Veld Type 5

(’Ngongoni Veld). In addition to mist the forest receives

a mean annual rainfall of about 1 600 mm (1 300—2 200

mm) (Rycroft 1944; Oatley 1978; pers. obs.) falling mostly

in summer. Predominant winds in the summer months

are southeasterly but in winter, as along most of the

escarpment, northwesterly berg winds predominate.

Rycroft (1944) attributes the stunted growth of forest trees

in the northwestern Karkloof and the deaths of numerous

young forest trees each year to these latter hot, dry winds.

* Dept, of Zoology & Entomology, University of Natal. RO. Box 375,

Pietermaritzburg 3200.

MS. received: 1990.02.20.

Little is published of temperatures within the forest but

Oatley (1978) recorded some data over a two year period.

From his data the mean annual maximum temperature is

18,0°C (range 15,2— 20,9°C) measured in February, and

the mean minimum is 8,4°C (range 4,6— 12,2°C) in July.

Most of the collecting for this survey was done near

the Farms Mbona and Chard which form part of the

Blinkwater forest patch. Other sites in the Karkloof were

also visited during the survey for the purpose of

completeness. All specimens are lodged at NU. In addition

to my specimens, any records of plants from other

collectors, which were found in NU, are included. Of

these, E.J. Moll has possibly contributed the most. Much

of his collecting was carried out in the forest near the

Farms The Start. Benvie and Ehlatini. Even though every

effort was made to make the checklist as comprehensive

as possible, there will be a number of species that have

been overlooked. Bryophytes were only occasionally

collected during this survey, and field observations indicate

that many more taxa occur in the forest than this checklist

suggests.

RESULTS AND DISCUSSION

Rycroft (1941) recorded 246 species in his preliminary

checklist of the forest plants but did not mention any

voucher specimens. He lists 45 species of pterido-

phytes, four gymnosperms, 25 monocotyledons and 172

dicotyledons.

Of the 400 taxa (266 genera, 115 families) recorded

in the Karkloof Forest during this survey, seven (1,8%)

are bryophytes, 69 (17,3%) are pteridophytes, four (1.0%)

are gymnosperms and 320 (80,0%) are angiosperms.

Of the angiosperms, 67 (16,8%) are monocotyledons and

253 (63,3%) are dicotyledons. Plant families containing

more than one percent of the total number of species are

listed in order of numerical importance in Table 1. The

genera of plants with four or more (> 1%) species are

Asplenium (11 spp.), Senecio, Solatium (7 spp.). Blech-

num, Cheilanthes, Plectranthus , Streptocarpus (5 spp.),

Canthium, Lycopodium, Maytenus, Protasparagus and

160

Bothalia 20,2 (1990)

TABLE 1. — Synopsis of the plant families of the Karkloof Forest with

four or more species (> 1%), listed in order of numerical

importance together with the number of genera in each family

Pteris (4 spp.). No comparative data are available tor other

mistbelt forests in Natal, but Acocks (1988) also notes the

dominance of ferns and other genera such as Plectranthus

in the forests of this region.

Although a crude attempt was made to include species

habitat preferences on the checklist, no indication of status

is shown [some quantitative data are available but will

appear elsewhere (Wirminghaus & Perrin in prep.)).

At least 17 of the species recorded from the forest were

found to be uncommon or to have localized distributions

within Natal. The ferns Arachnipdes foliosa, Asplenium

dregeanum, Asplenium preussii, Blechnum capense,

Elaphoglossum aubertii, Loxogramme lanceolata and

Trichomanes reptans are regarded as rare in Natal

(Jacobsen 1983) and are known from only one or two sites

in the forest. Other plants such as Hilliardia zuurbergen-

sis, Lobelia malowensis, Microstegiwn nudum, Mikaniop-

sis cissampelina, Plectranthus elegantulus ,

Polygonum nepalense and the trees Andrachne ovalis and

Suregada procera , are only known from a few localities

within Natal. The herbs Geranium natalense and

Plectranthus rehmannii are the only plants known to be

endemic to the Karkloof Forest (Codd 1985; Hilliard &

Burtt 1985).

ACKNOWLEDGEMENTS

The staff of the NU herbarium are thanked for helping

to identify much of the material listed here. I would also

like to thank Colleen Downs for her assistance in the field,

and Trevor Edwards for his comments on an earlier draft

of this paper.

REFERENCES

ACOCKS. J.P. H. 1988. Veld types of South Africa, 3rd edn. Memoirs

of the Botanical Survey of South Africa No. 57: 1—146.

CODD, L.E. 1985. Lamiaceae. In O.A. Leistner, Flora of southern Africa

28, part 4: 1-247.

COOPER, K.H. 1985. The conservation status of indigenous forests in

Transvaal, Natal and O.F.S., South Africa. Wildlife Society of

S.A., Durban.

GIBBS RUSSELL, G.E., REID, C., VAN ROOY, J. & SMOOK, L.

1985. List of species of South African plants. Part I. Memoirs

of the Botanical Survey of South Africa No. 51: 1—152.

GIBBS RUSSELL, G.E., WELMAN, W.G., RETIEF, E., IMMEL-

MAN, K.L., GERM1SHUIZEN, G., PIENAAR, B.J., VAN

WYK, M. & NICHOLAS, A. 1987. List of species of South

African plants. Part 2. Memoirs of the Botanical Survey of South

Africa No. 56: 1-270.

HILLIARD, O.M. & BURTT. B.L. 1985. A revision at Geranium in

Africa south of the Limpopo. Notes from the Royal Botanical

Garden. Edinburgh 42: 171—225.

JACOBSEN, W. B.G 1983. The ferns and Jem allies of southern Africa.

Butterworths, Durban.

LINDER, H P. 1989. Notes on southern African Angraecoid orchids.

Kew Bulletin 44: 317—319.

MOLL, E.J. 1976. The vegetation of the three Rivers region. Natal. Natal

Town and Regional Planning Commission, Pietermaritzburg.

OATLEY, T.B. 1978. The breeding biology and ecology of the slurred

robin Ponogocichla slellala. M.Sc. thesis. University of Natal,

Pietermaritzburg.

RYCROFT, H.B. 1941. The plant ecology of the Karkloof Forest, Natal.

M.Sc. thesis. Natal University College, Pietermaritzburg.

RYCROFT, H.B 1944 The Karkloof Forest, Natal. Journal of the South

African Forestry Association II: 14—25.

TAYLOR, H.C. 1961. The Karkloof Forest: a plea for its protection.

Forestry in South Africa I: 123 — 134.

WIRMINGHAUS, J.O. & PERRIN, M R. In prep. The plant ecology

of a portion of the Karkloof Forest, Natal.

CHECKLIST

The checklist is arranged according to Gibbs Russell el at. (1985) and Gibbs Russell et a!. (1987), and one orchid according to Linder (1989).

All voucher specimens cited are specimens collected by J.O. Wirminghaus and housed in NU unless otherwise stated or unless a literature citation

is given Introduced and alien species are indicated by an asterisk In order to make the list more meaningful, habitat annotations have been added

for each species recorded. These are derived either from herbarium specimen labels or from personal field observations and are as follows: I.

forest margins; 2, forest gaps or small clearings; 3, forest floor; 4, forest understorey; 5, forest canopy; 6, next to streams in forest.

BRYOPHYTA

RICCIACEAE

Riccia fluitans L. , 74!, 6

M ARCH ANTI ACEAE

Dumortiera hirsuta (Sw.) Nees, 655, 6

PALLAVICINI ACEAE

Symphyogyna podophylla (Thunb.) Nees i£ Mont., 855, 6

BRYACEAE

Bryum argenteum Hedw. , S.E. Wood 24, 3

Rhodohryum umbraculum {Hook.) Schimp. ex Par., 641, 3

Bothalia 20,2 (1990)

161

PTEROBRYACEAE

Calyptothecium hoehnellii (C. Muell.) Argent., Sim 585 , 3

NECKERACEAE

Porothamnium natalense (C. Muell.) Fleisch., Moll 2867, 3

PTERIDOPHYTA

LYCOPODIACEAE

Lycopodium

cernuum L. , 879, 1

clavatum L. , F. Buyer s.n., 1

gnidioides L. f, 801, 6

verticillatum L. {., Fisher 516, 6

SELAGINELLACEAE

Selaginella

kraussiana (Kunze) A. Braun, 630, 3, 6

mittenii Bak. , 654, 3

OPHIOGLOSSACEAE

Ophioglossum polyphyllum A. Br. ex Seub. , 903, 3

MARATTIACEAE

Marattia fraxinea J.E. Sin. ex J.F. Gmel. var. salicifolia (Schrad.) C.

Chr. , 609, 6

SCHIZAEACEAE

Anemia dregeana Kunze, Hilliard 2575, 3

Mohria caffrorum (L.) Desv. , 805, 1

GLEICHEN1ACEAE

Gleichenia umbraculifera (Kunze) T. Moore, 808. I

HYMENOPHYLLACEAE

Hymenophyllum tunbridgense (L. ) J.E. Sin., 1166, 6

Trichomanes

borbonicum V. d. Bosch, 800, 6

pyxidiferum L. var. melanotrichum (Schlechtd.) Schelpe, 799, 4, 6

reptans Swartz, 798, 6

CYATHEACEAE

Cyathea

capenxix ( L . f.) J.E. Sin., 770, 6

dregei Kunze. Ripley 57, 1

DENNSTAEDT1ACEAE

Blotiella glabra (Bory) A.F. Tryon, 1089. 4

Hixtiopteris incixa (Thunb.) J. Sin., 874. 1

Hypolepix xparxixora (Schrad.) Kuhn, 608, 2, 3, 6

Pteridium aquilinum (L.) Kuhn, Graham 83. 1, 2

VITTARIACEAE

Viltaria isoetifolia Bory, O. Seek' s.n., 4

AD1ANTACEAE

Adiantum

eapillux-venerix L , 856, 6

poiretii Wikstr. var. poiretii, 1151, 6

raddianum Pres I, 809, 6

Cheilanthex

bergiana Sclileclud. ex Kunze, 786, 3, 6

concolor ( Lungsd . & Fisch.) Schelpe c6 N.C. Anthony, 778, 3

eckloniana (Kunze) Mett. , 1152, 2

quadripinnata (Forssk.) Kuhn, Rycroft 548, 1

viridix (Forssk.) Swartz var. macrophylla (Kunze) Schelpe & N.C.

Anthony, 605, 3

Pellaea calomelanox (Swartz) Link, 1071, I

Pteris

buchananii Bak. ex Sim, Ripley 53, 3

catoptera Kunze, 611, 3

cretica L. , 973, 6

dentata Forssk. , Stanton 89, 3

POLYPOD1ACEAE

Loxogramme lanceolata (Swartz) Presl, 780, 4

Pleopeltis

macrocarpa (Bory ex Willd.) Kaulf. , 779, 4

schraderi (Mett.) Tardieu, 728, 4

Polypodium polypodioides (L. ) Hitchc. subsp. ecklonii (Kunze) Schelpe,

739, 4

ASPLENIACEAE

Asplenium

aethiopicum (Burm. f. ) Becherer, 776, 3, 6

boltonii Hook, ex Schelpe, 904, 6

dregeanum Kunze, 1082, 6

erectum Bory ex Willd. var. erectum. 769, 3, 6

lobatum Pappe & Rawson, 1033, 3

lunulatum Swartz, 1064, 3

monanthes L. , 863, 6

preussii Hieron. , 865, 6

rutifolium (Berg.) Kunze, 607, 4

splendens Kunze, 627, 3, 6

theciferum (Kunth) Mett. var. concinnum (Schrad.) Schelpe, 783, 4

THELYPTERIDACEAE

Thelypteris

dentata (Forssk.) E. St.John, 868, 6

gueinziana (Mett.) Schelpe, 867, 6

pozoi (Lag.) Morton, 748, 6

ATHYRIACEAE

Athyrium scandicinum (Willd.) Presl, 905. 6

Cystopteris fragilis (L.) Beritlt., 777. 6

Lunathyrium japonicum (Thunb.) Kurata* 784. 6

LOMARIOPSIDACEAE

Elaphoglossunr

acroxtichoidex (Hook. & Grew) Schelpe, 1140. 3

aubertii (Desv.) T. Moore, 768, 3. 6

ASPID1ACEAE

Arachniodes foliosa (C. Chr.) Schelpe, 864, 6

Ctenitis lanuginosa (Willd. ex Kaulf.) Cope!., 860, 4

Dryopteris

athamantica (Kunze) Kuntze, Graham 67, 3

inaequalis (Schlechtd.) Kuntze, 811, 3

Polystichunr

pungens (Kaulf.) Presl, 610, 3

transvaalense N.C. Anthony. 856, 6

BLECHNACEAE

Blechnum

australe L. , 1137, 3

capense Burm. f, 806, 1

giganteum (Kaulf. ) Schlechtd. , 775, 6

inflexum (Kunze) Kuhn. 880, 1

tabulare (Thunb.) Kuhn, 933, 1

GYMNOSPERMAE

ZAM1ACEAE

Encephalartos altensteinii Lehm., Rycroft (1941), 1

PODOCARPACEAE

Podocarpus

falcatus (Thunb.) R. Br. ex Mirb. , Hilliard 2026, 5

henkeli i Stapf ex Dallim. & Jacks., Sim 19014, 5

latifolius (Thunb.) R. Br. ex Mirb., Sim 19019 , 5

ANGIOSPERMAE — MONOCOTYLEDONEAE

POACEAE

Microstegium nudum (Trin.) A. Camus, 734, 3

Paspalum urvillei Steud.* 816, 6

Oplismenus hirtellus (L.) Beauv. , 725, 3

Panicum

aequinerve Nees, Moll 2872, 3

deustum Thunb., Mol! 1672, 3

Setaria

megaphylla (Steud.) Dtir. & Schinz, 750, 1, 2

sphacelata (Schumach.) Moss, 844, 1, 2, 6

Prosphytochloa prehensilis (Nees) Schweick. , 724 , 4

Agrostis lachnantha Nees, 751 , 3, 6

Pseudobromus silvaticus K. Schum. , 726, 3

162

Bothalia 20,2 (1990)

CYPERACEAE

Cyperus

albostriatus Schrad. , 861 , 3

distans L. f, 752, 3

Pycreus cimicinus (Pres!) Pfeiffer, 615, 6

Mariscus

keniensis (Kuekenth.) Hooper, 616, 6

thunbergii (Vahl.) Schrad., 823, 1, 2

Kyllinga elatior Kunth,- 738, 6

Isolepis

cernua (Vahl) Roem. & Schult. , 1097, t, 2

costata (Boeck.) A. Rich. var. macra (Boeck.) B.L. Bunt, 1099, 6

sepulcralis Steud., Moll 1076, 6

Schoenoxiphium

lehmannii (Nees) Steud. , 1060, 3

rufum Nees, 849, 1, 3

Carex

acutiformis Ehrh. , Bourquin 344, 3

mossii Ne lines, 909, 1, 2

ARACEAE

Zantedeschia aethiopica (L.) Spreng., 1118, 1, 2, 6

JUNCACEAE

Juncus

dregeanus Kunth, 1096, I, 2

effusus L. , 791, 6

lomatophyllus Spreng. . 794, 6

Luzula africana Drege ex Steud. , 1031, 1

LIL1ACEAE

Littonia modesta Hook. , 1153. 1

Chlorophytum

comosum (Thunk.) Jacq. , 1116, 3

krookianum Zahlhr. , Moll 3470, 3

Kniphofia

buchananii Bak. . Moll 3457, 2

tysonii Bak. subsp. tysonii, 795, 2

Dracaena hookeriana K. Koch, Rycroft (1941), I, 3

Protasparagus

aethiopicus (L.) Oberm., 970, 1, 2

natalensis (Bak.) Oberm., 1030, 5

setaceus (Kunth) Oberm. , 670, 3

sp., 561, 3

Myrsiphyllum asparagoides (L.) Willd., 995, I, 2

Behnia reticulata (Thunb.) Didr., 961. 4

HAEMODORACEAE

Barbcretta aurea Harr. , 1136, 6

AMARYLL1DACEAE

Scadoxus puniceus (L.) Friis & Nordal, 972, I, 6

Clivia miniata Regel, 993, 4

Nerine appendiculata Bak. , 843. I

VELLOZIACEAE

Talbotia elegans Balf, 651, 3, 6

DIOSCOREACEAE

Dioscorea

cotinifolia Kunth, 1161, I

dregeana (Kunth) Dur. & Schinz var. dregeana, 1075, 4

IRIDACEAE

Dietes

butcheriana Gerstner, Moll 3454, 6

iridioides (L.) Sweet ex Klatt, 1133, 3

Crocosmia aurea Planch., 556, 1, 2, 6

CANNACEAE

Canna indica L.*, 745, 6

ORCHIDACEAE

Stenoglottis fimbriata Lindl., 555, 4

Holothrix orthoceras (Harv. ) Reichb. f, O’Connor 349, 3

Disperis

fanniniae Harv., 554, 3

lindlcyana Reichb. f. , 668, 3

Liparis bowkeri Harv., 848, 3

Polystachya

ottoniana Reichb. f, 1050, 4, 5

pubescens Reichb. f. , 1081, 4, 5

Bulbophyllum scaberulum Rolfe, in cultivation at NU, 4

Angraecum

conchiferum Lindl., Moll 3368, 4, 5

pusillum Lindl., in cultivation at NU, 4, 5

sacciferum Lindl., in cultivation at NU, 4, 5

Tridactyle bicaudata (Lindl.) Schltr., in cultivation at NU, 4, 5

Diaphananthe caffra (H. Bol. ) Linder. 1117, 4

Mystacidium

flanaganii (H. Bol.) H. Bol., 543, 4

gracile (Reichb. f.) Harv., 1107, 1, 4

venosum Harv. ex Rolfe. 926, 4

ANGIOSPERMAE— DICOTYLEDONEAE

P1PERACEAE

Piper capense L. f, 871, 6

Peperomia

retusa (L. f.) A. Dietr. var. retusa, 916, 4

tetraphylla (G. Forst.) Hook. & Arn., 917, 4

ULMACEAE

Celtis africana Burin, f. , 1047, 5

MORACEAE

Ficus

craterostoma Warb. ex Mildbr. & Burr. . 581. 5

sur Forssk. , 969, 1

URT1CACEAE

Laportea

alatipes Hook, f, Moll 3522, 3

peduncularis (Wedd.) Chew subsp. peduncularis, 743, 6

Droguetia ambigua Wedd. , 1146, 2, 3

Didymodoxa caffra (Thunb.) Friis & Wibnot-Dear, Hilliard & Burtt

11887, I

VISCACEAE

Viscum nervosum Hochst. ex A. Rich. , 538, parasite on Monanthotaxis

caffra

SANTALACEAE

Osyridicarpos schimperianus (Hochst. ex A. Rich.) A. DC., 920, 2, 6

POLYGONACEAE

Rumex sp. cf. woodii N.E. Br. , 1080, 6

Polygonum

nepalense Meisn.*, 623, 6

salicifolium Willd. . 618, 6

AMARANTHACEAE

Achyranthes sicula (L.) All.*, 760, 6

CARYOPHYLLACEAE

Stellaria sennii Chiov* 990, 2, 6

Dry maria cordata (L.) Willd. subsp. diandra (Blume) J. Duke, 797, 3, 6

RANUNCULACEAE

Clematis brachiata Thunb., Moll 3526, 5

Ranunculus multifidus Forssk. , 1079, 6

Thalictrum rhynchocarpum Dill. & Rich. , Moll 3528, 6

MENISPERMACEAE

Cissampelos torulosa E. Mey. ex Harv. , 789, 4

ANNONACEAE

Monanthotaxis caffra (Sond.) Verde., 1093, 5

TRIMENIACEAE

Xymalos monospora (Harv.) Bail!., 629, 5

LAURACEAE

Ocotea bullata (Burch.) Baill., Moll 3478, 5

Cryptocarya

myrtifolia Stapf 583 , 4, 5

woodii Engl., 584, 4, 5

Bothalia 20,2 (1990)

163

PAPAVERACEAE

Papaver aculeatum Thunb .*, 1119, 6

BRASSICACEAE

Cardamine africana L. , 907, 6

CRASSULACEAE

Crassula

inandensis Schonl. & Bak.f, 1134, 3, 6

pellucida L. subsp. alsinoides (Hook.f) Toelken, 757, 6

sp. aff. alba Forssk. , 1174, 1

ESCALLONIACEAE

Choristylis rhamnoides Harv. , 761, 4, 6

PITTOSPORACEAE

Pittosporum viridiflorum Sims, Moll 3544, 1, 5

ROSACEAE

Rubus

cuneifolius Pursh*, 1073, 1, 2

rosifolius J.E. Sm* 1058, 1, 2

Duchesnea indica (Andr.) Focke*, 746, 6

Alchemilla kiwuensis Engl., 788, 6

Leucosidea sericea Eckl. & Zeyh. , 977, 1

Cliffortia

linearifol ia Eckl. & Zeyh., 1054, 1

nitidula (Engl.) R E. & Th. Fries Jr, 982 , 6

Prunus africana (Hook. f. ) Kalkm. , 1158, 5

CONNARACEAE

Cnestis natalensis (Hochst.) Planch. & Sond.. 1001, 5

FABACEAE

Acacia

ataxacantha DC. , Getliffe 219, 1

mearnsii De Wild 3, 981, I, 2, 6

melanoxylon R. Br*, Ross 2088, 1, 2

Calpurnia aurea (Ait.) Benth., 1040, 2, 5, 6

Crotalaria capensis Jacq., 983, 6

Argyrolobium tomentosum (Andr.) Druce, 821, 1, 2, 3, 6

Indigofera natalensis H Bol., 922, 4

Psoralea pinnata L. , 925, 1

Desmodium repandum (Vahl) DC., 612, I, 2, 6

Dalbergia obovata E. Mey. , 1041, 5

Dumasia villosa DC. var. villosa, 754, 2, 6

GERANIACEAE

Geranium

flanaganii Knuth, Hilliard & Burn 13485, 2

natalense Hilliard & Burn, 1051, 1

schlechteri Knuth, Hilliard & Burn 13489, 1, 2

OXALIDACEAE

Oxalis

corniculata L.*, 758, 6

obliquifolia Steud. ex Rich., 1052, 1

RUTACEAE

Zanthoxylum davyi (Verdoorn) Waterm. , 1002, 5

Calodendrum capense (L. f.) Thunb., 997, 5

Oricia bachmannii (Engl.) Verdoorn, 1090, 4

Vepris lanceolata (Lam.) G. Don, 1078, 5

Clausena anisata ( Willd .) Hook. f. ex Benth., 1087, 4, 5

PTAEROXYLACEAE

Ptaeroxylon obliquum (Thunb.) Radik., 979, 5

MELIACEAE

Ekebergia capensis Sparrm. , 1027, 5

POLYGALACEAE

Polygala confusa MacOwan, 1024, 1, 2 , 6

EUPHORBIACEAE

Andrachne ovalis (Sond.) Muell. Arg. , Bayer 687, 4

Drypetes

gerrardii Hutch., Bayer 1414, 1, 4

natalensis (Harv.) Hutch., 1076, 4

Micrococca capensis (Bail!. I Pram, 593, 4, 6

Clutia

katharinae Pax, 1039, 1

pulchella L. var. franksiae Prain, 803, 2, 4

Suregada procera (Prain) Croiz. , Smook 655, 4

Excoecaria simii (Kuntze) Pax, Moll 3535, 4

Euphorbia

dumosa E. Mey. ex Boiss., 1053, 1, 2

epicyparissias E. Mey. ex Boiss., Hilliard & Burn 10141, I, 2

kraussiana Bernh. var. kraussiana, 626, 1, 2, 6

CALL1TRICHACEAE

Callitriche sp. cf. compressa N.E. Br. , 1049, 6

ANACARDIACEAE

Harpephyllum caffrum Bernh. ex Krauss, 1148, 5

Protorhus longifolia (Bernh.) Engl., 964, 1

Rhus

chirindensis Bak. f, 1069, 5

dentata Thunb. , 1056, 1

pyroides Burch, var. gracilis (Engl.) Burtt Davy, 1059, 1

AQUIFOL1ACEAE

Ilex mitis (L.) Radik, var. mitis, 1067, 5, 6

CELASTRACEAE

Maytenus

acuminata (L. f.) Loes., 986, 4

heterophylla (Eckl. & Zeyh.) N.K.B. Robson, 1028, 1

mossambicensis (Klotzsch) Blakelock var. mossambicensis, 560, 4

undata (Thunb.) Blakelock, 1029, 1

Cassine

papillosa (Hochst.) Kuntze. 1004, 4

tetragona (L. f. ) Loes. , 590, 5, 6

ICACINACEAE

Apodytes dimidiata E. Mey. ex Arn. subsp. dimidiata, 591, 5

SAPINDACEAE

Allophylus

dregeanus (Sond.) De Winter. 585, 2, 4

melanocarpus (Sond.) Radik., 872, 5

MELIANTHACEAE

Bersama tysoniana Oliv., 1139, 1

GREY1ACEAE

Greyia sutherlandii Hook. & Harv, 1036, 1

BALSAM IN ACEAE

Impatiens hochstetteri Warb. subsp. hochstetteri, 540, 3

RHAMNACEAE

Scutia myrtina (Burm.f.) Kurz, 1066. 5

Rhamnus prinoides L'Herit., 927, I, 4

Phylica paniculata Willd., 1037, I

Helinus integrifolius (Lam.) Kuntze, R. McMahon s.n., 5

VITACEAE

Rhoicissus

rhomboidea (E. Mey. ex Harv. ) Planch. , 974, 5

tomentosa (Lam. ) Wild & Drum. , 1074, 5

Cyphostemma sp. c.f. cirrhosum (Thunb.) Desc. ex Wild & Drum.,

764, 4

TILIACEAE

Sparrmannia ricinocarpa (Eckl. & Zeyh.) Kuntze, 1155, 1

Grewia occidentals L. , 1122, 5

Triumfetta annua L. , 613, 6

MALVACEAE

Abutilon sonneratianum (Cav.) Sweet, 807, 2

Hibiscus pedunculatus L. f, 592, 1, 4

STERCULI ACEAE

Dombeya tiliacea (Endl.) Planch., 938, 1

OCHNACEAE

Ochna

arborea Burch, ex DC. , 968, 5

natalitia (Meisn.) Walp. , 978, 5

serrulata (Hochst.) Walp., 559, 4

164

Bothalia 20,2 (1990)

VIOLACEAE

Rinorea angustifolia (Thouars) Baill., 1032 , 4

FLACOURTIACEAE

Rawsonia lucida flarv. & Sond. , 1167, 4

Kiggelaria africana L. , Moll 3492, 5

Scolopia zeyheri (Nees) Harv. , Wells 1870, 5

Trimeria grandifolia (Hochsi.) Warb. , Moll 3493, 5, 6

Dovyalis

lucida Sim, 988, 4

rhamnoides (Burcli. ex DC.) Harv., 987, 4

Casearia gladiiformis Mast., 582 , 5

PASSIFLORACEAE

Passiflora edulis Sims*, 1073, 1

ACHARIACEAE

Ceratiosicyos laevis (Dumb.) Meeuse, Hilliard 5057, 4, 6

BEGONIACEAE

Begonia

geranioides Hook. f. , Hilliard 2583, 3, 6

sutherlandii Hook, f, 723, 6

OLINIACEAE

Olinia emarginata Burn Davy, Moll 3371. 5

THYMELAEACEAE

Peddiea africana Harv., 994, 4

Gnidia pulchella Meisn. , 1057, I

Englerodaphne pilosa Burn Davy, 600, 4, 6

Dais cotinifolia L. , 587, I, 2, 4

RHIZOPHORACEAE

Cassipourea

gerrardii (Schinz) Alston, 1070, 4

gummiflua Tut. var. verticillata (N.E. Br.) J. Lewis, 962, 1

COMBRETACEAE

Combretum

edwardsn Exell, 1 144, 5

kraussu Hochst., 1065, 5

Quisqualis parviflora Gerr. ex Harv., 998, 5

MYRTACEAE

Eugenia zuluensis Duernmer, 1068, 5

Syzygium gerrardii Harv. ex Hook, f.) Burn Davy, Moll 3539, 5

Eucalyptus grandis Hill ex Maid 7, 980. 1, 2

ONAGRACEAE

Oenothera erythrosepala Borbas*. 923, I

ARAL1ACEAE

Schefflera umbellifera (Sond.) Baill., Sim 19385, 5

Cussonia sphaerocephala Si rev, Moll 3436, 5

APIACEAE

Hydrocotyle americana L.*, 818, 2, 6

Centella asiatica (L. ) Urb. , 620, 3, 6

Sanicula elata Buch.-Ham. , 652, 3, 6

Conium fontanum Hilliard & Burn var. silvaticum Hilliard & Burn,

773, 6

Heteromorpha trifoliata (Wendt.) Eckl. <H Zeyh., 1156, I

MYRSINACEAE

Maesa lanceolata Forssk. var. rufescens (A. DC.) Talon, 588, I

Rapanea melanophloeos (L.) Mez, Wells 1923, 5

SAPOTACEAE

Bequaertiodcndron natalense (Sond.) Heine & J.H. Hems!., 1091, 4

EBENACEAE

Euclea crispa (Thunb.) Guerke var. crispa, Rycroft (1941), I, 2

Diospyros

lycioides Desf. subsp. sericea (Bernh.) De Winter, 1083, 1

whyteana (Hiern) F. White, 1006, 4

OLEACEAE

Chionanthus foveolata (E. Mey.) Stearn subsp. foveolata, Drewes s.n. , 5

Olea capensis L. subsp. macrocarpa (C.H. Wr.) Verdoom, Moll

3489, 5

Jasminium streptopus E. Me y. var. transvaalensis (S. Moore) Verdoom,

Moll 3547, I, 2, 4

LOGANIACEAE

Strychnos sp. c.f. usambarensis Gilg, 1094, 5

Nuxia floribunda Benth., 1077, 1, 5

Buddleja

dysophylla (Benth.) Radik., 1046, 4

pulchella N.E. Br. , 589, 4

salviifolia (L.) Lam., 978, 1, 2

APOCYNACEAE

Carissa bispinosa (L. ) Desf. ex Brenan var. acuminata (E. Me\.) Codd,

996, 4

Strophanthus speciosus (Ward & Harv.) Reber, 5377, 4

ASCLEPIADACEAE

Secamone gerrardii Harv. ex Benth., 999, 2, 4, 5

Riocreuxia torulosa Decne., 732, 4

CONVOLVULACEAE

Cuscuta campestris Yuncker, 1168, 1

VERBENACEAE

Verbena bonariensis L.*. 906. 6

Clerodendrunt glabrum E. Mey. var. glabrum, Rycroft (1941), 5

LAMIACEAE

Leonotis ocymitolia (Burm. f.) Iwarsson var. raineriana (Visiani)

Iwarsson, 819, 2

Stachys

aethiopica L. , 928, 3

caffra E. Mey. ex Benth. , Hilliard 2584, 1, 2

tubulosa MacOwan, 602. 3, 6

Mentha aquatica L. , 737, 1, 2

Plectranthus

dolichopodus Briq. , 604, 3, 6

elegantulus Briq. , 614, 3

fruticosus L'Herit., 601, 4

laxiflorus Benth., 603, 3, 6

rehmannii Guerke, 730, 6

SOLANACEAE

Physalis peruviana L.*, 965, 1, 2

Solanum

acanthoideum E. Mey. , 814, I

aculeastrum Dun. , 1142, 1

aculeatissimum Jacq. , 820. I. 2

didymanthum Dun. var. pluriflorum Dun., 813, 2

giganteum Jacq. , 812, 2

mauritianum Scop.*, 963. 1. 2, 6

nigrum L.*, 1132, 6

SCROPHULAR1ACEAE

Nemesia silvatica Hilliard & Burn, 1034, 6

Diclis reptans Benth., 991, 6

Halleria lucida L. , 967, 1, 5

Phygelius aequalis Harv. ex Hiern, 1038, 1

Bowkeria verticillata (Eckl. & Zeyh.) Schinz, 762, 6

Sutera floribunda (Benth.) Kuntze, Ripley 22, 1

SELAGINACEAE

Selago hyssopifolia E. Mey, Gordon-Gray 6371, 1

GESNERIACEAE

Streptocarpus

fanniniae Harv. ex C.B. Cl., 1127, 1

gardenii Hook., 765, 3

grandis N.E. Br. subsp. grandis, 766, 3, 6

polyanthus Hook, subsp. verecundus Hilliard, 931, 3, 6

silvaticus Hilliard, 669, 3, 6

ACANTHACEAE

Thunbergia

natalensis Hook., 541, 1, 2, 3

purpurata Harv. ex C.B. Cl., 919, 2

Sclerochiton harveyanus Nees, 763, 2

Bothalia 20,2 (1990)

165

Hypoestes

aristata (Vahl) Soland. ex Roem. & Schult., 882 , 1, 3

triflora (Forssk.) Roem. & Schult., 619, 3, 6

Isoglossa hypoestiflora Lindau, 1143, I

Justicia campylostemon (Nees) T. Anders., 562 , 4

RUBIACEAE

Conostomium natalense (Hochst.) Brem. var. glabrum Brem. , 558, 3

Burchellia bubalina (L. f. ) Sims, 975, 5, 6

Gardenia thunbergia L. f, 1044, 4

Hyperacanthus amoenus (Sims) Bridson, 1092, 4

Rothmannia

capensis Thunb. , Moll 3511, 5

globosa (Hochst.) Keay, 1043, 4

Oxyanthus speciosus DC. subsp. gerrardii (Sand.) Bridson, 1095, 2

Tricalysia

capensis (Meisn.) Sim, 976, 4

lanceolata (Sond.) Burn Davy, 1042 , 4

Canthium

ciliatum (Klotzsch) Kuntze, 1072, 4

mundianum Cham. & Schlechtd. , Moll 3494, 4

pauciflorum (Klotzsch) Kuntze, 1063, 4

spinosum (Klotzsch) Kuntze, 1138, 1

Psydrax locuples (K. Schum.) Bridson, Mol! 3485, 4

Pavetta lanceolata Eckl. , 856, 4

Galopina circaeoides Thunb., 621, 1, 2

CUCURBITACEAE

Zehneria scabra (L. f.) Sond. subsp. scabra, 932, 2 , 6

Momordica foetida Schum., Mrs T. Collins s.n., 1

Coccinia palmata (Sond.) Cogn., 847, 1, 2

CAMPANULACEAE

Wahlenbergia madagascariensis A. DC., 1169, I

LOBELIACEAE

Cyphia aspergilloides E. Wimm., Stewart 1789, 6

Lobelia malowensis E. Wimm., 653, 6

Monopsis stellarioides ( Presl ) Urb. subsp. stellarioides, 729, I, 2, 6

ASTERACEAE

Vernonia

anisochaetoides Sond. , 985, 1

mespilifolia Less., Moll 3525, 4

Conyza

scabrida DC. , 1154, 1

sumatrensis (Retz.) E.H. Walker*, 755, 6

Denekia capensis Thunb., 1086, 1, 2

Achyrocline stenoptera (DC.) Hilliard & Bunt, 1035, 1, 2

Helichrysum

cooperi Harv. , 747, 1, 2

mundii Harv., 845, 2

Printzia auriculata Harv., Hilliard 5059, 1

Spilanthes mauritiana (Pers.) DC., 822, 6

Galinsoga parviflora Cav.*, 736, 2

Phymaspermum acerosum (DC.) Kallersjo, Hilliard 4857, 1

Matricaria nigellifolia DC. var. tenuior DC., 753, I, 2

Hilliardia zuurbergensis (Oliv.) B. Nord., 787, 1, 2

Crassocephalum crepidioides (Benth.) S. Moore*. 727, 6

Senecio

deltoideus Less., 881, 2, 4

lygodes Hiern, 853, 6

macroglossoides Hilliard, 852, 2, 4

madagascariensis Pair., 790, 1, 6

panduriformis Hilliard, 721, 2

polyanthemoides Sch. Bip. , 625, 1, 2

tamoides DC., 804, I, 2

Mikaniopsis cissampelina (DC.) C. Jeffrey, 937, 1

Osteospermum herbaceum L. f. , Hilliard 2911 , I

Berkheya

bipinnatifida (Harv.) Roessl. subsp. bipinnatifida, 722, 1, 2

debilis MacOwan, 744, 6

Bothalia 20,2: 167-174 (1990)

Studies in the genus Riccia (Marchantiales) from southern Africa. 17.

Three new species in section Pilifen R. elongata , R. ampullacea and

R. trachyglossum

S. M. PEROLD*

Keywords: Marchantiales, Riccia ampullacea, R. elongata, R. trachyglossum, section Pilifer, southern Africa, taxonomy

ABSTRACT

Species in section Pilifer Volk (1983) are often very difficult to identify (Perold 1990b). Most of them require close examination

of the dorsal cell pillars in reasonably fresh collections, as these cells can seldom be reconstituted in long dried material.

The three species, R. elongata, R. ampullacea and R. trachyglossum, here described as new, have been maintained in

cultures for lengthy periods, during which their dorsal cells were studied. The spore ornamentation was also quite useful

in separating these species. R. elongata is known from eastern Transvaal, R. ampullacea from the Witteberg Mountains

of the eastern Cape Province and the Drakensberg Mountains of Lesotho and Natal, and R. trachyglossum is so far known

only from the highlands of Lesotho.

UITTREKSEL

Spesies in seksie Pilifer Volk (1983) is dikwels baie moeilik om te identifiseer (Perold 1990b). Die meeste vereis deeglike

ondersoek van die dorsale selpilare in redelik vars versamelings, aangesien die omvorming van selle in lank gedroogde materiaal

na hul oorspronklike toestand, selde moontlik is. Die drie spesies, R. elongata, R ampullacea en R trachyglossum, hier

as nuut beskryf, is lank in kulture gekweek, waartydens hul dorsale selle bestudeer is. Die spoorornamentasie was ook nut-

tig om tussen die spesies te onderskei. R elongata is bekend van Oos-Transvaal, R ampullacea van die Witteberge van

Oos-Kaapland en die Drakensberge van Lesotho en Natal, en R . trachyglossum is tot dusver slegs van die hoogland van

Lesotho bekend.

1. Riccia elongata Perold, sp. nov.

Thallus ?monoicus, perennis, mediocris, glaucus, nitens,

simplex vel furcatus, rami 8,0 mm longi, 1,1( — 2 ,0) mm

lati, 0,8 — 1,1( — 1,2) mm crassi, in sectione 1-2 plo latiores

quam crassi; squamae hyalinae, rotundatae, imbricatae,

undulatae, ultra margines thalli productae. Cellulae

dorsales epitheliales globosae, politae, hyalinae, 2 vel 3(vel

4) in columnis separatis, usque ad 200 pm longis

dispositae. Sporae : (70— )75 - 85(— 90) pm diametro,

triangulo-globulares, polares, alatae, imperfecte grosse

reticulatae, superficies distalis trans diametrum ±5-7

areolis, saepe umbone centrale. Chromosomatum numerus

n = 16 (Bornefeld 1989).

TYPE. — Transvaal, 2629 (Bethal); 5 km NE of Kriel

on road to Vandijksdrift, near disused bridge, on dry slope

(— AB), S.M. Perold 2018 (PRE, holo.).

Thallus ?monoicous, perennial, in gregarious patches

(Figure 2A), sometimes partly overlying each other, bluish

green to green, shiny to rather dull proximally, hyaline

scales extending beyond thallus margins (Figures IB; 2B,

E); medium-sized, simple or once to several times sym-

metrically or asymmetrically furcate, branches medium

to widely divergent, up to 8,0 mm long, segments 1,0— 4,0

x 1,1 mm (up to 2,0 mm wide when fully expanded

(Figure 2B)), 0,8— 1,1( — 1,2) mm thick, i.e. ± as wide as

thick, to nearly twice wider than thick in section (Figure

IF), ligulate to oblong, apex subacute, dorsally grooved

towards apex (Figure 2C), margins somewhat obtuse,

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

MS. received: 1989.07.07.

becoming subacute proximally, flanks steep to slightly

obliquely sloping, green, covered with scales (Figure 2D);

ventrally rounded, green; when dry (Figure 1A), margins

tightly inflexed, meeting along midline, with white, wavy

scales covering granular, greyish white dorsal face.

Anatomy of thallus: dorsal epithelium (Figures IE; 2F)

consisting of free-standing 2 or 3(or 4)-celled, fragile,

hyaline pillars, up to 200 /tm long, ± 'a the thickness of

thallus in section, apical cells globose, often wider than

long, 40-50(-60) x 45-65 pm, occasionally conical or

mammillose, and rather smaller, 35 x 45 ftm, middle cells

58 -75 x 50 -75 /xm, basal cells longer than wide,

62-80(-100) x 40-60 /xm; from above, cells glassy and

shiny, bulging, crowded together, top cell smallest, air

pores small, mostly 4-sided, occasionally triangular

(Figure ID); assimilation tissue ± 350 pm thick in section,

'/3 the thickness of thallus, mostly consisting of 6 cells,

35-47 x 37—40 pm, in vertical columns, enclosing

narrow, 4-6(-7)-sided air canals; storage tissue +

450-550 /xm thick, occupying ventral '/2 of thallus, cells

tightly packed, angular, up to 65 pm wide, containing

starch granules; rhizoids arising from ventral epidermal

cells and base of scales, mostly smooth, occasionally

tuberculate, 25 /xm wide. Scales (Figure 1G) rounded,

margins smooth, large, 850—1100 x 500—600 pm, pro-

jecting ± 200 pm (or more) beyond thallus margins,

imbricate, hyaline, base occasionally with some purple-

red cells, cells in body of scale long-hexagonal or oblong-

rectangular, 125-150 x 42-60 pm, in part of margin

brick-shaped, smaller, 25 x 62 pm. Antheridia not

seen. Archegonia only seen in immature state in sections.

Sporangia single, median in proximal part of thallus,

dorsally bulging, containing ± 250 spores each, but

168

Bothalia 20,2 (1990)

thalli rarely sporulating. Spores (70— )75 — 85(— 90) jum in

diameter, triangular-globular, polar, light brown, semi-

FIGURE I. — Riccia elongata. Morphology and anatomy. A, thallus,

dry; B, thallus actively growing and fully expanded; C, thallus

generally with partly indexed sides; D, dorsal cells and air pores

(hatched), assimilation tissue and air canals (stippled), as seen

from above; E, transverse section through dorsal cell pillars;

F, transverse section through branch with scales projecting

beyond margins; G, scale. A, D-G, S.M. Perold 2476\

B, C, S.M. Perold 2018. Scale bar on A-C, F = I mm; D,

E = 50 /im; G = 100 pm.

transparent; wing 3—5 pm wide, wider at perforated

angles, margin smooth to finely crenulate; ornamentation

irregularly and incompletely coarsely reticulate, similar

on both faces: distal face (Figure 3C, D, F) with 5—7

incomplete areolae across diameter, irregularly shaped and

variable in size, 10—25 pm wide, often with central boss

(Figure 3E), free-standing or attached, walls thick and

prominent, sparsely granular, occasionally raised at nodes,

extending onto wing; proximal face (Figure 3A, B) with

triradiate mark clearly defined, joined by some areolar

walls, areolae incomplete, ± 7 jum wide, occasionally with

central boss, walls nearly smooth, slightly raised at nodes.

Chromosome number n = 16 [Bornefeld 1989 (as R.

furfuracea, S.M. Perold 424)].

R. elongata has been named for, and can be recognized

by its longish, narrow, frequently simple branches, with

the sides tightly indexed when dry, and by large, wavy,

white scales. It is rather similar in habit to R. simii Perold

(1990a) (— R. albomarginata auct. non Bisch. sensu Sim)

(Volk pers. comm.), but differs from it by its scales being

less prominent, and less closely imbricate, by its lower

dorsal pillars, spore ornamentation and distribution.

The shiny, round, bulging cells in the dorsal pillars

are a character shared by a few other members in

section Pilifer Volk, e.g. R. concava Bisch. (Perold 1989),

R. furfuracea Perold (1990b) and R. trachyglossum

Perold (1990b), but these species frequently develop purple

colouration on exposure to the sun and differ from R.

elongata in habit, spore ornamentation and distribution.

R. elongata is rare and is so far known only from a

few localities in eastern Transvaal at altitudes of + 1 600—

2 000 m above sea level, with summer rainfall or 800-

1 000 mm p.a. It has been found growing on dry slopes

in association with grasses and Exormotheca sp., on a rock

‘island’ in a lake, (Elandsmeer) and at a seepage area at

the edge of weathered rock outcrops, in association with

R. volkii S. Arnell, R. natalensis Sim, and R. sorocarpa

Bisch. (Figure 4).

SPECIMENS EXAMINED

TRANSVAAL. — 2530 (Lydenburg): 29 km from Dullstroom, at

turnoff on dirt road to Boschhoek, near Marmerkop Station, on hillside

(-AB), S.M. Perold 424 (PRE). 2629 (Bethal): 5 km NE of Kriel on

road to Vandijksdrift, on dry slope near disused bridge (— AB), S.M.

Perold 2476 (PRE). 2630 (Carolina): near Chrissiesmeer, opposite lake,

near roadside, weathered rock outcrop and seepage (—AD), S.M. Perold

1058 (PRE); Chrissiesmeer, Farm Knock Dhu, Elandsmeer, on soil, on

rock ‘island’ above water level (—AD), Smook 4912 (PRE).

It is highly probable that S.M. Perold 303 and Volk

84-644 , both from Mooiriver, Natal, also belong to R.

elongata , but these gatherings are sterile and cannot be

placed here with certainty.

2. Riccia ampullacea Perold , sp. nov.

Tltallus monoicus, ?annuus, laete viridis vel glaucus,

nitens, proximaliter villosus; rami simplices vel 1—2

furcati, usque ad 8,0 mm longi, 1,5-2, 5 mm lati,

0.6— 0,9( — 1,1) mm crassi, 2— 2,5-plo latiores quam crassi

in sectione; squamae magnae, hyalinae. Epithelium dorsale

ex columnis liberis 3- vel 4-cellularibus, 200-250 pm lon-

gis constans, cellulis longioribus quam latis, saepe

Bothalia 20,2 (1990)

169

FIGURE 2. — Riccia elongata. Morphology and anatomy. A, thalli In cultivation; B, branch seen from above; C, apex with groove and scales;

D, apical scales seen from the side; E, marginal scales and dorsal cells; F, dorsal cell pillars. A-F, S.M. Perold2018. A, by A. Romanowski;

B-E, SEM micrographs. Scale bar on A-E = I mm; F = 50 /xm.

medio aliquantum constrictis, ampullaceis (inde nomen).

Sporae: 90-95(-105) pm diametro, triangulo-globulares,

polares, alatae, subtiliter reticulatae et in superficie distali

cum cristis pluribus crassis radiantibus. Chromosomatum

numerus n = 16 (Bornefeld 1989).

TYPE. — Lesotho, 2929 (Underberg): Sani Pass, moun-

tain slopes W of Border Post, on soil in small cave (— CB),

Van Rooy 3573 (PRE, holo.).

Thallus monoicous, ?annual, in crowded gregarious

patches (Figure 5A), bright green to bluish green.

FIGURE 3. — Riccia elongata. Spores. A, proximal face; B, proximal face, side view; C, F, distal face; D. distal face, side view; E, areolae

partly subdivided, one with central papilla. A— F, S.M. Perold 2018. A— E, SEM micrographs; F, LM photograph. Scale bar on A— E

= 50 pm: diameter of spore on F + 80 /xm.

170

Bothalia 20,2 (1990)

FIGURE 4. — Distribution map of R. elongata, • ; R ampullacea, ▲

and R. trachyglossum, □, in southern Africa.

glistening to dull and shaggy-haired proximally, with

hyaline scales extending beyond thallus margins (Figures

5A; 6B); medium-sized, branches simple or once or

twice furcate, branches variously divergent, up to 8,0 x

1,5— 3,0 mm, 0,6 — 0,9( — 1,1) mm thick, i.e. ± 2—3 times

wider than thick in section (Figure 5E), broadly oblong,

apex rounded, shortly emarginate, grooved apically

(Figure 5B), otherwise flat, margins acute, flanks sloping

obliquely outward and upward, green, covered with

hyaline scales; ventrally slightly rounded to flat, green;

when dry, whitish green, felt-like, concave dorsally,

margins incurved (Figure 5B), occasionally inflexed

and rarely meeting along midline, scales hyaline,

imbricate, slightly wavy.

Anatomy of thallus : dorsal epithelium (Figures 5C; 6E)

consisting of free-standing, 3— 4-celled pillars, 200—250

/xm long, '/4 — '/3 the thickness of thallus in section, cells

fragile, hyaline, longer than wide, often somewhat

constricted in the middle, top cell conical, 45— 67(-80)

x 30—37 /am, second cell 50 —70 x 35 — 52 /xm; third cell

80-110 x 37-50 /xm, basal cell with sides sometimes

bulging, 50 -75 x 52 /xm; from above, when fresh, cell

pillars distally inflated, erect, shiny, more proximally many

upper cells already collapsed, not in rows, air pores small,

up to 25 /xm wide, 4-5-sided; assimilation tissue 300-400

/xm thick in section, '/3 — '/2 the thickness of thallus,

consisting of 7 or 8 cells in vertical columns, 37— 42(— 50)

x 25—35 /am, enclosing 4— 6( — 8)-sided air canals (Figure

5D); storage tissue up to 400 /am thick, + '6 the

thickness of thallus, cells round or angular, + 50 /am wide;

rhizoids arising from ventral epidermal cells and base of

scales, mostly smooth, rarely tuberculate, 15—25 /xm

wide. Scales rounded, imbricate (Figure 6D), hyaline,

occasionally dark red toward base, large, 1 000—1 100 x

500 /xm, cells in body of scale 5- or 6-sided, 100—125 x

45 /xm, smaller and brick-shaped toward margin (Figure

5F), 50-62 x 25 /xm. Antheridia numerous, with

conspicuous hyaline necks, + 180 /xm long, at intervals

along middle of thallus, often in very close proximity to

archegonial necks. Archegonia with long thread-like,

purple necks. Sporangia bulging dorsally, overlying

tissue disintegrating and exposing dark spore mass

enclosed in sac (Figure 6F), often with archegonial

and juxtaposed antheridial necks still partly intact;

sporangium with + 480 spores each. Spores 90-95(— 105)

/xm in diameter, triangular-globular, polar, chestnut brown,

semi-transparent to nearly opaque, with wing ± 5 /xm

wide, margin crenulate, marginal angles perforated;

FIGURE 5 .—Riccia ampullacea. Morphology and anatomy. A, thallus

dry; B, thallus wet; C, transverse section through dorsal cell

pillars; D, paradermal section through assimilation tissue, air

canals stippled; E, transverse section through branch; F, scale.

A— F, Van Rooy 3573. Scale bar on A, B, E = 1 mm; C, D, =

50 /xm; F = 100 /xm.

Bothalia 20.2 (1990)

171

FIGURE 6. — Ricci a ampullaceu. Morphology and anatomy. A, thalli in cultivation: B. branch seen from above; C. apex with groove and scales;

D, apical scales seen from the side; E, dorsal cell pillars; F, sporangium emerging through dorsal cell covering, antheridial neck obscuring

archegonial neck. A — F, Van Rooy 3573. A. by A. Romanowski; B-E, SEM micrographs. Scale bar on A— D = 1 mm; E, F = 50 pm.

ornamentation finely reticulate and radiately ridged; distal

face with areolae ± 3—5 pm wide, but rarely complete,

mostly confluent and walls anastomosing into thick, high

ridges, radiating from the centre to the margin (Figure

1C — F); proximal face with triradiate mark distinct or

indistinct, numerous small, less than 5 /am wide, mostly

incomplete areolae on each facet, walls granulate, raised

at nodes, sometimes anastomosing into short, semi-

radiating ridges (Figure 7A, B). Chromosome number

n = 16 (Bornefeld 1989).

FIGURE 7. — Riccia ampullacea. Spores. A. proximal face; B, proximal face, side view; C. F. distal face; D. distal face, side view; E, radiating

ridges on distal face. A, B, D-F, Van Rooy 3573 ; C, Van Rooy 3164a. A-E, SEM micrographs; F, LM photograph. Scale bar on A-E

= 50 /xm; diameter of spore on F + 100 /xm.

172

Bothalia 20,2 (1990)

R. ampullacea is rather similar to R. parvo-areolata

Volk & Perold (1984), as both have wide, concave thalli

when dry, with large hyaline scales and dorsal cell pillars

consisting of 3-4 elongated cells. However, in R. ampul-

lacea the dorsal cells are frequently somewhat constricted

in + the middle and ampulla-shaped (ampulla = small

antique Roman glass phial, used for collecting and storing

tears, and variously shaped, but generally constricted

at the neck or in the middle), hence the specific

epithet. Furthermore, its numerous antheridial necks are

conspicuous and often in very close association with the

archegonial necks, resulting in the fertilization of many

archegonia. The spores generally have thick radiating

ridges on the distal face. Geographically the two species

are widely separated, as R. ampullacea appears to be

restricted to high altitudes of 2 000—3 000 m above sea

level, at mountainous localities in the Drakensberg of

Lesotho and Natal, and the Witteberg of the eastern Cape

Province, whereas R. parvo-areolata is known only from

the western Cape. R. ampullacea grows in association

with Riccia montana Perold, Plagiochasma sp. and with

the moss species, Bryum alpinum Huds. ex With, and

Brachymenium acuminatum Harv. in Hook., in damp

places on humus-rich soil overlying basalt outcrops.

SPECIMENS EXAMINED

NATAL. — 2929 (Underberg): Sani Pass, along basalt cliffs below

escarpment, E of Border Post (— CA), Van Rooy 3635 (PRE).

O.F.S. — 2927 (Maseru): Thaba Patswa, between Hobhouse and

Tweespruit, on top of plateau (-AC), Du Preez 2106a (PRE).

LESOTHO. — 2828 (Bethlehem): 5 km from New Oxbow Lodge, on

road to Mokhotlong, at waterfall over basalt cliff, in tributary of Fanana

River, near Maluti Club Ski Chalet, S aspect, alpine heath-grassland

(—DC), Van Rooy 2971 (PRE); 6 km from New Oxbow Lodge, on road

to Mokhotlong, at waterfall over basalt outcrops (—DC), Van Rooy 3045

p.p., 3050 (PRE). 2928 (Marakabei): Khubelu River crossing between

Tlokoeng and Mapholaneng, cliffs along river banks, SE aspect (— BB),

Van Rooy 3240 (PRE); 19 km from Mokhotlong to Tlokoeng, along small

tributary of Senqu River, wooded stream in grassland with cultivation

(— BD), Van Rooy 3164a (PRE); 35 km from Mokhotlong on road to

Butha Buthe, between Tlokoeng and Mapholaneng, cliffs overlooking

Khubelu River, S aspect (— BD), Van Rooy 3207 (PRE).

CAPE. — 3027 (Lady Grey): Witteberg Mountains, basalt cliffs at top

of Jouberts Pass, 10 km E of Lady Grey (— CB), Van Rooy 2724 (PRE).

3. Riccia trachyglossum Perold , sp. nov.

Thallus monoicus, ?annuus, glaucus, nitens, proximaliter

furfuraceus ut in lingua exasperata (inde nomen); rami

usque ad 5,0 mm longi, 1,0— 2,0 mm lati, 0,7— 0,9 mm

crassi, 1,5—2 plo latiores quam crassi in sectione; squamae

hyalinae, aliquantum ultra margines thalli productae.

Epithelium dorsalis ex columnis liberis 2 vel 3(vel 4)-

cellularibus ± 180 pm longis constans, cellulis globosis.

Sporae: (70 — )80 — 87( — 92) pm diametro, triangulo-

globulares, polares alatae, imperfecte reticulatae, super-

ficie distali trans diametrum cum ± 8 areolis irregulari-

bus. Chromosomatum numerus n = 16 (Bornefeld 1989).

TYPE. — Lesotho, 2929 (Underberg): Sani Top, moun-

tain slopes west of Border Post, on soil bank of jsmall pond

in bog (— CA), Van Rooy 3539 (PRE, holo.).

Thallus monoicous, ?annual, in crowded gregarious

patches or in partial rosettes or scattered, blue-green.

glistening, proximally dull and roughened, with hya-

ne scales extending slightly beyond thallus margins

Figures 8A; 9A); smallish, once to twice symmetrically

or asymmetrically furcate, branches narrowly to medium

divergent (Figure 9B), up to 5,0 x 1, 0-2,0 mm, 0,7— 0,9

FIGURE 8 .—Riccia trachyglossum. Morphology and anatomy. A, thallus

wet; B, thallus dry; C, dorsal cells seen from above, air pores

hatched, below, assimilation tissue with air canals stippled;

D, transverse section through dorsal cell pillars; E, transverse

section through branch; F, scale. A, C, D, F, S.M. Perold 2530:

B, Van Rooy 3539: E, J.M. Perold 33. Scale bar on A, B, E =

1 mm; C, D = 50 pm: F = 100 pm.

Bothalia 20,2 (1990)

173

mm-thick, i.e. 1,5 times to twice wider than thick in section

(Figure 8E), obcuneate to ovate, apex keeled (Figure 8A),

dorsal face distally grooved (Figure 9C), the sides raised,

tumid, margins subacute, flanks rather steep to sloping

obliquely, green, covered by hyaline scales; ventrally

gently rounded to almost flat , green; when dry (Figure

8B), margins apically inflexed, meeting along midline,

otherwise raised or incurved , dorsally white, scurfy,

scales only apically visible, flanks yellowish to reddish

brown occasionally.

Anatomy of thallus: dorsal epithelium in free-standing

cell pillars (Figures 8D; 9E), ± 180 /am tall, '/4 the

thickness of thallus in section, consisting of 2 or 3(or 4),

fragile, hyaline cells, with bulging sides, top cell ±

globose, rarely conical, 32—45 x 47—55 /am, second cell

55—62 x 47-62 /xm, basal cell 75—100 x 52 — 65 /a m;

from above, when fresh, dorsal cells irregular in size,

inflated, air pores 4-sided (Figure 8C); assimilation tissue

+ 350 /am thick, almost h2 the thickness of thallus in

section, generally consisting of 6 or 7 cells in vertical

columns, 50—65 x 58-62 /am, enclosing (3— )4(— 5)-

sided air canals (Figure 8C), ± 25 /am wide; storage tissue

up to 350 /am thick in section, cells angular, closely

packed, 37— 55 /am wide; rhizoids arising from ventral

epidermis and base of scales, mostly smooth, occasionally

tuberculate, 15 /am wide. Scales rounded, imbricate

(Figure 8F; 9D), hyaline, 750 x 500 — 550 /am, cells in

body of scale long-rectangular to short-hexagonal,

112 — 137( — 187) x 42 - 65 /am, smaller towards base, at

margin brick-shaped to irregularly shaped (Figure 8F).

Antheridia (Figure 9F) with hyaline necks + 125 /am long,

in 1 or 2 rows along middle of thallus (Figure 8A).

Archegonia with thin, purple necks. Sporangia bulging

dorsally along midline, numerous, containing + 580

spores each. Spores (70 — )80 — 87 ( — 92 ) /am in diameter,

triangular-globular, polar, light brown, semi-transparent.

with wing ± 5 /am wide, rather wider at perforated angles,

margin finely crenulate; ornamentation reticulate,

dissimilar on 2 faces: distal face (Figure 10 C— F), with

+ 8 angular to irregular areolae across diameter, 5—8 /am

wide, central ones often incomplete, walls sprinkled with

granules, raised at nodes; proximal face with triradiate

mark distinct, facets with areolae incomplete, ± 3—5 /am

wide, walls irregular, thin (Figure 10A, B). Chromosome

number n = 16 [Bornefeld 1989 (as R. furfuracea, J.M.

Perold 33, 34)\.

Due to the collapse of many dorsal cells, especially in

the proximal part of the thallus, the dorsal face has a rather

roughened or scurfy appearance. For this reason, the

specific epithet, trachyglossum has been chosen. It is

derived from a Greek phrase, meaning ‘rough tongue’; the

word is treated as a noun in apposition to the generic name,

and therefore has a neuter ending, even though the name

Riccia is feminine.

R. trachyglossum is distinguished from other species in

section Pilifer , which have globose to bulging dorsal cells,

by its somewhat smaller size, rather low hyaline scales and

raised, tumid margins toward the apex. The spores are

generally incompletely reticulate on both faces. It is so

far known only from Lesotho, at altitudes + 2 500— 3 000

m above sea level, where it grows on soil banks

in bogs, together with other Riccia spp. : R. stricta

(Lindenb.) Perold, R. crystallina L. emend. Raddi, R.

sorocarpa Bisch. and with Cyperaceae spp.

SPECIMENS EXAMINED

LESOTHO. — 2927 (Maseru): about 37 km E of Maseru, top of

Bushman’s Pass (Lekhale La Baroa), on soil at edge of exposed Hat rock

outcrop (-BD), J.M. Perold 33. 34 (PRE). 2929 (Underberg): Sam Top.

S of Border Post, between earth dam and bog, on soil banks (— CA).

S.M. Perold 2530. 2531 (PRE).

FIGURE 9 — Riccia trachyglossum. Morphology and anatomy. A, B, thallus from above; C. apex with groove; D. apical scales seen from the

side; E, dorsal cell pillars; F, antheridial neck and dorsal cells. A-F, S.M. Perold 2531. A-F. SEM micrographs. Scale bar on A-D

= 1 mm; E, F = 50 /am.

174

Bothaiia 20,2 (1990)

FIGURE 10. — Riccia trachyglossum. Spores. A, proximal face; B, proximal face, side view; C, F, distal face; D, distal face, side view; E. areolae

on distal face. A, C, J.M. Perold 33: B, S.M. Perold 2530: D-F, Van Rooy.3539. A-E, SEM micrographs; F, LM micrograph. Scale

bar on A-E = 50 ^m; diameter of spore on F ± 85 /tm. Drawings by J. Kimpton; SEM and LM micrographs by S.M. Perold.

ACKNOWLEDGEMENTS

The author wishes to thank Dr H.F. Glen, NBI, for

the Latin diagnoses and for his and the chaplain of

Irene Homes, Rev. M.H. de Lisle's help in choosing

specific epithets for two of the new species, R. ampullacea

and R. trachyglossum. Sincere thanks are also

due to Prof, (emer.) Dr O.H. Volk of Wurzburg University

for numerous discussions, for critically reading the

manuscript and for his generosity in sending me his

notes and drawings of R. ampullacea. A special word of

thanks to Mr J. van Rooy, NBI, and appreciation of my

late husband, J.M. Perold, for collecting specimens;

to Mrs A. Romanowski, photographer, NBI; Ms J.

Kimpton, artist, and to Mrs E. Bunton, typist, for their

contributions to this paper.

REFERENCES

BORNEFELD, T. 1989. The Riccia species of S and SW Africa.

Chromosome numbers and composition of the chromosome sets.

Nova Hedwigia 48: 371—382.

PEROLD, S.M. 1989. Studies in the genus Riccia (Marchantiales) from

southern Africa. 14. R. concava , section Pilifer. Bothaiia 19:

161-165.

PEROLD, S.M. 1990a. Studies in the genus Riccia (Marchantiales) from

southern Africa. 16. R. albomarginata and R. siitiii , sp. nov.

Bothaiia 20: 31-39.

PEROLD, S.M. 1990b. Studies in the genus Riccia (Marchantiales) from

southern Africa. 18. New species in section Pilifer from the NW

Cape: R. futfuracea, R. vitrea and R. namaquensis. Bothaiia 20:

175-183.'

VOLK, O.H. 1983. Vorschlag fur eine Neugliederung der Gattung Riccia

L. Mitteilungen der Botanischien Staatssammlung, Miinchen 19:

453-465.

VOLK, O.H. & PEROLD, S.M. 1984. Studies in the genus Riccia

(Marchantiales) from the south-west Cape. Bothaiia 15: 117-124.

Bothalia 20,2: 175-183 (1990)

Studies in the genus Riccia (Marchantiales) from southern Africa. 18.

New species in section Pilifer from the NW Cape: R. furfuracea,

R. vitrea and R. namaquensis

S. M. PEROLD*

Keywords: Marchantiales, Namaqualand, section Pilifer, Riccia , southern Africa, taxonomy

ABSTRACT

Another three new species of Riccia in section Pilifer Volk are described from Namaqualand, namely R. furfuracea,

R. vitrea and R namaquensis. There are certainly more undescribed species present in that region, but species from there

are often very difficult to distinguish: almost all have hyaline scales, the free-standing dorsal cell pillars need to be examined

in living plants and the spore ornamentation is quite variable.

(J1TTREKSEL

Nog drie nuwe Riccia- spesies, seksie Pilifer Volk, wat in Namakwaland voorkom, word beskryf: R. furfuracea, R. vitrea

en R. namaquensis. Daar is bepaald nog meer onbeskrewe spesies in die streek teenwoordig, maar dit is dikwels moeilik

om tussen spesies van daardie gebied te onderskei: feitlik almal het hialiene skubbe, die vrystaande dorsale selpilare moet

in lewende plante ondersoek word en die spoorornamentasie is redelik veranderlik.

INTRODUCTION

Namaqualand lies in the north-western and western Cape

Province; it comprises four regions: the Richtersveld, the

Sandveld, the Knersvlakte and the Namaqualand Klip-

koppe (domed, granite hills) which are geographically and

floristically quite distinct. The region is ± 100—200 m

above sea level, with its mountainous areas at an elevation

of less than 2 000 m. Namaqualand is arid with sporadic

sparse winter rain, 100—200 mm p.a., yet it is renowned

for its magnificent display of spring flowers.

The granite outcrops are particularly rich in Riccia

species. Several interesting and unique endemic species

have recently been described from here: R. schelpei

Volk & Perold (1986a), R. alboporosa Perold (1989a),

R. tomentosa Volk & Perold (1990) (with its spores per-

manently coherent in tetrads), R. parva-areolata Volk &

Perold (1984), R. alatospora Volk & Perold (1985), R.

hirsuta Volk & Perold (1986b) and R. hantamensis Perold

(1989b), the last four being members of section Pilifer Volk

(1983). Other Riccia species found here are: R. villosa

Steph., R. concava Bisch. ex Krauss, R. albomarginata

Bisch. ex Krauss (all three in section Pilifer ), R. limbata

Bisch. ex Krauss, R. nigrella DC., R. sorocarpa Bisch.,

R. bullosa Link ex Lin^denb. and, occasionally at tempo-

rary streams and seepages, annual species, R. cupulifera

A.V. Duthie, R. crystailina L. emend. Raddi, R. caver-

nosa Hoffm. emend. Raddi, R. curtisii (Aust.) Steph. and

R. purpurascens Lehm. & Lindenb.

In this area species of section Pilifer are common, in

contrast with the summer rainfall parts of southern Africa,

where they are much scarcer and belong to different

species. Mosses frequently growing in association with

Riccia spp. in this region are Barbula crinita Schultz,

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

MS. received: 1989.07.07.

Desmatodon convolutus (Brid.) Grout, Didymodon

australasii (Hook. & Grev.) Zander, D. xanthocarpus (C.

Mull.) Magill, Chamaebryum pottioides Ther. & Dix.,

Goniomitrium africanum (C. Miill.) Broth., Grimmia

laevigata (Brid.) Brid., G. pulvinata (Hedw.) J.E. Sm.,

Bryum argenteum Hedw., B. canariense Brid. and B.

torquescens Bruch ex De Not.

Three new species in section Pilifer from Namaqualand

are described in this paper: R. furfuracea, R. vitrea

and R. namaquensis. It is quite certain that there are

more species present, some probably with a restricted

distribution, but it is generally very difficult to distinguish

between them vegetatively, as differences are often subtle.

Nearly all have rounded hyaline scales and the fragile,

somewhat variable, dorsal cell pillars, which constitute a

most important character, need to be examined in

living plants. Under different conditions of growth and in

cultivation, these Riccia species display the same plasticity

of the gametophyte that makes the taxonomy of much of

the genus so difficult. Moreover, two or more species

commonly grow together in mixed stands, which can be

confusing, especially if the sporangia have disintegrated

and the spores are scattered.

Spores from all sporulating material of species in section

Pilifer held at BOL and PRE have been studied by SFM

and LM but, with some exceptions, the spore ornamenta-

tion shows a spectrum of variation which unfortunately

often makes it a less reliable and useful distinguishing

character in this section. It would appear that continued

observation over several seasons of marked populations

in the field, and the use of a fully equipped mobile

laboratory may be the only means toward clearly defining

the characters and limits of more species, but the long

travelling distances involved make it rather unpractical.

To isolate species, some other reliable vegetative characters

are needed, such as isozymes, which Dewey (1988)

separated by starch gel electrophoresis and found to be

176

Bothalia 20,2 (1990)

species specific, but once again, living plants are required.

Regrettably, it seems therefore that many collections,

especially the older ones, of species in section Pilifer are

unidentifiable, at least for the present.

1. Riccia furfuracea Perold , sp. nov.

Thallus monoicus, perennis, mediocris, glauco-viridis,

semel vel compluries furcatus, rami usque ad 8,0 mm

longi, (1,1 — )1,5 — 1,8( — 2,0) mm lati, 0,9— 1,2 mm crassi,

in sectione transversali minus quam duplo latiores quam

crassi, ovati vel late ovati, apice obtuse cuneati, squamis

hyalinis, basin versus purpureis, rotundatis, imbricatis,

magnis, supra marginem thalli eminentibus. Epithelium

dorsale ex columnis liberis vel satis indistinctis humilibus

2- vel 3-cellularibus constans, cellulis 75—150 /x m crassis,

cellula apicali mammosa vel globosa. Sporae (70 — )75 — 87

(—95) /xm diametro, deltoideo-globulares, polares, ala

parum undulata, imperfecte grosseque reticulata, pagina

distalis 7— 9(— 10) areolis incompletis, saepe cruce centrali,

pagina proximalis areolis indistinctis.

TYPE.— Cape, 3220 (Sutherland): Haashoogte, SW of

De Kom, Klein Roggeveld, damp east slope with dense

short scrub (—DA), Oliver 8957a (PRE, holo.).

Thallus monoicous, perennial, in crowded gregarious

patches (Figure 2A), glaucous green to green, often with

purple colouring along margins, rather dull to ± shiny,

large hyaline scales extending beyond thallus margins

(Figures 1A; 2B); medium-sized, once to several times

furcate, branches moderately to widely divergent, up

to 8,0 x (1,1 — )1,5 — 1,8( — 2,0) mm, 0,9 — 1,2 mm thick,

generally less than twice wider than thick in section

(Figure IE), ovate to broadly ovate, apex bluntly wedge-

shaped, emarginate, dorsally deeply grooved toward apex

(Figure 2C), gradually flattening out at ± midway along

length of branches, margins subacute, somewhat raised

distally, becoming acute and shortly winged, flanks erect

to steeply or obliquely sloping more proximally, green,

turning purple on exposure to intense sunlight, covered

with scales; ventrally rounded, green; when dry (Figure

IB), margins incurved to indexed, meeting along midline

over scurfy, yellowish green to glaucous green dorsal face,

scales ± opaque, at apex large, conspicuous (Figure 2D),

less prominent proximally.

Anatomy of thallus'. dorsal epithelium (Figures 1C; 2F)

75 — 150( — 180) ftm thick, consisting of two or three cells

in low, free-standing pillars, cells generally wider than

long, top cell mammillose or globose, rarely conical,

(32— )35— 47 x 40—52 /xm, second cell (30— )37— 40

(—50) X (40— )52— 60(— 75) /xm, third cell (if present)

(35-) 40—47 x (42 — )50— 62(— 75) /xm, cells mostly

collapsed toward margins and proximally; from above

(Figures ID; 2E), dorsal cells not very obviously arranged

in free-standing pillars, cells closely packed, not in rows

and not of uniform size, smaller cells wedged in between

larger ones, air pores 3—4-sided, up to 25 /xm wide; as-

similation tissue 350-450 fim thick, rather less than ‘/2

the thickness of thallus in section, consisting of columns

of 6— 8 cells; (37-)50— 62 x 32 — 40( — 45) /xm, enclosing

narrow air canals; storage tissue 400-550 /xm thick, +

'/2 of thickness of thallus, cells angular to rounded, ± 55

/xm wide, with numerous starch granules; rhizoids arising

FIGURE 1. — Riccia furfuracea . Morphology and anatomy. A, thallus,

wet; B, thallus, dry; C, transverse section through dorsal cell

pillars and assimilation tissue; D, dorsal cells and air pores

(hatched) seen from above; E, transverse section through branch;

F. scale. A, S.M. Perold 2180: B, Oliver 8910: C, D, S.M. Perold

1476: E, S.M. Perold 1398a: F, S.M. Perold 1475. Scale baron

A, B, F = 1 mm; C, D = 50 /xm; F = 100 /xm.

from ventral epidermal cells and from base of scales, some

smooth, others tuberculate, ± 25 /xm wide. Scales (Figure

IF) rounded, large, 750—1200 x 500—625 /xm, pro-

jecting ± 125 /xm beyond thallus margins, imbricate

Bothalia 20,2 (1990)

177

FIGURE 2. — Riccia furfuracea. Morphology and anatomy. A, thalli in cultivation; B, branches seen from above; C, apex with groove. D, scales

at apex; E, dorsal cell pillars seen from above; F, dorsal cell pillars seen partly in profile. A-F, S.M. Perold 2180. A, by A. Romanowski;

B— E, SEM micrographs. Scale bar on A-D = 1 mm; E, F = 50 ^m.

(Figure 2D), hyaline, base often partly purple to deep red,

cells in body of scale 50 — 85 X 37— 42 gm, oblong-

hexagonal, thin-walled, at almost smooth margin smaller,

in one or two rows. Antheridia in rows along midline, with

hyaline necks. Archegonia with purple-brown necks.

Sporangia bulging dorsally, up to 1000 pm wide, single,

medianly in proximal part, containing 470—510 spores

each. Spores (70— )75 — 87(— 95) ^m in diameter,

triangular-globular, polar, light brown to brown, semi-

transparent, wing 5,0— 7,5 pm wide, slightly undulating,

notched or with a pore at marginal angles, margin

finely crenulate; ornamentation incompletely and rather

coarsely reticulate, + dissimilar on two spore faces:

distal face (Figure 3C, D) with 7— 9( — 10) areolae across.

FIGURE 3. — Riccia furfuracea. Spores. A, proximal face; B, proximal face, side view; C, F, distal face; D, distal face, side view; E. areolae

and wing. A, B, D, E, S.M. Perold 2425: C, S.M. Perold 1890\ F, S.M. Perold 1489. A'-E. SEM micrographs; F, LM photograph. Scale

bar on A-E = 50 gm; diameter of spore on F + 85 pm.

178

Bothalia 20,2 (1990)

7,5—10,0 /x m wide, central ones incompletely separated,

some cross walls absent or poorly developed, others linked

up and forming an irregular, central cross (Figure 3F),

areolar walls thick, rounded, often dotted with granules

(Figure 3E), raised into large, blunt papillae at nodes;

proximal face (Figure 3A, B) with triradiate mark distinct,

narrow, sprinkled with granules which sometimes also

cover adjacent areas of each facet, areolae generally poorly

defined, incomplete, walls low, faintly granular, raised into

papillae at nodes.

The specific epithet, furfuracea , is derived from the

Latin furfuraceus meaning scurfy, and refers to the

roughened dorsal surface of the dry thallus. This species

is recognized by, and distinguished from other Riccia

species in section Pilifer , that have rather low dorsal pillars

with + bulging cells, such as R. concava Bisch. ex Krauss

(Perold 1989c), R. elongata Perold (1990b), R. trachyglos-

sum Perold (1990b) and R. pulveracea Perold (1990c), by

the mammillose top cells, closely packed together, neither

arranged in rows nor uniform in size, by large apical scales

and by coarsely reticulate spores with a central cross on

the distal face and low-walled, poorly defined areolae on

the proximal face.

It often grows together with other Riccia species on soil

at the margins of granitic rock outcrops, near seepages

or on stream banks. It is only known from the north-

western and south-western Cape (Figure 4).

SPECIMENS EXAMINED

CAPE. — 2917 (Springbok): Hester Malan Res., ± 4 km N of office

(— DB), S. M. Perold 1398 p.p., 1400\ Carolusberg, seepage area, near

old mine (— DB), S.M. Perold 2033-2035, 2045, 2049 (PRE). 3018

(Kamiesberg): 17-19 km NE of Kamieskroon, 3—5 km after turnoff on

road to Rooifontein, near seepage areas (— AA), S.M. Perold 1465 p.p.,

1476, 2140, 2155 p.p. , 2156 p.p. , 2171, 2180 (PRE); 1-2 km beyond Willem

Stone Bridge, Pedroskloof, on road to Rooifontein from Kamieskroon

(— AA), S.M. Perold 1489, 2176 (PRE); near village of Rooifontein, large

rock outcrop (-AB), S.M. Perold 1515 (PRE). 3119 (Calvinia): NE of

Nieuwoudtville, 7 km along road to Rondekop, near rocky stream bed

(—AC), S.M. Perold 2319 p.p., 2322 p.p. (PRE); Van Rhynshoek Farm,

16 km on road from Calvinia to Klipwerf, 6 km from FM tower (— BD),

S.M. Perold 1854 (PRE); Farm Daantjie-se-Kraal, 37 km along road

between Soetwater and Clanwilliam, 8 km before Botterkloof Pass, near

rock pool (— CB/CD), S.M. Perold 1869, 1870 (PRE); northern Roggeveld,

Vondelingsfontein, on damp ground in lee of scrub (— DD), Oliver 8910

(PRE). 3120 (Williston): northern Roggeveld, Knegtsbank, kloof N of

farm, southern slopes with dense renoster scrub (— CC), Oliver 8921

(PRE). 3219 (Wuppertal): 3 km before turnoff to Biedouw/Wuppertal,

on road R364 between Soetwater and Clanwilliam, at streamlet near road

(— AA), S.M. Perold 1879 (PRE); Biedouw Youth Camp, 19 km along

road to Wuppertal, at sandstone rock outcrop near waterfall (-AA), S.M.

Perold 1890, 1892, 1895 (PRE). 3220 (Sutherland): Farm Bergsig, 50

km S of Sutherland, 21 km along dirt road to Wolfhoek, at stream bank

behind farmhouse (-DA), S.M. Perold 2425, 2429 ( PRE); Haashoogte,

SW of De Kom, Klein Roggeveld, damp east slope with dense short scrub

(—DA), Oliver 8957a (PRE, holo.).

2. Riccia vitrea Perold , sp. nov.

Thallus ?monoicus, perennis, statura mediocri vel satis

magna, argenteo-viridis vel ferreus, proximale villosus,

semel vel compluries furcatus, rami usque ad 9,0 mm

longi, 1,2 — 1,8( — 2,3) mm lati, (0,9 — )1,2 — 1,5 mm crassi,

squamae hyalinae, interdum rotundatae, imbricatae,

undulatae, supra margines thalli eminentes. Epithelium

dorsalis ex columnis liberis 4- vel 5-cellularibus erectis

vel arcuatis 320-450(-500) /xm longis constans, columnae

fragmenta vitri simulantes, itaque nomen. Sporae (72—)

FIGURE 4.— Distribution map of R. furfuracea, # ; R vitrea, 0; and

R. namaquensis, □ .

75— 100(— 110) /xm diametro, deltoideo-globulares, polares,

alatae; pagina distalis, omnino vel imperfecte reticulata

vel radiate cristata, pagina proximalis areolis parvis valde

incompletis, parietibus ad papillas humiles reductis.

Chromosomatum numerus n = 8 (Bornefeld 1989).

TYPE. — Cape, 3018 (Kamiesberg); 19 km NE of

Kamieskroon, 5 km after turnoff on road to Rooifontein,

at large flat rocks, seepage area (— AA), S.M. Perold 1475

(PRE, holo.)

Thallus ?monoicous, perennial, in crowded, gregarious

patches (Figure 6A), distally steel-grey to silvery green,

shiny, proximally dull, shaggy-haired to cottony, with thick

white matted pile, hyaline scales extending beyond thallus

margins (Figure 6C); medium-sized to rather large, once

(Figure 6B) to several times furcate, branches narrowly

(Figure 5A) to moderately divergent, up to 9 mm long,

segments up to 4,0 x 1,2 — 1,8( — 2,3) mm and (0,9—)

1,2— 1,5 mm thick, i.e. almost as wide as thick, to l'/2

times wider than thick in section (Figure 5E), obovate,

apically acute, very fleshy and thick, deeply grooved

dorsally from apex to ± midway along length of branches,

but groove mostly obscured by tall dorsal cell pillars which

arch and interlock above it (Figure 6D), margins acute,

raised, flanks toward apex steep, becoming somewhat

obliquely sloping proximally, purplish; ventral face

rounded to almost flat, green;- when dry (Figure 5B)

margins distally inflexed, meeting along midline, flanks

densely covered with large, imbricate, wavy or billowing

scales, coarsely-celled, ± shiny, hyaline to pale cream,

sometimes with white streaks of encrusted salts, more

proximally scales dull.

Anatomy of thallus : dorsal epithelium (Figures 5C, D;

6F) consisting of 4 or 5(— 6)-celled, free-standing, fragile,

hyaline pillars, ± uniformly wide, to slightly wider toward

base, 320— 450(— 500) /im long, ± */3 the thickness of

thallus in section, cells 2(-3) times longer than wide, top

cell long-conical to bent, (60— )74— 92 x 25 —37 /am,

second cell 85—105 x 25—50 /xm, third cell 68— 125(— 137)

x 32-55 /xm, fourth cell 62— 125(— 150) x (37-)47-60

(—67) /xm, basal cell 65—100 x 37— 47(— 50) /xm; from

above, tall cell pillars conspicuous, fine, arched (Figure

6E), or erect (Figure 6F), and then rather like glass

Bothalia 20,2 (1990)

179

splinters, proximally collapsed into thick, matted, cotton-

woolly pile, air pores 4( — 6)-sided, obscured; assimilation

tissue (350— )400— 500 /xm thick, ± '/3 the thickness of

thallus; in section, consisting of up to 10 cells in vertical

columns, cells 32— 45(-50) x 28—32 /xm, enclosing

FIGURE 5. — Riccia vitrea. Morphology and anatomy. A, thallus wet;

B, thallus dry; C, transverse section through erect dorsal cell'

pillars and scales; D, transverse section through arched and

erect dorsal cell pillars; E, transverse section through branch;

F, scale. A, D, F, S.M. Perotd 2I49\ B, S.M. Perold 1475 ; C,

E, S.M. Perold 1419. Scale bar on A, B, E = 1 mm; C, D =

50 ix m; F = 100 |im.

narrow air canals; storage tissue occupying remaining

ventral ‘/3 of thallus in section (Figure 5E), 400(— 500)

/x m thick, cells closely packed, up to 50 /xm wide; rhizoids

arising from ventral epidermal cells, some smooth, others

tuberculate, 17—20 /xm wide. Scales (Figure 5F) rounded,

large, 1250—1750 x 600—850 /xm, projecting ± 200 /xm

above thallus margins, imbricate, billowing, hyaline to pale

cream, base sometimes reddish purple, cells in body of

scale up to 150 x 50 /xm, oblong-hexagonal, walls straight

to somewhat bulging, often appearing rather ‘stretched’,

at margins cells smaller, wider than long, in 1—4 rows.

Antheridia with long hyaline necks, obscured by tall dorsal

cell pillars. Archegonia with purple necks, scattered along

groove. Sporangia obscured or, toward base, bulging

dorsally, containing + 550 spores each. Spores (72—)

75 — 100( — 110) /xm in diameter, triangular-globular,

polar, brown to dark brown, mostly opaque, wing 5,0— 7,5

/xm wide, sprinkled with granules, perforated at wider

marginal angles, margin crenulate (Figure 7E); ornamenta-

tion completely or incompletely reticulate to radiately

ridged, dissimilar on two spore faces: distal face (Figure

7C, D) with up to 16 rather irregular areolae aqross

diameter, + 5 /xm wide, walls thin, granular, raised at

nodes, but frequently thickened and linked up to form short

radiating ridges with areolae confluent, especially in centre

of distal face (Figure 7F); proximal face with triradiate

mark well to poorly defined, areolae small, very incom-

plete, often only coarse granules or low papillae at the

nodes, intervening walls absent or very low (Figure 7A,

B). Chromosome number n = 8 (Bornefeld 1989).

R. vitrea has been so named because of the similarity

of the tall dorsal cell pillars in full grown, living plants,

to glass (Latin: vitrum ), or rather, slivers of shattered glass.

It can be recognized by the large billowing, hyaline scales,

sometimes turning a pale cream colour while remaining

transparent, and by the tall dorsal cell pillars, similar to

those in R. villosa Steph. and R. simii Perold (1990a)

(— R. albomarginata auct. non Bisch.) but not so ‘fine’,

hardly tapering and often interlocking. The spores are

variable in size and in ornamentation, especially on the

distal face which is completely to incompletely reticulate.

Spores from some specimens, Perold 1423, 1424 and 1475 ,

were repeatedly examined and photographed. A number

of specimens placed here were not fertile, however; others

had few sporangia, whereas in Perold 1423, they were

present in abundance, but seemingly from the previous

season, as many had disintegrated. Gemmae were found

in the type specimen, Perold 1475.

R. vitrea is so far known from only a few localities in

Namaqualand (Figure 4), where it grows in crowded stands

at seepages, on generally rather coarsely grained soil der-

ived from exfoliating granitic rock, together with R. bul-

losa Link ex Lindenb., R. albomarginata Bisch.

ex Krauss, R. schelpei Volk & Perold (1986a), R. nama-

quensis Perold (1990b) and R. furfuracea Perold (1990b).

SPECIMENS EXAMINED

CAPE. — 2917 (Springbok): Hester Malan Res., ± 4 km N of office

(— DB), S.M. Perold 1398 p.p. (PRE); Carolusberg, seepage area, near

old mine (-DB), S.M. Perold 1419. 1422-1425 p.p.; 2041. 2043. 2044.

2046, 2047 (PRE); Carolusberg (W), seepage area, 3800 ft., 14-9-1977

(-DB), Schelpe 7776 (BOL, P, PRE). 3018 (Kamiesberg): 19 km NE

of Kamieskroon, 5 km after turnoff on road to Rooifontein, at seepage

and large flat rocks (— AA), S.M. Perold 1475 (PRE, holo.).

180

Bothalia 20,2 (1990)

FIGURE 6. — Riccia vitrea. Morphology and anatomy. A, thalli in cultivation; B, branches seen from above; C, branches seen partly from the

side; D, apical scales and dorsal cell pillars arched over groove; E, arched dorsal cell pillars; F, erect dorsal cell pillars. A-F, S.M. Perold

2149. A, by A. Romanowski; B-E, SEM micrographs. Scale bar on A-D = 1 mm; E, F = 50 jxm.

3. Riccia namaquensis Perold , sp. nov.

Thallus ?monoicus, perennis, mediocris, purpureo-

viridis vel viridis, politus vel hebes, semel vel bis furcatus,

rami usque ad 8,0 mm longi, 1,8 — 2,3( — 2,5) mm lati,

1,2 — 1 ,4( — 1,6) mm crassi, in sectione sesqui- vel fere

duplo latiores quam crassi, oblongi vel obovati; squamae

hyalinae, pluristratae, arete imbricatae, ut videtur albae,

undulatae, supra marginem thalli eminentes. Epithelium

dorsalis ex columnis liberis 3- vel 4-cellularibus, (200—)

250— 350(— 400) ftm crassis constans. Sporae (65— )70-78

(—85) jtim diametro, deltoideo-globulares, polares, alatae,

(12 — )14— 16 areolis coarctatis trans paginam distalem;

pagina proximalis areolis parvis.

FIGURE 1.— Riccia vitrea. Spores. A, proximal face; B, proximal face, side view; C, F, distal face; D, distal face, partly in side view; E, areolae

toward wing. A, S.M. Perold 1423', B, D-F, S.M. Perold 1475', C, S.M. Perold 1424. A— E, SEM micrographs; F, LM photograph. Scale

bar on A— E = 50 //m; diameter of spore on F ± 90 ^m.

Bothalia 20.2 (1990)

181

TYPE. — Cape, 2918 (Gamoep): Carolusberg, Hester

Malan Res., near old mine, Hat granitic rock outcrop, at

seepage (— CA), S.M. Perold 1420 (PRE, holo.).

Thallus ?monoicous, perennial, in crowded gregarious

patches (Figure 9A), or occasionally in partial rosettes ±

25 mm across, becoming bare toward centre where basal

parts of thalli have died and disintegrated, dorsally purplish

green to bright green, shiny to rather dull proximally,

medium-sized, once or twice furcate, occasionally simple,

branches variously divergent (Figure 9B), up to 8,0 x

1,8 — 2,3( — 2,5) mm, 1,2 — 1,4( — 1,6) mm thick, generally

l'/2 times to nearly twice wider than thick in section

(Figure 8E), oblong to obovate (Figure 8A), apically

emarginate, grooved toward apex (Figure 9D), but soon

becoming flat to slightly concave; margins rather obtuse

to subacute, flanks distally nearly erect or slightly bulging,

toward base sloping steeply to more obliquely, often

turning deep purple below, covered by large, densely

imbricate, wavy, hyaline scales; ventral face rounded to

nearly flat, green; when dry (Figure 8B, C), margins

tightly inflexed, white scales often clasped together along

midline and covering dorsal face.

Anatomy of thallus: dorsal epithelium (Figures 8D; 9E,

F) (200— )250-350( -400) /zm thick, consisting of 3 or

4 (rarely 5) cells in densely crowded treestanding, hyaline

pillars, cells generally longer than wide, top cell variable,

often conical, up to 65 x 50( —60) /zm, rarely small and

rounded, 30 x 25 /zm, second cell 50—67 x 40 — 52( — 60)

/zm. very occasionally also small and rounded like some

top cells, third and fourth (basal) cells up to 100 X

37- 52(— 62) /zm, soon collapsing toward margins and

proximally; air pores from above, obscured by dorsal

pillars, shape generally 4-sided, sometimes irregular;

assimilation tissue 300-450 /zm thick, + 7,0 the thick-

ness of thallus in section, consisting of vertical columns

of 6 — 8( — 10) cells, 37— 52 x (30 — )37 — 45 /zm, separated

by narrow air canals; storage tissue occupying ventral 'o

of thallus, cells angular, closely packed, up to 60 /zm wide;

rhizoids, some smooth and others tuberculate, ± 20 /zm

wide, arising from ventral epidermal cells. Scales (Figures

8F; 9C) large, 1100-1350 x 650 /zm, wavy, closely

imbricate, projecting 150—250 /zm beyond thallus margins,

hyaline, but appearing white as several layers of scales are

superimposed, base sometimes with purple blotches, cells

(4— )5— 6-sided, in body of scale 65 —75—112 X 50 /zm,

smaller at margins, + 45 x 50 /zm. Antheridia in one

or two rows along midline of thallus, necks hyaline.

Archegonia scattered, necks purple. Sporangia situated

toward base, dorsally bulging, with ± 600 —700 spores

each. Spores (65-)70-78(-85) /zm in diameter,

triangular-globular, polar, light brown to deep brown,

semitransparent to opaque; wing ± 5 /zm wide, perforated

at angles, stippled with granules, margin crenulate;

ornamentation reticulate, rather dissimilar on the two

faces: distal face (Figure 10C— F) with (12 — )14 — 16

crowded areolae across diameter, up to 5,0 /zm wide, some

adjacent areolae, especially toward the centre, incomplete-

ly separated, walls irregular, with raised papillae at nodes;

proximal face (Figure 10A, B) with triradiate mark distinct,

sprinkled with fine granules, each tacet with ± 50 small

areolae, sometimes incomplete, walls low, often granulate.

R. namaquensis has been named for the region, where

it appears to be fairly common. Apparently it has a quite

wide ecological tolerance as it grows at seepages as well

as in drier areas on shallow soil at the edge of granite

FIGURE 8. — Riccia namaquensis. Morphology and anatomy. A. thal-

lus growing at seepage; B. thallus from drier area; C. thallus dry;

D, transverse section through dorsal cell pillars and scales; E.

transverse section through branch; F, scale. A, S.M. Perold 2136 ;

B, S.M. Perold 2036: C, S.M. Perold 1420: D. E. S.M. Perold

565: F. S.M. Perold 1832. Scale bar on A-C. E = 1 mm;

D = 50 /zm; F = 100 /zm.

182

Bothalia 20,2 (1990)

FIGURE 9. — Riccia namaquensis. Morphology and anatomy. A, thalli in cultivation; B, branches from above; C, scales at apex, seen from the

side; D, groove and scales at apex; E, dorsal pillars from above; F, dorsal pillars seen partly from the side. A — F, S.M. Perold 2136.

A, by A. Romanowski; B— F, SEM micrographs. Scale bar on A— D = 1 mm; E. F = 50 /x m.

outcrops. A somewhat wide species concept has been

adopted to accommodate plants from these rather different

habitats (thallus from seepage (Figure 8A); thallus from

drier region (Figure 8B) ), and it is possible that more than

one species has been included in this treatment of R. nama-

quensis. Under wetter conditions and in cultivation, the

dorsal cell pillars are generally longer, the glaucous

green colour of the thallus turns to bright green and in

cross section, the flanks slope obliquely upward and

outward. The reticulate spore ornamentation lacks a

distinctive pattern, and is also quite variable. Specimens

placed here, have dorsal epithelial pillars intermediate in

length between the lower pillars with bulging cells in R.

furfuracea and R. concava and the taller pillars with ‘tine’

FIGURE 10. — Riccia namaquensis. Spores. A, proximal face; B, proximal face, side view; C, F, distal face; D, distal face partly in side view;

E, areolae and wing. A-F, S.M. Perold 1420. A-E, SEM micrographs; F, LM photograph. Scale bar on A-E = 50 /xm; diameter of

spore on F + 75 fx m.

Bothalia 20,2 (1990)

183

cells in R. vitrea, although there is some overlap in length

at the lower limits of R. vitrea ; fortunately, the billowing

scales of R. vitrea help to distinguish it. Other species from

this region, which have pillars of ‘intermediate’ length,

are R. albomarginata Bisch. ex Krauss, not sensu Sim (see

Perold 1990a), which often turns brownish on drying and

has spores with coarse, radiating ridges on the distal face,

and R. parvo-areolata (Volk & Perold 1984) which has

narrowly winged spores with numerous small areolae and

concave thalli when dry. R. villosa and R. hirsuta (Volk

& Perold 1986b, 1990) have taller dorsal pillars and

triangular scales; R. alatospora (Volk & Perold 1985) and

R. hantamensis (Perold 1989b) have short, tapering pillars

and ornately ornamented, wide-winged spores.

SPECIMENS EXAMINED

CAPE. — 2918 (Gamoep): Hester Malan Res., Carolusberg, near old

mine, seepage area at flat granitic rocks (— CA), S.M. Perold 1420 (holo.),

1421, 2030, 2036, 2037 p.p., 2039 (PRE). 3018 (Kamiesberg): 18 km

NE of Kamieskroon, 3—4 km after turnoff to Rooifontein, large rock

outcrops near roadside (— AA), S.M. Perold 1464, 2139 (PRE); 12 km

from Kamieskroon on road to Leliefontein, rock outcrop, dry (— AA),

S. M. Perold 2095 (PRE); on road between Kamieskroon and Leliefon-

tein, 5 km before Leliefontein, seepage area at rock outcrop on right

side of road (—AC), S.M. Perold 2096, 2102 (PRE); Studer’s Pass, 23

km NE of Garies, on disturbed clayey soil at seepage area (—AC), S.M.

Perold 1613-1616, 2130-2133 (PRE); 3 km N of Leliefontein on road

to Bovlei, flat rocks (—AC), S.M. Perold 2136, 2137 (PRE); 29 km SE

from Platbakkies on road to Kliprand, Farm Banke, at dry base of rock

(— BC), S.M. Perold 1557, 1558 (PRE); on road between Platbakkies

and Kliprand, 1 km along turnoff to Frummelbakkies, at rock outcrop

(— BC), S.M. Perold 1580 (PRE). 3119 (Calvinia): 2 km from Nieuwoudt-

ville, on road from Van Rhyn’s Pass, on mud in ditch at roadside (—AC),

S.M. Perold 1753\ on dry soil between sandstone rocks above ditch

(-AC), S.M. Perold 1756 (PRE). 3219 (Wuppertal); 21-22 km S of Al-

geria Forest Station near ruins of buildings, at edge of sandstone rocks

(-AC), S.M. Perold 564, 565, 2372-2374, 2375-2377 ( PRE).

ACKNOWLEDGEMENTS

The author wishes to thank Dr H.F. Glen for the Latin

diagnoses; Ms J. Kimpton, artist, Mrs A. Romanowski,

photographer, and Mrs J. Mulvenna, typist, for their

contributions to this paper.

REFERENCES

BORNEFELD, T. 1989. The Riccia species of S and SW Africa.

Chromosome numbers and composition of the chromosome sets.

Nova Hedwigia 48: 371-382.

DEWEY, R.M. 1988. Electrophoretic studies in Riccia subgenus Riccia

(Hepaticopsida: Ricciaceae). The Bryologist 91: 344—353.

PEROLD, S.M. 1989a. Studies in the genus Riccia (Marchantiales) from

southern Africa. 11. Riccia montana and R. alboporosa, a further

two new white-scaled species of the group ‘Squamatae’. Bothalia

19: 9-16.

PEROLD, S.M. 1989b. Studies in the genus Riccia (Marchantiales) from

southern Africa. 13. A new species, R. hantamensis in section

Pi lifer, and a new record for R. alatospora. Bothalia 19: 157—160.

PEROLD, S.M. 1989c. Studies in the genus Riccia (Marchantiales) from

southern Africa. 14. R. concava section Pilifer. Bothalia 19:

161-165.

PEROLD, S.M. 1990a. Studies in the genus Riccia (Marchantiales) from

southern Africa. 16. R. albomarginata and R. simii, sp. nov.

Bothalia 20: 31—39.

PEROLD, S.M. 1990b. Studies in the genus Riccia (Marchantiales) from

southern Africa. 17. Three new species in section Pilifer: R.

elongata, R. ampullacea and R. trachyglossum. Bothalia 20:

167-174.

PEROLD, S.M. 1990c. Studies in the genus Riccia (Marchantiales) from

southern Africa. 19. Two new species: R. pulveracea, section

Pi life/ and R. bicolorata, section Riccia, group ‘Squamatae’.

Bothalia 20: 185-190.

VOLK, O H. 1983. Vorschlag fur eine Neugliederung der Gattung Riccia

L. Mitteilungen der Botanischen Staatssammlung, Munchen 19:

453-465.

VOLK, O.H. & PEROLD, S.M. 1984. Studies in the genus Riccia

(Marchantiales) from the south-west Cape. Bothalia 15: 117 —

124.

VOLK, O.H. & PEROLD, S.M. 1985. Studies in the genus Riccia

(Marchantiales) from southern Africa. 1. Two new species of the

section Pilifer: R. duthieae and R. alatospora. Bothalia 15:

531-539.

VOLK, O.H. & PEROLD, S.M. 1986a. Studies in the genus Riccia

(Marchantiales) from southern Africa. 3. R. schelpei, a new

species, in the new subgenus Chartacea. Bothalia 16: 19—33.

VOLK, O.H. & PEROLD, S.M. 1986b. Studies in the genus Riccia

(Marchantiales) from southern Africa. 6. R. hirsuta, a new species

in a new section. Bothalia 16: 187—191.

VOLK, O.H. & PEROLD, S.M. 1990. Studies in the genus Riccia

(Marchantiales) from southern Africa. 15. R. hirsuta and R.

tomentosa, sp. nov., two distinct species previously treated as

one. Bothalia 20: 23—29.

Bothalia 20,2: 185-190 (1990)

Studies in the genus Riccia (Marchantiales) from southern Africa. 19.

Two new species: R. pulveracea , section Pilifer and R. bicolorata ,

section Riccia , group ‘Squamatae’

S.M. PEROLD*

Keywords: Marchantiales, Riccia bicolorata, R. pulveracea, section Pilifer, section Riccia, southern Africa, 'Squamatae', taxonomy

ABSTRACT

R. pulveracea, specimens of which were collected by Duthie and tentatively referred to R. concava by her, is described

here, following the recent collection of fresh material. This species is distinguished from other members of section Pilifer

(Volk 1983) by low, generally two-celled, free-standing dorsal cell pillars, which when dry, appear powdery, hence the specific

epithet.

R. bicolorata, section Riccia, group 'Squamatae' occurs in the Cape but is rarely collected, and is characterized by bicoloured

scales, of which the wide hyaline margins are heavily encrusted with calcium deposits. It is somewhat similar to R. pott-

siana, but larger, and its scales are not so regularly arranged.

UITTREKSEL

R pulveracea, waarvan voorbeelde deur Duthie versantel en tentatief na R concava verwys is, word hier beskryf nadat

vars materiaal onlangs versamel is. Hierdie spesie word van die ander lede van seksie Pilifer (Volk 1983) onderskei deur

kort, vrystaande, tweesellige pilare wat in die droe toestand poeieragtig voorkom, vandaar die spesifieke epiteton.

R bicolorata wat tot die groep 'Squamatae', seksie Riccia behoort, kom in die Kaap voor maar word selde versamel en

word gekenmerk deur tweekleurige skubbe, waarvan die wye, hialiene rande bedek is met 'n neerslag van kalsiumsoute.

Dit kom in 'n mate ooreen met R. pottsiana, maar is groter, en die skubbe is nie so reelmatig gerangskik nie.

INTRODUCTION

Several specimens of the Riccia species, here described

as R. pulveracea , were collected at Bloemfontein by

Duthie, more than 50 years ago. She provisionally referred

them to R. concava Bisch. ex Krauss, but, as mentioned

in a previous paper (Perold 1989c), Duthie and Garside

never published anything on R. concava. Moreover, the

specimens do not form part of the main Riccia collection

at BOL, where the packets are mounted on herbarium

sheets. Instead, they are stored in boxes, 125 x 100 x

30 mm, mounted with glue on loose pieces of cardboard

and annotated by Duthie in pencil. On examination, the

dorsal cells could not be revived and measured, as was

to be expected, but spores were collected and repeatedly

photographed with SEM. Only with the collection,

cultivation and study of fresh gatherings and by

comparison of their spore ornamentation (fortunately quite

a useful character in this particular instance) could the old

Duthie specimens be identified and referred to the new

species, R. pulveracea.

R. bicolorata , the other species newly described in this

paper, can be recognized by bicoloured scales, their hyaline

margins heavily encrusted with calcium deposits, which

also cover the unistratose cells on the dorsal surface of

the thalli. It is rarely collected in the north-western,

southern and central Cape Province and is rather similar

to R. pottsiana Sim (1926), but larger, and the scales are

not so regularly arranged.

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

MS. received: 1989.11.27.

1. Riccia pulveracea Perold, sp. nov., a R. furfuracea

statura minore squamis parum minoribus, sporibus magis

granularibus et distributio geographica differt.

Thallus ?dioicus, perennis, gregarius, subparvus vel

magnitudine medius, viridis vel laete tlavo-virens, ramis

simplicibus vel semel vel bis furcatis, usque ad 6 mm

longis, 1,1 — 1,3( — 1,5) mm latis, parum latioribus quam

crassis, lingulatis, apice rotundatis. Squamae rotundatae,

imbricatae, hyalinae, cellulis nonnullis remotis purpureis,

apicem versus conspicuis. Columnae epithelii dorsalis

quaeque e cellulis ± 2 parvis compositae, 70-105 pm

longae, liberae, cellula apicali globosa vel valde mammil-

losa. Sporae (75— )80 — 87(— 93) gm diametro, triangulo-

globulares, polares, alatae, superficie distali areolis ±

12—14 forma subirregulari in diametro 2,5— 5,0(— 7,5) gm

latis, saepe incompletis; superficie proximali nota triradiali

distincta vel indistincta, areolis multis incompletis,

reticulum formantibus parietibus granularibus vel verrucu-

losis.

TYPE.— Cape, 3124 (Hanover): 18 km from Noupoort,

on road to Hanover, at bottom of slope, on ground between

bushes; false upper Karoo (-BB), Smook 3339 (PRE,

holo.; F, syn.).

Thallus ?dioicous, perennial, in gregarious patches

(Figure 2A), green to pale yellowish green, dorsal cells

inflated to collapsed proximally, scales hyaline and

conspicuous toward apex; smallish to medium-sized,

branches simple or once, occasionally twice, symmetri-

cally or asymmetrically furcate, medium to widely

divergent, up to 6 mm long, apical segments 2,5— 3,0 x

186

Bothalia 20,2 (1990)

1,1 — 1,3( — 1,5) mm, 0,9 mm thick, i.e. slightly wider to

V'2 times wider than thick in section (Figure IE),

ovate to lingulate, apex rounded (Figure 1A), slightly

emarginate, dorsal groove deep and sharp at apex (Figure

2B), soon shallow and wide, margins subacute, flanks

steep, green; ventral face rounded, green; when dry, rather

FIGURE 1 — Riccia pulveracea. Morphology and anatomy. A, thallus

wet and turgid; B, thallus, dry; C, dorsal pillars and air pores

seen from above; D, transverse section through dorsal cell pillars;

E, transverse section through branch; F, scale. A — F, Smook 6962c.

Scale bars on A, B, E = I mm; C, D = 50 /xm; F = 100 ^m.

concave, margins erect to indexed (Figure IB), sometimes

meeting in middle, revealing hyaline scales.

Anatomy of thallus'. dorsal epithelium (Figures ID;

2D-F) consisting of low, free-standing, mostly 2-celled,

hyaline pillars, 70—105 /xm long, + % the thickness of

thallus in section, apical cells globose to markedly mam-

millose, small, 35 — 55 x 37—42 /xm, basal cells 35-47

x 37— 52 /xm, soon collapsing and regular arrangement

of cells in underlying, rather compact assimilation tissue

then clearly visible, air pores (Figure 1C) mostly 4-sided,

small, ± 10 /xm wide; assimilation tissue 300-400 /xm

thick, less than '/2 the thickness of thallus in section,

consisting of vertical columns of 8—10 cells, (25— )32— 46

x 30-37 /xm, enclosing narrow, mostly 4-sided air canals;

storage tissue occupying ventral '/2 of thallus, + 450 /xm

thick, cells angular, 45—55 /xm wide; rhizoids 17—22 /xm

wide, some smooth, others tuberculate. Scales (Figure IF),

almost semilunar, margins mostly smooth, 750 —925 x

400—600 /xm, projecting 100—200 /xm above thallus

margins, and conspicuous toward apex (Figure 2C),

imbricate, wavy, hyaline, sometimes basal and scattered

cells higher up reddish purple, cells in body of scale

long-hexagonal, 50 — 65( — 80) x 25—35 /xm, marginal row

smaller, some brick-shaped. Antheridia in one or two

rows along middle of thallus, necks yellowish brown at

base, 110—200 /xm long. Archegonia with purple necks,

scattered. Sporangia 3 or 4 in a row, bulging dorsally, over-

lying tissue apparently remaining intact for some time, be-

fore thinning and disintegrating to liberate the spores, each

containing ± 470 spores. Spores (75-)80— 87(— 92) /xm

in diameter, triangular-globular, polar, light brown to

greyish brown, semitransparent to nearly opaque; wing

thin, rather undulate, width somewhat variable, 5, 0-7,5

/xm wide, broader at perforated marginal angles, margin

+ smooth; ornamentation different* on two spore faces:

distal face (Figure 3D— F) with + 12-14, rather irregu-

larly shaped areolae across diameter, 2,5— 5,0(— 7,5) /xm

wide, cross walls often incomplete and adjacent areolae

confluent, sometimes with thick knotted loops, or with

sinuating to shortly radiating ridges; proximal face (Figure

3A, B) with apex rather blunt, triradiate mark distinct to

indistinct, quite heavily sprinkled with granules, each facet

(Figure 3C) with numerous small, incomplete and rather

poorly defined areolae forming an open network, with low,

granular to verruculose walls.

R. pulveracea can be recognized by the low, generally

2-celled, dorsal pillars, with the top cell often markedly

mammillose when fresh and turgid; when dry, these cell

pillars collapse and form a fine, somewhat powdery

covering over the light green to yellowish green thallus.

The dorsal cells are arranged in free-standing pillars which

are not always obvious, especially in dry material. This

is demonstrated by Volk’s notes found with Duthie 5455

and 5484 ; ‘non R. concava. Epidermis (^epithelium)*

Reihen von Zeilen’. It appears that Volk was referring to

the regular rows of cells in the assimilation tissue under-

lying the + irregular pillars which had collapsed.

The proximal spore face is quite coarsely granular and

rather similar to that of R. sorocarpa Bisch. Occasionally

the distal face has 3—5 short radiating ridges. This may

* (= epithelium) added by SMP, in accordance with Volk 1983.

Bothalia 20,2 (1990)

187

FIGURE 2. — Riccia pulveracea. Morphology and anatomy. A, thalli in cultivation; B, branch seen from above; C, scales at apex; D, ± globular

apical cells; E, F, mammillose apical cells. A— F, Smook 6962c. A, by A. Romanowski; B— E, SEM micrographs. Scale bars on A— C

= 1 mm; D— F = 50 pm.

have prompted Duthie, who collected specimens of this

species near Bloemfontein more than 50 years ago, to refer

them to R. concava. Frequently the latter also has radiating

ridges, but the proximal spore face has numerous areolae

and is hardly granular; the dorsal cell pillars on the thallus

are taller, and the generally larger thalli often acquire a

mauve tinge on exposure to intense sunlight. Its distribu-

tion is also different (Perold 1989c). R. furfuracea Perold

(1990b) also has rather low dorsal cell pillars and is

somewhat similar to R. pulveracea, but here the spore

ornamentation is far less granular on the proximal face

and the distal face is usually marked with a central cross.

So far, R. pulveracea is known only from the Orange

Free State and central and eastern Cape (Figure 4) with

mostly summer rainfall of 200—800 mm p.a. It grows on

FIGURE 3. — Riccia pulveracea. Spores. A, B, proximal face; C, facet on proximal face; D, distal face with short radiating ridges; E, F, distal

face with areolae. A, D, Duthie 5484\ B, E, Smook 6962c\ C, Duthie 5461a\ F, Van Rooy 2598. A-E, SEM micrographs; F, LM photo-

graphs. Scale bars on A— E = 50 /im; diameter of spore on F = +85 pm.

188

Bothalia 20,2 (1990)

FIGURE 4. — Distribution map of R. pulveracea, • : and R. bicolorata,

□ , in southern Africa.

alkaline soil in between karroid bushes, in association

with other Riccia species, such as R. nigrella DC., R.

albornata Volk & Perold and occasionally with R. simii

Perold (1990a) [identified by Duthie as R. albomarginata

in Duthie 5461a, in acceptance of Sim’s misapplication of

the name]. Smook 6962 is a mixed gathering of R.

runssorensis Steph., R. nigrella DC., R. bicolorata sp.

nov. and R. pulveracea sp. nov. ; Duthie 5484 was mixed

with R. okahandjana S. Arnell (1963).

The name R. pulveracea was chosen because of the

somewhat powdery appearance of the dorsal surface in dry

thalli, due to the collapse of the low epithelial cell pillars.

SPECIMENS EXAMINED

O.F.S. — 2926 (Bloemfontein): Bloemfontein (— AA), Duthie 5455,

5461a, 5484, 5485, 5498 (BOL); Potts PRE-CH 1047. 3026 (Aliwal

North): Farm Olievenrand near Elandsberg, between Zastron and

Wesselsdale, mountain slopes, southern aspect (— BB), Van Roov 2451

(PRE).

CAPE. — 3027 (Lady Grey): 23 km S of Lady Grey, between farms

Rietfontein and De Kraal, sandstone outcrops in grassland (-CC), Van

Roov 2598 (PRE). 3123 (Victoria West): Farm Rietpoort, 34 km N of

Victoria West, on main road, flat top of koppie in damp area, with karoo

bushes (-AA), Smook 6962c (PRE). 3124 (Hanover): 15 km from

Noupoort on road to Hanover (—BA), Herman 549 p.p. (F; PRE); 18

km from Noupoort on road to Hanover, at bottom of slope, on soil between

karoo bushes ( — BB). Smook 3339 (F, syn.; PRE, holo.).

2. Riccia bicolorata Perold, sp. nov., R. pottsianae

Sim aliquanto similis sed statura majore squamisque minus

regulatim dispositis differt.

Thallus monoicus, perennis, gregarius vel in rosulis

partialibus, aliquantum parvus, flavo-virens vel albo-

virens, calcio incrustatus, ramis semel vel bis furcatis,

usque ad 4,5 x 1,0 — 1,3( — 1,5) mm, in sectione sesquiplo

vel ± duplo latioribus quam crassis, obovatis vel

ovatis, apice rotundatis. Squamae rotundatae, imbricatae,

bicolores (inde nomen specificum), basin versus atro-

purpureae, margine albo, calcio incrustato, basi purpureae

adhaerentes marginem atrum interruptum secus margines

thalli facientes. Epithelium dorsale unistratosum, cellulis

globosis vel conicis vel mammillosis, cito collabentibus.

Sporae (77— )85— 90(— 93) pm diametro, polares, alatae,

superftcie distali areolis 10 incompletis in diametro,

5,0— 7,5 )tm latis, papillis e nodis parietis eminentibus;

FIGURE 5. — Riccia bicolorata. Morphology and anatomy. A, thallus

wet; B, thallus dry; C, dark margin of thallus and projecting scales

seen from above; D, dorsal cells and air pores from above; E,

transverse section through dorsal epithelium and some assimilation

cells; F, transverse section through branch. G, scale. A— G, Smook

6990a. Scale bars on A, B, F = 1 mm; D, E = 50 /im; C, G

= 100 jim.

Bothalia 20,2 (1990)

189

superficie proximali nota triradiali sub-indistincta, areolis

incompletis parietibus granularibus.

TYPE. — Cape, 3123 (Victoria West): Central Karoo,

Farm Kalkfontein, 48,6 km NE of Victoria West, on flat

gravel plain, common in damp areas around bushes

(— AA), Smook 6990a (PRE, holo.).

Thallus monoicous, perennial, in gregarious patches

(Figure 6A), or in partial rosettes 8-10 mm across, green

to yellowish green, or whitish green, encrusted with

calcium deposits, scales mostly bicoloured, white and

purple; rather small, once or twice symmetrically furcate,

when young, primal branches closely associated, butterfly-

shaped (Figure 6B), but often tearing apart along middle

as growth continues and then mostly asymmetrically

furcate (Figures 5A; 6C), branches moderately divergent,

up to 4,5(— 5,0) mm long, terminal segments 1,0-2, 0 x

1,0 — 1,3( — 1,5) mm, 0,6— 0,8 mm thick, i.e. l'/2 times

to nearly twice wider than thick in section (Figure

5F), obovate to ovate, apex rounded, emarginate, groove

sharp and deep apically (Figure 6D), soon wide and

shallow; margins subacute, flanks steep to sloping

slightly obliquely, ventrally rounded, green; when dry

(Figure 5B), dorsally concave, margins raised or incurved,

sometimes indexed, flanks covered with imbricate,

appressed, calcium-encrusted, bicoloured scales, often

appearing ‘striped’.

Anatomy of thallus: dorsal epithelium (Figures 5D, E;

6E, F) unistratose, hyaline, cells inflated in and near

groove (Figure 6D, E), globose to conical or mammillose,

25-55 x 30-42 /im, soon collapsing and often becoming

covered with fine deposits of calcium salts; air pores

4—5-sided, rarely triangular, small in groove (Figure 6E),

15—25 pm wide, rapidly widening to ± 60 /xm (Figures

5D; 6F); assimilation tissue 280—350 )im thick, ±

!/3 — '/2 the thickness of thallus, consisting of vertical

columns of 6— 8(— 10) chlorophyllose cells, up to 45 x

42 p m, enclosing air canals, 37—75 pm wide, widening

toward margins; storage tissue occupying remaining

'/2-2/3 the thickness of thallus, cells ± 55 /on wide;

rhizoids 17—22 /im wide, some smooth and others

tuberculate. Scales rounded, 500 x 300 pm, appressed,

to slightly wavy, imbricate (Figure 6C), generally

bicoloured with a wide white margin encrusted with

calcium deposits and a deep purple, shiny base, the

adherent purple bases forming an interrupted dark border

along thallus margins (Figure 5C), cells in body of scale

short-hexagonal, up to 62 x 42 /un, marginal row some-

what smaller. Antheridia in a row along midline, hyaline

necks arising from small pits. Archegonia with purple

necks scattered. Sporangia toward base, single or in pairs,

adjacent or serially arranged, bulging dorsally, containing

180-190 spores each. Spores (77-)85— 90(— 93) /im in

diameter, triangular-globular, polar, light brown to brown,

semitransparent, wing ± 5 /im wide, thin, slightly

undulating, notched or perforated at marginal angles,

margin smooth; ornamentation reticulate, rather different

on the two spore faces: distal face (Figure 7C-F) with

± 10 incomplete areolae across the diameter, 5,0— 7,5 pm

wide, cross walls often undeveloped and radial walls

thickened, fading out toward margin, papillae projecting

from the nodes, especially over the centre; proximal face

(Figure 7A, B) with triradiate mark rather poorly defined,

dotted with granules, facets with incomplete areolae, walls

sprinkled with granules and raised into papillae at the

nodes.

R. bicolorata can be distinguished from other members

of section Riccia , group ‘Squamatae’ (Na-Thalang 1980),

by the appressed, bicoloured scales for which it has been

named, by the dark broken line along the thallus margins,

formed by the adherent purple bases of the scales, and by

FIGURE 6.— Riccia bicolorata. Morphology and anatomy. A, field-grown thalli; B, young branches in ‘butterfly’ shape; C, older branches; D,

apex with groove; E, dorsal cells in groove; F, inflated dorsal cells and pores. A, C, D, F, Koekemoer 300 ; B, E, S.M. Perold 1772a.

Scale bars on A— D = 1 mm; E, F = 50 /im.

190

Bothalia 20,2 (1990)

FIGURE 7. — Riccia bicolorata. Spores. A, proximal face; B, proximal face, side view; C, F, distal face; D, distal face, side view; E, ridges

and areolae toward margin. A— F, Smook 6990a. A— E, SEM micrographs; F, LM photograph. Scale bars on A— E = 50 /xm; diameter

of spore on F = ±85 p m.

its habit, as described above. It is somewhat similar to R.

argenteolimbata Volk & Perold, but the latter has a more

compact thallus, triangular pores and apolar spores; R.

albolimbata S. Arnell and R. albomata Volk & Perold are

larger plants with wavy scales (Volk et al. 1988);

R. montana Perold (1989a) has ligulate branches, and

R. alboporosa Perold (1989a) has a spongy appearance.

R. pottsiana is another species which bears some similarity

to R. bicolorata, but it is smaller and its dark red scales

are more regularly arranged.

R. bicolorata is known from a few collections in north-

western, central, southern and eastern Cape (Figure 4),

where it grows on alkaline soil, sometimes in association

with R. alboporosa, R. albomata and with R. pulveracea.

SPECIMENS EXAMINED

CAPE. — 2917 (Springbok): south of Springbok on road to

Kamieskroon, 36 km from Kokerboom Motel, on damp soil at granitic

rocky outcrop, on left side of road (— DD), S. M. Perold 1443 (PRE).

3025 (Colesberg): 5 km from Colesberg on road to Steynsburg, next

to river on soil under thicket, moist and shady (— CA), Smook 3215a

(F, PRE). 3119 (Calvinia): S of Loeriesfontein, Skietnes Kloof, E of Slag-

berg, rocky ledges facing south (— AB), Oliver 8849 p.p. (PRE);

Groothoek, 18 km along dirt road to Rondekop, at Soetlandsfontein River

below drift, on brackish soil between tillite rocks (-AD), S.M. Perold

1772a, 2318 (PRE). 3123 (Victoria West): Farm Kalkfontein, 48,6 km

NE of Victoria West, on flat, gravel plain, in damp areas between karoo

bushes, (— AA), Smook 6990a (PRE, holo.). 3321 (Ladismith): 8 km

from Ladismith, on road to Calitzdorp (-AD), Koekemoer 300 (PRE).

3421 (Riversdale): old bridge across Gourits River (-BA), S.M. Perold

2554 (PRE).

ACKNOWLEDGEMENTS

The author wishes to thank the curator of BOL for the

loan of specimens; Dr H.F. Glen for the Latin diagnoses;

Mrs L. Fish (nee Smook), Miss M. Koekemoer and

Messrs E.G.H. Oliver, P. Herman and J. van Rooy for

kindly collecting specimens. Sincere thanks to Mrs A.

Romanowski for developing and printing the micrographs;

to the artist, Ms J. Kimpton and to Mrs J. Mulvenna for

typing the manuscript. Sincere gratitude is expressed to

Dr E.O. Campbell, Massey University, New Zealand, and

Prof. Dr S. Jovet-Ast, Museum National de’Histoire

Naturelle, Paris, for refereeing this paper.

REFERENCES

ARNELL, S. 1963. Hepaticae of South Africa, pp. 441. Swedish National

Scientific Research Council, Stockholm.

NA-THALANG, O. 1980. A revision of the genus Riccia (Hepaticae)

in Australia. Brunonia 3: 61—140.

PEROLD, S.M. 1989a. Studies in the genus Riccia (Marchantiales) from

southern Africa. 11. Riccia montana and R. alboporosa, a further

two new white-scaled species of the group ‘Squamatae’. Bothalia

19: 9-16.

PEROLD, S.M. 1989b. Studies in the genus Riccia (Marchantiales) from

southern Africa. 12. Riccia albolimbata and the status of/?, al-

bosquamata, white-scaled species originally described by Arnell.

Bothalia 19: 17—25.

PEROLD, S.M. 1989c. Studies in the genus Riccia (Marchantiales) from

southern Africa. 14. R. concava, section Pilifer. Bothalia 19:

161-165.

PEROLD, S.M. 1990a. Studies in the genus Riccia (Marchantiales) from

southern Africa. 16. R. albomarginata and R. simii, sp. nov.

Bothalia 20: 31—39.

PEROLD, S.M. 1990b. Studies in the genus Riccia (Marchantiales) from

southern Africa. 18. New species in section Pilifer from the NW

Cape: R. furfuracea, R. vitrea and R. namaquensis. Bothalia 20:

175-183.

SIM, T.R. 1926. The bryophyta of South Africa. Transactions of the Royal

Society of South Africa 20: 15—17.

VOLK, O.H. 1983. Vorschlag fur eine Neugliederung der Gattung Riccia

L. Mitteilungen der Botanischen Staatssammlung, Munchen 19:

453-465.

VOLK, O.H., PEROLD, S.M. & BORNEFELD, T. 1988. Studies in

the genus Riccia (Marchantiales) from southern Africa. 10. Two

new white-scaled species of the group ‘Squamatae’: R. argenteo-

limbata and R. albomata. Bothalia 18: 155—163.

Bothalia 20,2: 191-196 (1990)

Studies in the genus Riccia (Marchantiales) from southern Africa. 20.

R. albovestita and its synonyms, R. duthieae and R. sarcosa

S. M. PEROLD*

Keywords: Marchantiales, Riccia albovestita, R. duthieae, R. sarcosa, Riccia section Pilifer, southern Africa, taxonomy

ABSTRACT

As has been repeatedly emphasized elsewhere, species in section Pilifer Volk are often very difficult to distinguish (Perold

1990b, 1990c). This is nowhere better illustrated than in the case of R. duthieae Volk & Perold and R. sarcosa Volk &

Perold, both initially described as new species, which are now regarded by me as synonyms of R. albovestita Volk. Comparisons

are drawn between the thallus dimensions, dorsal cell pillars, spore ornamention and other characters, in support of the

above conclusion. An updated distribution map and a list of the specimens examined are added, because several new collections

have been made since the original publications.

UITTREKSEL

Soos elders herhaaldelik beklemtoon, is spesies in seksie Pilifer Volk dikwels baie moeilik om te onderskei (Perold 1990b,

1990c). Dit word veral duidelik gel llustreer in die geval van R. duthieae Volk & Perold en R. sarcosa Volk & Perold. albei

oorspronklik as nuwe spesies beskryf, wat ek nou as sinonieme van R. albovestita Volk beskou. Vergelykings word tussen

tallusafmetings, dorsale selpilare, spoorornamentasie en ander kenmerke gemaak ter ondersteuning van bogenoemde

gevolgtrekking. ’n Nuwe verspreidingskaart en 'n lys van die eksemplare wat ondersoek is, word ingesluit, aangesien verskeie

nuwe versamelings sedert die oorspronklike publikasies gemaak is.

R. albovestita

R. albovestita was described from Namibia by Volk

(1981). It was only the fourth species in the by now quite

large, endemic section he was later to name ‘Pilifer’

(Volk 1983), in reference to the free-standing dorsal cell

pillars. Volk’s early collections of this species (Volk 12458

p.p. and 12462) had been identified by Arnell (1957, 1963)

as R. albomarginata (Volk 1981), a name which had been

generally misapplied since Sim (Perold 1990a). Volk (1981)

distinguished R. albovestita from R. simii Perold (= albo-

marginata sensu Sim), by the reticulate (not radiate)

ornamentation of the spores. His good spore drawings

(Abb. If) are clearly recognizable, but the SEM micro-

graphs (Abb.2) are perhaps less so, as the areolae on the

distal face are mostly complete and lack a central ‘knob’.

SEM micrographs of Volk 00484 (Figure 1A), 01164b

(Figure IB, D) and 12462 (Figure 1C, E) taken by me, show

low-walled or incomplete areolae, with irregularly raised

and thickened or toothed nodes on the proximal face; the

distal face of Volk 01164 has incomplete areolae, with only

the suggestion of a central ‘knob’; the two micrographs

of the distal face of Volk 12462 differ in that there are 10

incomplete areolae across the diameter in Figure IE and

only six on Figure IF. These spores were also illustrated

by Arnell (1963) but as being R. albomarginata spores

(Volk 1981).

The dorsal cells in the free-standing pillars of R. albo-

vestita as depicted by Volk, are of four cells, the top one

conical, with the others rounded and isodiametric, the

basal cell the same size as the others or smaller, but not

wider. An attempt to revive the dorsal cells achieved

limited success, yet it was found that some of the pillars

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

MS. received: 1989.11.27.

were tapering, with distinctly wider (up to 62 /tm) basal

cells.

Riccia duthieae

Subsequent gatherings, Volk 81-273, 81-274 from

Aberdeen in the central Cape, were tentatively also named

R. albovestita, but after further study, Volk became

convinced that this was yet another new species, later to

be named R. duthieae, although in litt. [5.5.83 (PRE)],

I had informed him of the close similarity between the

two species in the spore ornamentation on both faces, as

seen on SEM micrographs (Figures 1 & 2). Unfortunately,

I had no living material of R. albovestita from Namibia

to study and to compare the dorsal cell pillars with those

of the new collection. The cells of these pillars collapse

when dry (Volk & Perold 1984), and can hardly be recon-

stituted in herbarium material, in order to examine their

shape and size, which are very important taxonomic

characters.

R. duthieae was described (Volk & Perold 1985) as

bearing a close resemblance to R. albovestita and R. parvo-

areolata, but was thought to differ from them by having

3 (or rarely 4) cells in the dorsal pillars (Figure 3F), as

opposed to their 4 (or rarely 5)-celled pillars, and by

having a different spore ornamentation. It has since

become evident that the number of cells in the pillars and

their shape can vary within a species. Generally, however,

the pillars in these three species are relatively short and

tapering. The proximal face of the spores of R. duthieae

(Volk 81-273) (Figure 2 A) bears a strong resemblance to

that of R. albovestita (Volk 00484, 01164) (Figure 1A &

B); the distal face (Figure 2B, F) has fewer complete

areolae than R. albovestita, but the larger, central ones

contain a ‘knob’ as shown in the micrograph of S. M. Perold

1347 (Figure 2E). Two micrographs of less mature spores

192

Bothalia 20,2 (1990)

FIGURE 1 —Riccia albovestita. Spores. A, B, proximal face; C, triradiate mark; D-F, distal face. A, Volk 00484 ; B, D, Volk 01164b (type);

C, E, F, Volk 12462. A-E, SEM micrographs; F, LM photograph. Scale bar on A— E = 50 ftm; diameter of spore on F, ± 70 ^m.

of Smook 4036 (Figure 2C, D) (placed under R. duthieae)

have been included for purposes of comparison with R.

sarcosa spores.

R. sarcosa

Volk isolated specimens of the white-margined R.

sarcosa from a mixed gathering of R. simii Perold (= R.

albomarginata sensu Sim), Volk 81-292 , and from cultures

of R. duthieae, Volk 81-274. A sterile specimen, J.M.

Perold 35, was thought to also belong here. At that time,

it was not realized that with prolonged growth, the some-

what attenuate thallus margins of this species habitually

turn white (Figure 3A), nor that we might be dealing with

a widespread species (Figure 4) which was bound to show

some variation.

FIGURE 2. — Riccia duthieae. Spores. A, proximal face; B, distal face; C, young spores still in tetrads; D, distal face of young spore; E, F, distal

face. A, B, F, Volk 81-273 ; C, D, Smook 4036\ E, S.M. Perold 1347. A-E, SEM micrographs; F, LM photograph. Scale bar on A-E

= 50 /im; diameter of spore on F, ± 70 ^m.

Bothalia 20,2 (1990)

193

FIGURE 3. — Riccia albovestita and its synonyms. Morphology and anatomy. A, field-grown thalli, margins white; B, mature thalli in cultivation,

margins less markedly white; C, ‘open’ thallus with ventral scales not extending beyond thallus margins; D, thallus with apical margins

indexed, revealing large scales; E, dorsal cells from above; F, dorsal cells in cross section. A, S.M. Perold 955; B, Smook 6583', C, E,

R. duthieae, Volk 81-273', D, F, R. sarcosa, Volk 81-274b. A, B, by A. Romanowski; C— E, SEM micrographs; E, LM photograph. Scale

bars on A— D = 1 mm; E, F = 50 /xm.

R. sarcosa was regarded (Volk & Perold 1986) as being

distinct, on account of the white margin of the thallus

(Figure 3A), its mostly inconspicuous scales that do not

project above the thallus margins, the dorsal cell pillars

which have inflated basal and smaller terminal cells and

the spore ornamentation with deep-set, ringed areolae. As

mentioned above, the attenuate thallus margins develop

a white colouration with prolonged growth, and this is no

longer regarded as a really reliable distinguishing

character. In all three species the scales are apically

prominent, especially in dry specimens, where the margins

are incurved (Figure 3D), but along the body of the thal-

lus they hardly project beyond the margins (Figure 3C).

The dorsal cell pillars are relatively low and tapering, with

a wider basal cell (Figure 3F). The R. sarcosa spores

that I originally examined and photographed by SEM (the

only ones available to me then) were not fully mature and

nearly flat, hence the ornamentation and even the size were

misleading; nevertheless, comparison between the figures

of Volk & Perold (1985: fig. 2.5 and 1986: fig. 2.5) show

some similarity. Micrographs of more mature spores of

R. sarcosa specimens, Volk 81-274b (Figure 5A, C, D, F)

and 81-2 92b (Figure 5B, E), clearly have similar proximal

faces and their distal faces, especially of Volk 81-274b

(Figure 5C, D), when compared with young spores of

Smook 4036 (Figure 2C, D), have much in common, and

toward the centre, the ornamentation also appears to be

in two different levels; my LM photograph of R. sarcosa

(Volk & Perold 1986: Figure 2.7) has eight areolae across

the diameter on the distal face, as in R. albovestita.

FIGURE 4.— Distribution map of Riccia albovestita in southern Africa.

DISCUSSION

It has previously been shown that thallus morphology

can vary considerably within a species and that the

environment plays an important part in determining thallus

colour, width and thickness (Pande 1924; Abeywickrama

1945). Scale morphology, particularly pigmentation and

size are also affected (Berrie 1975; Na-Thalang 1969, 1980),

which could account for the possibly somewhat larger, hya-

line scales in R. albovestita from Namibia with its hot,

rather dry climate.

Spore size may vary quite widely under different

environmental conditions (Seppelt 1974, 1983), which

pould perhaps be another reason for the larger size of

/?. sarcosa spores, collected from specimens cultured

under ideal circumstances. The ornamentation appears to

be relatively stable, although Duthie & Garside (1936) had

in densely crowded gregarious patches or in incomplete gregarious or in incomplete rosettes 20 mm across

rosettes 10—40 mm across

194

Bothalia 20,2 (1990)

Measurements not given in original description were subsequently taken and added in square brackets.

Bothalia 20,2 (1990)

195

FIGURE 5. — Riccia sarcosa. Spores. A, B, proximal face; C-F. distal face. A, C, D, F, Volk 8! -274b \ B, E, Volk 81-2 92b ; A-E, SEM micro-

graphs; F, LM photograph (slide lent by Volk). Scale bar on A-E = 50 jim; diameter of spore on F, ± 100 pm.

suggested that environmental differences could be

correlated with variation in the ornamention in spores of

R. crystalline i (= R. plana). On the other hand, I could

not convincingly prove or disprove the effect of the

environment on spore sculpturing (Perold 1989b), as this

would require controlled experiments using sophisticated

equipment.

The shape and size of the cells in the dorsal pillars can

vary from markedly tapering with a small conical top cell

and wide basal cell to being almost uniformly wide.

R. albovestita, R. duthieae and R. sarcosa [as well as

R. alatospora Volk & Perold and R. hantamensis Perold

(1989a)] are the only species in section Pi lifer with

relatively short, generally tapering dorsal pillars. In

R. villosa Steph. and R. simii (= R. albomarginata sensu

Sim), the dorsal pillars are long (300—400 pm) and

tapering, whereas in all other species in the section they

are of ± uniform width. In Table 1, a comparison of

R. albovestita , R. duthieae and R sarcosa is given.

Several more species belonging to section Pilifer have

been identified and described in the last few years (Perold

1990b, 1990c) and hence more experience has been gained

with the members of this section. The adoption of a

somewhat wider species concept is clearly called for.

In cases where no clear breaks occur in the range of

characters such as the size, shape and number of cells in

the dorsal pillars and the ornamentation of the spores, as

shown in the above three species, their continued separa-

tion cannot be satisfactorily maintained. All three should

therefore be treated under R. albovestita and the reduction

of R. duthieae and R. sarcosa to synonymy of R. albo-

vestita is made as follows:

Riccia albovestita Volk in Mitteilungen der Botan-

ischen Staatssammlung, Miinchen 17: 245 (1981). Type:

Bezirk Windhoek No. 85 (Rietfontein), Volk 01164b (M,

holo.!).

Riccia duthieae Volk & Perold in Bothalia 15: 531 (1985). Type: Cape,

3224 (Graaff-Reinet): Aberdeen, next to road R57, 2 km north-east

of junction with R61, at shallow edges of vleis temporarily damp or

occasionally inundated (—AC), 1981.04.11, Volk 81-273 (M, holo.!; PRE,

iso.!).

Riccia sarcosa Volk & Perold in Bothalia 16: 23 (1986). Type: Cape,

3224 (Graaff-Reinet): Aberdeen, next to road R57, 2 km north-east

of junction with R61. at shallow edges of vleis temporarily damp or

occasionally inundated (-AC), 1981.04.11. Volk 81-274b (M, holo.!; PRE,

iso.!).

SPECIMENS EXAMINED

S.W. A. /NAMIBIA. -2217 (Windhoek): WIN 85 Rietfontein, feuchter

Granitzersatz, rostfleckig (—CD), Volk 01164 (M). 2314 (Sandwich

Harbour): WIN 15 Baumgartsbrunn, Kuiseb-Ufer (— BC), Volk 00484

(M). 2417 (Mariental): GIB 18 Haribes, Ufer des Rietriviers, leicht

beschattet (—DA), Volk 12462 p.p. (M).

TRANSVAAL. — 2228 (Maasstroom): 16 km W from Tolwe, at edge

of vlei, under thornbush, isolated thalli (—CD), S.M. Perold 784 (PRE).

2628 (Johannesburg): 11 km SE of Bapsfontein on road to Delmas, bridge

over stream, at edge of flat rock outcrop (—BA), S.M. Perold 2463—2465

(PRE). 2629 (Bethal): 5 km W of Kriel on road to Vandyksdrift. at depres-

sion with flat rock outcrop (— AB), S.M. Perold 342, 2473 (PRE).

O.F.S. — 2726 (Odendaalsrus): Odendaalsrus, in town on edge of marsh

(—DC), Stnook 6583 (PRE). 2728 (Frankfort): Petrus Steyn, 5 km E

of town, next to farm road in ditch (— CA), M. Crosby 520 (PRE). 2826

(Brandfort): Glen College (—CD). Zietsman 943 (PRE). 2827 (Senekal):

Willem Pretorius Game Reserve (—AC), Volk 81-292 (M, PRE); Paul

Roux, on flat rocks behind town, at watertank (— BD), S.M. Perold 1347

(PRE); 43 km from Marquard, ‘Boompie-alleen’ Wilderness Trail (—CD),

Koekemoer 102. 103 p.p. (PRE); 22 km E of Clocolan on road from

Ficksburg, Farm Holstein, at flat rock outcrop near stream (—DC), S.M.

Perold 1317 (PRE). 2926 (Bloemfontein): 30 km S of Bloemfontein, shal-

low depression at roadside (—AC), S.M. Perold 955 (PRE). 2927

(Maseru): 10 km S of Ladybrand, at rock outcrop (— AB), J.M. Perold

35, 39, 44 (PRE); Thaba Putswa, between Hobhouse and Tweespruit;

rock outcrop on plateau ( — AC), Du Preez 2105 (PRE)

196

Bothalia 20,2 (1990)

10 km S of Clocolan on road to Ladybrand, Hillcrest Farm, at flat rocks

(—BA), S.M. Perold 1319 (PRE). 3026 (Aliwal North): Farm Olieven-

rand, nr Elandsberg, between Zastron and Wesselsdale, soil on plateau

(— BB), Van Rooy 2419 (PRE).

CAPE.— 2724 (Taung): Farm Zoetvlei, 50 km W of Vryburg and 6

km from turnoff, on road to Louwna, on soil nr edge of dam (— AA),

S.M. Perold 2026 (PRE). 3125 (Steynsburg): Steynsburg Dist., Kaffers-

kraal Farm, W of Teebus, rocky koppie, on moist, dark brown mud

(-BC), Retief & Germishuizen 218 (PRE). 3222 (Beaufort West): Beaufort

West Dist., Farm Rystkuil, along margin of small pan (— DB), Retief

& Reid 294 (PRE). 3224 (Graaff-Reinet): Aberdeen, next to road R57,

2 km NE of junction with R61, at shallow edge of vlei (-AC), Volk 81-272,

81-273, 81-274b, 84-646 (M, PRE). 3319 (Worcester): Robertson (— DD),

Duthie 5182 (BOL); at Robertson Golf Links (— DD), Duthie 5193 (BOL).

3326 (Grahamstown): between Ulster and Mooi River, on mud at edge

of small earth dam (—BA), Smook 4036 (PRE).

ACKNOWLEDGEMENTS

The author wishes to thank Prof, (emer.) Dr O.H. Volk

of Wurzburg University and also the Curator of BOL for

the loan of specimens.

REFERENCES

ABEYWICKRAMA, B. A. 1945. The structure and life history of Riccia

crispatula Mitt. Ceylon Journal of Science A12: 145—153.

ARNELL, S. 1957. Hepaticae collected in South West Africa by Prof.

Dr O.H. Volk. Mitteilungen der Botanischen Staatssammlung,

Munchen 16: 262—272.

ARNELL, S. 1963. Hepaticae of South Africa, pp. 411. Swedish Natural

Science Council, Stockholm.

BERR1E, G.K. 1975. The biology of a West African species of Riccia

L. Journal of Bryology 8: 443—454.

BORNEFELD, T. 1989. The Riccia species of S and SW Africa.

Chromosome numbers and composition of the chromosome sets.

Nova Hedwigia 48: 371—382.

DUTHIE, A.V. & GARSIDE, S. 1936. Studies in South African

Ricciaceae I. Transactions of the Royal Society of South Africa

24: 93-133.

NA-THALANG, O. 1969 .Studies in the Australian Marchantiales. The

genus Riccia. Ph.D. thesis, University of Sydney.

NA-THALANG, O. 1980. A revision of the genus Riccia (Hepaticae)

in Australia. Brunonia 3: 61—140.

PANDE, S.K. 1924. Notes on the morphology and life history of Riccia

sanguinea Kash. Journal of the Indian Botanical Society 4:

117-128.

PEROLD, S.M. 1989a. Studies in the genus Riccia (Marchantiales) from

southern Africa. 13. A new species, R. hantamensis, in section

Pilifer, and a new record for R. alatospora. Bothalia 19: 157-160.

PEROLD, S.M. 1989b. Spore-wall ornamentation as an aid in identifying

the southern African species of Riccia (Hepaticae). Journal of

the Hattori Botanical Laboratory 67: 109 —201.

PEROLD, S.M. 1990a. Studies in the genus Riccia (Marchantiales) from

southern Africa. 16. R. albomarginata and R. simii, sp. nov.

Bothalia 20: 31—39.

PEROLD, S.M. 1990b. Studies in the genus Riccia (Marchantiales) from

southern Africa. 17. Three new species in section Pilifer. R.

elongata, R. ampullacea and R trachyglossum. Bothalia 20:

167-174.

PEROLD, S.M. 1990c. Studies in the genus Riccia (Marchantiales) from

southern Africa. 18. New species in section Pilifer from the NW

Cape: R. furfuracea, R. vitrea and R. namaquensis. Bothalia 20:

175-183.

SEPPELT, R. 1974. Riccia crystallina in South Australia. The Bryologist

IT. 224-229.

SEPPELT, R. 1983. The status of Riccia areolata and R. longiciliata,

two recently described species. Lindbergia 9: 117—120.

VOLK, O.H. 1981. Beitrage zur Kenntnis der Lebermoose (Hepaticae)

aus Siidwestafrika (Namibia). II. Mitteilungen der Botanischen

Staatssammlung, Munchen 17: 245—252.

VOLK, O.H. 1983. Vorschlag fur eine Neugliederung der Gattung Riccia

L. Mitteilungen der Botanischen Staatssammlung, Munchen 19:

453-465.

VOLK, O.H. & PEROLD, S.M. 1984. Studies in the liverwort genus

Riccia (Marchantiales) from the south-west Cape. Bothalia 15:

117-124.

VOLK, O.H. & PEROLD, S.M. 1985. Studies in the genus Riccia

(Marchantiales) from southern Africa. 1. Two new species of the

section Pilifer. R. duthieae and R. alatospora. Bothalia 15:

531-539.

VOLK, O.H. & PEROLD, S.M. 1986. Studies in the genus Riccia

(Marchantiales) from southern Africa. 2. A new species of the

section Pilifer. R. sarcosa. Bothalia 16: 23—27.

Bothalia 20,2: 197-206 (1990)

Studies in the genus Riccia (Marchantiales) from southern Africa. 21.

R. stricta, R. purpurascens and R. fluitans , subgenus Ricciella

S. M. PEROLD*

Keywords: Marchantiales, Riccia fluitans, R. purpurascens, R. stricta, southern Africa, subgenus Ricciella, taxonomy

ABSTRACT

Lindenberg (1836) regarded and published Riccia stricta as a variety of R. fluitans L. Subsequently, Nees (1838) and Gottsche

et al. (1846) also treated it as a variety. Trevisan (1877) raised its rank and published the epithet, Ricciella stricta Trevis.

Ricciella is, however, regarded as a subgenus; Ricciella stricta is, therefore, transferred to Riccia stricta (Lindenb.) Perold.

It is described in detail and illustrated. R. purpurascens Lehm. & Lindenb., a related endemic species, is also more fully

described than before and illustrated. R. fluitans L. apparently does not occur naturally in southern Africa. As far as is

known, a single local specimen of it was introduced.

UITTREKSEL

Lindenberg (1836) het Riccia stricta as 'n varieteit van R. fluitans L. beskou en gepubliseer. Vervolgens het Nees (1838)

en Gottsche et al. (1846) dit ook as -n varieteit beskou. Trevisan (1877) het dit tot die rang van spesie verhef en het die

naam Ricciella stricta Trevis. gepubliseer. Ricciella word egter as ’n subgenus beskou; derhalwe word Ricciella stricta na

Riccia stricta (Lindenb.) Perold oorgeplaas. Dit word hier volledig beskryf en gei llustreer. R purpurascens Lehm. & Lindenb.,

’n verwante endemiese spesie, word ook meer volledig as voorheen beskryf en word gei llustreer. R. fluitans L. , kom skynbaar

nie natuurlik in suidelike Afrika voor nie. Sover bekend, is die enkele lokale voorbeeld daarvan ingevoer.

1. Riccia stricta (Lindenb.) Perold , comb. nov.

R. fluitans L. var. 5 stricta Lindenb., Monographic der

Riccien 84 (1836).— var. 5 Nees, Naturgeschichte der

europaischen Lebermoose 4: 440 (1838). — var. e stricta

Gottsche et al.. Synopsis Hepaticarum 610 (1846).

Ricciella stricta (Lindenb.) Trevis. in Memorie de Reale

Istituto Lombardo Ser. 3,4: 62 (1877). Type: Cape,

Philipstown, Ecklon s.n. (BM-Herb. Lindenb. in Herb.

Hampe, holo.!). Philipstown, the locality given on the

label, is in the central Cape Province, and not in the south-

western part of it as indicated by Nees (1838): ‘Vorgebirge

der guten Hoffnung bei Phillipstown (sic) und Krakakam-

ma’, as well as by Gottsche et al. (1846): ‘Promontorio

Bonae Spei’. The substrate ‘haufig auch an Baumen

wachsend’ or ‘ad arborum truncos’, as reported by

Lindenberg (1836), Nees (1838) and Gottsche et al. (1846),

is highly suspect: it is not mentioned on the specimen

label, neither has this been observed for any Riccia

species; moreover, soil particles were found with the

specimen in the packet.

Invalidly published or unpublished synonyms:

Ricciella tenerrima Steph. ined. (leones Ineditae), Natal, dist.

Alexandra Sta., Dumisa, Rudatis 1291 (Ml).

Ricciella dinteri Steph. ined., Dinter: 136 (1926), South West Africa

[Namibia], Okozongomuinja, Dinter 1951.

Riccia stricta A.V. Duthie ined., in Arnell: 37 (1963).

Thallus monoicous, ?perennial, in long, narrow, to

somewhat wider ribbons (Figures 1A; 2A), forming

dense, tangled masses, small to medium-sized; branches

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

MS. received: 1989.11.27.

repeatedly symmetrically or asymmetrically furcate,

moderately to widely divergent, 15—20 mm long, segments

generally up to 5 mm long, occasionally longer, (0,2—)

0,5— 0,8(— 1,2) mm wide, 0,25— 0,35(— 0,5) mm thick, in

section (1 — )2 — 3( — 4) or more times wider than thick

(Figure IE, F), linear, apex often bulbous, especially when

forking imminent, to slightly narrower and somewhat

tapering, notched (Figure 2C) and only grooved toward

apex in living plants; bright green, often flecked or

streaked with purple along margins and flanks; margins

rounded, obtuse to subacute, flanks vertical to sloping

obliquely to almost parallel with ventral face, ventral face

gently rounded to flat, green, sometimes giving rise to

stolons (Figure IB); when dry, groove more pronounced

and seemingly longer, dorsally light green and flattened,

to not much altered.

Anatomy, dorsal epidermis forming flat cover over elon-

gated air chambers, cells long-hexagonal, 42—65 x 25

/xm (Figure 1C), smaller at margins and isodiametric, ±

25 jam wide; air pores small, ± 17 pm wide, more numer-

ous toward apex, surrounded by a ring of 5 or 6 smaller,

+ 12,5—17,5 /xm wide, thin-walled, oval companion cells,

partly overlying thicker-walled epidermal cells (Figure

1C); assimilation tissue 100-500 /xm thick, occupying less

than ‘/2, to most of thickness of thallus, air chambers

lengthwise elongated, in 1 or 2 layers, up to 65 pm wide,

separated by chlorophyllose cellular plates, cells in one

layer, isodiametric, 25-40 /xm; storage tissue 150—500

/xm thick, occupying ventral part of thallus, cells round-

ed, ± 25 /xm wide; ventral epidermal cells elongated,

17—20 /im wide, bearing rhizoids, 12—15 /xm wide, mostly

smooth, but some tuberculate, abundant at sporangia and

terminal bulbous part of occasional stolons (Figure IB),

otherwise rather scanty. Scales small, up to 250—500 x

150-400 /xm, present under apex and rarely protruding

slightly (Figure 2C), then at short intervals along ventral

198

Bothalia 20,2 (1990)

FIGURE 1. — Riccia stricta. Morphology and anatomy. A, habit; B,

ventral stolon; C, air pore with thin-walled companion cells and

thicker-walled epidermal cells; D, longitudinal section through

sporangium, showing long archegonial neck and spores still in

tetrads; E, transverse section through narrow thallus growing in

drier area; F, transverse section through thin, wide thallus from

wetter area; G, ventral face with scales; FI, single scale; I,

antheridial neck with basal collar of conical cells. A, Van Rooy

3539 ; B, Van Zinderen Bakker 7472 \ C, S.M. Perold 861 ; D, S.M.

Perold 365 ; E, G, S.M. Perold 354\ F, Magill 65 92 ; H, T.R. Sim

PRE-CH 1119', I, S.M. Perold 842. Drawings by J. Kimpton. Scale

bars on A, B = I mm; C, I = 50 /jm; D, F, G = 200 /xm; E,

FI = 100 pm.

face, up to five (Figure 1G), sometimes also at furcations,

obtusely triangular, concave, single (Figure 1H), or split

into two half-scales, but remaining connected at base,

fragile, hyaline or purple, cells 4—6-sided, isodiametric,

50-65 /xm, 1 or 2 rows toward apex wider than

long. Antheridia near apex, and more proximally, single,

at intervals medianly along branches, necks hyaline,

conspicuous, 150—200 /xm long, basally surrounded by

a collar of low, hyaline, conical cells, 37-50 x 30 /xm

(Figure II). Archegonia median, up to 3 per segment,

serially arranged. Sporangia frequently at wider and

always at thicker sites along thallus, protruding con-

spicuously on ventral face, subspherical and obliquely

orientated (Figure ID), ± 600 /xm wide, containing ±

270 spores each; neck purple, long, sloping at an angle

toward, and opening into an apically directed, shallow

furrow, its ‘blind’ end fringed with a few, erect, hyaline,

conical cells (Figure 2H). Spores (50— )62 — 70(— ' 75) /xm

in diameter, triangular-globular, polar, light brown, semi-

translucent; wing thick, + 7,5 /xm wide, usually wider

at perforated or notched marginal angles, with a row of

fine granules along edge, margin crenulate; ornamenta-

tion reticulate on both spore faces, but different: distal face

(Figure 3D— F) highly convex, with (4 — )5 — 6 large, deep

areolae across diameter of spore, 17—20 /xm wide, with

central pillar or boss, from which several low ridges radiate

outward, sometimes forming a network, areolar walls

rounded, thick, ±3—4 /xm wide and up to 7,5 /xm high,

extending onto wing, sometimes sparsely granulate; prox-

imal face (Figure 3A— C) with triradiate mark very promi-

nent, up to 5 /xm high and wide, frequently even wider

toward marginal angles at join with wing, each facet with

6—10 areolae, some incomplete, often subdivided by thin,

faint, radiating ridges, walls thin, ± 5 /xm high, marked-

ly raised at nodes and sometimes joined (Figure 3A). Chro-

mosome number n = 8 (Bornefeld 1989).

R. stricta grows on damp soil or mud, sometimes in

association with other Riccia species, e.g. R. natalensis

Sim, R. crystallina L. emend. Raddi, R. cavernosa Hoffm.

emend. Raddi, and with Anthoceros spp. Occasionally

it is aquatic and floats in masses on still water or is

submerged. The land form of R. stricta sometimes forms

bulbils at the apices of the thalli (Volk 1984) to survive

drought conditions, and also to store food reserves;

occasionally propagation is by ventral stolons (Figure IB).

R. stricta is widely distributed in the summer rainfall

area and is one of the • commonest Riccia species in

southern Africa (Figure 4). Its range extends further north-

wards into central Africa: Arnell (1956) reported it from

Masai Province in Kenya, and Stephani from Usambara

as R. fluitans (Stephani in Brunnthaler 1913) (see also

‘Specimens examined’).

Like the other members of the R. fluitans complex, R.

stricta is highly sensitive to the water supply and humidity.

It is therefore very variable in its morphology, the thalli

generally ranging from thicker and narrower (Figure IE)

in drier localities, to thinner and wider (Figure IF) in

wetter places.

It is well known that species in the R. fluitans complex

are very variable and notoriously difficult to distinguish

from one another, ideally requiring cultivation under

similar conditions (Berrie 1964). In Table 1, R. stricta is

Bothalia 20,2 (1990)

199

FIGURE 2. — Riccia stricta. Morphology and anatomy. A, terrestrial thalli in cultivation; B, apical part of branch with archegonial grooves, fringed

by conical cells; C, apex of thallus, with apical notch, scale protruding slightly behind notch; D, ventral scales, single above, split into

two below; E, air pores and dorsal cells; F, air pore with companion cells; G, sporangium protruding ventrally; H, apically directed ar-

chegonial groove with conical cells; I, antheridial neck with collar of conical cells at the base. A, S.M. Perold 2524\ B, Glen 1832\ C— I,

S.M. Perold 2499. A, by A. Romanowski; B— I, SEM micrographs. Scale bars on A,B = 1 mm; C-D, G-I = 100 pm; E, F = 10 pm.

compared to R. fluitans sensu stricto, R. canaliculata and

R. purpurascens.

Markham (pers. comm.) found a wide range of flavonoid

compounds in a specimen of R. stricta (S.M. Perold 2611),

most of which he had reported (Markham et al. 1978) for

R. fluitans ; in fact, no components were found in the

specimen by which it could be distinguished chemically

from R. fluitans. This serves to confirm the close

relationship of the two species.

R. fluitans L. sensu stricto does not appear to occur

naturally in southern Africa. A single specimen of it, leg.

Stephens BOL 25511, collected in a fishpond in

Rondebosch, Cape, was apparently introduced along with

other aquatic plants from Europe (note on packet in

Schelpe’s handwriting). None of the other southern

African specimens that were examined, had the same

appearance as this, which has somewhat thinner thalli,

shorter branches and smaller air chambers with more

distinctly visible walls, when viewed from above; dried

plants have a slightly ‘crinkled’, not smooth, appearance

(for differences between the species, see Table 1). Earlier

reports of R. fluitans from southern Africa by Krauss

(1846), Stephani (1913), Sim (1926) (as R. fluitans-

limicola), and Arnell (1963), have not been verified. The

checklist by Magill & Schelpe (1979) records it, and is

referred to by Mahu (1985) as proof of its presence in

southern Africa. This list, however, was a preliminary

account largely based on previous records.

Much of the tropical African material identified as R.

fluitans or R. fluitans sens. lat. may also belong to R.

stricta. Jones (1957) found that the spores of two of his

African collections ( 655 and 826) named R. fluitans sens,

lat. , differed from those of European material and Bizot

et al. (1978) concluded the same for material from Kiliman-

jaro. The spores illustrated by Jones (1985) ( Foster 55 from

Kampala, Uganda, and Jones 826 from Luki, Zaire) show

a pronounced triradiate mark on the proximal face and

thick areolar walls on the distal face, strongly reminis-

cent of those of R. stricta. Vanden Berghen (1972) who

examined some African collections of the complex (Sy-

moens 12436, 12774, Schmitz 7305 and Jean Louis

4410— all in BR) expressed the opinion that ‘R. stricta

Bothalia 20,2 (1990)

201

FIGURE 3. — Riccia stricta. Spores. A, B, proximal face; C, proximal face in side view; D, F, distal face; E, distal face from the side. A, Van

Rooy 2201 ; B, T.R. Sim PRE-CH 1090, C, Van der Bijl 10: D. Van Zinderen Bakker 7472: E, F, T.R. Sim 7590. Scale bars on A— E =

50 pm: diameter of spore on F = ± 70 fim.

A.V. Duthie in S. Arnell (1963) Hep. South Afr. , p. 37,

est peut-etre identique au taxon reconnu au Shaba’.

Schmitz 7305 and Jean Louis 4410 were also examined

by me and their spores studied with LM and SEM. In both,

the triradiate mark is pronounced and the areolar walls

on the distal face are thickened.

In spite of the uncertainty concerning the classification

of the R. fluitans complex, a number of new species have

been described in the group in recent years. Examples are:

R. rhenana Lorb. (1934) (possibly a diploid form of

R. fluitans ; R. duplex Lorb. (? a diploid form of R.

canaliculata (Berrie 1964) (see also Stottler & Crandall-

Stottler 1977); R. gamsiana Lorb. (= R. canaliculata );

FIGURE 4. —Distribution map of R. stricta , A ; and R. purpurascens,

O, in southern Africa.

R. media Klingmiiller (1957); R. abuensis Bapna (1962)

from India; R. limicola Jovet-Ast (1978) from Galapagos;

and from Australia, R. luticola Na-Thalang, as well

as two varieties of R. duplex and several of R. multifida,

all by Na-Thalang (1980). In a new species described

from Brazil, R. jovet-astiae (Vianna 1988), the SEM

micrographs of the spores are very similar to those of

R. stricta and the differences cited by Vianna between

this new species and R. stricta are not all significant,

for example, specimens of R. stricta are not dioicous,

but definitely monoicous, although in recently cultured

material, antheridia and archegonia were consistently

found in separate plants; the spore size exhibits a

wide range in the many fertile southern African specimens

examined (although Arnell’s (1963) figure at 40-50 /im,

seems a trifle on the small side); the thalli are certainly

not always markedly wider at the sporangia either.

The thallus branches in R. jovet-astiae, however, appear

to be rather shorter than in R. stricta.

R. stenophylla Spruce, one of the ‘older’ species in this’

group, also seems to be closely related to R. stricta. This

is confirmed by examination of the spores of Volk 82/895

from Rio, identified as R. stenophylla. These spores are

indistinguishable from three specimens in PRE identified

as R. fluitans : Schiffner 1873/74 (Java), Verdoorn IX 1930

(Java) and Wright CH 810 (? Cuba).

To conclude, perhaps one should bear in mind Meijer’s

(1958) words of caution, to postpone future splitting

of this cosmopolitan species complex, R. fluitans,

until more detailed studies, based on living material

from many parts of its whole range, have been made.

R. stricta, of course, is not a new species but merely a

new combination.

202

Bothalia 20,2 (1990)

SPECIMENS EXAMINED

ZIMBABWE. — 1725 (Livingstone): Victoria Falls, Kuun 5307. 5308

(BOL); T.R. Sim 9056. 9066 (PRE), PRE-CH 1114 (PRE). 1831

(Marandellas): Marandellas, Eyles 3885 (BOL). 2028 (Bulawayo):

Bulawayo hillside, submerged in stream and on streambank, Eccles BOL

25757 (BOL); Khama stream, T.R. Sim 9067 (PRE-CH 1095) (PRE).

MOZAMBIQUE. — 2532 (Lourenco Marques): Maputo, Junod 324

(PRE); Antioka, Magude, H.A. Junod (PRE). Exact localities not given.

Wager 10. 60 (PRE).

All of the above specimens from Mozambique had been

identified as R. fluitans or as R. purpurascens .

NAMIBIA. — 2016 (Otjiwarongo): Waterberg Sta., Groot Waterberg,

occasional on moist earth around reservoir (— BC), Schelpe 4807 (BOL);

Farm Otjihaenamaparero (OTJ 92), unter Wasser, kleine Polster bildend,

im Quellbach bei Dinosaurierspuren (-CD), Giess 15236 (PRE). 2416

(Maltahohe): Karab MAL23 (—DA), Volk 01261 (M, PRE); 2417

(Mariental): Haribes (—DA), Volk 12460 (M, PRE); Pavianskolk, Haribes

(—DA), Volk 5313 (M, PRE).

TRANSVAAL. — 2230 (Messina): Venda, Nwanedi Nat. Park, river

nr compound manager’s house, submerged (— CB) Glen 1832 (PRE);

Sebasa, + 3 miles [4,8 km] due S of Lake Funduzi, submerged in spring

and small stream (—CD), Meeuse 9411 (PRE). 2329 (Pietersburg): Letaba

Valley, 23 km along road R528 from Tzaneen to Haenertsburg, at seepage

area (— DD), S.M. Perold 2455 ( PRE). 2330 (Tzaneen): Lemana(— AA),

Junod PRE-CH69 (PRE); Lemonia Wood, Spelonken (— CA), Junod 3

(PRE-CFU049) (PRE); Woodbush For. Res. nr waterfall (— CC), Magill

6592 (PRE). 2331 (Phalaborwa): Kruger Nat. Park, east of Singwedzi

road, Letaba River (— AB), Brandt 5360 (BOL); Lebombo Mts, potholes

on Bangu Creek, just S of Olifants River (— DD), Magill 5039 (PRE).

2428 (Nylstroom): Doomdraai Nat. Res., 35 km NNW of Naboomspruit,

in vlei above dam, on mud (— BD), S.M. Perold 438 (PRE); Platrivier.

5 km W of Warmbaths, submerged in stream (-CC), S.M. Perold 2611

(PRE). 2430 (Pilgrims Rest): SW of Phalaborwa, Farm Parsons, along

Olifants River ( — BB), Venter 12199 (PRE). 2528 (Pretoria): Pretoria,

Rietfontein (— CA), H.A. Wager 18 (PRE); Malanspruit, on road from

Moloto to Cullinan, submerged, in dense masses (-DA), Mauve & Venter

5077 (PRE); 18 km NE of Cullinan, ‘Grotte’, above stream (-DA), S.M.

Perold 2600 (PRE). 2529 (Witbank): Loskop Dam Nat. Res.,

Rhenosterhoek, in damp gulley under vegetation on edge of exposed rock

sheet (—AD), Reid 1106 (PRE). 2530 (Lydenburg): Sabie, Bridal Veil

Falls, on sandy, muddy shallow bank on path to falls (-BA), Thompson

295 (PRE). 2531 (Komatipoort): Barberton (— CC), Hendry PRE-CH

1107 (PRE). 2627 (Potchefstroom): Gerhard Minnebron, by Oog, in

stromende water (-AC), Louw 1621 (PRE); same locality Ubbink 1154

(PUC); noord van Klerkskraal, by Eerste Oog (—AC), Ubbink 1004

(PRE); Pienaarkamp by permanente drinkgat op vertikale walle (-CA),

Ubbink 964 (PRE); Potchefstroom, north of Mooirivier (-CA), Goosens

PRE-CH 3672 (PRE). 2628 (Johannesburg): Johannesburg, Melville

(— AA), T.R. Sim PRE-CH 1088 (PRE); Eloff (—BA), T.R. Sim PRE-CH

1083 (PRE). 2629 (Bethal): 15 km along road R545, between Balmoral

and Ogies, pan at roadside, at turnoff to Roodepoortjie (— AA), S.M.

Perold 360 (PRE); 17 km W of Hendrina, on soil, at edge of pan (—BA),

S.M. Perold 354 (PRE); 5 km N of Hendrina, on soil next to road (—BA),

S. M. Perold 356 (PRE); 24 km from Bethal on road R29 to Ermelo,

at furrow in vlei nr dam (— BC), S.M. Perold 365 (PRE). 2630 (Carolina):

Chrissiesmeer, nr lake in damp furrow (-AD), S.M. Perold 1051 (PRE);

± 21 km from Panbult on road to Amersfoort (-CD), Germishuizen

2922a (PRE); 3 km S of Amsterdam on road to Piet Retief, on damp

soil ( — DA), S.M. Perold 1071 (PRE).

O.F.S. — 2729 (Volksrust): 31 km NE of Verkykerskop on road from

Memel, Farm Swiza, at seepage (—CD), S.M. Perold 1277 (PRE). 2828

(Bethlehem): 23 km W of Harrismith on road to Kestell, at turnoff to

Rydal Mount, on damp rock outcrop (-BD), S.M. Perold 1284 (PRE).

2926 (Bloemfontein): Bloemfontein, in small dry pan nr Lambton’s

plantation, SW corner of College grounds (-AA), Van Zinderen Bakker

7472 (BOL).

NATAL.— 2729 (Volksrust): Majuba Nek, on wet soil in shade of rocks

(-BD), Herschel PRE-CH 1089 (PRE). 2730 (Vryheid): Vryheid (-DD),

T. R. Sim PRE-CH 1074. PRE-CH 1120 (PRE); hill above Vryheid (— DD),

T.R. Sim PRE-CH 1123 (PRE). 2731 (Louwsburg): Ngoma State For. Res.,

along sides of damp pathway through Ntendeka Forest (-CD), Nicho-

las H94 (PRE). 2732 (Ubombo): Imbezane, on earth bank of stream

(-BC), Eyles 1405 (PRE); Manywana River, at crossing of southern dirt

road to Sodwana Bay, NW of False Bay (-CD), Magill 5385 (PRE);

Ubombo, Sodwana Bay, floating in river (—DA), Gerstner 705 (PRE).

2828 (Bethlehem): Mont-aux-Sources, Saxton, Tugela (— DD), no

collector's name or number (BOL). 2829 (Harrismith): Oliviershoek Pass,

between Harrismith and Bergville, at waterfall on wet cliff (— CA), Van

Rooy 1167 (PRE). 2831 (Nkandla): Nkandla, inheemse bos by swembad

en munisipale karavaanpark Eshowe op dooie drywende blare in poel

met stilstaande water (—CD), Botha & Van Wvk 1016 (PRE); same locality,

Nixon 109 (BOL); Ngoya (-DC), T.R. Sim PRE-CH 1105 (PRE). 2832

(Mtubatuba): St Lucia Res., Fanie’s Island, Umkhiwane trail, at side

of ditch (-AB), Glen 2029 (PRE). 2929 (Underberg): Giant's Castle

(-AB), Symons PRE-CH 1104 (PRE); bush below Cathkin Peak (— AB),

T.R. Sim PRE-CH 1103 (PRE); Sweetwaters, at stream (—BA), T.R. Sim

PRE-CH 1092 (PRE); Estcourt, along stream (— BB), West PRE-CH 3668

(PRE); Nottingham Road (— BD), Van der Bijl PRE-CH 1112 (PRE); Sani

Pass Hotel, 1 km beyond, streamlet at roadside, partly submerged (— CB),

S. M. Perold 2499 (PRE); Xumeni For. Res. (— DD), Doidge PRE-CH

3580 (PRE). 2930 (Pietermaritzburg): Tweedie (-AC), G.W. Sim 8093

(PRE); Fort Nottingham Commonage, Lion’s River Dist. (—AC), Hilliard

& Burn 10344 (PRE); Howick Falls (-AC), G.W. Sim PRE-CH 1087

(PRE); Muhwati, New Hanover (-BC), T.R. Sim PRE-CH 1118 (PRE);

Pietermaritzburg, stones in Umsundusi (— CA), T.R. Sim 7593 (PRE);

Town bush (— CB), T.R. Sim PRE-CH 1106. PRE-CH 1122. PRE-CH 1075

(PRE); nr top of Zwaartkop (— CB), T.R. Sim PRE-CH 1109 ( PRE); Scotts-

ville (— CB), T.R. Sim PRE-CH 1072 (PRE); Alexandra Park (— CB),

T. R. Sim PRE-CH 1115 (PRE); Carter’s Nurseries, under bank on dam

overflow (— CB), Wells 57 (PRE); Pinetown Dist., Everton, on damp

shady soil in garden (— DD), Hilliard & Burn 8151 (PRE); New Germany

(— DD), Moonsammy 14 (PRE), Van der Bijl 14, 19 (PRE); Durban,

Palmiet Nat. Res., wet vertical cliff lace above fast-flowing river (— DD),

G. Lambert 6 (PRE); Isipingo Flats, on moist, sandy clay along water

furrow (— DD), Ward PRE-CH 5354 (PRE). 2931 (Stanger): Tongaat Su-

gar Estate Gardens, on rock in pond (— CA), S.M. Perold 27 (PRE);

Verulam, Zwolle (-CA), Van der Bijl PRE-CH 1097 ( PRE). 3029(Kok-

stad): Kokstad (-CB), Mogg PRE-CH 1078 (PRE).

LESOTHO. — 2828 (Bethlehem): 38 km W of Oxbow Lodge, seeps

in shallow sandstone cave along road (— CB), Magill 4626 (PRE). 2929

(Underberg): in Hats S of Sani border post, alpine meadow (— CB), Magill

7067 (PRE); Sani Top, at disused airstrip, N of mountaineer's chalet,

on vertical earth wall of ditch (— CB), S.M. Perold 2521 (PRE); Sani

Top, S side of dam, nr border post, vertical earth wall of ditch (-CB),

S. M. Perold 2528 p.p., 2529 p.p. (PRE).

TRANSKEL— 3128 (Umtata): 4 km from Elliottdale on road to ’The

Haven’, disturbed, wooded stream (-DC), Van Rooy 2141 (PRE). 3129

(Port St Johns): Hluleka Nat. Res., Hluleka Forest, on sandy soil (—CD),

Van Rooy 2201 (PRE). 3130 (Port Edward): Mutentu River (-AA), Burn

Davy PRE-CH 110 (PRE). 3228 (Butterworth): Kentani Dist., growing

on waters' edge and vleis, common (-AD), Pegler 1354 (BOL, PRE);

9 km from Dwesa along road to Idutywa, wooded road cutting (-BD),

Van Rooy 2089 (PRE); Dwesa Nat. Res., Dwesa For., on shale on

streambank (— BD), Van Rooy 2123 (PRE).

CAPE. — 2722 (Olrfantshoek): Langeberg (— DD), Duthie 5384 (BOL).

3024 (De Aar): Phillipstown (sic) (—AD), Ecklon s.n. (BM, holo.). 3027

(Lady Grey): Barkly East Dist., 12 km from Lundin's Nek turnoff (— CA),

Thompson 260 (PRE). 3225 (Somerset East): Cradock, Fish River

(-BA), Duthie 5143 (BOL). 3226 (Fort Beaufort): Alice, in dam nr river

in village, aquatic, floating on surface (-DD), Potts 1758 (BOL); Alice,

plants on edge of dam, nr river in village (-DD), Potts 1760 (BOL).

3 2 27 (Stutterheim): Hogsback, indigenous forest (-CA), McDonald

PRE-CH 13422 (PRE); Kingwilliams Town, in water (—CD), T.R. Sim

i/i(PRE). 3319 (Worcester): Tulbagh, Farm Kleinberg, irrigation furrow

(-AC), Duthie 5022. 5391.5477 ( BOL), Theron 5379 (BOL); Worcester,

Hexrivier (-CB), Naude 5429 (BOL). 3323 (Willowmore): Diepwalle

For. Sta., nr Grootboom, on damp soil (— CC), S.M. Perold 913 (PRE).

3325 (Port Elizabeth): Zuurberg Nat. Park, upper pool (-BC). B-E.

& M. van Wyk 933. 2091. 2093 (PRE); springs at Uitenhage (-CD),

T. R. Sim 9041 (PRE). 3326 (Grahamstown): nearby Grahamstown (-BC),

Britten 2785 (PRE); Round Hill, Oribi Res., on surface of water (-BD),

Wirminghaus 247 (PRE). 3422 (Mossel Bay): nearby Knysna (-BB),

Davey 17009 (PRE). 3423 (Knysna): Knysna, Forest Hall (-AA), Duthie

5043. 5044. 5220. 5386 (BOL); Belvidere (-AA), Duthie 5388,5390

(BOL); Brenton (-AA). Duthie 5387 (BOL); Woodbourne (-AA),

Duthie 5389 (BOL); Plettenberg Bay (-AB), Duthie 6004 (BOL); Storms

River Mouth, Tsitsikama Nat. Park (-BB), Wagener 1, 2 (PRE).

Bothalia 20,2 (1990)

203

2. R. purpurascens Lehm. & Lindenb. in Linnaea

4 : 371 (1829); Lehmann: 23 (1832); Lindenberg: 451

(1836); Gottsche et al.\ 611 (1846); Stephani: 363

(1898); Sim: 15 (1926); S. Arnell: 36 (1963). Ricciella

purpurascens Trevis.: 62 (1877). Type: Cape, Crescit humi

in sylvula quercina ad latus boreale et ad radicem montis

Tafelberg in Promont. Bonae Spei, Ecklon s.n. (G! , lecto.,

selected here) (ex Herb. L., ex Herb. Rom.).

Thallus dioicous, annual, in thin, somewhat lax strands

or ribbons (Figure 5A), frequently overlapping and

becoming quite densely massed (Figure 6A), medium-

sized to rather large; branches repeatedly and irregularly

furcate, 9—17 mm long, segments 1-5 mm long, narrowly

to moderately divergent, 1,5 -2,0 mm wide, 0,4— 0,6 mm

thick medianly, over keeled ventral part, lateral wings up

to 0,3 mm thick, in section (Figure 5F) 3 — 5 ( — 7 ) times

wider than thick, linear, apex slightly narrowed (Figure

6B), not emarginate (Figure 6C); groove hardly distinct

toward apex, soon becoming wide and shallow or dis-

appearing altogether; light green, sometimes almost

translucent, occasionally purple along margins and over

gametangia; margins subacute, rather irregularly

undulating, winged or attenuate, flanks sloping obliquely

to almost flat, green; ventral face flat to narrowly keeled

medianly, frequently giving rise to stolons; when dry,

yellowish green, thin and flat.

Anatomy, dorsal epidermis forming a flat cover over

large air chambers (Figure 5C), cells 5-sided or oblong-

hexagonal, up to 110 X 35-60 pm, at apical margin

rectangular, smaller, 45 x 30 /xm; air pores small (Figure

6D), surrounded by (4— )5 radially arranged, thin-walled

cells (Figure 5D), 17—35 x 12-15 pm, partly overlying

thicker-walled epidermal cells; assimilation tissue ±

300-400 pm thick, occupying most of thickness of thallus,

air chambers polyhedral, in 2 layers medianly, uniseriate

laterally, separated by chlorophyllose plates, one cell thick;

storage tissue mostly only 1-4 layers of cells, angular,

50—62 /xm wide; rhizoids arising from ventral epidermis

medianly, numerous at sporangia and tips of stolons

(Figure 5B), ± 15 /xm wide, some smooth, others

tuberculate. Scales small, hyaline, ventral, toward apex

only, up to three pairs, remaining attached in the middle,

difficult to detect, evanescent. Antheridia serially arranged

along middle in groups of 2 or 3 with sterile areas in

between, bulging above and below (Figure 6F), antheridial

necks up to 200 /xm long, at base surrounded by collar

of hyaline, conical cells, ± 50 /xm long (Figure 6F).

Archegonia median, single or in pairs, sometimes

adjacent, necks vertically orientated, brownish purple with

hyaline tip, opening into a depression (Figure 6E), at base

surrounded by conical cells 30-40 /xm long (Figure 5E).

Sporangia ± 550 /xm wide, causing widening of thallus

at maturity, bulging ventrally with surrounding tissue

thickening into ± 6 layers of cells, containing up to 580

spores each. Spores (65 — )70 — 80( — 88) /xm in diameter,

triangular-globular, polar, yellow to bright brown, semi-

translucent; wing thick and up to 7,5 pm wide, at marginal

angles 2 small pores, one on either side of each arm of

triradiate mark, margin finely crenulate; ornamentation

incompletely reticulate and different on 2 spore faces:

distal face (Figure 7E, F) with 4-6 large, angular, mostly

incomplete areolae across diameter, ± 20 pm wide,

usually subdivided by secondary ridges or a central pillar

into smaller areolae, toward margin often reduced to short

ridges only, walls ± 5 pm high, densely fringed with gran-

ules, slightly raised at nodes; proximal face (Figure

FIGURE 5. —Riccia purpurascens. Morphology and anatomy. A, habit;

B, ventral face with stolons and sporangium; C, dorsal face with

air chambers and epidermal cells partly drawn in; D, air pore

with thin-walled companion cells and thicker-walled epidermal

cells; E, transverse section through sporangium, with conical cells

at depression; F, transverse section through thallus. A, B, F,

Morley 291 ; C, S.M. Perold 1941: D, E, S.M. Perold 1170.

Drawings by J. Kimpton. Scale bars on A, B, C = 1 mm; D

= 50 pm: E, F = 200 Mm.

204

Bothalia 20,2 (1990)

FIGURE 6. — Riccia purpurascens. Morphology and anatomy. A, field-grown thalli, arrow indicating row of gametangia; B, branch with sporangium

and rhizoids protruding ventrally; C, apex of thallus, not notched, without protruding scale; D, air pore and dorsal cells; E, thallus above

sporangium; F, cross section through antheridium, bulging above and below thallus, base of neck with conical cells. A, S.M. Perold 2386 ;

B— E, Koekemoer 284 p.p.; F, S.M. Perold 611. A, by A. Romanowski; B— E, SEM micrographs; F, LM photograph. Scale bars on A— C,

E, F = 1 mm; D = 100 pm.

7A— D) with triradiate mark prominent, ± 2,5 pm wide,

becoming wider toward marginal angles at juncture with

wing, row of superimposed papillae running along the

arms, on each of 3 facets rarely any complete areolae,

mostly short broken walls, straight or curved, edged with

tall uneven spines, warty papillae or low granules.

Chromosome number n = 8 (Bornefeld 1989).

R. purpurascens grows on damp, sandy soil or on mud,

sometimes on streambanks and rarely in light shade,

occasionally in association with other Riccia species, such

as R. cupulifera Duthie, R. bullosa Link ex Lindenb. , R.

crozalsii Lev. and R. limbata Bisch. ex Krauss as well as

with Fossombronia spp. and Gongylanthus spp. It is en-

demic to the north-western, south-western and southern

FIGURE 7. — Riccia purpurascens. Spores, A, B, D, proximal face; C, proximal face, partly from the side; E, distal face; F, distal face seen

from the side. A, S.M. Perold 1170', B, D— F, McLaughlin PRE-CH 4/97; C, S. Arnell 3321. Scale bars on A— C, E, F = 50 pm; diameter

of spore on D, ± 80 pm. All micrographs by S.M. Perold.

Bothalia 20,2 (1990)

205

Cape, which are winter rainfall areas (Figure 4). Arnell

(1963) reported R. purpurciscens from Victoria Falls,

Zimbabwe, but these specimens were actually R. stricta

and had been misidentified, as were several others at PRE.

The deep purple colouration that thalli of R. stricta develop

on exposure to the sun, may have led to this error.

R. purpurascens and R. stricta are two southern African

'linear’ species with ventral sporangia, and both produce

stolons. Although there is some overlap in their

distribution in the southern Cape, they generally occupy

distinct climatic areas, with R. stricta being far more

widespread in the summer rainfall areas; R. stricta is also

sometimes aquatic, whereas R. purpurascens is strictly

terrestrial. In addition, they differ in their apices, scales,

the strict dioicity of R. purpurascens, the vertically

positioned sporangia in the latter, its larger-sized thalli as

well as differences in the spore ornamentation, with the

areolar walls thinner and usually incomplete on the distal

face.

Oil cells were not found in the stolons of R.

purpurascens as reported by Sim (1926); numerous starch

granules are, however, present.

grounds (— AA), Duthie 5423 (BOL); Knysna, Woodbourne (— AA).

Duthie 5437 (BOL); Knysna, Erica Farm (-AA), McLaughlin 1211

(PRE).

ACKNOWLEDGEMENTS

I wish to thank the curators of BOL, BM and G for the

loan of specimens. I am most grateful to Prof, honoraire

S. Jovet-Ast for critically reading the manuscript, for

information concerning R. canaliculata and for the gift

of specimens; and thanks also to Dr E.C. Vianna. Sincere

thanks to Dr E.O. Campbell, Massey University, for

reading the manuscript and for mediating with Dr K.

Markham, who very kindly performed flavonoid analyses

on two specimens, which were much appreciated. Above

all, I wish to thank Prof, (emer.) Dr O.H. Volk, who also

very generously sent me specimens, literature and photo-

graphs, and with whom I have had numerous discussions.

Ms J. Kimpton, artist, Mrs A. Romanowski, photographer,

and Mrs J. Mulvenna, typist, are thanked for their valued

contributions.

REFERENCES

SPECIMENS EXAMINED

CAPE. — 3119 (Calvinia): 2 km from Nieuwoudtville, on mud in ditch

at roadside (-AC), S.M. Perold 1755 (PRE). 3217 (Vredenburg):

Witteklip, south of Vredenburg (-DD), Leighton 537 (BOL). 3218

(Clanwilliam): 17 km E of Clanwilliam, along Pakhuis Pass, at Leipoldt’s

grave, weathered sandstone rocks (— BB), S.M. Perold 1935 (PRE); 5

km along road to Cedarberg, after turnoff to Algeria For. Sta., damp

overhang nr road (— BD), S.M. Perold 2347 (PRE); Citrusdal, 22 km

N of, sandstone rock outcrops above Olifants River, Hex River Estates

(-BD). S.M. Perold 2386 (PRE); Weltevrede. 17 km W of Piketberg,

banks of Grootberg River (—DC). S.M. Perold 500 (PRE). 3219

(Wuppertal); Wuppertal, streambank (—AC), Malherbe & Davies 5377

(BOL). 3318 (Cape Town); Darling (—AD), Duthie 5425 (BOL);

Malmerbury Commons (—AD), Garside 26210 (BOL); 1 mile outside

Malmesbury on Piquetberg road, on soil (— BC), Schelpe 3901 (PRE);

Lion’s Head, nr Round House (—CD), 5. Arnell 240, 330 (PRE); Lion’s

Head above Clition (—CD), S. Arnell 494 (PRE); above Bakoven (—CD),

5. Arnell 922 (BOL); between Kloofnek and Round House (—CD), S

Arnell 248, 251 (PRE); Tafelberg in Promont. Bonae Spei (—CD), Ecklon

s.n. (G, lecto.); Table Mountain, rock shelf nr dripping water (—CD).

Esterhuysen 19232 (BOL); Table Mountain, streamside, on lower W

slopes above Camps Bay (—CD), Esterhuysen 21718 (BOL); Devil’s Peak

(-CD), T.R. Sim PRE-CH 1498 (PRE); Cape Town (-CD), Wager

PRE-CH253 (PRE); Stellenbosch (-DD), Duthie PRE-CH 1125, PRE-CH

1128 (PRE), Garside PRE-CH 1126 (PRE); Stellenbosch, riverbank

(-DD), Duthie 5012 (BOL); Stellenbosch. Platklip (— DD), Duthie 5325

(BOL); lower slopes above Brandwacht, damp, sandy soil at small stream

(-DD), Oliver 9027 (PRE); Stellenbosch, Papegaaiberg, partly dried

furrows (— DD), Malan s.n. (BOL); Papegaaiberg, on soil at foot of hill

(-DD). S.M. Perold 477 (PRE). 3319 (Worcester): Tulbagh (—AC),

Duthie 5381a (BOL); Tulbagh, waterfall in cave (—AC), Groenewald 5327

(BOL); Franschhoek, nr Waterfall Farm, on soil at bridge over stream,

(-CC), S.M. Perold 634a (PRE); Nuy, Rabiesberg. E of Worcester, on

soil ( — DA), Morley 360 (PRE); Farm Leipzig, Rabiesberg, E of Wor-

cester)—DC), S.M. Perold 587 (PRE). 3320 (Montagu): Montagu. Bath

Kloof (-CC), S. Arnell 724, 725, 755 (BOL); Cogman's Kloof, up in

kloof, on dried mud (-CC), S.M. Perold 590 (PRE). 3418 (Simonstown):

Clovelly Railway Sta. (— AA). S. Arnell 569 (PRE), 638 (BOL): Con-

stants slopes (— AA), S. Arnell 330, 332, 401 (BOL); Cape Point, moun-

tain slope (—AD), Duthie 5510 (BOL); Harmonystrand, flats NW of

Gordon’s Bay (— BB), Oliver 8778 (PRE). 3419 (Caledon): Bot River

area, Afdaksrivier, slopes E of Suikerboskop (—AC), Oliver 9224 ( PRE);

Greyton, kloof, on soil at rock wall next to footpath (—BA), S.M. Perold

611, 1170 (PRE); Greyton, kloof, on mud (—BA), Morley 291 (PRE);

Riviersonderend beyond dumping ground on mud at streambank (— BB),

S.M. Perold 594 (PRE). 3423 (Knysna): Knysna, Belvidere (-AA),

Duthie 5382, 5434 (BOL), 23 (PRE): Knysna, Gouna Forest track, Lily

Vlei ( — AA), S. Arnell 1734 (BOL); Knysna. Rectory

ARNELL, S. 1956. Hepaticae collected by O. Hedberg et al. Arkiv for

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ARNELL, S. 1957. Hepaticae collected in South West Africa by Prof.

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BAPNA, K.R. 1962. A new species of Riccia from Mount Abu (India).

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BIZOT. M., FRIIS. I., LEWINSKY, J. & POCS, T. 1978. East African

bryophytes IV. Danish collections. Lindbergia 4: 259—284.

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Nova Hedwigia 48 : 371-382.

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JONES, E.W. 1957. African hepatics XIII. The Ricciaceae in tropical

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208 -227.

JONES, E.W. 1985. African hepatics XXXV. Some new or little-known

species and some noteworthy records. Journal of Bryology 13:

497-508.

JOVET-AST, S. 1978. Riccia des lies Galapagos. Revue bryologique et

lichenologique 44: 411—428.

JOVET-AST, S. 1979. Recoite. en France, d'un Riccia fluitans L. fertile.

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togamie, Bryologique et Lichenologique 7: 283—431.

KLINGMULLER. W. 1957. Zur Kenntnis der hessischen Ricciaceen.

Berichte der Oberhessischen Gesellschaft fiir Natur- und Heil-

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KRAUSS, F. 1846. Pflanzen des Cap- und Natal-Landes gesammelt und

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Bothalia 20,2: 207-222 (1990)

Notes on African plants

VARIOUS AUTHORS

ANNONACEAE

UVARIA GRACIL1PES , A NEW SOUTHERN AFRICAN RECORD

During routine curation of material in the C.E. Moss

Herbarium, it was noticed that Hemm 434 did not match

material of Uvaria caffra E. Mey. ex Sond. or Uvaria

lucida Benth. subsp. virens (N.E. Br.) Verde. A name for

the anomalous specimen was sought in Flora Zambesiaca

(Robson 1960), where it was easily keyed to Uvaria

gracilipes N.K.B. Robson, a species reported at the time

only from the districts of Ndanga in Zimbabwe and

Inhambane in Mozambique. The identification was

confirmed at PRE. Two other collections from the

Transvaal and the duplicate of Hemm 434, all housed at

PRE, are also Uvaria gracilipes. Brynard & Pienaar 4317

(one of the Transvaal specimens) was matched with the

holotype Phelps 207, at Kew, and the identification was

thus confirmed. The species is not listed by Germishuizen

in Gibbs Russell’s List of species of southern African plants

(1987) nor its updates, and must therefore be considered

a new record for southern Africa.

Since Robson’s (1960) treatment of Uvaria gracilipes,

four more specimens have been accessioned into the

herbarium at Kew. Three are from the Nuanetsi District

in Zimbabwe and the fourth is from Mabalane in the

Gaza Province of Mozambique. These collections extend

the distributional range of the species to that shown in

Figure 1.

Key to the southern African species of Uvaria L. , adapted

from Robson (1960)

la Calyx cupular in bud, splitting into 3 lobes at anthesis or

remaining scarcely lobed; prolongation of anther connec-

tive thin or tapering U. lucida subsp. virens

lb Calyx with distinct lobes in the bud, separating into regular

sepals at anthesis, sepals free to the base; prolongation

of anther connective broadened and truncate:

2a Leaves attenuate at the base; petiole (2 — )3 — 6 mm long;

flowering pedicels about 10-20 mm long, stout,

thickenedabove, densely pubescent U. caffra

2b Leaves cuneate to rounded at the base; petiole 1—3 mm

long; flowering pedicels about 22—30 mm long, slender,

not thickened upwards, sparsely pubescent ... U. gracilipes

Uvaria gracilipes N.K.B. Robson in Boletim da

Sociedade Broteriana, ser. 2a, 32: 152 (1958); Robson:

111 (1960). Type: Zimbabwe, Ndanga District, on bank

of Lundi River, Phelps 207 (K!, holo.).

VOUCHER SPECIMENS

ZIMBABWE. — 2131 (Triangle): Nuanetsi District, Gona-Re-Zhou,

near tributary of Guluene River, north-east of Malugwe Pan, Mavi s.n.

FIGURE 1. — Distribution of Uvaria gracilipes N.K.B. Robson as

represented in J, K and PRE.

(K); Nuanetsi District, on Claredon Cliffs. Miller 652 (K); Nuanetsi

District. Lundi River, Fishans, Drummond 7703 (K); Ndanga District,

on bank of Lundi River, Phelps 207 (K, holo.).

MOZAMBIQUE.— 2332 (Massingire): Gaza de Mabalane para Mapai

(Pela estrada velha) 10 km depois de Mabalane, Grandvaux Barbosa &

De Lemos 8507 ( K). 2435 (Inharrime): Ponta Zavora, Grandvaux Barbosa

& De Lemos 8623 (K).

TRANSVAAL. — 2230 (Messina): Venda, Mutale (-BD), Hemm 434

(J, PRE); Venda, Klein Tshipise (-DA), Van Rooven 3424 (PRE); Kruger

National Park, Shingedzene, Brynard & Pienaar 4317 (K, PRE).

REFERENCES

GERMISHUIZEN, G. 1987. Annonaceae. In G.E. Gibbs Russell elal..

List of species of southern African plants, 2nd edn, part 2.

Memoirs of the Botanical Survey of South Africa No. 56.

ROBSON, N.K.B. 1958. New and little known species from the Flora

Zambesiaca area VI. Boletim da Sociedade Broteriana, ser. 2a,

32: 151-173.

ROBSON, N.K.B. 1960. Annonaceae. In A.W. Exell & H. Wild, Flora

Zambesiaca 1,1: 104—149. Crown Agents for Overseas Govern-

ments and Administrations, London.

K. BALKWILL* and A. NICHOLAS**

* C.E. Moss Herbarium, Department of Botany, University of the

Witwatersrand, P.O. Wits., 2050 South Africa.

** SALO, National Botanical Institute. Present address: The Herbarium,

Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE England.

MS. received: 1990.05. 17.

208

Bothalia 20,2 (1990)

CONVOLVULACEAE

THE CORRECT ORTHOGRAPHY AND AUTHOR CITATION OF FALCKIA

The genus Falckia Thunb. consists of three species and

is restricted to Africa. It is always cited as Falkia L.f.

Meeuse (1957: 658) discussed the historical background

of the publication of the genus, based on Thunberg’s col-

lection and observations. He followed the contemporary

assumption that Supplementum plantarum of the younger

Linnaeus had been published earlier than the first part of

Thunberg’s Nova genera plantarum , a dissertation defend-

ed by C.F. Homstedt. As to Falkia, this has been accept-

ed by Farr et al. (1979) and Gibbs Russell (1984) as well.

However, it has been established since, that Thunberg’s

publication dates from 24 November 1781, whereas Sup-

plementum plantarum was not published before April 1782

(Staflpu & Cowan 1981: 112). These publication dates re-

store historical ‘justice’ in the orthography and author ci-

tation of Falckia Thunb. Thunberg cited in the protologue

‘Act Lund.’, this probably refers to ‘Beskrifning pa et nytt

slagte uti ort-riket, kalladt Falckia', Physiogr. Sdlsk. Handl.

1.4: 303 t.4 f.3, which was only published in 1786.

Falckia Thunb. , Nova genera plantarum 1: 17. 14 Nov.

1781. Type species: Falckia repens Thunberg, ibidem

(descriptio generico-specifica); lectotype: Thunberg, Cape

of Good Hope, Convolvulus falckia, UPS-THUNB 4455.

Orth. var. : Falkia L. f., Supplementum plantarum 30: 211, April 1782.

REFERENCES

FARR, E.R., LEUSSINK. J.A. & STAFLEU, F.A. 1979. Index nominum

genericorum 2. Utrecht, The Hague.

GIBBS RUSSELL, G.E. 1984. List of species of southern African plants.

Memoirs of the Botanical Survey of South Africa No. 48.

MEEUSE, A.D.J. 1957. The South African Convolvulaceae. Bothalia

6: 641-792.

STAFLEU, F.A. & COWAN, R.S. 1981. Taxonomic literature 3. Utrecht,

The Hague.

DO. WIJNANDS* and A.D.J. MEEUSE**

* Botanical Gardens Wagepingen. Agricultural University, RO. Box 8010,

6700 ED Wageningen, The Netherlands.

**Harrelaers 1, 1852 KT Heiloo, The Netherlands.

MS. received: 1990.03.21.

POACEAE

A NEW SPECIES OF ERAGROST1S (CHLORIDEAE) IN THE E. CURVULA ALLIANCE

Eragrostis comptonii De Winter, sp. nov. , E.

planiculmi affinis sed culmis teretibus gracilibus nervosis,

non complanatis, caespite basin systemate ramoso

rhizomatum brevium, inflorescentia apertiori, ramulis

ultimis patentibus, spiculis (2)3 vel 4(5)-floris differt.

Gregi E. curvulae affinis, ut ab hybridis putativis inter

taxa duo indicatum.

TYPE. — Swaziland, 2631 (Mbabane), hill NE of Mba-

bane, alt. 4000’ (555,2 m), 13.3.1957, Compton 26766

(PRE, holo.; NBG, Swaziland Herbarium, iso.).

This species is named in honour of Prof. R.H. Compton

who has contributed more than any other person to the

expansion of the knowledge of the flora of Swaziland, and

by whom the type specimen as well as several other

specimens of this new species was collected.

Perennial up to 1,1 m high (usually about 700 mm),

forming dense, fine-leaved tufts from a system of short,

branched, knotty rhizomes, densely clothed by short

cataphylls. Culms thin to very slender, 1,0 to 1,5 mm in

diameter, glabrous, strictly erect, wiry. Nodes (4— )8 or

9 per culm, hidden by the leaf sheaths. Internodes 3 or

4, basal ones very short (4-6 mm) increasing abruptly

in length upwards, the upper about 100 mm long, glabrous.

Leaves 4—9, borne mainly on the culms. Leaf sheaths

increasing in length upwards on the culm, much longer

than the internode it subtends, glabrous. Collar incon-

spicuous, the blade rather gradually merging into the

sheath. Auricles very finely scabrid on the margin, pallid

or purplish. Ligule a very short fine line of bristly hairs.

Leaf blades up to 500 mm long, usually 300—400 mm;

when expanded up to 4 mm wide at the base; mostly

convolute, the rolled leaf very narrow, ending in a filiform

apex; glabrous and smooth abaxially, very finely and

densely scabrid adaxially; primary nerves up to 11,

alternating with (2)3 or 4 secondary nerves; strongly raised

adaxially and scabrid with very fine spines. Inflorescence

a diffuse open panicle, with green or purplish, filiform,

glabrous or finely scabrid branches; branches patent, the

spikelets borne terminally and discretely (not clustered).

Spikelets borne on filiform pedicels (3)4—8 mm long;

(2)3 or 4(-6)-flowered. Glumes somewhat unequal to

subequal; 1-nerved, firmly membranous, boat-shaped,

lanceate in outline, ± 1,5 mm long; apex acute. Lemmas

gunmetal-grey, 1,25—1,50 mm long, 3-nerved, boat-

shaped, broadly ovate in outline with a subacute apex.

Palea ± 1,25—1,50 mm long, greyish, submembranous,

bicarinate, the keels curved. Lodicules 2, ± 0,3 mm long,

fleshy, truncate. Stamens 3; anthers ± 1 mm long, linear,

yellow. Pistil ± 0,8 mm long; ovary ovoid; styles 2;

stigmas narrowly and sparsely plumose. Caryopsis ovoid

to ovoid-oblate, 0, 8-1,0 mm long, dark chestnut-brown,

semitranslucent; embryo nearly half the length of the

grain, broadly oblong in outline; hilum basal, puntiform.

SWAZILAND. —2631 (Mbabane): Malolotja Nature Reserve,

10.3.1986 (— AA), Braun 60 (PRE); 22 km NE of Mbabane on road to

Mapalaleni (— AA), De Winter 9816 (cult, in PHBG) (PRE); hill NE

of Mbabane, 2.3.1959 (—AC), Compton 28551 (PRE); hill NE of

Bothalia 20,2 (1990)

209

Mbabane, shade of rocks, 20.2.1958 (—AC), Compton 27551 (PRE); hill

NE of Mbabane, 5.4.1965 (—AC), Compton s.n. ; 7 km NE of junction

of Mbabane-Manzini roads on the Tea Road, 12.6.1985 (—AC), De Winter

9877 (cult, in PNBG) (PRE).

The presently known distribution of the species is

limited to Swaziland in the vicinity of Mbabane. Its

ecology is poorly known. Judging from the notes on

specimens it tends to favour somewhat shady places such

as at the foot of rocks or on forest margins.

There are strong indications from studies of populations

in the field that E. comptonii hybridizes with E. curvula

sens. lat. The presence of short knotty rhizomes in

‘typical’ specimens of E. comptonii, a feature which is

relatively rare in the genus and always absent in E. curvula,

first drew attention to the existence of this species. Many

of the specimens collected near the type locality are

intermediate between the two species. This may be part

of the explanation for the occurrence of forms of E. curvula

with thin wiry very erect culms which occur in the

Mbabane area in Swaziland. The habitats favoured by E.

comptonii, seem to be partially shady, relatively moist and

relatively stable, whereas the putative hybrids are found

in disturbed areas, as is usual for most populations of

Eragrostis curvula sens. lat. More intensive investigations

into the variable populations of E. curvula in the eastern

highlands are needed, particularly concerning certain E.

curvula forms which have lax open inflorescences.

Eragrostis planiculmis Nees, a species known to hybridize

with E. curvula sens, lat., has lax, open, nebulous

inflorescences with slender spikelets and very robust

flattened culms unlike those of the putative hybrids. The

distribution of the aberrant forms, moreover, is discordant

with the wider area of overlap of the distribution of E.

curvula sens. lat. and E. planiculmis. E. planiculmis is

therefore an unlikely candidate parent.

Another species regarded as a putative parent which

could have contributed to the lax open inflorescences

of E. curvula sens. lat. by hybridization, namely E.

chloromelas, has a present day distribution which makes

the possibility of a link with the eastern highland forms

very remote.

Before the discovery of E. comptonii the origin of these

forms with lax inflorescences and wiry culms was

therefore, very puzzling. E. comptonii, on the other

hand, possesses all the features necessary to explain

both the distribution and the particular pattern of variability

of E. curvula sens. lat. in certain parts of the eastern

highlands. The hypothesis that E. comptonii played a

role in the establishment of E. curvula types with wiry

culms and open inflorescences in the Mbabane and

adjacent areas of the eastern highlands should be tested

experimentally.

Examples of putative hybrids between E. curvula sens,

lat. and E. comptonii are De Winter 9813, 9811 and

9812. The first closely resembles E. comptonii but lacks

rhizomes, whereas 9811 and 9812 are progressively more

like E. curvula. Dlamini s.n.., collected on 5.4.1965, also

has intermediate characters.

The description of this species forms part of the study

of the Eragrostis curvula complex, which is being under-

taken by the author and a number of collaborators.

B. DE WINTER*

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

MS. received: 1990.02.06.

CUCURBITACEAE

A NEW SPECIES OF CITRULLUS (BENINCASEAE) FROM THE NAMIB DESERT, NAMIBIA

Citrullus rehmii De Winter sp. nov. , C. lanato

(Thunb.) Matsumura & Nakai affinis, a quo floribus

minoribus, seminibus minoribus atrobrunneis et fructibus

atroviridibus subtiliter salmoneo-maculosis differt. A C.

ecirrhoso Cogn. et C. colocynthidi (L.) Schrad. habitu

annuo non perenne, atque superficie fructuum subtiliter

maculosa differt.

TYPE.— -Namibia, near Ugab River on road from Uis

to Welwitschia [Khorixas] , near small ‘Inselberge’ of red

granite on sandy flats, 28.3.1963, De Winter & Hardy 8127

(PRE, holo.; K, NBG, WIND, iso.).

Monoecious, creeping, herbaceous annual producing

procumbent stems radiating from a central taproot. Matufe

vegetative parts coarsely scabrid with bristly hairs; young

parts densely covered with a mixture of bristly , bulbous-

based, glasslike, sharp-pointed, septate hairs and yellow-

ish, blunt, glandular hairs with a somewhat granular

appearance; bulbous bases of bristles increase in size as

vegetative parts mature, becoming opaque and white and

the glandular hairs largely disappear. Stems simple or

branched, 0,3—2 m long, sparsely to densely bristly, rough

to the touch. Leaves petiolate; petioles sparsely bristly,

(10 — )30( — 70) mm long; leaf blade triangular-cordate to

broadly cordate in outline, (50— )60(— 120) x (30— )50

(-60) mm, deeply dissected into 3—5 lobes, each lobe

strongly undulate and sinuous on the margin or deeply

lobed, usually very densely covered with bulbous-based

hairs on the lower surface, upper surface glabrous except

for bristles on the veins, on and near the leaf margin.

Tendrils very "slender, one at each node, bifurcate or

rarely simple, branches strongly coiled. Flowers solitary,

axillary, yellow, pedicellate. Male flowers articulated to

the peduncle at the base of the receptacle; receptacle semi-

hyaline, 2—3 mm deep. Calyx lobes firm, green, linear,

2—3 mm long, bristly on the outside. Corolla shallowly

saucer-shaped 17—20 mm in diameter when open, lobes

210

Bothalia 20,2 (1990)

7—8 mm long, ovate to broadly ovate; with 5—7 strong,

parallel main veins interconnected by a network of

secondary veins; main veins protruding on the lower

surface and bristly; margins and lower surface sparsely

covered with multicellular, blunt, shortly stalked,

glandular hairs. Stamens 3, filaments + 2 mm long;

adaxially bearded at the base; anther-thecae conduplicate.

Ovary totally absent. Female flowers with the receptacle

tube ± 1 mm long. Calyx lobes 5, linear, firm, green,

inserted on the receptacle, 2,5— 3,0 mm long, like the

receptacle covered with glassy, septate, bristly, sharp-

pointed hairs. Corolla lobes 5, demarcated from the

receptacle by a distinct rim linking the bases of the calyx

lobes, lobes ovate to broadly ovate, ± 6 mm long, apex

and margins densely covered with glandular hairs.

Staminodes 3 or 4, opposite the calyx lobes, reduced to

triangular, often fleshy scales, bearded at the base with

hyaline septate bristles and crowned by a sterile reduced

anther. Ovary inferior, subglobose, densely villous with

slender, hyaline, septate hairs up to 2 mm long; with 3

parietal placentae and numerous ovules. Fruit globose,

(80 — )100( — 120) mm in diameter, surface deep green,

finely mottled with irregular salmon-pink to orange-pink

spots, glabrous except for persistent, sparsely scattered,

hyaline, septate hairs; pulp white, very bitter. Seeds

straw-coloured to dark chestnut-brown, flattened, broadly

obovate in outline, ± 6,0 X ± 4,5 mm.

NAMIBIA.— 2014 (Khorixas): near Ugab River mouth on road from

Uis to Khorixas [Welwitschia] ( — DD), De Winter & Hardy 8127 (PRE,

holo.); between Uis and Khorixas (-BD), Toelken & Hardy 838 (K,

PRE, WIND). 2114 (Uis): Brandberg ( — BA). Giess 3712 (PRE, WIND);

112 km NE of Henties Bay, direction Uis (— BC), Chadwick 62 (K, NBG,

PRE). 2115 (Karibib): Klein Spitskoppe (— CC), Giess 8791 (WIND).

2819 (Warmbad): flats just east of Karasburg ( — BB), Oliver & Steenkamp

6255 (PRE).

Specific epithet

This species is named in honour of Prof. Sigmund Rehm

who, jointly with Dr P.R. Enslin, undertook intensive

studies of the chemistry and genetics of the South African

Cucurbitaceae including the genus Citrullus.

Distribution and ecology

Except for one record from east of Karasburg in the

arid south of Namibia, all collections of C. rehmii were

made in the Namib Desert from Khorixas [Welwitschia]

in the north to a little south of Klein Spitskoppe. A

photograph taken by P. Steenkamp at Sesriem on the

Namib Desert edge, in 1976, on which fruits of C. rehmii ,

C. ecirrhosus and C. lanatus are grouped together, seems

to indicate that all three species are found at this locality.

However, no records of any of the species from Sesriem

are known to me, so that this assumption could not be

verified. The record from Karasburg seems somewhat

doubtful and needs checking. Citrullus ecirrhosus , the

second desert species in southern Africa, is found in the

Namib Desert from north of the Brandberg to Kuboos in

the Richtersveld of the northern Cape. It is therefore

possible that the range of the two species may prove to

be very similar. C. rehmii apparently inhabits mainly the

gravelly to sandy-gravelly flats, avoiding the areas with

deeper sand.

Relationships

Little is known about the relationships of C. rehmii with

the three other species of Citrullus. It has been success-

fully hybridized with the wild form of C. lanatus as well

as with the bitter form of the watermelon cultivar

‘Hawkesbury’, also classified as C. lanatus, by Rehm &

Neethling (Unpublished Project Report 1967). Ellis in

1968 successfully hybridized C. rehmii with sweet tsam-

ma C. lanatus as well as C. ecirrhosus and C. colocynthis.

The FI hybrids of all crosses are fertile and strongly

intermediate in character, particularly in as far as

the mottling of the fruits is concerned. The degree of

sterility in the F2 progeny was not tested (R.P. Ellis

unpublished). Rehm & Neethling, however, found that the

F2 progeny of the crosses C. lanatus x colocynthis and

C. lanatus x ecirrhosus showed a high degree of sterility.

The F2 progeny of the crosses of C. colocynthis x ecir-

rhosus proved to be fertile. Because of lack of flowering

material no crosses with C. rehmii were made by Rehm

& Neethling. These unpublished experiments show that

C. lanatus is related to C. ecirhosus and C. colocynthis

but clearly specifically distinct, on morphological as well

as genetical grounds.

C. ecirrhosus and C. colocynthis are morphologically

somewhat similar but easily distinguished on the size and

mottling of the fruits, the presence or absence of tendrils

and the disjunct distribution.

As a matter of convenience the latter two species should,

on basis of morphological and phytogeographical differ-

ences, be retained as separate species, in spite of their

genetic compatibility.

As soon as material becomes available, crosses between

C. rehmii and the other species will be undertaken to

establish the degree of fertility of the progeny in the F2.

On morphological grounds it is predicted that C. rehmii

will prove to be genetically more closely related to C. ecir-

rhosus and C. colocynthis than to C. lanatus.

The fruits of all Citrullus species contain various

quantities of cucurbitacin E, an oxygenated tetracyclic

triterpene. C. rehmii contains cucurbitacin E and traces

of cucurbitacin B and I (Enslin & Rehm 1958). The

chemistry of the bitter principles contributes little to the

elaboration of relationships. The relationship with C.

lanatus is put forward mainly on the evidence of the annual

habit, low cucurbitacin E content and the presence of

tendrils. The leaf characters are similar in general features

to those of C. ecirrhosus and C. colocynthis and taken in

isolation, would favour a closer relationship with C. ecir-

rhosus. C. rehmii differs from all the other species by the

characteristic salmon-pink mottling on a dark green

background, of the rind of the fruit.

Uses

The fruits of hybrids made by R.R Ellis at the Botanical

Research Institute in 1967 were investigated by Enslin. The

cross between C. lanatus cultivar ‘Hawkesbury-bitter’ x

Bothalia 20,2 (1990)

211

C. rehmii yielded 200 mg/100 g glycoside, which in turn

yielded 91 mg/100 g cucurbitacin E. This was considerably

higher than the yield of hybrids of the non-bitter wild form

of C. lanatus x rehmii as well as that of C. colocynthis

X C. rehmii. Cucurbitacin E was used by Enslin to

synthesize alternative steroid hormones (P.R. Enslin pers.

comm. 1967).

REFERENCE

ENSLIN, P.R. & REHM, S. 1958. The distribution and biogenesis of

the cucurbitacins in relation to the taxonomy of the Cucurbitaceae.

Proceedings of the Linnean Society of London 169,3: 230—238.

B. DE WINTER

MS. received: 1990.02.06.

BRYOPHYTA

NEW AND INTERESTING RECORDS OF MOSSES IN THE FLORA OF SOUTHERN AFRICA AREA:

1. SPHAGNACEAE— GRIMMIACEAE

Since the publication of the first fascicle of the moss

Flora of southern Africa (Magill 1981), several new and

interesting records have been identified from geographical

regions referred to in the fascicle. Most of the new records

are a result of recent collecting by the authors and other

staff members of the National Botanical Institute. However,

large parts of the flora area remain under-collected,

especially the arid and semi-arid central and northern

regions.

New and interesting records are listed in the same

taxonomic order as in the first fascicle of the moss flora.

The quarter-degree grid reference (Edwards & Leistner

1971), name of the collector and collector’s number or

PRE-Cryptogamic Herbarium number are given for each

voucher specimen. All specimens cited are in PRE unless

otherwise indicated. New taxa, new records and name

changes for southern African mosses are reported in the

List of species of southern African plants (1985) and up-

dates to the list in Bothalia (Van Rooy 1985, 1986, 1988,

1989).

SPHAGNACEAE

Sphagnum

truncatum Hornsch. (Magill 1981:29)

Northern Transvaal (2230 CC: Bosnian PRE-CH 1365. 2329 CD:

Codd <6 Dyer PRE-CH 12235). eastern Transvaal (2430 DB: Van

der Schijff. 2530 BB: Sim 8014. 8580). Zululand (2831 DC: Sim

9671; Venter 2478) and the eastern Cape (3227 CB: Sim 7166).

capense Hornsch. (Magill 1981:31)

Northern Transvaal (2329 AA: Leeman PRE-CH 3649).

ANDREAEACEAE

Andreaea

subulata Harr, ex Hook. (Magill 1981:33)

Previously known only from Table Mountain, the species has

recently been identified from other localities in the south-western

Cape (3319 CA: Esterhuvsen 21806, 22270. 3319 CB: Esterhuysen

20031).

rupestris Hedw. (Magill 1981:35)

Transkei (3128 BC: Van Rooy 2277) and southern Cape (3324 DB:

Esterhuysen 28046).

FISSIDENTACEAE

Fissidens

usambaricus Broth. (Magill 1981: 43)

Transkei (3129 CB: Van Rooy 1928).

bryoides Hedw. (Magill 1981: 46)

Recently collected in the Transkei (3129 BA: Van Rooy 1826.

3228 BD: Van Rooy 2064) and the southern Cape (3323 DD:

Wagener PRE-CH 13328).

hoeegii P. Varde (Magill 1981:47)

Northern Transvaal (2329 DD: Brenan M3226) and the central Cape

(3224 BC: MacLea sub Rehm 588).

rufescens Hornsch. (Magill 1981:47)

Southern Transvaal (2628 CB: Perold 252) and the eastern Cape

(3027 CA: Van Rooy 2615, 2618. 2630. 3027 CC: Van Rooy 2665.

2668, 2677).

simii Schelpe (Magill 1981:50)

Now also known from the Orange Free State (3026 BD: Van Rooy

2591).

curvatus Hornsch. (Magill 1981:51)

Transkei (3128 DC: Van Rooy 2158. 3228 AB: Van Rooy 2031).

aciphyllus Dix. (Magill 1981: 53)

Zululand (2832 CC: Van Rooy 180).

submarginatus Bruch ex Krauss (Magill 1981:54)

Northern Cape (2722 AB: Magill 6401, 6402 , 6403, 6406. 2722

DC: Magill 6411).

parvilimbatus Sim (Magill 1981:57)

Southern Transvaal (2627 AC: Perold 1987).

borgenii Hampe (Magill 1981:58)

Central Transvaal (2527 CA: Magill 3005. 2529 CB: Magill 6387,

Van Rooy & Perold 652).

erosulus (C. Mull.) Par. (Magill 1981:58)

Northern Cape (2722 AB: Van Rooy 686. 2723 AD: Van Rooy 682.

2822 BA: Van Rooy 698) and Natal'(2930 AD: Wood 46. 3130 AA:

Van Rooy 961).

subobtusatus C. Mull. (Magill 1981:59)

Northern Cape (2722 AB: Magill 6405, Van Rooy 685. 2722 DC:

Van Rooy 697. 2822 BA: Magill 6420) and Natal (2729 DD: Bews

sub Sim 8267. 2828 DB: Van Rooy 1118. 2930 C'B: Sim 9945. 2930

DA: Sim 8158, Van Rooy 1515, 1522). A specimen collected in

southern Namibia (2718 CA: Magill 6426) is an additional record

for the region.

fontanus (B. Pyl.) Steud. (Magill 1981:67)

Eastern Cape (3027 CA: Net 21. 3027 CD: Sim 7096).

NANOBRYACEAE

Nanobryum dummeri Dix. (Magill 1981:69)

Previously known only from Port St Johns in the Transkei, the

species has recently been collected in a Zululand forest (2832

AB: Harrison 846).

ARCHIDIACEAE

Archidium

dinteri ( Irmsch .) Snider (Magill 1981: 72)

An additional record of this rare species is known from central

Namibia (2217 CB: Volk 84/722).

ohioense Schimp. ex C. Mull. (Magill 1981: 74)

Natal (2929 AA: Esterhuysen 34591), Lesotho (2929 CB: Magill

4401), the north-western Cape (3118 CD: Perold 1731) and Transkei

(3130 AC: Van Rooy 1745).

rehmannii Mitt. (Magill 1981: 77)

North-western Cape (3118 CD: Perold 1732. 3119 DD: Perold 1845).

julicaule C. Midi. (Magill 1981: 80)

Recently collected in Lesotho (2929 CB: Magill 7087).

DITRICHACEAE

Pleuridium

pappeanum (C. Mull. ) Jaeg. (Magill 1981: 84)

Lesotho (2929 CB: Magill 4398) and Transkei (3228 AB: Van Rooy

2028).

nervosum (Hook.) Mitt. (Magill 1981: 86)

Orange Free State (2827 DA: Van Rooy 447. 2827 DB: Van Rooy

421. 2828 AC: Perold 1351) and Lesotho (2929 CB: Magill 4378.

4389).

Eccremidium exiguum (Hook. f. & Wils.) Wils. ex Salm. (Magill 1981: 89)

Central Transvaal (2428 CA: Perold 810. 2528 DC: Perold 1373)7

212

Bothalia 20,2 (1990)

Ditrichum

strictum (Hook. f. & Wils.) Hampe (Magill 1981: 90)

Lesotho (2828 DD: Van Zanten et al. 7609922, 7609978. 2929 CA:

Van Rooy 3416. 3417).

difficile (Dub.) Fleisch. (Magill 1981: 93)

Now also known from the northern Transvaal (2329 DD: Brenan

M 3234; Magill 6552).

DICRANACEAE

Cladophascum gymnomitrioides (Dix.) Dix. ex Sim (Magill 1981: 110)

Northern Transvaal (2329 DB: Brenan M 3222), central Transvaal

(2529 CB: Volk 81/018), eastern Transvaal (2630 AD: Smook 4893.

2730 AC: Tolken 5718), north-western Cape (3018 AC: Perold 2129)

and south-western Cape (3319 AD: Perold 2408).

Dicranella

symonsii Dix. (Magill 1981: 122)

South-western Cape (3218 BB: Perold 1921. 3219 AC: Perold 1954).

rigida Dix. ex Sim (Magill 1981: 122)

North-western Cape (3118 DB: Oliver 7277).

Oreoweisia erosa (C. Miill.) Kindb. (Magill 1981: 127)

New from the Orange Free State (2729 CD: Perold 1272).

Leucoloma

syrrhopodontioides Broth. (Magill 1981: 136)

Northern Transvaal (2230 CC: Von Breitenbach 205).

rehmannii (C. Mull.) Rehm. ex Par. (Magill 1981: 136)

Recently collected in the Transkei (3129 BC: Van Rooy 1849. 3130

AC: Van Rooy 1780).

Microcampylopus perpusillus (Mitt.) Broth. (Magill 1981: 138)

Central Transvaal (2429 DD: Perold 984. 2529 CB: Van Rooy &

Perold 640).

Campylopus

delagoae (C. Miill.) Par. (Magill 1981: 140)

Northern Transvaal (2330 CC: Perold 2452).

stenopelma (C. Miill.) Par. (Magill 1981: 142)

Northern Transvaal (2330 CC: Brenan M 2811).

subchlorophyllosus C. Miill. ex Rabenh. (Magill 1981: 144)

Lesotho (2828 DA: Dealt & Killick 107, 109. 2929 CB: Deall &

Killick 161b).

pallidus Hook. f. & Wils. (Magill 1981: 147)

New from western Transvaal (2526 DD: Perold 889) and the

Transkei (3129 BC: Van Rooy 1874).

introflexus (Hedw.) Brid. (Magill 1981: 150)

Northern Transvaal (2230 CC: Bosman PRE-CH 1356. 2330 CC:

Von Breitenbach 101).

pilifer Brid. (Magill 1981: 152)

Southern Cape (3322 AC: Cholrtoky 1020. 3325 CD: Smook 3825a).

procerus (C. Miill.) Par. (Magill 1981: 153)

Northern Transvaal (2329 DD: Magill 6516).

CALYMPERACEAE

Calymperes rabenhorstii Hampe & C. Miill. (Magill 1981:164)

Previously known only from a single specimen collected in Zululand,

the species is now also known from the eastern Transvaal (2430 DB:

Retief et al. 1139a).

Syrrhopodon

uncinifolius C. Miill. (Magill 1981: 167)

Transkei (3128 BC: Van Rooy 2262, 2264. 3129 BC: Van Rooy 1882,

1843, 1908. 3130 AC: Van Rooy 1762, 1784).

obliquirostris C. Miill. (Magill 1981: 169)

Recently collected in the Transkei (3129 BC: Van Rooy 1880).

ENCALYPTACEAE

Encalypta vulgaris Hedw. (Magill 1981: 174)

New from the eastern Cape (3027 CD: Van Rooy 2792. 3027 DC:

Van Rooy 2751, 2762).

POTTIACEAE

Anoectangium wilmsianum (C. Miill.) Par. (Magill 1981: 181)

Eastern Cape (3027 CB: Van Rooy 2715. 3027 DD: Oliver 8357).

Hymenostylium recurvirostrum (Hedw.) Dix. (Magill 1981: 182)

Transkei (3129 BA: Van Rooy 1839).

Gymnostomum aeruginosum J.E. Sm. (Magill 1981: 183)

Recently collected in the eastern Transvaal (2630 CC: Perold 1082)

and the eastern Cape (3027 CA: Van Rooy 2622).

Aloina bifrons (De Not.) Delgadillo (Magill 1981:193)

Now also known from the northern Cape (2823 BC: Magill 6491a).

Acaulon rufochaete Magill (Magill 1981: 201)

Northern Cape (2824 DA: Magill 6501).

Phascum

peraristatum C. Miill. (Magill 1981: 202)

Northern Cape (2824 DA: Magill 6497).

leptophyllum C. Miill. (Magill 1981: 204)

Northern Transvaal (2329 BB: Ellis PRE-CH 13109), central

Transvaal (2528 CA: Steyn PRE-CH 12921. 2529 CB: Magill 6362;

Van Rooy & Perold 667), southern Transvaal (2628 AA: Brenan

M 3364), north-western Cape (2917 DB: Perold 1394a) and

southern Cape (3423 AA: Cholnoky 594, 616).

Tortula

porphyreoneura (C. Miill.) Townsend (Magill 1981: 213)

Central Transvaal (2528 CA: Perold 127), northern Cape (2624

DD: Perold 1381), north-western Cape (2817 CA: Oliver et al. 1651)

and south-western Cape (3318 DD: Perold 1197).

ammonsiana Crum & Anderson (Magill 1981: 216)

Northern Cape (2723 AD: Magill 6396, 6399), north-western Cape

(2919 AC: Magill 6461), Transkei (3129 CA: Van Rooy 1984, 2172)

and the eastern Cape (3027 CB: Van Rooy 2709. 3027 CD: Van

Rooy 2785).

pagorum (Milde) De Not. (Magill 1981: 217)

Western Transvaal (2526 CA: Magill 6388) and the northern Cape

(2625 AB: Magill 6392. 2822 BA: Magill 6421, 6422, 6423. 2822

CB: Van Rooy 746).

papillosa Wils. ex Spruce (Magill 1981: 218)

Recently found in the Orange Free State (2828 AB: Perold 1358).

chisosa Magill, Delgadillo & Stark (Magill et al. 1983)

Central Transvaal (2428 CC: Smook 2714), eastern Transvaal (2430

CC: Magill 3134), north-western Cape (2816 DB: Oliver et al. 548.

2817 AC: Van der Westhuizen PRE-CH 13115. 2917 BA: Magill 6436,

6440; Van Rooy 712, 715. 2917 DB: Perold 1389, 1392; Schelpe

7751, 7763; Smook 3593b. 2918 CA: Perold 2058) and the south-

western Cape (3218 BB: Magill & Schelpe 4035. 3219 AA: Schelpe

4970).

Weisiopsis involuta Magill (Magill 1981: 225)

Previously known only from the type locality in Botswana, the spe

cies has recently been collected on limestone in northern Namibia

(1816 DB: Brusse 4176).

Hyophila

involuta (Hook.) Jaeg. (Magill 1981: 228)

Northern Transvaal (2329 DB: Magill 6507) and central Transvaal

(2529 AD: Perold 965, 966, 967. 2529 CB: Magill 6384).

baginsensis C. Miill. (Magill 1981: 230)

Now also known from the central Transvaal (2527 CA: Venter 2018).

Didymodon

australasii (Hook. & Grev.) Zander (Magill 1981: 232, Van Rooy 1989)

Natal (2929 CB: Van Rooy 1365).

ceratodonteus (C. Miill.) Dix. (Magill 1981: 233)

Northern Cape (2724 BA: Van Rooy 681).

xanthocarpus (C. Miill.) Magill (Magill 1981: 235)

Recently collected in dry river valleys of eastern Lesotho (2928

BB: Van Rooy 3217. 2928 BD: Van Rooy 3191. 2929 AC: Van Rooy

3319).

Barbula

crinita Schultz (Magill 1981: 237)

Southern Transvaal (2627 BA: Mogg 35661. 2628 AD: Mogg

37590a).

acutata C. Miill. (Magill 1981: 239)

Central Cape (2922 BC: Smook 4484. 3025 CA: Smook 3239).

eubryum C. Miill. (Magill 1981: 245)

Northern Transvaal (2328 CC: Smook 4193).

afrofontana (C. Miill.) Broth. (Magill 1981: 246)

Central Transvaal (2527 DD: Perold 246) and southern Transvaal

(2627 AC: Perold 1985).

Bryoerythrophyllum jamesonii (Tayl.) Crum (Magill 1981: 248)

New for the eastern Cape (3027 DC: Russell 3761) and Transkei

(3129 BB: Van Rooy 1706).

Phasconica tisserantii P. Varde (Magill 1981: 251)

Southern Cape (3325 BB: Schrire 1889).

Timmiella pelindaba Magill (Magill 1981: 253)

Now also known from the eastern Cape (3027 CA: Van Rooy 2637,

2643).

Tortella

xanthocarpa (C. Mull.) Broth. (Magill 1981: 255)

Lesotho (2828 DC: Deall & Killick 85. 2929 CA: Van Rooy 3426.

2929 CB: Deall & Killick 149), the eastern Cape (3027 CB: Van

Rooy 2725. 3227 DD: Van Rooy 940) and Namibia (1823 AB: Hardy

5601).

fragilis (Hook. & Wils.) Limpr. (Magill 1981: 258)

Eastern Cape (3127 BB: Hilliard & Burtt 13146).

Bothalia 20,2 (1990)

213

Oxystegus cylindricus (Brid.) Hilp. (Magill 1981: 259)

Lesotho (2929 CC: Magill 4312).

Weissia

dieterlenii Ther. (Magill 1981: 263)

Eastern Cape (3027 CC: Van Rooy 2660. 3027 DC: Van Rooy 2765).

humicola C. Mull. (Magill 1981: 266)

Transkei (3128 BC: Van Rooy 2274).

latiuscula C. Mull. (Magill 1981: 267)

Now also known from the eastern Cape (3027 CB: Van Rooy 2710).

GRIMM1ACEAE

Grimmia

apocarpa Hedw. (Magill 1981: 274)

New from the Transkei (3228 BD: Van Rooy 2072. 2073).

ovalis (Hedw.) Lindb. (Magill 1981: HI)

Recently collected in the Transkei (3128 BC: Van Rooy 2282).

laevigatam (Brid.) Brul. (Magill 1981: 211)

Namibia (2816 BC: Van der Westhi(izen PRE-CH 13112a) and the

northern Cape (2822 CB: Van Rooy 747).

REFERENCES

EDWARDS, D. & LE1STNER, O.A. 1971. A degree reference system

for citing biological records in southern Africa. Mitteilungen der

Botanischen Staatssammlung, Miinchen 10: 501—509.

MAGILL, R.E. 1981. Bryophyta. In O.A. Leistncr, Flora of southern

Africa. Part. I Mosses, Fascicle I Sphagnaceae— Grimmiaceae.

Botanical Research Institute, Department of Agriculture and

Fisheries, Republic of South Africa.

MAGILL. R.E., DELGADILLO, C. & STARK, L.R. 1983. Tortula

chisosa sp. nov., a bistratose-leaved species from the United States,

Mexico and southern Africa. Annals of the Missouri Botanical

Garden 70: 200-202.

VAN ROOY, J. 1985. Bryophyta. In G.E. Gibbs Russell el al.. List of

species of southern African plants, Edn 2, Part 1. Memoirs of

the Botanical Survey of South Africa No. 51: 1—31.

VAN ROOY, J 1986. Bryophyta In Staff of the National Herbarium,

New taxa, new records and name changes for southern African

plants. Bothalia 16: 109, 110.

VAN ROOY, J. 1988. Bryophyta. In G.E. Gibbs Russell et al.. New taxa,

new records and name changes for southern African plants.

Bothalia 18: 294, 295.

VAN ROOY, J. 1989. Bryophyta. In B.C. de Wet et a I., New taxa, new

records and name changes for southern African plants. Bothalia

19: 276.

J. VAN ROOY* and S.M. PEROLD*

* National Botanical Institute, Private Bag X101, Pretoria 0001

LILIACEAE/ASPHODELACEAE

THE TYPE OF CHORTOURION BERGERIANUM (ALOOIDEAE)

Berger (1908) erected the genus Chortolirion to accom-

modate the four bulbous species of Haworthia known at

the time. Six years after this genus was established, Dinter

(1914) described a fifth species as C. bergerianum from

specimens which he collected near Windhoek in Namibia.

Obermeyer (1973) sank the four Chortolirion species recog-

nized by Berger (1908) in the synonymy of H. angolensis

Baker. However, she did not mention C. bergerianum in

her treatment of Chortolirion.

A revision of the smaller genera of the subfamily

Alooideae of the Asphodelaceae currently under way has

shown that Chortolirion represents a natural, monotypic

entity which should be afforded separate generic recog-

nition (Smith 1985, 1988). In an attempt to establish the

identity of C. bergerianum , I became aware that Dinter

failed to designate a type specimen when he described this

species. In this paper I present the results of my efforts

to typify the name C. bergerianum.

According to the International Code of Botanical

Nomenclature (Articles 7.1 and 7.2) all taxa of the rank

of family or below must have a nomenclatural type

(Greuter et al. 1988). However, no Dinter material

collected in January 1913 from the type locality (‘Voigt-

land’, 20 km east of Windhoek) could be traced in any of

the herbaria listed by Holmgren et al. (1988), Stafleu &

Cowan (1976) or Gunn & Codd (1981). The following

discussion should clarify the typification of C. bergeri-

anum'.

1, Dinter (1914) did not cite any herbarium specimens

nor did he mention any plant material which he used in

establishing this taxon; 2, no specimens collected in

January 1913 at the type locality and used (Article 7.3)

or annotated by Dinter in preparing the description, but

not cited or designated as type, could be found; 3, Dinter

(1914) included a single reference to an illustration (Fig.

12) which was published along with the original descrip-

tion of C. bergerianum in Neue und wenig bekannte

Pflanzen Deutsch-Sudwest-Afrikas (1914); 4, this figure,

which therefore forms part of the protologue, is of a sterile

specimen, but clearly agrees with Dinter’s description; 5,

Article 7.3 and Note 1 of this article clearly specify that

any designation made by the original author, if definitely

expressed at the time of the original publication of the

name of a taxon, is final . Furthermore, the type of a name

may be either a specimen or an illustration.

After considering the existing evidence, I concluded that

Dinter’s illustration, as an element included in the

protologue, is the type of C. bergerianum . Since it is the

only element cited, and no specimens used or annotated

by Dinter can be found, the illustration should be referred

to as the holotype and not treated as a lectotype. The type

of C. bergerianum is:

TYPE. — K. Dinter, Neue und wenig bekannte Pflanzen

Deutsch-Sudwest-Afrikas t. 12 (1914) (icono.).

ACKNOWLEDGEMENTS

I am grateful to Dr D.J.B. Killick for his detailed com-

ments on an earlier draft. The curators of the following

herbaria are thanked for the loan of herbarium specimens

and/or permission to study their collections during visits:

B, BM, BOL, HBG, J, K, KMG, M, NBG, PUC, PRE,

PRU, SAM, STE, Z.

REFERENCES

BERGER, A. 1908. Liliaceae-Asphodeloideae-Aloineae. In A. Engler.

Das Pflanzenreich 4.38.3.2 (Heft 33): 72 —74. Wilhelm Engel-

mann, Leipzig.

214

Bothalia 20,2 (1990)

DINTER, M.K. 1914. Neue und wenig bekannte Pflanzen Deutsch-

Slidwest-Afrikas unter besonderer Beriicksichtigung der Succulen-

ten. 1m Selbstverlag, Okahandja.

GREUTER, W. et al. 1988. International Code of Botanical Nomen-

clature. Koeltz Scientific Books, Konigstein. Regnum Vegetabile

118.

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

Africa. Balkema, Cape Town.

HOLMGREN, P.K., KEUKEN, W. & SCHOFIELD, E.K. 1981. Index

Herbariorum. Part 1. The herbaria of the world, 7th edn. Bohn,

Scheltema & Holkema, Utrecht. Regnum Vegetabile 106.

OBERMEYER, A. A. 1973. Aloe, Chamaealoe, Haworthia, Astroloba,

Poellnitzia and Chortotirion (Liliaceae). Bothalia 11: 119.

SMITH, G.F. 1985. Die taksonomiese status van die monotipiese genusse

Chortolirion Berger en Poellnitzia Uitewaal in die tribus Aloineae

(familie Asphodelaceae) . B.Sc. Hons, paper. University of

Pretoria.

SMITH, G.F. 1988. A scanning electron microscopic investigation of

the pollen morphology of Chortolirion Berger (Aloineae;

Liliaceae). South African Journal of Science 84: 428—430.

STAFLEU, F.A. & COWAN, R.S. 1976. Taxonomic literature , Vol. 1:

A— G, 2nd edn. Bohn, Scheltema & Holkema, Utrecht. Regnum

Vegetabile 94.

G.F. SMITH*

* Department of Plant Sciences, Potchefstroom University for Christi-

an Higher Education, Potchefstroom 2520.

MS. received: 1989.10.12.

MARCH ANTIALES

SPORE GERMINATION, EARLY PROTONEMA DEVELOPMENT AND VEGETATIVE REPRODUCTION IN RICCIA,

SECTION PHIFER

Spore germination in Riccia has been investigated by

a number of workers, commencing well over a hundred

years ago with the work done by Hofmeister (1851), Kny

(1867) and Fellner (1875). Not all researchers were,

however, in agreement about the site of exit of the germ

tube, for example, Campbell (1918) maintained that in R.

trichocarpa Howe, the exospore ruptures at the triradiate

mark. Subsequently, it has been repeatedly reported that

the spore wall dehisces on the convex distal face, with the

formation of a germ pore between the areolae and the

emergence of a germ tube through this pore.

Duthie & Garside (1936, 1939), studied spore germina-

tion in some species occurring in southern Africa, namely

R. cavernosa Hoffm. emend. Raddi (= R. rautanenii

Steph.), R. crystallina L. emend. Raddi (= R. plana Tayl.)

and the endemic, R. cupulifera A.V. Duthie, as well as

in the permanently coherent tetrad spores of R. compacta

FIGURE 2. — Riccia albovestita. Germination of spore and development of protonema: A, dehiscence of spore wall between two areolae; B, spore

with germination filament; C, D, spore with germination tube; E, quadrant stage; F, plateau stage; G, formation ot plateau into a column.

Smook 4036. A— G, LM photographs. Scale bars on A-G = 20 /xm.

Bothalia 20,2 (1990)

215

FIGURE 3. — Vegetative reproduction of R. albovestita. A, old thallus

with three young thalii, indicated by arrows; B, young thallus,

enlarged from A, with dorsal cell pillar, J.M. Perold 39. A, B.

LM photographs. Scale bars on A = 1 mm; B = 20 ftm.

Garside and R. curtisii (Aust.) T.P. James. The first three

species mentioned here belong to the subgenus Ricciella

(A. Braun) Bischo and the latter two to the subgenus

Thallocarpus (Aust.) Jovet-Ast.

Jovet-Ast (1975) also investigated spore germination as

well as the protonemal phase in several members of these

two subgenera, in addition to a few species in the subgenus

Riccia , namely R. crozalsii Lev., R. albolimbata S. Arnell,

and the European species R. sommieri Lev. and R.

beyrichiana Hampe ex Lehm. & Lindenb.

African section Pilifer Volk, subgenus Riccia. It is evident

that its germination is no different from those in the rest

of the genus and that this phenomenon is constant

throughout. Spores from a specimen of R. albovestita

Volk, leg. Smook 4036 , collected at the edge of a small

dam between Ulster and Mooi River in the eastern Cape,

germinated six years after collection, when several thalii

with sporangia were kept moist on filter paper in a covered

dish. Various stages can be seen in Figure 2A-G: the

spore wall dehisces between the areolae on the distal face

(Figure 2A) and soon a germinative pore appears, from

which a germinative filament emerges (Figure 2B); the

filament enlarges into a germinative tube (Figure 2C, D),

and at its tip, it produces a quadrant (Figure 2E), which

has 4 cells in 2 tiers; more cells are formed at the top of

the quadrant (Figure 2F) producing a plateau, which

elongates into a column (Figure 2G). At this stage the

protonema measures 180 x 60 ptm. Its subsequent growth

was not monitored.

Vegetative reproduction of young thalii from an old thallus

Figure 3A, B illustrates the vegetative reproduction of

new young thalii (indicated by arrows) from cells of the

gametophyte of R. albovestita. In Figure 3B, one young

thallus already has a dorsal cell pillar, here projecting

laterally.

The above developmental stages also agree with those

observed in other species of Riccia.

REFERENCES

CAMPBELL, D. H. 1918. The structure and development of mosses and

ferns. 3rd edn, pp. 708. New York.

DUTHIE, A.V. & GARSIDE, S. 1936. Studies in the South African Ric-

ciaceae. I. Transactions of the Royal Society of South Africa 24:

93-133.

DUTHIE, A.V. & GARSIDE, S. 1939. Studies in the South African Ric-

ciaceae. II. Transactions of the Royal Society of South Africa 27:

17-28.

FELLNER, F. 1875. Uber die Keimung der Sporen von Riccia glauca.

Jahresbuch der Akademie des Naturwissenschaftlichen Vereins

Graz. 41-47.

HOFMEISTER, W. 1851. Vergleichende Untersuchungen der Keimung,

Entfaltung und Fruchtbildung hoherer Kryptogamen , pp. 179.

Leipzig.

JOVET-AST, S. 1975. Germination et phase protonemique chez Riccia.

Revue bryologique et lichenologique 41: 263 —276.

KNY, L. 1867. Uber Bau und Entwicklung der Riccien. Jahrbuch fiir

Wissenschaftliche Botanik von Pringsheim V: 364—386.

VOLK, O.H. 1981. Beitrage zur Kenntnis der Lebermoose (Hepaticae)

aus Siidwestafrika (Namibia). II. Mitteilungen der Botanischen

Staatssammlung, Miinchen 17: 245—252.

VOLK, O.H. 1983. Vorschlag fur eine Neugliederung der Gattung Riccia

L. Mitteilungen der Botanischen Staatssammlung, Miinchen 19:

453-465.

S.M. PEROLD*

The present communication is, however, the first report

of spore germination in a species of the endemic southern

National Botanical Institute, Private Bag X101, Pretoria 0001.

216

Bothalia 20,2 (1990)

ORCHIDACEAE

NOTES ON A RARE CAPE DISA

Although the southern African ground orchids have

received extensive taxonomic study over the past two

decades, there are still some rare species which are only

known from a few poorly preserved herbarium specimens.

Consequently, critical details of the gynostemium structure

of these species are not known and often it is not clear

whether these 'species’ are mutants of some common

species, or in fact real, albeit rare, evolutionary species.

One such species is Disa neglecta Sond. The nomen-

clatural and taxonomic confusion surrounding this species

was clarified by Linder (1981), but the two collections (one

the type, the second consisting of one specimen from a

mixed collection) were too poor to study the detailed

structure of the flower. Recently, Miss Margaret Burger

located a population, and brought in fresh material which

allowed me to make the following observations.

The plants are 100—180 mm tall; leaves spirally

arranged, more or less U-shaped with rounded backs in

cross section, longest near base of plant, 90 x 8 mm,

narrowing to the point of insertion, imbricate, reaching

halfway up inflorescence, apices curved inwards. Inflores-

cence dense, 90—150 mm long, 30 mm in diameter at the

base and tapering upwards; lowermost bracts leaf-like,

much overtopping flowers, the uppermost bracts much

shorter and only marginally overtopping flowers; ovary

and pedicel 8—10 mm long and twisted, resupinating the

flower. Flowers facing downwards, galea brownish purple,

or densely brownish purple mottled, conspicuous; sepals

and lip lime green with faint mottling on margins. Sepals

unequal; dorsal sepal deeply galeate, 6x4 mm and 5

mm deep, apex slightly emarginate, subacute, base with

two shallow sacs less than 0,5 mm deep; laterals patent,

fleshy, concave above and apically very shortly apiculate,

3-nerved, 5x2 mm. Petals lunate, horizontal, 2 x 1 mm,

basal anticous lobe thin, main body of petal reduced to

basal portion, fleshy, with a posterior fleshy, downward

pointing lobe or tooth seated in sacs of galea. Lip linear,

pendent, fleshy, apex blunt, 4 x 1 mm. Anther erect with

a fleshy, tapering connective, 1 mm tall, connective 0,5

mm deep; anther cells obovate, 0,5 mm in diameter. Stigma

pulvinate, sessile, with three equal lobes. Rostellum

three-lobed, erect; central lobe conical, fleshy, 0,5 mm

tall, penetrating between anther cells; lateral lobes

spreading horn-like, terete, as tall as central lobe but more

slender, not canaliculate, bearing hemispherical viscidia

at apices, Figure 4.

Growing in full sun on a well drained, rocky sandstone

ridge, between 1 500 and 1 800 m altitude above sea lev-

el, in association with Disa micropetala Schltr. , D. vagina-

ta Harv. ex Lindl. and Satyrium pygmaeum Sond.

Flowering at the beginning of December, in the first year

after fire.

The general structure of the plant and the detailed flower

structure of Disa neglecta agree closely with those charac-

teristic for Disa section Disella Lindl., which is where

the species was placed by Linder (1981). The two sacs at

the base of the galea could represent the remnants of

the galea spur plus the dorsal groove characteristic of part

of section Disella. It is easy to imagine that reduction

under these circumstances would lead to such a structure,

which superficially approaches the double spur of

Satyrium.

However, in detailed structure the species shows sever-

al features which are unique in the genus. The high degree

of petal reduction is not found in any other species of the

subtribe, and the presence of a tooth that penetrates into

the spur (or in this case, the sacs) is only known in Disa

longicomis L. f. These petal teeth completely fill the spur,

and it is difficult to imagine their function. The erect

anther is equally strange and D. neglecta is the only species

in section Disella that has it. Erect anthers, however, do

occur occasionally in other members of the genus

(commonly in Disa section Micranthae , rarely in section

Disa). In D. neglecta the anther shows no evidence at all

of resupination. The rostellum structure is also unique

within the section. Although the rostellum is typically

three-lobed in the section, only in D. neglecta and in one

population of D. obtusa subsp. obtusa (also in section

FIGURE 4. — The structure of the flower of Disa neglecta. all from Burger

8. A, whole flower in front view, X 2,5; B, flower with sepals

and one petal removed, in side-on front view, X 6; C, flower

in side view, with galea cut away, and lateral sepals and lip re-

moved, X 6; D, gynostemium and petals in front view,

x 6. Cut surfaces shown hatched, viscidia black, s = stigma,

a = anther cell, c = connective, r = rostellum , p = petal.

Bothalia 20,2 (1990)

217

FIGURE 5.— Distribution of Disa negtecta.

Disella) from the summit of Table Mountain is the central

lobe taller than the lateral lobes. However, in D. neglecta

the lateral lobes are terete, rather than canaliculate as is

typical for the rest of the section.

The known distribution of the species is indicated in

Figure 5. The distribution range covers the mountains from

Worcester to the Cedarberg, but the species has only been

collected at the extremes of the range. Either this is a

remarkably disjunct distribution, or the species occurs in

the intervening Koue Bokkeveld Mountains, but has not

been collected there. As the Koue Bokkeveld Mountains

are generally poorly collected, the latter hypothesis is more

likely.

It is surprising how many rare species are found in

section Disella. Of the ten species, seven ( D . neglecta

Sond., D. longifolia Lindl., D. brevipetala Linder, D.

telipogonis Reichb. f. , D. micropetala Schltr. , D. subtenui-

comis Linder, D. ocellata H. Bol. and D. lineata H. Bol.)

are seen but rarely. Some of this rarity is clearly caused

by these species only appearing after fire. In addition, they

are usually of local occurrence, being restricted to moist,

black soils near the summits of mountains where they

receive moisture from the south-easter clouds during the

otherwise dry summer months. However, some of the

species (D. neglecta, D. subtenuicomis and D. longifolia )

appear to be genuinely rare, and are known from a single,

or very few populations. Clearly there is still scope for

an intensive botanical survey of the higher, more inacces-

sible mountain ridges and peaks.

SPECIMENS EXAMINED

CAPE.— 3219 (Wuppertal): between Breekkrans Twins and Apex Peak,

1 360-1 160 m (— CA), 20 December 1989, Burger 9 (BOL!). 3319 (Wor-

cester): Waaihoekskloof, below and south of Sybasberg, 1 500—1 800

m (-CB), 3 December 1989, Burger 8 (BOL!); Louwshoek Peak (—CD),

17 December 1944, Esterhuysen 11193B (BOL!); Tulbagh Mountains,

November, Eeklon & Zeyher s.n. (S!).

REFERENCE

LINDER, H P 1981. Taxonomic studies on the Disinae. III. A revision

of Disa Berg, excluding sect. Micranthae Lindl. Contributions

from the Bolus Herbarium 9: 1—370.

H.P LINDER*

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

7700.

AIZOACEAE

A NEW SPECIES OF GALENIA FROM THE GREAT KAROO

The genus Galenia L. is placed in the subfamily

Aizooideae (Aizoaceae s. str. ) and is generally character-

ized by androecial and gynoecial features (Bittrich &

Hartmann 1988; Bittrich 1990). The genus is now divided

into two subgenera, namely subgenus Galenia and

subgenus Kolleria Fenzl emend. Adamson. The descrip-

tions of the two subgenera, especially that of the subgenus

Galenia, as given by Adamson (1956), are partly incorrect,

and the monophyly of these taxa also still needs to be

shown. Subgenus Kolleria Fenzl emend. Adamson now

comprises about 20 species and, according to Adamson

(1956), is characterized by obovate or spathulate, often grey

or white leaves, secund cymose inflorescences, 4 or 5

perianth segments, 2-5 styles, and dehiscent fruits. These

characters, however, can also be observed in most species

of the closely related genus Aizoon L. They must there-

fore presumably be regarded as plesiomorphic, so that at

present no synapomorphy can be named which might

suggest the monophyly of subgenus Kolleria.

Galenia glandulifera Bittrich, sp. nov., planta herba

annua papulosa, species haec a G. papulosa (Eckl. &

Zeyh.) Sond. indumento pilis glandulosis differt (in G.

papulosa indumentum tantum eglandulosum). Chromo-

somatum numerus 2n — 16.

218

Bothalia 20,2 (1990)

FIGURE 6 .—Galenia glandulifera Bittrich. Flowering lateral branch

Plant raised from seeds of Gerbauler & Struck 24236. Scale bar

= 5 mm.

TYPE. — Cape 3323 (Willowmore): 1 km N Swartleegte

on foot of the Spioenkop (-BA), Gerbaulet & Struck

24236 (HBG, holo.; PRE).

Small annual glistening herb up to 100 mm tall and

100—600 mm in diameter, branching from the base; main

axis erect, compressed, without flowers, lateral branches

decumbent or ascending, elongate, ending in cymose

inflorescences; leaves and stems densely covered with large

water storing idioblasts (bladder cells) and more or less

sparsely covered with oblong ovoid, bottle-shaped or pyri-

form, apically secretory glands with a multiseriate stalk.

Leaves weakly succulent, opposite on main axis, alternate

on lateral branches, exstipulate, petiolate; lamina obovate,

obtuse, up to 20 x 8 mm on main axis and up to 8 x

5 mm on lateral branches, petioles 2-7 mm long. Inflores-

cence a dichasium, up to 300 mm long, terminating the

lateral branches, the two side branches of each node very

unequal in length, inflorescence thus appearing

monopodial; bracts recaulescent with their axillary shoot

up to the terminal flowers. Flowers protandrous, sessile,

perigynous, 5-merous, 5 mm in diameter; tepals 5,

triangular, green on outside, magenta or purple on inside.

Stamens 10, in pairs, alternating with the tepals, inserted

at the mouth of the perianth-stamen tube, light purple.

Styles 5, whitish papillose on outside, purplish hairy-

papillose on inside, free to the base. Ovary slightly con-

vex, surface with large papillae, 5-locular with one, rare-

ly two pendulous ovules per locule; placenta axile apical.

Disc a flat green ring at the base of the filaments. Fruit

a 5-angled loculicidal, hygrochastic capsule of up to 2 mm

diameter, with expanding keels and sheets on the inner

side of the valves, depressed centrally; valves slightly

opening outwards after wetting. Seed reniform, 0,75 x 0,5

x 0,4 mm, dark brown or black, with small tubercles.

Chromosome number 2n = 16. Figure 6.

CAPE. — 3121 (Fraserburg): 7 miles S of Fraserburg Station. Prince

Albert Drive (—DC), Zinn s.n. (SAM). 3123 (Victoria West): prope Mur-

raysburg (-DD), Tyson s.n. (SAM). 3221 (Merweville): Zwarts(-DD),

Pillans 16368 (SAM). 3223 (Rietbron): Nelspoort, Beaufort West Dist.

(— AA), Kingon s.n. (NBG). 3224 (Graaff-Reinet): SW of Aberdeen

(—AC), Maguire 749 (NBG). 3320 (Montagu): Whitehill (—BA),

Compton 9296 (NBG). 3321 (Ladismith): 15 miles E of Laingsburg on

road to Beaufort West (— AA), Lewis 4172 (SAM). 3322 (Oudtshoorn):

Karoo, Prince Albert (-AA), Stokoe s.n. (SAM); Tierberg (-AB), Dean

& Dean 677 (HBG). 3323 (Willowmore): 1 km N of Swartleegte on

foot of Spioenkop (—BA), Gerbaulet & Struck 24236 (HBG, PRE).

Galenia glandulifera occurs only in the Great Karoo.

Figure 7.

On account of its 5-merous flowers and broad leaves,

this species at present must be placed in subgenus Kolleria.

Specimens of G. glandulifera are often wrongly identified

as G. papulosa. G. glandulifera has glandular trichomes

(Figure 8), an extremely rare character in the Aizoaceae

s. str. These trichomes are most probably a modification

of the stalked bladder-hairs, which are typical for the

Aizooideae/Tetragonioideae and presumably a synapo-

morphy of the clade comprising these subfamilies (Bittrich

1990). The glandular trichomes of G. glandulifera secrete

very small droplets of a clear liquid at their apex, which

on drying becomes yellow and later brownish and smells

of coumarin.

In the plasma of the secretory cell at the tip of the

trichomes, a large nucleus and some chloroplasts are

visible (Figure 8). Chloroplasts (incidentally reported to

occur in the hairs of various vascular plants) can generally

also be found in the apical cells of the bladder hairs of

species belonging to the Aizooideae/Tetragonioideae. They

are also known to occur within the plasma of the epidermal

bladder cell idioblasts of other Aizoaceae (Luettge et al.

1978). It is noteworthy that in subgenus Galenia some

species, such as G. africana L., also have secretory

stalked trichomes on leaves and stems. In contrast to

G. glandulifera , these hairs are T-shaped (the most com-

mon and probably plesiomorphic hair form in the genera

FIGURE 7. — Distribution of G. glandulifera.

Bothalia 20,2 (1990)

219

«

FIGURE 8. — Stem epidermis of G. glandutifera in cross section with

bladder cell idioblasts and a glandular trichome. Scale bar =

0,1 mm.

Aizoon L., Galenia L., and Plinthus Fenzl emend.

Verdoorn). The secretion, which contains coumarin

derivates (G. africana is poisonous for cattle!), completely

covers the green parts of the plants with a yellow and sticky

layer which on warm days often smells strongly. These

differences may indicate on the one hand that the glandular

trichomes have evolved independently in both subgenera,

but on the other hand the extreme rarity of such trichomes

in Aizoaceae 5. str. suggests a close relationship between

species of Galenia with this character. If the latter is true,

subgenus Kolleria would be an unnatural taxon. A detailed

analysis of the secretion might provide further evidence

to decide between these two hypotheses.

ACKNOWLEDGEMENTS

I am very grateful to M. Gerbaulet and M. Struck for

providing seeds and pickled material of G. glandulifera .

I wish to thank M. Struck for critically reading the

manuscript. Dr J. Kadereit for improving the English text

and the directors and staff of the cited herbaria for the

loan of specimens. The investigations in Aizoaceae were

supported by a grant of the Deutsche Forschungsgemein-

schaft, for which I am very grateful.

REFERENCES

ADAMSON, R.S. 1956. The South African species of Aizoaceae. III.

Galenia L. South African Journal of Botany 22: 87—127.

BITTRICH, V. 1990. Systematic studies in Aizoaceae. Mitteilungen aus

dem lnstitut fur Allgemeine Botanik Hamburg 23b: 491—507.

BITTRICH, V. & HARTMANN, H.E.K. 1988. The Aizoaceae — a new

approach. Botanical Journal of the Linnean Society 97: 239—254.

LUETTGE, U., FISCHER, E. & STEUDLE, E. 1978. Membrane poten-

tials and salt distribution in epidermal bladders and photosyn-

thetic tissue of Mesembryanthemum crystallinum L. Plant, Cell

and Environment 1: 121—129.

V. BITTRICH*

* lnstitut fur Allgemeine Botanik und Botanischer Garten der Universitat

Hamburg, Ohnhorststr. 18, D-2000 Hamburg 52, Fed. Rep. Germany.

ANACARDIACEAE

A NEW SPECIES OF OZOROA FROM THE TRANSVAAL

Ozoroa barbertonensis Retief \ sp. nov. , O. albicanti

atfinis, sed foliis linearibus vel anguste oblongis, non

oblongis vel late ovato-oblongis, in rupibus serpentinis in

parte orientali terrae vulgo Transvaal dictae, non dolomiti-

cis, in parte boreali eiusdem terrae differt.

TYPE.— Transvaal, 2531 (Komatipoort): Barberton,

mountain slopes above the Agnes Gold Mine (— CC),

Balsinhas 3129 (PRE, holo.; K; MO). Figure 9.

A dioecious shrublet, 0,4— 1,5 m high, with several erect

stems from a woody rootstock. Stems brown, hairy, with

a milky latex. Leaves simple, spirally or subverticillate-

ly arranged, much longer than internodes, crowded;

petiolate; blade linear to narrowly oblong, (23 — )55 — 80

(-90) x (2 — )4 — 7( — 8,5) mm, coriaceous, discolorous,

upper surface shiny green, lower surface greyish green,

both surfaces with a hairy indumentum, hairs on upper

surface not as dense as on lower surface, those on lower

surface orientated towards the margin, apex acute, with

a mucro ± 1 mm long, base cuneate, margin revolute,

lateral veins numerous, parallel, unbranched or branched

near margin, less hairy; petiole 1, 5-3,0 mm long. Flowers

white, 5-merous, in axillary and terminal panicles, pedicels

articulated near apex. Male flowers : sepals 5-segmented,

1-1,5 mm long; petals 5, oblong, 2—2,5 mm long, apices

curved inwards; disc annular; stamens 5, filaments 0,5 mm

long, anthers 0,75 mm long. Female flowers : sepals 1 mm

long, petals 2 mm long, otherwise as in male flowers; disc

annular, staminodes ± 5 mm long; ovary symmetrical,

1-locular with a single basal ovule, styles 3, free, stigmas

somewhat capitate. Fruit a fleshy drupe, subglobose,

4,5-5 X 6-6,5 mm, irregularly rugulose. Seed pendulous

from the basal funicle.

220

Bothalia 20,2 (1990)

TRANSVAAL. — 2530 (Lydenburg): Agnes Mine (— DD), Balkwill &

Cadmatr 3604 (PRE, E, J, NU), Strey 4052 (PRE); Thorncroft Nature

Reserve ( — DD), Muller 2321 (PRE). 2531 (Komatipoort): Barberton,

mountain slopes above Agnes Gold Mine (— CC), Balsinhas 3129 (PRE.K,

MO), Barberton, Brommers (— CC), Onderstal 1228, 1229 (PRE).

FIGURE 9.— Holotype of Ozoroa barbertonensis Rebel, Balsinhas 3129.

FIGURE 10. — Ozoroa barbertonensis, Balsinhas 3129. SEM micro-

graph of upper surface of leaf blade, x 66.

FIGURE 11. — Ozoroa barbertonensis, Balsinhas 3129. SEM micro-

graph of lower surface of leaf blade, X 66.

FIGURE 12. — Distribution of Ozoroa barbertonensis, 0 : O. albicans ,

A ; O. schinzii, 0.

Ozoroa barbertonensis is apparently restricted to the

Barberton area of the Transvaal Lowveld, hence the

specific epithet. It occurs in Lowveld Sour Bushveld on

rocky slopes in grassland. Flowering time has been

recorded from January to March. An unusual feature of

the species is its association with serpentine soils (Balkwill

& Balkwill 1988), a characteristic shared with taxa such

as Berkheya coddii Roessler, Aloe thorncroftii Pole Evans

and Brachystelma dyeri K. & M-J. Balkwill (1988).

Ozoroa barbertonensis differs from all other southern

African species of Ozoroa by its linear to narrowly oblong

leaves with the upper surface shiny green and the lower

surface greyish green, caused by the dense indumentum

of hairs. A SEM study of the indumentum of the blade

revealed that the hairs on the upper surface (Figure 10)

are not as dense as those on the lower surface (Figure 11).

The leaves are also much longer than the internodes and

crowded. The habit of the species, a shrublet with several

Bothalia 20,2 (1990)

221

stems from a woody rootstock, is also a useful aid in its

identification as only two other described species of

Ozoroa in southern Africa share the same habit (other

members are trees or shrubs).

The nearest relatives of Ozoroa barbertonensis are O.

albicans R. & A. Fernandes and O. schinzii (Engl.) R.

& A. Fernandes. The three species are all many-stemmed

shrubs, sometimes up to 2 m high, but usually 0,4—1 m

in height. However, the shape and size of their leaf blades

differ markedly from one another. O. albicans has large

greyish green leaves with blades oblong to broadly ovate-

oblong, 75—155 x 30 -70 mm. The leaf blades of

O. schinzii are oblong to elliptic, 25—50 x 10—20 mm

with prominent lateral veins on the lower surface. Charac-

teristic of O. barbertonensis are its linear to narrowly ob-

long leaf blades, 55-80 x 4-7 mm. The geographical

distribution of these species is also different and does not

overlap, Figure 12.

REFERENCE

BALKWILL, K. & BALKWILL, M-J. 1988. Studies on serpentine flora:

a new species of Brachystelma (Asclepiadaceae). South African

Journal of Botany 54: 60—62.

E. RETIEF

ADIANTACEAE/PTERIDACEAE

DORYOPTERIS P1LOSA VAR. GEMMIFERA , A NEW FERN VARIETY ESTABLISHED

Doryopteris pilosa (Poir.) Kuhn var. gemmifera J.E.

Burrows & Strauss, var. nov. , a D. pilosa var. pilosa

frondibus gemmas ad juncturam stipitis rachidisque

ferentibus differt.

TYPE. — Transvaal, Thabazimbi District, Geelhout-

bosch, Strauss 17 (PRE, holo.). Figures 13; 14 A, B.

The publication of the new record of Doryopteris pilosa

(Poir.) Kuhn for the southern African subregion (Jacobsen

& Jacobsen 1988), makes no mention of this fern bearing

gemmae, although plants in the Geelhoutbosch population

frequently bear gemmae at or near the junction of the stipe

and the midrib. (Figure 13). This phenomenon also occurs

in D. pedata (L.) Fee from the New World, with the

proliferous form having been accorded varietal status, D.

pedata var. palmata (Willd.) Hicken.

In order to establish the relationship of D. pilosa from

southern Africa to the typical form from the Mascarene

Islands and Madagascar (Tardieu-Blot 1958), SEM micro-

graphs were made of the spores from two collections from

the Indian Ocean islands as well as from material from

the north-western Transvaal (Figure 14). There appear to

be no discernable differences in the spores from the two

regions. However, because of the disjunct geographical dis-

tribution of the two forms, together with the distinctive

gemmiferous habit, it was felt that the form from Geel-

houtbosch should be given varietal rank. Specimens from

the Penge area (Jacobsen & Jacobsen 1988) and other

regions of the Transvaal have not been reported to possess

gemmae but further field work will possibly result in their

inclusion within the new variety.

ACKNOWLEDGEMENTS

Our thanks are extended to Mrs S.M. Perold of the

National Botanical Institute, Pretoria for assistance with

the SEM micrographs, and to Dr A. Le Thomas of the

Museum National d’Histoire Naturelle, Paris for supplying

spore material. We also wish to thank the staff of the Bolus

Herbarium, Cape Town, the National Herbarium, Pretoria

and the Compton Herbarium, Kirstenbosch, for their

willing assistance.

FIGURE 13.— Doryopteris pilosa (Poir.) Kuhn var. gemmifera J.E.

Burrows & Strauss, Strauss 17: A, gemma at the base of the fertile

frond, X 0,9; B, gemmae at the base of sterile fronds, x 0,8.

222

Bothalia 20,2 (1990)

FIGURE 14. — Doryopteris pilosa (Poir.)

Kuhn. A, B, var. gemmifera J.E.

Burrows & Strauss, Strauss 17:

A, spore, distal face, x 786;

B, spore, side view, x 936.

C— F, var. pilosa. C, D, St Paul,

He de la Reunion, 1875, G. de

1'lsle 131 (P): C, spore, distal

face, X 948; D, spore, proximal

face, x 876. E, F, environs

dAntanimoro (Androy), Mada-

gascar, 6—9.2.1955, H. Humbert

et R. Capuron 28821 (P): E,

spore, proximal face, X 760; F,

spore, distal face, X 966.

REFERENCES

JACOBSEN, W.B.G. & JACOBSEN, N.H.G. 1988. Doryopteris pilosa,

a new record for South Africa. Bothalia 18: 90—92.

TARDIEU-BLOT, M.-L. 1958. Polypodiacees I In H. Humbert, Flore

de Madagascar et des Comores. Museum National d’Histoire

Naturelle, Paris.

J.E. BURROWS* and S.E. STRAUSS**

* Buffelskloof Nature Reserve, P.O.Box 236. Lydenburg 1120.

** P.O. Box 71670, Die Wilgers, Pretoria 0041

Bothalia 20,2: 223-227 (1990)

Ordination as a tool for substantiating and interpreting floristic

classifications: a case study

G.B. DEALL* and G.K. THERON**

Keywords: classification, Detrended Correspondence Analysis, ecological interpretation, ordination, PHYTOTAB

ABSTRACT

Detrended Correspondence Analysis ordination is used as a tool for substantiating the authenticity of hierarchical syntaxa

derived from a phytosociological classification of the vegetation of the Eastern Transvaal Escarpment in the Sabie area.

Ecological interpretation of syntaxa is facilitated by superimposing environmental variables on axes of ordination.

UITTREKSEL

Ordinasie deur middel van Ontneigde Ooreenstemmingsanalise is gebruik ter ondersteuning van die geldigheid van die

hierargiese sintaksons wat verkry is met fitososiologiese klassifikasie van die plantegroei van die Oos-Transvaalse Eskarp

in die omgewing van Sabie. Die ekologiese interpretasie van sintaksons is vergemaklik deur die aanbring van omgewings-

veranderlikes langs die asse van ordinasie.

INTRODUCTION

In phytosociological classification the possibility always

exists that groupings are controlled by the ecologist rather

than by structure internal to the data (Gauch & Whittaker

1981). Most community data sets have two to four gradients

and in this dimensional range hierarchical classification

tends to be weak. Ordination, on the other hand, is

especially strong in this range and may complement

classification for such data (Gauch 1982). Ordination is

therefore a useful adjunct to classification, facilitating an

a posteriori basis for determining whether Braun-Blanquet

syntaxa are real, naturally occurring entities, or whether

they are mere artefacts of data arrangement. Through its

ability to portray similarity relations geometrically,

ordination is also useful for emphasising inter-community

relationships (Mueller-Dombois & Ellenberg 1974). Fur-

thermore, by imposing environmental variables upon the

axes of ordination, the ecological interpretability of

vegetation can be enhanced.

A Braun-Blanquet phytosociological classification of

vegetation in the Sabie area by Deall (1985) using

PHYTOTAB (Westfall et al. 1982) revealed 62 basic

syntaxa (comprising communities and variants) distributed

through a hierarchy of 14 vegetation types and four

ecological-formation classes (Table 1).

The aim of this paper is to determine, by means of

ordination, whether these syntaxa (especially at vegetation-

type level) really represent the vegetational entities in

nature. Inter-community relationships and environmental

interpretations are also considered.

METHODS

The 62 syntaxa of Deall (1985) were ordinated on

floristic data using Detrended Correspondence Analysis

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

address: Africa School of Missions, P.O. Box 439, White River 1240.

** Department of Botany, University of Pretoria 0002.

MS. received: 1989.04.25.

(DCA) (Hill & Gauch 1980). The latter was effected using

the DECORANA program (Hill 1979), which is dependent

on inputs from various programs of the PHYTOTAB com-

puter package (Westfall et al. 1982). The ordination

involved the arrangement of plant communities by group

values (to the nearest 0,5 sd). Four different ordinations

were undertaken on the basis of four data subsets

corresponding to the four ecological-formation classes.

Relationships between communities and environmental

variables usually defy analysis by conventional statistical

TABLE 1.— Hierarchical arrangement of syntaxa in the Sabie area of

the Eastern Transvaal Escarpment (from Deall 1985)

224

methods and are mostly perceived by informal, visual

recognition of common patterns of distribution (Gauch

1982). One way of portraying such patterns is to super-

impose environmental variables on axes of community

ordination (Westfall 1983).

Environmental variables recorded during sampling were

accordingly superimposed on each ordination diagram.

Those variables exhibiting a definite gradient along

ordination axes were retained as being important in

community differentiation, whereas others were discarded.

Ordination axes were selected on the basis of their having

sufficient variance to represent inherent data structure

(Hill 1979) and their propensity to exhibit maximal

vegetational variation (based on ‘spread’ of syntaxa).

The ordination of plant communities (and variants) is

discussed in the context of their respective ecological-

formation classes, or data subsets. It must be noted that

the terms used for indicating environmental gradients in

the ordination diagrams (Figures 1—4) are not absolute,

but relative. That is, the scale of an environmental feature

on one axis is not necessarily the same as it is on the other

axis. For example, ‘deep’ on the x axis may be the same

depth as ‘shallow’ on the y axis.

RESULTS AND DISCUSSION

Forest and Mesic Thicket of the Mistbelt and Low

Country (4.2.1.)

Axes 1 and 3 in Figure 1 form the ordination axes of

the scatter diagram which represents relationships between

vegetation and environment in the 19 communities com-

prising the Forest and Mesic Thicket of the Mistbelt and

Low Country subset. Together these axes account for

69,4% of the variance. Gradient lengths of the first three

axes are slightly less than 4,0 sd, indicating a moderately

high beta diversity for Forest and Mesic Thicket of the

Mistbelt and Low Country [samples with a separation of

Sheltered

EXPOSURE

Exposed

t-OW ALTITUDE/CLIMATE High

Dry ^ ► Moist

Warm Cool

LITHOLOGY

Granite ◄ ► Quartzite

Deep ^ Shallow

Less rocky Rocky

OO 05 LO L5 2^0 2^5 3^

Axisl (sd units)

FIGURE I. —Detrended Correspondence Analysis (DCA) ordination of

the 19 plant communities comprising Forest and Mesic Thicket

of the Mistbelt and Low Country, Sabie area. Axes 1 and 3

represent the main environmental gradients as shown.

Bothalia 20,2 (1990)

greater than 4,0 sd will generally have no species in

common (Hill 1979)].

Community separation on the basis of vegetation type

is indicated in the ordination diagram by means of the

broken lines (Figure 1). Apart from the placement of

Community 17 into Vegetation Type 4. 2. 1.4 (Table 1), the

identity of the five vegetation types associated with Forest

and Mesic Thicket of the Mistbelt and Low Country is

faithfully maintained by the ordination. Thus the classifica-

tory basis of the vegetation-type category is, for the most

part, confirmed.

Some of the environmental gradients possibly

influencing community differentiation in Forest and Mesic

Thicket of the Mistbelt and Low Country are indicated

in relation to axes 1 and 3 of the ordination diagram

(Figure 1). Axis 1 corresponds predominantly to a

temperature/moisture/lithology gradient, whilst axis 3

appears to reflect a soil-depth gradient.

Forest vegetation is uniformly represented along both

axes (Figure 1). Whereas forest Communities 1 to 6 are

associated with the moist, cool conditions of higher

altitudes, forest Communities 17 to 19 are associated with

the dry, warm conditions of lower altitudes (axis 1, Figure

1). Thus forest ostensibly exhibits a wide moisture

tolerance. In reality, however, Communities 17 to 19

are confined to sheltered riparian sites where, despite

surrounding aridity, moisture is non-limiting.

The isolation on axis 1 of forest Community 3 is

probably due to both the shallow, rocky quartzite-derived

soil, and the exposed sites to which the Community is

confined (Figure 1). By contrast, forest Communities 1

and 2 occur on deeper dolomite-derived soils in more

sheltered sites, whereas Communities 4 to 6 occur on

deeper granite-derived soils, also in more sheltered sites.

Mesic Thicket (Vegetation Type 4. 2. 1.4) is confined to

the portion near the origin of each axis (Figure 1).

Constituent syntaxa (Communities 7 to 16) are conse-

quently grouped in the lower left sector of the ordination

diagram, corresponding to conditions that are not

sufficiently moist to sustain forest. The anomalous

occurrence of forest Community 17 in the ‘drier’ sector

of the diagram may reflect a moisture deficit relative to

Communities 18 and 19. Whereas the soils of the two latter

communities have a high clay fraction in the lower

horizons, those of Community 17 consist mainly of sand.

The water-holding capacity of soils in Community 17,

therefore, is likely to be less than in Communities 18 and

19.

Woodland and Xeric Thicket of the Low Country

(4.2.2)

Axes 1 and 2 in Figure 2 form the ordination axes of

the scatter diagram which represents relationships between

vegetation and environment in the 19 communities

comprising die Woodland and Xeric Thicket of the Low

Country subset.

These axes account for 74,3% of the variance. Gradient

lengths of the first three axes range from 2,35 to 3,08 sd,

Bothalia 20,2 (1990)

225

indicating a moderately low beta diversity for Woodland

and Xeric Thicket of the Low Country.

The identity of the three vegetation types associated with

Woodland and Xeric Thicket of the Low Country is faith-

fully maintained by the ordination (Figure 2). The clas-

sificatory basis of the vegetation-type category is therefore

confirmed.

Some of the environmental gradients possibly

influencing community differentiation in Woodland and

Xeric Thicket of the Low Country are indicated in relation

to axes 1 and 2 of the ordination diagram (Figure 2).

Axis 1 corresponds to a temperature/moisture/exposure

gradient, whereas axis 2 reflects a similar gradient com-

pounded with soil depth and slope. The net effect of these

gradients is a gradient corresponding to the positive

diagonal and operating from cool moist conditions, with

shallow rocky soils on exposed steep slopes (upper right);

to warm dry conditions, with deep, less rocky soils on

sheltered gentle slopes (lower left).

The low open/sparse woodland and the short sparse

shrubland communities of Vegetation Type 4. 2. 2. 3 are

clearly correlated with the shallow rocky soils on exposed

steep slopes (Figure 2). Vegetal cover in this part of the

gradient is accordingly low. As these habitats become

warmer, drier, and more sheltered, the vegetation becomes

more woody, resulting in the woodland and thicket com-

munities of Vegetation Type 4. 2. 2.1. On the more gentle

slopes with deepish, less rocky soils (lower left and right,

Figure 2), land is more arable, more inviting for human

settlement, and more susceptible to fire. Attendant vege-

tation is therefore likely to be disturbed. This may explain

the sparsely woody nature of Vegetation Type 4. 2. 2. 2.

Woodland of the Humid Mistbelt (4.2.3)

Axes 1 and 2 in Figure 3 form the ordination axes of

the scatter diagram which represents relationships between

vegetation and environment in the six communities

comprising Woodland of the Humid Mistbelt subset.

These axes account for 89,2% of the variance. Gradient

lengths of the first two axes are slightly higher than those

for Woodland and Xeric Thicket of the Low Country, but

less than those for Forest and Mesic Thicket of the Mistbelt

and Low Country. Beta diversity in Woodland of the

Humid Mistbelt is therefore relatively intermediate

between the mesic and the more xeric ecological-formation

class.

Clearly, the identity of the two vegetation types

associated with Woodland of the Humid Mistbelt is

faithfully maintained by the ordination (Figure 3). Thus

the classificatory basis of the vegetation-type category is

confirmed.

Some of the environmental gradients possibly

influencing community differentiation in Woodland of the

Humid Mistbelt are indicated in relation to axes 1 and 2

of the ordination diagram (Figure 3). Axis 1 corresponds

to a geomorphology/geology/exposure gradient, and axis

2 reflects a temperature/moisture gradient.

The low open woodland and low thicket associated with

Communities 39 and 40 apparently develops in response

to a high soil-moisture status occasioned by the deep clay

soils of sheltered midslope planes (Figure 3). Their

association with dolomite, moreover, is reminiscent of the

tall/short forest represented by Communities 1 and 2, and

it is therefore possible that Communities 39 and 40 are

serai to these forest communities.

On the shallow and sandy quartzite-derived soils of

exposed and rocky upper-pediment slopes, soil-moisture

is more limiting. Vegetation response is accordingly

towards a closed/open woodland with low vegetal cover

(Communities 41 to 44, Figure 3). Supplementary

moisture in the form of ‘fog-drip’ (Deall et al. 1989),

Sheltered

EXPOSURE

Exposed

Low ALTITUDE /CLIMATE High

Warm — — ► Cool

Dry Moist

High

Moist Shallow

Cool Rocky Steep

Low

Dry Deep Gentle

Warm Less rocky

FIGURE 2.— Detrended Correspon-

dence Analysis (DCA) ordina-

tion of the 19 plant

communities comprising

Woodland and Xeric Thicket of

the Low Country, Sabie area.

Axes 1 and 2 represent the

main environmental gradients

as shown.

226

Bothalia 20,2 (1990)

Upper

ped. slope

Quartzite

Shallow

Rocky

sand

Gentle

Exposed

GEOMORPHOLOGY

- Dolomite

Deep

- Less rocky

Clay

Steep

Sheltered

High

Moisl

Cool

Low

Dry

Warm

Axis 1 (Sd units)

FIGURE 3. — Detrended Correspondence Analysis (DCA) ordination of

the six plant communities comprising Woodland of the Humid

Mistbelt, Sabie area. Axes 1 and 2 represent the main environ-

mental gradients as shown.

especially at the higher altitudes of Communities 43 and

44, may provide some stimulus for the succession to

develop towards the elfin-like forests represented by

Community 3.

Grassland of the Humid Mistbelt (4.2.4)

Axes 1 and 2 in Figure 4 form the ordination axes of

the scatter diagram which represents relationships between

vegetation and environment in the nine communities

comprising the Grassland of the Humid Mistbelt subset.

These axes account for 89,7 % of the variance. Gradient

lengths of the first two axes range from 2,76 to 3,78 sd,

indicating a slightly higher beta diversity than other

ecological-formation classes.

Vegetation Type 4.2.4. 1 (Communities 45 and 46) is the

only one in Grassland of the Humid Mistbelt, the identity

of which is not maintained by the ordination. Rather, it

tends to merge with two others, namely Vegetation Types

4. 2. 4. 2 and 4. 2. 4. 3 (Figure 4). This suggests that Vegeta-

Upper ped slope

Dolomite *

Clay ««

High <

GEOMORPHOLOGY

LITHOLOGY

SOIL

ALTITUDE

Midslope

Quartzite

Sand

Low

tion Type 4. 2. 4.1 may be a classificatory artefact, with

lithology and physiography playing a much less signifi-

cant role in community differentiation than is indicated

by the phytosociological classification.

Some of the environmental gradients influencing com-

munity differentiation in Grassland of the Humid Mistbelt

are indicated in relation to axes 1 and 2 of the ordination

diagram (Figure 4). Axis 1 apparently corresponds to a

soil-texture/lithology/geomorpholgy gradient, whereas

axis 2 appears to reflect a soil-depth/lithology/geomor-

phology gradient. The net effect of these gradients is a

geomorphology gradient corresponding to the positive

diagonal, and operating from upper-pediment slopes

(lower left) to midslope planes (upper right); and also a

lithology gradient corresponding to the negative diagonal,

and operating from dolomite (upper left) to quartzite (lower

right).

On the positive diagonal (Figure 4), Communities 49

and 50 are possibly separated from the other communi-

ties on the basis of an improved soil-moisture status,

reflecting their lower position in the landscape. On the

negative diagonal (Figure 4), Communities 46 to 48 and

50 appear to be separated from the other communities by

virtue of their sandy quartzite-derived shallow soils, which

soils are in contrast with the deeper, dolomite-derived clay

soils of their counterparts (Communities 45, 49 and 51 to

53).

Community 50 appears on the diagram as an outlier

(Figure 4). This may reflect its disturbed character, being

situated in a municipal area where human influences are

manifold.

CONCLUSIONS

Only one out of 14 vegetation types derived from the

phytosociological classification of vegetation in the Sabie

area could not be substantiated by ordination. Syntaxa at

the vegetation-type level are therefore considered to be

authentic. In order to substantiate the validity of syntaxa

at the community and variant level, it would be necessary

to ordinate their component sample plots.

Deep Midslope

Less rocky plane Dolomite

Shallow Upper Quartzite

Rocky ped.

slope

Axis 1 (Sd units)

FIGURE 4. — Detrended Correspon-

dence Analysis (DCA) ordina-

tion of the nine plant com-

munities comprising Grassland

of the Humid Mistbelt, Sabie

area. Axes 1 and 2 represent

the main environmental gra-

dients as shown.

Bothalia 20,2 (1990)

227

Superimposition of environmental variables on com-

munity ordination axes enhances the ecological inter-

pretability of floristic classifications. Environmental

gradients affecting community differentiation are thus

identified and successional trends become more apparent.

ACKNOWLEDGEMENTS

Discussions with Dr J.C. Scheepers and Mr R.H.

Westfall helped crystallize some of the theoretical concepts.

Drawings were all done by Mrs J. Schaap.

REFERENCES

DEALL, G.B. 1985. A plant-ecological study of the Eastern Transvaal

Escarpment in the Sabie area. M.Sc. thesis, University of

Pretoria.

DEALL, G.B., SCHEEPERS, J.C. & SCHUTZ, C.J. 1989. The vege-

tation ecology of the Eastern Transvaal Escarpment in the Sabie

area. 1. Physical environment. Bothalia 19: 53—67.

GAUCH, H.G. Jr 1982. Multivariate analysis in community ecology.

Cambridge, Cambridge University Press.

GAUCH, H.G. Jr & WHITTAKER, R.H. 1981. Hierarchical classifica-

tion of community data. Journal of Ecology 69 : 537—557.

HILL, M .O. 1979. DECORANA — A FORTRAN program for detrended

correspondence analysis and reciprocal averaging. Unpublished

report. Ecology and Systematics, Cornell University, Ithaca, New

York.

HILL, M.O. & GAUCH, H.G. Jr 1980. Detrended Correspondence

Analysis: an improved ordination technique. Vegetatio 42: 47—58.

MUELLER-DOMBOIS, D. & ELLENBERG, H. 1974. Aims and

methods of vegetation ecology. John Wiley, New York.

WESTFALL, R.H. 1983. The plant ecology of the farm Groothoek,

Thabazimbi District. 1. Ordination. Bothalia 14: 785—789.

WESTFALL, R.H., DEDNAM, G., VAN ROOYEN, N. & THERON,

G.K. 1982. PHYTOTAB — a program package for Braun-Blanquet

tables. Vegetatio 49 : 35 — 37.

Bothalia 20,2: 229-231 (1990)

Miscellaneous notes

DESCRIPTIVE ECOLOGICAL ACCOUNT OF INTENSIVE SPRING FLOWERING OF EPHEMERAL VEGETATION IN THE

BOSHOF AREA, ORANGE FREE STATE, SOUTH AFRICA

INTRODUCTION

A farmer, Mr B. Slabbert alleged that he had never

before experienced vernal communities showing such

intense flowering in the Boshof area as during spring 1988.

The two authors visited the area in October 1988. The

ephemeral vegetation was still in full bloom and a total

of five sites were inspected. The key questions researched

were: 1, which species were forming these communities;

2, what were the ecological conditions and the structure

of the communities; 3, what could be the possible causes

of the phenomenon?

THE STUDY AREA

The Boshof District is situated in the Orange Free State,

South Africa. The approximate altitude of the study sites

is 1 200 m a.s.l. and Figure 1 indicates their location.

Geology, soils and climate

The mostly shallow Hutton Form soils are overlying

sediments (sandstone, siltstone, shales) of the Dwyka and

Ecca strata and dolerites of the Karoo Supergroup. Some-

times a calcareous hardpan was present.

The climate of Boshof is typically semi-arid, with very

hot summers and mild to cold winters. The hottest months

are December, January and February and the coldest

months June and July. Table 1 gives rainfall figures meas-

ured at Boshof.

Phytogeography

According to Rutherford & Westfall (1986), the area falls

into the Savanna Biome. They define savanna as vegeta-

tion with a herbaceous, usually graminoid layer, and an

upper layer of woody plants forming a widely spaced to

closed canopy cover. The area is part of Veld Type 16,

Kalahari Thornveld (Acocks 1988).

METHODS

Five sites were subjectively chosen and studied along

the Christiana-Boshof and Spioenheuvel-Warrenton roads,

on the Farms Biesenvalley, Kareekloof, Spioenheuvel and

Wintershoek (Figure 1). At each site a collection of the

flowering plants, notes on vegetation structure, composi-

tion and soils were made. Plants were identified at the

Botanical Research Institute and plant names follow Gibbs

Russell et al. 1985, 1987. Background information was

obtained from the extension officer Mr Nolly van Rensburg

and from the farmer, Mr Basie Slabbert.

RESULTS

Site 1

The first site (Figure 1) was situated on the side of the

Warrenton road at Biesenvalley Farm on a slightly raised

landscape on an upper pediment slope with shallow, red

soils of the Hutton Form and probably belonging to the

Mangano Series (N. van Rensburg pers. comm.). This type

of soil has fine sand in the B21 horizon and a clay content

between 6% and 15%. The slope is gentle and occasional

dolerite cropped out at the surface. Occasional termitaria

were also present, capacity is 13 ha per large stock unit.

This community is classed ‘sweet Fires are rare in this

TABLE 1. — Rainfall (in mm), preceding the flowering, for the station

at Boshof Prison. The mean averages over 53 years are given.

Source: Department of Agricultural Development, Kimberley

FIGURE 1. — Map indicating the positions of the five study sites

230

Bothalia 20,2 (1990)

is classed ‘sweet veld’, a grassland with occasional trees

(mainly Acacia spp.). The community had an estimated

total cover of 25 % , comprising a lower, grass-dominated

layer, with a conspicuous layer, up to 0,25 m tall, mainly

formed by Lobelia erinus (Figure 2) and Wahlenbergia

denticulata, and an emergent layer of grasses up to 0,8

m tall. In the shrub layer, Acacia tortilis was present with

a very low cover value. Grasses present were Eragrostis

superba, E. lehmanniana, Themeda triandra, Aristida

congesta, A. vestita, Heteropogon contortus and Cenchrus

ciliaris. In Table 2 all recorded species are listed.

Site 2

The second site was situated on a flat bottomland. The

red soils belong to the Hutton Form and probably to the

Shorrocks Series (N. van Rensburg pers. comm.). These

soils are about 0,5 m deep, have a high clay content of

15% to 35% and an eutrophic B21 horizon. The usual

vegetation is a sweet grassveld. This community showed

an estimated total cover of 20%, a low grass layer of up

to 0,03 m, a layer dominated by Lobelia erinus up to 0,25

m and a layer up to 0,5 m tall together with a shrub layer

up to 4 m tall with an estimated cover of only 1%. It is

known that these areas were pure grassveld before, and

the colonizing of the area by Acacia species is a recent

phenomenon. The dominant and most conspicuous plant

was Lobelia erinus. The same grasses as in Site 1 were

present, but in a poorer condition, owing to overgrazing.

All species recorded are listed in Table 2.

Site 3

This site was situated on the Farm Kareekloof (owner

Mr Botha), a flat bottomland, nearby a pond. The

vegetation cover was uneven, varying from very dense to

absent in patches. This could be related to local

overgrazing. There were only a few Acacia spp. shrubs,

and these were widely spaced. Among the flowering plants

Lobelia erinus, L. angolensis and Wahlenbergia denticula-

ta were the most common. All species recorded are listed

in Table 2.

Site 4

This site, on the farm Spioenheuvel, near the dwellings

of Mr Slabbert, is usually covered with sweet grassveld.

It is situated on a flat bottomland, with a soil belonging

to the Hutton Form and to the Mangano Series. It is a

sand with about 8% clay and has a depth of 0,3 m to 0,6

m. It is basic in reaction with calcrete layers at variable

depth.

The vegetation of the site consists of a dense forbland,

with occasional trees such as Acacia erioloba and Rhus

lancea. Total estimated cover was 45% with a lower forb

layer of 0-0,3 m (with 20% cover) and a higher layer from

0,03-0,5 m (15% cover). The dominating species, giving

the blue colour to this vegetation, was Wahlenbergia

androsacea. Species of Homeria, Lobelia, Felicia and

Nidorella were also recorded. Grasses present were

Anthephora pubescens, Stipagrostis uniplumis, Eragrostis

lehmanniana and Schmidtia pappophoroides. All species

recorded are listed in Table 2.

We were informed that this site had not been ploughed

before, but was only utilized for grazing.

Site 5

This site was situated on the roadside from Christiana

to Boshof, on the Farm Wintershoek of Mr van Wyk. The

usual vegetation is that of a sweet veld. The soils are

similar to Site 4. A strong woody element is prevalent in

TABLE 2. — Table of taxa (names after Gibbs Russell et al. 1985, 1987)

found on the five sites

Bothalia 20,2 (1990)

231

FIGURE 2. — Site 1 with abundant

herbaceous vegetation with

occasional shrubs of Acacia

spp. (e.g. Acacia tortilis, A.

karroo and A. erioloba). The

dominant flowering species in

this area were Lobelia erinus

and Wahlenbergia denticulata

(1988.10.11).

this community, represented by patches of Acacia spp. No

structural data were gathered from this site. The species

giving the blue colour to the vegetation was Wahlenbergia

androsacea. Other species recorded are listed in Table 2.

DISCUSSION

The preceding descriptions give an idea of the structure

and species composition of the vegetation. All species

found forming the communities are typical of the area.

The number and the degree of development and flowering

was extraordinary. It can be proposed that the unusually

conspicuous flowering could be related to several factors.

The very high rainfall of 450 mm in February, 116mm in

March and 183 mm in April 1988 allowed abundant water

storage in the soil which was then available in spring for

plant growth. The February rainfall surpassed the mean

annual rainfall. The increased water availability promoted

high germination and establishment rates of dormant seeds.

The additional spring rains during September 1988 of

29,5 mm maintained the soil moisture at high levels.

Overgrazing resulted in reduced competition from

perennial plants. Species such as Lobelia erinus and

Wahlenbergia androsacea are able to germinate and grow

during periods of relatively low temperature when grasses

are still mainly dormant. In this way there is little or no

competition for environmental resources by grasses.

REFERENCES

ACOCKS, J.P.H. 1988. Veld types of South Africa, 3rd edn. Memoirs

of the Botanical Survey of South Africa No. 57.

GIBBS RUSSELL, G.E. et al. 1985. List of species of southern African

plants. Edn 2, Part 1, pp. 152. Memoirs of the Botanical Survey

of South Africa No. 51.

GIBBS RUSSELL, G.E., et al. 1987. List of species of southern African

plants. Edn 2, Part 2, pp. 270. Memoirs of the Botanical Survey

of South Africa No. 56.

RUTHERFORD, M.C. & WESTFALL, R.H. 1986. Biomes of southern

Africa — an objective categorization. Memoirs of the Botanical

Survey of South Africa No. 54.

WEISSER, P.J. & GERMISHUIZEN, G. 1988. Ephemeral vegetation

in the Boshof area, Orange Free State, South Africa, pp.23.

National Botanical Institute, Pretoria. Unpublished.

P.J. WEISSER* and G. GERMISHUIZEN*

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

Bothalia 20,2: 233-240 (1990)

The vegetation of the north-western Orange Free State, South Africa.

1. Physical environment

M.S. KOOIJ* G.J. BREDENKAMP* and G.K. THERON*

Keywords: climate, geology, habitat, land types, physiography, soil, western Grassland Biome

ABSTRACT

The physiography, geology, soil, land types and climate of the north-western Orange Free State are described. The description

provides a contextual framework for the subsequent vegetation classification.

UITTREKSEL

Die fisiografie, geologie, grand, landtipes en klimaat van die noordwestelike Oranje-Vrystaat word beskryf. Die beskrywing

verskaf ’n gekoordineerde raamwerk vir die daaropvolgende plantegroeiklassifikasie.

INTRODUCTION

The north-western part of the Orange Free State is one

of the most important agricultural regions in South Africa.

The study area represents the south-western part of the

Highveld Agricultural Region. This region produces 80%

of the maize, 75% of the grain sorghum and 65% of the

sunflower seed output of the Republic of South Africa

(Scheepers 1975). This part of the Grassland Biome also

renders a large part of the Republic’s animal products

(Scheepers 1975; Mentis & Huntley 1982). Most of the

land has been ploughed, mainly for maize cultivation. The

remaining natural vegetation is restricted to non-arable

shallow or rocky soils, vertic clays in bottomland

situations, seasonally waterlogged vleys and along drainage

lines. The vegetation is often overgrazed by sheep and

cattle.

Little is known about the vegetation of the north-western

Orange Free State. The relevance of plant ecological

studies to land use planning and management is well

documented (Edwards 1967; Walker 1976; Bredenkamp

& Theron 1978; Muller 1983). A more detailed classifi-

cation of vegetation than that of Acocks (1988) is necessary

to meet the present needs for regional and subregional

planning (Deall, Scheepers & Schutz 1989). It was

therefore necessary to identify, classify and map the

vegetation in order to permit efficient land use planning

and also the compilation of management programmes for

optimal utilisation, without the degradation of vegetation.

In this account the fundamental physical environment of

the study area is described.

STUDY AREA

The study area represents the western parts of the 2726

Kroonstad map (1 :250000). The area is situated between

26°00' and 27°23' E longitude and 27°00' and 28°00' S

latitude. Towns situated in the study area are Kroonstad,

Welkom, Bothaville, Hennenman, Viljoenskroon,

Vredefort and Wesselsbron (Figure 1). The area covers

* Department of Botany, University of Pretoria, 0002, Pretoria.

MS. received: 1989.09.07.

approximately 1 437 000 ha and about 1 365 000 ha is

available for agriculture (Land Type Survey Staff 1984).

GEOLOGY

The geology of the study area is presented in Figure 2.

Archaic granite

Old Archaic granites are exposed in the vicinity of

Vredefort where they form the central part of the Vredefort

Dome. The round or castle-like koppies of stacked granite

blocks are typical of this area. These granites occur only

in the extreme north-eastern part of the study area. They

are grey-white in colour and consist of quartz, potassium

feldspar, plagioclase and biotite. The Archaic granite

weathers to form a coarse, sandy soil of the Glenrosa Form

in the uplands, and the Valsrivier and Sterkspruit Forms

in the lowlands (Harmse 1967).

Witwatersrand Supergroup

Rocks of the Witwatersrand Supergroup are exposed in

a small part of the study area. They occur adjacent to the

Archaic granites and, together with the Ventersdorp

Supergroup, represent the outer rim of the Vredefort

Dome.

The Witwatersrand Supergroup is of great economical

value, seeing that it is the source of gold-bearing ore in

South Africa. The Supergroup consists of alternating

groups of quartzite and shale or slate. The gold ore is

situated in fine layers in conglomerate. The resistance to

erosion of the various rock types differs greatly. Quartzite

is very resistant and therefore forms predominant parallel

ridges. It gives rise to coarse sandy shallow lithosols

( < 300 mm) and regosols represented by the Mispah Form

(Harmse 1967). The softer shale and slate weather easily

to form the valleys between the quartzite ridges. The shale

and slate erode to form dark clayey soils.

Ventersdorp Supergroup

This Supergroup is represented by outcrops along the

Vaal River and the adjacent outer layers of the Vredefort

Dome. A few outcrops are also found near Odendaalsrus.

234

Bothalia 20,2 (1990)

FIGURE 1 —The distribution of the different land types in the study area (Land Type Survey Staff 1984). Ae; Ai; Ba- Be Bd Db De Fb'

lb; hatched square is portion studied by Scheepers (1975), around Kroonstad.

The lavas of the Ventersdorp Supergroup cover the

Witwatersrand Supergroup. The Ventersdorp Supergroup

consists of dark blue-grey Andesitic lava with round

amygdaloids. The Andesitic lava consists of plagioclase

and pyroxene which weather easily, resulting in a flat

landscape with dark-coloured vertic soils with a high

intrinsic fertility. This Supergroup is often covered with

recent sand deposits.

Transvaal Sequence

Rocks of this Sequence are restricted to a small part in

the north of the study area. It occurs between the

Ventersdorp Supergroup outcrops adjacent to the Vaal

River and the Vredefort Dome (Figure 2). Two groups can

be identified in this Sequence namely the Chuniespoort

Group and the Pretoria Group.

Chuniespoort Group

This Group is situated between the Pretoria Group and

the Ventersdorp lavas. It is represented by the Malmani

Subgroup, previously known as the Dolomite Series.

It consists mainly of dolomitic limestone and chert.

Other minerals such as calcite and dolomite may be

present. The rocks are also rich in iron and magnesium

(Truswell 1977). Chert is a hard, extremely compact,

dull to semi-vitreous cryptocrystalline rock, consisting

dominantly of cryptocrystalline silica (Harmse etal. 1984).

These rocks were formed during a chemical deposition

of silica in water (Kruger 1971). Dolomite and chert

are both resistant to weathering. The result is a flat land-

scape with exposed dolomite and chert outcrops. The

soils of this area are mostly very shallow and rocky,

repesenting the Mispah, Glenrosa and shallow Hutton soil

forms.

Bothalia 20,2 (1990)

Pretoria Group

Within the study area this group is limited to a small

area north of Viljoenskroon. It consists mainly of quart-

zite and shale. Igneous rock occurs regularly and two types

are distinguished, namely Ongeluks lava that consists of

a dark green to grey Andesitic lava with quartz

amygdaloids and intrusive diabase plates. The soils derived

from the shale, lavas and diabase are usually clayey and

of the Bonheim, Arcadia or Rensburg Forms, whereas the

sandy soils derived from the quartzite are mostly of the

Mispah, Glenrosa and Hutton Forms.

Karoo Sequence

This Sequence occupies about 80% of the study area.

It comprises a thick shale layer, mudstone and sandstone

with tillite at the base and basaltic lava as a canopy. These

strata were not disturbed by earth movements. Four groups

can be distinguished, but only two of the four occur in

the study area, namely the Ecca and the Beaufort Groups.

235

Ecca Group

Sediments of this Group are approximately 200—300

m thick. A considerable amount of fossils can be found

in the sediments of this group. The Group can be

subdivided into three subdivisions, namely the Lower

Stage, consisting of soft dark blue shales; the Middle Stage,

consisting of grit, shale and coal and the Upper Stage, con-

sisting of soft, dark shale, usually covered with recent ae-

olian sand deposits.

The grit and sandstone are resistant to weathering and

form low hills and escarpments. These sediments are

usually covered with recent, deep, fine aeolian sands.

Beaufort Group

The Beaufort Group covers the south-eastern part of the

study area. The sediments are a sequence of shale and

mudstone with interbedded lenticular sandstone (Truswell

1977).

FIGURE 2. — The geology of the study area (Harmse 1967). Karoo Sequence: Beaufort Group: shale, mudstone arenaceous shale immature sandstone,

extensively invaded by dolerite dykes and sheets; Ecca Group: shale and mudstone (carbonaceous and calcareous), immature sandstone

and coal seams, extensively invaded by dolerite sheets and dykes. Transvaal Sequence: Pretoria Group: quartzite, banded ironestone, subor-

dinate shale and lava; Chuniespoort Group: dolomitic limestone, chert, thin band of quartzite and conglomerate at the base. Ventersdorp

Supergroup: andesitic lava, agglomerates and tuffaceous sediments. Witwatersrand Supergroup: quartzite, conglomerate shale, lava, slate

and tillite. Granite.

236

Bothalia 20,2 (1990)

The Karoo sandstone is soft and easily weathered to form

a variety of deep, red to yellow sandy soils. On the other

hand mudstone, shale and dolerite weather rapidly to give

rise to both sedentary and colluvial, base-rich, dark,

margalitic clays.

Recent deposits

Aeolian sand

The colour of the aeolian sand of the north-western

Orange Free State varies from red to grey. The colours

are a result of a fine layer of iron oxide that covers the

quartz and feldspar grains (Harmse 1967). The only

difference between the sands of the Kalahari area and the

north-western Orange Free State is that the sands of the

latter are covered with dense vegetation and that no

prominent dunes occur.

PHYSIOGRAPHY

The study area forms part of the Highveld inland plateau

region and consists of smoothly plained or gently rolling

land surfaces of the Tertiary and Miocene age (Mentis &

Huntley 1982).

The monotonous soft rolling landscape on Karoo

sediments is situated between 1 200 and 1 400 m above

sea level. This physiographic unit stretches from the Vaal

River south-eastwards to Theunissen and Ventersburg.

Ventersdorp lavas are responsible for the formation of

round hills whereas the uplands and mesas of Karoo

dolerite are formed by relics of Tertiary erosion surfaces.

Erosion within this physiographic unit is responsible for

the formation of peneplains with vertic, impermeable,

argillic soils of the Ecca Group. These clay peneplains are

usually unsuitable for agronomy.

In areas covered with aeolian sand, most of the surface

characteristics have been destroyed and therefore only an

undulating landscape is visible. The presence of Acacia

karroo is often the only indication of the existence of the

dolerite sills. In the waterlogged areas of this landscape

pans are a prominent characteristic. The pans can be

subdivided according to their size and age.

The occurrence of large pans is restricted to clay flats

situated on argillic sediment of the Upper Ecca. The form

of the pans is uneven. These pans have an inflow but no

outlet, and therefore the drainage is internal. The pan

floors consist of dark waterlogged alkaline soils. During

desiccation these soils form block fragments with gypsum

crystals. A sparse vegetation may cover the pan floor.

Small pans are situated on the grey sands with a high

water table. Iron and lime concretions occur frequently

on the pan floors. No vegetation is found on the floors

of these pans.

In the northern parts of the study area the gently

undulating landscape of the Vredefort Dome occurs. Rocky

hills and ridges of quartzite, shale and lavas of the

Witwatersrand Supergroup and Transvaal Sequence form

the rim of the Vredefort Dome. This landscape is steeply

dissected and the Vaal River cuts deeply through the rocky

ridges.

Drainage

The study area is situated in the catchment area of the

Vaal River. Drainage occurs along the Vaal, Renoster and

Vais Rivers and their tributaries, as well as flushes and

seepages into pans. The Karoo Sequence is deeply incised

by these rivers, gradually flattening towards the south-east.

Due to the gradual slopes in the area, erosion is

restricted to rivers and flushes. No marshy areas are

formed in the aeolian sand with a high permeability, but

the clayey flats usually form marshy areas during the rainy

season. This gives rise to the formation of the large pans

which are responsible for the internal drainage of the area

(Harmse 1967).

LAND TYPES AND SOILS

A land type denotes an area that can be shown on a

1:250 000 scale map and displays a marked degree of

TABLE 1.— The mean monthly rainfall for weather stations in the study area

B, Balkfontein-Bothaville No. 0399/894 4; H, Hennenman Police Station No. 0365/058 4; K, Kroonstad Municipality No. 0365/430 8; P, Plessisdraai-

Hoopstad No. 0363/239 5; Vk, Viljoenskroon Municipality No. 0400/792 9; Vf, Vredefort School No. 0437/660 7; W, Welkom No. 0364/300

1; **, Unreliable data; #, Weather Burea" 1986; +, Weather Bureau 1989.

Bothalia 20,2 (1990)

237

FIGURE 3. — Climatic diagrams for the study area. A, station; B, height above sea level; C, duration of observations in years (indicates precipita-

tion); D, mean annual temperature in °C; E, mean annual precipitation in mm; F, mean monthly precipitation; G, wet period; H, mean

monthly temperature; I, dry period; J, cold season (months with mean daily minimum below 0°C); K, months with absolute minimum

below 0°C; L, frost-free period (Larcher 1975).

uniformity with respect to terrain form, soil pattern

and climate. One land type differs from another in

terms of one or more of the following: terrain form,

soil pattern or climate (Land Type Survey Staff

1984).

Five different land types are distinguished in the

study area, namely the A, B, D, F and I land types

(Figure 1). Each of these land types can be further sub-

divided. They are discussed in detail in Land Type

Survey Staff (1984) and in the following papers: A land

type (Kooij et al. 1990c), B land type (Kooij et al.

1990b), D land type (Kooij et al. 1990a) and the Fb,

lb and Ba land types (Kooij et al. 1990d). Nomenclature

of soil forms follows MacVicar et al. (1977).

CLIMATE

The study area is situated in the H climate zone of the

Highveld region, according to the classification of the

Weather Bureau (1986). This region is characterised by

thunderstorms during the summer months. The winter

months are arid and cold.

Precipitation

Soil moisture is derived from precipitation mainly in

the form of rainfall, and to a lesser degree from mist, dew,

hail and snow (Deall, Scheepers & Schutz 1989).

Rainfall: the mean monthly rainfall for Balkfontein

(33 yrs), Kroonstad (33 yrs), Plessisdraai (10 yrs) and

238

Bothalia 20,2 (1990)

FIGURE 4.— Wind roses for two weather stations in the study area, representing wind directions and speed intervals (m/s)

for January, April, July and October (Weather Bureau 1988). A, 5% frequency intervals; B, percentage of calm in circle.

Bothalia 20,2 (1990)

239

TABLE 2. — The monthly number of days with frost

Weather station

Plessisdraai Balkfontein Welkom Kroonstad

The occurrence of frost is taken as days with a minimum temperature of <2,5°C (Weather Bureau 1989).

Welkom (23 yrs) is presented in Table 1 and Figure 3

(Weather Bureau 1986). Mean monthly rainfall figures for

a number of stations in the study area for the years

1987/1988, are given in Table 1 (Weather Bureau 1989).

Table 1 indicates the above average rainfall of the two years

(1987—1988) in which the vegetation study took place. The

study area clearly falls within the summer rainfall zone,

with most of the precipitation from October to March.

Annual totals average 611 mm at Kroonstad in the eastern

parts and 503 mm at Plessisdraai in the west. The rainfall

also increases from south to north, with an average of 526

mm at Welkom in the south and an average of 611 mm

at Kroonstad in the north. Thunderstorms are often violent

and associated with strong south-westerly gusting winds

and hail.

Frost is defined by the Weather Bureau as days with a

minimum temperature of 2,5°C and lower. Frost can be

expected from April to September in the study area (Ta-

ble 2).

Snow: due to the dryness of the winter, snow does not

often occur. Snowfalls were, however, recorded during

June and July in the study area in 1987 and 1988.

Temperature

Temperature alone may not be a significant factor in

determining major regional vegetation formations, it does

however, play a part in the determination of floristic

variations on a meso- and micro-scale (Schulze & McGee

TABLE 3. —The mean monthly maximum and minimum temperature (°C) for lour weather stations in the study area

Year

1951/84

Weather stations

Balkfontein Kroonstad Plessisdraai Welkom

1987/88

240

Bothalia 20,2 (1990)

TABLE 4. — The extreme temperatures (°C) recorded at four weather

stations in the study area

1978). Such variations result from differential effects of

temperature on plant growth rates, seed germination,

seedling survival and flowering phenology (Deall et al.

1989). The average temperature is given in Figure 3 and

the mean monthly maximum and minimum temperatures

of the Plessisdraai, Welkom, Balkfontein and Kroonstad

weather stations are represented in Table 3. There are no

big differences between the maximum and minimum tem-

peratures recorded at the different weather stations. The

extreme maximum and minimum temperatures recorded

at the four stations are represented in Table 4 (Weather

Bureau 1986).

Wind

Wind directions and velocities for the Welkom and

Kroonstad weather stations are represented in Figure 4.

The calmest days occur during May, June and July,

whereas August and September are the windiest months

(Weather Bureau 1989).

CONCLUSION

A knowledge of the physical environment of the study

area is necessary for the understanding and ecological

interpretation of the abstract plant communities identi-

fied during a phytosociological survey. Environmental

attributes largely determine the distribution of plant species

and plant communities. These differences in distribu-

tion patterns make the classification of vegetation into

ecological zones possible.

ACKNOWLEDGEMENT

This research was financially supported by the Foun-

dation for Research Development, CSIR.

REFERENCES

ACOCKS, J.RH. 1988. Veld types of South Africa, 3rd edn. Memoirs

of the Botanical Survey of South Africa No. 57: 1-146.

BREDENKAMP, G.J. & THERON, G.K. 1978. A synecological account

of the Suikerbosrand Nature Reserve. 1. The phytosociology of

the Witwatersrand geological system. Bothalia 12: 513-529.

DEALL, G.B., SCHEEPERS, J.C. & SCHUTZ, C.J. 1989. The vege-

tation ecology of the Eastern Transvaal Escarpment in the Sabie

area. 1. Physical environment. Bothalia 19: 53—67.

EDWARDS, D. 1967. A plant ecological survey of the Tugela Basin.

Memoirs of the Botanical Survey of South Africa No. 36: 1—285.

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South Africa. D.Sc. dissertation, Rijks University, Utrecht.

HARMSE, H.J. VON M., VAN DER WATT, H.v.H., VAN ROOYEN,

T.H. & BURGER, R.D.U.T. 1984. Glossary of soil science terms.

The Soil Science Society of South Africa, Pretoria.

KOOIJ, M S., BREDENKAMP, G.J. & THERON, G.K. 1990a. The

vegetation of the north-western Orange Free State, South Africa.

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KOOIJ, M.S., BREDENKAMP, G.J. & THERON, G.K. 1990b. Clas-

sification of the vegetation of the B land type in the north-western

Orange Free State. South African Journal of Botany 56: 309—318.

KOOIJ, M.S., BREDENKAMP, G.J. & THERON, G.K. 1990c. The

vegetation of the deep sandy soils of the A land type in the north-

western Orange Free State, South Africa. Botanical Bulletin of

Academia Sinica 31. In press.

KOOIJ, M.S., BREDENKAMP, G.J. & THERON, G.K. 1990d. The

plant communities of the hills and ridges in the north-western

Orange Free State, South Africa. Botanical Bulletin of Acade-

mia Sinica 31. In press.

KRUGER, J.A. 1971. 'n Ekologiese ondersoek van die plantegroei van

die plaas Somerville 53 en omgewing (Dist. Ventersdorp),

met besondere aandag aan die bodemkundige aspek. M.Sc.

thesis, Potchefstroom University for Christian Higher Education,

Potchefstroom.

LAND TYPE SURVEY STAFF 1984. Land types of the maps 2626

'Wesrand, 2726 Kroonstad. Memoirs on the Agricultural Natural

Resources of South Africa No.4: 1—441.

LARCHER, W. 1975. Physiological plant ecology. Springer-Verlag, Berlin.

MACVICAR, C.N., LOXTON, R.F., LAMBRECHTS, J.J.N., LE

ROUX, J., DE VILLIERS, J.M., VERSTER, E.,

MERRYWEATHER, F.R., VAN ROOYEN, T.H. & HARMSE,

H .J. VON M . 1977. Soil classification, a binomial system for South

Africa. Department of Agricultural Technical Services, Pretoria.

MENTIS, M.T. & HUNTLEY, B.J. 1982. A description of the Grassland

Biome Project. Co-operative Scientific Programme, Council for

Scientific and Industrial Research. Report No. 62. Graphic Arts

Division of the CSIR, Pretoria.

MULLER, T. 1983. A case for a vegetation survey in a developing country

based on Zimbabwe. Bothalia 14 : 721—723.

RUTHERFORD, M.C. & WESTFALL, R.H. 1986. Biomes of southern

Africa — an objective categorization. Memoirs of the Botanical

Survey of South Africa No. 54: 1—97.

SCHULZE, R E. & McGEE, O.S. 1978. Climatic indices and classifi-

cation in relation to the biography of southern Africa. In M.J. A.

Werger, Biogeography and ecology of southern Africa : 19—52.

Junk, The Hague.

TRUSWELL, J.F. 1977. The geological evolution of South Africa. Purnell,

London.

SCHEEPERS, J.C. 1975. The plant ecology of the Kroonstad and

Bethlehem areas of the Highveld Agricultural Region. D.Sc.

dissertation. University of Pretoria.

WALKER, B.H. 1976. An approach to the monitoring of changes in the

composition and utilization of woodland and savanna vegetation.

South African Journal of Wildlife Resource Management 6 : 1 — 32 .

WEATHER BUREAU 1986. Climate of South Africa. Climate statistics

up to 1984. Government Printer, Pretoria.

WEATHER BUREAU 1988. Climate of South Africa. Part 12. Surface

Winds. Government Printer, Pretoria.

WEATHER BUREAU 1989. Climatological data for 1987-1989 for the

Balkfontein, Plessisdraai, Kroonstad and Welkom weather sta-

tions. Computer printout, Weather Bureau. Pretoria.

Bothalia 20,2: 241-248 (1990)

The vegetation of the north-western Orange Free State, South Africa.

2. The D land type

M.S. KOOIJ*,' G.J. BREDENKAMP* and G.K. THERON*

Keywords: classification, habitat interpretation, plant communities, western Grassland Biome

ABSTRACT

As part of a research programme on the synthesis of the vegetation of the western Grassland Biome, the plant communities

of the D land type of the north-western Orange Free State were investigated. The results of a numerical classification

(TWINSPAN) of 59 releves were refined by Braun-Blanquet procedures. The analyses revealed eight plant communities

which may be grouped into two major communities. A hierarchical classification, description and ecological interpretation

ot the plant communities, and an ordination (DECORANA) with associated environmental gradients, are presented.

UITTREKSEL

Die ondersoek na die plantgemeenskappe van die D-landtipe in die noordwestelike Oranje-Vrystaat vorm deel van 'n

navorsingsprogram oor die sintese van die plantegroei van die westelike Grasveldbioom. Die resultate van ’n numeriese

klassifikasie (TWINSPAN) van 59 releves is met behulp van Braun-Blanquetprosedures verfyn. Agt plantgemeenskappe wat

in twee hoofgemeenskappe gegroepeer kan word, is onderskei. n Hierargiese klassifikasie, beskrywing en ekologiese inter-

pretasie van die plantgemeenskappe en ’n ordening (DECORANA) met geassosieerde omgewingsgradiente, word verskaf.

INTRODUCTION

The necessity to identify and describe the major vege-

tation types and subtypes within the Grassland Biome was

stated by Mentis & Huntley (1982) and Scheepers (1986).

One of the major goals of the vegetation classification and

mapping task group of the Grassland Biome Project is to

produce a vegetation classification and to map the region

on a 1:250 000 scale (Scheepers 1986). As mentioned by

Bredenkamp, Joubert & Bezuidenhout (1989), the first step

in the synthesis of the vegetation of the western Grassland

Biome, is to create a phytosociological data base for the

entire area. Little is known of the vegetation of the north-

western Orange Free State. Scheepers (1975) conducted a

semi-detailed survey of the vegetation in the Kroonstad

area, and Du Preez (1987) described some woody plant

communities from the Vredefort District. The north-

western Orange Free State was therefore identified as a

priority area for a phytosociological survey.

Extensive bottomland areas, representing the D land

type (Land Type Survey Staff 1984) occur in the north-

western Orange Free State. As the margalitic, vertic,

melanic or wet structured fersialitic soils of the D land

type are often not ploughed and mainly used as pasture

for sheep and game, this land type was chosen for the

initial vegetation survey.

THE STUDY AREA

The study area is situated between 26°00' and 26°23'

E longitude and 27°00' and 28°00' S latitude, in the Grass-

land Biome of the north-western Orange Free State (Kooij

et al. 1990: 234, fig. 1).

The Db and Dc map units accommodate land where

duplex soils with gleycutanic, pedocutanic and prisma-

* Department of Botany, University of Pretoria, 0002, Pretoria.

MS. received: 1989.09.07.

cutanic B horizons are predominant. These map units

comprise approximately 280 000 ha of the study area, with

approximately 35 700 ha unsuitable for agriculture (Land

Type Survey Staff 1984). Soils that display a duplex

character include the Estcourt, Sterkspruit, Swartland,

Valsrivier and Kroonstad Forms (MacVicar et al. 1977).

The Dc and Db map units are mostly located in bottom-

land situations, such as in the valleys of the Vaal, Vais and

Renoster Rivers, as well as in the Pan Veld situated in the

Wesselsbron and Welkom Districts. These map units are

not suitable for agronomy, due to the high clay (>35%)

content of these bottomland soils, and land use is mainly

restricted to sheep farming. The bottomlands are further-

more usually flooded during the rainy season.

The Db map unit refers to land where duplex soils with

non-red B horizons comprise more than half of the area.

The Dc map unit indicates land that qualifies for inclusion

in the D land type but, in addition to the duplex soils, more

than 10% of the land type comprise soil forms with one

or more of the following diagnostic horizons: vertic,

melanic or red-structured (Land Type Survey Staff 1984).

The geology of the Db map unit consists mainly of Ecca

sandstone, shale and mudstone with the sporadic intrusion

of dolerite sills. Mudstone, shale and sandstone of the

Tarkastad Subgroup occur in the eastern parts.

The Dc map unit representing the Pan Veld in the

Wesselsbron District consists mainly of the Sterkspruit

Form, whereas the soils of the Dc map unit in the vicinity

of the Vredefort Dome, mainly represents the Bonheim

Form. Pans occupy approximately 22% of this map unit.

The predominant geology of the area is Ecca sandstone,

mudstone and shale with the occasional intrusion of

dolerite plates. Calcrete occurs sporadically in these

bottomland pans. The plains between pans are mostly

covered with aeolian sand.

The soils of the Dc map unit adjacent to the rivers are

predominantly of the Bonheim and Mayo Forms. The

242

Bothalia 20,2 (1990)

TABLE 1. — A phytosociological table of the vegetation of the D land types of the north-western Orange Free State

Species with an occurrence of < 4 are omitted.

Bothalia 20,2 (1990)

243

TABLE 1. A phytosocioiogical table of the vegetation of the D land types of the north-western Orange Free State

Species with an occurrence of < 4 are omitted.

geology in these areas consists of Ecca shale, sandstone

and grit. In the vicinity of Kroonstad mudstone, shale and

sandstone of the Adelaide Supergroup are predominant,

with dolerite sills occurring here and there (Land Type

Survey Staff 1984).

METHODS

Releves were compiled in 59 stratified random sample

plots. As geology and land type are relatively uniform,

stratification was based on terrain units (De Beer 1988;

Land Type Survey Staff 1984). In accordance with

Scheepers (1975), plot sizes were fixed at 16 m2 for

grassland vegetation, while 100 m2 plots were used for

woody vegetation (Bredenkamp & Theron 1978). In each

sample plot total floristic composition, using the Braun-

Blanquet cover-abundance scale (Mueller-Dombois &

Ellenberg 1974) was noted. Taxon names generally

conform to those of Gibbs Russell et al. (1985 & 1987),

however, in accordance with Bredenkamp, Joubert &

Bezuidenhout (1989), Setaria flabellata Stapf and S.

sphacelata (Schumach.) Moss were treated as separate

species. Environmental data included identification of

terrain types, soil types and estimation of aspect, slope

and rockiness of the soil surface. Other soil properties were

obtained from Land Type Survey Staff (1984), Table 2.

Two way indicator species analysis (TWINSPAN)

(Hill 1979b) was applied to the floristic data set in order

244

Bothalia 20,2 (1990)

to derive a first approximation of the vegetation types of

the area. Refinement of this classification was done by the

application of Braun-Blanquet procedures (Behr &

Bredenkamp 1988; Bredenkamp, Joubert & Bezuidenhout

1989). The results are presented in a phytosociological

table (Table 1). In order to determine probable vegetation

gradients, an ordination algorithm, Detrended Correspon-

dence Analysis (DECORANA) (Hill 1979a) was applied

to the floristic data set.

RESULTS

Classification

In general the vegetation of the D land type can be

classified as a Panicum coloratum-Eragrostis eurvula

Grassland. Species constantly present include Panicum

coloratum, Pentzia globosa, Eragrostis eurvula, Eragrostis

obtusa and Bulbine narcissifolia (Species group L, Table

1). The vegetation is generally poor in species, with an

TABLE 2. — An analysis of the soil properties of the A and B horizons

Free State (adapted from L

average of 20 species per sample plot. The vegetation

shows some similarity to certain bottomland communities

described by Scheepers (1975). Direct comparisons are,

however, difficult due to the differences in techniques ap-

plied and also due to differences in the scale of the surveys.

The greater part of the vegetation is utilized as pasture

for sheep. The vegetation varies as a result of habitat

variations, for example in the topography, soil type,

drainage regime and vegetation management (grazing by

animals).

A hierarchical classification of the plant communities

is as follows:

1. Themeda triandra-Panicum coloratum Grassland

1.1 Themeda triandra-Elionurus muticus Grassland

1.2 Themeda triandra-Digitaria argyrograpta Grassland

1.3 Protasparagus laricinus-Acacia karroo Woodland

' the plant communities in the D land type of the north-western Orange

d Type Survey Staff 1984)

Bothalia 20,2 (1990

245

1.4 Themeda triandra-Hibiscus trionum Grassland

1.4.1 Themeda triandra-Felicia filifolia Grassland

1.4.2 Themeda triandra-Aristida bipartita Grassland

1.5 Themeda triandra-Echinochloa holubii Wetland

1.6 Setaria sphacelata-Eragrostis plana Wetland

2. Sporobolus ioclados-Lycium cinereum Pan Veld

Description of communities

1. Themeda triandra-Panicum coloratura Grassland

This major grassland type covers most of the D land

type, excluding the pans which occur scattered throughout

the area (Kooij et al. 1990: 234, fig. 1). The dominant

species in almost all the plant communities within this

major grassland type are Themeda triandra and Era-

grostis curvula. Within the study area, species group I

(Table 1) may be considered as diagnostic for this

Grassland.

1.1 Themeda triandra-Elionurus muticus Grassland

This community is found in upland situations on

relatively sandy and well drained soil, often with large

exposed rocks (Figure 1). The predominant soil types are

the Mispah and Glenrosa Forms and in non-rocky, sandy

areas the deep (>1 200) Hutton Form. No soil analysis

data on these soil types are available. The Themeda

triandra-Elionurus muticus Grassland is characterised by

species group A (Table 1). In typical situations Themeda

triandra dominates entirely, with canopy cover values of

up to 75%. Where Themeda triandra dominates, the

occurrence of other species is limited, with an average of

20 species per releve. Other grass species constantly

(>50% of the releves) found in this community are

the diagnostic species Elionurus muticus and Setaria

sphacelata and also Aristida congesta, Eragrostis superba,

Cymbopogon plurinodis, Heteropogon contortus, Panicum

coloratum and Eragrostis curvula. Non-grassy forbs occur

only scattered, but Commelina africana, Hermannia

depressa, Salvia radula, Crabbea acaulis, Lippia scaber-

rima, Oxalis sp., Hibiscus pusillus and the karoo dwarf

shrub Pentzia globosa occur constantly.

1.2 Themeda triandra-Digitaria argyrograpta Grassland

On slightly drier or overgrazed sites of the upland areas,

the total cover, and especially the cover of the grass

species, is less than in the general situation occupied by

the Themeda triandra-Elionurus muticus Grassland.

Although Themeda triandra is still dominant, Cymbopo-

gon plurinodis is locally conspicuous. An average of 33

species was recorded per sample plot. This community

is characterised by species group B (Table 1). Other species

constantly present in the releves representing the com-

munity, are the grasses Aristida congesta, Eragrostis

superba, Heteropogon contortus, Panicum coloratum and

the non-grassy forbs Commelina africana. Hibiscus pusil-

lus and Albuca sp.

1.3 Protasparagus laricinus-Acacia karroo Woodland

This Woodland is a bottomland community which

occurs along dry water courses or seasonal rivulets (Figure

FIGURE 1. — A schematic representation of the identified plant

communities in the Db and Dc map units. P. pan; R, river;

1, Themeda triandra-Panicum coloratum Grassland; 1.1, Themeda

triandra-Elionurus muticus Grassland; 1.2, Themeda triandra-

Digitaria argyrograpta Grassland; 1.3, Protasparagus laricinus-

Acacia karroo ’Woodland', 1.4, Themeda triandra-Hibiscus trionum

Grassland; 1.4.1, Themeda triandra-Felicia muricata Grassland;

1.4.2, Themeda triandra-Aristida bipartita Grassland; 1.5,

Themeda triandra-Echinochloa holubii Wetland; 1.6, Setaria

sphacelata-Eragrostis plana Wetland; 2, Sporobolus ioclados-

Lycium cinereum Pan Veld.

1) on vertic Rensburg and Arcadia soil Forms, on duplex

soils with clayey structured B horizons, e.g. the Sterk-

spruit, Valsrivier and Swartland Form, as well as on

younger alluvial soils such as the Oakleaf and Dundee

Forms (Table 2).

The vegetation is mostly severely overgrazed and in a

poor condition. The grass cover is usually low, associated

with an increase in density of Acacia karroo, Protaspara-

gus laricinus and Ziziphus zeyheriana and an enchroach-

ment of the karoo shrub Pentzia globosa. An average of

only 26 species was recorded per sample plot. The

community is characterised by species group C (Table 1).

Other constantly present species in the herbaceous layer

are Eragrostis superba, Corchorus asplenifolius, Lede-

bouria marginata, Panicum coloratum and Eragrostis

obtusa.

1.4 Themeda triandra-Hibiscus trionum Grassland

The extensive flat plains of the D land type are typically

covered by the Themeda triandra-Hibiscus trionum Grass-

land which represents the largest part of the Themeda

triandra-Panicum coloratum Grassland (Figure 1). The

vegetation is characterised by species group D (Table 1).

An average of 20 species was recorded per sample plot.

The vegetation is mostly dominated by Themeda triandra,

Eragrostis curvula and Panicum coloratum , with the karoo