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

A C. adiposa	+	+	?	7	2	©
B C. autographa	+ +	+	+	-	2	©	o
C C. fimbriata	+ +	+	+		2-4
D C. moniliformis	+ +	+	+	7	7		A
E C. paradoxa	+ ?	+	+	+	7
F O. ips	+ +	+	+	+	2	©	1B1
G O. multiannulatum	+ +	*	-	+	3-4		iffS
H O. piceae	+ +	**	+	7	2	©
I O. stenoceras	+ ?	+	?	7	2	o
J O. ulmi	+ +	+	7	+	3	©
A— E, ascospores with sheaths; F— J, ascospores without sheaths, except O. ips\ + fertile cell; ** = described as ascogenous hyphae.	= present; —	= absent; ?	= unknown; *	= described as

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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BAKSHI, B.K. 1951. Development of perithecia and reproductive structures in two species of Ceratocystis. Annals of Botany 15: 53—61

BANDONI, R.J., BISALPUTRA, A. A. & BISALPUTRA, T. 1967. Ascospore development in Hansenula anomala. Canadian Journal of Botany 45: 361—366.

BECKETT, A. 1981. Ascospore formation. In G. Turian & H R. Hohl, The fungus spore: morphogenetic control. Academic Press, New York.

BECKETT, A., ILLINGWORTH, R.F. & ROSE, A H. 1973. Ascospore development in Saccharomvces cerevisiae. Journal of Bacteriology II: 1054-1057.

BENNY, G.L., SAMUELSON, A. & KIMBROUGH, J.W. 1980. Ascocarp development in Chaetomidium arxii (Chaetomiaceae) with special emphasis on the ontogeny of its cephalothecoid peridium. Botanical Gazette 141: 347—352.

BLACK, S.H. & GORMAN, C. 1971. The cytology of Hansenula. III. Nuclear segregation and envelopment during ascosporogenesis in Hansenula wingei. Archives of Microbiology 79: 231—248.

BOYCE, J.S. 1961. Forest pathology. McGraw-Hill Book Company, New York.

CLARK, C.A. & MOYER, J.W. 1988. Compendium of sweet potato diseases. APS Press, St. Paul, Minnesota.

DADE, H.A. 1928. Ceratostomella paradoxa, the perfect stage of Thielaviopsis paradoxa de Seynes von Hohnel. Transactions of the British Mycological Society 13: 184—194.

DE HOOG, G.S. & SCHEFFER, R.J. 1984. Ceratocystis versus Ophiostoma: a re-appraisal. Mycologia 1C. 292—299.

DYBY, S.D. & KIMBROUGH. J.W. 1987. A comparative ultrastructural study of ascospore ontogeny in selected species of Peziza (Pezizales Ascomycetes). Botanical Gazette 148: 283—296.

ELLIOTT, J.A. 1925. A cytological study of Ceratostomella fimhriata . E. et H. Elliott. Phytopathology 15: 417-422.

ELLIS, D.H 1981a. Ultrastructure of thermophilic fungi I. Ascocarp morphology of Thermoascus aurantiacus. Transactions of the British Mycological Society 6: 457—465.

ELLIS, D.H. 1981b. Ultrastructure of thermophilic fungi III. Asco- carp morphology of Chaetomium thermophile and C. gracile. Transactions of the British Mycological Society 77: 165-177.

FURTADO, J.S. 1971 The septal pore and other ultrastructural features of the Pyrenomycete Sordaria fimicola. Mycologia 63: 104—113.

GARRISON, R.G., MARIAT, F., BOYD. K.S. & FROMENTIN, H. 1979. Perithecial ultrastructure and formation of ascospores of Ceratocystis stenoceras Robak C. Moreau. Annales de Micro- biologie 130: 3—21.

GWYNNE-VAUGHAN, H.C.I. & BROADHEAD, Q.E. 1936. Contri- butions to the study of Ceratostomella fimhriata. Annals of Botany L(CC): 747-758.

HASH1MOTO, T., GERHARDT, P, CONTI, S.F. & NAYLOR, H B. 1960. Studies on the fine structure of micro-organisms. V. Morphogenesis of nuclear and membrane structures during ascospore formation in yeast. Journal of Biophysical and Bio- chemical Cytology 7: 105—149.

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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.

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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.

REFERENCES

ANON. 1975. Forest tree species in Dwesa Forest and some of their uses.

Director of Forestry, Transkei. Unpublished.

B1GALKE, E. H . 1967. A list of plants used medicinally in the following 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, Botanical Research Unit, Grahamstown. Unpublished. -

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. Agricultural Journal of the Cape of Good Hope 16: 21—42, 104-114.

SIM , T.R. 1907. The forests and the forest flora of the colony of the Cape 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 poisonous plants of southern Africa. Livingstone, Edinburgh.

WATT, J.M. & BREYER-BRANDWIJK, M.G. 1962. Medicinal and poisonous plants in southern and eastern Africa. Edn 2. 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

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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

Family	No. species	No. genera
Asteraceae	26	15
Orchidaceae	16	8
Rubiaceae	16	9
Adiantaceae	13	4
Cyperaceae	13	7
Liliaceae	12	7
Aspleniaceae	11	1
Euphorbiaceae	11	7
Fabaceae	11	9
Poaceae	11	8
Lamiaceae	10	4
Rosaceae	8	6
Solanaceae	8	2
Acanthaceae	7	5
Flacourtiaceae	7	6
Aspidiaceae	6	4
Celastraceae	6	2
Scrophulariaceae	6	6
Anacardiaceae	5	3
Apiaceae	5	5
Blechnaceae	5	1
Gesneriaceae	5	1
Loganiaceae	5	3
Rutaceae	5	5
Dennstaedtiaceae	4	4
Hymenophyllaceae	4	2
Juncaceae	4	2
Lycopodiaceae	4	i
Polypodiaceae	4	3
Rhamnaceae	4	4
Thymelaeaceae	4	4
Urticaceae	4	3

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

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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

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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

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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

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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.

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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.

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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.

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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.

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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.

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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

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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

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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.

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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

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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.).

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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.

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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.

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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.

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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

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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.

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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.

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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.

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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.

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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

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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

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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)

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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