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. REFERENCES AINSWORTH, G.C., SPARROW, F.K. & SUSSMAN, A S. 1973. The Fungi. 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Ceratocystis ips associated with Orthotomicus erosus (Coleoptera: Scolytidae) on Pinus spp. in the Cape Province of South Africa. Phytophylactica 12: 65-69. WINGFIELD, M.J., VAN WYK, P.S. & MARASAS, W.F.O. 1988. Ceratocystiopsis proteae sp. nov. , with a new anamorph genus. Mycologia 80: 23—30. WISMER, C.A. 1961. Pineapple disease. Sugarcane Diseases of the World 1: 223-245. WOOD, F.A. & FRENCH, D.W. 1963. Ceratocystis fimbriata, the cause of stem canker of quaking aspen. Forest Science 9: 232—235. . Bothalia 20,2: 147-152 (1990) A preliminary checklist of Xhosa names for trees growing in Transkei COLIN T. JOHNSON* Keywords: ethnobotany, Transkei, trees, vernacular names ABSTRACT Xhosa names for 284 of the 392 tree taxa recorded for Transkei are listed. The semantics of the common names are discussed. The main moulding forces for these names were found to be: tree habit, characteristics shared with animals, medicinal uses and social or historic events. UITTREKSEL Xhosa-name vir 284 van die 392 boomtaksons wat in Transkei aangeteken is, word gelys. Die semantiek van die volksname word bespreek. Daar is bevind dat hierdie name hoofsaaklik op die volgende oorwegings gebaseer is: groeiwyse van boom, kenmerke in gemeen met diere. geneeskundige gebruike en sosiale of geskiedkundige gebeurtenisse. INTRODUCTION Over the years numerous people have collected and compiled various lists of Xhosa names for plants. Miller (1923) listed about 321 names of trees and shrubs. Palmer & Pitman (1972) in their publication give a multilingual glossary of Xhosa, Zulu and Sotho names. A publication from the Transkei Forestry Department (Anon. 1975) listed about 199 Xhosa names for trees in Dwesa Forest. Sim (1900) commented that ‘with careful handling these native names are much more intelligible than some of the European’. While one cannot advocate the use of common names in official reports, it appears that in Transkei, where botanical names are generally unknown and where a well recognised terminology already exists by which the different species can be distinguished, it is worthwhile recording these names along with their botanical equivalents. With the inception of the research project of compiling a checklist of the flora of Transkei (Johnson 1985) identification of species proved difficult. To alleviate this problem and to assist future botanists working in Transkei, a quick reference system was established and is presented. After the present paper had been accepted for publica- tion, F. von Breitenbach (1989) published a list of Xhosa tree names as part 6 of his series Standard names of trees in southern Africa. Most of the names chosen by Von Breitenbach are also given here. The present paper, however, provides introductory paragraphs on Xhosa nomenclature and covers a wider range of names and orthographic variants. METHODS The data for this paper were collected over a period of five years (1981—85) and present a summary of an unpublished report (Johnson 1985) and the work of Johnson & Cawe (1987). During the course of this survey extensive collecting and field studies were undertaken. * Botany Department, University of the Western Cape, Private Bag X17, Bellville 7535. MS. received: 1988.10.24. Herbarium specimens are housed at KEI. To ensure that Xhosa names previously recorded did not go unnoticed, a number of references were consulted (Anon. 1975; Bigalke 1967; Broster & Brown 1981; De Lange 1963; Dornan 1932; Ferreira 1949; Friede 1953; Kropf 1915; Miller 1923; Mzamane 1945; Palmer & Pitman 1972; Rose 1979; Sim 1900, 1907; Shaw 1938 and Von Breitenbach & Von Breitenbach 1983). Emphasis was placed on the collecting and recording of the Xhosa names and to a lesser extent research was carried out into their semantics, the reason for their application. RESULTS AND DISCUSSION Various moulding forces can be detected when analysing the Xhosa names. Herd boys learn to recognise trees from which they use sticks, women must know which trees they use for firewood and men must know what is best used for building huts, kraals, sledges etc. Herbalists have to recognise trees for their medical usage. Of the 392 tree taxa recorded for Transkei (Johnson & Cawe 1987), 119 taxa were found to lack a Xhosa equivalent. The 284 tree taxa having Xhosa names, are listed in alphabetical order in the checklist. XHOSA NOMENCLATURE Miller (1923) was the first to point out the presence of a generic and ‘species’ name in Xhosa botany. For a better understanding of Xhosa nomenclature, a list of Xhosa plant names, more complete than is presently available, is required. However, analysis of the names collected for trees support the findings of Miller and is to a degree in accordance with the rules of the Code of Botanical Nomen- clature regarding rank concepts. Superficial examination of the Xhosa names shows words that express a generic concept. This is illustrated by the use of iGalagala or umGalagala which refers to Buxus macowanii and B. natalensis while umGxam and umSintsana are applied to several species of Schotia and Erythrina respectively. 148 Bothalia 20,1 (1990) The Xhosa name conveys something characteristic about a particular species and thus helps to distinguish species. Commiphora harveyi , umHlanguthi, is distinguished from C. woodii , imiNyele, by an outstanding feature of the species. The first named species is described as a soft wooded tree that is used as a hedge round kraals while the second species is found growing on the edge of forests. As can be expected, Xhosa nomenclature does not necessarily correspond to that of botanical nomenclature. Some plant names are undoubtedly homonyms, that are given to more than one species. This is best exemplified by the use of umBomvane which refers to Cassine peraqua , C. papillosa, C. crocea, Cassipourea gerrardii, Eugenia capensis, Memecylon grandiflorum, Olinia radiata and Pleurostylia capensis. Other names are polynomials, several names given to one plant species, as is the case in Baphia racemosa which is recognized as isiFithi or uTshiphu. The greater proportion of Xhosa names must have evolved at a time when these people lived close to nature. Older names tend to refer to habitat or habit characters, newer ones to social events and uses. The moulding forces for Xhosa names can therefore be divided into a number of categories. Outstanding features of the tree or its habitat UmNukane (Ocotea bullata). The wood possesses an unpleasant smell when freshly cut. It is this, the same character, that is recorded in the English common name ‘Stinkwood’. UmSimbithi ( Milletia grandis), the Xhosa name quoted by Sim (1900), refers to the extremely heavy, hard and strong character of the wood. The heartwood is dark brown and the sapwood is yellow. The wood is currently used and sought after for knobkieries. The vernacular name umGomgom refers to the beautiful tree fern ( Cyathea dregei), that grows up to about 3,5 m, of which the stem with no durable wood is described as being ‘empty’. The dead branches of Podocarpus falcatus and P. latifolius are a common feature in the immediate surroundings of these trees and are not collected for firewood, the Xhosa name umKhoba refers to the lightness of the dead wood. The common name for Acacia cajfra, umNyamanzi , refers to the fact that this tree is frequently found along rivers and streams. Sharing characteristics with an animal UmNukambiba (Clausena anisata), the strong un- pleasant smell of the leaves is embodied in the Xhosa name which means ‘the bad smell of the striped field mouse’. UbuHlungu benyoka ( Acokanthera oppositifolia) . Except for the seeds, all parts of this plant contain latex that is highly toxic. An infusion made from the powdered bark, wood or leaves was used as a poison (Watt & Breyer- Brandwijk 1932, 1962). This feature must have been known to the early Xhosas as the common name compares the poison of this plant to snakebite. UmNukambila (Hippobromus pauciflorus) . The Xhosa name compares the odour of this tree to that of a wild animal, such as a rock-rabbit or striped field mouse. The smell is not as pungent as that of umNukambiba (Clausena anisata). UmGobandlovu (Pterocelastrus tricuspidatus) . With its rounded leaf canopy, this plant forms a major constituent of dune vegetation. The Xhosa name fancies a resemblance of the growth form of this plant with the back of an elephant. Functional characters IsiThungwa ( Cryptocarya woodii ), this plant is common- ly used as the plait when constructing the frame of storage huts or when building a sledge. The Xhosa name refers to this functional, ‘plaiting’ feature. Social or historic events A large number of names relate to rituals or social functions, thereby implying that the tree has symbolic values or magical powers (De Jager 1963). Some of the names have an obscure meaning which might be of con- siderable historical interest. UmNgquza (Biaus natalensis) relates to a feast where the bride prepares for marriage. Intshilo ( Calpumia sepiaria). The powdered bark is used by the warriors to wash their bodies, while the green leaves are burnt, creating smoke. This ritual is believed to pro- tect the warriors against their enemies. Imbulanyathi (Osyris lanceolata), refers to a mystical animal with the power and strength of a buffalo, which can assume the shape of a human. UmLungumabele (Zanthoxylum capense). This much branched tree is characteristically armed with thorny protuberances. These protuberances, which may be as much as 60 mm in diameter, are jokingly compared with the breasts of European women. Medicinal properties Plants are commonly used for medicinal purposes (Mzamane 1945) and of the 392 tree taxa that were recorded, 212 are believed to have healing power. The com- mon name seldom refers to the medicinal properties of the tree but rather describes the condition of the patient. The rural tribal people are greatly influenced by their natural surroundings and each family has its own ubuLawu or iyeza lasekhaya (medicine of the home) which can include Hippobromus pauciflorus, Acalypha glabra and Burchellia bubalina. Nomina nuda names Finally, some plant names are apparently empty names, derived from nothing specific, left over from old lore or obsolete forgotten phrases. This is best exemplified by the Bothalia 20,2 (1990) 149 use of umCandathambo and umCane which refer to Allophylus africanus and Sclerocarya birrea subsp. cajfra respectively. The generic name for Protea, isiQwane and the common name isiQwane-sehlati for Rapanea tnelanophloeos are to a degree bare, except that they are distinguished by the fact that Rapanea tnelanophloeos is referred to as occurring in a forest. ACKNOWLEDGEMENTS The author wishes to express his appreciation to the inhabitants and informants of the various districts of Transkei who were willing to share their knowledge. I owe special thanks to Messrs W. Dutton, T.M. Sokutu and H. Kalala who provided valuable comments on the manuscript. A special word of thanks to Sharon Slingers and Belinda Feris for typing the manuscript. 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 150 Bothalia 20,2 (1990) Calpurnia aurea (Ail.) Benth., inDlole, iNdloli, umHlahlambedu, umSiphane, umSitshana Canthium inerme (L.f.) Kuntze, ubuChotshana, umNyushulube, iliTyelehlathi, umVuthwamini mundianum Cham. & Schlechtd., umNqayimbila. umSantulane, umSan- dulane, umSuntulane obovatum Klotzsch, umBombemfene, umBonemfene spinosum (Klotzsch) Kuntze. isiPhambatha, isiPambato Capparis sepiaria L., inTshilo, inTsihlo, uPasimani tomentosa Lam., umFihlo, imFihlo, umPasimani, inTsihlo Carissa bispinosa (L.) Desf. ex Brenan, isiBetha-nkunzi, isaBetha-inkunzi, isiBetha-umtunzi, isiBetha-nkunzi, umVusankunzi macrocarpa (Eckl.) A. DC., amaThungula, umThungula, umTungula, umTungulu Cassine aethiopica Thunb., umNqayi, utnGxube, umNguyi, umBomvane crocea (Thunb.) Kuntze, umMaqundeni, umThunyalele papillosa (Hochst.) Kuntze, umBomvane, iKhukhuzi, umMakhankatha peragua L., umBovane, umBomvane, iKhukhuzi. uMaphipha Cassinopsis ilicifolia (Hochst.) Kuntze. isiBethankunzi-sehlathi, iCegceya, iGcegceya tinifolia Harv., umHlaza Cassipourea flanaganii (Schinz) Alston, umMemezi gerrardii (Schinz) Alston, umBomvane. umHlamalala, umKatane, umKhathane, umMaphiphe, isiThushana gummiflua TuL, umBiko, umKanga, iQonga Catha edulis (Vahl) Forssk. ex End/., iGqwaka, umHIwazi Celtis africana Bunn, f, umVumvu gomphophylla Baker, umVumvu Chaetacme aristata Planch.. umKaboti, umKambothi. umKhovothi. um- Kombota, umKovoti Chrysanthemoides monilifera (L. ) Nor!.. iTholonja. ulwAinfiti Clausena anisata (Willd. ) Hook. f. ex Benth., isiFutu. umFuto, umNukam- biba, umNukambele, umNukambile, umTuto, isiTutu Clerodendrum glabrum E. Mey. , isiDwadwa. umKangazani. uNukisi- qaqa, umQangazani, umQwaqu. Qwaqwana Cliffortia nitidula (Engl.) R E. & T.C.E. Fries, umBele, uNwele Clutia pulchella L., uFiyo, umSipane, iQadi Cola natalensis Oliv, imBalonyathi, umNqayana. umTenenda. um- Thenenenda Combretum bracteosum (Hochst.) Brandis, uQota, uQoto calf rum (Eckl. & Zeyh.) Kuntze. umDubu, umDubi. erythrophyllum (Burch.) Sond., umDubu-wehlathi, umDubu kraussii Hochst., umDubu-wehlathi, umDuba-wehlathi Commiphora harveyi (Engl.) Engl., umHIanguthi, umHlunguthi wtxxlii Engl. , umHlunguthi. umHlunguthi weHlathi. imiNele, imiNyele, iNyele Cordia caffra Sond , umHIovu hlovu, umLovu-lovu. umNovu-novu Croton sylvaticus Hochst., umEgwaqane, untEgwaqanisa, untFeze, um- Fezu. umGwaqane, umGwaqanisa, uMagwacane, uMagwaqane Cryptocarya latilolia Sond., umGxaleba, umGxobothi, umNcatyana, umThungwa, umThunywa, umTungwa, umXaleba, iXoboti myrtifolia Stop], umGqebe, umNcatyana, umThungwa, isiThungwa, umThungwane, umThungwe, umTungwa, umXaleba, iXoboti woodii Engl., umNcatyana, umThungwa, isiThungwa, isiTungwa, um- Xaleba, iXoboti wyliei SiapJ, umNcatyana, iNqanyana, umXaleba, iXobothi Cunonia capensis L., umQwashube Curtisia dentata (Burnt J.) C.A. Sm., isiBande, umGuna. umGxima, umG- zina, umGxina, umHIebe, umLahleni Cussonia sphaerocephala Strey, umSenge spicata Thunb., umSenge Cyathea capensis (L.f.) J.E. Sin.. isiHihi, umKhoma-khoma dregei Kuntze, umGomgom, isiHihi Dais cotinifolia L., iNtozani, umTozane, inTozani Dalbergia armata E. Mey., luBobo, uBobo, uBobu, iTozane, umZungulu obovata E. Mey., uZungu, noZungu, umZungulu Deinbollia oblongifolia (E. Mey. ex Arn.) Radik., umAsibe, umAsibele, umBangabanga, uMasibele, umSibe Diospyros lycioides Desf, umBhongisa, umQokolo, umTshiwane, umTshicwane natalensis (Harv.) Brenan, umQokolo, umSitshana, umSitshane, uTshi- wane villosa (L.) De Winter, umBhongisa, umCandane, umTshekisane simii (Kuntze) De Winter, umBhongisa whyteana (Hiern) E White, umKhaza, umKhaze, iNtsazimane, um- Tenatena, umTenatene, inTsazimane, umTshikivane, umTshi- kivane Dombeya cymosa Harv, umSengathi, umZingathi tiliacea (Endl.) Planch., iTyibe, iTyibo Dovyalis caffra (Hook. f. & Harv) Hook, f, umQokolo, iNgali rhamnoides (Burch, ex DC.) Harv, umNyezana, umNyezane, um- Qaqoba Duvernoia adhatodoides E. Mey. ex Nees, isiBiko, iHlwehlwe, isiPheka Dracaena hookeriana K. Koch, umKhomakhoma, umKobakoba Dry petes arguta (Muell. Arg.) Hutch., umGudlamfene, umHlagela, iNyanzane gerrardii Hutch., umHIakela, umKhiwane natalensis (Harv) Hutch., umKhiwane Ehretia rigida (Thunb.) Druce, umBotshani, umHlele, umHIeli, iNqo- bogobana Ekebergia capensis Sparrm., umGwenya, umGwenye wezinja, um- Gwenyezinja, umNyamati, umQwehle, umQwengazinja, um- Qwenga-izinja Encephalartos sp., umPanga Englerodaphne subcordata (Meisn.) Engl., umHIoza, iNtozane, inTonzane Erythrina caffra Thunb., umSintsi humeana Spreng., umSintsana latissima E. Mey., umGqwane, umKuwane, umNqwane, umSintsana, umQwane lysistemon Hutch., umSinsi, umSintsana, umSintsi Erythroxylum emarginatum Thonn., uQadansube, uTimani pictum E. Mey. ex Sond., umDwibi, umGcamashe, iNtozani Euclea crispa (Thunb.) Guerke, umGwali, umHlungu, inTshekisane natalensis A. DC., iDungampunzi, umKhaza Eugenia capensis (Eckl. & Zeyh.) Harv & Sond., umBelvane, umBom- vane, isiDuli, isiDuli sehlathi, isiDul-sehlathi, umHela-hela, uMaphipha, iTolowa, umTsheka, umTsheqa Euphorbia grandidens Haw., umHIontlo, umHIonhlo triangularis Desf, umHIontlo, umHIonhlo tetragona Haw., umHIontlo Faurea macnaughtonii Phill., iCuba lethole, Cuba latole, iSafo, iSefi, isiFo, isiQwane saligna Harv, iCuba lethole Ficus bizanae Hutch. & Burtt Davy, umKovoti, umThombe craterostoma Warb. ex Mildbr. & Burr., umThombe sur Forssk., umKhiwane natalensis Hochst., uLuzi, umThombe, umTombi, umZombi Garcinia gerrardii Harv ex Sim, umBande, umBandi, umBinda, um- Bindi, umNebelele wehlathi Gardenia amoena Sims, umThongathi, umThongothi thunbergii L.f, umKancaza, umKangaza, umKangaze, umKhangazi, um Khangaza Grewia lasiocarpa E. Mey. ex Harv, uHlolo, umHIolo, uLuzi occidentalis L., umNqabaza, umVilani, umVileni Greyia flanaganii H. BoL, uSinya, umVilani-wamaxhegokazi sutherlandii Hook. & Harv, umBerebere, inDula, umDalu Halleria lucida L., umBinta, umBinza, umBiza, uBiza Harpephyllum caffrum Bemh., umGwenye, umGwenyezinja, umGwenye- wezinja, umGwenye-hangula Heteromorpha trifoliata (Wendl.) Eckl. & Zeyh., umBangandlatho, um- Bangandlele, umBanganolala, umBongondlala Heywoodia lucens Sims, umNebelele, umNebelele wehlathi Hibiscus tiliaceus L., miLolo, umLolwa Hippobromus pauciflorus (L.f.) Radik., ulwAtile, ulwAthile, umFazoneng- xolo, Futa, isiFuta, isiFutha, umHlwathile, uLathile, ubuLawu, uLwathile, umLungumabele, umNukambila, umNungumabele, Bothalia 20,2 (1990) 151 umNquma, ulWTile Homalium dentatum (Harv.) Warb., iGqabela, iGqabile, iKomanci, iKomanzi Ilex mitis (L.) Radik., umDuma, isiDumo, isiDuma Indigofera cylindrica DC., umSipane natalensis H. Bol., uSitshana Jubaeopsis caffra Becc. , iKomba, iNkomba, Kiggelaria africana/,., umDuma, umHIandlela, umHlandela, umHIin- zanyathi, umHIinziyati, umKhokonkho, umKokoko, umLuvuluvu, umVethi, umVeti Leucosidea sericea Eckl.& Zeyh., isiDwadwa, iNtshitshi , inTshitshi, um- Tyityi Linociera foveolata (E. Mey.) Knobl., umDlebe, umHIebe, umNqumaswele, um- Qumaswele, umQumaswile peglerae (C.H. Wright ) Gilg <6 Schellenb., umDlebe Macaranga capensis (Baill.) Benth., umBengele Maerua cafra (DC.) Pax, umQomoqomo racemulosa (DC.) Gilg & Ben., umBhonziso, umMphunziso, umPhun- ziso, umPunziso lanceolata Eorssk., iNtendekiwane, inTendekiwane. umTendekwane, inTendekiwane, umTentekwane, umTentekwani, isiThende Maytenus acuminata (L.f.) Loes., umNama, iNqayi, umNyama, umZungulwa, umZungulwo heterophylla (Eckl. & Zeyh.) N.K.B. Robson, umHIongwe, umKoko- hoha, umQaqoba, umQaqoha, umQuqoba mossambicensis (Klotzsch) Blakelock, inGqwanganeyehlathi, iNgqwan ganeyehlathi, nemorosa (Eckl. & Zeyh.) Marais, umHIangwe, peduncularis ( Sond .) Loes., umNgqi. umNqayi procumbens (L.f.) Loes., umPhonophono undata (Thunb.) Blakelock, umGorane, umGovane, umGurane, um- Kokane, umKokusa, umNqayimpofu, umQaqoba wehlathi Memecylon grandiflorum R. & A. Fernandes, umBande, umBomvane, umBondi Micrococca capensis (Baill.) Prain, uBubu Millettia grandis (E. Mey.) Skeels, umKunye, umSimbithi sutherlandii Harv., umGunye, umQunye Mimusops caffra E. Mey. ex A. DC., umHlungulu, umHlope, umNweba, um- Thunzi, umTunzi, umYoyiga, obovata Sond., amaSetole, amaSethole, amaSithole, amaSitole, um- Thunzi, umTunzi, umTunzi wehlathi Monanthotaxis caffra (Sond.) Verde., isiDwaba Noltea africana (L.) Reichb. f, umGlindi, umGlinidi, umKhuthula, um- Khuthuhla, maKutula, amaLuleka, iPalode, iPhalode, iPhalo elide, iYeza lesidiya Nuxia congesta R. Br. ex Fresen., umKhobeza, umKobess floribunda Benth., isiKali, isiKhali, inGqota, iNgqotha Ochna arborea Burch, ex DC., umTensema, inTensema, umTentsema, um- Thentsema, umTentsenda, umVithi natalitia (Meisn.) Walp., umBomvane-ncici, umBomvu, isiBomvu, um- Thentsama serrulata (Hochst.) Walp., iliTye Ocotea bullata (Burch.) Baill., umHlungulu, umNimbithi, umNimbiti, um- Nukane kenyensis (Chiov.) Robyns, umNukani Olea africana Mill., umNquma, umQuma capensis L., iGqwanxe, iGqwanci, iGqwanxe, iGqwangxe, iGxanci, iG- qwhanxi, iGxwanxi, umHIebe, umNquma-swile, umSinjane woodiana Knobl., umGqukunqa, umNqurunqa, umNququnya, um- Qoqunya Olinia emarginata Burn Davy, umNgona-lahlo, umNgene-lahle, uQudu radiata J. Hofmeyer & Phill., umBomvane, umPhanzi ventosa (L.) Cufod., umPhanzi ongenalahle, umNonono, oNqena-lahle Osyris lanceolata Hochst. & Steud., umBulunyathi, uMbulunyathi, iN- tekaza Oxyanthus speciosus DC., isiBika, umBinda, umBindi, iHlwehlwe, iM- pekane Pachystigma macrocalyx (Sond.) Robyns, inTzane, umVilo-wehlathi Pappea capensis Eckl. & Zeyh., iliLitye, iLitye Passerina rigida Wikst., iNwele Pavetta lanceolata Eckl., umDIesa, umGonogono, umHIeza, iLitoba, um- Ponyane, iliToba revoluta Hochst., umCilikishe, isiFufufu, usKolpati, umPonyane, iNtozane, uSkolpati, iriTozane Peddiea africana Harv., isiFufufu, Gcamche, uLuzi, iNtozane, um- Ponyane, inTozane Phoenix reclinata Jacq., iDama, uSundu Phyllanthus discoides (Baill.) Mull. Arg., umPanzite, umPhanzithe Pittosporum virdiflorum Sims, umGqengqwe, umKwenkwe Pleurostylia capensis (Turcz.) Oliv., umBomvane, umBomvani, uMaphipha, umMaphipha, umNgqangqa, umNqwanqwa, um- Thunyelo Portulacaria afra Jacq. , iGanishe, iGwanishe Podocarpus falcatus (Thunb.) R. Br. ex Mirb., umCeya, umKhoba, umKoba, um- Kolaya, umKoleya, umSonti henkelii Stapf ex Dallim. & Jacks., umSonti latifolius (Thunb.) R. Br. ex Mirb., umCeya, umGeya, umKhoba, um- Koba, umSonti Premna mooiensis (H.H.W. Pearson) Pieper, umCacabane, umKakabane, umTyetyembane, umTyetyembani, umTyintyambani Protea sp., isiAdlunge, isiQwane Protorhus longifolia (Bemh.) Engl., isiFuce, umHluthi, umHluthi- wezinja, umKhambathi, iKhubalo, umKomiso, umKumbati, um- Kumbat, umKupati, umLuthi, uZintlwa Prunus africana (Hook, f.) Kalkman, umKakase, iTywina-elikhulu Psychotria capensis (Eckl.) Vatke, umGonogono, unoGqiwathi-wehlathi, uGqiwathi-wehlathi, umGubhe, iHwehlwe, umSalamiya, iliTye- lehlathi Ptaeroxylon obliquum (Thunb.) Radik., umTali, umTati, umThathi, um- Thati, umThothe Pterocelastrus echinatus N.E. Br., iBolo tricuspidatus (Lam.) Sond., iBholo, iBolo, umGobandlovu, iTywina, uTwina Putterlickia pyracantha (L.) SzyszyL, umHIangwe, umKokhoba, iNtlang- wana, inThlangwana Quisqualis parviflora Gerr. ex Harv., uQhotho Rapanea melanophloeos (L.) Mez, isiQalaba-sehlathi, isiQalaba-hlathi, isiQwane, isiQwane-sehlathi, isiQwane-wehlathi, isiQwan- dashube, isiQwand-weshube Rauvolfia caffra Sond., umHlambamase, umHlambamasi, umlela, um- Jele, umJelo, umTelo, umThondisa, umThundisa, umTundisa Rawsonia lucida Harv. & Sond., umLongo, umNqagi-maphuthi, umNqayi-wephuthi, umNqayi-masende, umNqeyi-masende, um- Pitshi-wehlathi Rhamnus prinoides L'Herit., umGilindi, umGlindi, umNonye, umNyenye Rhizophora mucronata Lam., umHluma, umKhangathi, isiKhangathi, isiKhungathi Rhoicissus digita (L.f.) Gilg & Brandt, iSaqoni-esincinci rhomboidea (E. Mey. ex Harv.) Planch., umThwazi Rhus chirindensis Bak., umHlakothi, umHlokothi, umHlakotini, umHla- kothi-omkhulu, iNcakotsa, iNtlolokotshane, iNtlokotshane- enkulu, iNyangane, isiNyangane, iTlokotshane-enkulu fastigiata Eckl. & Zeyh., iNtlokolotshane-encinci, inNtlokotshane- encinane, iNtlokolotshane-yedobo, inNtolokotshane-enkulu, inTlolokotshane-yedobo laevigata L., umHlakoti, iNcakotsa, isiNyangani, umTlakoti lucida L., iNtlokotshane-ebomvu, inTlokotshane-bomvu , inTlakotshane- bomvu macowanii Schonl., umHlakothi natalensis Bernh., uGana, umGwele, uGwele, unGwele, iNgwele nebulosa Schonl., umHlokotshane Rinorea angustifolia (Thouars) Baill., uGudhla-mfeni, uGudla-mfene, ukuTyakwemfene Rothmannia capensis Thunb., iBolo, umGuba, umGubhe, isiGubhu, umGupa, um- Sugusu, umZukuza globosa (Hochst.) Keay, umGube, umSugusu Salix capensis Thunb. , umBenya, umBhenya, umGculube, umGcunuba, um- Gcunubi, umNceunch, umNculuba, umNcunube, umNcunubi, umNgcunube, umSwi, umThentsema mucronata Thunb., umBenya, umGcwimbe, umNgcwimbe, umNgcu- nube, umNgcunubi, umYezana Sapium ellipticum (Hochst.) Pax, umHongolo 152 Bothalia 20,2 (1990) Schcfflera'umbellifera (Sond.) Bail!., umBengele, umBumba, umGezisa, umKisiso, umRongo, umSengane, umSenge, umSenge-mbuzi, umSenge-wempunzi, umSengu-mbute Sclerochiton harveyanus Nees , umHlalani, isiThibothi Scholia afra (L.) Thunb. var. afra, umGxam brachypetala Sond., umGxam, umQonci latifolia Jacq., umGxam, umXamo Sclerocarya birrea (A. Rich.) Hochst. subsp. caffra (Sond.) Kokwaro., umCane Scolopia mundii (Eckl. & Zeyh.) Warb., umKokoko, isiNqandazembe, isiNqandi- zembe, umNqanqa, iQumza, zeyheri (Nees) Harv., iDumza-elinameva, umKhakboda, umKhakhoba, umQaqoba, iQumza-elinameva Scutia myrtina (Bunn f.) Kurz, isiBhingo, isiBhingwa, uBobo, umKapula, umKokwane, isiNcumncum, isiPhingo, isiPinga, isiPingo, isiPingwa, umQapuma, uQapuma, umQaphula, uQapula, umQaqoba-omnyama, umQokwane, umSondezo, iTywina Seemannaralia gerrardii (Seemann) Vig. , uMaweni Sideroxylon inerme L., umGqwashu, aMasethole, uMasethole, uMaze- tole, Mazetole uMazetoli, Mazetoli, umQwashu Smodingium argutum E. Mey. ex Sond., umThomvane, uThovane, um- Tomvane, umTovane, Tovane, uTovani, Tovani Solanum giganteum Jacq., iCuba-lasendle Strelitzia nicolai Regel & Koern., isiGceba, isiGude, iKamanga Strophanthus speciosus (Ward & Harv.) Reber, umKhukhumeza Strychnos decussata (Pappe) Gilg, umHlamalala, umKhangaza, umKhangele, um- Nama, iNama henningsii Gilg, umCaloti, umKaloti, umNonono, umNomo mitis S. Moore, umNdwibi, umNgqongqothi, umNgqungquthi, uMngqungquthi, umNqunquthi, umNqunquti spinosa Lam., umHlaba, umHlala, umHleli, iNhalo, inSala usambarensis Gilg, uNdlunye Suregada africana (Sond.) Kuntze, isiThubi Syzygium cordatum Hochst., umDoni, umJom, umJomi, umSibi, umSwi gerrardii (Harv. ex Hook, f.) Burn Davy, umBomva, umJome, umJome- wehlathi, umJomi, uManzani, isiTholothi Tarchonanthus camphoratus L., isiDuli, isiDuli-sehlathi, isiDuli-wehlathi, isiDuli- selinate, isiDuli-sasendle, isiDuli-selindle trilobus DC., isiDuli sehlathi Teclea gerrardii Verdoorn, umBoza natalensis (Sond.) Engl., umGani, umSane, umSenge-wesane, um- Senge-mzani, umSingomzane, umSingo, umZane Tecomaria capensis (Thunb.) Spach, iCakatha, iCakata, iDywadi, umKoto Trema orientalis (L.) Blume, uBatima, umBengele, Pak, umVangazi, umVumvu Tricalysia capensis (Meisn.) Sim, iNdulwane lanceolata (Sond.) Burn Davy, umBonyane, Dhlesa, umDlesa, umHleza, isiNyana, umPonyane, umPonyone Trichilia dregeana Sond. , umHlakele, umKhuhlu, umKuhlu, umKuhlwa Trichocladus crinitus (Thunb.) Pers., iTambo, iThambo ellipticus Eckl. & Zeyh. ex Walp., uGqonci, umVa wenyathi, umNqundu- wenyati Trimeria grandifolia (Hochst.) Warb., iliDhlebe-lendhlovu, iDlebe- lendlovu, inDlebe-yendlovu, inDlebezendlovu, iNdlebe-yendlovu, iNdlebezendlovu, iGqabela, uGqabela, iGqabi, uGqabile, Nqa- bane, umNqabane, iTabatane, iQabela Turraea floribunda Hochst., umaDlozana, umHIatholana, umLahlana, umVuma Umtiza listeriana Sim, umThiza Urera cameroonensis Wedd., umBabazane Uvaria caffra E. Mey. ex Sond., iDwaba Vangueria infausta Burch., umVilo, umViluka Vepris undulata (Thunb.) Verdoorn & C.A. Sm., umDlebe, umNgama- zwele, umNqumaswile, umSane, umZani, umZane Vitellariopsis marginata (N.E. Br.) Aubrev., umTunzi Voacanga thouarsii Roem. & Schult., umHlambamaas, umHIambamasi, umThofi, umTomvi, umThondisa, umThundisa, umTundisa Xymalos monospora (Harv) Baill., umBokoboko, iGxwanci, iKomanci, iKomanzi, uVete, uVethe, uVeti, umVeti, uVeto Zanthoxylum capense (Thunb.) Harv., umLungumabele, umNungumabele. um- Nungwane davyi (Verdoorn) Waterm., umLungu mabele, umNungu mabele, um- Nungwane Ziziphus mucronata Willd., umPafa, umPhafa Bothalia 20,2: 153-157 (1990) Nomenclature in Mesembryanthema (Aizoaceae): the generic names by Rappa and Camarrone H.E.K. HARTMANN* and V. BITTRICH* Keywords: Aizoaceae, Mesembryanthema, nectaries, Pentacoilanthus, Tetracoilanthus ABSTRACT A re-examination of the seven generic names used by Rappa and Camarrone reveals that three of them were validly published (Pentacoilanthus, Tetracoilanthus, Pteropentacoilanthus)', the first one is lectotypified here. The other four names are invalidly published. Their synonymy is given nevertheless where possible. UITTREKSEL 'n Herondersoek van die sewe genusname wat Rappa en Camarrone gebruik het, toon dat drie van hulle geldig gepubliseer is ( Pentacoilanthus , Tetracoilanthus, Pteropentacoilanthus ); die eerste een word hier gelektotipifiseer. Die ander vier name is ongeldig gepubliseer. Hul sinonimie word nietemin aangegee waar moontlik. INTRODUCTION In a survey of all genera of the Aizoaceae (including the Mesembryanthema) (H.E.K.H.) it was found that the genera described by Rappa and Camarrone (1953, 1955, 1960) have never been examined critically, and their validity and synonymy have never been established. Based on recent investigations in the group involved (V.B.), it is now possible to settle the nomenclatural uncertainties. Early investigations of flowers of Mesembryanthema led Rappa (1912) to the description and distinction of two different types of nectaries: lophomorphic ones, which are crest-shapkl (Figure 1A,B), and coilomorphic ones, which are conchiform or shell-shaped (Figure 2A, B). Nectaries may also be absent, and consequently Rappa (1912) names three groups: Lofomorfi, Anettari, and Coilomorfi, which were later classified as subfamilies (Rappa & Camarrone 1953, 1960). They lack Latin descriptions, however, and therefore do not conform with article 36 of the Interna- tional Code of Botanical Nomenclature (Greuter et al. 1988, abbreviated ICBN subsequently). The names are therefore not validly published. The fundamental distinction between lophomorphic and coilomorphic types of nectaries has been confirmed repeatedly (e.g. Ihlenfeldt 1960). Bittrich (1987) stresses the fact that the subfamily Mesembryanthemoideae (— Aptenioideae Schwant. ex Bittrich & H. Hartm.) is characterized by coilomorphic nectaries, and Bittrich & Hartmann (1988) mention them as a synapomorphic character for the subfamily. It is also worth noting that in Aptenioideae the number of carpels agrees with the number of perianth lobes as well as with the number of nectaries. N.E. Brown (1925) already used this character set in his keys to various genera. It must be remembered, though, that neither the number of carpels nor that of the nectaries is always constant within populations (e.g. Phyllobolus subg. Prenia , Bittrich 1987). Rappa & Camarrone (1953, 1955, 1960) use the number and shapes * Institut fur Allgemeine Botanik, Ohnhorststr. 18, D-2000 Hamburg 52, Federal Republic of Germany. MS. received: 1989.08.24. of nectaries for further subdivision of the subfamilies, but only in the Aptenioideae (Coilomorphioideae, Rappa FIGURE 1 .—Leipoldtia weigangiana (Dinter) Dinter & Schwant. ex H. Hartm. & Rust. (Ruschioideae) Hartmann 8476 (HBG). A. view of upper ovary surface, white scale bar = 1 mm; B, enlarged section of lophomorphic holonectary, the amorphous particles on the surface are remains of nectar, white scale bar = 0,1 mm. a, androecium, cut off; c, calyx, cut off; g, gynoecium raised subapically, with deep fissures between lobes; n, lophomorphic nectary which forms a complete ring (holonectary); s, styles, cut off. SEM micrograph: H. Golling. 154 Bothalia 20,2 (1990) FIGURE 2. — A, Brownanthus kuntzei (Schinz) Ihlenfeldt & Bittrich (Aptenioideae) Ihlenfeldt & Warda 12158 (HBG), view of ovary surface with five separate coilomorphic nectaries. B, Aridaria umbelliflora (Jacq.) Schwant., Jurgens 22261 (HBG), closer view of a single coilomorphic nectary, a, androecium, cut off; b, prominent bladder cells; c, calyx, cut off; cn, coilo- morphic nectary; g, gynoecium; o, ovary; s, styles, cut off. SEM micrograph: H. Golling. White scale bar = 1 mm. & Camarrone 1960) do they draw nomenclatural conse- quences, i.e. name genera. The guiding principle of Rappa & Camarrone (1953) is primarily the number of nectaries, complemented later (1955, 1960) by the absence or presence of valve wings. In the first step, two genera based on five, respectively four, nectaries are distinguished: Pentacoilanthus and Tetracoilanthus (Rappa & Camarrone 1953). Four more genera are added later after the importance of valve wings is recognized: Perapentacoilanthus (Rappa & Camarrone 1955), Pteropentacoilanthus, Pterotetracoilanthus , and Peratetracoilanthus (Rappa & Camarrone 1960). The names indicate that a certain pattern is followed predicting particular character combinations for the genera. This approach can be traced from the recognition of informal groupings (Rappa & Camarrone 1955) to the formal descriptions of taxa (Rappa & Camarrone 1960). The mechanical and even predictive procedure followed is well illustrated by the description of Pterotetracoilanthus , with a character combination which is theoretically possible but which the authors failed to find in any real plant (see 6. below). This unconventional approach is certainly one reason why the descriptions of Rappa & Camarrone have been neglected. Another reason is that, in most cases, the authors cite several species for their genera taken from various other genera previously described. Also, the new genera were not typified, making identification extremely difficult. Nevertheless, none of these shortcomings alone invalidate the names as such, and a detailed re-investigation was carried out to establish a reliable taxonomy for future use. Since each genus poses its own problems, the genera are discussed separately in chronological order. DISCUSSION 1. Pentacoilanthus Rappa & Camarrone (1953) The description conforms with the relevant requirements (art. 32—36 ICBN), and the name is therefore validly published. Four species are included, Mesembryanthemum aitonis, M. granulicaule, M. splendens and M. crystalli- num. Unfortunately no type species is mentioned and a lectotype must therefore be chosen. (1) Mesembryanthemum aitonis Jacq. can be excluded because, in contrast to the genus description of Rappa & Camarrone (1953) the nectaries of this species are not shell-shaped but tubular; these tubes may extend as dee- ply into the receptacle as the locules (Figure 3). It seems unlikely that Rappa & Camarrone studied longitudinal sec- tions and it is therefore assumed that they were unaware of the different nectary morphology of this species. Sur- face views do not permit the assessment of the depth of the nectary (Figure 2A, B). M. aitonis is therefore not available for selection as a lectotype. At present, the spe- cies is included in Mesembryanthemum L. subg. Mesem- bryanthemum (Bittrich 1987). FIGURE 3. — Mesembryanthemum aitonis Jacq. (Aizoaceae) Ihlenfeldt & Bittrich 13857 (HBG). Longitudinal section of flower. CA, prominent dorsal appendage of CL, calyx lobe; N, one of the five very deep coilomorphic nectaries. Bothalia 20.2 (1990) 155 The remaining three species do possess shell-shaped nectaries, and the choice between them is not easy. (2) Mesembryanthemum granulicaule Haw. (belonging to the genus Psilocaulon N.E. Br.) is a doubtful species. It is often synonymized with Psilocaulon articulation (Thunb.) N.E. Br. (the type species of the genus Psilocaulon) because of the rough papillose surface of leaves and stem; the latter species has white to pink flowers. The drawing of M. granulicaule, however, which is kept at Kew and is assumed to have been seen by Haworth (it would have to be chosen as the lectotype of the species in a revision), shows yellow flowers. The very few herbarium specimens with yellow flowers were all collected in a small area in the Orange Free State. They may well represent the true M. granulicaule as studied by Haworth. It seems extremely unlikely, though, that Rappa & Camarrone have investigated living or pickled material of this very rare species and herbarium material is useless for the investigation of nectaries. Furthermore, in species of Psilocaulon, four- and five-locular ovaries can occur within one species or even on one plant. Species of Psilocaulon can therefore not be associated unambiguously with either Pentacoilanthus or Tetracoilan- thus. Considering the care with which Rappa and Camarrone led their investigations, it seems highly unlikely that they saw species, or even specimens, with both 4- and 5-lobed ovaries. (3) Mesembryanthemum splendens |now included in Phyllobolus subg. Aridaria (N.E. Br. ) Bittrich, Bittrich (1987)| is a doubtful species because its identity is very difficult to determine. It belongs to a group of about twenty described species in the subgenus Aridaria most of which are synonymous. Due to the insufficient knowledge of the group, however, the exact number and delimitation of the species is still uncertain. It is scarcely possible to identify any species of this group with the help of literature only. The most useful characters are form and sculpture of seeds, rarely mentioned in descriptions. It is unlikely therefore, that Rappa & Camarrone investigated the true M. splendens , particularly since it is very rare in cultivation. Material kept under this name often belongs to the closely allied M. umbeUijlorus Jacq. It would seem highly unwise to choose this species as the lectotype. (4) Mesembryanthemum erystullinum L. is the only species figured by Rappa & Camarrone (1953). This species has not only been well known in Europe for a long time, it is also naturalized around the Mediterranean and is sometimes used as a vegetable. The figure mentioned above shows with great certainty a flower of M. crystal- linum, because the extremely large bladder cells of the epidermis of the receptacle are highly characteristic of this species. Therefore, and in contrast to the two other species with shell-shaped nectaries discussed above, it is most probable that M. crystallinum is the plant studied by Rappa & Camarrone. Consequently, M. crystallinum is chosen as the lectotype of the genus Pentacoilanthus. At the same time, M. crystallinum is the type species of the genus Cryophytum (N.E. Brown 1926). By this lectotypification, the name Pentacoilanthus becomes superfluous and is illegitimate (art. 63.1 ICBN). It should be stressed, though, that the choice was not influenced by this consequence but is based on agreement of characters with the descrip- tion and the probable identity of the original material. It should be mentioned further that the name Pentacoilan- thus would in any case be a synonym of another generic name, since all relevant species had been described earlier. The name Pentacoilanthus crystallinum , is not validly published because the basionym was not cited clearly and fully as required (art. 33.2 ICBN). At present, the species is considered to belong to Mesembryanthemum L. subg. Cryophytum (N.E. Br.) Bittrich (Bittrich 1987). 2. Tetracoilanthus Rappa & Camarrone (1953) The description conforms with the relevant requirements (art. 32—36 ICBN), and the name is therefore validly published. Since only one species is included, this is accepted as the indication of the type (art. 37.2 ICBN). Tetracoilanthus is based on the same type as the genus Aptenia N.E. Br. (1925), namely A. cordifolia (L.) Schwant. The name is therefore superfluous and conse- quently illegitimate (art. 63.1 ICBN). The combination T. cordifolius (L. f.) Rappa & Camar- rone is not validly published, because the basionym is not indicated clearly and fully (art. 33.2 ICBN). At present, the species is included in the genus Aptenia N.E. Br. (Bittrich 1987). 3. Perapentacoilanthus Rappa & Camarrone (1955) The description of this genus is almost identical with that of Pentacoilanthus Rappa & Camarrone 1953 (see 1. above) except for the addition of ‘valvis... quae, septis deficientibus, circa nudum axem consistunt’. This statement probably refers to the columella, the formation of which is very variable and changes from low to high with all intermediates. In Aptenioideae, the septa are always transformed into expanding keels in their upper parts, and consequently a remaining collumella will appear naked in the open capsule. These inconsistencies detract from the taxonomic value of the character. The two descriptions are consequently considered to be identical. Although Rappa and Camarrone did not state it explicitly, there is no doubt that they coined the name Perapentacoilanthus to replace Pentacoilanthus Rappa & Camarrone (1953). They had realised in the mean- time that the absence or presence of valve wings is an important character. Consequently they were now planning to group species and name genera not only according to the number of nectaries, but also according to the absence or presence of valve wings, as described in the introduction above. This is borne out by the following: a, under the heading ' Pentacoilanthus . Genere privo di umenoprosteci’ (genus deprived of valve wings), Rappa & Camarrone (1955) indicate that they intend to alter the diagnostic characters of Pentacoilanthus, which indeed they did later (Rappa & Camarrone 1960: 13); b, in the 1960 publication they include all the species originally mentioned under Pentacoilanthus Rappa & Camarrone (1953) under Perapentacoilanthus. This action was obviously initiated with the aim to name the genera with five, respectively four, coilomorphic nectaries in parallel fashion ( Pentacoilanthus matching Tetracoilanthus without valve wings; Pteropentacoilanthus—Pterotetra- coilanthus with free valve wings; Perapentacoilanthus— 156 Peratetracoilanthus with valve wings forming pockets; compare list in Rappa & Camarrone 1955: 10). To declare Perapentacoilanthus a superfluous and therefore illegitimate name (art. 63.1 ICBN) requires the definite inclusion of the holotype of the earlier synonym Pentacoilanthus. This is not possible literally, since Pen- tacoilanthus 1953 was published without the indication of a type, but the inclusion of all four species of the original Pentacoilanthus 1953 under Perapentacoilanthus (Rappa & Camarrone 1960) makes it clear that the type would be included, whichever species had been chosen. As a consequence, Pentacoilanthus 1953 and Perapentacoilan- thus 1955 must be considered synonymous, being names for the same genus. 4. Pentacoilanthus Rappa & Camarrone (1960) As stated above, the characterization of the genus differs from the Pentacoilanthus of 1953 in an important diagnostic character, namely the absence of valve wings, and an entirely different set of species is accordingly assigned to the genus. This name therefore represents a later homonym of Pentacoilanthus Rappa & Camarrone 1953 (although no type is mentioned in either genus) and is consequently illegitimate (art. 64.1 ICBN). Further- more, the lack of citation of a type at this date makes the name an invalidly published one (art. 37.1 ICBN ). All species included in Pentacoilanthus 1960 were taken from the genus Sceletium N.E. Br., and they are at present assigned to Phyllobolus N.E. Br. subg. Sceletium (N.E. Br.) Bittrich (Bittrich 1987). 5. Pteropentacoilanthus Rappa & Camarrone (1960) Only one species is cited as an example with the description of the genus and this validates the name (art. 37.2 ICBN). The genus is based on the same species as Halenbergia Dinter, namely H. hypertrophica (Dinter) Dinter. The name Pteropentacoilanthus is therefore a superfluous name and illegitimate (art. 63.1 ICBN). The combination P hypertrophicum (Dinter) Rappa & Camarrone is neither validly published, because the basionym is not cited fully (art. 33.2 ICBN), nor legitimate, since it includes the type of an earlier named taxon (art. 63.1 ICBN). At present, the species is considered to belong to Mesembryanthemum L. subg. Opophytum (N.E. Br.) Bittrich (Bittrich 1987). 6. Pterotetracoilanthus Rappa & Camarrone (1960) No type is named with the description, and the name is therefore invalid (art. 37.1 ICBN); no species are mentioned at all. 7. Peratetracoilanthus Rappa & Camarrone (1960) Eight species are cited with the description, but no type is chosen, and the name is therefore invalidly published (art. 37.1 ICBN). The new combinations are invalid as well because no basionyms are given (art. 33.2 ICBN), and Bothalia 20,2 (1990) the name of the genus is invalidly published (art. 43.1 ICBN). TAXONOMY 1. Pentacoilanthus Rappa & Camarrone in Lavori dell’Istituto Botanico e Giardino Coloniale Palermo 14: 32 (1953), nom. illeg. Lectotype species: Mesembryanthe- mum crystallinum L.: 480 (1753) (here designated). Type: Dillenius: 231, t. 211 (1732). Cryophytum N.E. Br.: 412 (1925). Mesembryanthemum L. subg. Cryophytum (N.E. Br.) Bittrich: 72 (1987). Type: C. crystallinum (L.) Schwant., fide N.E. Brown: 245 (1926). Perapentacoilanthus Rappa & Camarrone 1960, nom. illeg., nom. invalid (see 3 below). 2. Tetracoilanthus Rappa & Camarrone in Lavori dell’Istituto Botanico e Giardino Coloniale Palermo 14: 34 (1953), nom. illeg. Type: Mesembryanthemum cordifolium L. f. : 260 (1782). Holotype: Thunberg s.n. 1773 in Herb. Montin (S!) [= Aptenia cordifolia (L. f.) Schwant.: 69 (1928)]. Note on the holotype: there is no doubt that the sheet cited and seen is the holotype since the description on the back refers to the original publication. The specimen agrees well with the description. The holotype was already recognized by both Norlindh and Glen in 1976, but there seems to be no published reference to this. In any case, the existence of the holotype supersedes the selection of a neotype by Preston & Sell (1988). Aptenia N.E. Br.: 412 (1925). Litocarpus L. Bol.: t. 261 (1927), nom. illeg. 3. Perapentacoilanthus Rappa & Camarrone in Lavori dell’Istituto Botanico e Giardino Coloniale Palermo 15: 6 (1955), nom. illeg., nom. invalid. No type given (see 1 above). 4. Pentacoilanthus Rappa & Camarrone in Lavori dell’Istituto Botanico e Giardino Coloniale Palermo 18: 13 (1960), nom. illeg., nom. invalid. No type given (see 1 above). Sceletium N.E. Br. : 412 (1925). Phyllobolus N.E. Br. subg. Sceletium (N.E. Br.) Bittrich: 75 (1987). 5. Pteropentacoilanthus Rappa & Camarrone in Lavori dell’Istituto Botanico e Giardino Coloniale Palermo 18: 14 (1960), nom. illeg. Type species: Mesembryanthemum hypertrophicum Dinter [= Halenbergia hypertrophica (Dinter) Dinter: 200 (1937)]. Type: Dinter 3875 (B!). Halenbergia Dinter: 200 (1937). Opophytum N.E. Br.: 412 (1925). Mesembryanthemum L. subg. Opophytum (N.E. Br.) Bittrich: 73 (1987). 6. Pterotetracoilanthus Rappa & Camarrone in Lavori dell’Istituto Botanico e Giardino Coloniale Palermo 18: 14 (1960), nom. invalid. No type given. No species mentioned. 7. Peratetracoilanthus Rappa & Camarrone in Lavori dell’Istituto Botanico e Giardino Coloniale Palermo 18: 14—15 (1960), nom. invalid. No type given. Bothalia 20,2 (1990) 157 ACKNOWLEDGEMENTS The basic field work on Mesembryanthema was supported by the German Science foundation and the National Botanic Gardens of South Africa; to both we are most grateful. We would like to thank H. Golling for help with SEM micrographs, and Prof. P. Hiepko, Berlin, for critically reading the manuscript. REFERENCES BITTRICH, V. 1987. Untersuchungen zu Merkmalsbestand, Gliederung und Abgrenzung der Unterfamilie Mesembryanthemoideae (Mesembryanthemaceae Fenzl). Mitteilungen des lnstituts fur Allgemeine Botanik Hamburg 21: 5-116. BITTRJCH, V, & HARTMANN, H E.K. 1988. The Aizoaceae — a new approach. Botanical Journal of the Linnean Society 97: 239 - 254. BOLUS, L. 1927. Litocarpus cordifolius. The Flowering Plants of South Africa 7: t. 261. BROWN, N.E. 1925. Aptenia, Cryophxtum. Opophytum, Sceletium. Gardeners' Chronicle ser. 3,78: 412. BROWN, N.E. 1926. Cryophxtum. In E.P Phillips, The genera of South African flowering plants: 245. Cape Town. DILLENIUS, J.J 1732. Hortus elthamensis: 231, t. 211. London D1NTER, K, 1923. Succulentenforschung in Sudwcstafrika Feddes Repertorium, Beiheft 23: 63. Berlin. DINTER, K. 1937. Halenbergia hypertrophica. Kakteen und andere Sukkulenten: 200. Berlin. GREUTER, W, et al. 1988. International Code of Botanical Nomenclature. Regnum vegetabile 118. Konigstein. IHLENFELDT, H.-D. 1960. Entwicklungsgeschichtliche, morphologische und systematische Untersuchungen an Mesembryanthemen Feddes Repertorium 63: 1 — 104 LINNAEUS, C. 1753. Species plantarum. Stockholm. LINNAEUS, C. 1782. Supplementum plantarum Brunswick. PHILLIPS, E P 1926. The genera of South African flowering plants. 1st edn. Pretoria. PRESTON, C D. & SELL, P.D. 1988. The Aizoaceae naturalized in the British Isles. Watsonia 17: 221. RAPPA, F. 1912. Per una classificazione naturaledei Mesembrianthemi. Bulletin Reale Orto Botanico Giardino Colonial e Palermo II: 21-36. RAPPA, F. & CAMARRONE, V. 1953. Lavori dell' Istituto Botanico e Giardino Coloniale Palermo 14: 1 — 39. RAPPA, F. & CAMARRONE, V. 1955. Lavori dell' Istituto Botanico e Giardino Coloniale Palermo 15: 1 — 16. RAPPA, F. & CAMARRONE, V. I960. Lavori dell' Istituto Botanico e Giardino Coloniale Palermo 18: 1—24. SCHWANTES, G. 1928. Mesembriaceen unserer Kulturen in neuer Benennung. Gartenflora 77: 69. Bothalia 20,2: 159-165 (1990) A checklist of the plants of the Karkloof Forest, Natal midlands J.O. WIRMINGHAUS* Keywords: checklist, Karkloof. forest. Natal ABSTRACT A checklist of the plants of the Karkloof Forest is presented. Of the 400 taxa recorded, seven are bryophytes. 69 are pteridophytes. four are gymnosperms and 320 are angiosperms. At least 17 of the plants recorded are localized or uncommon in Natal. Two of these are endemic to the area. UITTREKSEL 'n Kontrolelys van plante van die Karkloofwoud word verskaf. Van die 400 taksons wat aangeteken is. is sewe briofiete, 69 pteridofiete, vier gimnosperme en 320 angiosperme. Minstens 17 van die aangetekende plante is tot sekere lokaliteite beperk of is seldsaam in Natal. Twee hiervan is endemies in die gebied. INTRODUCTION Rycroft (1941) carried out the first detailed study of the Karkloof Forest’s vegetation and provides a preliminary checklist of the plants. Taylor (1961) gives a brief account of the plant communities of the Karkloof Forest although his paper is directed at its conservation. More recently Moll (1976) included the Karkloof Forest in his vegetation survey of the Three Rivers region. Natal. Other than the above, little has been published on the area, despite the fact that it is one of the largest remaining patches of mistbelt forest in Natal. The absence in regional herbaria of records of species commonly seen in the area highlighted the need for a comprehensive ch ecklist of the forest plants. STUDY AREA AND METHODS The Karkloof Forest lies in the Natal midlands and covers an estimated area of 6 000 ha (Cooper 1985). It extends in length for 40 km, from about 22 km north of Howick (29°17’S. 30°09’E) to 13 km northeast of New Hanover (29°14’S, 30°28’E), and comprises a number of forest patches restricted mainly to the steep eastward side of the Karkloof Range. The forest ranges in altitude from 800 m near the Karkloof and Umgeni River confluence up to I 700 m on the upper slopes of Mt Gilboa. Most of the forests in the midlands region, including the Karkloof. fall under Acocks’s (1988) Veld Type 5 (’Ngongoni Veld). In addition to mist the forest receives a mean annual rainfall of about 1 600 mm (1 300—2 200 mm) (Rycroft 1944; Oatley 1978; pers. obs.) falling mostly in summer. Predominant winds in the summer months are southeasterly but in winter, as along most of the escarpment, northwesterly berg winds predominate. Rycroft (1944) attributes the stunted growth of forest trees in the northwestern Karkloof and the deaths of numerous young forest trees each year to these latter hot, dry winds. * Dept, of Zoology & Entomology, University of Natal. RO. Box 375, Pietermaritzburg 3200. MS. received: 1990.02.20. Little is published of temperatures within the forest but Oatley (1978) recorded some data over a two year period. From his data the mean annual maximum temperature is 18,0°C (range 15,2— 20,9°C) measured in February, and the mean minimum is 8,4°C (range 4,6— 12,2°C) in July. Most of the collecting for this survey was done near the Farms Mbona and Chard which form part of the Blinkwater forest patch. Other sites in the Karkloof were also visited during the survey for the purpose of completeness. All specimens are lodged at NU. In addition to my specimens, any records of plants from other collectors, which were found in NU, are included. Of these, E.J. Moll has possibly contributed the most. Much of his collecting was carried out in the forest near the Farms The Start. Benvie and Ehlatini. Even though every effort was made to make the checklist as comprehensive as possible, there will be a number of species that have been overlooked. Bryophytes were only occasionally collected during this survey, and field observations indicate that many more taxa occur in the forest than this checklist suggests. RESULTS AND DISCUSSION Rycroft (1941) recorded 246 species in his preliminary checklist of the forest plants but did not mention any voucher specimens. He lists 45 species of pterido- phytes, four gymnosperms, 25 monocotyledons and 172 dicotyledons. Of the 400 taxa (266 genera, 115 families) recorded in the Karkloof Forest during this survey, seven (1,8%) are bryophytes, 69 (17,3%) are pteridophytes, four (1.0%) are gymnosperms and 320 (80,0%) are angiosperms. Of the angiosperms, 67 (16,8%) are monocotyledons and 253 (63,3%) are dicotyledons. Plant families containing more than one percent of the total number of species are listed in order of numerical importance in Table 1. The genera of plants with four or more (> 1%) species are Asplenium (11 spp.), Senecio, Solatium (7 spp.). Blech- num, Cheilanthes, Plectranthus , Streptocarpus (5 spp.), Canthium, Lycopodium, Maytenus, Protasparagus and 160 Bothalia 20,2 (1990) TABLE 1. — Synopsis of the plant families of the Karkloof Forest with four or more species (> 1%), listed in order of numerical importance together with the number of genera in each family 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 Bothalia 20,2 (1990) 163 PAPAVERACEAE Papaver aculeatum Thunb .*, 1119, 6 BRASSICACEAE Cardamine africana L. , 907, 6 CRASSULACEAE Crassula inandensis Schonl. & Bak.f, 1134, 3, 6 pellucida L. subsp. alsinoides (Hook.f) Toelken, 757, 6 sp. aff. alba Forssk. , 1174, 1 ESCALLONIACEAE Choristylis rhamnoides Harv. , 761, 4, 6 PITTOSPORACEAE Pittosporum viridiflorum Sims, Moll 3544, 1, 5 ROSACEAE Rubus cuneifolius Pursh*, 1073, 1, 2 rosifolius J.E. Sm* 1058, 1, 2 Duchesnea indica (Andr.) Focke*, 746, 6 Alchemilla kiwuensis Engl., 788, 6 Leucosidea sericea Eckl. & Zeyh. , 977, 1 Cliffortia linearifol ia Eckl. & Zeyh., 1054, 1 nitidula (Engl.) R E. & Th. Fries Jr, 982 , 6 Prunus africana (Hook. f. ) Kalkm. , 1158, 5 CONNARACEAE Cnestis natalensis (Hochst.) Planch. & Sond.. 1001, 5 FABACEAE Acacia ataxacantha DC. , Getliffe 219, 1 mearnsii De Wild 3, 981, I, 2, 6 melanoxylon R. Br*, Ross 2088, 1, 2 Calpurnia aurea (Ait.) Benth., 1040, 2, 5, 6 Crotalaria capensis Jacq., 983, 6 Argyrolobium tomentosum (Andr.) Druce, 821, 1, 2, 3, 6 Indigofera natalensis H Bol., 922, 4 Psoralea pinnata L. , 925, 1 Desmodium repandum (Vahl) DC., 612, I, 2, 6 Dalbergia obovata E. Mey. , 1041, 5 Dumasia villosa DC. var. villosa, 754, 2, 6 GERANIACEAE Geranium flanaganii Knuth, Hilliard & Burn 13485, 2 natalense Hilliard & Burn, 1051, 1 schlechteri Knuth, Hilliard & Burn 13489, 1, 2 OXALIDACEAE Oxalis corniculata L.*, 758, 6 obliquifolia Steud. ex Rich., 1052, 1 RUTACEAE Zanthoxylum davyi (Verdoorn) Waterm. , 1002, 5 Calodendrum capense (L. f.) Thunb., 997, 5 Oricia bachmannii (Engl.) Verdoorn, 1090, 4 Vepris lanceolata (Lam.) G. Don, 1078, 5 Clausena anisata ( Willd .) Hook. f. ex Benth., 1087, 4, 5 PTAEROXYLACEAE Ptaeroxylon obliquum (Thunb.) Radik., 979, 5 MELIACEAE Ekebergia capensis Sparrm. , 1027, 5 POLYGALACEAE Polygala confusa MacOwan, 1024, 1, 2 , 6 EUPHORBIACEAE Andrachne ovalis (Sond.) Muell. Arg. , Bayer 687, 4 Drypetes gerrardii Hutch., Bayer 1414, 1, 4 natalensis (Harv.) Hutch., 1076, 4 Micrococca capensis (Bail!. I Pram, 593, 4, 6 Clutia katharinae Pax, 1039, 1 pulchella L. var. franksiae Prain, 803, 2, 4 Suregada procera (Prain) Croiz. , Smook 655, 4 Excoecaria simii (Kuntze) Pax, Moll 3535, 4 Euphorbia dumosa E. Mey. ex Boiss., 1053, 1, 2 epicyparissias E. Mey. ex Boiss., Hilliard & Burn 10141, I, 2 kraussiana Bernh. var. kraussiana, 626, 1, 2, 6 CALL1TRICHACEAE Callitriche sp. cf. compressa N.E. Br. , 1049, 6 ANACARDIACEAE Harpephyllum caffrum Bernh. ex Krauss, 1148, 5 Protorhus longifolia (Bernh.) Engl., 964, 1 Rhus chirindensis Bak. f, 1069, 5 dentata Thunb. , 1056, 1 pyroides Burch, var. gracilis (Engl.) Burtt Davy, 1059, 1 AQUIFOL1ACEAE Ilex mitis (L.) Radik, var. mitis, 1067, 5, 6 CELASTRACEAE Maytenus acuminata (L. f.) Loes., 986, 4 heterophylla (Eckl. & Zeyh.) N.K.B. Robson, 1028, 1 mossambicensis (Klotzsch) Blakelock var. mossambicensis, 560, 4 undata (Thunb.) Blakelock, 1029, 1 Cassine papillosa (Hochst.) Kuntze. 1004, 4 tetragona (L. f. ) Loes. , 590, 5, 6 ICACINACEAE Apodytes dimidiata E. Mey. ex Arn. subsp. dimidiata, 591, 5 SAPINDACEAE Allophylus dregeanus (Sond.) De Winter. 585, 2, 4 melanocarpus (Sond.) Radik., 872, 5 MELIANTHACEAE Bersama tysoniana Oliv., 1139, 1 GREY1ACEAE Greyia sutherlandii Hook. & Harv, 1036, 1 BALSAM IN ACEAE Impatiens hochstetteri Warb. subsp. hochstetteri, 540, 3 RHAMNACEAE Scutia myrtina (Burm.f.) Kurz, 1066. 5 Rhamnus prinoides L'Herit., 927, I, 4 Phylica paniculata Willd., 1037, I Helinus integrifolius (Lam.) Kuntze, R. McMahon s.n., 5 VITACEAE Rhoicissus rhomboidea (E. Mey. ex Harv. ) Planch. , 974, 5 tomentosa (Lam. ) Wild & Drum. , 1074, 5 Cyphostemma sp. c.f. cirrhosum (Thunb.) Desc. ex Wild & Drum., 764, 4 TILIACEAE Sparrmannia ricinocarpa (Eckl. & Zeyh.) Kuntze, 1155, 1 Grewia occidentals L. , 1122, 5 Triumfetta annua L. , 613, 6 MALVACEAE Abutilon sonneratianum (Cav.) Sweet, 807, 2 Hibiscus pedunculatus L. f, 592, 1, 4 STERCULI ACEAE Dombeya tiliacea (Endl.) Planch., 938, 1 OCHNACEAE Ochna arborea Burch, ex DC. , 968, 5 natalitia (Meisn.) Walp. , 978, 5 serrulata (Hochst.) Walp., 559, 4 164 Bothalia 20,2 (1990) VIOLACEAE Rinorea angustifolia (Thouars) Baill., 1032 , 4 FLACOURTIACEAE Rawsonia lucida flarv. & Sond. , 1167, 4 Kiggelaria africana L. , Moll 3492, 5 Scolopia zeyheri (Nees) Harv. , Wells 1870, 5 Trimeria grandifolia (Hochsi.) Warb. , Moll 3493, 5, 6 Dovyalis lucida Sim, 988, 4 rhamnoides (Burcli. ex DC.) Harv., 987, 4 Casearia gladiiformis Mast., 582 , 5 PASSIFLORACEAE Passiflora edulis Sims*, 1073, 1 ACHARIACEAE Ceratiosicyos laevis (Dumb.) Meeuse, Hilliard 5057, 4, 6 BEGONIACEAE Begonia geranioides Hook. f. , Hilliard 2583, 3, 6 sutherlandii Hook, f, 723, 6 OLINIACEAE Olinia emarginata Burn Davy, Moll 3371. 5 THYMELAEACEAE Peddiea africana Harv., 994, 4 Gnidia pulchella Meisn. , 1057, I Englerodaphne pilosa Burn Davy, 600, 4, 6 Dais cotinifolia L. , 587, I, 2, 4 RHIZOPHORACEAE Cassipourea gerrardii (Schinz) Alston, 1070, 4 gummiflua Tut. var. verticillata (N.E. Br.) J. Lewis, 962, 1 COMBRETACEAE Combretum edwardsn Exell, 1 144, 5 kraussu Hochst., 1065, 5 Quisqualis parviflora Gerr. ex Harv., 998, 5 MYRTACEAE Eugenia zuluensis Duernmer, 1068, 5 Syzygium gerrardii Harv. ex Hook, f.) Burn Davy, Moll 3539, 5 Eucalyptus grandis Hill ex Maid 7, 980. 1, 2 ONAGRACEAE Oenothera erythrosepala Borbas*. 923, I ARAL1ACEAE Schefflera umbellifera (Sond.) Baill., Sim 19385, 5 Cussonia sphaerocephala Si rev, Moll 3436, 5 APIACEAE Hydrocotyle americana L.*, 818, 2, 6 Centella asiatica (L. ) Urb. , 620, 3, 6 Sanicula elata Buch.-Ham. , 652, 3, 6 Conium fontanum Hilliard & Burn var. silvaticum Hilliard & Burn, 773, 6 Heteromorpha trifoliata (Wendt.) Eckl.