156 Bothalia 33,2(2003)

BRENAN, J.RM. 1949. Check-lists of the forest trees and shrubs of the RADCLIFFE-SMITH, A. 1996. Euphorbiaceae. In G.V. Pope, Flora British Empire 5. Tanganyika Territory. Imperial Forestry zambesiaca 9,4. Royal Botanic Gardens, Kew. Institute, Oxford. RENDLE, A.B.. BAKER, E.G., MOORE. S. & GEPP, A. 1911. A con­ COATES PALGRAVE, M. 2002. Keith Coates Palgrave Trees of south­ tribution to our knowledge of the flora of Gazaland; being an ern Africa, edn 3. Struik, Cape Town. account of the collections made by C.F.M. Swynnerton. Journal DE CANDOLLE, A.P. 1866. Prodromus systematis naturalis regni of the Linnean Society, Botany 40: 1-245. vegetabilis, vol. 15. Treuttel & Wiirtz, Paris. THULIN, M. 1993. Excoecaria. Flora of Somalia 1: 305-307. Royal MULLER ARGOVIENSIS, J. 1866. Euphorbiaceae (in part). In A.P. de Botanic Gardens, Kew. Candolle, Prodromus 15: 1219. Treuttel & Wiirtz, Paris. PAX, F.A. 1890. Euphorbiaceae. In A. Engler & K. Prantl, Die natiir- lichen Pflanzenfamilien 3,5:1-119. Engelmann, Leipzig. J.E. BURROWS*, S.M. BURROWS*, P. LOFFLER+ & L. LOFFLER+ PRAIN. D. 1913. Euphorbiaceae. In W.T. Thiselton-Dyer, Flora of tropical Africa 6,1. Reeve, London. * Buffelskloof Herbarium, P.O. Box 710, 1120 Lydenburg, South Africa. RADCLIFFE-SMITH. A. 1987. Euphorbiaceae (part 1). In R.M. Polhill. + P.O. Box 764. Mbabane H100, Swaziland. Flora of tropical East Africa. Balkema. Rotterdam. Boston. MS. received: 2003-01-28.

LYCOPERDACEAE-GASTEROMYCETES

CALVATIA SECT. MACROCALVATIA REDEFINED AND A NEW COMBINATION IN THE

INTRODUCTION neither accommodate all of the species assigned to by Kreisel (1989) nor all of those assigned to Whilst accepting Kreisel’s (1992) incorporation of the Calvatia sect. Handkea by Lange (1993). Instead of genus Langermannia Rostk. into Calvatia Fr„ we, like establishing yet another new section to accommodate the others (e.g. Calonge & Martin 1990; Demoulin 1993; excluded species, however, we here emend the circum­ Lange 1993; Calonge 1998), are not convinced of the scription of Calvatia sect. Macrocalvatia to accommo­ generic status of his segregate genus Handkea (Kreisel date also those Handkea species currently excluded by 1989). Kreisel (1989) referred to Handkea those members the original diagnosis. previously placed in Calvatia, but characterized by ‘asep- tate, slit-like, pitted capillitium', and included species with sterile bases (compact or cellular) and without. Emended sectional description

For the same reasons as expressed in Lange (1993), and Calvatia sect. Macrocalvatia Kreisel in Feddes from conclusions based on our own comparative morpho­ Repertorium 64: 163 (1962), emend. J.C.Coetzee, Eicker logical and anatomical studies of the Lycoperdaceae of & A.E.van W\k. Type species: Calvatia excipuliformis southern Africa (to be reported elsewhere), we agree with (Scop.: Pers.) Perdeck. Lange (1993) in treating Handkea as a section of Calvatia. To that purpose Lange (1993) relegated the genus Handkea Kreisel: 282 (1989); Calvatia sect. Handkea Handkea Kreisel to sectional rank under the name (Kreisel) M.Lange: 143 (1993), nom. inval. Calvatia sect. Handkea (Kreisel) M.Lange. Lange's sec­ tional name was not validly published, however, since the Fruit bodies with sterile base or not. Capillitial septa requirements of ICBN Art. 33.3 (Greuter et al. 2000) were extremely rare to essentially absent, easily missed; capil­ not met. Moreover, Lange apparently overlooked the fact litial threads fragmenting at septa or more commonly by that, long before the establishment of the genus Handkea, irregular rupturing of walls; capillitial walls fragile, with Kreisel (1962) had established Calvatia sect. Macro- small perforations and conspicuous slit-like fissures, calvatia Kreisel, to which he assigned Calvatia excipuli- often between perforations. formis (Scop.: Pers.) Perdeck [= (Scop.: Pers.) Kreisel] and Calvatia utriformis (Bull.: New' combination in Calvatia Pers.) Jaap [= (Bull.: Pers.) Kreisel], the latter being the type species of the genus Handkea. The inclusion of Handkea into Calvatia necessitates Calvatia sect. Macrocalvatia is thus also pertinently cited the following new combination (description based on dry as an earlier synonym of Handkea by Kreisel (1989). herbarium material; colour codes and colour terms fol­ Therefore, if the latter two species, (and a number of other low Komerup & Wanscher 1981): species of Handkea) are to be accommodated in Calvatia, there already exists a section for them, namely Calvatia Calvatia capensis (LloydI J.C.Coetzee, Eicker & sect. Macrocalvatia. A.E.van W\k, comb. nov. Type: South Africa, Stellenbosch. A.V. Duthie 403 (Lloyd Myc. Coll. 7567 in BPI 706162, A problem arises, however. The original diagnosis for holo.!; Herb. v.d. Byl in PREM 31472, iso.!). Calvatia sect. Macrocalvatia, as defined by Kreisel (1962: 163), is unambiguous: 'Subgleba distincte cel- Lanopila capensis Lloyd in Mycological Writings 7: 1177 (1923); losa. Capillitium non septatum.’ This diagnosis clearly Verwoerd: 25 (1925); Bottomley: 579 (1948). provides for species with fruit bodies characterized by cellular sterile bases only and excludes those with com­ Handkea capensis (Lloyd) Kreisel & G.Moreno: 84 (1996) pact bases as well as those without sterile bases. Calvatia Illustrations: Lloyd: t. 230. fig. 2352 (1923); Kreisel & G.Moreno: sect. Macrocalvatia in the sense of Kreisel can therefore 86. figs 1-7(1996). Bothalia 33,2 (2003) 157

Basidiocarp epigeous, with prominent rooting base, ± Duthie from Stellenbosch. South Africa. In his original globose (sensu Verwoerd 1925: Bottomley 1948), holotype description Lloyd made no mention of the subgleba, a appearing to have been ± 30 mm wide and 25-30 mm high, feature first described from the other half (the isotype) by dehiscing by irregular fragmentation of peridium. Peridium Verwoerd (1925). In a footnote to his often overlooked very thin, 0.09-0.2 mm. rigid but brittle and extremely work (in Afrikaans) on South African Gasteromycetes, fragile, differentiated into an exo- and endoperidium (two Verwoerd (1925) had the following to say regarding this layers not discernible from type material, however). (our translation): ‘According to Lloyd ... it does Exoperidium dark brown with reddish tinge (7F6) to paler not have a sterile base. The half in my possession, how­ and concolorous with endoperidium, fugacious, remaining ever. clearly shows one. Lloyd described it from the as tiny, weft-like mycelial patches on some specimens. other half.” The half described by Lloyd does have a ster­ Endoperidium consisting of a thin, amorphous crust over­ ile base, however, and it is difficult to understand how he laying a slightly thicker hyphal layer, surface colour vari­ could have overlooked this structure. Contrary to the ous shades of pale to darker brown [± 5C5 (topaz) and 5D5 statement by Kreisel & Moreno (1996), Bottomley (clay) to 6E6 (cocoa brown/leather brown/tan) to almost (1948) also acknowledged the presence of a sterile base. 7D5], with an evanescent metallic gloss, disappearing with The holotype is in a very poor state, consisting only of time (persistent only in folds on holotype), leaving surface the sterile base with very little still attached to it; dull brown, colour of inside surface ± 5D6 (honey yel­ the peridium has disintegrated almost completely into low/oak brown) to 5E6 (mustard brown), outer surface of tiny fragments. holotype ornamented with numerous, tiny, off-white, stel­ late ridges. Hyphal layer of endoperidium composed of Although Verwoerd (1925) did not recognize the cyanophilic, branched hyphae mostly up to 5 pm diam.. but numerous, very conspicuous slit-like perforations in the inflated in places, especially at branches and apices, mod­ capillitial walls of Lanopila capensis for what they real­ erately thick-walled (mostly 0.75-1 pm), not tapering but ly were, he did notice the resultant appearance of the ending in rounded to often inflated tips, true septa infre­ capillitium, which he described as ‘... with a marbled sur­ quent but not uncommon, often perforated with slit-like face'. emphasizing the diagnostic value of this character. pits but much less conspicuously so compared to capilli­ Bottomley (1948) described the capillitium as having 4a tium. Gleba cottony to powdery, very fragile, brown (5D5 watered appearance'. to 6E6) consisting of and capillitium. Capillitium golden brown in clear lactophenol. strongly cyanophilic in Numerous long. thin, angular, needle-shaped crystals, lactophenol with aniline blue, inamyloid in Melzer’s solu­ not visible with the naked eye but very conspicuous under tion. dichotomously branched, commonly up to 7 (rarely the SEM. occur on the peridial surface of the holotype. up to 11) pm diam., gradually tapering to thin-walled, These crystals were not observed on the isotype, howev­ rounded tips, as little as 1 pm diam., often undulating er. and are assumed to be an artefact of unknown origin. towards apices, very fragile, breaking up into fragments of varying length, septa not observed: capillitial wall smooth, Although Ponce de Léon (1981) excluded this moderately thickened, mostly varying between 0.5 and from Lanopila Fr.. he did not designate it to another 1.25 pm. L/H averaging 0.69 (n = 18) for hyphae 5-7 pm genus. On a slip dated 1991. inserted with the type speci­ diam., with numerous, very conspicuous, small to large fis­ men. he assigned it to Calvatia Fr.. however, albeit with sures or slit-like pits. Paracapillitiwn absent a question mark. Calonge. also on a herbarium slip dated golden brown in clear lactophenol. cyanophilic reaction 1992. placed it in Langermannia. Based on Bottomley’s variable, inamyloid in Melzer's solution, globose, with description of the gleba being 'septate but fragmenting at short hyaline apiculus mostly less than 1 pm long, but up the septa'. Kreisel (1992, 1994) reduced Lanopila capen­ to ± 1.3 pm not uncommon, uniguttulate. radial symmetric, sis to synonymy under Calvatia flava (Massee) Kreisel. isopolar, 3-5 pm diam., generally appearing to have a After having examined the holotype. however, he recog­ diameter slightly less than much of capillitium; wall nized it as a distinct species and placed it in the genus ± 0.5 pm thick, glabrous under light microscope but dis­ Handkea Kreisel on the basis of its slit-like capillitial tinctly verruculose under SEM, verrucae not more than 0.2 perforations (Kreisel & Moreno 1996). We prefer to pm high. Subgleba present, small (up to 10 mm high), yel­ retain it in Calvatia, however, assigning it to sect. lowish brown in holotype (5D5) to brown in isotype (6E6). Macrocalvatia emend. compact, composed of yellow-brown, cyanophilic, branched, much contorted and bent hyphae. commonly up Prior to this study Calvatia capensis was known from to 7.5 pm diam.. tapering like capillitial hyphae. moderate­ the type collection only, but a re-examination of the puff­ ly thick-walled (mostly between 1.0-1.5 pm), apparently balls in the E.L. Stephens collection, recently transferred aseptate. false septa observed but rare; diaphragm absent, from BOL to PREM. as well as some specimens from the boundary with gleba poorly defined. Lloyd collection in BPI. brought to light at least five more collections of this fungus. Distribution: Western Cape. South Africa. Specimens examined Habitat: soil in a temperate climatic zone with mild, wet winters and hot, dry summers (Mediterranean cli­ WESTERN CAPE —3318 (Cape Town): Rietvlei. 10 June 1951. mate). Fynbos Biome. (-CD or DC), herb Stephens 925 (PREM): Rietvlei. (-CD or DC). herb. Stephens 193! (PREMi: Devil's Peak near wattles. 7 July 1962. (-CD). Chapnuin s.n. sub herb. Stephens 4301 (PREM): Stellenbosch. Discussion: the material from which Lloyd (1923) Papegaaiberg. 19 June 1921. (-DD). A.V. Duthie 304 sub herb. Uoyd first described this fungus consists of one half of a longi­ 7567(RP1706I62. holo.. PREM3I472. iso.l. 3418 (Simonstown): Smits- tudinally bisected specimen sent to him by Miss A.V. winkel Bay. by roadside. 28 Apnl 1937. (-AD). R.S. Adamson s.n. sub 158 Bothalia 33.2 (2003) herb. Stephens 452 (PREM). Locality unknown: South Africa. A.V. GREUTER. W.. MCNEILL, J., BARRIE. F.R.. BURDET. H.M.. DE­ Duthie s.n. sub herb. Lloyd 51765 (BPI709920). MOULIN, V.. FILGUEIRAS, T.S., NICOLSON. D.H.. SILVA. PC.. SKOG. J.E.. TREHANE. P., TURLAND, N.J. & HAWKS- WORTH. D.L. 2000. International Code of Botanical Nomen­ The following specimens, all with slit-like capillitial clature (Saint Louis Code). Regnum vegetabile 138. Koeltz pores and semblances of sterile bases are very similar to Scientific Books. Konigstein. C. capensis, but the material is too scanty to allow for KORNERUP. A. & WANSCHER. J.H. 1981. Methuen handbook of colour. definite identification: edn 3, 1st reprint. Eyre Methuen, London. KREISEL, H. 1962. Die Lycoperdaceae der Deutschen Demokrati- WESTERN CAPE—3318 (Cape Town): Glen, in grass. 12 May schen Republik. Feddes Repertorium 64: 89-201. 1954. (-CD), Chapman 424 sub herb. Stephens 1387 (PREM): between KREISEL. H. 1989. Studies in the Calvatia complex (Basidiomycetes). Klipheuwel and Bellville. 17 July 1955, (-DA to DC), herb. Stephens Nova Hedwigia 48: 281-296. 1500 (PREM). Locality unknown: herb. Stephens 2032 (PREM). KREISEL. H. 1992. An emendation and preliminary surv ey of the genus Calvatia (Gasteromycetidae). Persoonia 14: 431^439. KREISEL. H. 1994. Studies in the Calvatia complex (Basidiomycetes) ACKNOWLEDGEMENTS 2. Feddes Repertorium 105: 369-376. KREISEL. H. & MORENO. G. 1996. The genus Handkea Kreisel (Basi­ The curators of BPI and PREM are thanked for the diomycetes, Lycoperdaceae) in the southern hemisphere. Feddes Repertorium 107: 83-87. loan of specimens in their care. LANGE, M. 1993. Classifications in the Calvatia group. Blyttia 51: 141-144. REFERENCES LLOYD, C.G. 1923. The genus Lanopila. Mvcological Writings 7: 1177. PONCE DE LEON, P. 1981. Langermannia bicolor (Lév.) Demoulin & BOTTOMLEY. A.M. 1948. Gasteromycetes of South Africa. Bothalia Dring. Phytologia 48: 373-383. 4: 473-810. VERWOERD, L. 1925. Suid-Afrikaanse Lycoperdaceae en Nidulari- CALONGE. F.D. 1998. Flora mycologica Iberica 3. Gasteromycetes I. aceae. Annale van die Universiteit van Stellenbosch. Reeks A, Lxcoperdales, Nidulariales, Phallales, Sclerodermatales, wis- en natuurkunde 3: 1—45. Tulostomatales. Cramer. Stuttgart. CALONGE, F.D. & MARTIN. M.P 1990. Notes on the taxonomical J.C. COETZEE* and A.E. VAN WYK** delimitation in the genera Calvatia. Gastropila, and Langer- mannia (Gasteromycetes). Boletín Sociedad Micológica de * Department of Horticulture and Food Technology, Peninsula Madrid 14: 181-190. Technikon. P.O. Box 1906, 7535 Bellville. DEMOULIN. V. 1993. Calvatiapachyderma (Peck) Morg. and Gastro­ ** H.G.W.J. Schweickerdt Herbarium. Department of Botany, Uni­ pila fragilis (Lév.) Homrich & Wright, two possible names for versity of Pretoria, 0001 Pretoria. the same fungus. Mycotaxon 46: 77-84. MS. received: 2002-12-05.

ASTERACEAE

A NEW COMBINATION IN THE GENUS MACLED1UM (MUTISEAE)

Klatt (1866) described a new variety of Dicoma zey- Dicoma kirkii Harv. var. microcephala S.Moore: 231 (1921). Type: heri, var. thyrsiflora, for a taxon collected in Barberton South Africa, Transvaal [Mpumalanga], Barberton. Thomcroft 1074 (BM. holo.; PRE!, iso.). (Figure 2). Thellung (1921) subsequently raised this vari­ ety to the species level. Dicoma thyrsiflora. In the same According to K. Balkwill (pers. comm.), Macledium year Moore (1921) described the same taxon as Dicoma zeyheri subsp. thyrsiflorum is a serpentine endemic. kirkii var. microcephala. Pope (1991) considered this taxon to be an example of variation in Dicoma zeyheri The subspecies of Macledium zeyheri are keyed out as subsp. zeyheri. While revising the genus Dicoma. the first follows: author studied the variation in Dicoma zeyheri. and var. thyrsiflora was found to be distinct and morphologically la Involucral bracts extending 60-280 mm down the capitulum different from the typical subspecies. Ortiz (2001) rein­ stalk; leaves located basally or absent...... stated the genus Macledium Cass, and Dicoma zeyheri ...... M zeyheri subsp. argyrophxllum was transferred to this genus. He, like Pope (1991), also lb Involucral bracts confined to the head or extending less than recognized only two subspecies: M. zeyheri subsp. argy- 60 mm down the capitulum stalk; plants leafy: rophyllum and M. zeyheri subsp. zeyheri and included the 2a Heads very large, more than 30 mm wide, solitary or up to above discussed taxon under the typical subspecies. It is three on a peduncle; involucral bracts silvery tinged purple, with narrow whitish margins; leaves up to 35 here proposed that the name of this taxon be reinstated mm w ide...... M. zeyheri subsp. zeyheri with the correct nomenclatural combination as follows: 2b Heads relatively small, less than 20 mm wide, often numerous in a panicle; involucral bracts light green with broad, whitish margins; leaves relatively narrow, Macledium zeyheri (Sond.) S.Ortiz subsp. thyrsi- up to 20 mm wide...... M. zeyheri subsp. thyrsiflorum florum (Klatt) N.C.Netnou. comb. nov.

ACKNOWLEDGEMENTS Dicoma zeyheri Sond. var. thyrsiflora Klatt in Bulletin de I'Herbier Boissier 4: 844 (1866). Dicoma thyrsiflora (Klatt) Thell. in Thell. & Schinz: 25 (1921). Type: South Africa, Transvaal [Mpumalanga]. This note is part of an M.Sc. thesis submitted to Rand Kaapriver Valley. Barberton. Galpin 911 (PRE!. holo.; K. BOL!. iso.). Afrikaans University. Financial assistance from the NBI.