Polyporaceae): an African Lentinoid Fungus with an Unusual Combination of Both Skeleto-Ligative Hyphae and Pleurocystidia

Home , Heliocybe, Lentinus, Neolentinus, Panus

Plant Ecology and Evolution 146 (2): 240–245, 2013 http://dx.doi.org/10.5091/plecevo.2013.792

SHORT COMMUNICATION

Lentinus cystidiatus sp. nov. (Polyporaceae): an African lentinoid fungus with an unusual combination of both skeleto-ligative hyphae and pleurocystidia

André-Ledoux Njouonkou1,*, Roy Watling2 & Jérôme Degreef3

1Department of Biological Science, Faculty of Sciences, University of Bamenda, P. Box 39, Bamenda, Cameroon 2Caledonian Mycological Enterprises, 26 Blinkbonny Avenue, Edinburgh EH4 3HU Scotland, U.K. 3Department of Cryptogamy, National Botanic Garden, Domain of Bouchout, Nieuwelaan 38, B-1860 Meise, Belgium *Author for correspondence: [email protected]

Background and aims – Lentinus species are a major component of the agaricoid flora of tropical Africa where fifteen species have been documented with few studies in Cameroon. This work aims to contribute to the taxonomy of the genus Lentinus by describing a putative new species collected in south-western Cameroon. Methods – A unique lentinoid fungi specimen collected in Korup National Park in the South-West region of Cameroon and preserved in the Edinburgh herbarium (E) was examined macro- and microscopically following classical mycological description methods. Key results – The specimen examined possesses squamules on the pileus and stipe surface, no annulus, furcated branching dichotomous lamellae, oblong-cylindrical basidispores, basidia generally bearing four sterigmata (sometimes two or one) reaching 5 µm long, skeleto-ligative hyphae and pleurocystidia. The simultaneous presence of both pleurocystidia and skeleto-ligative hyphae has never been encountered in the genus Lentinus. Due to this unusual combination and other specific features of this specimen, it is considered as a representative of a new species within the genus Lentinus. Discussion – Lentinus cystidiatus exhibits some similar characteristics with Lentinus squarrosulus that possesses squamules on the pileus and stipe surface and mostly with species of the sub-genus Lentinus section Dicholamellatae that are characterized by furcated lamellae. As skeleto-ligative hyphae is characteristic of the sub-genus Lentinus and pleurocystidia characteristic of sub-genus Panus, the simultaneous presence of elements brings into question the taxonomic position of this new species and the systematics of the genus Lentinus in general. Conclusions – The existence of this new species in Cameroon opens a door on the necessity of a taxonomic revision of the genus Lentinus and allied genera.

Key words – Lamellate Polyporaceae, pleurocystidia, skeleto-ligative hyphae, Korup National Park, Cameroon, Central Africa.

INTRODUCTION raceae, Aphyllophorales) is the most popular genus as some of its representatives are edible or medicinal (Pegler 1983, Lentinoid mushrooms are characterised macroscopically by Rammeloo & Walleyn 1993). a depressed umbilicate to deeply infundibuliform pileus and decurrent lamellae. Species that portray such characteristics The taxonomy of the genus Lentinus has been preoccupy- mainly belong to genera Heliocybe Redhead & Ginns, Len- ing for decades. Pegler (1983) distinguished two subgenera tinula Earle, Lentinus Fr., Panus Fr., Neolentinus Redhead within Lentinus on the basis of their common tough, coria- & Ginns, Trogia Fr. or Pleurotus (Fr.) P.Kumm. Many of ceous-fleshy texture. He restricted the subgenus Lentinus to them are xerophitic, saprobic and lignicolous mainly caus- species combining skeleto-ligative hyphae and, sometimes, ing white and brown rot thanks to the enzymes they produce hyphal pegs but no pleurocystidia (metuloids and gloeocys- such as laccase, peroxidase, cellulase and ligninase (Caval- tidia); the subgenus Panus was restricted to species with skel- lazzi et al. 2004, Mossebo et al. 2007). Lentinus (Polypo- etal hyphae, without hyphal pegs and, sometimes, pleurocys-

tidia. Corner (1981) and Singer (1986) considered Lentinus Botanic Garden of Edinburgh (E) and the herbarium of the and Panus as two different genera. They limited Lentinus to Royal Botanic Gardens of Kew [K(M)]. So far, many papers those species with ligative hyphae, separating those species have been published on fungi of the KNP, mainly on non- with skeletal hyphae in Panus. Despite the comprehensive gill mushrooms (Hjortstam et al. 1993, Masuka & Ryvarden morphological investigations of Corner (1981) and Pegler 1999, Roberts & Ryvarden 2006, Roberts 1999, 2000, 2001, (1972, 1975, 1983), many disagreements on the limits of the Douanla-Meli et al. 2007) and ectomycorrhizal taxa (Watling genus Lentinus (s. l.) were remaining. Based on wood rot 1993). These studies have led to the description of one new types, Redhead & Ginns (1985) segregated the genera Neo- genus Korupella P.Roberts & Hjortstam and many new fun- lentinus and Heliocybe both responsible for brown rot wood gal species. from Lentinus that causes white rot. Phylogenetic studies While studying Lentinus materials collected in the KNP inferred from molecular characteristics using ribosomal 28S and deposited in E, we noted that morphological and ana- and ITS sequence data (Hibbett & Vilgalys 1993, Douanla- tomical characters of one collection identified as ‘Lentinus Meli & Langer 2010, Grand et al. 2012) showed that Lenti- squarrosulus’ differed from the description of this species nus, Panus, Neolentinus, and Heliocybe must be considered made by Pegler (1971, 1972, 1977, 1983). Furthermore, this as distinct genera. Recently Karunarathna et al. (2011) on specimen showed unusual characters that did not match any basis of morphological characters and ITS sequence data de- previously known species of Lentinus. It is described here as scribed three new species of Lentinus sensu stricto (Lentinus a new species. roseus Karunar., K.D. Hyde & Zhu L.Yang, L. concentricus Karunar., K.D.Hyde & Zhu L.Yang and L. megacystidiatus Karunar., K.D.Hyde & Zhu L.Yang) from northern Thailand. MATERIALS AND METHODS Nevertheless, due to the fact that phylogenetic and enzymatic The materials studied in this work were collected during studies of Lentinus (sensu Pegler 1983) concerned very few field trip by one of us (Roy Watling) in the KNP in January species (Mossebo 2002), Manimohan et al. (2004) still used 1989. After collection basidiomata were dried before being the classification of Pegler (1983). kept in the herbarium. Macroscopic description was made on Lentinus species are mostly confined to the tropics and fresh material. This description was completed by later ob- sparsely distributed in the temperate regions (Pegler 1983). servations on dried materials. Free hand sections were made As several species of Lentinus (as well as Panus) are xero- with the aid of a razor blade. For microscopic observations, morphic and long-lived, many mycologists and botanists the sections were firstly mounted in 10% KOH to assess the have collected specimens during their field trips and material natural colour of microscopic features. Then, for a suitable is well represented in various herbaria (Pegler 1983). Despite observation of elements, sections were stained in Congo Red this fact, species identification and delimitation still remain or Phloxine B in 10% KOH. Melzer’s reagent was used to difficult because of the complex microstructure and the high test the amyloidy or dextrinoidy of basidiospores and other degree of polymorphism in this group (Pegler 1983). structures. A binocular microscope (Olympus BH2) equipped The Korup National Park (KNP) in the South West Re- with a drawing tube and an ocular micrometer was used for gion of Cameroon, is a natural lowland rainforest covering observations, drawing and measurements. an area of 127 000 ha. It is abundantly watered, with a mean The dimension of spores are given as follow: L –(L)–L × annual rainfall reaching more than 5000 mm, monthly av- 1 2 l1–(l)–l2 µm; Q = Q1–(Qm)–Q2 where L1 is the smallest length, erage temperature 24–30°C and 86% average humidity. The (L) the average length, L the greatest length, l the smallest forest soils are sandy, low in nutrients, and acid (Newbery et 2 1 width, (l) the average width and l2 the greatest width. Q is al. 1988). The Park hosts an Atlantic coastal forest dominat- the ratio of length and width; Q the smallest ratio, (Q) the ed by representatives of the Caesalpiniaceae many of them 1 average ratio and Q2 the greatest ratio. N is the number of being ectomycorrhizal (Newbery et al. 1988). It also includes spores measured. The dimensions of basidia do not include part of the Guinea-Congolian refugium into which equatorial those of sterigmata. forest was reduced during last Glacial Maximum in the Pleis- Species and genera authors and nomenclatural criteria tocene (Leal 2004, Nicolas et al. 2012). The natural settings and the poor accessibility have allowed the natural rainforest are from Index Fungorum (http://www.indexfungorum.org/ to flourish in the area and enhanced its preservation. Names/Names.asp). The herbarium codes were retrieved from sweetgum (http://sweetgum.nybg.org/ih/). Korup is characterised by high levels of endemism and is reputed to be the best remaining example of the most wide- spread Atlantic Biafran forest of western central Africa and RESULTS richer than any other studied African forest. More than 620 Lentinus cystidiatus Njouonkou, Watling & Degreef, sp. species of trees and shrubs have been recorded from Korup, nov. almost 30% of them are endemic and at least 480 species of herbs and climbers are also present (Anonymous 2008). The Pileus < 101 mm latus, cyathiformis vel infundibuliformis; mycoflora of KNP is the most explored and the best known superficies ferrens squamulos contraetos; (spuma) pallida quantitatively and qualitatively in Cameroon (Douanla-Meli lutea leviter atra ad medium et pallens ad externum ; umbili- 2007). Many fungal collections made mainly by Roy Wa- cus tenuis subtiliter striatus. Lamellae decurrentes, pallida et tling of the Royal Botanic Garden, Edinburgh (from 1989 to (spuma) atra, moderate densae sine liberis lamellulis sed for- 1991) and Peter Roberts of the Royal Botanic Gardens, Kew mans furcatas cum integris lamellis. Stipes 23–51 × 5–9 mm, (in 1996 and 1997) are kept in the herbarium of the Royal centralis vel leviter excentricus, cylindricus leviter attenu-

241 Pl. Ecol. Evol. 146 (2), 2013

B C

Figure 1 – Lentinus cystidiatus: A, habit of the basidiome; B, basidiospores; C, basidia; D, cheilocystidia; E, pleurocystidia; F, generative hyphae; G, skeleto-ligative hyphae. Scale bar = 2 cm (A), 10 µm (B–F). atus, solidus, superficies colore similis pilei coloris cum basi radiata. Systema hypharum dimitica; hyphae generatoriae aterrima, fibrillosa cum aliquis squamilosis et sine annulo. 2–4 µm latae hyaline, ramosae, fibulatae; hyphae skeleto- Basidiosporae 5–7 × 2–3 µm, cylindricae, hyalinae. Basidia -ligativae 3–5 µm latae, paries crassus cum central luce. Pilei 20–33 × 4–5.5 µm, clavato-cylindrica, 40-sporigera interdum crusta epicutis hypharum 3–5 µm latus, tenuis paries et fer- 2 (duo) et 1 (unis). Pleurocystidia 19–31 × 6–17 µm, clavata rens fibulas. – Type: Cameroon, South-West region, Ndiang vel clavato, cylindrica vel fusiforma, paries tenuis, hyalina. division, Korup National Park, transect Q to base camp 2, 30 Cheilocystidia 10–22 × 5–7,5 µm, clavata vel clavato-cylin- Jan. 1989, Watling 216442 (holo-: E, barcode E00297138). drica, hyalina, paries tenuis. Trama hymenophoralis enormis, Mycobank number: MB 803425

242 Njouonkou, Watling & Degreef, A new Lentinus (Polyporaceae) from Cameroon

Basidiomata lentinoid (fig. 1A). Pileus 69–101 mm 3–5.5 µm diameter hyphae, thin-walled, with clamp-connec- diam., soon depressed, finally shallowly cyathiform even tions. Fig. 1B–G. infundibuliform; surface cream-colour to yellow-ochre, Distribution – Known only from Korup National Park in slightly darker towards centre, drying darker even with rust- south west Cameroon. like hues; surface glabrous but with fuscous brown, widely dispersed squamules distributed in concentric rings, finely Habitat – Single or clustered in small troop distributed on sulcate-striate at margin, incised and soon lacerate at thinned exceedingly rotten, moss-covered, fallen, large diameter margin. Lamellae deeply decurrent, pale cream-colour to branch, on forest floor along track, in deep shade. dark cream-colour, paler than pileus, drying more ochra- Conservation status – Unknown. ceous, moderately crowded, furcate with 3 to 4 dichotomic Etymology – The epithet cystidiatus refers to the presence forks before reaching the stipe, without free lamellules; edge of pleurocystidia which, while not common in Lentinus, are apparently entire. Stipe 23–51 × 5–9 mm, slightly excentric present in this species. or central, elongate and tapering downwards to a swollen in- sertion into wood, solid, surface concolorous or paler than the pileus although darker at base, slightly furfuraceous/fi- DISCUSSION brillose with some scurfy darker squamules, without annu- Lentinus cystidiatus is characterized by the following fea- lus. Context white, weak smell, thick at the disk, much thin- tures: (1) umbilicate to infundibuliform pileus with squamu- ner elsewhere. Fig. 1A, electronic appendix 1. lose surface and striate margin; (2) furcated lamellae without Basidiospores 5–(6.3)–7× 2–(2.7)–3 µm; Q = 1.8–(2.3)– free lamellulae; (3) central to eccentric stipe; (4) oblong- 2.8 [N = 25]; oblong-cylindric (fig. 1B), hyaline, smooth sur- cylindric basidiospores; (5) dimitic hyphae system, skeleto- face, thin-walled, with few contents; inamyloid, not dextri- ligative hyphae with a wide lumen; and (6) presence of pleu- noid. Basidia 20–33 × 4–5.5 µm; clavate-cylindric, bearing rocystidia in the hymenium. The first five characters set this four sterigmata, sometimes two or one reaching 5 µm long species in Lentinus; section Dicholamellatae Pegler that in- (fig. 1C).Pleurocystidia 19–41 × 6–17 µm; clavate, cylindric cludes L. badius (Berk.) Berk., L. araucariae Har. & Pat. and or fusiform (fig. 1D); thin-walled and sometimes with hya- L. brunneofloccosus Pegler. Amongst the species of this sec- line contents, originating from the sub-hymenial layer and tion, it is differentiated by the absence of a ring, by larger ba- rising very rarely above the basidia. Cheilocystidia 10–22 × sidia (sometimes mono- or bisporic), and by the presence of 5–7.5 µm; clavate to clavate-cylindric, thin-walled (fig. 1E). pleurocystidia but it also possesses skeleto-ligative hyphae, Hymenophoral trama irregular, of radiate construction, hya- an unsual combination of microscopic structures (table 1). line, consisting of a dimitic hyphal system with skeleto-ligative hyphae. Generative hyphae 2–4 µm diameter, hyaline, Lentinus cystidiatus and L. squarrosulus share the same thin-walled (fig. 1F), branched, with clamp-connections. habit and the presence of squamules on the pileus surface. Skeleto-ligative hyphae 3–5 µm diam., hyaline, slightly However, in L. squarrosulus lamellae are free with lamel- thickened wall (2 µm) but leaving a broad lumen, abundantly lulae while in L. cystidiatus they are furcated, branching branched, with branches of diameter decreasing towards the dichotomously without lamellulae. Microscopically, L. extremities becoming sometimes thread-like (fig. 1G). Sub- squarrosulus possesses hyphal pegs without pleurocystidia hymenium well-developed, pseudoparenchymatous. Context in contrast to L. cystidiatus which possesses pleurocystidia composed of a dimitic hyphae system similar to that of the without hyphal pegs. In the field, identification could be a hymenophoral trama. Pileal surface formed by an epicutis of challenge due to the fact that L. squarrosulus has great vari-

Table 1 – Comparison between Lentinus cystidiatus and other members of the Section Dicholamellatae.

Species L. araucariae L. badius L. brunneofloccosus L. cystidiatus sp. nov. Pileus colour Chestnut brown or paler Dark sepia to fuliginous Chestnut brown to Cream-colour to yellow- umbrinous or greyish ochre brown Lamellae Branching dichotomously Branching dichotomously Branching dichotomously Branching dichotomously arrangement without free lamellulae without free lamellulae without free lamellulae without free lamellulae Stipe Subannulate Subannulate Annulate Not annulate Spore form and Ellipso-cylindric Oblong-cylindric Oblong-cylindric Oblong-cylindric average dimensions 5–7 × 3–3.5 µm 4.7–6.5 × 2–3µm 5.5–8.5 × 2–3.2 µm 5–7 × 2–3 µm Q = 1.87 Q = 2.2 Q = 2.59 Q = 2.3 Basidia dimensions 16–20 × 4–5 µm 17–20 × 3.5–4.5 µm 13–17 × 4–5 µm 20–33 × 4–5.5 µm Vegetative hyphae Skeleto-ligative hyphae Skeleto-ligative hyphae Skeleto-ligative hyphae Skeleto-ligative hyphae Hyphal pegs Present Present Absent Absent Pleurocystidia Absent Absent Absent Present

243 Pl. Ecol. Evol. 146 (2), 2013

ation of basidiome colour and is the most common and best on eucalyptus bark-based medium. Food, Agriculture & Envi- known species of Lentinus in tropical Africa. ronment 2: 291–297. A phylogenetic study of Lentinus subg. Lentinus and re- Corner E.J.H. (1981) The genera Lentinus, Panus and Pleurotus lated taxa based on ITS1-5.8 S-ITS2 rDNA sequences made with particular reference to Malaysian species. Beihefte Zur by Grand et al. (2011) showed similarity between the topol- Nova Hedwigia 69. ogy of major clades and morphological sections summarized Douanla-Meli C. (2007) Fungi of Cameroon. Ecological diversity by Pegler (1975, 1983). The only species of the section Di- with emphasis on the taxonomy of non-gilled Hymenomycetes cholamellatae (to which L. cystidiatus belongs) included in from the Mbalmayo forest reserve. Bibliotheca Mycologica 202. their study was L. badius that poorly supported the phylogenetic position of this section. Douanla-Meli C., Langer E. (2010) Reassessment of phylogenetic relationships of some lentinoid fungi with velutinate basidi- As far as lentinoid fungi (especially in the genus Lenti- omes based on partial 28S ribosomal RNA gene sequencing. nus) are concerned, this is the first time that the combination Sydowia 62: 23–35. of pleurocystidia and skeleto-ligative hyphae are found in the Douanla-Meli C., Ryvarden L., Langer E. (2007) Studies of tropi- same species. According to Corner (1981), Pegler (1983) and cal African pore fungi (Basidiomycota, Aphyllophorales): three Singer (1986), pleurocystidia coupled with skeletal hyphae new species from Cameroon. Nova Hedwigia 84: 409–420. are characteristic of the genus (or sub-genus) Panus while http://dx.doi.org/10.1127/0029-5035/2007/0084-0409 skeleto-ligative hyphae without any form of pleurocystidia Grand E.A., Hughes K.W., Petersen R.H. (2011) Relationships are encountered in the genus (or sub-genus) Lentinus. This within Lentinus subg. Lentinus (Polyporales, Agaricomyce- new species combines both features and can be considered tes), with emphasis on sects. Lentinus and Tigrini. Mycological as intermediate between the two taxa. Such species having Progress 10: 399–413. http://dx.doi.org/10.1007/s11557-010- intermediate position and characters suggests that a revision 0711-4 of lentinoid fungi is urgently needed as suggested already by Hibbett D., Vilgalys R. (1993) Phylogenetic relationships of Lenti- other authors (Hibbett & Vilgalys 1993). However, it is here nus (Basidiomycotina) inferred from molecular and morpholog- classified in the genus Lentinus because of its strong mor- ical characters. Systematic Botany 18: 409–433. http://dx.doi. phological likeness to other species of Dicholamellatae sec- org/10.2307/2419417 tion. Its systematic position could be confirmed after collect- Hjortstam K., Ryvarden L., Watling R. (1993) Preliminary check- ing additional specimens, improving the species identity and list of non-agaricoid macromycetes in the Korup National Park, undertaking enzymatic and molecular studies. Cameroon and surrounding area. Edinburgh Journal of Botany 50: 105–119. http://dx.doi.org/10.1017/S0960428600000743 Karunarathna S.C., Yang Z., Zhao R.-L., Vellinga E.C., Bahkali SUPPLEMENTARY DATA A.H., Chukeatirote E., Hyde K.D. (2011) Three new species of Lentinus from northern Thailand. Mycological Progress 10: Supplementary data are available in pf format at Plant Ecol- 389–398. http://dx.doi.org/10.1007/s11557-010-0701-6 ogy and Evolution, Supplementary Data Site (http://www. ingentaconnect.com/content/botbel/plecevo/supp-data), and Leal M.E. (2004) The African rain forest during the Last Glacial Maximum, an archipelago of forests in a sea of grass. PhD the- consists of a photograph of the basidiomes of the holotype sis, Wageningen University, Wageningen, Netherland. of Lentinus cystidiatus Njouonkou, Watling & Degreef (Wa- tling 216442) from Korup National Park, Cameroon. Manimohan P., Divia N., Arun Kumar T.K. (2004) The genus Lenti- nus in Kerala State, India. Mycotaxon 90: 311–318. Masuka A.J., Ryvarden L. (1999) Dichomitus in Africa. Myco- ACKNOWLEDGEMENT logical Research 103: 1126–1139. http://dx.doi.org/10.1017/ S0953756299008436 Special thanks go to Mapon Mombagnam Rose Christel Mossebo D.C. (2002) Growth of wood-inhabiting Lentinus species from the University of Yaoundé I for her help with Latin de- from Cameroon in laboratory culture. Mycologist 16: 168–171. scription and to Dr. Nkengazong Lucia and Mr. Mekou Yous- http://dx.doi.org/10.1017/S0269915X02004068 souf Bele of CIFOR in Yaoundé for improving the English Mossebo D.C., Njouonkou A.L., Courtecuisse R., Amougou A. language in the first manuscript. We are also very grateful to (2007) Enzymatic activities and decay characteristics in some the reviewers whose comments have substantially improved wood-rotting Basidiomycetes from Cameroon and determina- the quality of this paper. Financial support to the first author tion of time dependent activity of syringaldazine in spot tests. was provided by IAPT, IRD through project SEP (Sud Ex- Cryptogamie, Mycologie 28: 107–121. pert Plante 304). He also thanks the Belgium Focal Point of Newbery D.M., Alexander I.J., Thomas D.W., Gartlan J.S. (1988) the Global Taxonomy Initiative (GTI) who gave him suitable Ectomycorrhizal rain-forest legumes and soil phosphorus in condition to finalize the paper. Korup National Park, Cameroon. New Phytologist 109: 433– 450. http://dx.doi.org/10.1111/j.1469-8137.1988.tb03719.x REFERENCES Nicolas V., Missoup A.-D., Colyn M., Cruaud C., Denys C. (2012) West-Central African Pleistocene lowland forest evo- Anonymous (2008) Management plan of Korup National Park and lution revealed by the phylogeography of Misonne’s soft- its peripheral zone (2009–2013). Yaoundé, Ministère des Forêts furred mouse. African Zoology 47: 100–112. http://dx.doi. et de la Faune. org/10.3377/004.047.0119

Cavallazzi J.R.P., Brito M.S., Oliveira M.G.A., Villas-Bôas S.G. & Pegler D.N. (1971) Lentinus Fr. and related genera from Congo- Kasuya M.C.M. (2004) Lignocellulolytic enzymes profile of Kinshasa (Fungi). Bulletin du Jardin botanique national de Bel- three Lentinula edodes (Berk.) Pegler strains during cultivation gique 41: 273–281. http://dx.doi.org/10.2307/3667639

244 Njouonkou, Watling & Degreef, A new Lentinus (Polyporaceae) from Cameroon

Pegler D.N. (1972) Lentineae (Polyporaceae), Schizophyllaceae Robert P. (2000) Corticioid fungi from Korup National Park, et espèces Lentinoïdes et pleurotoïdes des Tricholomataceae. Cameroon. Kew Bulletin 55: 803–842. http://dx.doi. Flore Illustrée des Champignons d’Afrique Centrale 1. org/10.2307/4113628 Roberts P. (2001) Heterobasidiomycetes from Korup National Pegler D.N. (1975) The classification of the genus Lentinus Fr. (Ba- Park, Cameroun. Kew Bulletin 56: 163–187. http://dx.doi. sidiomycota). Kavaka 3: 11–20. org/10.2307/4119434 Pegler D.N. (1977) A preliminary agaric flora of East Africa. Kew Roberts P., Ryvarden L. (2006) Poroid fungi from Korup National Bulletin, Additional Series 6. Kew, Royal Botanical Gardens. Park, Cameroon. Kew Bulletin 61: 55–78. Pegler D.N. (1983) The genus Lentinus. A world monograph. Kew Singer R. (1986) The Agaricales in modern taxonomy. Koenigstein, Bulletin, Additional Series 10. Koeltz. Rammeloo J., Walleyn R. (1993) The edible fungi of Africa South Watling R. (1993) Comparison of the macromycete biotas in select- ed tropical areas of Africa and Australia. In: Issac S., Frankland of Sahara: a literature survey. Scripta Botanica Belgica 5. J.C., Watling R., Whalley A.J.S. (eds) Aspects of tropical my- Redhead S.A., Ginns J.H. (1985) A reappraisal of agaric genera as- cology: 171–191. New York, Cambridge University Press. sociated with brown rots of wood. Transactions of the Myco- logical Society of Japan 26: 349–381. Manuscript received 30 Aug. 2012; accepted in revised version 6 Roberts P. (1999) Clavarioid fungi from Korup National Park, Mar. 2013. Cameroon. Kew Bulletin 54: 517–539. http://dx.doi. org/10.2307/4110853 Communicating Editor: Elmar Robbrecht.

245