New Lichenicolous, Muscicolous, Corticolous and Lignicolous Taxa of Burgoa S
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Mycol Progress (2007) 6:61–80 DOI 10.1007/s11557-007-0523-3 ORIGINAL ARTICLE New lichenicolous, muscicolous, corticolous and lignicolous taxa of Burgoa s. l. and Marchandiomyces s. l. (anamorphic Basidiomycota), a new genus for Omphalina foliacea, and a catalogue and a key to the non-lichenized, bulbilliferous basidiomycetes Paul Diederich & James D. Lawrey Received: 26 July 2006 /Revised: 10 January 2007 /Accepted: 12 January 2007 /Published online: 1 March 2007 # German Mycological Society and Springer-Verlag 2007 Abstract A catalogue and a key to the non-lichenized, andiomyces, M. buckii and M. nothofagicola, are described. bulbilliferous basidiomycetes are given. The new genus As Marchandiomyces aurantiacus is phylogenetically more Burgella is described for the lichenicolous B. flavoparme- close to Erythricium than to Marchandiomyces,itis liae, phylogenetically close to Sistotrema oblongisporum and proposed to exclude it from that genus and to use the Multiclavula. The genera Pneumatospora and Tricellulortus holomorphic generic name Marchandiobasidium for both are placed in synonymy of Minimedusa, the new combina- anamorph and teleomorph of this species. The new genus tion M. obcoronata is proposed, and the new facultative Marchandiomphalina is introduced for the lichenized lichenicolous M. pubescens is described. The new facultative Omphalina foliacea, a taxon phylogenetically close to lichenicolous Burgoa angulosa is phylogenetically close to Marchandiobasidium. the generic type B. verzuoliana, whilst the new B. moriformis and B. splendens are provisionally included in Keywords Bulbils . Cantharellales . Minimedusa . the genus Burgoa.ABurgoa-like species in the Ceratoba- Sclerotia . Sistotrema sidiaceae is left unnamed. Two new species of March- Introduction Taxonomic novelties Burgella Diederich & Lawrey; B. flavoparmeliae Diederich Most lichenicolous fungi belong to the ascomycetes or & Lawrey; Burgoa angulosa Diederich, Lawrey & Etayo; heterobasidiomycetes, and only a few lichenicolous homo- B. moriformis Diederich, Ertz & Coppins; B. splendens basidiomycetes are known. These include agarics belonging Diederich & Coppins; Marchandiomphalina Diederich, to the genera Arrhenia and Fayodia, and sclerotial fungi Lawrey & Binder; Marchandiomphalina foliacea (P. M. Jørg.) Diederich, Lawrey & Binder; Marchandiomyces buckii belonging to Athelia, Leucogyrophana and Marchandiomy- Diederich & Lawrey; M. nothofagicola Diederich & Lawrey; ces (Lawrey and Diederich 2003). Sclerotia are frequently Minimedusa pubescens Diederich, Lawrey & Heylen; M. obcoronata regarded as being of several types, including true sclerotia, (B. Sutton, Kuthub. & Muid) Diederich & Lawrey pseudosclerotia and bulbils (Clémençon et al. 2004). True P. Diederich (*) sclerotia are known from the lichenicolous Athelia arach- Musée national d’histoire naturelle, noidea and Leucogyrophana lichenicola, and bulbils from 25 rue Munster, the lichenicolous Marchandiomyces aurantiacus and M. L-2160 Luxembourg, Luxembourg e-mail: [email protected] corallinus. Hotson (1912, 1917) presented a survey of all known J. D. Lawrey bulbilliferous fungi, mainly described in the genus Pap- Department of Environmental Science and Policy MSN 5F2, ulaspora (= Papulospora), and in his key he grouped those George Mason University, 4400 University Drive, with basidiomycetous affinities, i.e. either presenting Fairfax, VA 22030-4444, USA clamps or producing basidia in culture. These basidiomy- 62 Mycol Progress (2007) 6:61–80 cetous, bulbilliferous fungi were later accommodated in the gamic communities, frequently overgrowing or killing other new genus Burgoa (Goidánich 1938), whilst most other cryptogams, and as they are particularly interesting in species with ascomycetous affinities were left in Papulas- phylogenetic and evolutionary studies. pora. Weresub and LeClair (1971) gave a thorough review of the different definitions of bulbils and papulaspores (= papulospores), and the authors concluded that the term Materials and methods bulbil should be restricted to basidiomycetes, and papulas- pores to ascomycetes. They further described the new Specimens from the herbaria BR, E, GMUF, MA, NMW, NY, genus Minimedusa for a species with basidiomycetous STU and UPS, and from the private collections of A. Aptroot, affinities, in which bulbils develop from a multihyphal M.Cole,P.Diederich,J.Etayo, O. Heylen-Walraevens, B. base. As Sistotrema-like basidia have occasionally been Schultheis and P. van den Boom, were examined. observed in cultures of Burgoa species (e.g., Weresub and Dry herbarium specimens were examined and measured LeClair 1971), and Burgoa-like bulbils were encountered in under a binocular microscope Leica MZ 7.5 (magnification some Sistotrema specimens or cultures (Clémençon et al. up to 50×) and photographed using a Nikon Coolpix 4500. 2004; Eriksson et al. 1984; Hallenberg 1984), it is generally Entire unsectioned bulbils were studied in water, 5% KOH, assumed that Burgoa is the anamorphic state of Sistotrema. or lactophenol cotton blue (LCB), either without or with During the past 20 years, we regularly collected or pressure on the coverslip. Microscopic drawings were done received material of fungi overgrowing corticolous crypto- using a drawing tube, and photographs of bulbils in LCB gams (lichens or mosses) or growing directly over bark or were prepared using a Zeiss Photomikroskop III with a lignum, with bulbils resembling Marchandiomyces species, Canon PowerShot G5. but pale colored (whitish to brownish, yellowish or pale orange). Morphological and anatomical studies revealed the existence of at least six apparently undescribed species, Results amongst which we were able to grow four in pure culture, some of them from up to 5-year-old herbarium specimens. New lichenicolous, muscicolous, corticolous DNA sequences were obtained from these four cultured and lignicolous taxa of Burgoa s. l. and Marchandiomyces s. l. species, and from one additional species by direct PCR. Phylogenetic analyses suggest that the five species belong Burgella Diederich & Lawrey, gen. nov. to the Cantharellales (Binder et al. 2005,as“cantharelloid clade”), and most of them group with species of Sistotrema. Fungus anamorphicus bulbillis lichenicola superficialibus We were also able to get DNA sequences of the type vel parve immersis, melleis, subsphaericis ad ellipsoideis cultures of the type species of Burgoa and Minimedusa, and vel irregularibus, glabris, cellulis subsphaericis adhaerenti- we found that several of our new species group with them. bus; in cultura bulbillis agglomeratis gelatinosis corallifor- Furthermore, we had the opportunity to study, culture mibus, septis fibulatis. and sequence Marchandiomyces specimens belonging to Type: Burgella flavoparmeliae Diederich & Lawrey. undescribed species. Finally, our phylogenetic results Colonies appearing as dispersed or rarely agglomerate suggested that Omphalina foliacea is closely related to bulbils. Bulbils honey-colored, ±shiny, without hairs, Marchandiomyces s. l. roundish to shortly ellipsoid or irregular in form, partly The phylogenetic results will be published in a separate immersed to superficial, externally without specialized paper that will include almost all lichenicolous species cells, cells in surface view polyhedral, internally of strongly forming sclerotia or bulbils, a selection of other bulbilli- adherent, more or less roundish to ellipsoid or polyhedral ferous species, and a selection of lichenized basidiomycetes cells separating only with difficulty; clamps not observed. (Lawrey et al. 2007). A summary tree of these results is Colonies in liquid culture showing abundant white aerial given in Fig. 1. hyphae, from which orange agglomerations of bulbils arise. The aim of the present taxonomic paper is to formally Hyphae hyaline, septate, straight, rarely branched or describe the new, lichenicolous, muscicolous, corticolous anastomosed; septa with or without clamp connections. and lignicolous bulbilliferous species studied by us, to Agglomerations of bulbils gelatinous in appearance, almost describe a new genus for Omphalina foliacea, to give a coralloid, composed of irregularly shaped bulbils; bulbils catalogue of all known non-lichenized, bulbilliferous externally and internally composed of irregular, roundish or basidiomycetes, and a key allowing their identification. elongate cells with clamped septa. Although all these new species lack both teleomorphic and Following Lawrey et al. (2007), Burgella flavoparmeliae anamorphic stages, it is important to formally name them, belongs to the subclade of the Cantharellales containing the as they are important in corticolous or lignicolous crypto- Clavulinaceae. These results place B. flavoparmeliae and Mycol Progress (2007) 6:61–80 63 Fig. 1 Phylogenetic relation- Pterulaceae ships of lichenized, lichenico- Burgoa turficola Stephanospora caroticolor lous, and bulbilliferous fungi Phyllotopsis nidulans discussed in the text. A truncat- Typhula phacorrhiza Agaricales Arrhenia spp. ed version of a tree from Lawrey Pseudoarmillariella ectypoides Lichenomphalia spp. et al. (2007) inferred from multi- Lichenomphalia umbellifera gene data using Bayesian anal- Xeromphalina spp. Amylocorticium subincarnatum yses. Focal species are Plicaturopsis crispa Myxomphalia maura highlighted in bold Fayodia gracilipes Sarcomyxa serotina Camarophyllus pratensis various mushroom-forming agarics Atheliales Boletales Jaapia argillacea Tylospora asterophora Russulales Polyporales (includes