Hymenochaetales: a Molecular Phylogeny for the Hymenochaetoid Clade
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Mycologia, 98(6), 2006, pp. 926–936. # 2006 by The Mycological Society of America, Lawrence, KS 66044-8897 Hymenochaetales: a molecular phylogeny for the hymenochaetoid clade Karl-Henrik Larsson1 the Hymenochaetaceae forms a distinct clade but Department of Plant and Molecular Sciences, Go¨teborg unfortunately all morphological characters support- University, Box 461, SE 405 30 Go¨teborg, Sweden ing Hymenochaetaceae also are found in species Erast Parmasto outside the clade. Other subclades recovered by the Institute of Agricultural and Environmental Sciences, molecular phylogenetic analyses are less uniform, and Estonian University of Life Sciences, 181 Riia Street, the overall resolution within the nuclear LSU tree 51014 Tartu, Estonia presented here is still unsatisfactory. Key words: Basidiomycetes, Bayesian inference, Michael Fischer Blasiphalia, corticioid fungi, Hyphodontia, molecu- Staatliches Weinbauinstitut, Merzhauser Straße 119, D-79100 Freiburg, Germany lar systematics, phylogeny, Rickenella Ewald Langer INTRODUCTION Universita¨t Kassel, FB 18 Naturwissenschaft, FG ¨ Okologie, Heinrich-Plett-Straße 40, D-34132 Kassel, Morphology.—The hymenochaetoid clade, herein also Germany called the Hymenochaetales, as we currently know it Karen K. Nakasone includes many variations of the fruit body types USDA Forest Service, Forest Products Laboratory, known among homobasidiomycetes (Agaricomyceti- 1 Gifford Pinchot Drive, Madison, Wisconsin 53726 dae). Most species have an effused or effused-reflexed Scott A. Redhead basidioma but a few form stipitate mushroom-like ECORC, Agriculture & Agri-Food Canada, CEF, (agaricoid), coral-like (clavarioid) and spathulate to Neatby Building, Ottawa, Ontario, K1A 0C6 Canada rosette-like basidiomata (FIG. 1). The hymenia also are variable, ranging from smooth, to poroid, lamellate or somewhat spinose (FIG. 1). Such fruit Abstract: The hymenochaetoid clade is dominated body forms and hymenial types at one time formed by wood-decaying species previously classified in the the basis for the classification of fungi. Thus the artificial families Corticiaceae, Polyporaceae and hymenochaetoid clade, as it is defined here, draws its Stereaceae. The majority of these species cause a white members from several families as circumscribed in rot. The polypore Bridgeoporus and several corticioid premolecular classifications: Agaricaceae, Polypora- species with inconspicuous basidiomata live in associ- ceae, Corticiaceae, Stereaceae and Hymenochaeta- ation with brown-rotted wood, but their nutritional ceae but includes only the type genus for the last strategy is not known. Mycorrhizal habit is reported family name. for Coltricia perennis but needs confirmation. A Micromorphological characteristics are exceedingly surprising element in the hymenochaetoid clade is variable. Three basic kinds of hyphae involved in a group of small white to brightly pigmented agarics construction of basidiomycete basidiomata (viz. gen- earlier classified in Omphalina. They form a subclade erative hyphae, skeletal hyphae and binding hyphae) together with some similarly colored stipitate stereoid are present although most species have only the and corticioid species. Several are associated with generative type. Spores are mainly smooth but vary in living mosses or one-celled green algae. Hyphoderma shape from the large globose ones found in pratermissum and some related corticioid species have Globulicium hiemale to the extremely narrow and specialized organs for trapping and killing nematodes strongly bent spores in Hyphodontia (Chaetoporellus) as a source of nitrogen. There are no unequivocal latitans. A few species have finely ornamented spores morphological synapomorphies known for the hyme- (viz. Coltriciella spp. and Hyphodontia (Rogersella) nochaetoid clade. However almost all species ex- griseliniae). amined ultrastructurally have dolipore septa with Most species have some kind of vegetative (sterile) continuous parenthesomes while perforate parenthe- cells in the fruit body tissue, often sharing the space somes is the normal condition for other homobasi- with the basidia in the hymenium (SUPPLEMENTARY diomycete clades. The agaricoid Hymenochaetales FIG. 1). They collectively could be called cystidia but have not been examined. Within Hymenochaetales because some of them have a distinctive form, unique terms have been introduced for them. The majority of Accepted for publication 23 October 2006. species in Hymenochaetaceae have a characteristic 1 Corresponding author. E-mail: [email protected] kind of cystidia called setae (FIG. 1J). These thick- 926 LARSSON ET AL:HYMENOCHAETOID CLADE PHYLOGENY 927 FIG. 1. Macro and micro characters in Hymenochaetales. A–I. Basidiome and hymenophore types. A, Cotylidia pannosa, stipitate with smooth hymenophore (photo David Mitchel, www.nifg.org.uk/photos.htm). B. Coltricia perennis, stipitate with poroid hymenophore. C. Contumyces rosella, stipitate with lamella. D. Clavariachaete rubiginosa (photo Roy Halling) E. Phellinus robustus, sessile to effuse-reflexed with poroid hymenophore (photo Andrej Kunca, Forest Research Institute, Slovakia, www.forestryimages.org). F. Coltricia montagnei, stiptate with contrical lamella (photo Dianna Smith, www.mushroo- mexpert.com). G. Hydnochaete olivacea, resupinate to effuse-reflexed with coarse, compressed aculei. H. Resinicium bicolor, resupinate with small, rounded aculei. I. Hyphodontia arguta, resupinate with acute aculei. J, setal cystidium in Hymenochaete cinnamomea. Bars: A, D 5 10 mm, B 5 2 mm; G–I 5 1 mm, J 5 10 mm. 928 MYCOLOGIA walled, dark, brown and usually acutely pointed cells basidia from the same apical cell and not, as usual, can be observed with a hand lens and give the species through hyphal branching (SUPPLEMENTARY FIG. 1). a beard-stubble look. Their function is possibly to Each new basidium bursts through the old, empty protect the hymenium from insects. Thin-walled and basidium leaving a progressively longer row of hyaline cystidia characterize the hymenium in all sheathing basidia walls along the subtending hypha. species of Hyphodontia, many other small genera of Basidial repetition is not unique for Repetobasidium, corticioid fungi and some of the agaricoid and although this is the genus where it first was observed stereoid genera. Often these cystidia are pestle- and described (Eriksson 1958). shaped with a globular apex. The exact function is Ultrastructural characteristics as observed with not known but a general idea is that leptocystidia a scanning or transmission electron microscope are emerging beyond the basidial layer function as of limited importance for the taxonomy and classifi- excretory organs because in living specimens the cation of higher fungi. The prime exception is the apex of such cystidia often are covered by a droplet septal pore apparatus that differs markedly among that disappears or dissolves when material is mounted various groups, and these anatomical differences for observation microscopically. In some cases the correlate well with basidiomycete higher order classi- apical droplet seems encased in a vesicle and resists fications. Agaricomycetidae is characterized by a septal dissolution. The most well developed vesicle-bearing pore apparatus called a dolipore. In a dolipore the cystidia, called halocystidia, are found in Resinicium septal pore is surrounded on each side of the septum bicolor and related species. Crystal-covered cystidia are by a half-dome-shaped membrane called parenthe- other indications of excretion capacity. In Hyphodon- somes because in a TEM picture it looks like the tia and Resinicium some species have lagenocystidia septum is placed within parentheses. Most homo- that are hypha-like but with a needle-like termination. basidiomycetes have parenthesomes that are perfo- At the apex these cystidia carry a rosette of crystals, rated by a number of small openings and appear as presumably composed of calcium-oxalate. Thin- to dashed marks in the TEM; however a small number of thick-walled cystidia with an apical crystal cap charac- species instead have nonperforate parenthesomes. terize species in the polypore genera Trichaptum and Because species in Auriculariales and Tulasnellales Oxyporus and thick-walled, strongly encrusted cystidia have the nonperforate type, this is regarded as the (metuloids, lamprocystidia) can be seen in Hypho- ancestral condition from which the perforated de- derma puberum and a few other species. In Tubulicri- veloped. In Hymenochaetales as well as in Cantha- nis all species have lyocystidia, a hallmark of the rellales both types occur, which indicate that the genus. These cystidia have a thick-walled ‘‘stem’’ and evolution of parenthesome type is more complicated a thin-walled variously shaped apex. The thick-walled than initially understood. part is usually more or less amyloid and dissolves easily in 5% KOH. Gloeocystidia (enclosed cystidia Ecology.—Saprotrophy is the dominating life strategy with more or less refractive contents) are not in Hymenochaetales. Most species live in deadwood common but occur in Hyphoderma praetermissum and satisfy their energy needs by decaying the and related species and in Physodontia. Hyphoderma polysaccharides cellulose, hemicellulose and lignin. praetermissum also is known for its stephanocysts. When all these molecules are degraded at roughly These are one- or two-celled hemispherical or globose equal rates the resulting decay is called white-rot as structures that occur on hyphae in the