Hymenochaetales: a molecular phylogeny for the hymenochaetoid clade

Karl-Henrik Larsson1

Department of Plant and Molecular Sciences, GoteborgUniversity, Box 461, SE 405 30 Goteborg, Sweden

Erast ParmastoInstitute of Agricultural and Environmental Sciences,Estonian University of Life Sciences, 181 Riia Street,51014 Tartu, Estonia

Michael FischerStaatliches Weinbauinstitut, Merzhauser Straße 119,D-79100 Freiburg, Germany

Ewald LangerUniversitat Kassel, FB 18 Naturwissenschaft, FGOkologie, Heinrich-Plett-Straße 40, D-34132 Kassel,Germany

Karen K. NakasoneUSDA Forest Service, Forest Products Laboratory,1 Gifford Pinchot Drive, Madison, Wisconsin 53726

Scott A. RedheadECORC, Agriculture & Agri-Food Canada, CEF,Neatby Building, Ottawa, Ontario, K1A 0C6 Canada

Abstract: The hymenochaetoid clade is dominatedby wood-decaying species previously classified in theartificial families Corticiaceae, Polyporaceae andStereaceae. The majority of these species cause a whiterot. The polypore Bridgeoporus and several corticioidspecies with inconspicuous basidiomata live in associ-ation with brown-rotted wood, but their nutritionalstrategy is not known. Mycorrhizal habit is reportedfor Coltricia perennis but needs confirmation. Asurprising element in the hymenochaetoid clade isa group of small white to brightly pigmented agaricsearlier classified in Omphalina. They form a subcladetogether with some similarly colored stipitate stereoidand corticioid species. Several are associated withliving mosses or one-celled green algae. Hyphodermapratermissum and some related corticioid species havespecialized organs for trapping and killing nematodesas a source of nitrogen. There are no unequivocalmorphological synapomorphies known for the hyme-nochaetoid clade. However almost all species ex-amined ultrastructurally have dolipore septa withcontinuous parenthesomes while perforate parenthe-somes is the normal condition for other homobasi-diomycete clades. The agaricoid Hymenochaetaleshave not been examined. Within Hymenochaetales

the Hymenochaetaceae forms a distinct clade butunfortunately all morphological characters support-ing Hymenochaetaceae also are found in speciesoutside the clade. Other subclades recovered by themolecular phylogenetic analyses are less uniform, andthe overall resolution within the nuclear LSU treepresented here is still unsatisfactory.

Key words: Basidiomycetes, Bayesian inference,Blasiphalia, corticioid fungi, Hyphodontia, molecu-lar systematics, phylogeny, Rickenella

INTRODUCTION

Morphology.—The hymenochaetoid clade, herein alsocalled the Hymenochaetales, as we currently know itincludes many variations of the fruit body typesknown among homobasidiomycetes (Agaricomyceti-dae). Most species have an effused or effused-reflexedbasidioma but a few form stipitate mushroom-like(agaricoid), coral-like (clavarioid) and spathulate torosette-like basidiomata (FIG. 1). The hymenia alsoare variable, ranging from smooth, to poroid,lamellate or somewhat spinose (FIG. 1). Such fruitbody forms and hymenial types at one time formedthe basis for the classification of fungi. Thus thehymenochaetoid clade, as it is defined here, draws itsmembers from several families as circumscribed inpremolecular classifications: Agaricaceae, Polypora-ceae, Corticiaceae, Stereaceae and Hymenochaeta-ceae but includes only the type genus for the lastfamily name.

Micromorphological characteristics are exceedinglyvariable. Three basic kinds of hyphae involved inconstruction of basidiomycete basidiomata (viz. gen-erative hyphae, skeletal hyphae and binding hyphae)are present although most species have only thegenerative type. Spores are mainly smooth but vary inshape from the large globose ones found inGlobulicium hiemale to the extremely narrow andstrongly bent spores in Hyphodontia (Chaetoporellus)latitans. A few species have finely ornamented spores(viz. Coltriciella spp. and Hyphodontia (Rogersella)griseliniae).

Most species have some kind of vegetative (sterile)cells in the fruit body tissue, often sharing the spacewith the basidia in the hymenium (SUPPLEMENTARY

FIG. 1). They collectively could be called cystidia butbecause some of them have a distinctive form, uniqueterms have been introduced for them. The majority ofspecies in Hymenochaetaceae have a characteristickind of cystidia called setae (FIG. 1J). These thick-

Accepted for publication 23 October 2006.1 Corresponding author. E-mail: karl-henrik.larsson@dpes.gu.se

Mycologia, 98(6), 2006, pp. 926–936.# 2006 by The Mycological Society of America, Lawrence, KS 66044-8897

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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 withporoid 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 Hymenochaetecinnamomea. Bars: A, D 5 10 mm, B 5 2 mm; G–I 5 1 mm, J 5 10 mm.

LARSSON ET AL: HYMENOCHAETOID CLADE PHYLOGENY 927

walled, dark, brown and usually acutely pointed cellscan be observed with a hand lens and give the speciesa beard-stubble look. Their function is possibly toprotect the hymenium from insects. Thin-walled andhyaline cystidia characterize the hymenium in allspecies of Hyphodontia, many other small genera ofcorticioid fungi and some of the agaricoid andstereoid genera. Often these cystidia are pestle-shaped with a globular apex. The exact function isnot known but a general idea is that leptocystidiaemerging beyond the basidial layer function asexcretory organs because in living specimens theapex of such cystidia often are covered by a dropletthat disappears or dissolves when material is mountedfor observation microscopically. In some cases theapical droplet seems encased in a vesicle and resistsdissolution. The most well developed vesicle-bearingcystidia, called halocystidia, are found in Resiniciumbicolor and related species. Crystal-covered cystidia areother indications of excretion capacity. In Hyphodon-tia and Resinicium some species have lagenocystidiathat are hypha-like but with a needle-like termination.At the apex these cystidia carry a rosette of crystals,presumably composed of calcium-oxalate. Thin- tothick-walled cystidia with an apical crystal cap charac-terize species in the polypore genera Trichaptum andOxyporus and thick-walled, strongly encrusted cystidia(metuloids, lamprocystidia) can be seen in Hypho-derma puberum and a few other species. In Tubulicri-nis all species have lyocystidia, a hallmark of thegenus. These cystidia have a thick-walled ‘‘stem’’ anda thin-walled variously shaped apex. The thick-walledpart is usually more or less amyloid and dissolveseasily in 5% KOH. Gloeocystidia (enclosed cystidiawith more or less refractive contents) are notcommon but occur in Hyphoderma praetermissumand related species and in Physodontia. Hyphodermapraetermissum also is known for its stephanocysts.These are one- or two-celled hemispherical or globosestructures that occur on hyphae in the substrate, andthey are not always detectable in the basidiomata (forexcellent illustrations see Hallenberg 1990). Hallen-berg (1990) showed that stephanocysts can developon germinating spores, at least when they aredispersed on an artificial medium such as malt agar.Hallenberg concluded that stephanocysts were essen-tial for adsorption. On the other hand Tzean andLiou (1993) suggested that stephanocysts werenematode-catching organs and that the fungus usesnematodes as a nitrogen source. Related species havemorphologically similar but one-celled structurescalled echinocysts.

Basidium shape can be a useful taxonomicalcharacter. The corticioid genus Repetobasidium owesits name to the ability for repeated formation of new

basidia from the same apical cell and not, as usual,through hyphal branching (SUPPLEMENTARY FIG. 1).Each new basidium bursts through the old, emptybasidium leaving a progressively longer row ofsheathing basidia walls along the subtending hypha.Basidial repetition is not unique for Repetobasidium,although this is the genus where it first was observedand described (Eriksson 1958).

Ultrastructural characteristics as observed witha scanning or transmission electron microscope areof limited importance for the taxonomy and classifi-cation of higher fungi. The prime exception is theseptal pore apparatus that differs markedly amongvarious groups, and these anatomical differencescorrelate well with basidiomycete higher order classi-fications. Agaricomycetidae is characterized by a septalpore apparatus called a dolipore. In a dolipore theseptal pore is surrounded on each side of the septumby a half-dome-shaped membrane called parenthe-somes because in a TEM picture it looks like theseptum is placed within parentheses. Most homo-basidiomycetes have parenthesomes that are perfo-rated by a number of small openings and appear asdashed marks in the TEM; however a small number ofspecies instead have nonperforate parenthesomes.Because species in Auriculariales and Tulasnellaleshave the nonperforate type, this is regarded as theancestral condition from which the perforated de-veloped. In Hymenochaetales as well as in Cantha-rellales both types occur, which indicate that theevolution of parenthesome type is more complicatedthan initially understood.

Ecology.—Saprotrophy is the dominating life strategyin Hymenochaetales. Most species live in deadwoodand satisfy their energy needs by decaying thepolysaccharides cellulose, hemicellulose and lignin.When all these molecules are degraded at roughlyequal rates the resulting decay is called white-rot asopposed to brown-rot, which leaves most of the ligninintact (Rayner and Boddy 1988). The capacity tocause brown-rot is a derived condition that hasdeveloped repeatedly from white-rot ancestors onseveral occasions during evolution (Gilbertson 1980,Hibbett and Donoghue 2001). In Hymenochaetalesonly one case of suspected brown-rot is reported (viz.the gigantic polypore Bridgeoporus nobilissimus [Red-berg et al 2003]) but experimental data is lacking.

Many corticioid species with thin and inconspicu-ous basidiomata occur on strongly brown-rotted woodwhere most cellulose already is removed by a brown-rot fungus such as Fomitopsis pinicola (polyporoidclade). Examples from Hymenochaetales includeSphaerobasidium minutum, Tubulicrinis spp. Repetoba-sidium spp. and Hyphoderma involutum. The life

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strategy of such secondary rot fungi is not known, andalthough they live in close connection to brown-rotted wood it is too simplistic to classify them asactively performing brown-rot.

Several polyporoid Hymenochaetales genera, suchas Phellinus, Inonotus, Fomitiporia, Porodaedalea andTrichaptum, are strong primary decayers. Somecolonize living trees, thus blurring the distinctionbetween saprotrophic and parasitic strategies. Mostspecies invading living trees attack the dead tissue inthe center of stems (i.e. the heartwood) and thereforemay not directly harm the host, except to weaken thetrunks making them vulnerable to strong winds. Someheart-rot fungi (e.g. Inonotus obliquus on Betula spp.)can break through the sapwood and form blackcankers on stems.

Few species actually kill their host but these canbecome serious pathogens for forestry or urbanlandscaping (Inonotus ulmicola). Phellinidium weirii(; Phellinus weirii) causes laminated root rot inDouglas-fir and other conifers of western NorthAmerica. In infected stands there is a long-termimpact on stand structure and the fungus is one ofthe most important disturbance agents in Pacificconifer forests (Hansen and Goheen 2000). Anotherexample is Phellinus tremulae that occurs almosteverywhere aspen species grow and is reported to beable to spread through the sapwood. Among theHymenochaetales, species causing the greatest lossesto forestry are Porodaedalea pini (; Phellinus pini)growing on pines and Phellinus igniarius growing onvarious hardwoods. Both species destroy the heart-wood.

A distinct group of brightly colored to whitishHymenochaetales fruit directly on or in associationwith bryophytes. Basidiomata are either mushroom-like, with lamellae and central stipes (e.g. Rickenellaand Cantharellopsis) or more stereoid, with smooth towrinkled hymenia (e.g. Cyphellostereum and Cotylidia).Colonized bryophytes appear healthy, but it has beenshown that living rhizoids of mosses can be penetrat-ed by Rickenella fibula (Redhead 1981) and thatanother species, R. pseudogrisellum, forms claspingdigitate appresoria on the rhizoids of the liverwortBlasia (Redhead 1980, 1981), sometimes beingdispersed by infecting gemmae of Blasia (Redhead1980) together with a symbiotic Nostoc (Redheadunpubl). In general it is not known whether theseHymenochaetales parasitize their host or whether theconnection is of a more mutualistic nature. Separa-tion of the Rickenella subclade from the Agaricales ingeneral where Omphalina and Gerronema are placed,and interspersion of other taxa, led to the recognitionof several small agaric genera (Cantharellopsis, Con-tumyces, Loreleia and Sphagnomphalia, which more

correctly is named Gyroflexus) for agaricoid species(Redhead et al 2002).

Several species forming corticioid basidiomatafrequently contain one-celled green algae in the basalbasidioma layer. Examples from Hymenochaetalesinclude Resinicium bicolor and Globulicium hiemale.The algal connection in Resinicium was studied byPoelt and Julich (1969) but they were unable toestablish any direct hyphal invasion of algal cells.However they noted a more proliferous hyphalbranching close to the algae. Similar connectionswith algae are known also in other homobasidiomy-cete orders but a direct parasitism of algae is knownonly with certainty in the corticioid genus Athelia(Poelt and Julich 1969). At least one Hymenochae-tales species is lichenized (Palice et al 2005). It bearsthe curious name Omphalina foliacea and was de-scribed based on sterile thalli only. Although lackingbasidiomes it was placed in Omphalina with otherlichenized omphalinoid agarics currently classified asLichenomphalia in the Agaricales. The lichenizedfungus, which is not necessarily agaricoid, lacksa unique generic name.

Two more life strategies should be mentionedbriefly. Danielson (1983) studied the ectomycorrhizaformed by Pinus banksiana (jack pine) both in vivoand in vitro. One of the supposed symbionts was thestipitate poroid Coltricia perennis, which forms basi-diomata on dry sandy forest soils. Danielson was ableto synthesize a mycorrhizal association with pineseedlings in the laboratory and even managed to getbasidiomata. However living mycorrhizal root tipsformed with Coltricia have not been detected innature. Umata (1995) studied the ability of aphyllo-phoralean fungi to induce germination in seeds ofthe achlorophyllous orchid Galeola altissima. Hefound that several wood-decaying fungi, includingPhellinus sp., induced germination in the laboratory,but for only one of these fungi, Erythromyces crocicreas,has a connection to orchids been demonstrated in thewild.

The nematode-capturing ability established forstephanocyst and echinocyst producing Hyphodermaspecies is yet another nutritional mode shown byspecies in Hymenochaetales (Tzean and Liou 1993).Stephanocysts and echinocysts are covered by anadhesive mucilage and attach easily to the nematodecuticle. Captured nematodes are killed and the bodiespenetrated by hyphae. Tzean and Liou (1993) testeda number of corticioid fungi for nematode-destroyingcapacity. All species with stephanocysts and echino-cysts could kill nematodes but a number of otherHyphoderma species lacking these structures alsoseemed to kill nematodes by being toxic. Nematodesfeeding on hyphae from the latter group of

LARSSON ET AL: HYMENOCHAETOID CLADE PHYLOGENY 929

Hyphoderma species died within 2 h. The fungus thenproduced hyphae that coiled around the nematodeand penetrated the body. Toxic Hyphoderma speciesare not related to those carrying the specializednematode-catching organs. Only the latter groupbelongs to Hymenochaetales while the toxic speciesbelong to the polyporoid clade (Hibbett and Thorn2001). A nematode-killing feeding behavior also hasdeveloped independently among Pleurotus and Ho-henbuehelia species in Agaricales (Thorn et al 2000).

Economic importance.—The enzymes and secondarymetabolites produced by fungi have received consid-erable interest for their potential use as drugs or forbiotechnological applications. Several species ofPhellinus and Inonotus are used in Asian folkmedicine and the products are commercially avail-able. One example is Phellinus baumii (often errone-ously called Phellinus linteus) that is known for its usein traditional Chinese medicine (Ying et al 1987) andin several countries marketed as a drug againstcancer, diabetes and toxicity (Shon et al 2003).Phellinus rimosus is reported as a drug used by tribesin Kerala in India (Ajith and Janardhanan 2003).Indigenous people in Siberia use chaga as a cleansingand disinfecting substance but the same substancealso has been used against liver and heart ailmentsand in cancer therapy. Chaga is produced fromInonotus obliquus and commercially available. Thereis a rich scientific literature reporting the identifica-tion of substances and the effects they may have.

The cellulolytic and ligninolytic enzymes producedby wood-decaying fungi have been studied intensivelywith the aim of bringing them into practical use in thepulp and paper industry or for cleaning industrialwaste products. One of the best studied modelorganism is Phanerochaete chrysosporium that belongsin the polyporoid clade (Hibbett and Thorn 2001),but some interest also has been devoted to species ofHymenochaetaceae (Wesenberg et al 2003).

History and classification.—Hymenochaetales was in-troduced by Oberwinkler 1977. His circumscriptionwas more or less the same as for Hymenochaetaceaeby Donk (1964) and Patouillard (1900) who recog-nized what he called Serie des Igniaires. Thecharacters emphasized by Patouillard and his succes-sors were the uniformly brown hyphae and basidio-mata, the simple-septate hyphae, the setae, theblackening of tissue when mounted in KOH (xantho-chroic reaction) and the association with white rot.Donk included, with some hesitation, the corticioidgenera Vararia and Asterostroma and the clavarioidLachnocladium because he regarded the dicho- andasterohyphidia occurring in these taxa as modifiedsetae. However Oberwinkler (1977) noted that other

characters (spore amyloidity, gloeocystidia) pointedthese latter three genera in the direction of Russulalesand molecular phylogenetic studies have confirmedthat conclusion (Larsson and Larsson 2003). Donk(1964) also included Phaeolus schweinitzii in Hyme-nochaetaceae although it has no setae and isassociated with a brown rot. Parmasto and Parmasto(1979) indicated that its brown pigment is differentfrom the hymenochaetoid fungi and that a xantho-chroic reaction is not specific for Hymenochaetaceae.Molecular data confirm that Phaeolus does not belongto Hymenochaetales but rather has its place in thevicinity of Laetiporus in Polyporales (Binder et al2005).

Already the first comprehensive molecular study ofthe homobasidiomycetes indicated that Hymenochae-tales sensu Oberwinkler should be interpreted morebroadly (Hibbett and Donoghue 1995). The samplingof 62 mainly polyporoid taxa showed that Oxyporusand Trichaptum were closely related to Hymenochae-tales despite lacking setae, xanthocroic reaction,brownish hyphae and, in the case of Trichaptum,having nodose-septate hyphae. However Trichaptumwas known to have imperforate parenthesomes, whichpointed to a relationship with Hymenochaetales andalso set the dolipore morphology in focus for furtherstudy. Langer and Oberwinkler (1993) already hadascertained that several corticioid species (viz. Hypho-dontia spp., Basidioradulum radula and Schizoporaparadoxa) have imperforate parenthesomes. In a sub-sequent molecular investigation (Hibbett et al 1997)these three genera were included and found tocluster with Hymenochaetales, Oxyporus and Trichap-tum. However the strain of ‘‘Hyphodontia alutaria’’(GEL 2071) used as a DNA source actually representsResinicium bicolor (cf. Binder et al 2005).

Hibbett and Thorn (2001) provided the firstdescription of what here is called Hymenochaetales(as hymenochaetoid clade) taking into account all thenew results received from molecular phylogeneticstudies. Moncalvo et al (2002) showed that speciesfrom the stipitate stereoid genus Cotylidia and theagaricoid genera Cantharellopsis, Omphalina andRickenella also had their place in or close toHymenochaetales. Redhead et al (2002) reclassifiedthese agarics in the Hymenochaetales. Larsson et al(2004) provided a second overview of the group anda third was published recently (Binder et al 2005),both with an emphasis on corticioid taxa.

The Hymenochaetales in its original sense (i.e.Phellinus, Inonotus, Hymenochaete and related genera,with setae, xanthochroic reaction, simple-septatehyphae etc.) here will be referred to as Hymenochae-taceae. The family includes close to 400 species. Theyare found in all parts of the world, and because all

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species form quite conspicuous basidiomata they havebeen extensively collected and studied. Recentmorphological descriptions and keys to the poroidtaxa can be found in Gilbertson and Ryvarden (1986,1987), Larsen and Cobb-Poulle (1990), Nunez andRyvarden (2000) and Ryvarden (2004). Hymenochaeteand related genera with a nonporoid hymenophoreare treated by Leger (1998) and Parmasto (2001,2005).

Species in Hymenochaetaceae contain a group oforganic compounds called styrylpyrones. Similarcompounds also are known from various plantfamilies and they probably form part of a defenseagainst infections and browsing. The distribution ofstyrylpyrones within Hymenochaetaceae has beenused in the classification of the group (Fiasson1982, Fiasson and Bernillon 1983). Chemical char-acters together with detailed morphological studiesprompted Fiasson and Niemela (1984) to accept 10genera for the poroid species in Europe as a re-placement for the prevailing classification with onlytwo genera: Inonotus for monomitic and annualspecies and Phellinus for species with dimitic andperennial basidiomata.

Hymenochaete originally was introduced for specieswith effused to effused-reflexed basidiomata anda smooth hymenophore. For similar taxa witha hydnoid hymenophore the artificial genus Hydno-chaete is available. Two neotropical species withstipitate basidiomata and a smooth hymenophorewere placed in Stipitochaete and those with a clavarioidbasidioma in Clavariachaete. There are several othergenera within Hymenochaetaceae that are morpho-logically distinct and therefore kept separate by mostauthors. The laterally stipitate genus Cyclomycescarries its name because the type species hasa concentrically lamellate hymenophore. The samehymenophore configuration sometimes can be foundalso in specimens of Coltricia montagnei whileneighboring fruit bodies may be strictly poroid.Onnia and Coltricia species have stipitate basidiomataand a poroid hymenophore, but the latter genus isregarded as distinct because setae are lacking. This isalso true for Aurificaria that differs by an olivaceousdiscoloring of the basidiospores in KOH. Setae alsoare lacking in Coltriciella but its spores are finelyornamented, quite unique within Hymenochaeta-ceae. Finally Pyrrhoderma has laterally stipitate basi-diomata with the cap covered by a shiny crust.

The phylogeny of Hymenochaetaceae has beenthoroughly studied by molecular methods (e.g.Wagner and Fischer 2001, 2002a, b). These studiesindicated that Hymenochaetaceae, as it formerly wascircumscribed, was not a monophyletic group. Somecorticoid species (viz. Basidioradulum radula and

Hyphodontia quercina) and two polypore genera,Schizopora and Trichaptum, were intermixed amongtypical hymenochaetoid species. On the other handthe molecular data gave support to the work ofFiasson and Niemela (1984) who divided the poroidgenera Inonotus and Phellinus in several smallergroups based on morphology and chemical charac-ters. The new classification for the poroid species is ingeneral well supported by morphological, physiolog-ical and ecological characters but does not supporta division between monomitic annual taxa anddimitic perennial ones.

The situation within Hymenochaete and othergenera with a smooth or hydnoid hymenophoreappears less resolved when molecular data areanalyzed. Wagner and Fischer (2002b) found thatall species they sampled belonged to one clade exceptHymenochaete tabacina that clustered among some ofthe poroid species. Consequently a new genus,Pseudochaete, was introduced (Wagner and Fischer2002b).

The mushroom-forming species in Hymenochae-tales are all of the omphalinoid type (viz. small fruitbodies with rather thick, shallow and strongly de-current lamellae). The first hint that not all ompha-linoid agarics belonged to the same clade came ina paper on the phylogeny of agaric fungi (Moncalvoet al 2000) and a subsequent paper with a broadersampling, which established that some species aremembers of the hymenochaetoid clade (Moncalvo etal 2002). In a companion paper Redhead et al (2002)studied the phylogeny and classification of thesehymenochaetoid agarics and showed that they all areassociated with bryophytes. The last two studies alsowere the first to expose that the stipitate stereoidgenus Cotylidia seemed related to the omphalinoidspecies in Hymenochaetales. Cotylidia formerly wasclassified together with other stipitate stereoid speciesin Podoscyphaceae. Now it has been shown thatPodoscypha has its place in the clade called polyporoidby Hibbett and Thorn (2001) and far removed fromCotylidia (Kim and Jung 2000).

Hymenochaetales includes a number of specieswith thin, effused basidiomata and with a smooth tohydnoid hymenophore. Such fungi traditionally havebeen placed in a single family Corticiaceae, widelyacknowledged as an artifical family (Donk 1964,Julich 1982). Julich (1982) published a comprehen-sive classification for corticioid fungi using morphol-ogy only. The corticioid taxa included in Hymeno-chaetales are drawn from four of the orders in Julich’sclassification, which further emphasizes the difficul-ties that have plagued all attempts to fit corticioidfungi into a morphology-based classification. It wasnot until DNA characters became available that the

LARSSON ET AL: HYMENOCHAETOID CLADE PHYLOGENY 931

phylogeny of corticioid fungi could be investigatedreliably. Now we know that corticoid fungi occur inevery major homobasidiomycete clade (Larsson et al2004, Binder et al 2005).

Hyphodontia is the largest genus of corticioid fungiin Hymenochaetales with ca. 90 species. Descriptionsand keys to most Hyphodontia species are found inLanger (1994).

MATERIALS AND METHODS

ITS and nLSU sequences available at GenBank werecompiled and complemented with sequences generated forthis study. The GenBank sequences originate from a numberof studies, not all of them published. In many cases one labhas deposited a sequence from one of the regions in questionwhile another lab has contributed another, both beinglabeled with the same species name but generated fromdifferent specimens. We have combined sequences fromdifferent sources when they are identified as conspecific. Wetook a calculated risk that a compilation trusting the namesattached to sequences might introduce ambiguities as tosequence homogeneity. We consider that risk to be out-weighed by the advantages of a denser sampling and a datamatrix with fewer missing data. The dataset was trimmed toremove duplicates, unidentified specimens and some se-quences that were deemed too short to give a reliable signal.After preliminary analyses (not shown) a number ofsequences from densely sampled subclades within Hymeno-chaetaceae (e.g. Inonotus and Phellinus) were removed toreceive a more manageable dataset without sacrificingresolution at and above the genus level. Thirty-fivesequences, mainly of corticioid species, have not beenpublished before. Two species from Cantharellales (Sisto-trema) and two species from Auriculariales (Exidiopsis,Protodontia) are included as outgroup. All information onspecimens made use of in this study along with GenBankaccession numbers is available in SUPPLEMENTARY TABLE I.

The dataset included 174 ingroup sequences and 1552nucleotide positions including introduced gaps. HoweverITS1 and ITS2 were deemed too variable and were excludedfrom analyses together with several variable regions withinLSU. Protocols for DNA extraction, PCR and sequencingfollowed Larsson and Larsson (2003) and Larsson et al(2004).

Heuristic maximum parsimony analyses were performedwith PAUP*4.0 (200 random taxon addition replicates,keeping 50 trees per replicate, MAXTREES 5 15 000). Theanalysis used 1025 characters of which 529 were constant,135 variable but parsimony uninformative and 361 parsi-mony informative. Branch support was estimated withnonparametric bootstrapping as implemented in PAUP*4.0(100 replicates, 10 random addition sequences per repli-cate, keeping 50 trees per replicate, MAXTREES 5 15 000).

Bayesian inference of phylogeny was performed withMrBayes 3.0B4 (Ronquist and Huelsenbeck 2003). MrMo-delTest 2.2 (Nylander 2004) was used to estimate separatebest-fit models of evolution for 5.8S and LSU. A heteroge-neous Bayesian inference was set up with model parameters

estimated separately for each partition. Eight Metropolis-coupled Markov chain Monte Carlo (MCMCMC) chainswith a temperature of 0.2 C were initiated; these were run3 000 000 generations with tree and parameter samplingevery 1500 generations (2000 trees). The initial burn-in wasset to 50% (1000 trees). A 50% majority-rule consensuscladogram was computed from the remaining trees; theproportions of this tree correspond to Bayesian posteriorprobabilities (BPP).

RESULTS AND DISCUSSION

MP analysis of the nLSU+5.8S dataset resulted in 200equally parsimonious trees with little support fordeeper nodes. The MP tree file with all shortest treesand the bootstrap tree file are available as supple-mentary material. The models GTR+I+G (LSU) andSYM+G (5.8S) were supported as the best fit modelsfor the two data partitions and employed in MrBayesanalyses. Chain convergence was attained well aheadof the initial burn-in threshold and chain mixing wasfound to be satisfactory. A 50% majority-rule consen-sus cladogram with Bayesian posterior probabilities(FIG. 2); branch lengths reflect estimated number ofchanges per site.

Bayesian inference produced a fairly well resolvedtree with high posterior probability values for severalclades. Major monophyletic clades are discussed brieflybelow in alphabetical order as indicated on the tree.

(A) Oxyporus clade.—Oxyporus is a genus of white-rotting polypores that often attack living trees of bothhardwoods and conifers. Mostly it is only theheartwood that is decayed but at least one speciesalso can invade the sapwood, which of course isdetrimental for the tree. Bridgeoporus nobilissimuspreviously was classified in Oxyporus but recentlysegregated because it is suggested to be a brown rotagent (Burdsall et al 1996). The phylogenetic positionof Bridgeoporus within Hymenochaetales was notinvestigated here.

(B) Rickenella clade.—This group is a mixture ofspecies with effused, stipitate stereoid and stipitatelamellate basdiomata. No morphological charactersare diagnostic of the group together, but it isinteresting to note that nutritional modes includevarious interactions with other living organisms(association with bryophytes, association with greenalgae and predation on nematodes, all of which couldserve as nitrogen sources). Within the larger clade aresome well defined groups and at least three of themcan be recognized by morphology. Resinicium ina restricted sense emerges as a well supported genuscharacterized by large halocystidia. Hyphodermapraetermissum together with related nematode-catch-

932 MYCOLOGIA

FIG. 2. Phylogenetic relationships of Hymenochaetales inferred from 5.8S and nucLSU rDNA sequences with Bayesiananalysis. A 50% majority rule consensus cladogram Bayesian posterior probabilities $ 0.94 are shown above internodes; branchlengths reflect estimated number of changes per site. Closed horizontal parentheses indicate that the species has doliporeswith continuous parenthesomes. Broken horizontal parentheses indicate presence of the perforated parenthesome type. A.Oxyporus clade. B. Rickenella clade. C. Kneifiella clade. D. Hyphodontia clade. E. Coltricia clade. F. Hymenochaetaceae clade.

LARSSON ET AL: HYMENOCHAETOID CLADE PHYLOGENY 933

ing species are also supported as monophyletic. Forthis group the name Peniophorella is available (Lars-son in press). The Skvortzovia subclade unites severalcorticioid species that until now have been placed indifferent genera. Burdsall and Nakasone (1981)pointed out the similarities exhibited by cultures ofMycoacia meridionalis and Odontia furfurella, andNakasone (1990) placed the two together in Resini-cium. Hjortstam and Bononi (1987) erected Skvortzo-via to encompass Odontia furfurella. It seems appro-priate to refer also the other species to the samegenus. All have small hymenial cystidia with an apicalcap of exudated material. Leifia, Odonticium andRepetobasidium are three corticioid genera thatmorphologically do not seem to have anything incommon, and the possibility that the group is anartifact caused by the analysis must be considered.Odonticium romellii (type species) has an odontioidhymenium, thick-walled simple-septate hyphae andnarrowly allantoid spores. Zmitrovich (2001) recentlysuggested a connection with Leifia flabelliradiata andalso made a combination to Odonticium. Althoughthe agaricoid taxa now placed in the Hymenochae-tales appeared to be congeneric in an early analysis(Lutzoni 1997) and only later were suggested torepresent several genera (Redhead et al 2002), it isonly after combining data on additional taxa (Rick-enella fibula and Cyphellostereum laeve, Larsson et al2004) and other corticioid taxa in the current analysisthat the diversity is revealed further. Each of thegenera stands alone save for Rickenella, and there theappresoria-forming Blasia parasite, R. pseudogrisella,appears to be separable from other Rickenella thatpenetrate rhizoids directly and therefore should bereclassified as is here proposed:

Blasiphalia Redhead, gen. nov., a Rickenello appresso-riis praesentibus differt. Differs from Rickenella bypresence of appresoria. Etymology: Latinized non-sense word from fragments of Blasia and Omphalia(f.). Type: Blasiphalia pseudogrisella (AH Sm)Redhead.

Blasiphalia pseudogrisella (AH Sm) Redhead, comb.nov.; basionym: Mycena pseudogrisella AH Sm,North American Species of Mycena, p 124, 1947.

(C) Kneiffiella clade.—Most species in this cladeeither have long, tubular cystidia originating in thesubiculum (pseudocystidia; SUPPLEMENTARY FIG. 1E)or thin-walled, tubular hymenial cystidia. The groupseems quite natural and was recovered also in the MPstrict consensus tree but generated no significantbootstrap support. Kneiffiella is an old genus nameavailable for this group and most combinations arealready in place (Julich and Stalpers 1980).

(D) Hyphodontia clade.—The Hyphodontia speciesin this clade includes the type species, H. pallidula.Hyphodontia sensu stricto apparently will becomea quite small genus with 5–6 closely related speciesonly. They all are characterized by septate cystidia incombination with lagenocystidia (SUPPLEMENTARY FIG.1I). With its spectacular lyocystidia, Tubulicrinis ismorphologically well defined but in this analysis thegenus is split in two groups. The phylogeny presentedhere is certainly not the final word on the composi-tion of the Hyphodontia clade.

(E) Coltricia clade.—This clade holds a mixture ofcorticioid species and two genera that earlier wereclassified with Hymenochaetaceae. A sample of Hypho-dontia species form two clades, a weakly supported onecentered on Hyphodontia aspera and a moderatelysupported clade that includes H. crustosa, H. sambuciand H. pruni. The latter group also includes Pyr-rhoderma adamantinum. This is a stipitate poroidspecies that usually is placed in Hymenochaetaceae,and the phylogenetic position shown here needsconfirmation. All Hyphodontia species have varioustypes of hymenial, little differentiated and oftencapitate cystidia. Coltricia and Coltriciella form a strong-ly supported clade. These genera of stipitate polyporesshow most of the traits that characterize Hymenochae-taceae but they mostly lack setae. However amongAsian and South American species of Coltricia thereare species with setae or setal hyphae (viz. C. hamata,C. duostratosa, C. tomentosa and C. vallata). Coltricielladiffers clearly from Coltricia by finely ornamentedspores but the phylogenetic analysis gave no supportfor a separation in two genera.

(F) Hymenochaetaceae clade.—The Hymenochaeta-ceae in its traditional sense is not supported asmonophyletic in our analysis. This is in accordancewith the results received by Wagner and Fischer(2002b). However the Bayesian inference supportsa monophyletic Hymenochaetaceae that excludesonly Coltricia and Coltriciella. The subdivision ofPhellinus and Inonotus into smaller genera in generalis supported strongly here. Exceptions include Onnia,Phellinidium and Pseudoinonotus and these generamight have to be revised. The segregation ofPseudochaete from Hymenochaete also is supportedwhile genera erected on account of a specific fruitbody type or hymenium configuration (Stipitochaete,Hydnochaete) are not.

In molecular analyses with a homobasidiomycete-wide sampling the hymenochaetoid clade has re-ceived mainly low or moderate support values (e.g.Hibbett et al 1997, Larsson et al 2004, Binder et al2005). Binder and Hibbett (2002) managed to raisebootstrap values to 95% when four gene regions and

934 MYCOLOGIA

three representatives for the clade were included. Thefour genes studied (nuclear SSU and LSU, mitochon-drial SSU and LSU) also were analyzed separately andin combinations of two and three genes. The datashows that most phylogenetic signal seems to emanatefrom the mitochondrial genes and especially the SSUregion. Future phylogenetic investigations in Hyme-nochaetales should take advantage of that result.

No unequivocal morphological synapomorphiessupport Hymenochaetales, and the order presentlycan be defined only in terms of molecular data. Theoccurrence of dolipores with continuous parenthe-somes and the possibility that this structure isa synapomorphy for Hymenochaetales have gainedconsiderable interest. However Hyphoderma praeter-missum has perforate parenthesomes (Langer andOberwinkler 1993, Keller 1997). The tree topology(FIG. 2) still indicates that continuous parenthesomesmight define a monophyletic group consisting ofclades C–F. Further exploration of septal ultrastruc-ture for species here referred to clades A and B isdesirable.

ACKNOWLEDGMENTS

We thank Ellen Larsson and Henrik Nilsson for great helpwith lab work and phylogenetic analyses respectively. K-HLarsson was financially supported by the Swedish SpeciesInformation Centre, Swedish Agricultural University, Upp-sala. We also acknowledge support from NSF 0090301,Research Coordination Network: A Phylogeny for KingdomFungi to M. Blackwell, J.W. Spatafora and J.W. Taylor.

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936 MYCOLOGIA

SUPPLEMENTARY FIG. 1. Microscopic characters in Hymenochaetales. A, Tubulicrinis accedens, lyocystidium. B, Hymenochaetecinnamomea, hymenial seta. C, Inonotus cuticularis, branched and hooked seta from the upper pileus surface. D, Tubulicrinissubulatus, lyocystidium. E, Hyphodontia subalutacea, pseudocystidium originating in the subiculum. F, Resinicium granulare,halocystidium with partly collapsed halo. G, Hyphoderma puberum, thickwalled, encrusted cystidium (metuloid). H,Repetobasidium vestitum, basidia with rows of remnants of old basidia walls anda capitate hymenial cystidium. I, Hyphodontiaarguta, lagenocystidia with apical incrustation. Scale bar 5 10 mm.

SUPPLEMENTARY FIG 2. Maximum parsimony tree for Hymenochaetales with bootstrap values indicated.

SUPPLEMENTARY TABLE I. GenBank numbers for all sequences used in analyses

Species ITS1 5.8S ITS2 Nuc LSU

Asterodon ferruginosum AY463380 AY586631Aurificaria luteoumbrina AY059033Coltricia cinnamomea AF311003Coltricia montagnei AY039683Coltricia perennis DQ234559 DQ234559 DQ234559 AF287854Coltriciella dependens AY059059Coltriciella navispora AY059062Coltriciella oblectabilis AY059061Coltriciella pusilla AY059060Cyclomyces fuscus AF385163Cyclomyces tabacinus AF385164Fomitiporella caryophylli AY558611 AY558611 AY558611 AY059021Fomitiporella cavicola AY059052Fomitiporia hartigii AY558621 AY558621 AY558621 AF311005Fomitiporia hippophaeicola AY558622 AY558622 AY558622 AF311006Fomitiporia mediterranea AY854080 AY854080 AY854080 AY684157Fomitiporia punctata AF515563 AF515563 AF515563 AF311007Fomitiporia robusta AY558645 AY558645 AY558645 AF311008Fulvifomes fastuosus AY558615 AY558615 AY558615 AY059057Fulvifomes kawakamii AY059028Fulvifomes nilgheniensis AY558633 AY558633 AY558633 AY059023Fulvifomes robiniae AF411825Fuscoporia ferrea AY558617 AY558617 AY558617 AF311030Fuscoporia ferruginosa AY189700 AY189700 AY189700 AF311032Fuscoporia gilva AF250932 AF250932 AF250932 AF518636Fuscoporia torulosa AY558649 AY558649 AY558649 AF311041Fuscoporia viticola AY558653 AY558653 AY558653 AY885166Hydnochaete duportii AY635770Hydnochaete japonica AY558596 AY558596 AY558596 AF385153Hydnochaete olivacea AY293185Hymenochaete acanthophysata AF385144Hymenochaete adusta AY558594 AY558594 AY558594 AF385161Hymenochaete carpatica AF385158Hymenochaete cervinoidea AF385157Hymenochaete cinnamomea AY463416 AY586664Hymenochaete corrugata AF518620Hymenochaete cruenta AF385152Hymenochaete denticulata AY558595 AY558595 AY558595 AF385155Hymenochaete fuliginosa AF385154Hymenochaete pinnatifida AF385149Hymenochaete rhabarbarina AJ406468Hymenochaete rubiginosa AY463417 AY586665Hymenochaete separabilis AF385146Inocutis dryophilus AF311012Inocutis jamaicensis AY072029 AY072029 AY072029 AY059048Inocutis rheades AF237731 AF237731 AF237731 AF311019Inocutis tamaricis AY558604 AY558604 AY558604 AF311021Inonotopsis subiculosus AF311020Inonotus anderssonii AY558599 AY558599 AY558599 AY059041Inonotus baumii AF200230 AF200230 AF200230 AY059058Inonotus cuticularis AF237730 AF237730 AF237730 AF311010Inonotus glomeratus AF247968 AF247968 AF247968 AY059032Inonotus hispidus AY558602 AY558602 AY558602 AF518623Inonotus linteus AF153010 AF153010 AF153010 AF458461Inonotus obliquus AY251310 AY251310 AY251310 AY279001

Species ITS1 5.8S ITS2 Nuc LSU

Inonotus pachyphloeus AY558635 AY558635 AY558635 AY059020Inonotus porrectus AY558603 AY558603 AY558603 AY059051Inonotus quercustris AY072026 AY072026 AY072026 AY059050Inonotus tropicalis AY641432 AY641432 AY641432 AY598826Inonotus weirianus AY558654 AY558654 AY558654 AY059035Mensularia hastifer AF311013Mensularia nodulosa AF311016Mensularia radiata AF237732 AF237732 AF237732 AF311018Onnia leporina AF311022Onnia tomentosa AY558607 AY558607 AY558607 AF311023Onnia triqueter AF311024Phellinidium ferrugineofuscum AF311031Phellinidium fragrans AY558619 AY558619 AY558619 AY059027Phellinidium pouzarii AF311039Phellinidium sulphurascens AY829344 AY829344 AY829344 AY059016Phellinidium weirii AY829342 AY829342 AY829342 AY829346Phellinus bicuspidatus AY189699 AY189699 AY189699 AY059022Phellinus cinereus AY340048 AY340048 AY340048 AF311027Phellinus conchatus AY558614 AY558614 AY558614 AF311028Phellinus ingniarius AF110991 AF110991 AF110991 AY839834Phellinus (Fuscoporia) johnsonianus AF250931 AF250931 AF250931 AF458458Phellinus laevigatus AF053226 AF053226 AF053226 AF311034Phellinus occidentalis AY558634 AY558634 AY558634 AY059019Phellinus populicola AY558638 AY558638 AY558638 AF311038Phellinus (Inonotus) rhabarbarinus AY558642 AY558642 AY558642 AF458466Phellinus spiculosus AY189702 AY189702 AY189702 AY059055Phellinus tremulae AY340064 AY340064 AY340064 AF311042Phellinus tuberculosus AY558652 AY558652 AY558652 AF311043Phellopilus nigrolimitatus AY558632 AY558632 AY558632 AF311036Phylloporia chrysita AF411821Phylloporia ephedrae AF411826Phylloporia pectinata AF411823Phylloporia ribis AY558643 AY558643 AY558643 AF311040Porodaedalea canchriformans AF200242 AF200242 AF200242 AY059029Porodaedalea chrysoloma AY189706 AY189706 AY189706 AF311026Porodaedalea niemelaei AY059054Porodaedalea pini AY558636 AY558636 AY558636 AF311037Pseudochaete tabacina AY558598 AY558598 AY558598 AF385145Pseudoinonotus chondromyelus AF311009Pseudoinonotus dryadeus AY558601 AY558601 AY558601 AF311011Pyrrhoderma adamantinum AY059031Pyrrhoderma scaurum AY059030Stipitochaete damicornis AF385162Blasiphalia pseudogrisella U66437 U66437 U66437 U66437Cantharellopsis prescotii AF261461Contumyces rosella U66452 U66452 U66452 U66452Cotylidia aurantiaca AF261460Cotylidia aurantiaca var alba AF261458Cotylidia diaphana AF261459Cotylidia sp. AY854079 AY854079 AY854079 AY629317Cyphellostereum laeve AY745705Gyroflexus brevibasidiatus U66441 U66441 U66441 U66441Loreleia marchantiae U66432 U66432 U66432 U66432Rickenella fibula DQ241782 DQ241782 DQ241782 AY700195Rickenella mellea U66438 U66438 U66438 U66438Atheloderma mirabile DQ873592 DQ873592 DQ873592

SUPPLEMENTARY TABLE I. Continued

Species ITS1 5.8S ITS2 Nuc LSU

Athelopsis lunata DQ873593 DQ873593 DQ873593 DQ873593Basidioradulum radula DQ234537 DQ234537 DQ234537 AY700184Fibricium rude AY700202Globulicium hiemale DQ873595 DQ873595 DQ873595 DQ873595Hyphoderma echinocystis DQ677494 DQ681200Hyphoderma guttuliferum AY463419 AY586667Hyphoderma praetermissum DQ873597 DQ873597 DQ873597 DQ873597Hyphoderma puberum DQ873599 DQ873599 DQ873599 DQ873599Hyphoderma capitatum DQ677491 DQ677491 DQ677491Hyphoderma orphanellum DQ677500 DQ677500 DQ677500Hyphoderma sibiricum DQ677503 DQ677503 DQ677503Hyphodontia abieticola DQ873601 DQ873601 DQ873601 DQ873601Hyphodontia alienata AY466401 AY466401 AY586727Hyphodontia alutacea AJ406453Hyphodontia alutaria DQ873603 DQ873603 DQ873603 DQ873603Hyphodontia arguta DQ873605 DQ873605 DQ873605Hyphodontia aspera DQ873606 DQ873606 DQ873606 DQ873607Hyphodontia barbajovis DQ873608 DQ873608 DQ873608 DQ873609Hyphodontia borealis AY463429 AY586677Hyphodontia breviseta DQ873612 DQ873612 DQ873612 DQ873612Hyphodontia cineracea AJ406450Hyphodontia crustosa DQ873614 DQ873614 DQ873614 DQ873614Hyphodontia curvispora DQ873615 DQ873615 DQ873616Hyphodontia detritica DQ677507 DQ677507 DQ677507Hyphodontia floccosa DQ873618 DQ873618 DQ873618 DQ873618Hyphodontia hastata DQ873619 DQ873620Hyphodontia nespori DQ873622 DQ873622 DQ873622 DQ873622Hyphodontia niemelaei AJ406462Hyphodontia nudiseta AJ406460Hyphodontia pallidula AJ406594Hyphodontia paradoxa AF145571 AF145571 AF145571 AY059067Hyphodontia pruni DQ873624 DQ873624 DQ873624 DQ873625Hyphodontia quercina AY463430 AY586678Hyphodontia radula AF145570 AF145570 AF145570 AJ406466Hyphodontia rimosissima DQ873627 DQ873627 DQ873627 DQ873628Hyphodontia sambuci AJ406461Hyphodontia serpentiformis AJ406465Hyphodontia subalutacea DQ873633 DQ873633 DQ873634Hyphodontia sp. DQ873633 DQ873633 DQ873633 DQ873634Leifia flabelliradiata DQ873635 DQ873635 DQ873635Mycoacia pinicola DQ873637 DQ873637 DQ873637 DQ873637Odonticium romellii DQ873639 DQ873639 DQ873639Oxyporus corticola DQ873641 DQ873641 DQ873641 DQ873641Oxyporus latemarginatus AF163047 AF163047 AF163047Oxyporus populinus AJ406467Palifer verecunda DQ873642 DQ873642 DQ873642 DQ873643Phlebia georgica DQ873645 DQ873645 DQ873645 DQ873645Repetobasidium conicum DQ873647 DQ873647 DQ873647 DQ873647Repetobasidium mirificum AY293208Resinicium bicolor AY463463 AF518645Resinicium chiricahuaensis DQ863692Resinicium friabile DQ863690Resinicium furfuraceum DQ873648 DQ873648 DQ873648 DQ873648Resinicium meridionalis DQ863693Resinicium saccharicola DQ863691Rogersella griselinae DQ873651 DQ873651 DQ873651 DQ873651

SUPPLEMENTARY TABLE I. Continued

Species ITS1 5.8S ITS2 Nuc LSU

Skvortzovia furfurella DQ873649 DQ873649 DQ873649Sphaerobasidium minutum DQ873652 DQ873652 DQ873653Trichaptum abietinum AF347104 AF347104 AF347104Tubulicrinis globisporus DQ873655 DQ873655 DQ873655 DQ873655Tubulicrinis gracillimus AF518661Tubulicrinis hirtellus DQ873657 DQ873657 DQ873657 DQ873657Tubulicrinis inornatus DQ873659 DQ873659 DQ873659Tubulicrinis subulatus AY463478 AY586722Sistotrema brinkmannii X X X XSistotrema resinicystidium X X X XExidiopsis calcea AY463406 AY586654Protodontia piceicola DQ873660 DQ873660 DQ873660 DQ873660

SUPPLEMENTARY TABLE I. Continued