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Clark University, Biology Department, 950 Main Street, Worcester, Massachusetts 01610
Else C. Vellinga
Department of Plant and Microbial Biology, University of California at Berkeley, Berkeley, California 94720-3102
Neale L. Bougher
Department of Environment and Conservation, Western Australian Herbarium, Locked Bag 104, Bentley Delivery Centre, Perth, Western Australia 6983, Australia
Oluna Ceska
P.O. Box 8546, Victoria, British Columbia, V8W 3S2, Canada
Pierre-Arthur Moreau
Laboratoire de Botanique, Faculté de Sciences Pharmaceutiques et Biologiques, 3 rue du Professeur Laguesse, BP 83, F-59006 Lille Cédex, France
Maria Alice Neves
Institute of Systematic Botany, New York Botanical Garden, Kazimiroff Boulevard at 200th Street, Bronx, New York 10458-5126
Joseph F. Ammirati
University of Washington, Biology Department, Box 355325, Seattle, Washington 98195 USA
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTSKEY TO VINACEOUS OR...TAXONOMYDISCUSSIONLITERATURE CITED |
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A taxonomic treatment of vinaceous and reddish species of Tubaria (Agaricales) is presented based on morphology and nucleotide sequences. Accessions from western North America, Europe, Central America, the Caribbean and Australia are compared. Phylogenetic analysis of the 25S rRNA gene and internal transcribed spacer (ITS) regions demonstrates that Tubaria is not monophyletic. However the autonomy of six brown-spored, saprotrophic species—Naucoria vinicolor, Pholiota serrulata, Phaeomarasmius confragosus, Pholiota punicea, Tubaria rufofulva and T. bispora sp. nov.—is affirmed. All six species form a strongly supported monophyletic group that we label section Confragosae in the genus Tubaria. This section is related to T. dispersa, T. albostipitata and numerous representatives of the T. furfuracea complex. Tubaria minima is related to other lineages of dark-spored Agaricales. Taxonomic descriptions, illustrations and a key to confirmed species of section Confragosae are provided. Four new combinations are proposed, and one new species, T. bispora, is described as new from Costa Rica and Martinique. A lectotype is designated for Pholiota serrulata.
Key words: Agaricales, Basidiomycota, new combinations, species, systematics
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSKEY TO VINACEOUS OR...TAXONOMYDISCUSSIONLITERATURE CITED |
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, Ammirati 1989, Liu et al 1995) and others documented in Crepidotus (Luther and Redhead 1981, Pöder and Ferrari 1984, Bandala, Montoya, Horak 2006) and Stropharia (Shaw and Kibby 2001, Shaw, Butlin, Kibby 2004). Descriptions of several lesser-known but conspicuously colored agarics are scattered in North American, South American and Australian literature in the genera Tubaria (Murrill 1917, Bougher and Syme 1998), Phaeomarasmius (Singer 1953), Pholiota (Smith and Hesler 1968) and Naucoria (Peck 1909, Arora 1986). We became interested in the identification of several vinaceous pigmented species of brown-spored Agaricales that inhabit wood in natural habitats or mulched landscapes in suburban areas because these species are encountered so rarely and their taxonomic affiliations poorly understood. Co-authors have studied such material from Australia, Costa Rica, Europe and the west coasts of Canada and the United States and determined these accessions to represent unclarified species of the genus Tubaria (W.G. Sm.) Gillet (Moser 1983, Horak 2005).
Tubaria encompasses 8–16 species in Europe (Bon 1992, Volders 2002, Horak 2005), but the total number of names of taxa admitted into the genus is 114 (http://www.indexfungorum.org). Some combinations (e.g. T. decurrens [Peck] Murrill) have not been catalogued by Index Fungorum. Nevertheless Moreau (2005) suggests there are more than 60 taxa worldwide. Family placement has been controversial with some authors treating Tubaria in the Crepidotaceae (Singer 1986), Cortinariaceae (Romagnesi 1940) or Strophariaceae (Kühner 1980, Horak 2005) based on morphological data. Recent molecular phylogenetic evidence supports the exclusion of Tubaria from Crepidotaceae s. str. (Aime, Vilgalys, Miller 2005). A multigene study by Matheny et al (2006) affirms this but also provides evidence that Tubaria is not closely related to Cortinariaceae s. str. or to Strophariaceae s. str. Rather Tubaria and exemplars of Flammulaster Earle and Phaeomarasmius Scherffel are nested with significant Bayesian support in a clade joining the Inocybaceae, Crepidotaceae, Panaeoleae and Agrocybe erebia.
Most species of Tubaria are characterized by small basidiomes (pileus diameter often less than 5 cm), brownish coloration, presence of a central stipe, subdecurrent to adnate lamellae, ochraceous or brown spore deposits, presence of cheilocystidia, often smooth basidiospores that lack a germ pore, a filamentous pileipellis and saprotrophic habit. At least three species from the subtropics or Asia have been described in the genus with verrucose spores— T. thermophila Singer, T. verruculospora Pegler and T. lithocarpicola M. Zang (Singer 1948, Pegler 1977, Zang 2001)—and two others with rugose to punctate spores from Europe or North America—T. dispersa (L. : Fr.) Singer and T. decurrens (Peck) Murrill (Kühner and Romagnesi 1953, Moreau unpublished data). These five and other species described from non-European regions (e.g. Peck 1895, Murrill 1917, Singer and Digilio 1952, Singer 1957, 1969, Grgurinivoc 1997) have not been synthesized into any coherent systematic treatment of the genus (Singer 1986), although T. thermophila recently was transferred to Crepidotus based on molecular and morphological findings (Aime, Baroni, Miller 2002). Bon (1992) established an infrageneric classification for European species of Tubaria, but species such as T. confragosa were left unclassified. Here we apply morphological and molecular methods to identify our materials to species, confirm their generic status and produce a taxonomic treatment for annulate or vinaceous-colored species of Tubaria. In so doing we obtained type materials of species of Tubaria, Pholiota and Naucoria to assist in identification and clarify several taxonomic and nomenclatural issues.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSKEY TO VINACEOUS OR...TAXONOMYDISCUSSIONLITERATURE CITED |
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, Kornerup and Wanscher (1967) and Munsell (1954). References to Ridgway (1912) are capitalized (e.g. Tawny). References to Munsell Soil Color (1976) and the Munsell Soil Color Charts (1954) are by chroma and hue (e.g. 10YR 7/6) and from Kornerup and Wanscher (1967) by plate and row number (e.g. 6D5). All other color notations are approximations. "L" refers to the number of lamellae that reach the stipe. Microscopic descriptions of spores and basidiome tissues were made from fresh or dried material mounted in 3% KOH or 10% NH4OH. Extreme spore measurements are placed in parentheses. Typically 20–30 basidiospores were measured per collection. Line drawings were made with a drawing tube at 1000x for spores and 400x for other features. All materials are deposited in herbaria at AD, CUW, LIP, NY, PERTH, UC, WTU and ZT and the personal herbarium of O. Ceska (Vancouver Island, British Columbia, Canada). Herbarium acronyms follow Holmgren, Holmgren, Barnett (1990).
DNA extraction, PCR, and sequencing.— –
Dried material was ground in a 1.5 mL Eppendorf tube with a micropestle after freezing in liquid nitrogen. DNA extractions relied on an E.Z.N.A Fungal Miniprep Kit (Omega Bio-tek, Doraville, Georgia) following the protocol of the manufacturer. In the final step DNA was eluted with 70 µ L of elution buffer. PCR amplification, purification and sequencing of the 25S ribosomal RNA gene (25S rRNA) and the 5.8S ribosomal RNA plus internal transcribed spacers (hereafter referred to as ITS) follows protocols outlined in Yang et al (2005).
DNA alignments and phylogenetic analyses.— –
Sequence chromatograms were edited in Sequencher 4.1 (Gene-Codes, Ann Arbor, Michigan). Sequences were aligned with Clustal X 1.83 (Thompson et al 1997) and adjusted manually in MacClade 4.0 (Maddison and Maddison 2000). We performed an initial screening of 25S sequences to determine clade affinities of our tubarioid sequences in a dataset of 105 dark-spored Agaricales. Taxon selection was based on family/clade designations in the Agaricoid clade of Matheny et al (2006), Strophariaceae sensu Kühner (Gulden et al 2005), Crepidotaceae sensu Singer (Aime, Vilgalys, Miller 2005) and the/tubaria clade in Bodensteiner et al (2004). Previously published sequences from these studies were used in addition to others drawn from Binder and Bresinsky (2002); Matheny et al (2002); Moncalvo et al (2002); Frøslev, Matheny, Hibbett (2005); Matheny (2005); Walter, Garnica, Weiß (2005); Yang et al (2005); and Matheny and Bougher (2006). Results from this phylogenetic screening determined taxon selection for a combined 25S and ITS analysis. In the combined analysis, 39 25S and 34 ITS sequences of Tubaria and outgroups were assembled in interleaved format in PAUP* 4.0 (Swofford 2003). The final number of taxa analyzed was 49. Ten taxa lacked 25S sequences, and 15 others lacked ITS sequences. Twenty-one new 25S sequences have been deposited at GenBank (DQ986293
[GenBank]
-DQ986303
[GenBank]
, DQ987902
[GenBank]
-DQ987906
[GenBank]
, EF051051
[GenBank]
-EF051055
[GenBank]
) along with 30 new ITS sequences (DQ987907
[GenBank]
-13, DQ989327
[GenBank]
-34, DQ991348
[GenBank]
-56, EF051056
[GenBank]
-EF051060
[GenBank]
). GenBank accessions DQ494697
[GenBank]
/AF041544 for Stropharia rugosoannulata and AY8183450/AY646102 Stropharia ambigua were used as outgroups in the combined analysis. Outgroup choice is based on Matheny et al (2006).
All datasets were analyzed with Bayesian inference using the parallel version of MrBayes 3.1.2 with default priors (Ronquist and Huelsenbeck 2003, Altekar et al 2004) and maximum parsimony (MP) in PAUP*. These alignments are available at TreeBASE (S1750). Gaps were scored as missing data. PAUP* analyses were done on a Macintosh 1.33 GHz PowerPC G4. Bayesian analyses were implemented on a Linux cluster with AMD Opteron 246 processors. Separate partitions were modeled with a general-time-reversible (GTR) model (nst = 6) and rate heterogeneity parameters (gamma-distributed substitution rate and a proportion of invariable sites) following the AIC criterion in Modeltest 3.06 (Posada and Crandall 1998). Bayesian analyses entailed two independent runs each using four chains for 5 000 000 generations and sampling trees every 500 generations. Likelihood scores were inspected to determine stationary phases in a spreadsheet application, and multiple runs were subject to convergence diagnostics (examination of average standard deviation of split frequencies) to assess convergence of tree samples. With this information in hand we determined the number of initial trees to discard to construct 50% majority-rule consensus trees and calculate posterior probabilities (PP). PP values greater than or equal to 0.95 are considered significant. One thousand bootstrap replicates were done in PAUP* to determine parsimony support values or bootstrap proportions (BP). For bootstrapped tree searches 10 random stepwise additions with TBR were used but with MULTREES turned off. Values above 70% are considered significant.
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RESULTS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSKEY TO VINACEOUS OR...TAXONOMYDISCUSSIONLITERATURE CITED |
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). Two independent lineages of Naucoria vinicolor, one lineage of Pholiota serrulata and an unidentified Phaeomarasmius sequence were placed within a significantly supported clade (0.96 PP/73% BP) of 14 Tubaria sequences, including the type T. furfuracea.
The tribe Tubarieae (Tubaria and Flammulaster/Phaeomarasmius allies in Matheny et al 2006) is not closely allied with Crepidotaceae (including Pleuroflammula), Cortinariaceae or Strophariaceae sensu Gulden et al (2005). 25S rRNA sequences of Phaeosolenia densa, a reduced or cyphelloid brown-spored species regarded as a member of the/tubaria clade in Bodensteiner et al (2004) clustered with the outgroup Agaricaceae sequence in both MP and Bayesian analyses of 25S sequences (data not shown).
Combined 25S rRNA and ITS sequences support recognition of six species in Tubaria, section Confragosae.— –
The combined 25S rRNA and ITS super-matrix is composed of 1735 included characters (268 parsimony informative) and 49 taxa and was used to investigate species level relationships within Tubaria. Posterior probabilities were drawn from the last 8000 trees (burn-in of 2001 trees) sampled from two independent runs for a total of 16 000 trees. A 50% majority-rule consensus tree is shown, including branch lengths (FIG. 1).
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, Volders 2002), and six species of section Confragosae. Isolates of the T. furfuracea complex form an unresolved polytomy, whereas Confragosae is strongly supported as monophyletic (0.98 PP/87% BP). The section includes the wine-red Naucoria vinicolor, maroon T. rufofulva and a second clade of vinaceous specimens, to which we apply the name Pholiota punicea A.H. Sm. & Hesler (see taxonomy below). Also included are the annulate species Pholiota serrulata from Australia, T. confragosa from north temperate regions, and the newly described species T. bispora from Costa Rica and Martinique. Below we present a key to aid identification of these six species. |
KEY TO VINACEOUS OR ANNULATE SPECIES OF TUBARIA, SECT. CONFRAGOSAE |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSKEY TO VINACEOUS OR...TAXONOMYDISCUSSIONLITERATURE CITED |
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TAXONOMY |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSKEY TO VINACEOUS OR...TAXONOMYDISCUSSIONLITERATURE CITED |
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Basionym. Phaeomarasmius, section Confragosi Singer, Sydowia 15:75. 1962.
Pholiota, section Confragosae (Singer) A.H. Sm. & Hesler, North Am. Species of Pholiota: 39. 1968.
Comments. –
Six species are confirmed by molecular analyses to belong to this strongly supported mono-phyletic group in Tubaria: T. bispora, T. confragosa, T. punicea, T. rufofulva, T. serrulata and T. vinicolor (FIG. 1). The group is distinguished morphologically from other species of Tubaria by darker yellowish brown spore deposits and resistance of spore walls to collapse in microscopic mounts. The development of the partial veil is variable among the species in this section ranging from an evanescent cortina (T. punicea and T. vinicolor), a cortinate ring (T. bispora) or typically a membranous annulus (T. confragosa, T. rufofulva, and T. serrulata).
Both 25S rRNA and ITS sequences distinguish the Confragosae from the T. furfuracea complex, T. dispersa and T. albostipitata with significant measures of branch support. The T. furfuracea complex is distinguished by the lighter pigmented spore deposits (ochraceous) and spore walls that fail to revive (appearing collapsed) for some spores in microscopic mounts. Species in this group generally do not produce an annulus. The only brick-red species of Tubaria we suspect to belong outside the Confragosae is T. pallescens Peck, originally described from southern California (Peck 1895) (see comments under T. punicea and T. vinicolor below).
Smith and Hesler (1968) considered two additional species, Pholiota anomala Peck and P. canescens A.H. Sm. & Hesler, as members of section Confragosae and reported each having spores with a minute germ pore. This character is reported for a few species of Flammulaster (Vellinga 1986), but none of the species of the Confragosae and other typical species of Tubaria we examined possess a germ pore. Nevertheless if P. canescens A.H. Sm. & Hesler (non-Peck) proves to be an autonomous species of Tubaria a new name will be required because Tubaria canescens Peck (see Murrill 1917) has nomenclatural priority. Tubaria strophosa Singer (Singer 1969) is a temperate southern hemisphere species from Nothofagus forests that might belong to the Confragosae by virtue of the presence of an annulus and brown spores. In total Singer (1986) admitted 14 species in sect. Confragosi (in his concept of the genus Phaeomarasmius). Thus sect. Confragosae (Tubaria) might contain up to as many as 20 species worldwide.
Tubaria bispora Matheny, P.-A. Moreau, Neves, et Vellinga, sp. nov. FIGS. 2, 9, 10
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Pileus 0.5–1.5 cm diam, hemispherical to convex, margin recurved to slightly so in age, disk flattened or sometimes depressed, smooth or minutely furfura-fceous under a hand lens, (faintly) striate one-third to one-half to center; dark rusty brown with slight purplish tone when moist becoming reddish brown (8D6), hygrophanous, first fading at the center (8A3), elsewhere fading to pale cinnamon buff when dry (7A3-8A3); marginal veil fibrillose, discontinuous, light ochraceous; odor mild, taste fungoid; tissues exuding no pigments in 3% KOH solution. Lamellae adnate-subdecurrent to uncinate, subdistant to distant (12 L with several tiers of lamellulae), cinnamon to reddish brown (7D6); edges fimbriate under a lens. Stipe 1.0–2.0 cm x 1 mm, equal; surface longitudinally silky-striate, dark rusty brown when moist to shiny reddish brown (8D6) but concolorous with the pileus, with a delicate ochraceous cortinate ring on all basidiomes observed, densely pruinose at extreme apex, base with white mycelium at attachment with substrate, elsewhere somewhat fibrillose becoming smooth below; hollow. Basidiospores in side view 7.0–9.0 x 5.5–6.0 µ m, a few 10.0 x 7.5 µ m from hymenium, on average 7.6 x 5.9 µ m, Q = 1.2–1.4, on average 1.3, slightly wider in frontal view smooth, elliptic to broadly elliptic or occasionally subphaseoliform, smooth, apices rounded, apiculus indistinct, wall slightly thickened, generally not collapsing in alkali mounts, brownish yellow to yellowish, germ pore absent, often with one or two guttules. Basidia 25–30 x 5–7 µ m, 2-sterigmate, occasionally 1-sterig-mate, with long straight sterigmata 5–9 µ m long, subclavate, hyaline. Pleurocystidia none. Cheilocystidia in clusters on lamella edge, many 25–45 x 7–9 µ m, but some up to 85 µ m long, narrowly fusiform to cylindric with obtuse to swollen apices, at times subclavate, thin-walled or slightly thickened, hyaline. Lamellar trama regular, hyphae cylindric, generally 5–12 µ m diam, weakly incrusted, yellowish brown in mass. Pileipellis a discontinuous trichoderm of numerous upright terminal elements, these composed of chains of cells 30–60 x 15–25 µ m, elsewhere a cutis of prostrate cylindric hyphae mostly 7–15 µ m diam, incrusted to coarsely so, walls slightly thickened, "Ochraceous-Tawny" in mass. Stipitipellis with few terminal cells similar to cheilocystidia at the extreme apex. Clamps present.
Habitat and distribution. – Growing on wood in montane neotropical forest of Quercus insignis Martens & Galeotti, Costa Rica, also on soil along path rich in dead and decaying roots in mesophilic tropical forest, Martinique.
Specimens examined. – COSTA RICA. Puntarenas Coto Brus, Zona Protectoroa Las Tablas. Sitio Tinieblas. 1450 m elev, N8° 54' , W82° 46' , 11 Jun 2004, leg. M.A Neves, MAN 45 (HOLOTYPE, USJ; ISOTYPE, NY). MARTINIQUE. Case Pilote, arête Jean-Louis, 21 Aug 2005, leg. C. Lécuru and P.A. Moreau, PAM-Mart55 (LIP; CUW).
Commentary. –
The combination of bisporic basidia, small basidiomes, dark reddish brown (when moist), presence of a cortinate annular zone and tropical distribution support the autonomy of T. bispora. Two isolates, one from montane Quercus forests in Costa Rica and a second from mesophilic tropical forest in the Lesser Antilles, share almost identical 25S and ITS sequences. No species of Tubaria were recorded previously in floristic treatments of Agaricales and other fungi in the Lesser Antilles (Pegler 1983, Courtecuisse 2006), and no descriptive information is provided for unclarified Tubaria spp. in a checklist of Caribbean fungi by Minter et al (2001).
Phylogenetic analyses suggest T. bispora is most closely related to the north temperate T. confragosa (FIG. 1), which has a well developed annulus, flocculose pileal surface and tetrasporic basidia. Tubaria confragosula (Singer) Contu, described from subtropical Argentina, appears similar to T. bispora. It is reported to bear both tetra- and bisporic basidia, with the larger spores produced by bisporic basidia, and a pileipellis as "ex hyphis trichodermii palisadici" becoming a cutis with age. Contu’s interpretation of this species (Contu 2000) collected on introduced Eucalyptus from Sardinia is not entirely consistent with Singer’s protolog (Singer 1953). A comparison between T. confragosula sensu Contu and the type of Phaeomarasmius confragosulus Singer requires attention.
Tubaria confragosa (Fr.) Harmaja, Karstenia 18:55. 1978. FIG. 3
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Agaricus confragosus Fr., Epicr. Syst. Myc.: 169. 1838.
Pholiota confragosa (Fr.) P. Karst., Hattsvampar Bidr. Finlands Natur och Folk: 304. 1879.
Fulvidula confragosa (Fr.) Singer, Rev. Mycol. 2:239. 1937.
Naucoria confragosa (Fr.) Singer, Not. Syst. Sect. crypt. Inst. bot. Acad. Sci. U.S.S.R. 5 (‘1941’):94. 1945.
Phaeomarasmius confragosus (Fr.) Singer, Lilloa 22:577. (‘1949’) 1951.
Tubaria confragosa (Fr.) Kühner, Trav. Labor. La Jaysinia Samoëns 3:67. 1969 (Inval. Art. 33.3).
Pileus 3.0–6.0 cm, broadly convex to plano-convex with flattened disk, margin decurved; surface dry, not striate, flocculose or furfuraceous, margin with sparse remnants of membranous veil; reddish brown (5YR 4/3 or Rood’s Brown) when moist, hygrophanous, fading at the center first, becoming pale brown (10YR 7/4 or Cinnamon-Buff), disk fading to pinkish-cinnamon, the tiny scales not white but pale brown or pinkish-cinnamon to cinnamon-buff; odor strong, like Suillus, flavor similar to odor, context pinkish cinnamon or pallid with pinkish tinges when faded, firm, not changing where cut or bruised; flesh negative with PDAB, slowly green in tincture of guaiac. Lamellae adnate, close, 35–44 L with numerous lamellulae, Hazel (5YR 4/6 or 7E8) or rusty cinnamon; edges faintly pallid and obscurely fimbriate, concolorous with faces in age; even, medium breadth to somewhat narrow, 3–4 mm broad. Stipe 4.0–7.5 cm x 4–6 mm, even, straight or flexuous, tough, fibrous like Laccaria, not fragile; veil forming a superior membranous annulus that is easily removed, in age remnants of veil may be sparse; surface fibrillose to coarsely fibrillose, with a thin pallid fibrillose superficial layer; light reddish brown (5YR 6/3-6/4-5/3 or Light Vinaceous-Cinnamon to Mikado Brown) throughout except for the white base, not at all pruinose; hollow. Basidiospores in side view 6.0–7.5 x 4.5–5.0 µ m, on average 6.6–6.7 x 4.5 µ m, Q = 1.3–1.7, on average 1.45–1.5, slightly wider in frontal view smooth, elliptic to oblong, smooth, thick-walled, Ochraceous-Tawny or rusty to cinnamon-brown (darker than T. furfuracea complex), unreactive in Melzer’s; germ pore absent; spore deposit brown (Buckthorn Brown to Cinnamon-Brown or 10YR 5/8). Basidia 24–30 x 5–6 µ m, 4-spored, cylindric, hyaline. Pleurocystidia none. Cheilocystidia in clusters on lamella edge, 38–70 x 4–8 µ m, cylindric with swollen, obtuse, or less often indistinctly subcapitate apices, thin-walled or at times slightly thickened, hyaline. Lamellar trama regular, tawny cinnamon in mass, hyphae mostly 5–13 µ m diam, cylindric to inflated. Pileipellis composed of trichodermial tufts of thin- to thick-walled hyphae, these incrusted with pigment, 8–25 µ m diam, end cells undifferentiated. Stipitipellis composed of cylindric hyphae, mostly 5–10 µ m diam, yellowish brown in mass, extreme apex with (sub)cylindric terminal cystidioid cells, at times flexuous, these slightly thick-walled or thin-walled. Clamps present.
Habitat. –
In northern Europe reported on dead deciduous wood, primarily of Betula and Populus and less frequently on Alnus and Fagus (Moser 1983). In North America on hardwood and conifer logs (Smith and Hesler 1968) ranging from east coast to west coast and as far south as North Carolina.
Specimens examined. – USA. WASHINGTON: Kitsap County, Seabeck, along Hood Canal, on moss covered rotten hardwood (probably Alnus), 10 Jun 2001, leg. S. Clark, PBM 2105 (WTU). Olympic National Park, Whiskey Bend Trail, on wood and needles of Pseudotsuga, 17 Jun 1999, leg. C. Ammirati, MTS 4646 (WTU).
Comments. –
Sequence analysis supports the morphological findings of Kühner (1969) and Harmaja (1978), who placed Phaeomarasmius confragosus in Tubaria. Tubaria confragosa appears most closely related to T. bispora from Central America and the Caribbean. In our judgment the description of Pholiota canescens (Smith and Hesler 1968) is similar to T. confragosa except for the spores with a small germ pore. Excellent photos of North American T. con-fragosa can be found in Bessette, Bessette, Fischer (1997) and Lincoff (1981). The combination of the overall pinkish cinnamon coloration, well developed membranous annulus, and typically flocculose pileal surface easily distinguish the species morphologically. One European isolate differs from two western North American isolates by five 25S nucleotide positions and 12 ITS nucleotide positions.
Tubaria punicea (A.H. Sm. & Hesler) Ammirati, Matheny, et P.-A. Moreau, comb. nov. FIGS. 4, 11, 12 Basionym: Pholiota punicea A.H. Sm. & Hesler, North Am. Species of Pholiota: 40. 1968.
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Habitat and distribution. on rotten wood or base of Arbutus (Pacific Madrone, Ericaceae) typically in hollowed bases of large trees, sometimes damaged by fire, in mixed forest of Pseudotsuga, Arbutus or Quercus; distributed along coastal areas of British Columbia south to Marin County, California, possibly more widespread. The type was collected from a burned stump at the California-Oregon state line.
Specimens examined. – CANADA. BRITISH COLUMBIA: (All Canadian materials are split between the personal herbarium of O. Ceska and CUW). Vancouver Island, Miniskirt Mountain, near Victoria, N48° 27.794' , W123° 31.211' , 18 Dec 2005, leg. O. and A. Ceska, OC1, OC2. Vancouver Island, Mary Hill, DND property off Williams Head Road, Metchosin, N48° 20.931' , W123° 32.650' and N48° 28.894' , W123° 32.771' , 21 Dec 2005, leg. O. and A. Ceska, OC3, OC4. Vancouver Island, Albert Head, DND property, Metchosin, N48° 23.341' , W123° 28.735' , 23 Dec 2005, leg. O. and A. Ceska, OC5. Vancouver Island, Heals Rifle Range, DND property, Saanich, Victoria, N48° 32.938' , W123° 28.109' , 24 Dec 2005, leg. O. and A. Ceska, OC6, N48° 32.979' , W123° 28.135' , OC7. Vancouver Island, Cobble Hill near Shawnigan Lake, N48° 40.897' , W123° 37.918' , 25 Dec 2005, leg. O. and A. Ceska, OC8 and OC9. Vancouver Island, Observatory Hill, Saanich, Victoria, N48° 31.280' , W123° 25.228' , 26 Dec 2005, leg. O. and A. Ceska, OC10 & OC11. Vancouver Island, Yew Point, Esquimalt Lagoon, Victoria, DND property, N48° 26.292' , W123° 27.062' , 27 Dec 2005, leg. O. and A. Ceska, OC12. UNITED STATES. CALIFORNIA: Marin County, Mount Tamalpais, along Trucker Trail and on the shady side of Bon Tempe Lake, 3 Mar 2003, leg. E.C. Vellinga, ECV 3160 (UC). San Mateo County, Hubbard Park, 1 Dec 2006, collected for MSSF Fungus Fair, ECV 3522 (UC). San Francisco Watershed, 1 Dec 2006, collected for MSSF Fungus Fair, ECV 3523 (UC). OREGON. California-Oregon state line, 6 Nov 1937, Smith 8507 (HOLOTYPE, MICH).
Comments. –
The gross morphological description is based on material from British Columbia and merged with minor details from Smith and Hesler (1968). The microscopic description of Smith and Hesler (1968) is consistent with our examination of the type and reproduced here after some reformatting. It is apparent however that Smith did not fully document the type collection. Only brief notes occur on the label, which indicate blood red in all parts with a white fibrillose zone on the stipe. Our study of the holotype does not disagree much with the original description. Basidia are 1-, 2- and 4-sterigmate, which probably accounts for a larger range in spore size. Whether P. punicea is the same as N. vinicolor is unclear based on the types.
We initially designated vinaceous tubarioid collections from British Columbia, Washington and California as Naucoria vinciolor Peck (see below). However analysis of sequence data (FIG. 1) confirms two separate clades, one of which is ruderal in its distribution (occurring in landscaped areas with introduced vegetation) the other specific to mature (sometimes burned) bases of Arbutus in natural habitats. To distinguish the two clades we apply the name Tubaria punicea to the species that occurs on Arbutus. The original habitat description of P. punicea is more consistent with our application of the name than to the ruderal clade.
In his Tubaria monograph Murrill (1917) makes reference to the brick-red T. pallescens Peck (Peck 1895). We examined the type of T. pallescens (NYS) but conclude it may be more closely related to the T. furfuracea complex given the pale brown spores, many of which are collapsed. The stipe of T. pallescens is also described as yellowish, not red.
It might appear that the citation of 12 collections from British Columbia would indicate T. punicea is common, but this is not the case. For example, at the Cobble Hill site, more than 100 mature trees were examined before two trees were found with the fungus. Other localities were comparable to Cobble Hill. No materials of T. punicea and Naucoria vinicolor have been deposited at SFSU or WTU before this study, which further supports their rare status.
Collections OC6 and OC4 demonstrate length heterogeneity in the second spacer (ITS2). Both sequences share the same nucleotide polymorphic insertion/deletion (indel) event ( James et al 2001).
Tubaria rufofulva (Cleland) D.A. Reid & E. Horak, Austral. J. Bot., Supp. Ser. 10:29. 1983. FIGS. 5, 14 Basionym: Pholiota rufofulva Cleland, Trans. Proc. R. Soc. S. Austr. 51:303. 1927.
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Habitat and distribution. –
On woody debris and logs in Karri (Eucalyptus diversicolor) forest in Western Australia; on sandy soil and at the base of Eucalyptus in South Australia (Cleland 1934). Also reported on Nothofagus from New Zealand and Papua New Guinea (Horak 1983).
Specimens examined. –
AUSTRALIA. SOUTH AUSTRALIA. Burnside, AD 9930, 25 Jul 1925 (HOLOTYPE, AD). Kinchina, 8 Jun 1925, AD 9928. Para Wirra Cons. Park, 25 Jul 1999, AD-C 52053/PSC 319. Flinders Case National Park, 28 Jun 2004, AD-C 49169/PSC 1924. Murray Danggali Cons. Park, AD-C 52052/PSC 890. WESTERN AUSTRALIA. Near Manjimup, Pine Creek Road, 13 Jun 1985, ZT 2641 (ZT). Kellerberrin in dry open woodland, 20 Jul 1995, E5557 (PERTH). Denmark, in wet tall, forest, illustrated in Bougher and Syme (1998), 22 May 1995, E5662 (PERTH). Pemberton, in wet tall forest, 19 Aug 2001, E7038/PBM 2191 (PERTH, WTU).
Comments. –
The maroon color of T. rufofulva, also illustrated in Bougher & Syme (1998), distinguishes it from the more common T. serrulata (described below), at least in Western Australia (see FIGS. 13, 14). This difference also is supported by rRNA sequences (FIG. 1). Pinkish tinges characteristic of the annulus and stipe in T. rufofulva are not evident in T. serrulata. Cleland (1933) cited colors such as brown, tawny and buff in his original description of Pholiota serrulata. For T. rufofulva (as P. rufo-fulva) Cleland (1927) described the colors as Pecan Brown, Russet, Tawny when old, and dark maroon-brown when moist. For these reasons T. rufofulva and T. serrulata should not be considered synonyms of each other as done by Grgurinovic (1997).
Pholiota imperfecta Cleland also was listed as a synonym of T. rufofulva in Grgurinovic (1997). It originally was described by Cleland (1933) with smaller basidiomes and differently shaped spores (oblique vs. elliptic) compared to P. serrulata. However Cleland’s protocol describes the pileus of P. imperfecta as "ceraceo-flavobrunneus", the stipe as "pallidus vel rufobrunneus", and the annulus as "albidus". Because no vinaceous or maroon colors are mentioned in the protolog, it is doubtful P. imperfecta represents T. rufofulva. However none of the collections of P. imperfecta at AD provide a clear picture of what species this name represents. All collections are in poor condition and of variable identity. For example AD-C 22420, which is labeled on the voucher packet as "Type of P. imperfecta", consists mainly of an Inocybe species with metuloid cystidia. AD-C 22421 is labeled on the voucher packet as "Co-type (of P. imperfecta)" and contains probably a Galerina species (spores with a distinct plage). AD 9920, AD 9921, AD 9922 and AD 9923, listed as syntypes of P. imperfecta by Grgurinovic (1997), are determined as matching T. rufofulva, except for AD 9923, which matches T. serrulata. Because of these ambiguities we consider P. imperfecta a taxon of doubtful interpretation.
Accessions of T. rufofulva fall into two spore-group categories. One group (including the type) has larger spores on average (8.0–8.9 x 4.9–5.5 µ m) than a second group (including PBM 2191), which on average has smaller spores (6.7–7.3 x 4.3–5.0 µ m). We have not studied material for OKM 24681, for which an autonomous 25S rRNA branch is shown (FIG. 1). The two spore groups of T. rufofulva do not appear to have any other consistent morphological differences, and there is no apparent geographical pattern distinguishing the groups. The significance of the two spore groups is unknown at this time.
Notes of material collected in Papua New Guinea cited in Horak (1983) indicate basidiomes of a smaller stature with a veil that is fragmentary or lacking and occurrence on decaying wood of Nothofagus. The spore size for this group is consistent with the smaller spore size of the second group referred to above.
Tubaria serrulata (Cleland) Bougher et Matheny, comb. nov. FIGS. 6, 7, 13 Basionym: Pholiota serrulata Cleland, Trans. Roy. Soc. South Australia 57:190. 1933.
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Habitat and distribution. – On ground on woody debris including buried remnants of Banksia cones in urban bushland. The type is recorded on ground from Mount Lofty, South Australia. Known from temperate Western Australia, South Australia and Victoria.
Specimens examined. – AUSTRALIA. SOUTH AUSTRALIA: Belair National Park, 12 Apr 1917, AD 9917 (AD); Belair National Park, 19 May 1925, AD 9918 (AD); Belair National Park, 16 May 1931, AD 9919 (AD); Mount Lofty, 29 Apr 1921, AD 9924 (AD); Mount Lofty, 25 Apr 1924, AD 9925 (AD); Mylor, 16 May 1925, AD 9929 (LECTOTYPE here designated, AD); Encounter Bay, 9 May 1926, labeled "co-type", AD 9931 (AD). WESTERN AUSTRALIA: Walpole, in wet tall forest, E4637 (PERTH); Kellerberrin, dry open woodland, E5382 (PERTH); Plavins Forest Block, east of Dwellingup, in Jarraah (Eucalyptus marginata) forest, 9 Jul 1996, E5631 (PERTH); Kings Park, Perth, under Allocasuarina sp., 2 Jun 1997, E5837 (PERTH); Dwellingup, in Jarrah forest, 28 Jul 1997, E5928 (PERTH); Valema Farm, Corrigin, in Eucalyptus loxophleba woodland, 19 Jul 2000, E6633 (PERTH); Corner of Ludlow and Hithergreen Road, 15 km northeast of Busselton, under Eucalyptus gomphocephala, E. marginata and Agonis flexuosa, 26 Jun 2001,E6798 (PERTH); Perth, Warwick Open Space, 1 Jul 2001, PBM 2142 (E6899) (WTU, PERTH); Brookton Highway, south of Perth, near Epacridaceae on granite outcrop, 3 Jul 2001, PBM 2149 (E6931); Perth, Shenton Bushland, 22 Jul 2001, PBM 2160 (E7008) (WTU, PERTH); same locality as above, PBM 2161 (E7009) (WTU, PERTH); Kensington, Perth, urban bushland, under mixed planted eucalypts and other Myrtaceae, 13 Jul 2004, E8069 (PERTH); Bold Park, Perth, in Jarrah woodland, 14 Jul 2004, E8071 (PERTH); Bold Park, Perth, under Marri (Eucalyptus calophylla), 14 Jul 2005, E8209 (PERTH). VICTORIA: Snowy River National Park, 29 Apr 1995, E5270 (PERTH).
Comments. –
We apply the name Pholiota serrulata to material collected in Western Australia and Victoria after comparison with type materials of this species at AD. Grgurinovic (1997) treated P. serrulata as a synonym of Tubaria rufofulva, a disposition we do not accept mainly due to the differences in pileal color—reddish brown for P. serrulata and maroon for T. rufofulva. In the field (FIGS. 13, 14) the colors clearly distinguish the two species, which is also confirmed by rRNA sequences (FIG. 1). Lectotypification of the collection AD 9929 from Mylor, a place name mentioned by Cleland (1933) in his protolog, justifies its selection as lectotype. Other collections cited as syntypes in Grgurinovic (1997) are inappropriate selections because they are from locations other than Mylor, including AD 9919 (from Belair National Park), despite annotations on the voucher packet to the contrary written 27 May 1999.
Tubaria serrulata appears close to T. confragosa of the northern hemisphere given the annulus but lacks the flocculose pileus and thick-walled hyphae of the pileipellis of that species and has a differently colored pileus. A striate reddish brown and annulate species, T. strophosa Singer (Singer 1969), is recorded from the Nothofagus zone in Argentina. Based on the protolog it would appear similar to T. serrulata.
Tubaria vinicolor (Peck) Ammirati, Matheny, et Vellinga, comb. nov. FIGS. 8, 15 Basionym: Naucoria vinicolor Peck, Bull. Torrey Bot. Club 36:334. 1909.
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Habitat and distribution. – Solitary, gregarious or caespitose, in grass covering soil and rotting wood, around stump of Eriobotrya japonica (Japanese plum, Rosaceae), producing basidiomes in October in Washington and during December, January and March in California.
Specimens examined. – UNITED STATES. CALIFORNIA: Claremont, on open ground, Jan 1909, leg. C.F. Baker (HOLOTYPE of Naucoria vinicolor, NYS). Alameda County, Berkeley, 861 Keeler Ave, in front garden, growing in several clusters around stump of Eriobotrya japonica, 3 Mar 2005, leg. E.C. Vellinga, ECV 3340 (UC). Berkeley, same locality as previous, 23 Dec 2005, leg. E.C. Vellinga, ECV 3449 (UC). Berkeley, same locality as previous, 23 Nov 2006, leg. E.C. Vellinga, ECV 3501 (UC). WASHINGTON. King County, Redmond, Rose Hill, 80th St. and 139th NE, leg. D. Winkler, 17 Oct 2003, JFA 12905 (WTU). Redmond, same collector and locality as previous, 9 Oct 2005, JFA 13103 (WTU).
Comments. –
Tubaria vinicolor is a North American vinaceous or wine red species found in ruderal landscapes with introduced vegetation. To date it is known only from the west coast. It appears closely related to two other vinaceous species, T. punicea, also from the west coast of North America, and T. rufofulva, from temperate Australia (FIG. 1). Arora (1986) reports Naucoria vinicolor on Quercus in California, as far south as Los Angeles County where it is encountered rarely. However he was not aware of T. punicea at the time, with which T. vinicolor could be confused. We were unable to study any material produced on Quercus. Nevertheless ecology and sequence data appear to readily distinguish T. vinicolor from T. punicea.
The type (NYS, Peck 1909) contains what appear to be two specimens, the stipes of which are in one piece, but the pilei have been broken into several pieces. One stipe base is attached directly to a piece of wood. The pileipellis structure is difficult to interpret, although it is clearly a cutis. Naucoria in its modern sense as Alnicola (see Moreau 2005) is not an appropriate generic designation for N. vinicolor. Species of Alnicola possess verrucose spores and are ectomycorrhizal.
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DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSKEY TO VINACEOUS OR...TAXONOMYDISCUSSIONLITERATURE CITED |
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and Kirk et al (2001) recognized about 15 species in the genus, to which we add T. bispora, T. confragosa, T. punicea, T. rufofulva, T. serrulata and T. vinicolor. Of these six species, four have conspicuous vinaceous or maroon basidiomes and three have a membranous annulus or distinct annular cortinate ring-zone. All have spores that are more strongly pigmented and resist collapsing when revived in microscopic mounts compared to members of the T. furfuracea complex.
One hundred fourteen records of epithets and infraspecific taxa have been used in Tubaria. A small percentage of these records are invariably synonyms (e.g. T. autochthona [Berk. & Broome] Sacc. and T. dispersa [L. : Fr.] Singer) or represent species now classified in other genera such as Psilocybe, Galerina, Phaeogalera, Crepidotus and Flammulaster; and many taxa probably fall in the T. furfuracea complex. However some species such as Tubaria ferruginea Maire ex E. Horak & P.-A. Moreau and Flammulaster siparia (Fr. : Fr.) Watling have an ambiguous position by sharing morphological characters of both genera (Vellinga 1986, Horak and Moreau 2005). Yet as we have shown arguably just as many species, if not more, remain poorly known and displaced in other Friesian genera, e.g. Pholiota s. lat. (Smith and Hesler 1968), Naucoria s. lat. (Moreau 2005), and in a broadly circumscribed but nonmonophyletic Phaeomarasmius (Singer 1986). Others have been subsumed under broad morphological species (Grgurinovic 1997) or remain undescribed in areas poorly studied by systematic mycologists (e.g. Australia and the neotropics).
Our results suggest that Tubaria is a paraphyletic group (excluding T. minima, not shown) in which at least one questionable isolate of Flammulaster is derived. However we were able to recover a strongly supported subclade of at least 14 Tubaria species, including one possibly mislabeled Phaeomarasmius isolate. At this time the most inclusive clade of Tubaria is composed of taxa with pileate-stipitate basidiomes, central stipe, a filamentous pileipellis incrusted with pigment, smooth or faintly roughened pale brown, ochraceous or yellowish-brown spores with a thin or thick wall, no germ pore, distinctive cheilocystidia, subdecurrent to adnate lamellae, a partial veil (cortinate or annulate), clamp connections and saprotrophic habit. No cyphelloid taxa have been established unequivocally as related to Tubaria based on our phylogenetic analyses. However Bodensteiner et al (2004) recovered Phaeosolenia as a sister taxon of one Tubaria sequence but with poor support. Several non-European species have been described with verrucose or ornamented spores, but sequence data for most of these are not available (see Aime, Baroni, Miller 2002, who transferred T. thermophila to Crepidotus).
Several authors have accepted a generic distinction between Tubaria and Flammulaster Earle and Phaeomarasmius Scherffel (Orton 1960, Watling 1967, Horak 1980, Moser 1983, Vellinga 1986, Horak 2005). Phylogenetic analyses of 25S rRNA (Aime, Vilgalys, Miller 2005) and multigene studies (Matheny et al 2006) indicate that exemplars of these genera indeed are closely related. However neither Flammulaster nor Phaeomarasmius is currently monophyletic (FIG. 1) and Singer (1986) treated them as a single genus under the name Phaeomarasmius. A new genus, Phaeomyces Horak (Horak 2005), has been described that might be affiliated with these groups as well. Tubaria and allies clearly merit more attention to address their generic boundaries and putative relationship to cyphelloid taxa.
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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1 Corresponding author. E-mail: pmatheny{at}clarku.edu
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSKEY TO VINACEOUS OR...TAXONOMYDISCUSSIONLITERATURE CITED |
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