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Sharma, Gräser & Singh ARTICLE Auxarthronopsis have more than 10 septa. The phylogenetic interrelationships of Auxarthron and Amauroascus have been studied by Sole et al. (2002a) using ITS sequences, who found the former genus to be monophyletic and the latter polyphyletic. Our nrSSU and nrLSU trees also showed Auxarthron as monophyletic. The new taxon was excluded from the Auxarthron clade (Figs 4– 5), and the ITS sequence comparisons also showed Auxarthronopsis as distinct from Auxarthron, but closer to Amauroascus purpureus, Nannizziopsis albicans, and Amauroascus volatilis-patellis. The ITS similarity of Auxarthronopsis bandhavgarhensis to the nearest neighbour, Amauroascus purpureus, was less than 85 % (119 bp out of 620 bp; 92 substitutions and 27 indels). Morphologically, A. purpureus is also distinct in possessing purple ascomata that lack knuckle joints and peridial appendages, and forming globose, frilled ascospores (Ito & Nakagiri 1995). The other phylogenetically closest neighbour, Nannizziopsis albicans (Fig. 6), differs from the new taxon in having a peridium of undifferentiated peridial hyphae without appendages, and in lacking a well-developed reticuloperidium (Guarro et al. 1991). Phylogenetically these two species differ at 128 of 627 nucleotide positions, including 74 substitutions and 54 indels. Another phylogenetically related species, Amauroascus volatilis-patellis, differs from Auxarthronopsis bandhavgarhensis in forming a peridium of undifferentiated hyphae lacking appendages, and in producing ascospores which are punctate-reticulate. It also differs at 166 of 623 nucleotide positions, 102 of which are substitutions and 64 of which are indels. In the nrLSU and nrSSU trees (Figs 5–6), Auxarthronopsis was placed close to Amauroascus, Renispora, and Neogymnomyces. However, the new taxon is morphologically inconsistent with these genera in the distinctness of the ascomata, with a mesh-like reticuloperidium with swollen septa. Renispora flavissima has ascomata of poorly differentiated hyphae, which lack appendages, and forms finely pitted reniform or bacilliform ascospores. The type species of Neogymnomyces, N. demondreunii, forms discrete, spherical golden-yellow ascomata with hyaline, thick-walled, smooth and septate peridial hyphae, along with long, hyaline, peridial appendages. Recently, a new species of Neogymnomyces, N. virgineus, was described by Doveri et al. (2011) based on morphological and molecular data; however, it could not be included in our analysis as its ITS and LSU sequences (JN038187, JN038186) were not available to us. Neogymnomyces virgineus forms irregularly globose to pulvinate ascomata with pale yellow, fairly thick walled, verruculose septate hyphae along with two kinds of peridial appendages: one long, tapering and smooth, and the other comparatively short, often branched, verruculose, and rounded at the tips. Neogymnomyces also differs in lacking swollen septa in the peridial hyphae and the appendages, and in possessing an arthro- and aleurioconidial asexual morph characterised by frequent swollen cells. Another genus that possesses knuckle joints is the monotypic Pectinotrichum (Varsavsky & Orr 1971). Pectinotrichum llananse forms a reticuloperidium with long appendages and knuckle joints, but differs from Auxarthronopsis in having pectinate hyphae, large tubercules on the appendages and peridial hyphae, a Chrysosporium asexual morph, and almost smooth ascospores. Currah (1994) transferred Pectinotrichum llanense to Auxarthron, but its distance from Auxarthron was substantiated by the molecular studies of Sugiyama et al. (1999). Phylogenetically that species is distant from Auxarthronopsis as it separates out from the main Onygenaceae clade in SSU analysis (Fig. 5) and clusters with Ctenomyces serratus. Varsavsky & Orr (1971), who originally noted that the pectinate hyphae of P. llanense resemble the ctenoid appendages of Ctenomyces serratus, suggested that genetic studies of these genera would provide more evidence of a close relationship between them (Varsavsky & Orr 1971). Knuckle-joints are a common feature in species with well developed reticuloperidia, viz. Gymnoascus reesii, Pectinotrichum llananse, and almost all Auxarthron species. Their purpose may be to provide strength to the mesh-like spherical peridium which protects the central ascospore mass from mite grazing in nature (Summerbell 2000). The swollen septa in Auxarthronopsis are slightly different in that they have a ring of tubercles visible under SEM (Fig. 3c, d) making knuckle joints relatively prominent in LM even though they are found on thin, smooth or sparsely asperulate, peridial hyphae. Currah (1985) suggested that the morphological discontinuity seen in several genera of Onygenales might be bridged by as yet unrecorded forms. Perhaps Auxarthronopsis represents one such intermediate genus. The order Onygenales now comprises 18 genera that have peridial appendages. These are usually borne on either a membranous cleistoperidium or a completely hyphal, mesh-like reticuloperidium. They may also be formed on an incomplete hyphal mesh-like incompositoperidium. A key to all appendaged genera of the order is provided, including Ctenomyces, the peridium of which includes membranous as well as hyphal elements. Key to the genera of Onygenales with peridial appendages 1 Ascomata cleistoperidium type ......................................................................................................................................... 2 Ascomata reticuloperidium or incompositoperidium type .................................................................................................. 3 2(1) Ascomata made of only cleistoperidium ................................................................................................. Castanedomyces Ascomata made of inner layer cleisto-type, outer layer reticulo-type ............................................................. Ctenomyces 3(1) Peridial appendages straight with simple or curved apices, branched or unbranched, uncinate, never coiled ................ 4 Peridial appendages coiled .............................................................................................................................................. 11 100 ima fUNGUS
Auxarthronopsis gen. sp. nov. 4(3) Peridial hyphae or appendages with knuckle joints .......................................................................................................... 5 Peridial hyphae or appendages without knuckle joints ..................................................................................................... 7 5(4) Peridial appendages without or up to 3 septa .................................................................................................. Auxarthron Peridial appendages multi-septate .................................................................................................................................... 6 ARTICLE 6(5) Peridium with short pectinate appendages ............................................................................................... Pectinotrichum Peridium lacking pectinate appendages, swollen septa on appendages ............................................... Auxarthronopsis 7(4) Peridial appendages straight, rarely branched .................................................................................................................. 8 Peridial appendages, if present, branched or uncinate ..................................................................................................... 9 8(7) Peridial appendages with acute apices, ascospores smooth ............................................................................... Acitheca Peridial appendages with blunt apices, ascospores ornamented ............................................................... Nannizziopsis 9(7) Ascospores smooth ......................................................................................................................................................... 10 Ascospores pitted, pitted-reticulate ........................................................................................................ Neogymnomyces 10(9) Peridial appendages if present, smooth, roughened (uncinate or boat-hook shaped apices) ...................... Gymnoascus Peridial appendages ornamented with numerous short fine hairs, dichotomously branched, with blunt apices ....................................................................................................................................................................... Bifidocarpus 11(3) Peridial appendages coiled ............................................................................................................................................ 12 Peridial appendages uncinate ..................................................................................................................... Uncinocarpus 12(11) Ascospores hyaline ......................................................................................................................................................... 13 Ascospores yellow to orange ......................................................................................................................................... 14 13 (12) Ascospores smooth ....................................................................................................... Histoplasma (syn. Ajellomyces) Ascospores irregularly ridged ........................................................................................................................... Kuehniella 14(12) Ascospores ellipsoidal or oblate ...................................................................................................................................... 15 Ascospores globose, finely echinulate .................................................................................................................. Apinisia 15(14) Peridial hyphae made up of ossiform cells .................................................................................................... Arthroderma Peridial hyphae not made up of ossiform cells ................................................................................................................ 16 16(15) Peridial appendages thin walled, ascospores minutely roughened, pitted ........................................................ Shanorella Peridial appendages thick walled .................................................................................................................................... 17 17(16) Ascospores smooth ......................................................................................................................................... Spiromastix Ascospores regularly punctate-muricate ........................................................................................................... Polytolypa ACKNOWLEDGEMENTS REFERENCES R.S. is indebted to Tiger Trust, India, for organising a two- day Workshop on Tiger Conservation at Tiger Lodge, near Bhandhavgarh National Park, that made the collection of soil samples possible, to Madan L. Sharma (Regional Sophisticated Instrumentation Centre, Panjab University, Chandigarh) and Sugat V. Shende (SEM Facility, Department of Physics, University of Pune, Pune) for assistance with the scanning electron microscopy. Ram C. Rajak and Akhilesh K. Pandey are acknowledged for support during the initial stages of this study. We thank the Department of Science and Technology (DST), Government of India, New Delhi for financial support ( No. SR/FT/L-36/2005 and No. SP/SO/PS-55/2005), and the Director, MACS’Agharkar Research Institute, Pune, for extending facilities. Currah RS (1985) Taxonomy of the Onygenales: Arthrodermataceae, Gymnoascaceae, Myxotricaceae, and Onygenaceae. Mycotaxon 24: 1–216. Currah RS (1988) An annonated key to the genera of the Onygenales. Systema Ascomycetum 7: 339–350. Currah RS (1994) Peridial morphology and evaluation in the prototunicate ascomycetes. In: Ascomycete Systematics: problems & perspectives in the nineties (Hawksworth DL, ed.): 281–293. [NATO ASI Series no. 269.] New York: Plenum Press. Doveri F, Pecchia S, Vergara M, Sarrocco S, Vannacci G (2011) A comparative study of Neogymnomyces virgineus, a new keratinophilic species from dung, and its relationships with the Onygenales. Fungal Diversity 52: 13–34. volume 4 · no. 1 101
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