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Phylogenetic Relationships and an Assessment of Traditionally Used Systematics and Biodiversity ISSN: 1477-2000 (Print) 1478-0933 (Online) Journal homepage: https://www.tandfonline.com/loi/tsab20 Phylogenetic relationships and an assessment of traditionally used taxonomic characters in the Sporormiaceae (Pleosporales, Dothideomycetes, Ascomycota), utilising multi‐gene phylogenies Asa Kruys & Mats Wedin To cite this article: Asa Kruys & Mats Wedin (2009) Phylogenetic relationships and an assessment of traditionally used taxonomic characters in the Sporormiaceae (Pleosporales, Dothideomycetes, Ascomycota), utilising multi‐gene phylogenies, Systematics and Biodiversity, 7:4, 465-478, DOI: 10.1017/S1477200009990119 To link to this article: https://doi.org/10.1017/S1477200009990119 Published online: 11 Mar 2010. Submit your article to this journal Article views: 192 View related articles Citing articles: 29 View citing articles Full Terms & Conditions of access and use can be found at https://www.tandfonline.com/action/journalInformation?journalCode=tsab20 Systematics and Biodiversity 7 (4): 465-478 Issued 1 December 2009 doi:io.ioi7/Si4772oooo999OU9 © The Natural History Museum Phylogenetic relationships and an assessment of traditionally used taxonomic characters in the Sporormiaceae (Pleosporales, Dothideomycetes, Ascomycota), utilising multi-gene phylogenies Asa Kruys1,* & Mats Wedin2 1Department of Systematic Biology, Evolutionary Biology Centre, Uppsala University, Norbyvägen 18D, SE-752 36 Uppsala, Sweden 2Department of Cryptogamic Botany, Swedish Museum of Natural History, Box 50007, SE-104 05 Stockholm, Sweden submitted February 2009 accepted June 2009 Contents Abstract 465 Introduction 466 Materials and methods 467 Results 469 Discussion 469 Conclusions and suggestions for the future 475 Taxonomy 476 Preussia alloiomera comb. nov. 476 Preussia antarctica comb. nov. 476 Preussia bipartis comb. nov 476 Preussia borealis comb. nov 476 Preussia dubia comb. nov. 476 Preussia lignicola comb nov. 476 Preussia longisporopsis comb. nov. 476 Preussia minipascua comb. nov. 476 Preussia octomera comb nov. 476 Preussia splendens comb nov. 476 Preussia tenerifae comb nov. 476 Preussia tetramera comb. nov. 476 Westerdykella angulata comb nov. 476 Westerdykella aurantiaca comb nov. 476 Acknowledgements 476 References 476 Abstract The family Sporormiaceae (Pleosporales, Dothideomycetes, Ascomycota) occur worldwide and a majority of the species are coprophilous. The taxonomy and classification of the family are based on a small number of morphological and ecological characters. Several taxa are easily confused by their shared morphological features, and the relationships between genera are poorly known and in need of critical study. The aims of this study were to resolve the phylogenetic relationships within the Sporormiaceae, test the current generic classification, and study the utility of traditional characters for the taxonomy in the group. To resolve these questions, we analysed combined data sets of ITS-nLSU rDNA, mtSSU rDNA and ß-tubulin sequences with parsimony and Bayesian methods. * Corresponding author. Email: [email protected] 465 Published online 11 Mar 2010 466 Asa Kruys and Mats Wedin The results showed that many characters, which previously have been used in the taxonomy and classification of the Sporormiaceae, such as the substrate choice, presence or absence of an ostiole, and presence or absence of germ slits, were all homoplastic and not useful for circumscribing monophyletic groups. A number of characters may be useful for circumscribing smaller clades if used in combination with other features, such as the shape of the ascus with the shape of the ascospores. Our phylogenetic analyses show that Preussia and Sporormiella are non-monophyletic, and a constrained analysis forcing these genera into monophyly resulted in significantly worse trees. Spororminula is nested in Preussia s. lat., and Eremodothis and Pycnidiophora are nested within Westerdykella. Finally, we suggest a new generic classification for the family Sporormiaceae, including Sporormia, Preussia (including Sporomiella and Spororminula) and Westerdykella (including Eremodothis and Pycnidiophora). We also propose 14 new combinations: Preussia alloiomera (S.I. Ahmed & Cain) Kruys, Preussia antarctica (Speg.) Kruys, Preussia bipartis (Cain) Kruys, Preussia borealis (I.Egeland) Kruys, Preussia dubia (S.I. Ahmed & Cain) Kruys, Preussia lignicola (W. Phillips & Plowr.) Kruys, Preussia longisporopsis (S.I. Ahmed & Cain) Kruys, Preussia minipascua (S.I. Ahmed & Cain) Kruys, Preussia octomera (Auersw.) Kruys, Preussia splendens (Cain) Kruys, Preussia tenerifae (Arx & Aa) Kruys, Preussia tetramera (S.I. Ahmed & Cain) Kruys, Westerdykella angulata (A.C. Das) Kruys and Westerdykella aurantiaca (J.N. Rai & J.P. Tewari) Kruys. Keywords ß-tubulin, bitunicate, coprophilous, fungi, loculoascomycete, phylogeny Introduction odothis, has been included in the family since then (Kruys et al., 2006; Lumbsch & Huhndorf, 2007). Also, the hypho- The classification of many groups of Fungi has traditionally mycete Amorosia should be transferred to the Sporormiaceae been based on a few easily observed characteristics, which of- according to Mantle et al. (2006), but its closest relatives are ten reflect aspects of their ecology and thus may be subject to unclear. parallel evolution or extreme specialisation in highly derived The type genus, Sporormia, is characterised by globose groups. There are numerous examples of non-monophyletic pseudothecia, which open with an ostiole. The 16-celled fungal groupings that have remained in classifications un- spores are typically joined together in a bundle with one com- til very recently; like the well-known basidiomycetes in the mon gelatinous sheath, when released from the ascus (Ahmed Aphyllophorales that have been shown to comprise highly un- & Cain, 1972; Dissing, 1992). Germ slits in the spore walls, related groups which all have the crust-like fruiting bodies otherwise common and characteristic for the family, are absent in common (Larsson et al., 2004; Binder et al., 2005), and in the type species Sporormiafimetaria (Ahmed & Cain, 1972). the mazaedia-producing Caliciales among the ascomycetes, Sporormia, Sporormiella and Preussia are considered closely which independently have acquired prototunicate, evanescent related and are easily confused by their shared morphological asci and passive spore dispersal (Tibell, 1984; Wedin & Tibell, features (Munk, 1957; von Arx & Muller, 1975; Eriksson, 1997; Wedin et al., 1998). With the progress of molecular 1981). Sporormiella, the most species-rich genus in the fam- phylogeny, the monophyly of these groups and the potentially ily, includes species with perithecioid ascomata and 4-celled synapomorphic traits defining them can be tested with inde- spores with germ slits (Ahmed & Cain, 1972). Preussia differs pendent means and classifications updated accordingly (e.g. morphologically from 4-celled Sporormiella species only by Hibbet et al., 2007; Lumbsch & Huhndorf, 2007). Still, po- the ascomata being cleistothecioid (Cain, 1961; Barr, 2000). tentially non-monophyletic groups defined on singular ecolo- The taxonomic importance of the ostiole has, however, been gical or morphological traits prevail, particularly among taxa questioned, since its presence in Preussia and Sporormiella on lower systematic levels. One such example is the generic seem to depend on growth conditions (von Arx, 1973; Guarro delimitation within the Sporormiaceae (Pleosporales, Dothi- et al., 1997a). According to experiments in vitro, different diomycetes, Ascomycota). morphs can even be found within the same culture (Guarro The Sporormiaceae occur worldwide and live as sap- etal., 1997a). robes on various substrates including dung, plant debris, soil Substrate choice has been another important diagnostic and wood. Recently, endophytic representatives have also been character for separating Preussia and Sporormiella util- reported (Arenal et al., 2007). Most taxa, however, are copro- ised by von Arx and van der Aa (1987). Preussia, in philous and their ascospores are useful tools in paleoecological their sense, includes species on plant debris, wood or soil, studies, when reconstructing animal abundances in the past while Sporormiella is restricted to coprophilous species (Burney et al., 2003; van Geel et al., 2003, 2007). The charac- with perithecioid ascomata. Doveri (2004) also considered teristic ascospores are thick-walled and dark brown, strongly the preference for a certain substrate an important fea- constricted at septa and often fragmenting into part-spores at ture, together with the shape of the ascus where Sporormi- maturity (Barr, 2000). Germ slits in the spore walls are also ella should have cylindric or cylindric-claviform asci and common features (Fig. 1). The spores are developed in fis- Preussia more or less clavate asci. He also believed that situnicate asci and pseudothecioid ascomata (Fig. 1). In the Preussia in general have more superficial ascomata. Typ- latest treatment (Barr, 2000), the Sporormiaceae comprises ical species of Sporormiella sensu von Arx & van der Aa eight genera, and c. 100 species. One more genus, Erem- occur on substrates other than dung, while for instance Phylogeny of Sporormiaceae 467 à. Figure 1 Ascomata (a) with ostiole (Sporormiella sp.) and (b) without ostiole (Preussia funiculata). (c) Spore with germ slits (Sporormiella sp.). Asci and spores of taxa representing major monophyletic groups in Sporormiaceae. (d) Westerdykella sp. (the Westerdykella
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