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Pleosporales - CBS - KNAW

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Fungal Diversity<br />

substrates and many occur in freshwater or has been<br />

isolated from air. The polyphyletic nature of T. aristata<br />

has been well documented (Tanaka et al. 2009). Anamorphic<br />

stages can serve as a diagnostic character for this<br />

family.<br />

Diademaceae, Massariaceae, Sporormiaceae<br />

and Teichosporaceae<br />

The Sporormiaceae is coprophilous having Phoma or<br />

Phoma-related anamorphic states (Cannon and Kirk 2007).<br />

Comoclathris (Diademaceae) is linked with Alternaria-like<br />

anamorphs (Simmons 1952). Myxocyclus links to Massaria<br />

(Massariaceae) (Hyde et al. 2011). The anamorphic stage of<br />

Chaetomastia (Teichosporaceae) isAposphaeria- orConiothyrium-like<br />

(Barr 1989c).<br />

Generally speaking, the morphologically simple conidiophores<br />

are usually considered phylogenetically uninformative<br />

(Seifert and Samuels 2000). Phoma-like anamorphs commonly<br />

occur in <strong>Pleosporales</strong>, while their colorless and unicellular<br />

conidia are also not phylogenetically informative (Seifert and<br />

Samuels 2000).<br />

All of the above mentioned anamorphic taxa of<br />

<strong>Pleosporales</strong> have phialidic, annellidic or sympodial conidiogenous<br />

cells, representing apical wall-building type<br />

(compared to ring wall-building and diffused wallbuilding)<br />

(Nag Raj 1993), which may indicate that the<br />

wall-building type probably has phylogenetic significance.<br />

Molecular phylogeny of <strong>Pleosporales</strong><br />

Numerous genes have been applied in phylogenetic<br />

studies of <strong>Pleosporales</strong>, mostly including LSU, SSU,<br />

mtSSU and ITS as well as the protein genes, such as<br />

RPB1, RPB2, TEF1, β-tubulin (TUB1) and actin (ACT1).<br />

A single gene such as ITS or LSU, has been used to study<br />

phylogenetic relationships between Leptosphaeria and<br />

Phaeosphaeria (Câmara et al. 2002) or Pleosporaceae<br />

and Tubeufiaceae (Kodsueb et al. 2006a, b) (Table2). The<br />

use of these phylogenetic markers, although making<br />

important contributions, has not been successful in<br />

resolving numerous relationships in single gene dendrograms.<br />

One exception is the use of SSU sequences to<br />

demonstrate the phylogenetic significance of pseudoparaphyses<br />

(Liew et al. 2000) whilst rejecting the phylogenetic<br />

utility of pseudoparaphyses morphology (cellular or<br />

trabeculate). Analyses with combined genes have had<br />

more success. For instance combined analyses with LSU<br />

and SSU sequence data could be used to define family<br />

level classification in a few cases (Dong et al. 1998; de<br />

Gruyter et al. 2009; Lumbsch and Lindemuth 2001; Pinnoi<br />

et al. 2007; Zhang et al. 2009b) (Table2). The addition of<br />

more than two genes has been used to determine relationships<br />

between orders. For instance, genes such as LSU,<br />

SSU and mtSSU have been used to analyze ordinal<br />

relationships in Loculoascomycetes (Lindemuth et al.<br />

2001), and to analyze phylogenetic relationships of<br />

coprophilous families in <strong>Pleosporales</strong> (Kruys et al.<br />

2006). Phaeocryptopus gaeumannii (T. Rohde) Petr. was<br />

showntobelonginDothideales based on LSU, SSU and<br />

ITS sequence analysis (Winton et al. 2007), while Schoch<br />

et al. (2006) used four genes, i.e. LSU, SSU, RPB2 and<br />

TEF1 to evaluate the phylogenetic relationships among<br />

different orders of the Dothideomycetes. Fivegenes,viz.<br />

LSU, SSU, TEF1, RPB1 andRPB2, were used to study the<br />

phylogenetic relationships of different orders within<br />

Dothideomycetes (Schoch et al. 2009) and of different<br />

families within <strong>Pleosporales</strong> (Zhang et al. 2009a)<br />

(Table 2). It is clear that even more genes will be required<br />

to address the remaining issues and the promise of genome<br />

analyses is within reach (www.jgi.doe.gov/sequencing/<br />

why/dothideomycetes.html) forDothideomycetes.<br />

The importance of generic type specimens<br />

The type specimen (collection type) is a fundamental<br />

element in the current Code of Botanical Nomenclature at<br />

familial or lower ranks (Moore 1998). A type specimen<br />

fixes the name to an exact specimen at family, genera,<br />

species and variety/subspecies rank and is ultimately based<br />

on this single specimen, i.e. a family name is based on a<br />

genus, the genus name is based on a species, and the<br />

species name is based on a specimen (Kirk et al. 2008).<br />

The generic type is of great importance in defining<br />

generic circumscriptions in fungal taxonomy. The generic<br />

types of <strong>Pleosporales</strong> have been studied previously by<br />

many mycologists. For instance, Müller and von Arx<br />

(1962) studied the generic types of “Pyrenomycetes”, and<br />

described and illustrated them in detail. Sivanesan (1984)<br />

described and illustrated the generic representatives of<br />

Loculoascomycetes for both their teleomorphs and anamorphs,<br />

and their links were emphasized. A large number<br />

of pleosporalean genera have been studied by Barr (1990a,<br />

b). Almost all of the previous work was conducted more<br />

than 20 years ago, when no molecular phylogenetic studies<br />

could be carried out and thus had been carried out in a<br />

systematic fashion.<br />

Aim and outline of present study<br />

The present study had two principal objectives:<br />

1. To explore genera under <strong>Pleosporales</strong> based on the<br />

generic types and provide a detailed description and<br />

illustration for the type species of selected genera,<br />

discuss the study history of those genera, and explore<br />

their ordinal, familial, and generic relationships;

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