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Pleosporales - CBS - KNAW

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Fungal Diversity<br />

Table 1 Major circumscription changes of <strong>Pleosporales</strong> from 1955 to 2011<br />

References<br />

Luttrell 1955<br />

Müller and von Arx 1962<br />

Luttrell 1973<br />

Barr 1979a<br />

Barr 1987b<br />

Kirk et al. 2001, 2008<br />

Boehm et al. 2009a, b; Mugambi<br />

and Huhndorf 2009b; Schoch et<br />

al. 2009; Shearer et al. 2009;<br />

Suetrong et al. 2009; Tanaka et<br />

al. 2009; Zhang et al. 2009a<br />

Circumscriptions distinguishing <strong>Pleosporales</strong> from other orders of Dothideomycetes<br />

Pleospora-type centrum development.<br />

Ascomata perithecoid, with rounded or slit-like ostiole; asci produced within a locule, arranged regularly<br />

in a single layer or irregularly scattered, surrounded with filiform pseudoparaphyses, cylindrical,<br />

ellipsoidal or sac-like.<br />

Ascocarps perithecioid, immersed, erumpent to superficial on various substrates, asci ovoid to mostly<br />

clavate or cylindrical, interspersed with pseudoparaphyses (sometimes form an epithecium) in mostly<br />

medium- to large-sized locules.<br />

Saprobic, parasitic, lichenized or hypersaprobic. Ascomata perithecioid, rarely cleistothecioid or<br />

hysterothecioid, peridium pseudoparenchymatous, pseudoparaphyses cellular, narrow or broad,<br />

deliquescing early at times, not forming an epithecium, asci oblong, clavate or cylindrical, interspersed<br />

with pseudoparaphyses, ascospores mostly asymmetric.<br />

Saprobic, biotrophic or hemibiotrophic. Ascomata globose, subglobose or conical, asci bitunicate, oblong,<br />

clavate or cylindrical, cellular pseudoparaphyses, ascospores hyaline or pigmented, asymmetric or<br />

symmetric, with or without septa.<br />

Ascomata perithecioid or rarely cleistothecioid, sometimes clypeate, mostly globose, thick-walled,<br />

immersed or erumpent, black, sometimes setose, peridium composed of pseudoparenchymatous cells,<br />

pseudoparaphyses trabeculate or cellular, asci cylindrical, fissitunicate, with a well-developed ocular<br />

chamber, rarely with a poorly defined ring (J-), ascospores hyaline to brown, septate, thin or thickwalled,<br />

sometimes muriform, usually with sheath, anamorphs hyphomycetous or coelomycetous.<br />

Hemibiotrophic, saprobic, hypersaprobic, or lichenized. Habitats in freshwater, marine or terrestrial<br />

environment. Ascomata perithecioid, rarely cleistothecioid, immersed, erumpent to superficial, globose<br />

to subglobose, or lenticular to irregular, with or without conspicuous papilla or ostioles. Ostioles with or<br />

without periphyses. Peridium usually composed of a few layers of cells with various shapes and<br />

structures. Hamathecium persistent, filamentous, very rarely decomposing. Asci bitunicate, fissitunicate,<br />

cylindrical, clavate to obclavate, with or without pedicel. Ascospores hyaline or pigmented, ellipsoidal,<br />

broadly to narrowly fusoid or filiform, mostly septate.<br />

Morpho-characters used in taxonomy of <strong>Pleosporales</strong><br />

Sexual characters<br />

According to the Linnean classification system, reproductive<br />

structures are the most important criteria in plant<br />

taxonomy, and this proposal is widely applied in fungal<br />

taxonomy (Gäumann 1952). In the classification of Dothideomycetes,<br />

reproductive characters such as the uni- or<br />

multilocular nature and shape of ascomata, presence and<br />

shape of ostioles/papillae, shape and apical structures of<br />

asci and shape, pigmentation and septation of ascospores<br />

play important roles at different ranks (Clements and Shear<br />

1931; Luttrell 1951, 1955, 1973). Besides the common<br />

morphological characters possessed by Dothideomycetes<br />

(bitunicate and fissitunicate asci as well as the perithecioidlike<br />

ascostromata), most pleosporalean fungi also have<br />

pseudoparaphyses among their well-arranged asci (Zhang et<br />

al. 2009a). Currently, classification of <strong>Pleosporales</strong> at the<br />

family level focuses mostly on morphological characters of<br />

ascomata (such as size, shape of ostiole or papilla), presence<br />

or absence of periphyses, characters of centrum (such as asci,<br />

pseudoparaphyses and ascospores) as well as on lifestyle or<br />

habitat (Barr 1990a; Shearer et al. 2009; Suetrong et al.<br />

2009; Tanaka et al. 2009; Zhang et al. 2009a), whilst relying<br />

extensively on DNA sequence comparisons.<br />

Ascomata<br />

Most species of <strong>Pleosporales</strong> have uniloculate ascomata.<br />

The presence (or absence) and forms of papilla and ostiole<br />

are the pitoval character of ascomata, which serve as<br />

important characteristics in generic or higher rank classification<br />

(Clements and Shear 1931). The vertically flattened papilla has<br />

recently been shown as an effective criterion for familial level<br />

classification, e.g. in the Amniculicolaceae and the Lophiostomataceae<br />

(Zhang et al. 2009a). Papillae and ostioles are<br />

present in most species of <strong>Pleosporales</strong>, except in the<br />

Diademaceae and Sporormiaceae. Members of Diademaceae<br />

have apothecial ascomata, and some genera of Sporormiaceae<br />

have cleistothecioid ascomata. Another coprophilous pleosporalean<br />

family, Delitschiaceae, can be distinguished from<br />

Sporormiaceae by the presence of periphysate ostioles.<br />

Pseudoparaphyses<br />

Presence of pseudoparaphyses is a characteristic of<br />

<strong>Pleosporales</strong> (Kirk et al. 2008; Liew et al. 2000). Although<br />

pseudoparaphyses may be deliquescing in some families<br />

when the ascomata mature (e.g. in Didymellaceae), they are<br />

persistent in most of other pleosporalean members. According<br />

to the thickness, with or without branching and density<br />

of septa, pseudoparaphyses were roughly divided into two<br />

types: trabeculate and cellular, and their taxonomic significance<br />

need to be re-evaluated (Liew et al. 2000).

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