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Introduction to Fungi, Third Edition

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460 LOCULOASCOMYCETES<br />

Fig17.1 Sexualreproductionin Pleosporascirpicola. (a) Young ascus withundifferentiated cy<strong>to</strong>plasm.Note the thick ascus wall.<br />

(b)Maturingasciwithcy<strong>to</strong>plasmcleavedin<strong>to</strong>ascospores.(c)Matureascuswithmuriformascospores.Theascusis shownafewseconds<br />

before discharge; the ec<strong>to</strong>tunica hasruptured (arrow), and themore flexible endotunica hasbeenforced <strong>to</strong> expandby theincreasing<br />

turgor pressure. (d) Discharged ascospore mountedin Indian ink <strong>to</strong> show the mucilaginous extensions. Allimages <strong>to</strong> same scale.<br />

(Berbee, 1996; Dong et al., 1998; Silva-Hanlin &<br />

Hanlin, 1999). Molecular analyses based on a relatively<br />

small number of representatives of ascomycete<br />

groups suggest that the most closely related<br />

groups are Pyrenomycetes (perithecioid fungi)<br />

belonging <strong>to</strong> Diaporthales, Hypocreales, Microascales<br />

and Sordariales (see Chapter 12; Spatafora,<br />

1995). Several orders are currently included in<br />

the Loculoascomycetes (Barr & Huhndorf, 2001;<br />

Kirk et al., 2001) but we shall consider representatives<br />

only of the two most important, the<br />

Pleosporales and Dothideales. Because the<br />

arrangement of orders and especially families<br />

is still in a state of flux, we shall not discuss it.<br />

17.2 Pleosporales<br />

This is a large group of ascomycetes believed <strong>to</strong><br />

be monophyletic on the basis of molecular<br />

evidence (Berbee, 1996). Several economically<br />

important genera of plant pathogens are<br />

included, such as Cochliobolus, Phaeosphaeria and<br />

Pyrenophora, parasitic on grasses and cereals.<br />

Pleospora, Lewia and Lep<strong>to</strong>sphaeria are common<br />

endophytes, saprotrophs or parasites of other<br />

herbaceous plants. They and their anamorphs<br />

may also have significance as human allergens,<br />

as human pathogens and in the production<br />

of myco<strong>to</strong>xins. Sporormiella fruits on herbivore<br />

dung.<br />

The development of pseudothecia in these<br />

forms conforms <strong>to</strong> the Pleospora type of Luttrell<br />

(1951). Ascogonia arise within a stroma and,<br />

in the region of the ascogonia, a group of<br />

vertically arranged septate hyphae appears,<br />

each hypha arising as an outgrowth from a<br />

stromatal cell. These hyphae are capable of<br />

elongating by intercalary growth and are<br />

termed pseudoparaphyses (Luttrell, 1965).

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