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Introduction to Fungi, Third Edition

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332 HYMENOASCOMYCETES: PYRENOMYCETES<br />

Fig12.9 Chae<strong>to</strong>mium globosum. (a) Perithecium showing<br />

cirrhus of ascospores. (b) Asci. (c) Ascospores.<br />

other arthropods (Wicklow, 1979). When the<br />

perithecia are ripe, a column-like mass (cirrhus)<br />

of dark ascospores is extruded through the<br />

ostiole, supported by the perithecial hairs<br />

which surround it (Fig. 12.9a). The spore<br />

column results from the breakdown of the asci<br />

within the body of the perithecium, i.e. the asci<br />

do not discharge their spores violently. The<br />

young asci are cylindrical <strong>to</strong> club-shaped, but<br />

this stage is very evanescent and is only found<br />

in young perithecia (Fig. 12.9b), often before<br />

the spores have become pigmented and sometimes<br />

before the perithecium has developed an<br />

ostiole. Spores of most species are pale brown <strong>to</strong><br />

grey and lemon-shaped, with a single terminal<br />

germ pore.<br />

The development of perithecia in<br />

Chae<strong>to</strong>mium shows some variation between<br />

species (Whiteside, 1957, 1961). The ascogonia<br />

are coiled and lack antheridia. Investing hyphae<br />

arise from the ascogonial stalk or from vegetative<br />

cells surrounding the ascogonium.<br />

Perithecial hairs develop early from the external<br />

cell layer. The centrum is at first filled with<br />

hyaline pseudoparenchyma. At the apex of the<br />

perithecium, certain of these cells become<br />

meristematic and give rise <strong>to</strong> the elongate<br />

periphyses which line the inside of the ostiole.<br />

Ascogenous hyphae develop from the ascogonium<br />

at the base of the centrum and, at about<br />

this time, the surrounding pseudoparenchyma<br />

cells of the centrum deliquesce <strong>to</strong> form a cavity.<br />

In some species croziers develop, but in others<br />

they are absent. Paraphyses of two types have<br />

been described. In C. brasiliense, lateral paraphyses<br />

arise from the pseudoparenchyma cells<br />

outside the hymenium, whilst hymenial paraphyses<br />

have been illustrated in C. globosum<br />

(Whiteside, 1961). The paraphyses are evanescent<br />

and disappear as the asci mature.<br />

Most species of Chae<strong>to</strong>mium are homothallic,<br />

but a few, e.g. C. cochliodes, are heterothallic.<br />

Both homo- and heterothallic strains of C. elatum<br />

have been reported. Conidial states are rare in<br />

Chae<strong>to</strong>mium, but simple phialides and phialospores<br />

occur in C. elatum and C. globosum, whilst<br />

C. piluliferum forms both phialospores and<br />

globose thalloconidia of the Botryotrichum type<br />

(Daniels, 1961).<br />

12.3 Xylariales<br />

The order Xylariales is probably polyphyletic and<br />

comprises about 800 species in 8 families (Kirk<br />

et al., 2001). Members of this group produce dark<br />

perithecial stromata with asci which have an<br />

iodine-positive apical apparatus. Ascospores are<br />

typically melanized. Xylariales are saprotrophs<br />

or plant pathogens and are associated especially<br />

with the bark and wood of trees.<br />

We shall only consider one family, the<br />

Xylariaceae, which is by far the most important.<br />

The dark-coloured ascospores are generally<br />

smooth-walled and inaequilateral (one face<br />

being more strongly curved than the other),<br />

with a hyaline germ slit. Anamorphs are sometimes<br />

also stromatic, producing hyaline or<br />

lightly pigmented conidia holoblastically from<br />

a sympodially proliferating conidiogenous<br />

region, the sympodula (Fig. 12.13). They have

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