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Introduction to Fungi, Third Edition

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HYPOCREALES<br />

337<br />

Fig12.13<br />

serpens.<br />

The conidial (Geniculosporium)stateofHypoxylon<br />

Fig12.12 Hypoxylon. (a) Perithecial stromata of H. fragiforme<br />

on beech (Fagus sylvatica).One stroma has been broken open<br />

<strong>to</strong> show the perithecia embedded in the outer layer.<br />

(b) Perithecial stromata of H. multiforme on birch (Betula<br />

pendula).<br />

12.4 Hypocreales<br />

The Hypocreales are a large group of fungi,<br />

although estimates vary as <strong>to</strong> the number of taxa<br />

contained in it. Kirk et al. (2001) included 117<br />

genera and 654 species, whilst Rossman (1996)<br />

stated that there may be 2000 5000 holomorphic<br />

taxa. Some are known only as anamorphs.<br />

Hypocreales are characterized by pale or brightly<br />

coloured perithecia (or rarely cleis<strong>to</strong>thecia)<br />

which may be single or borne on or embedded<br />

in a fleshy stroma. The asci are unitunicate, with<br />

or without a well-defined apical apparatus.<br />

Perithecial development conforms <strong>to</strong> the<br />

‘Nectria’ type of Luttrell (1951). The ascogonia,<br />

which are formed within a stroma, become<br />

surrounded by concentric layers of vegetative<br />

hyphae which form a true perithecial wall. The<br />

cells of the inner wall layer in the apical region<br />

of the young perithecium produce a palisade of<br />

inward-growing hyphal branches. These hyphal<br />

branches grow downward <strong>to</strong> form a vertically<br />

arranged mass of hyphae with free ends termed<br />

apical paraphyses (Luttrell, 1965). Pressure<br />

exerted by the elongation of the apical paraphyses,<br />

accompanied by expansion of the wall,<br />

creates a central cavity within the perithecium.<br />

The free tips of the apical paraphyses ultimately<br />

push in<strong>to</strong> the lower portion of the wall so that<br />

they become attached at both the <strong>to</strong>p and<br />

bot<strong>to</strong>m of the perithecial cavity. Ascogenous<br />

hyphae arising from the ascogonium spread out<br />

across the floor and sides of the cavity and<br />

produce asci by means of croziers. The asci grow<br />

upward among the apical paraphyses and form

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