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Introduction to Fungi, Third Edition

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420 HYMENOASCOMYCETES: PEZIZALES (OPERCULATE DISCOMYCETES)<br />

Fig14.4 Oedocephalum conidial state of Peziza subviolacea<br />

(¼ P. praetervisa). (a) Conidiophores terminating in a<br />

club-shaped vesicle bearing numerous dry blas<strong>to</strong>conidia.<br />

(b) Details of developing conidia (<strong>to</strong>p) and a vesicle from which<br />

the conidia have been detached (bot<strong>to</strong>m). (c) Conidia, two of<br />

which are germinating. Scale bar ¼ 40mm (a,b) and 20 mm(c).<br />

From Webster et al. (1964), with permission from Elsevier.<br />

coprophilous, growing on the dung of herbivorous<br />

animals, but A. carbonarius grows on old<br />

bonfire sites. Common coprophilous species<br />

are A. furfuraceus (¼ A. stercorarius) which is very<br />

commonly found on old cattle dung, often<br />

along with A. immersus (Figs. 14.5 and 14.6,<br />

respectively). Whilst these species are heterothallic,<br />

some others, e.g. A. crenulatus (¼ A. viridulus),<br />

are homothallic. Characteristic features of all<br />

species are the purple colour of the ascospores<br />

and the protruding, operculate asci. Ascobolus<br />

furfuraceus forms yellowish saucer-shaped apothecia<br />

up <strong>to</strong> 5 mm in diameter, and when mature<br />

the surface of the apothecium is studded with<br />

purple dots which mark the ripe asci (Plate 6d).<br />

As the asci mature they elongate above the<br />

general level of the hymenium. The ascus tips<br />

are pho<strong>to</strong>tropic and this ensures that when they<br />

explode the spores are thrown away from<br />

the dung. The ascospores have a mucilaginous<br />

perispore which aids attachment. Ascobolus<br />

immersus has yellow globose apothecia about<br />

1 2 mm in diameter, with very large ascospores<br />

(about 70 30 mm). The perispores cause all<br />

the eight ascospores <strong>to</strong> adhere <strong>to</strong> form a single<br />

projectile about 250 mm long, capable of being<br />

discharged for up <strong>to</strong> 30 cm horizontally. In<br />

general, multi-spored projectiles have a lower<br />

surface-<strong>to</strong>-volume ratio and are projected further<br />

than single spores (see p. 317 for Sordaria).<br />

There is a general trend among coprophilous<br />

fungi <strong>to</strong>wards multi-spored projectiles. In the<br />

genus Saccobolus, which also belongs <strong>to</strong> the<br />

Ascobolaceae, all eight spores are firmly cemented<br />

<strong>to</strong>gether by their perispores.<br />

The spores of Ascobolus become attached <strong>to</strong><br />

herbage and, when eaten by a herbivore, germinate<br />

in the faeces. It is likely that digestion<br />

stimulates spore germination. Most spores fail<br />

<strong>to</strong> germinate on nutrient media but can be<br />

triggered <strong>to</strong> do so by treatment with 0.4% NaOH<br />

or bile salts, and incubation at 37°C. The purple<br />

pigment in the spore wall develops late and is<br />

deposited within the perispore from the ascus<br />

epiplasm. Immature spores are colourless. The<br />

spore wall bears longitudinal colourless striations<br />

in some species, e.g. A. immersus (Fig. 14.5)<br />

and A. furfuraceus (Fig. 14.6). Both species can be<br />

grown and induced <strong>to</strong> form apothecia in culture<br />

(see Webster & Weber, 2001).<br />

14.5.1 Mating behaviour<br />

There is variation in the mating behaviour of<br />

different species of Ascobolus. A single ascospore<br />

culture of A. scatigenus (¼ A. magnificus) does not<br />

produce apothecia. Sex organs (coiled ascogonia<br />

and antheridia) are formed only when mycelia<br />

of different mating types are grown <strong>to</strong>gether.<br />

Each strain is hermaphroditic, i.e. is capable<br />

of developing both ascogonia and antheridia.<br />

However, A. scatigenus is self-incompatible,<br />

i.e. the antheridia of one strain do not fertilize<br />

the ascogonia borne on the same mycelium.<br />

The ascospores of this fungus are of two types, A

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