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Introduction to Fungi, Third Edition

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ASCOSPHAERALES<br />

287<br />

ascocarps, and it is thus an exception <strong>to</strong> the<br />

generalization which places such fungi in the<br />

Archiascomycetes (Chapter 9) or Hemiascomycetes<br />

(see Eremothecium, Galac<strong>to</strong>myces, Saccharomycopsis;<br />

Chapter 10). Support for the inclusion of<br />

Eremascus in the Euascomycetes comes not only<br />

from molecular data (Berbee & Taylor, 1992a;<br />

Anderson et al., 1998) but also from the presence<br />

of typical Euascomycete septa with one central<br />

pore and associated Woronin bodies (Kreger-van<br />

Rij et al., 1974). Further, the arthroconidia are<br />

delimited by a double-septum (Harrold, 1950)<br />

and are released by rhexolytic secession, which is<br />

typical of certain Plec<strong>to</strong>mycetes (see Fig. 11.3f).<br />

Two species are known, E. albus and E. fertilis<br />

(Fig. 11.1). Both are associated with sugary substrates<br />

such as mouldy jam, but several collections<br />

of E. albus have been made from powdered<br />

mustard. Harrold (1950) has shown that both<br />

fungi grow best on media with a high sugar<br />

content (e.g. 40% sucrose), but do not grow well<br />

in a water-saturated atmosphere. The mature<br />

mycelium consists of uninucleate segments.<br />

Both species are homothallic. On either side of<br />

a septum, short gametangial branches arise<br />

which are swollen at their tips and, in the case<br />

of E. albus, coil around each other. The gametangial<br />

tips of E. albus are usually uninucleate and,<br />

following breakdown of the wall separating the<br />

tips of adjacent gametangia, nuclear fusion<br />

occurs. This is followed by meiosis and mi<strong>to</strong>sis<br />

so that eight nuclei result, each one being<br />

surrounded by cy<strong>to</strong>plasm <strong>to</strong> form a uninucleate<br />

ascospore (Fig. 11.1m). The ascospores are<br />

dispersed passively following breakdown of the<br />

ascus wall. On germination a multinucleate germ<br />

tube emerges, but the uninucleate condition is<br />

soon established by the formation of septa.<br />

11.2.2 Ascosphaera<br />

Ascosphaera spp. are associated with bees and<br />

related insects, growing saprotrophically in their<br />

nests on the gathered pollen and nectar. They can<br />

be maintained in pure culture but commonly<br />

Fig11.1 Eremascus.(a d) Eremascus fertilis, stages in the development of asci. (e g) Eremascus albus, stages in the development<br />

of asci. Note the coiling of the gametangia and the globose ascospores of E. albus.(h m) Eremascus albus, nuclear behaviour<br />

during ascus formation (after Harrold,1950). (h) Uninucleate gametangia. (i) Plasmogamy and karyogamy. (k m) Nuclear divisions<br />

preceding ascospore formation.

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