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Introduction to Fungi, Third Edition

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EUAGARICS CLADE<br />

551<br />

Control of witches’ broom disease is difficult<br />

but crop losses can be reduced by sanitation,<br />

i.e. the regular removal of brooms and diseased<br />

pods. Other measures involve the use of<br />

Trichoderma isolates as agents of biological<br />

control. Some are antagonistic, but T. stromaticum<br />

is mycoparasitic on mycelium and basidiocarps<br />

(Samuels et al., 2000; Sanogo et al., 2002).<br />

Systemic fungicides are also used. Further,<br />

there is an ongoing search for hosts resistant <strong>to</strong><br />

the pathogen which could be used for breeding<br />

purposes. The origin of the cocoa pathogen is<br />

thought <strong>to</strong> be native (wild) species of Theobroma<br />

growing in the forests, but other C. perniciosa<br />

strains attack members of the Solanaceae, Bixa<br />

spp., lianas (climbers) and woody debris on the<br />

forest floor.<br />

A closely related fungus, C. roreri<br />

(Myceliophthora roreri), causes the damaging<br />

frosty pod rot disease of cacao. Infected pods<br />

are covered by white powdery masses of winddispersed<br />

spores, originally regarded as conidia.<br />

These spores are borne on a dikaryotic mycelium<br />

which can be grown in culture. Dikaryotic<br />

hyphae swell and branch <strong>to</strong> form sporophore<br />

initials. The nuclei in the dikaryon fuse and septa<br />

are laid down, resulting in chains of diploid cells<br />

which have been interpreted as the equivalent of<br />

probasidia. The diploid cells develop thick walls<br />

<strong>to</strong> become spores and their nuclei undergo<br />

meiosis, but division may be arrested at a<br />

binucleate state (after the first meiotic division)<br />

or proceed <strong>to</strong> the formation of nuclear tetrads.<br />

On germination, there are indications that a<br />

four-celled metabasidium may be produced, with<br />

sterigmata which function as infective hyphae.<br />

As in C. perniciosa, this monokaryophase is biotrophic<br />

and can only be grown on living cocoa<br />

tissue. The life cycle of C. roreri is thus considerably<br />

modified, and a recognizable mushroomlike<br />

basidiocarp stage probably does not occur<br />

(Evans et al., 2002, 2003; Griffith et al., 2003).<br />

19.4.10 Mycenaceae<br />

Mycena (about 150 spp.)<br />

This is a large polyphyletic genus of fungi, with<br />

most species clustering in phylogenetic analyses<br />

around the type-species, Mycena galericulata. This<br />

group has been called Mycenaceae by Moncalvo<br />

et al. (2002). Mycena spp. produce rather small,<br />

delicate basidiocarps which have long slender<br />

stipes and conical or bell-shaped caps. Fruit<br />

bodies may emerge singly or in clusters from<br />

wood, leaf litter and other debris such as twigs,<br />

pine cones and bracken petioles in woodland and<br />

pastures. Some species exude latex when the<br />

stipe is broken, e.g. M. sanguinolenta with blood<br />

red latex and M. galopus which produces a<br />

milk white exudate. Much is known about the<br />

autecology of M. galopus, which has a perennial<br />

mycelium when growing in coniferous litter and<br />

also fruits on a wide range of twiggy debris but<br />

does not grow in bulky wood masses, possibly<br />

because of restricted aeration there (Frankland,<br />

1984; Dix & Webster, 1995). It is capable of<br />

growing on most of the constituents of leaf litter<br />

and its ability <strong>to</strong> break down lignin and cellulose<br />

enables it <strong>to</strong> function as a typical white-rot decay<br />

fungus. Mycena galopus is a key decomposer of<br />

oak leaves and, within two years of leaf fall,<br />

its mycelium may be present on 80% of fallen<br />

leaves. This species is regarded as a secondary<br />

colonizer, growing on plant material which is<br />

already well-colonized by other fungi. In this<br />

context, it is worth mentioning that many<br />

Mycena spp. produce antifungal metabolites,<br />

including strobilurins, and these may aid in the<br />

displacement of other wood-rotting fungi.<br />

Despite this, moribund fruit bodies of various<br />

Mycena spp., including strobilurin producers,<br />

may be parasitized by the zygomycete Spinellus<br />

fusiger (Plate 3e).<br />

Basidiocarp development in M. stylobates<br />

growing on beech leaves has been described<br />

and well-illustrated by Walther et al. (2001).<br />

An irregular arrangement of interwoven<br />

hyphae within the leaf bursts through <strong>to</strong> form<br />

an ovoid structure at the surface, composed<br />

mainly of vertically arranged hyphae. Cells at the<br />

margin increase in diameter and enclose the<br />

early stages of the primordium entirely.<br />

Separation of this large-celled wrapping tissue<br />

from internal hyphae results in the formation of<br />

a ring-like groove at the base of the primordium<br />

and a layer of protective hyphae covering a<br />

central bulb. The cells of the outer layer of<br />

vertically arranged hyphae increase in diameter

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