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Introduction to Fungi, Third Edition

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566 HOMOBASIDIOMYCETES<br />

and their potential biotechnological applications.<br />

There is a conidial state called<br />

Sporotrichum pulverulentum (Burdsall, 1981). Little<br />

is known about the ecological role of P. chrysosporium<br />

because this species is rarely found in<br />

nature.<br />

19.7 Russuloid clade<br />

The russuloid clade is probably the most confusing<br />

group in the eight-clade system of Hibbett<br />

and Thorn (2001), containing the complete range<br />

of hymenophore types shown in Table 19.1 and<br />

Fig. 19.1. The phylogeny within this clade has<br />

been examined in detail by Larsson and Larsson<br />

(2003), who found that the main character uniting<br />

all members is the presence of gloeocystidia,<br />

i.e. cystidia filled with light-refractile (lipidrich)<br />

contents (Fig. 19.29b). In Stereum, these are<br />

modified as laticiferous hyphae (Fig. 19.27). Other<br />

characters commonly used <strong>to</strong> identify russuloid<br />

fungi are diagnostic only at lower taxonomic<br />

ranks, e.g. the ornamented walls of basidiospores<br />

and their starch-positive (amyloid) staining<br />

with Melzer’s iodine. We shall study representatives<br />

of agaricoid basidiocarps (Russulaceae),<br />

polyporoid fruit bodies (Heterobasidion), the<br />

spine-bearing Auriscalpium and corticioid forms<br />

(Stereaceae). The family arrangement is still<br />

tentative, and we give family names only where<br />

these have a fair chance of survival in future<br />

classification schemes.<br />

19.7.1 Russulaceae<br />

There are two important genera, Lactarius and<br />

Russula, and they share many features such<br />

as being ec<strong>to</strong>mycorrhizal species, and having<br />

a characteristically brittle texture <strong>to</strong> the gills<br />

and general fruit bodies due <strong>to</strong> the presence<br />

of sphaerocysts embedded in the tissue (see<br />

Fig. 19.4). Basidiospores are typically amyloid<br />

and ornamented with warts or ridges.<br />

Russula (c. 750 spp.)<br />

The genus Russula presents a similar taxonomic<br />

nightmare <strong>to</strong> Cortinarius, and it is unlikely that<br />

this genus is monophyletic. Species are widely<br />

distributed and ec<strong>to</strong>mycorrhizal. The basidiocarps<br />

(Fig. 19.26a) are moderate <strong>to</strong> large, often<br />

with a brightly coloured upper cap surface<br />

(white, yellow, green, red, purple or black). The<br />

gills are straight, arranged in a crowded but<br />

regular pattern, and vary from white <strong>to</strong> strawcoloured.<br />

The spores are ornamented with a<br />

network of branched ridges or plates. Some<br />

common species are R. ochroleuca (edible),<br />

R. fellea (bitter <strong>to</strong> the taste and inedible) and the<br />

sickener, R. emetica (poisonous). Russula ochroleuca<br />

fruits under coniferous and deciduous trees,<br />

R. fellea under beech (Fagus sylvatica) and R. emetica<br />

under pines.<br />

Lactarius (c. 400 spp.)<br />

The common name for Lactarius is milk cap,<br />

referring <strong>to</strong> the milky juice which exudes when<br />

the flesh of the cap is broken. The juice varies in<br />

colour from white <strong>to</strong> yellow, orange or violet<br />

and may change after exposure <strong>to</strong> the air. For<br />

example, the juice of L. deliciosus (saffron milk<br />

cap; Plate 10c) is carrot-coloured at first, but<br />

turns bright green upon prolonged exposure <strong>to</strong><br />

air. The juice also varies in taste, being mild <strong>to</strong><br />

faintly bitter in L. quietus, hot and acrid in<br />

L. pyrogalus, or first mild and then acrid in L.<br />

rufus. The juice is contained within broad<br />

laticiferous hyphae (see Fig. 19.4). As in Russula,<br />

the flesh also contains clusters of sphaerocysts.<br />

The gills are generally decurrent and the spores<br />

are ornamented like those of Russula. Some<br />

species of Lactarius have a narrow range of<br />

mycorrhizal partners, e.g. L. <strong>to</strong>rminosus (woolly<br />

milk cap) and L. turpis (ugly milk cap) are<br />

associated with birch (Betula), and L. deliciosus<br />

and L. deterrimus with Pinus and Picea. There<br />

are several species with edible basidiocarps,<br />

but some are poisonous or their edibility is<br />

unknown. Fruit bodies of L. deliciosus are especially<br />

valued, and it is possible <strong>to</strong> produce them<br />

in plantations of artificially inoculated<br />

host trees.<br />

19.7.2 Bondarzewiaceae<br />

Heterobasidion, a seemingly typical polyporoid<br />

wood-degrading bracket fungus, has been<br />

placed in the russuloid clade in several phylogenetic<br />

analyses. Some authors have assigned it <strong>to</strong>

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