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Introduction to Fungi, Third Edition

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558 HOMOBASIDIOMYCETES<br />

fungus is actually parasitic, and quite possibly it<br />

is merely stimulated <strong>to</strong> fruit by the presence of<br />

the earth ball.<br />

Leccinum (75 spp.)<br />

Like Boletus spp., the genus Leccinum has large<br />

fleshy basidiocarps. A characteristic feature is<br />

that the stem is covered with scales composed of<br />

cystidia. Leccinum scabrum (brown birch bolete)<br />

and L. versipelle (orange birch bolete) are common<br />

ec<strong>to</strong>mycorrhizal associates of birch (Betula).<br />

Basidiocarps of Leccinum are generally edible.<br />

19.5.3 Suillaceae<br />

Suillus (90 100 spp.)<br />

Suillus species have medium-sized fleshy basidiocarps<br />

forming ec<strong>to</strong>mycorrhizal associations with<br />

conifers. Their fruit bodies are usually of yellowish<br />

colours due <strong>to</strong> the abundance of pigments<br />

derived from the shikimic acid pathway, especially<br />

grevillins (Fig. 19.22d; Besl & Bresinsky,<br />

1997). The cap may be dry and scaly but is more<br />

usually viscid or slimy. There are species with a<br />

ring on the stem (Fig. 19.21d), e.g. S. grevillei<br />

(larch bolete) and S. luteus (slippery jack), whilst<br />

in many others (e.g. S. bovinus, S. granulatus) there<br />

is no ring (Plate 9g). The spores are smooth<br />

and elongate, and pale brown <strong>to</strong> brown. The<br />

presence of bundles of cystidia in the hymenium<br />

is a feature which distinguishes this genus<br />

from other boletes, and the classification in<strong>to</strong> a<br />

separate family is supported by phylogenetic<br />

studies (e.g. Grubisha et al., 2001). Also included<br />

in the Suillaceae are the gasteromycete genus<br />

Rhizopogon (see p. 581) and the gill-bearing<br />

Gomphidius.<br />

The distribution of species mirrors that of<br />

their mycorrhizal hosts, coniferous trees which<br />

are largely confined <strong>to</strong> North-temperate regions,<br />

only extending <strong>to</strong> other areas where introduced.<br />

There is a fairly high degree of host specificity.<br />

For example, in nature S. grevillei (Fig. 19.21d) is<br />

almost exclusively associated with Larix spp.<br />

(larch) whilst S. bovinus and S. luteus are associated<br />

with Pinus spp. This ecological specificity is<br />

probably associated with the effects of competition<br />

because in the labora<strong>to</strong>ry, under aseptic<br />

conditions, a wider range of hosts has been<br />

infected experimentally. Many species of<br />

Suillus have edible basidiocarps and some areas<br />

of planted pine (P. radiata) forests in South<br />

America are devoted <strong>to</strong> production of S. luteus,<br />

with as<strong>to</strong>nishing annual productivity values of<br />

up <strong>to</strong> 1 t dry weight ha 1 reported (Dahlberg &<br />

Finlay, 1999). Suillus bovinus and S. variegatus are<br />

common in Swedish pine plantations. Whereas<br />

S. bovinus is an early-stage mycorrhizal fungus,<br />

S. variegatus is often found in older stands.<br />

In open communities dominated by seedling<br />

pines, S. bovinus first develops a large number of<br />

genets (genetically defined mycelial individuals)<br />

derived from basidiospores. Once established,<br />

a colony extends by mycelial spread and by<br />

rhizomorphs <strong>to</strong> nearby host roots. The extent of a<br />

genet can be estimated by evidence of somatic<br />

incompatibility between mycelial isolates made<br />

from basidiocarps. As the original genets expand,<br />

they compete with each other. The number of<br />

genets decreases, but as the genets increase in<br />

size they may fragment so that parts of the same<br />

genet may become separated. The maximum<br />

rate of mycelial extension has been estimated <strong>to</strong><br />

be about 20 cm per annum and the age of the<br />

largest genet <strong>to</strong> be about 75 years (Dahlberg &<br />

Stenlid, 1990, 1994; Dahlberg, 1997).<br />

19.5.4 Coniophoraceae<br />

This group includes wood-rotting fungi such<br />

as Serpula and Coniophora which form<br />

spreading, crust-like resupinate fructifications.<br />

Morphological characteristics suggested that the<br />

Coniophoraceae have an affinity with Boletales<br />

(Pegler, 1991), and Hibbett and Thorn (2001)<br />

have included them in their bolete clade. This is<br />

supported by the presence of pulvinic acids in<br />

both Serpula and Coniophora (Gill & Steglich,<br />

1987). Here we shall consider only Serpula.<br />

Serpula<br />

Serpula lacrymans (Figs. 19.21e,f) causes dry rot<br />

and is one of the most serious agents of timber<br />

decay in buildings. There is a very extensive<br />

literature (see Rayner & Boddy, 1988; Jennings &<br />

Bravery, 1991). Both hardwoods and softwoods<br />

are attacked but, because softwoods are more<br />

commonly used in building construction, it is on

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