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while the plant was still alive. Classic examples of this
are the stem base fungi such as the take-all fungus
(Gaeumannomyces graminis, Chapter 12) and the eyespot
fungus (Tapesia yallundae) of cereal crops. The inocu-
lum surviving from the previous growing season can
then initiate infection of a subsequent cereal crop.


Pioneer saprotrophic fungi

Several species of Mucor, Rhizopus and other
Zygomycota are commonly found in soils or fecal-
enriched materials, and also in the rhizosphere (root
zone) of plants, where they utilize sugars and other
simple soluble nutrients. Several Pythiumspecies also
fall into this category because they colonize fresh
plant residues in soil, although they are best known
as pathogens of living root tips and other fleshy plant
tissues (Chapter 12).
The spores of these and similar fungi germinate and
grow rapidly in response to soluble nutrients, and
within a few days they produce a further batch of asex-
ual dispersal spores or sexual resting spores (see Fig. 2.8).
They usually cannot degrade the more complex struc-
tural polymers (cellulose, etc.). In laboratory culture they
are found to be intolerant of the antibiotics or growth
metabolites of other fungi, and they do not themselves
produce antibiotics. So, these pioneer fungi typically
have a short exploitative phase and a high competit-
ive ability.
The sapstain fungi that rapidly colonize newly
felled trees are typical pioneer saprotrophs. They grow
in the nonwoody (parenchymatous) medullary rays
and they discolor the wood by their darkly pigmented

hyphae (Fig. 11.8). Although they cause no structural
damage to the timber, they can seriously reduce its
marketable value. These staining fungi include sev-
eral species of Ophiostoma(related to Ophiostoma ulmi
which causes Dutch elm disease; Chapter 10).
Vanneste et al. (2002) describe how natural products
and biological control agents are being developed to
control the sapstain fungi in New Zealand. However,
similar “bluestain” fungi such as Chlorosplenium aerug-
inascens (Ascomycota) cause an attractive turquoise
pigmentation of wood. This used to be highly valued
for producing inlaid veneers in decorative objects, termed
Tunbridge Ware because the industry was based in the
Kentish town of Tunbridge Wells.

Polymer-degrading fungi

Several fungi have an extended phase of growth on the
major structural polymers such as cellulose, hemicellu-
loses, or chitin. Once established, these fungi tend to
defend the resource against potential invaders, either
by sequestering a critically limiting nutrient such
as nitrogen or by producing inhibitory metabolites
(Chapter 6).
The large number of fungi in this category differ
in the types of substrate they utilize, their different
environmental requirements, and their growth in
different phases of a decomposition sequence. The
following examples illustrate this diversity.
When cereal straw and other cellulose-rich materials
are buried in soil they are colonized by fungi such
as Fusarium, Chaetomium, Stachybotrys, Humicola, or
Trichodermaspp. The Fusariumspp. tend to be favored

220 CHAPTER 11

Fig. 11.8(a,b) Darkly pigmented sapstain fungi growing in the medullary rays of felled trees.

(a) (b)
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