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Introduction to Fungi, Third Edition

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OPHIOSTOMATALES<br />

367<br />

Fig12.37 Asexual reproduction in Ophios<strong>to</strong>ma ulmi. (a,b) Synnema<strong>to</strong>us conidiophores (Graphium ulmi). Parallel bundles of dark<br />

hyphae branch at their tips <strong>to</strong> produce holoblastic conidia which accumulate in a sticky drop. (c,d) Mononema<strong>to</strong>us synanamorph<br />

terminating in conidiogenous cells with a succession of holoblastic conidia which, on detachment, leave a protruding scar or denticle.<br />

In d some of the conidia show yeast-like budding. (e) An older hypha within which a new hypha and endoconidia have developed<br />

(from Harris & Taber,1973).<br />

based on their vegetative incompatibility characteristics,<br />

showed that they can be separated<br />

in<strong>to</strong> two distinct vc (vegetative compatibility)<br />

supergroups. One biotype, centred on the<br />

Romania Moldova Ukraine region of Europe,<br />

was designated the Eurasian or EAN vc supergroup,<br />

whilst the other, centred on the Southern<br />

Great Lakes region of North America, has<br />

been designated the North American or NAN vc<br />

supergroup. They have been formally recognized<br />

as distinct subspecies, O. novo-ulmi subsp. novoulmi<br />

for the EAN strains and subsp. americana<br />

for the NAN strains. There are morphological<br />

differences between the perithecia of the two<br />

subspecies, those of subspecies novo-ulmi having<br />

longer necks than those of subspecies americana<br />

(Brasier & Kirk, 2001). Naturally occurring<br />

hybrids between the two subspecies have been<br />

detected. Rare interspecific hybrids between<br />

O. ulmi and O. novo-ulmi also occur where the<br />

former species is being replaced by the latter<br />

(Brasier et al., 1998). Hybridization between introduced<br />

pathogens permits their rapid evolution<br />

(Brasier, 2001). Ophios<strong>to</strong>ma himal-ulmi is endemic<br />

<strong>to</strong> the Himalayas and has been distinguished as<br />

a third species (Brasier & Mehrotra, 1995).<br />

Fertilized females of bark-boring beetles,<br />

possibly carrying ascospores or conidia, excavate<br />

tunnels beneath the bark of living trees, often<br />

those already weakened by the disease, <strong>to</strong> lay<br />

a cluster of eggs. After hatching, the developing<br />

larvae also make tunnels beneath the bark<br />

radiating outwards from the egg chamber. They<br />

feed on the infected wood, and the galleries<br />

which they excavate are often lined by synnemata<br />

or perithecia, so that conidia and ascospores<br />

become attached <strong>to</strong> their mouthparts and<br />

bodies. Young, sexually immature adults emerge<br />

in the spring and summer. They fly <strong>to</strong> the<br />

crotches (branch points) of young twigs where<br />

they feed on bark before maturation and<br />

mating. During this twig-feeding stage spores<br />

of the fungal pathogen may be introduced in<strong>to</strong><br />

the host sapwood, and from these multiple<br />

inoculation points the fungus may spread in<strong>to</strong><br />

host tissues by mycelial growth or by movement

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