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Introduction to Fungi, Third Edition

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THE ‘TRUE’ SMUT FUNGI (USTILAGINOMYCETES)<br />

639<br />

entire infected host organs or galls may be<br />

dispersed, and teliospores or mycelium of smut<br />

fungi may be borne on or in seeds.<br />

The most common dispersal route in the<br />

smut fungi is by wind-blown teliospores, and<br />

the transatlantic wind-borne passage of African<br />

sugarcane smut, Sporisorium scitamineum, has<br />

been proposed as the most likely route of entry<br />

in<strong>to</strong> the Caribbean. Teliospores of many smuts<br />

may adhere <strong>to</strong> the seeds of their host plants<br />

and germinate with them. Certain species causing<br />

loose smuts of cereals (see p. 639) systemically<br />

infect the living embryo of the cereal grain.<br />

Grains contaminated by external spore dusts or<br />

systemic infections can be spread by human<br />

transport. Insect dispersal is important in the<br />

anther smut, Microbotryum violaceum, which is<br />

related <strong>to</strong> the rust fungi (see p. 652).<br />

Fig 23.2 Teliospore surfaces of Ustilago spp. (a) Smooth<br />

surface in U. hordei. (b) Spiny surface in U. nuda.From<br />

Va¤nky (1994); original prints kindly provided by K.Va¤nky.<br />

23.2.3 Teliospore release and dispersal<br />

Piepenbring et al. (1998c) have written an<br />

excellent review of the teliospore as a dispersal<br />

unit in a wide range of smut fungi, and we can<br />

only summarize a few salient features here.<br />

Teliospores of most species are produced in a<br />

sorus enclosed by a thin layer of host tissue. They<br />

are generally released dry. Spore release is often<br />

by the simple rupture of the host epidermis<br />

surrounding the sorus, possibly as a result of<br />

pressure exerted by the expanding teliospores.<br />

The host epidermis may aid in the release of<br />

teliospores, e.g. if it is hit by water drops. The<br />

mechanism in this case is similar <strong>to</strong> that found<br />

in puffballs (see p. 578). Gusts of wind which<br />

shake infected host organs may also be effective<br />

in releasing smut spores over time. Alternatively,<br />

23.2.4 Ustilago species on<br />

monocotyledonous hosts<br />

Several Ustilago spp. cause diseases on grasses<br />

and cereals. Sori of the agriculturally most<br />

important species, U. hordei, U. nuda, U. tritici<br />

and U. avenae, are produced in place of the<br />

developing seeds. Other Ustilago spp. affect the<br />

leaves of their hosts, e.g. U. filiformis (formerly<br />

U. longissima), which causes leaf stripe smut<br />

on Glyceria spp. Ustilago maydis infects both<br />

vegetative and reproductive organs of its host<br />

(Plate 12h), causing gall-like deformations. This<br />

species is considered in detail on pp. 643 647.<br />

Many Ustilago spp. have also been described from<br />

dicotyledonous hosts, but it now seems that all<br />

of them belong <strong>to</strong> Microbotryum (Vánky, 1998,<br />

1999; Almaraz et al., 2002; see p. 652).<br />

Phylogenetic studies have shown U. hordei,<br />

U. nuda, U. tritici and U. avenae <strong>to</strong> be closely<br />

related <strong>to</strong> each other, <strong>to</strong> the point where it<br />

becomes difficult <strong>to</strong> distinguish individual<br />

species by DNA sequences and microscopy. They<br />

are regarded as the core species of Ustilago (S<strong>to</strong>ll<br />

et al., 2003). Since these species can hybridize<br />

with each other, it has been proposed that they<br />

should be merged in<strong>to</strong> one taxon, U. segetum<br />

(see Bakkeren et al., 2000). However, because<br />

the existing names are so well-established and<br />

because well-known plant diseases are caused by

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