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Introduction to Fungi, Third Edition

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ONYGENALES<br />

293<br />

disturb the soil. One hot spot of infection is the<br />

San Joaquín Valley in Southern California, where<br />

coccidioidomycosis is known as ‘valley fever’. The<br />

Californian population of C. immitis is reproductively<br />

isolated from populations elsewhere and<br />

has recently been given the status of a separate<br />

species, C. posadasii (Fisher et al., 2002). A readable<br />

account of the his<strong>to</strong>ry of C. immitis has been given<br />

by Odds (2003), and Cox and Magee (2004) have<br />

covered general aspects of its interactions with<br />

the mammalian host.<br />

Onygena<br />

This is perhaps the most unusual yet least<br />

researched member of the Onygenaceae. Its<br />

fructification (Fig. 11.4a) is interpreted as a<br />

stalked ascostroma, i.e. an aggregate of several<br />

gymnothecia at the tip of a sterile stalk which<br />

may be 1 cm or more in length. The stalks are<br />

pho<strong>to</strong>tropic during growth. At maturity, the<br />

peridium of the stroma ruptures, thereby exposing<br />

the ascospores. There are two species,<br />

O. corvina which is associated with animal hair<br />

and bird feathers, and O. equina growing on the<br />

hooves and horns of herbivorous mammals<br />

(Currah, 1985). Both species are strongly keratinolytic,<br />

and although little work has been<br />

published on their biological features, we can<br />

assume that keratinolysis proceeds as in other<br />

Onygenales. According <strong>to</strong> Kunert (2000), the key<br />

feature is the ability <strong>to</strong> use keratin as the sole<br />

source of both carbon and nitrogen. The vast<br />

surplus of nitrogen is released in<strong>to</strong> the environment<br />

as ammonia, thereby generating an alkaline<br />

pH of 9.0 or higher. The cystein-rich keratin also<br />

contains sulphur in excess of the growth requirements<br />

of keratinolytic fungi. This is often released<br />

as sulphate, thereby buffering the pH increase<br />

caused by the release of ammonia.<br />

Like many protein-degrading fungi, Onygena<br />

produces a cadaverous smell in culture, and<br />

Currah (1985) has suggested that this might<br />

attract carrion flies if produced in nature. This<br />

would make Onygena an insect-dispersed fungus.<br />

11.3.2 Ar throdermataceae<br />

There are only two genera in this small family.<br />

Ctenomyces (Fig. 11.5), with only one species<br />

(C. serratus), has a Chrysosporium anamorph<br />

in which one-celled microconidia are formed as<br />

terminal or intercalary cells of hyphae. Species of<br />

Arthroderma also sometimes produce Chrysosporium-like<br />

microconidia (Fig. 11.6b) but are better<br />

known by their macroconidial synanamorphs<br />

Epidermophy<strong>to</strong>n, Microsporum and Trichophy<strong>to</strong>n.<br />

These are multicellular with transverse septa,<br />

and are spindle-shaped or cylindrical. They are<br />

typical and readily recognized (Fig. 11.6a). The<br />

perfect state has not been found for many<br />

of these anamorphs, but they are suspected <strong>to</strong><br />

be phylogenetically close <strong>to</strong> Arthroderma (Gräser<br />

et al., 1999; Hirai et al., 2003). Howard et al. (2003)<br />

have given a useful summary of Arthroderma and<br />

its associated anamorphs, listing 47 species. The<br />

teleomorph of the Arthrodermataceae is a typical<br />

gymnothecium with a basket of branching and<br />

anas<strong>to</strong>mosing hyphae enclosing spherical asci<br />

which release their spores passively. Characteristic<br />

appendages are often present, e.g. in<br />

Ctenomyces (Fig. 11.5), a keratinolytic species associated<br />

with feathers. The combed appendages<br />

may serve <strong>to</strong> attach the gymnothecium <strong>to</strong> bird<br />

feathers for dispersal (Currah, 1985).<br />

Members of the Arthrodermataceae are generally<br />

keratinolytic, i.e. they degrade skin and hair.<br />

Howard et al. (2003) distinguished between species<br />

primarily associated with man (anthropophilic),<br />

animals (zoophilic) or the soil (geophilic). Because<br />

of their ability <strong>to</strong> grow on the skin, hair and nails<br />

of animals, the Arthrodermataceae are collectively<br />

called derma<strong>to</strong>phytes. Diseases caused by<br />

derma<strong>to</strong>phytes are colloquially known as ‘ringworm’,<br />

whereas they are called tinea within the<br />

medical profession, with descriptive terms such<br />

as capitis, barbae, corporis and pedis added <strong>to</strong><br />

describe mycoses of the scalp, beard, general<br />

body, or feet, respectively (Howard et al., 2003).<br />

These infections are usually confined <strong>to</strong> the outer<br />

(dead) skin regions and are relatively easily<br />

controlled either by the superficial (<strong>to</strong>pical)<br />

application of creams containing a wide variety<br />

of drugs, or by oral treatment especially with<br />

triazoles or terbinafine (see p. 278; Weitzman &<br />

Summerbell, 1995; Gupta et al., 1998). Griseofulvin,<br />

produced by Penicillium griseofulvum (see<br />

p. 302), was one of the first oral and <strong>to</strong>pical<br />

drugs and is still in use <strong>to</strong>day, especially in

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