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Introduction to Fungi, Third Edition

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OTHER MEMBERS OF THE PUCCINIACEAE<br />

631<br />

have been mentioned at the beginning of this<br />

chapter.<br />

Uromyces appendiculatus is a macrocyclic au<strong>to</strong>ecious<br />

rust on French bean (Phaseolus vulgaris) and<br />

many other leguminous plants. Several varieties<br />

have been distinguished. The fungus survives the<br />

winter as teliospores on plant debris or, in milder<br />

conditions, as urediniospores (McMillan et al.,<br />

2003). As one would expect of a situation in which<br />

a rust fungus undergoing sexual reproduction is<br />

being controlled by the breeding of resistant<br />

cultivars, numerous (more than 200) physiological<br />

races of U. appendiculatus are known. Control is<br />

by means of resistant cultivars or, if these fail,<br />

with protectant fungicides (maneb and chlorothalonil).<br />

Strobilurin-type compounds are also<br />

now used against U. appendiculatus. Since beans<br />

are a higher-value crop than cereals, fungicide<br />

applications are generally more profitable.<br />

Uromyces viciae-fabae has a similar life cycle <strong>to</strong><br />

that of U. appendiculatus, but it infects broad bean<br />

(Vicia faba) and numerous other plants. It is a<br />

cosmopolitan species but seems <strong>to</strong> cause lesser<br />

damage than U. appendiculatus.<br />

Uromyces pisi is heteroecious, producing spermogonia<br />

and aecia on Euphorbia cyparissias and<br />

uredinia and telia on various leguminous hosts,<br />

especially pea (Pisum sativum). Infections on the<br />

alternate host are of biological interest because<br />

the fungus infects systemically and causes its<br />

alternate host <strong>to</strong> produce pseudoflowers, at the<br />

expense of its real flowers (Pfunder & Roy, 2000).<br />

The name U. pisi is now known <strong>to</strong> cover a<br />

complex of several biologically distinct species<br />

which are united by their alternate host but<br />

infect different members of the Fabaceae as<br />

principal hosts (Pfunder et al., 2001).<br />

22.6 Other members of the<br />

Pucciniaceae<br />

22.6.1 Phragmidium<br />

All Phragmidium spp. are au<strong>to</strong>ecious and confined<br />

<strong>to</strong> the Rosaceae. The two best-known species,<br />

both extremely common in Europe, are P. violaceum<br />

on leaves of the Rubus fruticosus species<br />

aggregate (bramble or blackberry; Plate 12c) and<br />

P. mucronatum on roses. Both are macrocyclic. The<br />

uredinial and telial stages are readily recognized<br />

from above as chlorotic or purple lesions, and<br />

the spores are formed on the underside of the<br />

host leaves. They are yellowish-orange (urediniospores)<br />

or violet <strong>to</strong> black (teliospores). The<br />

teliospores contain mostly 4 (P. violaceum) or<br />

6 8 (P. mucronatum) cells with very dark and<br />

rough walls, and are borne on a long stalk<br />

(pedicel) which becomes easily detached from the<br />

lesions. When teliospores are mounted in water,<br />

the base of the pedicel breaks and exudes a large<br />

amount of mucilage (Figs. 22.13c,d). In nature,<br />

this may facilitate the attachment of the teliospore<br />

<strong>to</strong> adjacent surfaces during overwintering<br />

(Ingold et al., 1981). The teliospores of P. mucronatum<br />

were among the first fungus spores illustrated<br />

by Robert Hooke in 1667 soon after the<br />

invention of the microscope (see Large, 1940).<br />

Phragmidium violaceum shows promise as a<br />

biocontrol agent against the spread of brambles<br />

which were introduced in<strong>to</strong> Australia and are<br />

spreading there in their typically uncontrollable<br />

manner (Mahr & Bruzzese, 1998). Phragmidium<br />

mucronatum is not <strong>to</strong>lerated by rose breeders or<br />

hobby gardeners; it is controlled by fungicide<br />

applications. Removal of fallen leaves in autumn<br />

can also help <strong>to</strong> control the disease.<br />

22.6.2 Gymnosporangium<br />

There are about 60 species of Gymnosporangium<br />

which produce their spermogonia and aecia<br />

on members of the Rosaceae, and telia on<br />

Cupressaceae. Uredinia are not normally<br />

formed. One of the most commonly encountered<br />

species is G. fuscum (¼ G. sabinae), the pear trellis<br />

rust, which alternates between Juniperus spp. of<br />

the sabina group (principal host) and pear trees.<br />

The dikaryotic mycelium can survive for many<br />

years in the principal host, and repeated production<br />

of telia is associated with a spindle-shaped<br />

swelling or canker of the infected trunk or twig.<br />

Spore-bearing telia are produced in the spring<br />

as horn-like outgrowths which greatly expand<br />

during rainfall due <strong>to</strong> swelling of the long teliospore<br />

stalks (Plate 12d; Fig. 22.13f). Teliospores at<br />

the surface of a swollen telial horn germinate<br />

<strong>to</strong> release basidiospores at the time of leaf

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