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Introduction to Fungi, Third Edition

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464 LOCULOASCOMYCETES<br />

Fig17.4 Stagonospora nodorum. (a) Pycnidia seen from above. (b) Pycnidia in section in agar culture. (c) Portion of wall of pycnidium<br />

showing origin of conidia. (d) Conidia. (a,b) <strong>to</strong> same scale; (c,d) <strong>to</strong> same scale.<br />

(Pedras & Biesenthal, 1998; Howlett et al., 2001).<br />

In addition, L. maculans is able <strong>to</strong> de<strong>to</strong>xify phy<strong>to</strong>alexins<br />

produced by the host plant (Pedras et al.,<br />

2000). Since P. lingam is not closely related <strong>to</strong><br />

most other Phoma spp., it has been re-named<br />

Plenodomus (Reddy et al., 1998).<br />

Diverse anamorphic states are known in<br />

other species of Lep<strong>to</strong>sphaeria and Phaeosphaeria,<br />

and some examples of important pathogens are<br />

summarized in Table 17.1.<br />

17.2.3 Ascochyta and Phoma<br />

Several related Ascochyta-type anamorphs infect<br />

legumes, causing diseases such as blight of<br />

chickpea (A. rabiei), foot rot and blight of peas<br />

(A. pinodes), and leaf- and pod-spot of broad beans<br />

(A. fabae) and peas (A. pisi). The pseudothecial<br />

state, where present, is now called Didymella<br />

(formerly Mycosphaerella). These species usually<br />

overwinter on crop residues, infecting new crops<br />

in spring as ascospores, but they are also seedborne.<br />

During the growing season, the infection<br />

is spread by rainsplash of pycnidiospores. An<br />

excellent review of the biology of Ascochyta,<br />

highlighting the severity of diseases which can<br />

be caused, has been written by Pande et al. (2005)<br />

for A. rabiei. Pycnidia contain hyaline, two-celled<br />

conidia which, according <strong>to</strong> Brewer and Boerema<br />

(1965), arise by septation from the conidiogenous<br />

cell. A few conidia with no, two or three<br />

septa may be produced occasionally (Fig. 17.5).<br />

Extensive work has been carried out on characterizing<br />

the mating type idiomorphs of<br />

Ascochyta spp. These data and analyses of other<br />

sequences have shown that Ascochyta spp. belong<br />

<strong>to</strong> the Pleosporales and are not closely related<br />

<strong>to</strong> Mycosphaerella, which is in the Dothideales<br />

(Barve et al., 2003).<br />

Phoma medicaginis (Fig. 17.6) has also been<br />

shown <strong>to</strong> be related <strong>to</strong> Ascochyta spp. (Fatehi et al.,<br />

2003). Unfortunately, little information is available<br />

<strong>to</strong> circumscribe the very large genus Phoma<br />

which is almost certainly polyphyletic, being<br />

associated with several different teleomorphs.<br />

However, a valuable and much-needed monograph<br />

of this difficult group has been written by<br />

Boerema et al. (2004). The conidial fructification<br />

of Phoma is a pycnidium which usually has only<br />

one ostiole, rarely two (Figs. 17.6a and 17.7b).<br />

Like most pycnidia, its outermost wall layer is<br />

pigmented and consists of cells of distinct shape,<br />

which is a useful feature in species identification.<br />

In the case of Phoma spp., it is called a<br />

textura angularis (Fig. 17.6a). The cavity of the<br />

pycnidium is lined by a hymenium of conidiogenous<br />

cells from which one-celled hyaline<br />

pycnidiospores develop. The absence of filiform

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