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Phytopathologia Mediterranea Basidiomycetes and Esca Linked with the Occurrence of Basidiomycetes

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Phytopathologia Mediterranea Basidiomycetes and Esca Linked with the Occurrence of Basidiomycetes Phytopathol. Mediterr. (2006) 45, S30–S42 Biodiversity and geographic distribution of basidiomycetes causing esca-associated white rot in grapevine: a worldwide perspective MICHAEL FISCHER Staatliches Weinbauinstitut, Merzhauser Str. 119, D-79100 Freiburg, Germany Summary. An overview is provided of the biodiversity, taxonomic classification and geographic distribution of basidiomycetes isolated from white rot in esca-affected grapevine all over the world. A total of 10 species have been recovered, all belonging to the Hymenochaetales and five of them of uncertain identity. Identified taxa include Fomitiporia mediterranea (Europe, Asia), F. polymorpha (North America), F. australiensis (Australia), and Inocutis jamaicensis (South America). One species, not yet formally described, is Fomitiporella vitis (South America). Uncer- tain taxa occur in Australia and South Africa. Existence of the fungi is widely restricted to vegetative mycelium. Fruit bodies are rare in the field and molecular data are usually necessary for differentiation. Key words: Fomitiporia, Hymenochaetales, molecular systematics. In spite of some progress achieved in recent iv) in several wine-producing countries basidio- years, information is still sparse on the taxonomic mycetes are not considered serious pathogens classification, geographic distribution and life of grapevine, and they have not attracted par- strategies of basidiomycetes on esca-diseased ticular attention from researchers. grapevine. There are several reasons for this: It was at the 2nd workshop of the International i) very often the occurrence of a particular fun- Council on Grapevine Trunk Diseases (ICGTD), gus is limited to the vegetative state, i.e., to held in Lisbon, Portugal in September 2001, that the mycelium living inside the infected plant; white rot was presented as one of the crucial com- ii) if fruit bodies develop, they are often inconspic- ponents of so-called ‘esca proper’, evident in older uous, forming crust-like layers only on the less vines (Graniti et al., 2000; Surico, 2001). In this obvious parts of infected trunks; way esca proper was separated from the dieback iii) frequently, fruit bodies are in poor condition, phenomena observed in younger vines, which are making proper identification time-consuming now called Petri disease. Before this it had already or even impossible; been shown that esca-related wood deterioration, including white rot as well as other symptoms such as brown-red wood and dark stripes, may be caused by a number of fungal organisms, including mem- To correspond with the Author: bers of the mitosporic genera Phaeomoniella and/ Fax: +49 761 40165 70 E-mail: [email protected] or Phaeoacremonium, but that white rot is always S30 Phytopathologia Mediterranea Basidiomycetes and esca linked with the occurrence of basidiomycetes. was also reported from French vineyards (Vinet, Esca and its possible precursor, Petri disease, 1909; Viala, 1926). In their ‘Hyménomycètes de have been reported from wine-growing countries France’, Bourdot and Galzin (1927) mentioned a worldwide (Larignon and Dubos, 1997; Ferreira et resupinate basidiomycete as a strong parasite on al., 1999; Mugnai et al., 1999; Chiarappa, 2000; Vitis in the Midi region of southern France, but Pascoe and Cottral, 2000). In several areas, most- this observation was not repeated in later publica- ly in Europe, white-rot fungi are considered major tions. contributors to esca-related problems. In other re- While studies in esca had intensified by the end gions however, these fungi play only a subordinate of the 1990’s, no significant progress was made in role, although they have been shown to occur vir- the classification and distribution of white rot ba- tually everywhere in northern and southern-hem- sidiomycetes. The taxa most mentioned were still ispheric vineyards (Fischer, 2001). P. igniarius, S. hirsutum, and also Trametes hirsu- Since the data are incomplete, the taxonomic ta (Wulf. : Fr.) Pilát (Mugnai et al., 1996; Larignon classification and phylogenetic relationships of and Dubos, 1997; Chiarappa, 2000). In the years esca-related basidiomycetes are uncertain for many following, P. igniarius was omitted from this list countries, and only a few defined names have so as it was shown to be a European-based species far been introduced (Fischer, 2002; Fischer et al., occurring only on species of Salix (Fischer, 1995, 2005). It is interesting to note that the spectrum of 2002; Fischer and Binder, 1995, 2004). However, fungi associated with precursor diseases of esca another species, Fomitiporia punctata (P. Karst.) seems to be fairly uniform, while for the basidio- Murrill (= P. punctatus (P. Karst.) Pilát) now gained mycetes the situation is more complex. At present, increasing prominence and came to be viewed as it is not known how far white rot fungi in general the main causal agent of white rot in both Italian are an essential part of esca and whether injured and German vineyards (Mugnai et al., 1999). F. wood should also be seen as an important source punctata was the first esca-related basidiomycete of nutrients attracting a wide range of more or less to be thoroughly studied using both molecular and well-adapted fungal organisms. epidemiological data, though in a limited geograph- This paper provides an overview of the biodi- ic area (Cortesi et al., 2000; Fischer, 2000). Shortly versity, taxonomic classification and geographic afterwards, as a result of pairing tests of single- distribution of basidiomycetes isolated from white spore isolates, growth data of the vegetative myc- rot in esca-affected grapevines all over the world. elium and molecular sequence data of the nuclear Several colleagues, by supplying fungal isolates encoded ribosomal ITS region, a new species, F. and sharing valuable information, have contribut- mediterranea M. Fischer (designated Fmed in what ed to this paper. I gratefully acknowledge the sup- follows), was separated from F. punctata and spec- port of Laura Mugnai, Francesca Mela, Walter imens collected from decayed wood of grapevine Gams and Hanns-Heinz Kassemeyer (see chapter that had originally been identified as F. punctata on Europe), Douglas Gubler and his working group were now re-classified as the closely related Fmed (North America), Marta Gatica and Jaime Auger (Fischer, 2002). (South America), Jacky Edwards, Ian Pascoe and During the last few years, grapevine-inhabit- James Cunnington (Australia), and Francois Hal- ing basidiomycetes have attracted increasing at- leen and Paul Fourie (South Africa). tention also outside Europe, and valuable informa- tion on their geographic distribution, life strategy, Basidiomycetes on grapevine - from past as well as the symptoms associated with them have to present been provided by several working groups (Gatica et al., 2000, 2004; Pascoe and Cottral, 2000; Ed- Ravaz (1909) was probably the first researcher wards et al., 2001). At the 2nd ICGTD workshop a to study basidiomycete mycelium and fruit bodies preliminary overview was presented illustrating on diseased vines. He identified the fruit bodies as the genetic diversity of esca-related basidiomycetes belonging to Phellinus igniarius (L.) Quél. (at the worldwide, without however making any state- time: Fomes igniarius (L. ex Fr.) Kickx.). Besides ments about taxonomic classification (Fischer, P. igniarius, Stereum hirsutum (Willd.: Fr.) Pers. 2001). Nevertheless, more detailed studies, al- Vol. 45, Supplement, 2006 S31 M. Fischer though conducted in a limited period of time, clearly and Fischer (2001, 2002) for taxonomic details. The showed that a considerable number of basidiomyc- reference taxa were as follows: etes occur on grapevine. Mensularia radiata (Sow. : Fr.) W.B. Cooke (= Inonotus radiatus (Sow. : Fr.) P. Karst.); P. igni- Why only white rotters on grapevine? arius (see Fischer, 1995; Fischer and Binder, 2004, To our knowledge, no brown rot species have so for geographic distribution and host range); Ino- far been reported from Vitis vinifera (for central notus hispidus (Bull. : Fr.) P. Karst. (while some- Europe, see checklists in Kreisel, 1961; Fischer and times found on grapevine, this species has no path- Kassemeyer, 2003). On the other hand, of the sev- ogenic significance on this host; see Fischer and eral thousand basidiomycetes living on wood, Kassemeyer, 2003); I. cuticularis (Bull. : Fr.) P. brown rot fungi probably constitute less than 10% Karst.; Inocutis rheades (Pers.) Fiasson & Niemelä of the total (Gilbertson, 1981; Gilbertson and Ry- (= I. rheades (Pers.) Pil.); Fomitiporia hesleri M. varden, 1986–1987; Ryvarden and Gilbertson, Fischer (see Fischer and Binder, 2004); F. polymor- 1993–1994) and as a strong tendency they occur pha M. Fischer (see Fischer and Binder, 2004); F. primarily on conifer wood; those on hardwood are robusta (P. Karst.) Fiasson & Niemelä, and F. punc- mostly restricted to members of the Fagaceae and tata. Betulaceae (Nobles, 1958, 1971; Gilbertson, 1981). In all, 35 isolates were included in the survey (Fig. 1), 22 of which came from grapevine (in ital- Geographic distribution and taxonomic ics in Fig. 1). Two more isolates, while collected classification of basidiomycete species on from non-Vitis hosts, have been shown previously grapevine to belong to species associated with white rot of grapevine. These isolates are 99-105 (Fmed; Esca-related isolates included in the present Fischer, 2002) and VPRI 22409b (F. australiensis; survey originated from Europe, North and South Fischer et al., 2005).
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