The xylariaceous way of life

A.J.S. Whalley

This paper outlines current positions on systematic arrangements of the Xylariaceae, discusses the value of the different characters used in attaining these arrangements, and reviews the activities of members of the family in nature. The taxonomic significance and potential ecological role of the many secondary metabolites produced by xylariaceous taxa is evaluated. The importance of the Xylariaceae in wood decomposition and as agents of disease is discussed. Aspects of their distribution in relation to host, habitat and climate are presented.

Mycal. Res. 100 (8): 897-922 (I996) 897 Printed in Great Britain Presidential address The xylariaceous way of life A. J. S. WHALLEY School of Biomolecular Sciences, Liverpool John Moores University, Byrom Street, Liverpool L3 3AF, UK This paper outlines current positions on systematic arrangements of the Xylariaceae, discusses' the value of the different characters used in attaining these arrangements, and reviews the activities of members of the family in nature. The taxonomic significance and potential ecological role of the many secondary metabolites produced by xylariaceous taxa is evaluated. The importance of the Xylariaceae in wood decomposition and as agents of disease is discussed. Aspects of their distribution in relation to host, habitat and climate are presented. The study of fungi provides as great, and as interesting, a challenge as any to be found in the living world. At a time when biodiversity is in vogue the fungi in their broadest sense excel. According to popular consensus there are more than l' 5 million species (Hawksworth, 1991; 1993) and they exhibit an impressive range of diversity of form, biochemical and physiological activities, and adaptation to habitat. They rank as one of the world's most important living resources. In a mycological lifetime it is unrealistic to expect an individual to understand in any depth this diversity and I have unashamedly concentrated my efforts on a single ascomycete family; the Xylariaceae. This is not an admission of lack of interest in other groups but more a tribute to what a single ascomycete family can offer. The Xylariaceae is a large and relatively well known family which has representatives in most countries of the world (Dennis, 1956, 1957, 1961, 1963, 1970; Miller, 1961; Martin, 1967a, b; 1968a, b; 1969a-c, 1970; Rogers, 1979a; Rogers, Callan & Samuels, 1987; Rogers et aI., 1988; Granmo et al., 1989; San Martin & Rogers, 1989; Van der Gucht, 1992; Van der Gucht & Van der Veken, 1992; Gonzalez & Rogers, 1993; Whalley, 1993, 1996; Whalley et al., 1995). Eriksson & Hawksworth (1993) recognised 35 genera plus three others which they considered might belong there. Most members of the family are wood inhabitants but there are also representatives found in litter, soiL dung and associated with insects (Rogers, 1979a; Whalley, 1985). In spite of their widespread distribution and occurrence in a wide range of environmental situations Rogers, in his Presidential Address to the Mycological Society of America, drew special attention to our lack of knowledge about the activities of the Xylariaceae in nature (Rogers, 1979a). We have made considerable progress since then - hence the title of my address. SYSTEMA TIC BASE Genera of the Xylariaceae The acceptable number of known genera within the Xylariaceae, in the absence of a clear circumscription of the family, is still open to debate. In their current outline of the ascomycetes Eriksson & Hawksworth (1993) recognized 35 genera and indicated a further 3 which might prove to belong there. In an index to the genera of the family Lress0e included 37 genera but admitted that a few of these are uncertain A.J.S. Whalley President, British Mycological Society, 1994 The xylariaceous way of life 898 Table 1. Genera of Xylariaceae Eriksson & Hawksworth (1993) Lil'ssoe (I994) Whalley (this review) Anthostomelia Sacco Ascotricha Berk. ?Ascotrichelia Validos. & Guarro ?Astrocystis Berk. & Broome Biscogniauxia Kuntze Calceomyces Udagawa & S. Ueda Camillea Fr. Anfhosfomelia Anfhosfomelia ?Ascofricha ?Ascotrichelia Asfrocysfis Biscogniauxia Calceomyces Camillea ?Chaenocarpus ?Coliodiscula Creosphaeria Daldinia Engleromyces Enfonaema ?EuepixyIon Daldinia Ces. & De Not. Engleromyces Henn. Entonaema A. Moller Fassia Dennis Helicogermslita Lodha & D. Hawksw. Hypocopra (Fr.) I. Kickx f. Hypoxylon Bull. Induratia Samuels, E. Miill. & Petrini Kretschmaria Fr. Leprieuria Lil'ssoe, I. D. Rogers & Whalley Lopadostoma (Nitschke) Traverso ?Paucithecium Lloyd Penzigia SacCo Phaeosporis Clem. Phylacia Lev. Podosordaria Ellis & Holw. Poroconiochaela Udagawa & Furuya Poronia Willd. Pulveria Malloch & Rogerson Rhopalostroma D. Hawksw. Rosellinia De Not. Sarcoxylon Cooke Sfilbohypoxylon Henn. Sfromafoneurospora S. C. long & E. E. Davis Thamnomyces Ehrenb. Theissenia Maubl. Thuemenelia Penz. & SacCo Usfulina Tul. & C. Tul. Versiomyces Whalley & Watling Wawelia Namysl. Xylaria Hill ex Schrank Asfrocystis Biscogniauxia Calceomyces Camillea Chaenocarpus Fr. Collodiscula I. Hino & Katum. Creosphaeria Theiss. Engleromyces Enfonaema Euepixylon Fiiisting Helicogermslifa Holttumia Lloyd Hypocopra Hypoxylon Induratia Krelschmaria Leprieuria Lopadosfoma ?Myconeesia Kirschst. Nemania Gray emend. Pouzar Obolarina Pouzar Phaeosporis Phylacia Podosordaria Poronia (as Pyrenomyxa Morgan) Rhopalosfroma Rosellinia Sarcoxylon ?Seynesia Sacco Helicogermslifa ?Holttumia Hypocopra Hypoxylon Indurafia Krelschmaria Leprieuria Lopadosfoma Nemania Obolarina ?Penzigia Phaeosporis Phylacia Podosordaria Poronia Pulveria Rhopalosfroma Rosellinia Sarcoxylon Sfromafoneurospora Thamnomyces Theissenia Thuemenelia ?Sfilbohypoxylon Sfromafoneurospora Thamnomyces Theissenia Thuemenella ?Wawelia Xylaria Versiomyces Wawelia Xylaria (Lcess0e, 1994). However, comparison of these two recent listings of xylariaceous genera reveals a number of important differences (Table 1). Eriksson & Hawksworth surprisingly did not recognise Nemania and Obolarina although these are widely accepted (e.g. Candoussau & Rogers, 1990; Whalley & Edwards, 1995) whilst Lcess0e (1994) assigned Daldinia and Versiomyces to Hypoxylon. There appear to be no good reasons to include Daldinia in Hypoxylon although they are clearly closely related. Indeed Daldinia is structurally and distinctively different when examined by light and scanning electron microscopy (Gaskell, 1995) and in many aspects is more closely related to Entonaema than to Hypoxylon sensu stricto (Rogers, 1982; Whalley & Edwards, 1987, 1995). Versiomyces also appears sufficiently distinctive to warrant separate generic status (Rogers, pers. comm.). Taking these differing views into account and allowing for the erection of new genera in the future an upper figure of 40 or more genera seems realistic (Table 1). Systematic foundation A sound systematic foundation is an essential prerequisite for specialist investigations into any aspect of the life of members of the family. The Xylariaceae has been the focus of considerable attention concerning taxonomic relationships and as a result a genuine understanding of the associations between species and genera is evolving (Rogers, 1993, 1994; Whalley & Edwards, 1995). It is well known that many A. J. S. Whalley 899 Figs 1-6. Pigmentation, diversity and pathology in Xylariaceae. Fig. 1. Hypoxylon haemaloslroma Mont. an example of a highly pigmented species. Fig. 2. Biscogniauxia nolhofagi. Fig. 3. Camillea leprieurii. Fig. 4. Chromatogram of secondary metabolites produced by selected Xylariaceae. Lane 1 = Hypoxylon fragiforme, 2 = H. howieanum, 3 = H. mammalum, 4 = H. truncatum, 5 = H. atropuncialum, 6 = H. multiforme, 7 = Biscognianlia marginata. Fig. 5. Beech trees in Sicily infected by Biscogniauxia nummularia. Fig. 6. Apples trees infected by Rosellinia necatrix. The xylariaceous way of life 900 Figs 7-12. Fig. 7. Camillea selangorensis, stroma. (Bar = 5 mm). Fig. 8. Hypoxylon hians, stromata. (Bar = 1 em ). Fig. 9. Rhopalostroma kanyae Whalley & Thienhirun, stroma. (Bar = 1 mm). Fig. 10. Rosellinia bunodes, stroma. (Bar = 1 mm). Fig. 11. Hypoxylon megannulatum Talig. & Whalley, stromata. (Bar = 1 mm). Fig. 12. Hypoxylon stygium Lev., stromata. (Bar = 2 mm). representatives of the family exhibit an exceptional range of morphological diversity (e.g. Dennis, 1956, 1957; Martin, 1967 a; Rogers, 1979a, 1993; Petrini & Muller, 1986; Whalley, 1993; Figs 1-3, 7-14) and that characteristics other than traditional ones are necessary to resolve many of the taxonomic difficulties, or the confusion arising, as a result of this diversity (Martin, 1967a; Rogers, 1979a, 1993; Whalley & Edwards, 1995). Morphological variation in Xylaria Hill ex A. J. S. Whalley 901 1981, 1985), stromal pigmentation (Greenhalgh & Whalley, 1970; Whalley & Whalley, 1977) the nature of the apical Stromata leathery or woody. never Section Hypoxylon apparatus and ascus tip (Griffiths, 1973; Beckett & Crawford, carbonaceous, bright coloured, usually 1973; Stiers, 1977; Rogers, 1979a; Pouzar, 1985 a, b), and some shade of red, purple or brown, globose, subglobose, pulvinate, or widely cytological data (Rogers, 1964 ; 1965, 1967 a, b, 1968 a-c, effused; ostioles umbilicate. 1969, 1970, 1972, 1973, 1975a, b; Rogers & Stiers, 1974) Stromata globose, subglobose, pulvinate, Section Papillata have all provided varying degrees of discriminating inforor widely effused; ostioles papillate. mation. The type of apical apparatus which usually presents Stromata coloured at least when young subsection Papillata a typical blue amyloid reaction with Melzer's reagent has becoming brown to black at maturity. Stromata white or light grey when young subsection Primo-cinerea proved to be especially useful at the generic level (Griffiths, becoming black and very carbonaceous at 1973; Nannfeldt, 1976; Rogers, 1979a; LiESS0e, Rogers & maturity. Whalley, 1989; San Martin & Rogers, 1989; Gonzalez & Ostioles papillate and surrounded by an Section Annulata Rogers, 1993; Figs 19-21). The length and position of the annular disk; stromata usually coloured. germination slit on the ascospore is also important, particularly Stromata restricted or indefinitely effused, Section Applanata carbonaceous and applanate; ostioles in Xylaria and its allies (Beckett, 1979a, b; Rogers, 1979b, umbilicate or papillate. 1983, 1984a, b; Rogers & Callan, 1986a, b; Ju & Rogers, 1990; Figs 14 & 15). Application of SEM to study surface ornamentation of ascospores has proved to be extremely valuable (Figs 27-31). At the generic level Camillea is Schrank is especially challenging and in certain species characterized by its light-coloured ascospores with no visible complexes knowledge of the anamorph is required in order to germ slits by light microscopy but by SEM they are separate the closely related taxa (Martin, 1970; Rogers, 1985, distinctively ornamented with warts, spines, pits, reticulations 1993; San Martin & Rogers, 1989; Callan & Rogers, 1990). or are longitudinally ribbed (Rogers, 1977 b; LiESS0e et al., The danger of over reliance on gross morphological features 1989; Gonzalez & Rogers, 1993; Whalley, 1995; Whalley, is well illustrated by reference to the dimorphic Camillea Whalley & Jones, 1996). In Stromatoneurospora the ascospores leprieurii Mont. (Fig. 3). In its expanded or applanate form it are striate (Jong & Davis, 1973). At the species level certain was placed in Hypoxylon as H. melanaspis Mont. but in its individuals possess characteristically ornamented ascospores. elongated state it was accepted in Camillea (Dennis, 1957; In Hypoxylon sensu stricto a number of taxa are characterized Miller, 1961). Examination of ascospores, anamorphic form, by ascospores ornamented with faint transverse striations and chemical profiles proved that the two states are merely oriented perpendicular to the long axis of the spore morphological forms of the same taxon (LiESS0e, Rogers & (Rogers & Candoussau, 1982; Van der Gucht & Van der Veken, 1992; Fig. 28). Similar spore ornamentation is found Whalley, 1989). Traditionally the form of the stroma, pigmentation, shape in a number of Daldinia species (Van der Gucht, 1993). In and dimensions of the ascospores have featured strongly in H. weldenii J. D. Rogers and Nemania chestersii (J. D. Rogers & the earlier accounts e.g Hypoxylon (Miller, 1961), Xylaria Whalley) Pouzar the ascospores have longitudinal ridges (Dennis, 1956, 1957) and other xylariaceous genera (Martin, running parallel to the long axis which give them a striate 1967a; Rogers, 1979a; LiESS0e & Spooner, 1994) In many appearance (Rogers, 1977 a; Rogers & Whalley, 1978; Fig. cases the' classical' approach although convenient has resulted 29). There is insufficient data on peridial anatomy to draw in the formation of unnatural groupings where species or conclusions and at present the cytological data obtained does genera appear to be taxonomically closely related when in not appear to be taxonomically useful. reality they are distinct and distant. In his monograph of It was, however, the realization that the anamorphic state Hypoxylon Miller (1961) divided the genus into four sections provides a previously unused source of taxonomic characters on the basis of stromal form, texture and colour, and nature of useful in the separation of closely related species, species the ostiole (Figs 1-3, 7-8, Il-12, 15-16, 18; Table 2). groups or often genera (Chesters & Greenhalgh, 1964; Subsequent studies employing other characteristics show that Martin, 1967a, 1968a, b; 1969a-c, 1970; Greenhalgh & these divisions are artificial and fail to recognise taxonomic Chesters, 1968; Jong & Rogers, 1972) which stimulated affinity or distance between the taxa involved (Pouzar, 1979, considerable impetus for the development of a more 1985 a, b; Rogers, 1979a; Whalley & Edwards, 1987, 1995; satisfactory systematic arrangement in the Xylariaceae (Rogers, Granmo et aI, 1989; Whalley & Greenhalgh, 1973). Thus 1979a). It can be seen that specific anamorphic forms are Miller's subsection Primocinerea of the section Papillata linked with certain genera (Table 3). Thus Xylocladium appears contains elements now distributed in a range of genera e.g. exclusive to Camillea (Rogers, 1975 c, 1979 a; LiESS0e, Rogers Nemania (Pouzar, 1985a, b), Rosellinia (Petrini, 1992) and & Whalley, 1989; Gonzalez & Rogers, 1993) whilst Euepixylon (LiESS0e & Spooner, 1994). Section Applanata Geniculosporium links with Nemania (Whalley & Rogers, 1980; (Miller, 1961) is currently redistributed between Camillea and Pouzar, 1985 a, b; Petrini & Miiller, 1986) and Lindquistia with Biscogniauxia (Pouzar, 1986; LiESS0e, Rogers & Whalley, the dung inhabiting Podosordarill and Porania (Jong & Rogers, 1989; Whalley, LiESS0e & Kile, 1990a, b; Van der Gucht, 1969; Rogers & Russell, 1973; Furuya & Udagawa, 1977; 1992; Gonzalez & Rogers, 1993). Undoubtedly other genera Rogers, 1985; Rogers & LiESS0e, 1992). Nodulisporium is e.g. Xylaria, Rosellinia, will require considerable rearrangement characteristic for Hypoxylon sensu stricto, Daldinia, and many as more data becomes available. Peridial anatomy (Jensen species of Biscogniauxia (Greenhalgh & Chesters, 1968; Jong & Table 2. Classification of Hypoxylon after Miller (1961) The xylariaceous way of life Figs 13-18. Fig. 13. Poronia punetata, stromata. (Bar = 1 em ). Fig. 14. Rosellinia aquila, stromata. (Bar = 1 mm). Fig. 15. Hyporylon stygium, stroma with papillate ostiole surrounded by ostiolar disc. (Bar = 1 mm). Fig. 16. Hypoxylon mammatum, papillate ostioles. (Bar = 2 mm). Fig. 17. Rhopalostroma kanyae, stromal head with sunken ostioles (arrowed). (Bar = 1 mm). Fig. 18. Camillea fusiformis M. A. Whalley, ostioles (arrowed). (Bar = 100 ｾｭＩＮ 902 A. J. S. Whalley 19 903 20 21 22 Figs 19-26. Fig. 19. Kretzschmaria clavus, ascus tip with massive amyloid apical apparatus. (Bar = 10 ｾｭＩＮ Fig. 20. Camillea fusiformis, ascus tip with rhomboid amyloid apical apparatus. (Bar = 10 ｾｭＩＮ Fig. 21. Hypoxylon rubiginosum, ascus tip with reduced discoid Fig. 22. Biscogniauxia anceps (Sacc.) j. D. Rogers, Y. M. ju & Cand., biCelled amyloid apical apparatus (arrowed). (Bar = 10 ｾｭＩＮ Fig. 23. Rhopalostroma kanyae, ascospores with germination slit. (Bar = 2 ｾｭＩ Fig. 24. ascospore with germination slit. (Bar = 10 ＮＩｭｾ Hypoxylon investiens (Schwein.) Curt., ascospore with sigmoid germination slit. (Bar = 3 ｾｭＩＮ Fig. 25. Rosellinia evansii Lress0e & Spooner, ascospore with helical germination slit. (Bar = 5 ｾｭＩＮ Fig. 26. Rosellinia thelena (Fr.) Rabenh., ascospores with germination slits gaping open prior to germination. (Bar = 2 ｾｭＩＮ Rogers, 1972; Callan & Rogers, 1986; Petrini & Muller, 1986; Figs 33-34.). Furthermore, the routine cultivation of a wide range of taxa, representing a diverse range of xylariaceous genera, opened the way for chemical investigations of the secondary metabolites they produced (Whalley & Edwards, 1987, 1995; Fig. 4). Compounds isolated and characterized include dihydroisocoumarins and related compounds (Anderson, The xylariaceous way of life 904 Fig. 28. Hypoxylon rubiginosum, ascospore ornamentation. Fig. 27-32. Fig. 27. Camillea tinetor, ascospore ornamentation. (Bar = 1 ｾｭＩＮ (Bar = 1 ｾＩＮ Fig. 29. Nemania chestersii, ascospore ornamentation. (Bar = 1 ｾｭＩＮ Fig. 30. Camillea selangorensis, part of ascospore Fig. 31. Camillea fusiformis, ascospores with ribbed ornamentation. (Bar = 10 ｾＩＮ Fig. showing warted ornamentation. (Bar = 1 ｾＩＮ 32. Rosellinia bunodes, ascospores with long tapering ends. (Bar = 10 ＮＩｭｾ Edwards & Whalley, 1983), butyrolactones (Edwards & Whalley, 1979; Anderson, Edwards & Whalley, 1982), sesquiterpene alcohols (Anderson et a/., 1984a; 1984b; Poyser et al., 1986; Edwards, Poyser & Whalley, 1988; Edwards et ai., 1989), cytochalasins (Edwards, Maitland & Whalley, 1989; Dagne et ai., 1995) and succinic acid derivatives (Anderson, Edwards & Whalley, 1985). The distribution of these and other compounds, such as cubensic acid (Edwards, Maitland & Whalley, 1991), globoscinic acid and globoscin (Adeboya et ai., 1995a; berteric, carneronic and rnalaysic acids (Adeboya et ai., 1995 b), are proving to be valuable taxonomic pointers at both generic and species level (Whalley & Edwards, 1995). A. J. S. Whalley 905 Table 3. Anamorph-teleomorph connections Teleomorph Anamorph Anlhoslomella = Nodulisporium type 2a), Nodulisporium Preuss and Virgariella S. Hughes (Francis, Minter & Caine, 1980) Dicyma Boulanger (Hawksworth, 1971) 7Humicola-like (Valldosera & GuaTTo, 1988) Acanthodochium Samuels, j. D. Rogers & Nagas. (Samuels, Rogers & Nagasawa, 1987; ju & Rogers, 1990) Geniculosporium (Eckblad & Granmo, 1978; Whalley & Edwards, 1985), Nodulisporium (Greenhalgh & Chesters, 1968; Callan & Rogers, 1986; Gonzalez & Rogers, 1993), Periconiella (Petrini & Muller, 1986) Nodulisporium (Udagawa & Ueda, 1988) Xylocladium Syd. (Crane & Dumont, 1975; L""ss0e, Rogers. & Whalley. 1989; Gonzalez & Rogers. 1993). Unknown Acanthodochium (Samuels, Rogers & Nagasawa, 1987) Nodulisporium (Chesters & Greenhalgh, 1964; Petrini & Muller, 1986) Unknown Nodulisporium (Rogers, 1982). Geniculosporium (Whalley. 1976) Unknown Unknown Unknown Nodulisporium. Virgariella, Hadrotrichum Fuckel, Rhinocladiella Nannf. (Martin, 1967 a; Greenhalgh & Chesters, 1968; jong & Rogers, 1972; Petrini & Muller, 1986) Nodulisporium (Samuels. Muller & Petrini, 1987) Hadrotrichum (Petrini & Muller. 1986) Geniculosporium (Samuels & Muller, 1980) scolecosporous anamorph, Liberiella-like Gu, Gonzalez & Rogers, 1993) Geniculosporium (Chesters & Greenhalgh. 1964; Petrini & Muller, 1986) Rhinocladiella-like (Candoussau & Rogers, 1990) Unknown Sporothrix Hektoen & C. F. Perkins Gong & Davis, 1974) Geniculosporium (Rodrigues & Samuels, 1989) Lindquistia Subram. & Chandrash. (Subramanian & Chandrashekara, 1977; Rogers & L""ss0e, 1992) Lindquistia (Subramanian & Chandrashekara, 1977; Stiers, Rogers & Russell, 1973) Unknown Nodulisporium (Hawksworth & Whalley, 1985) Geniculosporium, Dematophora R. Hartig, Nodulisporium (Petrini, 1992) Unknown Unknown Unknown Nodulisporium (Samuels & Muller, 1980) Unknown Nodulisporium (Samuels, 1989; Samuels & Rossman, 1992) Unknown Anamorph described by Minter & Webster (1983) as being geniculate but not assigned to a form genus Typically produced on developing stromata but no form genus yet assigned. Xylocoremium flabelliforme (Schwein.: Fr.) ). D. Rogers is associated with X. cubensis (Rogers, 1984 b; 1985) 7Ascolricha 7Ascolrichella Astrocyslis Biscogniauxia Calceomyces Camillea 7Chaenocarpus Collodiscula Daldinia Engleromyces Entonaema Euepixylon Helicogermslita Holttumia Hypocopra Hypoxylon Induratia Krelzschmaria Leprieuria Lopadostoma Nemania Obolarina 7Penzigia Phaeosporis Phylacia Podosordaria Poronia Pulveria Rhopalostroma Rosellinia Sarcoxylon Slilbohypoxylon Stromaloneurospora Thamnomyces Theissenia Theumenella Versiomyces Wawelia Xylaria Geniculosporium Chesters & Greenh. (Martin 1969 a, ACTIVITIES OF THE XYLARIACEAE These can be broadly grouped under decomposition of natural substrata, their role as phytopathogens, and as endophytes. The boundaries between these activities are often ill defined and subject to change depending on interactions especially between the host, or substratum, and the environment. Thus taxa which are present in living hosts as endophytes can become pathogenic if the host becomes stressed, and pathogenic species may linger after killing the host by leading a saprotrophic existence (Chesters, 1950; Whalley, 1985; Boddy, 1994). Role in wood decomposition Although members of the Xylariaceae can colonize a wide range of substrata the vast majority are wood inhabitants (Rogers, 1979a). Earlier studies on their occurrence and host specificity were through necessity related to the presence of the teleomorph in the field and led to a general belief that the Xylariaceae are only involved in the decomposition of woody structures (Merrill. French & Wood, 1964; Rogers, 1979a; Whalley, 1985). Furthermore their appearance in the teleomorph stage is often closely linked to the stage of wood decomposition (Whalley, 1985). Thus Hypoxylon fragiforme and Biscogniauxia nummularia develop on dying or recently dead branches and stems where the wood is still in a relatively undecayed condition (Chesters, 1950; Chapela & Boddy, 1988a-c). In contrast Euepixylon udum and Nemania conj1uens are always associated with well decomposed, decorticated wood, which is often water soaked (Whalley, 1985). The ability of members of the Xylariaceae to participate in wood decomposition is without question (Rogers, 1979a; Whalley, 1985; Rayner & Boddy, 1986, 1988). Selected species have The xylariaceous way of life 906 Figs 33, 34. Nodulisporium anamorph of Biscogniauxia nummularia. (Fig. 33. Bar = 1 ｾｭ［ been shown to degrade lignin (Sutherland & Crawford; 1981) and others exhibit impressive produdion of cellulolytic enzymes (Wei et al., 1992). In early studies on wood decay in the laboratory involving various Xylariaceae, Blaisdell (1939) found that D. concentrica, a species of Hypoxylon, and a Xylaria caused significant weight losses in three different hardwoods. Cartwright & Findlay (1946) then demonstrated that X. polymorpha (Pers.: Fr.) Grev. when inoculated on to beech wood blocks caused a 14 % weight loss over a 4 month period. Weight losses from 10 to 26 % in aspen and red oak woods in 3 months were recorded for D. concentrica, B. atropunetata B. mediterranea, H. mammalum (as H. pruinatum (Klotzsch) Cooke) and C. punetulata (Merrill et aI., 1964) whilst Xylaria digitata (L.: Fr.) Grev. and H. rubiginosum caused a 16 % weight loss in aspen wood after 3 months (Rajagopalan, 1966). In a study of the chemical and microscopic features of wood decay caused by a seledion of Xylariaceae D. concenlrica was found to be particularly adive causing a weight loss in birch of 62'9% after 2 months (Nilsson et al., 1989). The Xylariaceae tested were found to degrade preferentially the syringylpropane units of lignin and Nilson et al. (1989) suggested this might explain their general inability to degrade pine. Interestingly few Xylariaceae colonize coniferous wood (Miller, 1961; Rogers, 1979a; Whalley, 1985). In a study of decay of oak and beech wood blocks by xylariaceous and diatrypaceous fungi Abe (1989) found that they all demonstrated greater tolerance to low water potentials than the three Basidiomycotina species included in the investigation and that they caused the greatest weight losses in those wood blocks maintained at a low moisture content. Abe also found that the xylariaceous and diatrypaceous isolates, with the exception of H. howeianum Peck, removed 27-37% of the total oak wood lignin achieving a 35-51 % weight loss after six months: he concluded that they have great abilities for decaying wood and adapt well to dry environments (Abe, 1989). The Xylariaceae have, however, received considerable attention recently in their role as latent decay fungi which Fig. 34. Bar = 10 ｾｭＩＮ cause small pockets of decay or decay columns, sometimes extensive, depending on the host and the fungal species involved (e.g. Rayner & Boddy, 1986, 1988; Boddy, 1994). Investigations of attached branches of ash (Fraxinus excelsior L.) and beech (Fagus sylvatica L.) have shown that natural colonization of sapwood occurs in the absence of major wounds (Boddy, Gibbon & Grundy, 1985; Boddy, Bardsley & Gibbon, 1987; Chapela & Boddy, 1988 a, b). Although differences were observed between patterns of early colonization of beech and ash branches these might be attributed to differences in branching patterns, location of dying lateral branches and in the speed of main branch death (Chapela & Boddy, 1988a). In ash, extensive individuals of various Basidiomycotina and Ascomycotina, including D. concentrica and H. rubiginosum, formed very rapidly in sedors of branches, often in less than a single growing season (Boddy et al., 1985, 1987). In beech, early colonizing fungal communities were shown to have a distind spatial organization where decay often started in distal regions with stain-associated fungi which had developed from latent propagules in the wood and bark (Chapela &...Boddy, 1988a). Hypoxylon fragiforme and B. nummularia were frequently found to be early colonists (Chapela & Boddy, 1988a). Although there were differences in the pattern of colonization observed, in branches of beech and ash, the water content of the sapwood in both trees was strongly implicated in the eventual outcome of colonization (Chapela & Boddy, 1988 b). In beech, H. fragiforme and B. nummularia were found to develop from pockets within the wood where each pocket contained a genetically different individual. Chapela & Boddy (1988b) concluded that for these fungi infedion is a common event and that they do not require large wounds in order to colonize the tree. They were therefore viewed as being present as inconspicuous fungal propagules within the wood which subsequently developed as 'latent invaders' once the high water content of the wood was reduced (Chapela & Boddy, 1988 b). The rapid appearance of teleomorphs of H. fragiforme and B. nummularia in freshly felled beech branches (Chapela & Boddy, 1988c) is consistent with these findings. It A. J. S. Whalley is now apparent that a significant number of xylariaceous taxa fit this pattern and furthermore, they can, if suitable conditions prevaiL become invasive and cause canker disease in their host (see below). Xylariaceae as phytopathogens Although the Xylariaceae are not usually considered to be important as plant pathogens a growing number of species are now acknowledged to cause considerable economic loss in natural ecosystems or under agricultural conditions (Rogers, 1979a; Whalley, 1985). Representatives of the family cause canker in trees, root rots in a wide range of plant species, and needle blights in conifers (Table 4). Canker diseases. Hypoxylon mammatum is the best known and economically the most important cause of a canker disease and has long been recognized as the cause of serious stem canker in aspen (Povah, 1924; Manion & Griffin, 1986; Fig. 16). It infects mainly Populus tremuloides Michx. in North America and P. tremula L. in Europe (French, Hodges & Froyd, 1969; Pinon, 1979; Manion & Griffin, 1986). It was calculated that in 1971 in Minnesota, eastern Upper Michigan, and south western Wisconsin Hypoxylon canker was responsible for an annual loss of $4'4 million at the time of harvest (Marty, 1972). An estimated 1-2% of aspen grown in the u.s.A. is killed by H. mammatum each year (Anderson, 1964). The fungus also occurs on Acer, Alnus, Betula, Carpinus, Fagus, Picea, Pyrus, Salix, Sorbus, and Ulnus but only appears to be an important pathogen of Populus (Manion & Griffin, 1986). In North America the disease is prevalent throughout the northeast, the Great Lakes region, and the northwestern prairies. It is, however, absent from the northern Rocky Mountains and Alaska in spite of abundant aspen and the presence of the fungus (Juzwik, Nishijima & Hinds, 1978). In Europe it is believed that the disease has been present in the Alps for over 30 years (Pinon, 1979). Ascospores are assumed to be the source of infection but ascospore inoculations have proved unreliable and considerable research has been undertaken on the conditions governing germination in attempts to determine the exact conditions for infection (Rogers & Berbee, 1964; Manion & Griffin, 1986). An impressive amount of data has been accumulated on the site of infection and the growing conditions of the trees which then become infected. Thus good correlation has been observed between natural cankers and one or two year old lateral branches (Manion, 1975) and with branch galls caused by the insect Saperda inornata Say (Anderson, Ostry & Anderson, 1979). In spite of conflicting information there is evidence to relate water stress in trees with higher incidence of Hypoxylon canker (Day & Strong, 1959; Bier & Rowat, 1962; Bagga & Smalley, 1969; Bruck & Manion, 1980). It has also been shown that under water stress conditions the proline content of aspen increased dramatically and that radial growth rates of H. mammatum isolates were stimulated much more by proline than other amino acids (Griffin, Quinn & McMillen, 1986). It was later shown that there is clonal variation in the amino acid contents of aspen induced by diurnal drought stress and this may influence susceptibility to Hypoxylon 907 canker (Griffin et al., 1991). If the conclusion of Chapela (1989), that H mammatum is an endophyte of aspen, proves to be the norm then the importance of water stress becomes highly significant Hypoxylon mammatum is the most damaging of the canker causing Xylariaceae and it is also noteworthy for the production of phytotoxins. Hubbes (1964) first suggested that a toxin or toxins might be involved in the pathological process. Initial attempts to isolate and purify the toxic principle, known as mammatoxin, revealed the presence of several compounds which were active in bioassays (Schipper, 1978; Stermer, Scheffer & Hart, 1984). Recent investigation of culture filtrates of H. mammatum has revealed the occurrence of two families of toxins. One has been identified as a diterpene (hymatotoxin) with a structure similar to momilactones and the second as tetralones (Bodo et al., 1987; Pinon & Manion, 1991). The momilactones were originally isolated from rice (Oryza sativa cv. Koshihikan) husk and inhibit the growth of rice roots and lettuce seeds (Kato et al., 1973, 1977). Tetralones are similar to juglone, a compound toxic to many plants as well as bacteria, fungi, fish, and mice (Soderquist, 1973). Hypoxylon mammatum has also been shown to produce cytochalasin D (Whalley & Edwards, 1995) but this does not cause the same phytotoxic response induced by culture filtrates or pure hymatoxins (Pinon & Manion, 1991) although the cytochalasins are well known for their tissue toxicity in plants (Sawai, Okuno & Ito, 1982; Betina, 1989). There is an intricate host-parasite relationship between H. mammatum and its aspen host with pyrocatechol and other bark substances proving inhibitory to growth (Hubbes, 1962a, b, 1964, 1966) and with the post-infection production of phytoalexins which inhibit ascospore germination but not mycelial growth (Flores & Hubbes, 1979.) Hypoxylon mammatum canker in aspen has attracted the greatest attention but a number of species of Camillea and Biscogniauxia cause significant canker diseases and die back (Table 4) if only in hosts which are stressed through injury, drought or fire damage (Rogers, 1979a; Whalley, 1985). Cork oak (Quercus suber L.) has been declining steadily in Europe for many years and Biscogniauxia mediterranea, the causative agent of coal canker in oak, has been identified as an important factor for this decline (Macara, 1974, 1975). In a 1974 survey of cork oak stands in Portugal 41'5% of trees were recorded as diseased, with B. mediterranea being the principal pathogen (Macara, 1975). Surveys over the past five years indicate that Biscogniauxia is associated with around 30 % of the dying trees (Santos, pers. comm.). In North Africa the disease may reach epidemic proportions (Barbosa, 1958). It is also evident that oak canker is both more prevalent and serious in trees suffering from water stress and there have been increases in the disease following hot, dry spells (Macara, 1974, 1975; Santos, pers. comm.). Biscogniauxia mediterranea is, however, not the only reason for the decline of the cork oak since in Catalonia (NE Spain) Botryosphaeria stevensii Shoemaker hasbeen found to be the main fungal pathogen involved (Luque & GirbaL 1989). Spooner (1986) recorded B. mediterranea from Britain growing on Castanea sativa Mill. He noted that the stromata The xylariaceous way of life 908 Table 4. Examples of diseases caused by members of the Xylariaceae Species Principal hosts Canker diseases Biscogniauxia alropunclala (Schwein.) Pouzar B. medilerranea (De Not.) Kuntze B. nolhofagi Whalley, LreSS0e & Kile B. nummularia (Bull.: Fr.) Kuntze CamiIIea punclulala (Berk. & Rav.) Lress0e, j. D. Rogers & Whalley C. linclor (Berk.) Lress0e, j. D. Rogers & Whalley Hypoxylon mammalum (Wahlenb.) j. H. Mill. H. rubiginosum (Pers.: Fr.) Fr. Xylaria arbuscula Sacco Quercus Quercus Nothofagus. cunninghamii Fagus Quercus Acer, Plalanus Populus CameIIea Macadamia Root rot diseases Krelzschmaria clavus (Fr.) Sacco K. deusla (Hoffm.: Fr.) P. M. D. Martin RoseIIinia bunodes (Berk. & Broome) Sacco R. desmazieresii (Berk. & Broome) Sacco R. necalrix Prill. R. pepo Pat. Xylaria mali Fromme Macadamia Hevea Coffea Salix many Coffea Malus Needle blights R. herpolrichioides Hepting & R. W. Davidson R. minor (Hahn.) S. M. Francis were widely effused and that the bark exhibited extensive peeling and shedding, It was also reported that 'there was no evidence that the fallen Castanea on which the British collection was made had been subjected to fire damage, although the tree grew on well-drained, sandy soil on a hillside and may well have suffered from drought stress during previous dry summers' (Spooner, 1986). A good correlation was also obtained between water stress and ability of B. mediterranea to cause canker in Quercus cerris L. where following inoculation experiments only those seedlings maintained under water stress conditions became diseased (Vannini & Scarascia Mugnozza, 1991). In 1990 the Forestry Commission of Messina (Sicily) reported a serious decline of beech (Fagus sylvatica L.) in the lowest region of a beech wood on the Nebrodi mountain range. Subsequent field surveys and laboratory investigations identified B. nummularia as the associated fungus (Granata & Whalley, 1994; Figs 5, 33-34). They concluded that the prolonged drought and high temperatures experienced in Sicily over the previous ten years had caused conditions of water stress in the beech trees and made them susceptible to the disease (Granata & Whalley, 1994). The greatest damage was observed on the southern slopes of the Nebrodi mountains where exposure greatly influences the amount of water in the soil. No diseased trees were observed in areas where the soil moisture was high (Granata & Whalley, 1994). The relationship between water stress in beech and strip canker development caused by B. nummularia and similar fungi has also been emphasized by Lonsdale (1983), who queried whether fungi such as B. nummularia and Eutypa spinosa (Pers.) Tul. & C. Tul. are directly pathogenic or just colonize tissue which had become debilitated by another factor. Hendry, Boddy & Lonsdale (1993) observed interactions between species of Xylariaceae and Diatrypaceae isolated from beech with beech callus material. They found that when B. nummularia and E. spinosa were grown in dual Tsuga Conifers culture with callus significant stimulation of radial growth rate occurred and they concluded that this 'suggests that they possess some pathogenic ability and do not simply invade dead tissue' (Hendry et al., 1993). Additionally they demonstrated that undiluted culture filtrate from B. nummularia was lethal to beech callus although at a dilution of 10- 2 growth of the callus was stimulated (Hendry et aI., 1993). However, B. nummularia produces 5-methylmellein in culture and there is no evidence that this or any other, as yet unidentified, compounds are phytotoxic or are implicated in canker formation (Anderson et aI., 1983; Whalley & Edwards, 1995). Another Biscogniauxia species, B. nothofagi is associated with dieback and canker in Nothofagus cunninghamii Oerst. in Tasmania, Australia (Whalley, Lress0e & Kile, 1990a; Fig. 2). It was found that the Biscogniauxia was most frequent on trees which had been attacked by Chalara australis Kile & J. Walker. The Chalara causes severe vascular disease resulting in crown wilt and general water stress (Kile & Walker, 1987) thus providing the necessary conditions for the Biscogniauxia to develop and become invasive. The association between water stress and tree diseases caused by xylariaceous fungi has, however, been recognized for many years. Biscogniauxia atropunctata causes a serious drought related disease on oaks in the southern states of North America (Thompson, 1963; Van ArsdeL 1972; Bassett, Fenn & Mead, 1982; Tainter, Williams & Cody, 1983; Bassett & Fenn, 1984). Also in the United States, Camillea punctulata causes a stem canker of Quercus (Barnett, 1957) and C. tinctor is associated with canker of American sycamore (McAlpine, 1961) and plane (Platanus occidentalis L. and P, acerifolia (Ait.) Willd.) (Hepting, 1971). Camillea punctulata is frequently linked with trees water stressed through prior infection with the oak wilt fungus, Ceratocystis fagacearum (Bretz) J. Hunt (Davis, 1966). Undoubtedly water status in the host plant is of the utmost importance concerning the development and A. J. S. Whalley subsequent presentation of disease caused by members of the Xylariaceae. Onsando (1985) suggested that tea bushes (Camellia sinensis (L.) O. Kuntze) are susceptible to wood rot by Hypoxylon serpens (Pers.: Fr.) J. Kickx (= Nemania serpens (Pers.: Fr.) Gray) following sun scorch, and Agnihothrudu (1967; 1978) reported nine species of xylariaceous fungi, mainly Hypoxylon, as traumatic parasites of tea. The most unusual association, however, must be that observed between H. rubiginosum and catalpa (Catalpa bignonioides Walt.) which was reported from the campus of the University of Georgia, U.S.A. (Weidell, 1942). Local fishermen obtain the Catalpa worm (Ceratomia catalpae) for bait by beating the trunks of the trees with clubs to dislodge them, thus causing localized injury leading to the development of canker caused by the Hypoxylon (WeidelL 1942). A recent study in Poland on alder (Alnus incana (L.) Moench) which had been weakened by industrial emissions found that under these conditions H. fuscum (Pers.) Fr. became a serious pathogen causing a very intensive white pocket rot in wood of dead alders (Domanski & Kowalski, 1987). Although Miller (1961) considered H. fuscum to be a saprotroph on members of the Betulaceae, Rogers (1968d) suspected that it might be parasitic on Alnus tenuifolia Nutt. growing in North America. He did, however, stress that infection of trees in nature takes place under very specific conditions (Rogers, 1968d) In the majority of cases the xylariaceous species causing canker diseases exhibit the characteristics of endophytic fungi which then develop as stress imposed on the fungus by the healthy tree is alleviated as the tree itself becomes stressed, usually water stress. This type of infection is often referred to as latent (Rayner & Boddy, 1986). Root rot infections. A very diverse range of Xylariaceae causes root rot diseases (Table 4). Rosellinia and Kretzschmaria species are the most widespread and economically important although occasionally, and under specific conditions, some species of Xylaria can become damaging. Rosellinia necatrix is outstanding as a plurivorous pathogen and apparently has a worldwide distribution (Sivanesan & Holliday, 1972 b; Rogers, 1979a; Teixeira de Sousa & Whalley, 1991). According to Sztejnberg & Madar (1980) R. necatrix can infect over 170 plant species from 63 genera and 30 families. In a report on the differences in susceptibility of a range of plant species Teixeira de Sousa (1985) noted that herbaceous plants were also infected and could be regarded as a potential source of inoculum. Since R. necatrix rarely produces ascocarps in nature and most identifications are through necessity based on vegetative characters there is some uncertainty concerning the published outstandingly wide host range and geographical distribution (Teixeira de Sousa & Whalley. 1991). It has been suggested that R. necatrix is probably more prevalent in temperate areas with the closely related species, R. bothrina (Berk. & Br.) SacCo (= R. arcuata Petch) being more usual in the tropics (Francis, 1985; Sivanesan & Holliday, 1972 b). Although a method for the induction of mature ascocarps in R. necatrix has been developed (Teixeira de Sousa & Whalley, 1991), long term experiments with isolates from different hosts and from temperate and tropical regions still need to be 909 undertaken to determine precisely the differences, if any, between R. necatrix and R. bothrina. There are many reports of serious problems caused by R. necatrix on a variety of economically important plants in Europe, e.g. apple, grape vine, pear, plum, sweet cherry, poplar, jasmine and scented geranium (Cellerino, 1973; Cellerino & Anselmi, 1980; Guillaumin, Mercier & Dubois, 1982; Delatour & Guillaumin, 1985; Teixeira de Sousa, 1985; Cellerino, Anselmi & Giorcelli, 1988; Teixeira de Sousa et al., 1995). In the Alcobac;:a region of Portugal 42 % of the orchards are infected with Rosellinia and 14 % of the apple trees exhibit advanced disease symptoms and are either dead or dying (Teixeira de Sousa et aI., 1995; Fig. 6) and in Italy 'R. necatrix represents one of the most dangerous agents of root-rot in Poplars' (Cellerino & Anselmi, 1980) and can cause a loss of about 0'5-1 % in total production. However, in some regions this loss may reach 5-10 % and in some plantations it can be as high as 20--25 % (Cellerino et al., 1988). In a study on factors influencing the incidence of R. necatrix in poplars Anselmi & Giocelli (1990) found that R. necatrix spread readily on loose soil with a high sand content. Furthermore they noted that soil moisture content near field capacity encouraged mycelial spread from tree to tree but dry conditions rendered the trees liable to attack (Anselmi & Giocelli, 1990). The need to remove from the soil any woody or organic material which might have had contact with Rosellinia was also seen to be an important aspect of a control strategy (Anselmi & Cellerino, 1986; Anselmi & Giocelli, 1990). In New Zealand where R. necatrix causes a serious white root rot in walnut Uuglans regia L.) the importance of removing remains of dead trees, especially their roots, was recognized to be necessary for the restriction of the disease (BoesewinkeL 1977). Solarization of soil prior to planting or exposure of infected roots to air, light and summer heat coupled with treatment of the soil with 0'1-0'2 % suspensions of benomyl or thiabendazole compounds seem to offer the best chance of control at present (Teixeira de Sousa, 1985; Sztejnberg et al., 1987; Anselmi & Giocelli, 1990). LiUle is known about the pathogenicity of R. necatrix but the role of toxins are strongly implicated. It produces in culture a number of interesting metabolites including rosellinic acid (Chen, 1960, 1964), cytochalasin E (Aldridge, Burrows & Turner, 1972; Whalley & Edwards, 1995) and rosellichalasin (Kimura, Nakajima & Hamasaki, 1989). Preliminary findings show that cytochalasin E induces a number of the typical disease symptoms caused by R. necatrix infection when applied to cut branches of apple (Whalley, unpublished). Rosellinia bunodes is a truly tropical species and is reported to be widespread in tropical America, Central African Republic, India, Indonesia, Malaysia, Philippines, Sri Lanka, Zaire and parts of Central America (Sivanesan & Holliday, 1972 a). It causes black root rot of tropical and subtropical woody hosts but is most frequent on cacao (Theobroma cacao L.), quinine (Cinchona spp.), coffee (Coffea spp.), rubber (Hevea brasiliensis Muel!.) and tea (Camellia sinensis) (Sivanesan & Holliday, 1972a). Unlike R. necatrix it normally produces perithecia in nature and these develop a distinctive wart-like ornamentation (Fig. 10) and have ascospores with characteristically long tapering ends (Fig. 32). Transmission is via mycelium The xylariaceous way of life impregnated organic litter or woody debris (Sivanesan & Holliday, 1972a). There is no evidence for the production of cytochalasin E or any of the other phytotoxic compounds known from R. necatrix (Edwards & Whalley, unpublished). Rosellinia pepo is another tropical species causing black root rot. It is however, apparently restricted to central America, the West Indies and West Africa (Booth & Holliday, 1972). Coffee is the most important host although it is described as being plurivorous (Booth & Holliday, 1972). Recently an interesting ring-die back in creeping willow (Salix repens L.) has been attributed to R. desmazieresii which attacks the roots and underground stems causing chlorosis. wilting and death of plants (Barrett & Payne, 198].; Ofong. Pearce & Barrett, 1991). There also appears to be good correlation between environmental conditions occurring at the time of development of new disease rings and conditions favourable for the growth and development of the Rosellinia in vi/ro (Ofong et al., 1991). It is known to produce cytochalasin D in laboratory culture but it is not known if it is produced in the field or if it has pathological implications (Whalley & Edwards, 1995). Kretzschmaria deusta (= Ustulina deusta (Hoffm.: Fr.) Lind) has long been recognized as an important pathogen of various tree species (Wilkins, 1934). It is associated with the base of trunks but infects through the roots (Wilkins, 1943). More recent accounts show that in Czechoslovakian forests 11-20% of beech (Fagus) trees were infected by this fungus (Cerny, 1970, 1975) and in the Sara mountains of Yugoslavia Prijincevic (1982) recorded an infection rate of up to 42 % of trees in some localities. In Britain K. deusta is a common cause of decay in broadleaved trees, especially beech (Fagus) and elm (Ulmus) (Burdekin, 1977). Recently Greig (1989) reported on the decay of horse chestnut (Aesculus hippocastanum L.) caused by this fungus. He concluded that infection was probably via the roots and that K. deusta is much more widespread than had previously been assumed (Greig, 1989). A tropical version of this fungus, often referred to as UstuIina zonata (Lev.) Sacco (Miller, 1961), is responsible for serious root rot in rubber (Hevea brasiliensis) but it also infects a number of other commercially important plants in West Malaysia (Varghese, 1971). Recently the disease has been on the increase in the rubber plantations of South India although the application of fungicides with a petroleum wound dressing has proved effective in controlling the disease provided treatment is undertaken at an early stage of the infection (Idicula et aL 1990). The closely related K. clavus is now proving to cause a very damaging root decay in macadamia (Macadamia intergrifoIia Maiden & Betche) in Hawaii (Ko, Kunimoto & Maedo, 1977) and Taiwan (Ann & Ko, 1988). Kretzschmaria clavus is widespread in the tropics and subtropics (Whalley, 1993, 1995) and occurs on a variety of forest trees (Ko, Tomita & Short, 1986). In Hawaii, most macadamia orchards with serious Kretzschmaria root rot problems are located near forests of Metrosideros collina (Forst.) Gray subsp. polymorpha (Gang.) Rock. The orchard areas were originally covered by forest and the discovery that K. clavus is a common inhabitant of M. collina subsp. polymorpha led Ko et al. (1986) to speculate that forest trees provided the source of the infection in macadamia. Kretzschmaria clavus is not the only xylariaceous 910 species to infect macadamia in Hawaii. Recently Ko & Kunimoto (1991) demonstrated the pathogenicity of Xylaria arbuscula to macadamia but concluded that in this case infection occurs via the trunk and not the roots. There are, however, a number of species of Xylaria which cause root infections. Xylaria mali is the cause of damaging black root rot of apple in the southern Appalachian states of America (Fromme, 1928; Dozier et al., 1974; Clayton, Julis & Sutton, 1976) but is localized. According to Sivanesan & Holliday (1972c) X. polymorpha can act as a weak pathogen which enters through wounds and has been associated with Acer rubrum, Coffea arabica L., Platanus acerifolia and several other hosts. Needle blight diseases. Needle blights of conifers caused by species of Rosellinia have been reported from Europe and N. America on a number of occasions (Francis, 1986). Rosellinia herpotrichioides has usually been judged to be responsible causing outbreaks on young plants of Douglas fir (Pseudotsuga menziesii (Mirb.) Franco) in forest nurseries (Salisbury & Long, 1956; Smith, 1966), on Sitka spruce seedlings (Picea sitchensis (Bong.) Carr), (Shea, 1964) and on three-year old plants of Picea abies (L.) Karsten (Vegh, 1984). However taxonomic studies on these and related Rosellinia species led Francis to conclude that all of these infections in young conifer seedlings were caused by R. minor and not R. herpotrichioides (Francis, 1986). She also maintained separate taxonomic status for R. herpotrichioides noting that apart from sound taxonomic differences it had been found causing needle blight on lower branches of mature hemlock (Tsuga canadensis (L.) Carr) growing nearest to the stream banks along coves in the Pisgah National Forest in N. Carolina; U.s.A. (Hepting & Davidson, 1937), and therefore a very different situation to the nursery beds where R. minor occurred (Francis, 1986). A number of other reports of Rosellinia causing disease in conifers (e.g. Georgescu & Ga§met, 1954; Petrak, 1961; Burmeister, 1966) can be attributed to R. minor (Francis, 1986). There is no evidence to link other species of Rosellinia, e.g. R. aquila (Fr.) De Not. and R. thelena (Fr.) Rabenh. with needle blight diseases and current evidence indicates that R. minor can cause serious loss in forest nursery beds (petrak, 1961); especially where there is overcrowding, prolonged high humidity and a growth form giving a dense lower canopy of foliage (Georgescu & Gasmet, 1954; Burmeister, 1966; Francis, 1986; Cech, 1990). The Xylariaceae as endophytes Endophyte fungi have been described as those organisms that live inside the plant tissue for at least part of their life cycle without causing any disease symptoms in the host (Petrini, 1991). Over the past 20 years an increasing number of investigations of endophytic fungi have drawn attention to the almost ubiquitous, and some times dominant, presence of members of the Xylariaceae especially in tropical plants (e.g. Petrini & Petrini, 1985; Rodrigues, 1994; Rodrigues & Samuels, 1990; Rogers, Stone & Iu, 1994; Sieber-Canavesi & Sieber, 1987; Whalley, 1993; Petrini, Petrini & Rodrigues, 1995). A total of eleven genera of Xylariaceae have been A. J. S. Whalley recorded as endophytes (Table 5) with Camillea predided to be a future addition (Petrini, Petrini & Rodrigues, 1995). Identification of xylariaceous endophytes is often difficult since they fail to produce suitable diagnostic features and only very infrequently form their teleomorph in culture. In spite of this the pioneering work of Petrini and colleagues (e.g. Petrini, 1992; Petrini & Petrini, 1985; Petrini, Petrini & Fisher, 1987; Petrini & Rogers, 1986; Petrini, Petrini & Rodrigues, 1995) has resulted in the development of keys and the necessary information to enable confident identification to be made to generic level for temperate isolates. The situation regarding tropical endophytic Xylariaceae is much more complex however, as a result of their abundance and impressive diversity (Rodrigues & Samuels, 1990; Whalley, 1993). It is doubtful whether differentiation of species on the basis of cultural and anamorphic features alone will ever be possible since differences between individual species are often insufficient to allow for absolute identifications to be made (Petrini, Petrini & Rodrigues, 1995). However, studies with Xy/aria indicate that a combination of morphological charaders and biochemical analyses might enable satisfadory identifications to be made (Brunner & Petrini, 1992; Rodrigues, 1992; Rodrigues, Leuchtmann & Petrini, 1993). There are also indications that secondary metabolite profiles from endophytic isolates might be 'matched' with those obtained from cultures derived from teleomorphic material thus enabling identity to be established (Whalley & Edwards, 1995). A preliminary study of X. cubensis (Mont.) Fr. comparing secondary metabolites e.g. cubensic acid (Adeboya et al., 1995 a) obtained from teleomorphic derived cultures with those produced by endophytic Xy/aria isolates from Euterpe o/eracea Mart. leaves confirmed the findings of Rodrigues that they belonged to X. cubensis (Edwards & Whalley, unpublished; Rodrigues, 1992). Ongoing research on secondary metabolites of the Xylariaceae will include endophytic isolates to determine the suitability of this approach for the identification of endophytic members of the family. Inoculation experiments to produce the teleomorphs provide another, but long term, alternative. Although Xylariaceae are frequently isolated from living tissues of many plants (Petrini, Petrini & Rodrigues, 1995) their isolation can be dependent upon the isolation method used. Chapela & Boddy (1988b) and Chapela (1989) reported on an increase in frequency of recovery of isolates of xylariaceous species using seledive isolation methods. Application of different drying regimes to Fagus grandifolia Ehrh. and Populus tremu/oides resulted in the isolation of Xylariaceae at levels of 32 % and 41% of all of the endophytic species in these hosts. Their recovery using non-seledive methods was rare (Chapela, 1989). It is perhaps noteworthy that H. mammatum was recovered from P. tremuloides and H. fragiforme from Fagus and therefore both species were isolated from the host tree species on which they usually produce their teleomorph in nature (Miller, 1961). In the case of H. fragiforme there is an intricate host/fungus recognition mechanism whereby ascospore germination is mediated by monolignol glucosides, present in the host, ading as specific recognition messengers (Chapela et a/., 1991). Ascospores are primed to germinate through the process of eclosion in the presence of the messenger and therefore germinate rapidly on 911 their desired host (Chapela et a/., 1990; 1993). There appears to be a similar mechanism operating between D. concentrica and Fraxinus (Gaskell, 1995). An increasing number of studies now show that individual xylariaceous species form a dominant part of the endophytic biota in certain tropical plant leaves. Thus X. cubensis was the second most frequent species isolated from leaves of Licua/a ramsayi (Muel!.) Domin. (Rodrigues & Samuels, 1990) and an unidentified species of Xy/aria was a frequent inhabitant of Sty/osanthes guianensis Sw. leaves (Pereira, Azevedo & Petrini, 1993). In her study of leaf endophytes of the tropical palm, E. olemcea, Rodrigues (1992) found X. cubensis to be numerically the most important species although other xylariaceous genera were present. These included species of Anthostomella, Da/dinia and Hypoxy/on (Rodrigues, 1992). It would be a mistake to think that aiL or even the majority, of the Xylariaceae live as endophytes. It could be predided that those species of Xylaria which form their teleomorph on fallen leaves and fruits are endophytic. However, current information suggests that they are not and that the leaves or fruits ad as 'baits' in the litter layer. Xy/aria carpophi/a (Pers.) Fr. only occurs on fallen beech (Fagus) cupules (Rogers, 1979 b) but it is widely distributed in the British Isles and Europe (Dennis, 1981; Watling & Whalley, 1977) and elsewhere e.g. North America (Rogers, 1979b) and Japan (Yokoyama & Shidei, 1972). Incubation of beech cupules taken diredly from the tree or 'captured' in suspended litter nets always failed to develop stromata of X. carpophila although the fungus was consistently present on fallen cupules in the litter. However, incubation of cupules from the litter which showed no signs of the Xy/aria frequently developed typical stromata after several weeks incubation in a damp chamber (Whalley, 1987; Whalley & Gaskell, unpublished). Preliminary studies with fruit inhabiting Xy/aria species in Thailand indicate a similar pattern. Similarly Lress0e & Lodge (1994) could find no evidence that x. axifera Mont., a species specifically associated with fallen Araliaceae petioles, could invade attached petioles. They failed to isolate X. axifera from either diseased or healthy senescent petioles collected shortly before or after abscission or from healthy non-senescent petioles obtained from the canopy. They concluded that it is likely that the petioles are colonized by X. axifera once they reach the ground (Lress0e & Lodge, 1994). In the same study they suggested that X. guareae Lress0e & Lodge, a species host specific to Guarea guidonia (L.) Sleumer, might be endophytic. Investigation of other host specific species and those occurring on leaves, petioles or fruits should prove interesting and rewarding. In spite of these unpredicted findings, the Xylariaceae occur as endophytes of many plants on a worldwide basis although current evidence shows them to occur more frequently in the tropics (Rodrigues, 1994; Rodrigues & Samuels, 1990). The outstanding questions remaining unanswered concern their presence in healthy plant tissue, frequently in a host species on which their teleomorph is never formed in nature, and how they get there? In a comparison by means of somatic incompatibility testing and gel electrophoresis of isoenzymes of thirty strains of X. cubensis, isolated from leaves of the Brazilian rainforest palm E. oleracea, a high degree of genetic The xylariaceous way of life 912 Table 5. Genera of the Xylariaceae with known endophytic representatives Genus Authentication Anlhaslamella Biscogniauxia Daldinia Hypaxylan Krelzschmaria Nemania Rasellinia Xylaria Petrini Petrini Petrini Petrini Petrini Petrini Petrini Petrini & Petrini, 1985; Petrini el aI., 1987 & Miiller, 1986 & Petrini, 1985; Petrini & Miiller, 1986 Miiller, Petrini. Petrini, Petrini, & Petrini, & & & & 1986 1985; Petrini & Miiller, 1986 1985; Petrini & Rogers, 1986 1985; Petrini, 1992 1985 diversity was shown to occur (Rodrigues, Leuchtmann & Petrini, 1993). They concluded that this diversity indicated that the endophytic thalli of X. cubensis had been established by ascospores. Since both teleomorphic and anamorphic stromata were found to occur in the vicinity they suggested that a conidial inoculum might also be involved and concluded that colonization of the leaves probably occurred via airborne propagules (Rodrigues et al., 1993). In a study of ribosomal DNA length polymorphisms within populations of X. magnoliae (Gowan & Vilgalys, 1991) concluded that the populations are not highly clonal and that it seemed likely that conidia or ascospores serve for longdistance dispersal. This would account for the high level of variability found within narrow populations, and the occurrence of the same phenotype in distant localities (Gowan & Vilgalys, 1991). Xylaria magnoliae is, however, a fruit inhabiting species and on current evidence is unlikely to be an endophyte but the implication that conidia may provide an infection inoculum in this and X. cubensis, a common endophyte, is significant. Greenhalgh & Roe (1984) reported on the presence of differing nuclei in the conidia of a range of xylariaceous taxa, including species of Xylaria. They found considerable differences in germination percentages between species but they could not correlate this with the presence of normal or irregular nuclei and they concluded that 'it seems likely that the conidia of these fungi possess the ability to germinate but that certain conditions, as yet unknown, are required' (Greenhalgh & Roe, 1984). It is tempting to suggest that a specific host fungus recognition occurs for conidia in a similar fashion to the eclosion phenomenon observed by Chapela et al. (1990, 1991, 1993), for ascospores of some species of Hypoxylon, HOST, HABITAT, AND DISTRIBUTION The distribution of members of the Xylariaceae as for other fungi is influenced by a number of factors; usually interacting. These include host specificity or preference, nature of the habitat, and climatic interactions. Host preference There are few examples of absolute specificity amongst the Xylariaceae but many where a particular taxon has a strong host preference (Table 6; Rogers, 1979a; Whalley, 1985). Thus, Rosellinia buxi has only been found growing on Buxus sempervirens L. (Petrini, 1992; Whalley & Hammelev, 1988), H. fraxinophilum is almost exclusive to Fraxinus (Pouzar, 1972) and B. nummularia appears restricted to Fagus (Whalley & rather than selectivity is Edwards, 1987), Host ｰｲ･ｦｾｮ｣ preferred since an appraisal of host relationships in the Xylariaceae reveals a number of misconceptions (Granmo et aI., 1989). In a survey of Biscogniauxia and Hypoxylon sensu stricto in the Nordic countries they found host range to be much wider than generally believed. A total of 1113 specimens of H. multiforme Fr. was examined and, although Betula (37%) and Alnus (23 %) were clearly the dominant trees involved, 13 host species were identified. Similarly H. fuscum occurred on a total of 8 tree species but Alnus (35 %) and Corylus (29%) were strongly preferred hosts. Biscogniauxia nummularia (Fagus), B. marginata (Fr.) Pouzar (Sorbus), B. cinereolilacina (J. H. Mill.) Pouzar (Tilia) and H. vogesiacum var. macrosporum (Salix) all appeared to have specific host associations (Granmo et aI., 1989). In Daldinia concentrica, one of the largest and more spectacular members of the family, host varies according to geographical situation (Whalley & Watling, 1982). This taxon is widely distributed throughout the world and is wellrepresented in Britain (Child, 1932; Whalley & Watling, 1982). Its frequency, however declines towards the North and in Scotland it is uncommon, In the South of Britain Fraxinus is the preferred host but this eventually becomes replaced by Betula in the North of England and Scotland (Whalley & Watling, 1980b, 1982). According to Winter (1887) and von Arx & Muller (1954) Fraxinus and Alnus are the most common substrata in Great Britain and on the continent but in Norway Alnus and Betula are the usual host trees (Eckblad, 1969). Whalley & Watling (1982) pointed out that the ability to spread north on trees other than the most frequent host is well-known for basidiomycetes. Thus Fomes fomentarius (Fr,) J, Kickx is common on Betula in Scotland but frequents Fagus south of the border (Watling, 1978) whilst Inonotus obliquus (Pers,: Fr.) Pilat has a wide host range in continental Europe and in Britain but although widespread in Scotland, especially in the northern regions, it is restricted to Betula, even in mixed communities (pegler, 1964). Daldinia concentrica, along with an increasing number of the Xylariaceae, is a common endophyte which occurs in a diverse range of host plants in which it fails to produce a teleomorph (Petrini et al., 1995). It is likely therefore that this and other species can exist as endophytes well outside of their normal host range but can only be detected if selective isolation techniques are applied. This creates serious problems when mapping the distribution of these fungi. Although host association is fairly well documented for many of the temperate taxa for most tropical species there is only spasmodic host data with the exception of many of the fruit and seed infecting species of Xylaria, those taxa confined to palms or bamboo and the manglicolous species (Miller, 1961; Rogers, 1979b; San Martin & Rogers, 1989; Whalley, Jones & Alias, 1994), Biscogniauxia fuscella (Rehm) San Martin & J. D. Rogers on Celtis laevigata Willd., C. obularia (Fr.) Lress0e, J, D. Rogers & Lodge on Delonix regia (Bojer ex Hook) Raf. and Mangifera indica L., C. tinctor on Platanus sp. A. J. S. Whalley (Gonzalez & Rogers, 1993) and Cordia subcordata Lam. (Van der Gucht, 1992), H. dieckmanii Theiss. on Planchonella obovata (R. Br.) Pierre, H. oodes Berk. & Broome on Diospyros maritima BI. and Excoecaria agal/ocM L. (Van der Gucht & Van der Veken, 1992), X. guareae on Guarea guidonia, X. meliacearum Lcess0e on fine litter of trees in the Meliaceae, including species of Thiehilia and Guarea, and X. axifera on fallen petioles of Araliaceae, especially Schefflera spp. (Lcess0e & Lodge, 1994) are examples of a select band for which host substratum details are available. Habitat selectivity The Xylariaceae are not confined to wood but occupy a wide range of habitats including dung, litter and associations with insects (Table 7). Although the vast majority are wood inhabitants there are differences in the nature of the wood which is suitable for particular species. Thus a distindion can be made between those which are associated with bark, those with decorticated wood, and those with no apparent preference. Hypoxylon fragiforme, H. fuscum and B. nummularia are examples of taxa which occur on freshly fallen branches or even on branches still attached to the parent tree. They are considered to be endophytic which explains their rapid appearance in dying or recently dead branches (see above). Euepixylon udum and N. confluens in contrast only occur on decorticated wood which is also well decomposed. In both species there is strong host preference for Quercus and they usually occur in wet oakwoods appearing mainly on water sodden wood (Whalley & Watling, 1980a; Whalley, 1985). The physical environment for the development of many of the wood inhabiting Xylariaceae is also important. In the tropics most species of Camillea, Biscogniauxia and Hypoxylon are found in open to sun-exposed sites often occurring in clearings or tree fall sites (Whalley, 1993; Van der Gucht & Whalley, 1996). In contrast Kretzschmaria and Xylaria are more usually well represented in dense, more wet, forest situations (Whalley, 1993; Van der Gucht & Whalley, 1995) Poronia, Podosordaria, Hypocopra and Wawelia are specifically dung inhabitants (Rogers, 1979a; Whalley, 1985). With the exception of Wawelia they possess ascospores with gelatinizing outer walls which are important in their attachment to herbage (Krug & Cain, 1974a, b; Rogers 1970). All four genera are adapted for dry conditions (Rogers, 1979a; Minter & Webster, 1983; Lundqvist, 1992). In Hypocopra the stromata are more or less rudimentary, frequently with an external clypeus and the dung itself probably ads as a substitute stroma. Poronia and Podosordaria form stalked stromata which raises their perithecia above the substratum and improves the efficiency of ascospore dissemination. Wawelia species appear to be genuinely rare and are strongly associated with leporid droppings in dry habitats (Lundquist, 1992). Minter & Webster (1983) demonstrated the need for low relative humidity for the development of stromata by W. oetospora Minter & J. Webster. The appearance of a Wawelia species at several localities in Devon following the unusually hot summer of 1995 is therefore unlikely to be coincidental (Webster, pers comm.). Little is known about the adivities of these dung fungi in nature but Poronia punctata (L.: Fr.) Fr. produces a 913 Table 6. Examples of host selectivity Species Host Biscogniauxia nummularia (Bull.: Fr.) Kuntze B. mediterranea (De Not.) Kuntze B. nothofagi Whalley. ｌｾｳｓＰ･ 8< Kile Daldinia vemicosa (Schwein.) Ces. 8< De Not. Euepirylon udum (Pers.: Fr.) ･Ｐｳｾ 8< Spooner Hyporylon cohaerens Pers.: Fr. H. cohaerens var. microsporum J. D. Rogers 8< Cando Hypoxylon fragiforme (Pers.: Fr.) J. Kickx H. frarinophilum Pouzar H. rutilum Tul. 8< C. Tul. H. voges;acum var. macrospora J. H. Mill. Nemania conf/uens (Tode: Fr.) ･Ｐｳｾｌ 8< Spooner Rosel/inia buxi Fabre R. desmazieresii (Berk. & Broome) Sacc. Xylaria longipes Nitschke Fagus Quercus Nothofagus Wex Quercus Fagus Quercus Fagus Fraxinus Fagus Salix Quercus Buxus Salix Acer number of antibiotics, the pundaporonins, in culture (Edwards et aI., 1988). Wicklow & Hirschfield (1979) had earlier shown that P. punctata and Hypocopra merdaria (Fr.) Fr. are late colonists of cattle faeces and that they were antagonistic to all earlier-sporulating colonists in cultural tests. The association between certain Xylariaceae and litter is now well established with the growing number of reports on species restrided to this habitat (Whalley, 1985; Lcess0e & Lodge, 1994). Rogers (1979b) reported on a number of Xylaria species inhabiting fallen' fruits', sometimes with clear taxon specificity, and Lcess0e & Lodge (1994) recently discussed host and habitat association for three litter inhabiting species of Xylaria from the tropics. These, together, with H. terrieola which frequents coniferous needles in the Atlantic Pyrenees (Candoussau, 1977) form part of a discrete group of Xylariaceae associated with this specific habitat. The frequent occurrence of X. oxyacanthae on Crataegus berries in newly planted Crataegus plantations in Holland was attributed to situations where there is the right balance between shade and moisture once the fruits have reached a certain stage of decomposition (Reynders, 1983). The association between the Xylariaceae and insects, usually termites, is restrided to a few species of Xylaria (Rogers, 1979a). Xylaria nigripes is probably the most common and Widespread species and is associated with recently abandoned termite nests where it often occurs along side the basidiomycete, Termitomyees (Dennis, 1961; Sands, 1969; Hiem, 1977). When freshly removed combs of the termite, Maerotermes ukuzii Fuller, were incubated at 28°C at high humidity Xylaria spp. grew from the surface after 2-3 days and over a 2-3 week period stromata up to 30 em high developed (Rohrmann & Rossman, 1980). Surprisingly the Xylaria was found not to function in lignin decomposition in the comb and it was therefore suggested that it might provide a necessary growth factor or serve to condition the substrate for optimal growth of the associated Termitomyces sp. (Rohrmann & Rossman, 1980). These termite associated Xylaria species would appear to be intricately linked although the role of the Xylaria remains unknown. A critical examination of their taxonomic relationships and ecological activities would certainly prove interesting and worthwhile. The xylariaceous way of life 914 180 '180 --- ＭｴｲｌＮＺｾＫ｟ｽｖ］ＴＱｬ -- - --- --Fig. 35. Distribution map of Camillea tinetor (closed circle) and Hypoxylon vogesiacum vaL macrospora (star). Dashed line = palm line. Distribution In considering the distribution of members of the Xylariaceae the fad that records are almost exclusively based on their teleomorphic state requires cautious interpretation. In a study of Biscogniauxia and Hypoxylon in the Nordic countries Granmo et al. (1989) found H. fragiforme to be restricted to the temperate zone with distribution seemingly exclusively linked to the distribution of Fagus, its preferred host. Hypoxylon fragiforme, however, although considered to have a narrow host range on the basis of teleomorph presence occurs widely as an endophyte on many plant taxa and therefore has the ability to extend its range well outside of its recognized distribution (Petrini & Petrini, 1985). Distribution range might be considerably extended by ability to grow on a wide host range e.g. H. multiforme (Granmo et al., 1989) or as for D. concentrica by switching host preference in the northern latitudes (Whalley & Watling, 1982). 'Examination of the distribution of selected species or genera indicates that while many taxa are global in their distribution others genuinely seem to be restricted' (Whalley, 1993). Hypoxylon Dogesiacum var. macrospora exhibits an arctic-alpine distribution (Whalley & Petrini, 1984; Whalley & Knudsen, 1985; Granmo et al., 1989; Fig. 35) and in Europe R. diathrausta (Rehm) L. E. Petrini only occurs at high altitude in the Alps and is then limited to branches of Pinus montana Miller ssp. prostata Tubeuf or closely related taxa. Furthermore, only trees located in positions of extreme exposure are involved and it appears that branches subjected to wind burn provide the highly selective habitat for this species and, therefore, its restricted geo- graphical range (Whalley, 1985). Interestingly Ouellette & Ward (1970) found that R. diathrausta requires freezing temperatures for ascospore germination or pretreatment at _3° prior to germination at 12°. Growth is possible at 0° with an optimum of 12-15° and as a result the species is welladapted for its situation (Ouellete & Ward, 1970; Whalley, 1985). At the generic level there are several examples of restricted distribution. For example it is usually stated that Camillea is almost exclusively confined to the Americas with the largest concentration of species occurring in the Amazon region, including the Guianas, although the genus is quite well represented in Central America and Mexico (Lressoe et al., 1989; Gonzalez & Rogers, 1993). Although a clear majority of species fit this pattern C. tinctor is one taxon with wide distribution throughout the tropics (Fig. 35). It occurs in West Africa (Lressoe et aI., 1989) and in Singapore (Miller, 1961) and although Rogers et al. (1987) failed to record this, or any other Camillea species, from Sulawesi C. tinctor regularly occurs in Papua New Guinea (Van der Gucht, 1992), in Malaysia (Whalley, Whalley & Jones, 1996) and in Thailand (Thienhirun & Whalley, unpublished). The recent discovery of a new Camillea, C. selangorensis M. A. Whalley, Whalley & E. B. G. Jones, in Malaysia suggests that other Camillea species may occur outside of their traditionally recognized area (Whalley et a/., 1996). Thamnomyces is another genus known mainly from lowland tropical America with just a single species, T. camerunensis Henn. recorded from Africa (Dennis, 1961; Fig. 36). It has been suggested that the spores may be unfit for air-borne dispersal and that this might explain their limited A. J. S. Whalley 915 fig. 36. Distribution map of Rhopalostroma (closed star), Thamnomyces (open star) and Hypoxylon hians (closed circle). Dashed line palm line. Table 7. Examples of habitat selectivity Bark Biscogniauxia medi/erranea (De NoL) Kuntze Hypoxylon cohaerens Pers.: Fr. H. fragiforme (Pers.: Fr.) j. Kickx H. fraxinophilum Pouzar Decorticated wood Euepixylon udum (pers.: Fr.) LiESS0e &. Spooner Nemania caries (Schwein.) Pouzar N. confluens (T ode: Fr.) la'ss0e &. Spooner Litter Xylaria carpophila (Pers.) Fr. X. magnoliae j. D. Rogers X. oxyacan/hae Tul. &. C. Tul. x. persicaria (Schwein.: Fr.) Berk. &. M. A. Curtis H. /erricola j. H. Mill. Dung Poronia Willd. Hypocopra (Fr.) j. Kickx f. Podosordaria Ellis & Holw. Wawe1ia Namysl. Insects Xylaria brasiliensis (Theiss.) Lloyd X. furca/a Fr. X. nigripes (Klotzsch) Cooke distribution (Dennis, 1957). If Camillea and Thamnomyces are mainly South American then the monotypic Engleromyces is considered to be African (Dennis, 1961; Whalley, 1993). Its sole representative E. goetzii Henn. occurs in Uganda, Zaire, = Kenya and Tanzania (Dennis, 1961; Kokwaro, 1983) where it is classed as a parasite which is only found on the upper stems of the mountain bamboo Anlndinaria alpina K. Schum. (Kokwaro, 1983). However its discovery in the mountains of Nepal (Otani, 1982) indicates a much wider geographical range and examination of mountain bamboo in other regions could well prove rewarding. Thamnomyces is classed as predominantly South American and Engleromyces as mainly African but Rhopalosfroma, a genus of xylariaceous fungi with stipitate dark brown or black stromata usually possessing abruptly expanded convex heads, has a strong Indian and South East Asian connection. Eight of the ten known species are found there with the other two occurring in Africa (Hawksworth, 1977; Hawksworth & Whalley, 1985; Whalley & Thienhirun, 1996; Fig. 36). Another curious and distinctive member of the family, Leprieuria bacillum (Mont.) ｌｾｳＰ･Ｌ J. D. Rogers & Whalley is widely distributed in South and Central America and has not yet been reported from outside that region in spite of its unique form ･ＰｳｾｌＨ et al., 1989). Hypoxylon hians Berk. & Cooke also possesses a very distinctive form and seems to be genuinely restricted to Tasmania and Victoria in Australia and South Island, New Zealand (Miller, 1961; Whalley, 1993; Fig. 36). It therefore seems probable that regardless of the lack of intensive collecting, especially in the tropics, a number of xylariaceous species, or genera, exhibit recognizable distribution patterns or trends. Comparing different regions of the world current data indicates that the Xylariaceae are generally more numerous in the tropics with South America and Mexico The xylariaceous way of life possessing the highest species richness (Whalley, 1993; Van der Gucht & Whalley, 1996). Camillea is an important component of the xylariaceous mycota in South America and Mexico with 15 and 14 species respectively compared with & Whalley, 1996; only 2 from South East Asia (Van der ｇｾ｣ｨｴ Whalley et ai., 1996). Xylaria is much better represented in South and Central America than in Africa, South East Asia, Papua New Guinea or Europe whilst Nemania exhibits a stronger presence in Europe than elsewhere (Van der Gucht & Whalley, 1996). Annulate species of Hypoxylon are absent from Europe and Kretzschmaria has a strong presence in tropical and subtropical habitats (Van der Gucht & Whalley, 1996). The distribution of members of the Xylariaceae is a result of complex interadions including availability of suitable host species, appropriate substratum conditions, and overall climatic influences. Our understanding of their distribution can only improve as more data is accumulated but the severe lack of colleding by specialists, especially in the tropics, is a major problem at the present time. The tendency for many of the Xylariaceae to occur as endophytes, and therefore have an extended range, imposes further restridions on our ability to map their distribution with accuracy. CONCLUSIONS This consideration of the adivities of the Xylariaceae in nature highlights their different lifestyles by examining their ability to occupy different habitats, to decompose wood, to cause disease in plants, and to develop under different climatic conditions and in different geographical regions. (i) The Xylariaceae are worldwide in their distribution and are exceptionally well represented in the tropics especially South and Central America. (ii) The Xylariaceae are notable for their produdion of a wide range of novel secondary metabolites, including antibiotics and phytotoxic compounds. (iii) The Xylariaceae show adaptation to life in dry habitats well illustrated by the xerophilic Wawelia. (iv) Most wood inhabiting species have the ability to cause white rot. (v) The Xylariaceae have a remarkable presence as endophytes often appearing dominant in tropical plant species. (vi) A number of xylariaceous species cause damaging disease in plants but often only when the host is water stressed. I am grateful to many colleagues and former students for their valuable discussions and contributions to our work on the Xylariaceae, in particular Dr R. L. Edwards (University of Bradford), Professor G. Granata (University of Catania), Dr T. A. Gaskell (Liverpool John Moores University), Dr G. Hadley (University of Aberdeen), Professor D. L. Hawksworth (LM.L), Professor E. B. G. Jones (University of Portsmouth), Dr N. J. Hywel-Jones (N.C.G.E.B. Bangkok), Dr G. A. Kile (CSIRO), Dr H. Knudsen (University of Copenhagen), Dr T. Lress0e (University of Copenhagen), Dr D. J. Lodge (USDA, Puerto 916 Rico), Dr D. J. Maitland (University of Bradford), Dr L. Manoch (Kasetsart University), Professor A. Nawawi (University of Malaysia), Professor R. H. Petersen (University of Tennessee), Dr L. E. Petrini (Comano), Professor J. D. Rogers (Washington State University), Dr N. M. S. Santos (Oerias), Dr G. P. Sharples (Liverpool John Moores University), Eng. A. J. Teixeira de Sousa (ENFVN, Alcoba.;a), Dr B. C. Sutton (LM.L), Dr H. K. Taligoola (Botswana), Khun S. Thienhirun (Royal Forest Department, Bangkok), Dr K. Van der Gucht (University of Gent), Dr R. Watling (RBG, Edinburgh), Professor J. Webster (University of Exeter), Mrs M. A. Whalley (Liverpool John Moores University). REFERENCES Abe, Y. (1989). Effect of moisture on decay of wood by xylariaceous and diatrypaceous fungi and quantitative changes in the chemical components of decayed woods. Transactions of the Mycological Society of Japan 30. 169-181. Adeboya. M..Edwards. R. L.. LiESS0e. T. &:Maitland. D. j. (1995 a). Metabolites of the higher fungi. Part 27. Berteric acid. cameronic acid and malaysic acid. three new polysubstituted fatty acids related to cubensic acid from species of the fungus genus Xylaria. Journal of Chemical Research(s) 1995.356-357. Adeboya. M.. Edwards. R. L.. LiESS0e. T.. Maitland. D. j. &: Whalley. A. j. S. (1995 h). Metabolites of the higher fungi. Part 28. Globoscinic acid and globoscin. a labile acid-lactone system from Xylaria glohosa and Xylaria ohovata. Journal of the Chemical Society. Perkins Transactions 1. 2067-2072. Agnihothrudu. V. (1967). Some probable traumatic parasites on tea and rubber in Southern India. Hypoxylon serpens (tea) and H. ruhiginosum (rubber). Indian Phytopathology 20. 196-198. Agnihothrudu, V. (1978). Species of Hypoxylon occurring on some plantation crops in India. In Taxonomy of Fungi (ed. C. V. Subramanian). pp. 278-281. University of Madras: India. Aldridge. D. c., Burrows. B. F. &: Turner. W. B. (1972). Structures of the fungal metabolites cytochalasin E and cytochalasin F (Rosellinia mcatrix). Journal of the Chemical Society. Chemical Communications Section D 3. 148-149. Anderson. j. R.. Spencer, H. &: X-ray crystal skeleton from Briant. C. E.. Edwards. R. L.. Mabelis. R. P.• Poyser. j. P.• Whalley. A. j. S. (1984a). Punctatin A (antibiotic M95464): structure of a sesquiterpene alcohol with a new carbon the fungus Poronia punctata. Journal of the Chemical Society. Chemical Communications. 405-406. Anderson, j. R., Edwards. R. L.. Freer. A. A.. Mabelis. R. P.. Poyser. j. P.. Spencer. H. &: Whalley. A. j. S. (1984 b). Punctatins B and C (antibiotics M95154 and M951560: further sesquiterpene alcohols from the fungus Poronia pune/ata. Journal of the Chemical Society. Chemical Communications. 917-919. Anderson. j. R.• Edwards. R. L. &: Whalley. A. j. S. (1982). Metabolites of the higher fungi. Part 19. Serpenone. 3-Methoxy-4-methyl-5 prop-l-enylfuran2(5H)-one. A new y-Butyrolactone from the fungus Hypoxylon serpens (Barron's strain) (Persoon ex Fries) Kickx. Journal of the Chemical Society. Perkins Transaclions 1. 215-221. Anderson. j. R.. Edwards. R. L. &: Whalley. A. J. S. (1983). Metabolites of the higher fungi. Part 21. 3-Methyl-3,4-dihydroisocoumarins and related compounds from the ascomycete family Xylariaceae. Journal of the Chemical Society, Perkins Transaclions 1. 2185-2192. Anderson. j. R., Edwards. R. L. &: Whalley, A. j. S. (1985). Metabolites of the higher fungi. Part 22. 3-Butyl-3-methylsuccinic acid and 2-hexylidene-3methylsuccinic acid from XyJariaceous fungi. Journal of the Chemical Society, Perkins Transaclions 1. 1481-1485. Anderson. N. A.. Ostry, M. E. &: Anderson. G. W. (1979). Insect wounds as infection sites for Hypoxylon mammatum on trembling aspen. Phytopathology 69. 476-479. Anderson. R. L. (1964). Hypoxylon canker impact on aspen. Phytopathology 54. 253-257. Ann. P. J. &: Ko. W. H. (1988). Root rot of macadamia caused by Ganoderma A.I.S. Whalley lucidum and Kretzschmaria clavus. Journal of Agricultural Research of China 37, 424-429. Anselmi, N. & Cellerino, G. P. (1986). Influence of plant residues in soil on the diffusion of Rosel/inia necatrix Prill. IUFRD World Congress, 52.06.00, Ljubljana (Yugoslavia), September 7-21, 17pp. Anselmi, N. & Giorcelli, A. (1990). Factors influencing the incidence of Rosel/inia necatrix Prill. in poplars. European Journal of Forest Pathology 20, 175-183. Arx, j. A. von & Muller, E. (1954). Die Gattungen der amerosporen Pyrenomyceten. Beitriige Kryptogamenflora Schweiz 11, 1-434. Bagga, D. K. & Smalley, E. B. (1969). Factors affecting canker development on Populus rremuloides artificially inoculated with Hypoxylon pruinatum. Canadian Journal of Botany 47, 907-914. Barbosa, M. A. de F. (1958). 0 carvao do enttrecasco Hypoxylon mediterraneum (De Not.) Ces. et De Not. Publicacois direcCiio Geral dos Serviros florestais e Agricolas 25, 95-132. Barnett, H. L. (1957). Hypoxylon punctulatum and its conidial state on dead oak trees and in culture. Mycologia 49, 588-595. Barrett, D. K. & Payne, K. (1982). Ring-dying of Salix repens caused by Rosellinia desmazieresii. Transactions of the British Mycological Society 78, 566--569. Bassett, E. N. & Fenn, P. (1984). Latent colonization and pathogenicity of Hypoxylon atropunctatum on oaks. Plant Disease 68, 317-319. Bassett, E. N., Fenn. P. & Mead, M. A. (1982). Drought-related oak mortality and incidence of Hypoxylon canker. Arkansas Farm Research 31, 8. Beckett, A. (1979a). Ultrastructure and development of ascospore germ slit in Xylaria longipes. Transactions of the British Mycological Sociely 72, 269-276. Beckett, A. (1979b). Comparative ultrastructure of the ascospore germ slit in Xylaria and Daldinia. Transactions of the British Mycological Society n. 320--322. Beckett, A. & Crawford, R. M. (1973). The development and fine structure of the ascus apex and its role during spore discharge in Xylaria longipes. New Phytologist n, 357-369. Betina, V. (1989). Mycotoxins: Chemical, Biological and Environmental Aspects. Elsevier: Amsterdam. Bier, j. E. & Rowat. M. H. (1962). The relation of bark moisture to the development of canker diseases caused by native facultative parasites. VIII. Ascospore infection of Hypoxylon pruinatum (Klotzsche) Cke. Canadian Journal of Botany 40, 897-901. BlaisdelL D. j. (1939). Decay of hardwoods by Ustulina vulgaris and other ascomycetes. Phytopathology 29, 2. BoesewinkeL H. j. (1977). Walnut diseases. New Zealand Journal of Agriculture, November 1977, 21-22. Boddy, L. (1994). Latent decay fungi: the hidden foe. Arboricultural Journal 18, 113-135. Boddy, L., Bardsley, D. & Gibbon, O. (1987) Fungal communities in attached ash branches. New Phytologist 107, 143-154. Boddy, L., Gibbon, O. & Grundy, O. M. (1985). Ecology of Daldinia concentrica: effect of abiotic variables on mycelial extension and interspecific interactions. Transactions of the British Mycological Society 85, 201-211. Bodo, B., Davoust, D., Lecommandeur, D., Rebuffat, 5., Genetet, I. &: Pinon, j. (1987). Hymatoxin A. a diterpene sulfate phytotoxin of Hypoxylon mammatum, parasite of aspen. Tetrahedron Letters 28, 2355-2358. Booth, C. & Holliday, P. (1972). Rosel/inia pepo. C. M. I. Descriptions of Pathogenic Fungi and Bacteria. No. 351. Bruck, R. I. & Manion, P. D. (1980). Interacting environmental factors associated with the incidence of hypoxylon canker on trembling aspen. Canadian Journal of Forest Research 10, 17-24. Brunner, F. & Petrini, O. (1992). Taxonomy of some Xylaria species and xylariaceous endophytes by isoenzyme electrophoresis. Mycological Research 96, 723-733. Burdekin, D. A. (1977). Gale damage to amenity trees. Arboricultural Journal 3, 181-189. Burmeister, P. (1966). Beobachtungen uber einige wichtige Pilzkrankheiten an Zierkoniferen im oldenburgischen Baumschulgebiet. Die Gartenbauwissenschaft 31. 469-506. Callan, B. E. & Rogers, j. D. (1986). Cultural characters and anamorphs of Biscogniauria (= Nummularia) marginata, B. dennisii and B. repanda. Canadian Journal of Botany 64, 842-847. 917 Callan, B. E. & Rogers, j. D. (1990). Teleomorph-anamorph, connections and correlations in some Xylaria species. Mycotaxon 36, 343-369. Candoussau, F. (1977). Hypoxylon terricola Miller dans Ie midi de la France, espece nouvelle pour I'Europe. Mycotaxon 6, 173-177. Candoussau, F. & Rogers, j. D. (1990). Notes on Dbolarina dryophila from· France. Mycoiaxon 39, 345-349. Cartwright. K. St G. & Findlay, W. P. K. (1946). Decay of timber and its preservation. H.M.s.o.: London. Cech, 1. (1990). Rosellinia minor (Hahn.) Francis in Christbaumkultieren. European Journal of Forest Pathology 20, 113-117. Cellerino, G. P. (1973). Sur la diffusion des pourridilfs dans les pe.upleraies de la vallee du Po (note preliminaire). FAO/CIPlOIS, Casale M. - Rome, 23 mai-l juin, 1973, 5pp. Cellerino. G. P. & Anselmi, N. (1980). Poplar disease situation in 1979-1980. FAO/IPC/DISIUFRO 52.05.03 Kornik. Poland (1-5 Sept.), 7 pp. Cellerino, G. P., Anselmi, N. & Giorcelli, A. (1988). Repartition en Italie et conditions favorisant les attaques de Rosellinia necatrix Prill. sur Peuplier. Annales of the Faculty of Science and Agriculture University of Torino 15, 165-176. Cerny, A. (1970). Bionomie houby Hypoxylon deustum (Hoffm. et Fr.) Grev. jeji hospodarsky vyznam a roz = sireni v Ceskoslovensku. Lesnielvi 10. Praha. Cerny, A. (1975). Najskodlivejsi paraziticke a saprofyticke drevokazne houby buku. Soubor referatu. Pestovani, teiba a zpracovani buku c CSR. Libero. Chapela, I. H. (1989). Fungi in healthy stems and branches of American beech and aspen: a comparative study. New Phytologist 113, 65-75. Chapela, I. H. & Boddy, L. (1988a). Fungal colonization of attached beech branches 1. Early stages of development of fungal communities. New Phytologist 110, 39-45. Chapela, I. H. & Boddy, L. (1988 b). Fungal colonization of attached beech branches n. Spatial and temporal organization of communities arising from latent invaders in bark and functional sapwood, under different moisture regimes. New Phytologist 110, 47-57. Chapela, I. H. & Boddy, L. (1988c). The fate of early fungal colonizers in beech branches decomposing on the forest floor. FEMS Microbiology Ecology 53, 273-284. Chapela, I. H., Petrini, O. & Blelser, G. (1993). The physiology of ascospore eclosion in Hypoxylon fragiforme: mechanisms in the early recognition and establishment of an endophytic symbiosis. Mycological Research 97, 157-162. Chapela, I. H., ｾｩｮｲｴ･ｐ O. & Hagmann, L. (1991). Monolignol glucosides as specific recognition messengers in fungus I plant symbioses. Physiological and Molecular Plant Pathology 39, 289-298. Chapela, I. H., Petrini, O. & Petrini, L. E. (1990). Unusual ascospore germination in Hypoxylon fragiforme: first steps in the establishment of an endophytic symbiosis. Canadian Journal of Botany 68, 2571-2575. Chen, Y-S. (1960). Studies on the metabolic products of Rosellinia necafrix Berlese 1. Isolation and characterisation of several physiologically active neutral substances. Bulletin of the Agricultural Chemical Society of Japan 24, 373-381. Chen, Y-S. (1964). Studies on the metabolic products of Rosellinia necairix Berlese Part n. The structure of rosellinic acid. Agricultural and Biological Chemistry 28, 431-435. Chesters, C. G. C. (1950). On the succession of microfungi associated with the decay of logs and branches. Transactions of the Lincolnshire Naturalis!'s Union 12, 129-135. Chesters, C. G. G. & Greenhalgh, G. N. (1964). Geniculosporium serpens gen. et sp. nov., the imperfect state of Hypoxylon serpens. Transactions of the British Mycological Society 47. 393-401. Child, M. (1932). The genus Daldinia. Annals of the Missouri Botanical Garden 19, 429-496. Clayton, C. N., lulis, A. j. & Sutton, T. B. (1976). Root rot diseases of apple in North Carolina. Agricultural Experiment Station, North Carolina State University at Raleigh, Bulletin 455, 3-11. Crane, j. L. & Dumont, K. P. (1975). Hyphomycetes from the West Indies and Venezuela. Canadian Journal of Botany 53, 843-851. Dagne, E., Gunatilaka, A. A. L.. Asmellash, 5., Abate, D., Kingston, D. G.I., Hofmann, G. A. & johnson, R. K. (1995). Two new cytochalasins from Xylaria obovata. Tetrahedron 50, 5615-5620. The xylariaceous way of life Davis, 1. C. (1966). Appraisal of Hypoxylon punctulatum as a biological control agent of Ceratocystis fagacearum in oak-wilt trees. Phytopathology 66, 772-775. Day, M. W. & Strong, F. C. (1959). A study of hypoxylon canker on aspen. Michigan Agriculture Experiment Station Quarterly Bulletin 41, 87G-877. Delatour, C. & Guillaumin, J. J. (1985). Importance des pourridies dans les regions temperees. European journal of FOrest Pathology 15, 258-263. Dennis, R. W. G. (1956). Some Xylarias of tropical America. Kew Bulletin 1956, 401-444. Dennis, R. W. G. (1957). Further notes on tropical American Xylariaceae. Kew Bulletin 1957, 297-332. Dennis, R. W. G. (1961). Xylarioideae and Thamnomycetoideae of Congo. Bulletin jardin Botanique de tEtat (Bruxelles) 31, 109-154. Dennis, R. W. G. (1963). Hypoxyloideae of Congo. Bulletin jardin Botanique de tEtat (Bruxelles) 33, 317-343. Dennis, R. W. G. (1970). Fungus flora of Venezuela and adjacent countries, (Kew Bulletin, Additional Series 3). HMSO: London Dennis, R. W. G. (1981). British Ascomycetes. J. Cramer: Vaduz. Domanski, S. & Kowalski, 1. (1987). Fungi occurring on forests injured by air pollutants in the Upper Silesia and Cracow industrial regions. European journal of Forest Pathology 17, 337-348. Dozier Jr. W. A, Latham, A L Kouskolekas, C. A & Mayton, E. L. (1974). Susceptibility of certain apple rootstocks to black root rot and woolly aphids. Hortiscience 9, 35-36. Eckblad, F.-E. (1969). The genera Daldinia, Ustulina and Xylaria in Norway. Norwegian journal of Botany 16, 139-145. Eckblad, F.-E. & Granmo, A (1978). The genus Nummularia (Ascomycetes) in Norway. Norwegian journal of Botany 25, 69-75. Edwards, R. L., Maitland, D. J., Poyser, J. P. & Whalley, A J. S. (1989). Metabolites of the higher fungi. Part 25. Punctaporonin G from the fungus Poronia punctata (Linnaeus: Fries) Fries. journal of the Chemical Society, Perkins Transactions 1, 1939-1941. Edwards, R. L., Maitland, D. J. & Whalley, A J. S. (1989). Metabolites of the higher fungi. Part 24. Cytochalasin N, 0, P, Q, and R. New cytochalasins from the fungus Hypoxylon terricola Mill. journal of the Chemical Society, Perkins Transactions 1, 57-65. Edwards, R. L., Maitland, D. J. & Whalley, A J. S. (1991). Metabolites of the higher fungi. Part 26. Cubensic acid, 3,7,11,15-tetrahydroxy-18(hydroxymethyl)-2,4,6,10,14,16,20-heptamethyldocosa-4E, 8E, 12E, 16Etetraenonic acid, a novel polysubstituted C22 fatty acid from the fungus Xylaria cubensis (Mont.) Fr. with substituents and substitution pattern similar to the macrolide antibiotics. journal of the Chemical Society, Perkins Transactions 1, 1411-1417. Edwards, R. L., Poyser, J. P. & Whalley, A J. S. (1988). Metabolites of the higher fungi. Part 23. The punctaporonins. Novel bi-, tri-, and tetra-cyclic sesquiterpenes related to caryophyllene from the fungus Poronia punctata (Linnaeus: Fries) Fries. journal of the Chemical Society, Perkins Transactions 1, 823-831. Edwards, R. L. & Whalley, A J. S. (1979). Metabolites of the higher fungi. Part 18. 3-butyl-4-methylfuran-2(5H)-one and 3-butyl-4-methylenefuran-2(5H)one. New y-butyrolactones from the fungus Hypoxylon serpens (Persoon ex Fries) Kickx. journal of the Chemical Society, Perkins Transactions 1, 803-806. Eriksson, O. E. & Hawksworth, D. L. (1993). Outline of the ascomycetes 1993. Systema Ascomycetum 12, 51-257. Flores, G. & Hubbes, M. (1979). Phytoalexin production by aspen (Populus tremuloides Michx.) in response to infection by Hypoxylon mammatum (Wahl.) Mill. and Alternaria spp. European journal of Forest Pathology 5, 28G-288. Francis, S. M. (1985). Rosellinia necatrix - fact or fiction? Sydowia 38, 75-86. Francis, S. M. (1986). Needle blights of conifers. Transactions of the British Mycological Society 87, 397-400. Francis, S. M., Minter, D. W. & Caine, 1. S. (1980). Three new species of Anthostomella. Transactions of the British Mycological Society 75, 201-206. French, D. W., Hodges, C. S. & Froyd Jr, J. D. (1969). Pathogenicity and taxonomy of Hypoxylon mammatum. Canadian journal of Botany 47, 223-226. Fromme, F. D. (1928). The black root rot disease of apple. Virginia Agriculture Experiment Station Technical Bulletin 34. Furuya, K. & Udagawa, S. (1977). Coprophilous pyrenomycetes from Japan. IV. Transactions of the Mycological Society of japan 17, 243-261. 918 GaskelL T. A (1995). Development and stromal structure in Daldinia concentrica (Bolt: Fr.) Ces. & De Not. PhD. Thesis, Liverpool John Moores University, U.K. Georgescu, C. & Ga§met, V. (1954). Un atac de Rosellinia byssiseda (T ode) Schroet., la puietii de Molid. Revista Padurilor 68, 31-34. Gonsalez, F. S. M. & Rogers, J. D. (1993). Biscogniauxia and Camillea in Mexico. Mycotaxon 47, 229-258. Gowan, S. P. & Vilgalys, R. (1991). Ribosomal DNA length polymorphisms within populations of Xylaria magnoliae (Ascomycotina). American journal of Botany 78, 1603-1607. Granata, G. & Whalley, A J. S. (1994). Decline of beech associated to Biscogniauxia nummularia in italy. Petria 4, 111-116. Granmo, A, Hammelev, D., Knudsen, H., Lzess0e, T., Sasa, M. & Whalley, A J. S. (1989). The genera Biscogniauxia and Hypoxylon (Sphaeriales) in the Nordic countries. Opera Botanica 100, 59-84. Greenhalgh, G. N. & Chesters, C. C. G. (1968). Conidiophore morphology in some British members of the Xylariaceae. Transactions of the British Mycological Society 51, 57-82. Greenhalgh, G. N. & Roe, G. M. (1984). Conidial structure in Xylaria and related genera. In Taxonomy of Fungi (ed. C. V. Subramanian), pp. 365-377. University of Madras: India. Greenhalgh, G. N. & Whalley, A J. S. (1970). Stromal pigments of some species of Hypoxylon. Transactions of the British Mycological Society 55, 89-96. Greig, B. J. W. (1989). Decay in an avenue of horse chestnut (Aesculus hippocastanum L) caused by Ustulina deusta. Arboricultural journal 13, 1-6. Griffin, D. M, Quinn, K. & McMillen, B. (1986). Regulation of hyphal growth rate of Hypoxylon mammatum by amino acids: stimulation by proline. Experimental Mycology 10, 307-314. Griffin, D. M, Schaedle, M, Manion, P. D. & De Vit, MJ . (1991). Clonal variation in amino acid contents of roots, stems, and leaves of aspen (Populus tremuloides Michx.) as influenced by diurnal drought stress. Tree Physiology 8, 337-350. Griffiths, H. B. (1973). Fine structure of seven unitunicate pyrenomycete asci. Transactions of the British Mycological Society 60, 261-271. Guillaumin, J. J., Mercier, S. & Dubos, B. (1982). Les pourridies 11 Arrnillariella et Rosellinia en France sur vigne, arbres fruitiers et cultures florales. 1. Etiologie et symptomatologie. Agronomie 2, 71-80. Hawksworth, D. L. (1971). A revision of the genus Ascotricha Berk. Mycological Papers 126, 1-28, 5 pI. Hawksworth, D. L. (1977). Rhopalostroma, a new genus in the Xylariaceae s.I. Kew Bulletin 31, 421-431. Hawksworth, D. L. (1991). The fungal dimension of biodiversity: magnitude, significance, and conservation. Mycological Research 95, 641-655. Hawksworth, D. L. (1993). The tropical fungal biota: census, pertinence, prophylaxis, and prognosis. In Aspects of Tropical Mycology (ed. S. Isaac, J. c. Frankland, R. Watling & A J. S. Whalley), pp. 265-293. Cambridge University Press: Cambridge, U.K. Hawksworth, D. L. & Whalley, A J. S. (1985). A new species of Rhopalostroma with a Nodulisporium anamorph from Thailand. Transactions of the British Mycological Society 84, 56G-562. Heim, R. (1977). Termites et Champignons. Societe Nouvelle des editions, Boubee: Paris. Hendry, S. J., Boddy, L. & Lonsdale, D. (1993). Interactions between callus cultures of European beech, indigenous ascomycetes and derived fungal extracts. New Phyt%gist 123, 421-428. Hepting, G. H. (1971). Diseases of forest and shade trees of the United States. USDA-Forest Service. Agriculture Handbook 386, 658. Hepting, G. H. & Davidson, R. W. (1937). A leaf and twig disease of hemlock caused by a new species of Rosellinia. Phytopathology 27, 305-310. Hubbes, M. (1962a). Inhibition of Hypoxylon pruinatum by pyrocatechol isolated from bark of aspen. Science 136, 156. Hubbes, M. (1962 b). Two glycosides from aspen fungistatic against Hypoxylon pruinatum (Klot.) Cke. Canadian Department of Forestry, Forest Entomology and Pathology Branch, Bi-monthly Progress Report 18, 2-3. Hubbes, M. (1964). New facts on host-parasite relationships in the Hypoxylon canker of aspen. Canadian journal of Botany 42, 1489-1494. Hubbes, M. (1966). Inhibition of Hypoxylon pn<inatum by aspen bark meal and the nature of active extractives. Canadian journal of Botany 44, 365-386. Idicula, S. P., EdathiL 1. 1., Jayarathnam, K. & Jacob, C. K. (1990). Dry rot A. J. S. 919 Whalley disease management in Hevea brasiliensis. Indian journal of National Rubber Research 3, 35-39. Jensen, J. D. (]98]). The developmental morphology of Hypoxylon serpens in culture. Canadian journal of Botany 59, 40-49. Jensen, J. D. (]985). Peridial anatomy and pyrenomycete taxonomy. Mycologia 77, 688-70]. Jong, S. C. & Davis, E. E. (1973). Stromatic Neurosporas. Mycologia 65, 458-464. Jong, S. C. & Davis, E. E. (]974). Areolospora, a new humicolous genus in the Xylariaceae. Norwegian journal of Botany 21, 23-30. Jong, S. C. & Rogers, J. D. (] 969). Poronia oedipus in culture. Mycologia 60, 973-976. Jong, S. C. & Rogers, J. D. (]972). Illustrations and descriptions of conidial states of some Hypoxylon species. Washington State University Agricultural &periment Station Technical Bulletin 71, ]-49. Ju, Y-M., Gonzalez, F. S. M. & Rogers, J. D. (]993). Three xylariaceous fungi with scolecosporous conidia. Mycotaxon 47, 2]9-228. Ju, Y-M. & Rogers, J. D. (1990). Astrocystis reconsidered. Mycologia 82, 342-349. Juzwik, J., Nishijima, W. T. & Hinds, 1. E. (]978). Surveys of aspen cankers in Colorado. Plant Disease Reporter 62, 906-910. Kato, T, Kabuto, C, Sasaki, N., Tsunagawa, M., Aizava, H., Fujita, K., Kato, Y & Kitahara, Y (]973). Momilactones, growth inhibitors from rice, Oryza sativa L. Tetrahedron Letters 39, 3861-3864. Kato, 1., Tsunakawa, M., Sasaki, N., Aizawa, H., FUjita, K., Kitahara, Y & Takahashi, N. (1977). Growth and germination inhibitors in rice husks. Phytochemistry 16, 45-48. Kile, G. A. & Walker, J. (]987). Chalara australis sp. nov. (Hyphomycetes), a vascular pathogen of Nothofagus cunninghamii (Fagaceae) in Australia and its relationship to other Chalara species. Australian journal of Botany 35, ]-32. Kimura, Y, Nakajima, H. & Hamasaki, T. (1989). Structure of rosellichalasin, a new metabolite produced by Rosellinia necatrix. Agricultural and Biological Chemistry 53, ]699-]701. Ko, W. H. & Kunimoto, R. K. (]99]). Quick decline of macadamia trees: association with Xylaria arbuscula. Plant Pathology 40, 643-644. Ko, W. H., Kunimoto, R. K. & Maedo, I. (]977). Root decay caused by Kretzschmaria clavus: its relation to macadamia decline. Phytopathology 67, 18-21. Ko, W. H., Tomita, J. & Short, R. L. (1986). Two natural hosts of Kretzschmaria clavus in Hawaiian forests. Plant Pathology 38, 254-255. Kokwaro, J. O. (1983). An African knowledge of ethnosystematics and its application to traditional medicine, with particular reference to the medicinal use of the fungus Engleromyces goetzei. Bothalia 14, 237-243. Krug, J. C. & Cain, R. F. (]974a). A preliminary treatment of the genus Podosordaria. Canadian journal of Botany 52, 589-605. Krug, J. & Cain, R. F. (]974 b). New species of Hypocopra. Canadian journal of Botany 52, 809-843. Lressoe, T. (]994). Index Ascomycetum. 1. Xylariaceae. Systema Ascomycetum 1,43-112. Laessoe, T & D. I. Lodge (1994). Three host specific Xylaria species. Mycologia 86, 436-446. Lressoe, T., Rogers, J. D. & Whalley, A. J. S. (1989). Camillea, jongiella and light spored species of Hypoxylon. Mycological Research 93, 121-155. Lressoe,1. & Spooner, B. M. (1994). Rosellinia and Astrocystis (Xylariaceae): new species and generic concepts. Kew Bulletin 49, ]-70. Lonsdale, D. (1983). Some aspects of the pathology of environmentally stressed trees. Yearbook of the International Dendrology Society 1982, 90-97. Lundqvist, N. (]992). Wawelia effusa Lundqvist, spec. nov. (Xylariaceae). Persoonia 14, 4]7-423. c. Luque, J. & Girbal, J. (1989). Dieback of cork oak (Quercus suber) in Catalonia (NE Spain) caused by Botryosphaeria stevensii. European journal of Forest Pathology 19, 7-13. Macara, A. M. (1974). Principais do sobreiro. Boletim do Instituto dos Produtos Florestais, Cortica 430, 149-156. Macara, A. M. (1975). Estimativa em 1975. Dos prejuizos causados pelas principais doen<;as do sobreiro num montado da regiiio ribatejana. Boletim do Instituto dos Produtos Florestais, Cortica 444, 205-212. Manion, P. D. (]975). Two infection sites of Hypoxylon mammatum in trembling aspen (Populus tremuloides). Canadian journal of Botany 53, 2621-2624. Manion, P. D. & Griffin, D. M. (1986). Sixty-five years of research on Hypoxylon canker of aspen. Plant Disease 70, 803-808. Martin, P. (1967a). Studies in the Xylariaceae: 1. New and old concepts. journal of South African Botany 33, 205-28. Martin, P. (1967 b). Studies in the Xylariaceae: II. Rosellinia and the Primocinerea Section of Hypoxylon. journal of South African Botany 33, 3 ]5-328. Martin, P. (1968a). Studies in the Xylariaceae: III. South African and foreign species of Hypoxylon Section Entoleuca. South African journal of Botany 34, 153-199. Martin, P. (]968 b). Studies in the Xylariaceae IV. Hypoxylon. Sections Papillata and Annulata. South African journal of Botany 34, 303-330. Martin, P. (1969a). Studies in the Xylariaceae. V. Euhypoxylon. journal of South African Botany 35, 149-206. Martin, P. (1969 b). Studies in the Xylariaceae. VI. Daldinia, Numulariola and their allies. journal of South African Botany 35, 267-320. Martin, P. (1969c). Studies in the Xylariaceae VII. Anthostomella and Lopadostoma. journal of South African Botany 35, 393-4]0. Martin, P. (1970). Studies in the Xylariaceae VIII. Xylaria and its allies. journal of South African Botany 36, 73-138. Marty, R. (1972). The economic impact of hypoxylon canker on the Lake States resource. In Aspen Symposium Proceedings USDA Forest Service General Technical Report, NC-], pp. 2]-26. McAlpine, R. G. (1961). Hypoxylon tinctor associated with a canker on American sycamore trees in Georgia. Plant Disease Reporter 45, 196-]98. Merrill, W., French, D. W. & Wood, F. A. (1964). Decay of wood by species of the Xylariaceae. Phytopathology 54, 56-58. Miller, J. H. (1961). A monograph of the World Species of Hypoxylon. University of Georgia Press: Athens, U.S.A. Minter, D. W. & Webster, J. (1983). Wawelia octospora sp. nov., a xerophilous and coprophilous member of the Xylariaceae. Transactions of the British Mycological Society 80, 370-373. Nannfeldt, J. A. (]976). Iodine reactions in ascus plugs and their taxonomic significance. Transactions of the British Mycological Society 67, 283-287. Nilsson, T., Daniel, G., Kirk, 1. K. & Obst, J. R. (1989). Chemistry and microscopy of wood decay by some higher ascomycetes. Holzforschung 43, 11-18. Ofong, A. U., Pearce, R. B. & Barrett, D. K. (1991). Biology and pathogenicity of Rosellinia desmazieresii. Mycological Research 93, 189-]94. Onsando, J. M. (]985). Wood rot disease of tea (Hypoxylon serpens) - a review. Tea 6, 39-42. Otani, Y (1982). Engleromyces goetzii P. Henn. collected in Nepal. Reports on the cryptogamic study in Nepal 93-96. National Science Museum: Tokyo. Ouellette, G. I. B. & Ward, E. W. B. (1970). Low temperature requirements for ascospore germination and growth of Hypoxylon diathrauston. Canadian journal of Botany 48, 2223-2225. Pegler, D. N. (1964). A survey of the genus Inonotus (Polyporaceae). Transactions of the British Mycological Society 47,175-195. J. 0., Azevedo. J. L. & Petrini, O. (1993). Endophytic fungi of Stylosanthes: a first report. Mycologia 85, 362-364. Petrak. F. (1961). Ober das Auftreten von Rosellinia herpotrichioides Hepting Pereira, and Davidson auf jungen Fichten im Pflanzgarten Rauris der Forstverwaltung Lend in Salzburg. Sydowia 15, 242-246. Petrini, L. E. (] 992). Rosellinia species of the temperate zones. Sydowia 44, 169-281. Petrini, L. E. & Miiller, E. (1986). Haupt- und Nebenfruchtformen europiiischer Hypoxylon-Arten (Xylariaceae, Sphaeriales) und verwandter Pilze. Mycologia Helvetica I, 501-627. Petrini, L. E. & Petrini, O. (1985). Xylariaceous fungi as endophytes. Sydowia 38, 2]6-234. Petrini, L. E., Petrini, O. & Fisher, P. J. (1987). Anthostomella calligoni, an endophyte of Suaeda fruticosa in Dorset. Transactions of the British Mycological Society 89, 387-389. Petrini, L. E. & Rogers, J. D. (]986). A summary of the Hypoxylon serpens complex. Mycotaxon 26, 409-436. Petrini, O. (1991). Fungal endophytes of tree leaves. In Microbial Ecology of Leaves (ed. J. H. Andrews & S. S. Hirano), pp. 179-197. Springer Verlag: New York. The xylariaceous way of life Petrini, 0., Petrini, L. E. & Rodrigues, K. F. (1995). Xylariaceous endophytes: an exercise in biodiversity. Fitopatologia Brasi/eira in press. Pinon, j. (1979). Origine et principaux charaderes des souches francaise d'Hypoxylon mammatum (WahL) Miller. European Journal of Forest Pathology 9, 129-142. Pinon, j. & Manion, P. D. (1991). Hypoxylon mammatum and its toxinsrecent advances in understanding their relationships with canker disease of poplar. European Journal of Forest Pathology 21, 202-209. Pouzar, Z. (1972). Hypoxylon fraxinophi/um spec. nov. and H. moravicium spec. nov., two interesting species found on Fraxinus angustifolia. Ceskti Mykologie 26,129-137. Pouzar, Z. (1979). Notes on the taxonomy and nomenclature of Nummularia (Pyrenomycetes). Ceskti Mykologie 33, 207-219. Pouzar, Z. (1985 a). Reassessment of Hypoxylon serpens -complex I. Ceskti Mykologie 39, 15-25. Pouzar, Z. (1985 b). Reassessment of Hypoxylon serpens -complex II. Ceskti Mykologie 39, 129-134. Pouzar, Z. (1986). A key and conspectus of Central European species of Biscogniauxia and Obolarina (Pyrenomycetes). Ceskd Mykologie 40, 1-10. Povah, A. (I 924). Hypoxylon poplar canker. Phytopathology 14, 140-145. Poyser, j. P., Edwards, K L., Anderson, j. K, Hursthouse, M. B., Walker, N. P. C, Sheldrick, G. M. & Whalley, A. j. S. (1986). Punctatins A. D, E, and F (antibiotics M95464, M167906, M171950 and MI89122), isomeric allylic alcohols from the fungus Poronia punctata: X-ray crystal structures of D and E acetonide. Journal of Antibiotics 39, 167-169. Prijincevic, M. B. (I982). Economic significance of the infection of beech forests by Hypoxylon deustum (Hoffm. et Fr.) Grev. at Sara Mountain. European Journal of Forest Pathology 12, 7-10. Rajagopalan, C (1966). Studies on four species of woodrotting fungi. Transactions of the Kansas Academy of Sciences 68, 541-552. Rayner, A. D. M. & Boddy, L. (I986). Population structure and the infection biology of wood decay fungi in living trees. Advances in Plant Pathology 5, 119-160. Rayner, A. D. M. & Boddy, L. (1988). Fungal Decomposition of Wood. Its Biology and Ecology. john Wiley: Chichester. UK Reynders, j. (I983). Xylaria oxyacanthae en X. carpophila in het Amsterdamse Bos. Coolia 26, 60--61. Rodrigues, K. F. (1992). Endophytic fungi in the tropical palm Euterpe oleracea Mart. Ph.D. Thesis, City University of New York. U.s.A. Rodrigues, K. F. (1994). The foliar fungal endophytes of the Amazonian palm Euterpe oleracea Mart. Mycologia 86, 376-385. Rodrigues, K. F., Leuchtmann, A. & Petrini, O. (1993). Endophytic species of Xylaria: cultural and isozymic studies. Sydowia 45, 116-138. Rodrigues, K. F. & Samuels, G. j. (1989). Studies in the genus Phylacia (Xylariaceae). Memoirs of the New York Botanical Garden 49, 290-297. Rodrigues, K. F. & Samuels, G. j. (1990). Preliminary study of endophytic fungi in a tropical palm. Mycological Research 94, 827-830. Rogers, j. D. (1964). Hypoxylon pruinatum: the chromosome number. Mycologia 56, 369-373. Rogers, j. D. (1965). Hypoxylon fuscum.l. Cytology of the ascus. Mycologia 57, 789-803. Rogers, j. D. (1967 a). Hypoxylon multiforme: cytology of the ascus. Mycologia 59, 295-305. Rogers, j. D. (1967 b). Cytological aspects of the ascospores of Hypoxylon grenadense var. macrospora. Canadian Journal of Botany 45, 1166-1168. Rogers, j. D. (I968 a). Hypoxylon deustum: the chromosome number. Mycopathologia et Mycologia Applicata 35, 249-255. Rogers, j. D. (1968 b). Nuclear phenomena in the ascospores of Hypoxylon punctulatum. Canadian Journal of Botany 46, 865-866. Rogers, j. D. (1968 c). Xylaria curta: cytology of the ascus. Canadian Journal of Botany 46, 1337-1340. Rogers, j. D. (1968d). Hypoxylon fuscum: a review of the fungus and its relationship with Alnus in the Northwest. In Biology of Alder (ed. j. M. Trappe, j. F. Franklin, R. F. Tarrant & G. M. Hansen), pp. 251-258. Portland Press: Oregon, U.S.A. Rogers, j. D. (1969). Hypoxylon rubiginosum: cytology of the ascus and surface morphology of ascospores. Mycopathologia et Mycologia Applicata 38, 215-233. Rogers, j. D. (1970). Cytology of Poronia oedipus and Poronia punctata. Canadian Journal of Botany 48, 1665-1668. 920 Rogers, j. D. (1972). Hypoxylon cohaerens: cytology of the ascus. Mycopathologia et Mycologia Applicata 48, 161-165. Rogers, j. D. (I973). Cytology of Podosordaria leporina. Canadian Journal of Botany 51, 791-793. Rogers, j. D. (1975 a). Hypoxylon serpens: cytology and taxonomic considerations. Canadian Journal of Botany 53, 52-55. Rogers, j. D. (1975 b). Xylaria polymorpha II. Cytology of a form with typical robust stromata. Canadian Journal of Botany 53, 1736-1743. Rogers, j. D. (1975 c). Nummularia broomeiana: conidial state and taxonomic aspects. American Journal of Botany 62, 761-764. Rogers, j. D. (1977 a). A new Hypoxylon species with appendaged, ornamented ascospores. Canadian Journal of Botany 55, 2394-2398. Rogers, j. D. (1977 b). Surface features of the light colored ascospores of some applanate Hypoxylon species. Canadian Journal of Botany 55, 2394-2398. Rogers, j. D. (1979a). The Xylariaceae: Systematic, biological and evolutionary aspects. Mycologia 71, 1-42. Rogers, j. D. (1979b). Xylaria magnoliae sp. nov. and comments on several other fruit-inhabiting species. Canadian Journal of Botany 57, 941-945. Rogers, j. D. (1982). Entonaema liquescens: description of the anamorph and thoughts on its systematic position. Mycotaxon 23, 429-437. Rogers, j. D. (1983). Xylaria bulbosa, Xylaria curta, and Xylaria longipes in Continental United States. Mycologia 75, 457-467. Rogers, j. D. (1984a). Xylaria acuta, Xylaria cornu-damae, and Xylaria mali in Continental United States. Mycologia 76, 23-33. Rogers, j. D. (1984 b). Xylaria cubensis and its anamorph Xylocoremium /labelliforme, Xylaria allantoidea and Xylaria poitei in Continental United States. Mycologia 76, 912-923. Rogers, j. D. (1985). Anamorphs of Xylaria: taxonomic considerations. Sydowia 38, 255-262. Rogers, j. D. (1993). Teleomorph, anamorph, and holomorph considerations in the Xylariaceae. In The Fungal Holomorph: Mitotic, Meiotic and Pleomorphic Speciation in Fungal Systematics (ed. D. K Reynolds & j. W. Taylor), pp. 179-182. CA.B. International: Wallingford, UK Rogers, j. D. (1994). Problem genera and family interfaces in the eupyrenomycetes. In Ascomycete Systematics: Problems and Perspectives in the Nineties (ed. D. L. Hawksworth), pp. 321-331. Plenum Press: New York. Rogers, j. D. & Berbee, j. G. (1964). Developmental morphology of Hypoxylon pruinatum in bark of quaking aspen. Phytopathology 54, 154-162. Rogers, j. D. & Callan, B. E. (1986a). Xylaria poitei: stromata, cultural description, and structure of conidia and ascospores. Mycotaxon 26, 287-296. Rogers, j. D. & Callan, B. E. (1986 b). Xylaria polymorpha and its allies in Continental United States. Mycologia 78, 391-400. Rogers, j. D., Callan, B. E., Rossman, A. Y. & Samuels, G. j. (1988). Xylaria (Sphaeriales, Xylariaceae) from Cerro de la Neblina, Venezuela. Mycotaxon 31, 103-153. Rogers, j. D .. Callan, B. E. & Samuels, G. j. (1987). The Xylariaceae of the rain forests of North Sulawesi (Indonesia). Mycotaxon 31, 113-172. Rogers, j. D. & Candoussau, F. (I982). Hypoxylon gillesii, a new species with ornamented ascospores from Madagascar. Mycotaxon 15, 507-514. Rogers, j. D. & Lress0e, T. (I992). Podosordaria ingii sp. nov. and its Lindquistia anamorph. Mycotaxon 44, 435-443. Rogers, j. D. & Russell, D. W. (I973). Conidial state of Poronia punctata. Canadian Journal of Botany 51, 481-484. Rogers, j. D. & Stiers, D. L. (1974). Cytology of Rosellinia mammiformis and R. aquila. Canadian Journal of Botany 52, 5-10. Rogers, j. D., Stone, j. & ju, Y.-M. (1994). Anthostomella formosa var. abietis var. nov. and notes on Apiorhynchostoma. Mycologia 86, 700-703. Rogers, j. D. & Whalley, A. j. S. (1978). A new Hypoxylon species from Wales. Canadian Journal of Botany 56, 1346-1348. Rohrmann, G. F. & Rossman, A. Y. (I980). Nutritional strategies of Macrotermes ukuzii (Isoptera: Termitidae). Pedobiologia 20, 61-73. Salisbury, P. j. & Long, j. R. (1956). A new needle blight of Douglas fir seedlings caused by Rosellinia herpotrichioides Hepting & Davidson. Proceedings of the Canadian Phytopathological Society 23, 19. Samuels, G. j. (1989). Thuemenella cubispora, a xylariaceous ascomycete and its biogeography. Mycological Society of America Newsletter 40, 46. Samuels, G. j. & Muller, E. (1980). Life history studies on Brazilian ascomycetes. 8. Thamnomyces chordalis (anam.: Nodulisporium) and Camillea bacillum A. J. S. Whalley (anam.: Geniculosporiurn) with notes on taxonomy of the Xylariaceae. sydowia 33, 274-281. Samuels, G. j., Muller, E. 111; Petrini, O. (1987). Studies in the Amphisphaeriaceae (sensu lato) 3. New species of Monographella and Pestalosphaeria and two new genera. Mycotaxon 28, 473-499. Samuels, G.).. Rogers, j. D. & Nagasawa, E. (1987). Studies in the Amphisphaeriaceae (sensu lata). I. Collodiscula japonica and its anamorph, Acanthodochiurn collodisculae. Mycotaxon 28, 453-459. Samuels, G. j. & Rossman, A. Y (1992). 1Iluernenella and sarawakus. Mycologia 84,26-40. Sands, W. A. (1969). The association of termites and fungi. In The Biology of Termites (ed. K. Krishna & F. Weesner), pp. 495-524. Academic Press: New York and London. San Martin, F. & Rogers, j. D. (1989). A preliminary account of Xylaria of Mexico. Mycotaxon 34, 283-373. Sawai, K., Okuno, T. & Ito, T. (1982). The toxicity of cytochalasin E on plants. Annales Phytopathological Society of japan 48, 529-531. Schipper, A. L., Jr. (1978). A Hypoxylon mammaturn pathotoxin responsible for canker formation in quaking aspen. Phytopathology 68, 866-872. Shea, K. R. (1964). Rosellinia herpotrichioides on Sitka spruce seedlings in Washington. Plant Disease Reporter 48, 512-513. Siever-Canavesi, F. & Sieber, T. N. (1987). Endophytiscche Pilze in der Tanne (Abies alba Mill.) - Vergleich zweier Standorte im Schweizer Mittelland (Naturwald-Aufforstung). Sydowia 40, 250--273. Sivanesan, A. & Holliday, P. (1972 a). Rosellinia bunodes. CM.1. Descriptions of Pathogenic Fungi and Bacteria. No. 351. Sivanesan, A. & Holliday, P. (1972b). Rosellinia necamx. CM.1. Descriptions of Pathogenic Fungi and Bacteria. No. 352. Sivanesan, A. & Holliday, P. (1972c). Xylaria polymorpha. CM.!. Descriptions of Pathogenic Fungi and Bacteria. No. 354. Smith, R. S. (1966). Rosellinia needleblight of Douglas fir (Pseudotsuga menziesii) in California. Plant Disease Reporter 50, 249--250. Spooner, B. M. (1986). New or rare British microfungi from Esher Common, Surrey. Transactions of the British Mycological Society 86, 401-408. Soderquist, C ]. (1973). juglone and allelopathy. journal of Chemical Education 50, 782-783. Stermer, B. A., Scheffer, R. P. & Hart, j. H. (1984). Isolation of toxins of Hypoxylon mammatum and demonstration of some toxin effects on selected clones of Populus tremuloides. Phytopathology 74, 654--658. Stiers, D. L. (1977). Fine structure of the ascus apex in Hypoxylon serpens, Poronia punctata, Rosellinia aquila, and R. mammiformis. Cytologia 42, 697-702. Stiers, D. L., Rogers, ]. D. & Russell, D. W. (1973). Conidial state of Poronia punctata. Canadian journal of Botany 51, 481-484. Subramanian, C V. & Chandrashekara, K. V. (1977). Lindquistia, a new hyphomycete genus. Boletin de la Sociedad Argentina de Botanica 18, 145-151. Sutherland, j. B. & Crawford, D. L. (1981). Lignin and glucan degradation by species of Xylariaceae. Transactions of the British Mycological Society 76, 335-337. Sztejnberg, A., Freeman,S., Chet, I. & Katan, j. (1987). Control of Rosellinia necatrix in soil and in apple orchard by solarization and Trichoderma harzianum. Plant Disease 71, 365-369. Sztejnberg, A. & Madar, Z. (1980). Host range of Dematophora necatrix, the cause of white root rot disease in fruit trees. Plant Disease 64, 662--664. Tainter, F. H., Williams, T. M. & Cody,). B. (1983). Drought as a cause of oak decline and death on the South Carolina coast. Plant Disease 67,195-197. Teixeira de Sousa, A. j. (1985). Lutte contre Rosellinia necatrix (Hartig) Berlese, agent du pourridie laineux: sensibilite de quelques especes vegetales et lutte chimique. European journal of Forest Pathology 15, 323-332. Teixeira de Sousa, A. j., Guillaumin, j. j., Sharples, G. P. & Whalley, A. j. S. (1995). Rosellinia necatrix and white root rot of fruit trees and other plants in Portugal and nearby regions. Mycologist 9, 31-33. Teixeira de Sousa, A. j. & Whalley, A. j. S. (1991). Induction of mature stromata in Rosellinia necatrix and its taxonomic implications. svdowia 43, 281-290. Thompson, G. E. (1963). Decay of oaks caused by Hypoxylon atropunctatum. Plant Disease Reporter 47, 202-205. Udagawa, S.-1. & Ueda, S. (1988). Calceomyces, a new genus of the Xylariaceae with shoe-shaped ascospores. Mycotaxon 32. 447-455. 921 Valldosera, P. & Guarro, G. (1988). Some coprophilous ascomycetes from Chile. Transactions of the British Mycological Society 90, 601--605. Van Arsdel. E. P. (1972). Some cankers on oaks in Texas. Plant Disease Reporter 56, 300--304. Van der Gucht. K. (1992). Contribution towards a revision of the genera Camillea and Biscogniauxia (Xylariaceae, Ascomycetes) from Papua New Guinea. Mycotaxon 45, 259-273. Van der Gucht. K. (1993). Spore ornamentation makes a nice difference: Daldinia eschscholzii and Daldinia concentrica. In Aspects of Tropical Mycology (ed. S. Isaac, j. C Frankland, R. Watling & A. j. S. Whalley), pp. 309--310. Cambridge University Press: Cambridge, U.K. Van der Gucht, K. & Van der Veken, P. (1992). Contribution towards a revision of the genus Hypoxylon s. str. (Xylariaceae, Ascomycetes) from Papua New Guinea. Mycotaxon 44, 275-299. Van der Gucht. K. & Whalley, A. j. S. (1996). Notes on the ecology of the Xylariaceae of Papua New Guinea. sydowia, in press. Vannin;' A. & Scarascia Mugnozza, G. (1991). Water stress: A predisposing factor in the pathogenesis of Hypoxylon mediterraneum on Quercus cems. European journal of Forest Pathology 21, 193-201. Varghese, G. (1971). Infection of Hevea brasiliensis by Ustulina zonata (Lev.) Sacco journal of the Rubber Research Institute of Malaya 23, 157-163. Vegh, I. (1984). Identification en France de Rosellinia herpotrichioides Hepting et Davidson sur Picea exce/sa Link ornamental. Revue Horticole 247, 35-37. Watling, R. (1978). The distribution of larger fungi in Yorkshire. Naturalist 103, 39--57. Watling, R. & Whalley, A. j. S. (1977). Provisional Maps of Xylariaceous Fungi. Biological Records Centre: Monks Wood. Wei, D.-L., Chang, Y-H., Lin, Y-W., Chuang, C-L. & jong, S.-C (1992). Production of cellulolytic enzymes from the Xylaria and Hypoxylon species of Xylariaceae. World journal of Microbiology and Biotechnology 8, 141-146. Weidel!. D. j. (1942). Damage to catalpa due to recreational use. journal of Forestry 40, 807. Whalley, A. J. S. (1976). Notes on the conidial state of Hypoxylon udum. Transactions of the British Mycological Society 67, 515-517. Whalley, A. j. S. (1985). The Xylariaceae: some ecological considerations. sydowia 38, 369-382. Whalley, A. j. S. (1987). Xylaria inhabiting fallen fruits. Agarica 8, 6&-72. Whalley, A. J. S. (1993). Tropical Xylariaceae: their distribution and ecological characteristics. In Aspects of Tropical Myc%gy (ed. S. Isaac, j. C Frankland, R. Watling & A. j. S. Whalley), pp. 103-II9. Cambridge University Press: Cambridge, UK Whalley, A. j. S. (1996). Biscogniauxia, Camillea and Hypoxylon (ascomycetes) in South East Asia. In Tropical Mycology (ed. K. K. janardhanan, D. L. Hawksworth, C Rajendran & K. Natarajan), in press. Oxford & IBH Publishing Co.: New Delhi, India. Whalley, A.]. S. & Edwards, R. L. (1985). Nummulariella marginata: its conidial state, secondary metabolites and taxonomic relationships. Transactions of the British Mycological Society 85, 385-390. Whalley, A. j. S. & Edwards, R. L. (1987). Xylariaceous fungi: use of secondary metabolites. In The Evolutionary Biology of Fungi (ed. A. D. M. Rayner, C M. Brasier & D. Moore), pp. 423-434. Cambridge University Press: Cambridge, U.K. Whalley, A. j. S. & Edwards, R. L. (1995). Secondary metabolites and systematic arrangement within the Xylariaceae. Canadian journal of Botany 73 (Supplement I), S802-S81O. Whalley, A.]. S. & Greenhalgh, G. N. (1973). Numerical taxonomy of Hypoxylon. 1. Comparison of classification of the cultural and perfect states. Transactions of the British Mycological societv 61, 435-454. Whalley, A. j. S. & Hammelev, D. (1988). Some Xylariaceae from Georgia and the Russian Soviet Federative Socialist Republic. Agarica 9, 67-70. Whalley, A. j. S., Hywel-jones. N. L., jones, E. B. G. & Whalley, M. A. (1995). A preliminary account of the genera Biscogniauxia and HypoxVlon in the Chanthaburi and Chon Buri Provinces of South East Thailand. sydowia 47, 70--81. Whalley, A. j. S., jones, E. B. G. & Alias, S. A. (1994). The Xylariaceae (ascomycetes) of mangroves in Malaysia and South East Asia. Nova Hedwigia 59, 207-218. Whalley, A. J. S. & Knudsen, H. (1985). HVpoxy/on vogesiacum var. macrospora, a new record for Greenland, Transactions of the British Mycological Society 84,563. The xylariaceous way of life Whalley, A j. S., Lressoe, T. & Kile, G. A (1990a). A new species of Biscogniauxia with appendaged ascospores from Tasmania. Mycological Research 94, 27-239. Whalley, A j. S., Lressoe, T. & Kile, G. A (1990b). Validation of Biscogniauxia Sect. Appendiculata. Mycological Research 94, 864. Whalley, A j. S. & Petrini, L. E. (1984). Hypoxylon vogesiacum var. macrospora from Europe. Transactions of the British Mycological Society 83, 364-368. Whalley, A j. S. & Rogers, j. D. (1980). The geniculate conidial state of Hypoxylon chestersii. Transactions of the British Mycological Society 74, 439-440. Whalley, A j. S. & Thienhirun, S. (1996). Rhopalostroma kanyae sp. nov. from Thailand. Mycological Research 100, in press. Whalley, A j. S. & Watling, R. (1980a). Hypoxylon udum Pers. ex Fr. Transactions of the Botanical Society of Edinburgh 43, 217-220. Whalley, A j. S. & Watling, R. (1980 b). Daldinia concentrica versus Daldinia vernicosa. Transactions of the British Mycological Society 74, 399-406. Whalley, A j. S. & Watling, R. (1982). Distribution of Daldinia concentrica in the British Isles. Transactions of the British Mycological Society 78, 47-53. Whalley, A j. S. & Whalley, M. A (1977). Stromal pigments and taxonomy of Hypoxylon. Mycopathologia 61, 99-103. (Accepted 8 March 1996) 922 Whalley, M. A (1995). Camilleafusiformis sp. nov. from Ecuador. Sydowia 47, 82-88. Whalley, M. A, Whalley, A j. S. & jones, E. B. G. (1996). Camilleaselangorensis sp. nov. from Malaysia. Sydowia 48, 145-151. Wicklow, D. T. & Hirschfield, B. j. (1979). Evidence of a competitive hierarchy among coprophilous fungal populations. Canadian Journal of Microbiology 25, 855-858. Winter, G. (1887). Die Pilze Deutschlands, Osterreichs und der Schweiz. 2. A6. Ascomyceten, Gymnoasceen und Pyrenomyceten. In Rabenhorst's Kryptogramen Flora, 2, Auf!, 1, Pilze. Leipzig. Wilkins, W. H. (1934). Studies in the genus Ustulina with special reference to parasitism. I. Introduction, survey of previous literature and host index. Transactions of the British Mycological Society 18, 320-346. Wilkins, W. H. (1943). Studies in the genus Ustulina with special reference to parasitism. VI. A brief account of heart rot of beech (Fagus sylvatica L.) caused by Ustulina. Transactions of the British Mycological Society 26, 169-170. Yokoyama, K. & Shidei, Y. (1972). Two species of fungi on buried seeds of Fagus crenata Blume. Transactions of the Mycological Society of Japan 13, 149-152.

RELATED PAPERS

RELATED TOPICS

Log In

The xylariaceous way of life

The xylariaceous way of life

Related Papers

RELATED PAPERS

RELATED TOPICS