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What is Fumago vagans?
[ 371
]
Trans. Brit. mycol. Soc. 48 (3), 37 1-375 (1965) Printed in Great Britain
WHAT IS FUMAGO VAGANS? By R.
J.
FRIEND
Department of Botany, University of Exeter, Devon The literature concerning Fumago, especially as it involves the hypothesis of the polymorphism of sooty moulds and the relation of Fumago to Capnodium, is briefly discussed. The relevant results of a cultural investigation ofsooty mould on Tilia, and of an examination of herbarium specimens labelled Fumago are recorded. The conclusion is reached that the genus Fumago is invalid on the grounds that presumed type material of the lectotype species, F. vagans, consists of two entirely discordant elements.
Fungi in the genus Fumago Pers., and particularly the lectotype species F. vagans Pers., have been commonly mentioned in the literature concerning sooty moulds ever since the genus was first described (Persoon, 1822). In brief, Fumago was said to have dark, septate hyphae forming muriform bodies in chains and upright conidiophores bearing Cladosporium-like conidia. It has been commonly reported that it forms a black, superficial growth (sooty mould) on various parts of many different living plants and occasionally its growth on inanimate objects has been noted. It is usually put in the dictyosporous Dematiaceae near Alternaria on the basis of the muriform bodies ascribed to it (Clements & Shear, 1931), though Saccardo (I886) and Ferraris (1910) considered it allied to Cladosporium, Ferraris even classifying it in the Phaeodidymae. There are many references in the literature to a definite or to a probable connexion between Fumago and certain Capnodiaceae (especially Capnodium), though Per soon himself did not suggest any such connexion in his original diagnosis. The writer, while carrying out investigations on sooty mould on lime trees (Tilia x vulgaris) in Exeter, also studied the position of Fumago because sooty mould on limes has commonly been ascribed to Fumago spp., and especially to F. vagans. REVIEW OF LITERATURE
Polymorphism
if soory moulds
Acceptance of the hypothesis that sooty moulds are essentially polymorphic had for many years a considerable influence on the way in which many mycologists thought about Fumago. They seem to have regarded sooty mould on anyone host species as consisting of only one fungus, so that when a number of different fungus fruiting bodies were found in one specimen of sooty mould, these were all considered to be different forms of a single polymorphic fungus rather than evidence for the presence of a number of different fungi. Some selected publications in which the polymorphic view of sooty moulds has been maintained are: Tulasne &
372
Transactions British Mycological Society
Tulasne (1863), Rabenhorst (1887, 1910), Gaumann (1928) and Butler (1918). The first three of these publications ascribed the sooty mould either to F. vagans or to a synonym of F. vagans. The composite nature of sooty mould was first unequivocally demonstrated by Neger (1918), who carried out detailed cultural work, isolating over a dozen different fungi. Since his time many other mycologists have carried out similar investigations with comparable results. Fraser (1933), who worked for a number ofyears on sooty moulds in Australia, stated that she obtained no cultural evidence that any member of the Capnodiaceae had an imperfect state in the Moniliales, though she did find that certain of them could have a pycnidial as well as a perithecial state. Batista & Ciferri (1963) have also connected with perithecial states certain fungi in the Sphaeropsidales but not any in the Moniliales. Now that the hypothesis maintaining the polymorphism ofsooty moulds has been discredited, and their composite nature established beyond any doubt, the place of Fumago in sooty moulds needs to be reassessed.
The relation oj Fumago to Capnodium The numerous references in the literature to species of Fumago (including F. vagans) as the imperfect states of species of Capnodium and other genera of the Capnodiaceae, apparently originated from Montagne (1849) who described F. vagans as having an ascigerous state which he named Capnodium salicinum Mont. The grounds on which he connected what is classified as a dematiaceous hyphomycete with an ascomycete seem to be only that the characteristic structures of each were found in close proximity in a single growth of sooty mould. This assumption made by Montagne was widely accepted until early this century and has even been perpetuated to the present day, though more recent authors usually consider the connexion as probable rather than definite (e.g, Bender, 1931; Barnett, 1960). No evidence from cultural experiments has ever been quoted in support of this supposed connexion. As yet there is no sound evidence for connecting any member of the Capnodiaceae with any hyphomycete, and Fumago as described by Persoon, must be classified as a hyphomycete. The obvious conclusion is that the supposed connexion between Fumago and Capnodium arose from the false premise that sooty moulds are polymorphic, and therefore can no longer be reasonably maintained. Fumago in descriptions oj sooty moulds The records and descriptions of Fumago in the literature have been, and still are, extremely confused and confusing, and little would be gained by giving a detailed review of the publications involved. The most commonly recurring features in such descriptions are: (i) the presence of dark, muriform structures; (ii) the presence of Cladosporium-like conidia; (iii) the statement that it is a very variable fungus; (iv) the fact that it is very common and widespread, occurring on many different kinds of living plants. It seems obvious that very often the name Fumago (especially F. vagans) has been used rather loosely for any superficial, dark, fungus growth on living plants where perithecia cannot be found.
Fumago vagans. R. J. Friend
373
It is considered highly significant that, in spite of all the cultural investigations carried out on sooty moulds, no fungus which exhibits the main characters of Fumago as described by Persoon has yet been isolated in pure culture. All the fungi that have been isolated have been referred to other genera which almost invariably fall into the Capnodiaceae, Sphaeropsidales, or Moniliales, with those in the last group being by far the most common in sooty mould on plants in temperate climates, and often quite common in sooty mould on plants in tropical regions. CULTURAL INVESTIGATION OF SOOTY MOULD ON TILIA
Isolations of the components of sooty mould on lime leaves were attempted and the results (Friend, 1965) indicated that Aureobasidium pullulans (de Bary) Am. (= Pullularia pullulans (de Bary) Berkh.), and Cladosporium herbarum (Pers.) Link ex Fr. were the fungi most commonly present, making up the bulk of the growth of sooty mould. A number of other common saprophytic hyphomycetes were also isolated but they are considered to be of only minor components of sooty mould on lime. No fungus was isolated that could be referred to either Fumago or Capnodium. EXAMINATION OF HERBARIUM MATERIAL
Selected specimens of a variety of plants bearing sooty mould in the herbarium of the Royal Botanic Gardens, Kew, were examined. They included two Tilia leaves bearing a heavy deposit of epiphyllous sooty mould and labelled in the handwriting of Persoon, 'Fumago Joliorum', a name later regarded as a synonym of F. vagans. Three other Tilia leaves bearing a similar growth of sooty mould, labelled F. vagans and dated October 1819, are also probably authentic Persoon material. Microscopical examination of these specimens indicated that the sooty mould was very similar to that prevalent on lime leaves in Exeter some 144 years later. Large numbers of conidia very similar to those of Cladosporium herbarum were present and also many dark, muriform bodies that may well be referable to Aureobasidium pullulans which can form such structures (Hoggan, 1924). Though not designated' type' material, these old specimens are considered important because they certainly indicate the sort of fungus growth on which Persoon based his description of F. vagans. Other specimens labelled as various species of Fumago (or synonyms of Fumago) usually bore muriform structures, probably of A. pullulans, and also conidia referable to C. herharum. A specimen labelled F. rivieriana on myrtle leaves, dated 1865, had abundant conidia of Cladosporium, whereas another of F. rivieriana on Camellia japonica, dated January 1883, had no Cladosporium conidia. A specimen labelled F. humuli on Humulus, dated 1879, bore both conidia of Cladosporium and muriform structures (? A. pullulans). On a specimen labelled CladosporiumJumago Link ( =F. vagans) on a twig of? Salix sp., dated summer 1854, there were abundant muriform structures, but no conidia of Cladosporium. Selected specimens ofsooty mould on a variety of plants, in the herbarium of the Commonwealth Mycological Institute, Kew, were also examined, 24
~yc.
48
374
Transactions British Mycological Society
including ten labelled F. vagans. Conidia of Cladosporium were frequently found but were absent from some specimens, and muriform structures (? A. pullulans) were present in most, though not all. On one, labelled 'Changa manga' from Pakistan, dated April 1961 (IMI 87063), conidia of Cladosporium were numerous. On a specimen of Prunus domestica from Essex, England, dated September 1947 (IMI 17480), and on Tilia sp. from Brno, Czechoslovakia, dated November 1926 (IMI 10423), both conidia of Cladosporium and muriform structures (?A. pullulans) were present. On a specimen ofPolygonatum multijlorum from Hampshire, England, dated August 1954 (IMI 57709), on a specimen of Cydonia vulgaris from Brno, dated May 1924 (IMI 10424), and on a specimen of Asarum europaeum marked 'from F. Petrak, Teplitz, Bohemia', dated August 1897 (IMI 16523), the sooty mould was probably predominantly A. pullulans. On a specimen of Salixcordata from Ottawa, Canada, dated September 1944 (IMI 25939), only dark, septate hyphae were present with no distinct spore forms. The conclusions from this examination could well be summarized as follows: Sooty moulds, hitherto referred to as Fumago tiagans or closely related species, are very variable, common and widely distributed, occurring on many different types of living plants, and often, but not always, having Cladosporium-like conidia and dark, muriform bodies. This summary has precisely the same content as that used earlier to summarize the most common features in published descriptions of Fumago spp. DISCUSSION
Both Arnaud (1910) and Neger (1918) reported that the fungi of greatest predominance in sooty moulds in Europe were A. pullulans and C. herbarum. It is considered that sound evidence in support of their findings is provided by the cultural investigation of sooty mould on lime (Friend, 1965) and the examination of herbarium material. Because F. vagans was formerly said to be such a common cause of sooty mould, one obvious question which follows is, What, then, is Fumago vagans? Nearly 40 years ago Woronichin (1926) considered F. oagans, as generally known, to be a mixture of A. pullulans and C. herbarum, and the evidence presented in this paper is considered to verify his contention. In conclusion, it is maintained that the name Fumago vagans Pers. must be rejected because it is based on type material consisting of two entirely discordant elements (Lanjouw et al. 1956). Clements & Shear (1931) selected F. vagans as the lectotype of the genus, and therefore Fumago Pers. must also be rejected. This paper is based on part of an investigation carried out in fulfilment of the degree of M.Sc. in the University of Exeter. The writer gratefully acknowledges the assistance rendered by the Royal Botanic Gardens, Kew, and the Commonwealth Mycological Institute, Kew, in making available for examination relevant herbarium material, and also by Dr S. A. J. Tarr of the University of Exeter for his helpful advice and criticism.
Fumago vagans. R. J. Friend
375
REFERENCES
*ARNAUD, G. (1910). Contribution a l'etude des Fumagines. Annls 11J1col. 7, 470-476. BARNETT, H. L. (1960). Illustrated genera of imperfect fungi, pp. 124, 125. Minneapolis: Burgess Publishing, Co. BATISTA, A. C. & CIFERRI, R. (1963). The sooty moulds of the family Asbolisiaceae. Quad. 1st. bot. Pavia, p. 31. (Publ. 1st. Micol. Univ. Recife 163.) BENDER, H. B. (1931). Genera of fungi imperfecti: North American species and hosts. Ph.D. Dissertation, Yale, p. 567. BUTLER, E. ]. (19 I8). Fungi and disease in plants, pp. 30, 3 1; 4 1I, 412; 461-463; 487-489. Calcutta and Simla: Thacker, Spink and Co. CLEMENTS, F. E. & SHEAR, C. L. (1931). Thegeneraoffungi, p. 217. New York: Wilson. FERRARIS, T. (1910-14). Flora Italica cryptogama-i-Hyphales, p. 358. Societa Botanica Italiana. FRASER, L. (1933). An investigation of the sooty moulds of N.S.W. 1. Proc, Linn. Soc. NS.W. 58, 375-395· FRIEND, R.]. (1965). A study of sooty mould on lime trees (Tilia x vulgaris). Trans. Br. mycol. Soc. 48,367-37°. GXUMANN, E. A. (1928). Comparative morphology of the fungi, p. 265. (Translated by C. W. Dodge.) New York. HOGGAN, I. A. (1924). On Dematium pullulans de Bary. Trans. Br. mycol. Soc. 9, 100-107. LANJOUW,]. et al. (1956). International Code of Botanical Nomenclature, p. 43, Article 66. Utrecht, Netherlands. *MONTAGNE,]. P.F. C. (1849)' De Capnodio, nov.gen. Annls Sci. nat. Bot. (Ser. 3), 11,233. *NEGER, F. W. (1918). Experimentelle Untersuchungen tiber Russtaupilze. Flora, N.F. 10,67- 139. PERSOON, C. H. (1822). Mycologia europea. 1, p. 9. RABENHORST, L. (1887). Kryptogamen-Flora. 2, 74-77. Pilze, RABENHORST, C. H. (1910). Kryptogamen-Flora, 9, 266-269. Pilze. SACCARDO, P. A. (1886). Sylloge Fungorum. IV, p. 547. TULASNE, L. R. & TULASNE, C. (1863). Selecta fungorum carpologia. II, pp. 279-284. Paris: Imperial Press. (Translated by W. B. Grove, Oxford: Clarendon Press, 1931.) *WORONICHIN, N. N. (1926). Zur Kenntnis der Morphologie und Systematik der Russtaupilze Transkaukasiens. Annls mycol. 24, 231-265. (* Abstracts only available to the writer.)
(Accepted for publication 9 November 1964)
]
Trans. Brit. mycol. Soc. 48 (3), 37 1-375 (1965) Printed in Great Britain
WHAT IS FUMAGO VAGANS? By R.
J.
FRIEND
Department of Botany, University of Exeter, Devon The literature concerning Fumago, especially as it involves the hypothesis of the polymorphism of sooty moulds and the relation of Fumago to Capnodium, is briefly discussed. The relevant results of a cultural investigation ofsooty mould on Tilia, and of an examination of herbarium specimens labelled Fumago are recorded. The conclusion is reached that the genus Fumago is invalid on the grounds that presumed type material of the lectotype species, F. vagans, consists of two entirely discordant elements.
Fungi in the genus Fumago Pers., and particularly the lectotype species F. vagans Pers., have been commonly mentioned in the literature concerning sooty moulds ever since the genus was first described (Persoon, 1822). In brief, Fumago was said to have dark, septate hyphae forming muriform bodies in chains and upright conidiophores bearing Cladosporium-like conidia. It has been commonly reported that it forms a black, superficial growth (sooty mould) on various parts of many different living plants and occasionally its growth on inanimate objects has been noted. It is usually put in the dictyosporous Dematiaceae near Alternaria on the basis of the muriform bodies ascribed to it (Clements & Shear, 1931), though Saccardo (I886) and Ferraris (1910) considered it allied to Cladosporium, Ferraris even classifying it in the Phaeodidymae. There are many references in the literature to a definite or to a probable connexion between Fumago and certain Capnodiaceae (especially Capnodium), though Per soon himself did not suggest any such connexion in his original diagnosis. The writer, while carrying out investigations on sooty mould on lime trees (Tilia x vulgaris) in Exeter, also studied the position of Fumago because sooty mould on limes has commonly been ascribed to Fumago spp., and especially to F. vagans. REVIEW OF LITERATURE
Polymorphism
if soory moulds
Acceptance of the hypothesis that sooty moulds are essentially polymorphic had for many years a considerable influence on the way in which many mycologists thought about Fumago. They seem to have regarded sooty mould on anyone host species as consisting of only one fungus, so that when a number of different fungus fruiting bodies were found in one specimen of sooty mould, these were all considered to be different forms of a single polymorphic fungus rather than evidence for the presence of a number of different fungi. Some selected publications in which the polymorphic view of sooty moulds has been maintained are: Tulasne &
372
Transactions British Mycological Society
Tulasne (1863), Rabenhorst (1887, 1910), Gaumann (1928) and Butler (1918). The first three of these publications ascribed the sooty mould either to F. vagans or to a synonym of F. vagans. The composite nature of sooty mould was first unequivocally demonstrated by Neger (1918), who carried out detailed cultural work, isolating over a dozen different fungi. Since his time many other mycologists have carried out similar investigations with comparable results. Fraser (1933), who worked for a number ofyears on sooty moulds in Australia, stated that she obtained no cultural evidence that any member of the Capnodiaceae had an imperfect state in the Moniliales, though she did find that certain of them could have a pycnidial as well as a perithecial state. Batista & Ciferri (1963) have also connected with perithecial states certain fungi in the Sphaeropsidales but not any in the Moniliales. Now that the hypothesis maintaining the polymorphism ofsooty moulds has been discredited, and their composite nature established beyond any doubt, the place of Fumago in sooty moulds needs to be reassessed.
The relation oj Fumago to Capnodium The numerous references in the literature to species of Fumago (including F. vagans) as the imperfect states of species of Capnodium and other genera of the Capnodiaceae, apparently originated from Montagne (1849) who described F. vagans as having an ascigerous state which he named Capnodium salicinum Mont. The grounds on which he connected what is classified as a dematiaceous hyphomycete with an ascomycete seem to be only that the characteristic structures of each were found in close proximity in a single growth of sooty mould. This assumption made by Montagne was widely accepted until early this century and has even been perpetuated to the present day, though more recent authors usually consider the connexion as probable rather than definite (e.g, Bender, 1931; Barnett, 1960). No evidence from cultural experiments has ever been quoted in support of this supposed connexion. As yet there is no sound evidence for connecting any member of the Capnodiaceae with any hyphomycete, and Fumago as described by Persoon, must be classified as a hyphomycete. The obvious conclusion is that the supposed connexion between Fumago and Capnodium arose from the false premise that sooty moulds are polymorphic, and therefore can no longer be reasonably maintained. Fumago in descriptions oj sooty moulds The records and descriptions of Fumago in the literature have been, and still are, extremely confused and confusing, and little would be gained by giving a detailed review of the publications involved. The most commonly recurring features in such descriptions are: (i) the presence of dark, muriform structures; (ii) the presence of Cladosporium-like conidia; (iii) the statement that it is a very variable fungus; (iv) the fact that it is very common and widespread, occurring on many different kinds of living plants. It seems obvious that very often the name Fumago (especially F. vagans) has been used rather loosely for any superficial, dark, fungus growth on living plants where perithecia cannot be found.
Fumago vagans. R. J. Friend
373
It is considered highly significant that, in spite of all the cultural investigations carried out on sooty moulds, no fungus which exhibits the main characters of Fumago as described by Persoon has yet been isolated in pure culture. All the fungi that have been isolated have been referred to other genera which almost invariably fall into the Capnodiaceae, Sphaeropsidales, or Moniliales, with those in the last group being by far the most common in sooty mould on plants in temperate climates, and often quite common in sooty mould on plants in tropical regions. CULTURAL INVESTIGATION OF SOOTY MOULD ON TILIA
Isolations of the components of sooty mould on lime leaves were attempted and the results (Friend, 1965) indicated that Aureobasidium pullulans (de Bary) Am. (= Pullularia pullulans (de Bary) Berkh.), and Cladosporium herbarum (Pers.) Link ex Fr. were the fungi most commonly present, making up the bulk of the growth of sooty mould. A number of other common saprophytic hyphomycetes were also isolated but they are considered to be of only minor components of sooty mould on lime. No fungus was isolated that could be referred to either Fumago or Capnodium. EXAMINATION OF HERBARIUM MATERIAL
Selected specimens of a variety of plants bearing sooty mould in the herbarium of the Royal Botanic Gardens, Kew, were examined. They included two Tilia leaves bearing a heavy deposit of epiphyllous sooty mould and labelled in the handwriting of Persoon, 'Fumago Joliorum', a name later regarded as a synonym of F. vagans. Three other Tilia leaves bearing a similar growth of sooty mould, labelled F. vagans and dated October 1819, are also probably authentic Persoon material. Microscopical examination of these specimens indicated that the sooty mould was very similar to that prevalent on lime leaves in Exeter some 144 years later. Large numbers of conidia very similar to those of Cladosporium herbarum were present and also many dark, muriform bodies that may well be referable to Aureobasidium pullulans which can form such structures (Hoggan, 1924). Though not designated' type' material, these old specimens are considered important because they certainly indicate the sort of fungus growth on which Persoon based his description of F. vagans. Other specimens labelled as various species of Fumago (or synonyms of Fumago) usually bore muriform structures, probably of A. pullulans, and also conidia referable to C. herharum. A specimen labelled F. rivieriana on myrtle leaves, dated 1865, had abundant conidia of Cladosporium, whereas another of F. rivieriana on Camellia japonica, dated January 1883, had no Cladosporium conidia. A specimen labelled F. humuli on Humulus, dated 1879, bore both conidia of Cladosporium and muriform structures (? A. pullulans). On a specimen labelled CladosporiumJumago Link ( =F. vagans) on a twig of? Salix sp., dated summer 1854, there were abundant muriform structures, but no conidia of Cladosporium. Selected specimens ofsooty mould on a variety of plants, in the herbarium of the Commonwealth Mycological Institute, Kew, were also examined, 24
~yc.
48
374
Transactions British Mycological Society
including ten labelled F. vagans. Conidia of Cladosporium were frequently found but were absent from some specimens, and muriform structures (? A. pullulans) were present in most, though not all. On one, labelled 'Changa manga' from Pakistan, dated April 1961 (IMI 87063), conidia of Cladosporium were numerous. On a specimen of Prunus domestica from Essex, England, dated September 1947 (IMI 17480), and on Tilia sp. from Brno, Czechoslovakia, dated November 1926 (IMI 10423), both conidia of Cladosporium and muriform structures (?A. pullulans) were present. On a specimen ofPolygonatum multijlorum from Hampshire, England, dated August 1954 (IMI 57709), on a specimen of Cydonia vulgaris from Brno, dated May 1924 (IMI 10424), and on a specimen of Asarum europaeum marked 'from F. Petrak, Teplitz, Bohemia', dated August 1897 (IMI 16523), the sooty mould was probably predominantly A. pullulans. On a specimen of Salixcordata from Ottawa, Canada, dated September 1944 (IMI 25939), only dark, septate hyphae were present with no distinct spore forms. The conclusions from this examination could well be summarized as follows: Sooty moulds, hitherto referred to as Fumago tiagans or closely related species, are very variable, common and widely distributed, occurring on many different types of living plants, and often, but not always, having Cladosporium-like conidia and dark, muriform bodies. This summary has precisely the same content as that used earlier to summarize the most common features in published descriptions of Fumago spp. DISCUSSION
Both Arnaud (1910) and Neger (1918) reported that the fungi of greatest predominance in sooty moulds in Europe were A. pullulans and C. herbarum. It is considered that sound evidence in support of their findings is provided by the cultural investigation of sooty mould on lime (Friend, 1965) and the examination of herbarium material. Because F. vagans was formerly said to be such a common cause of sooty mould, one obvious question which follows is, What, then, is Fumago vagans? Nearly 40 years ago Woronichin (1926) considered F. oagans, as generally known, to be a mixture of A. pullulans and C. herbarum, and the evidence presented in this paper is considered to verify his contention. In conclusion, it is maintained that the name Fumago vagans Pers. must be rejected because it is based on type material consisting of two entirely discordant elements (Lanjouw et al. 1956). Clements & Shear (1931) selected F. vagans as the lectotype of the genus, and therefore Fumago Pers. must also be rejected. This paper is based on part of an investigation carried out in fulfilment of the degree of M.Sc. in the University of Exeter. The writer gratefully acknowledges the assistance rendered by the Royal Botanic Gardens, Kew, and the Commonwealth Mycological Institute, Kew, in making available for examination relevant herbarium material, and also by Dr S. A. J. Tarr of the University of Exeter for his helpful advice and criticism.
Fumago vagans. R. J. Friend
375
REFERENCES
*ARNAUD, G. (1910). Contribution a l'etude des Fumagines. Annls 11J1col. 7, 470-476. BARNETT, H. L. (1960). Illustrated genera of imperfect fungi, pp. 124, 125. Minneapolis: Burgess Publishing, Co. BATISTA, A. C. & CIFERRI, R. (1963). The sooty moulds of the family Asbolisiaceae. Quad. 1st. bot. Pavia, p. 31. (Publ. 1st. Micol. Univ. Recife 163.) BENDER, H. B. (1931). Genera of fungi imperfecti: North American species and hosts. Ph.D. Dissertation, Yale, p. 567. BUTLER, E. ]. (19 I8). Fungi and disease in plants, pp. 30, 3 1; 4 1I, 412; 461-463; 487-489. Calcutta and Simla: Thacker, Spink and Co. CLEMENTS, F. E. & SHEAR, C. L. (1931). Thegeneraoffungi, p. 217. New York: Wilson. FERRARIS, T. (1910-14). Flora Italica cryptogama-i-Hyphales, p. 358. Societa Botanica Italiana. FRASER, L. (1933). An investigation of the sooty moulds of N.S.W. 1. Proc, Linn. Soc. NS.W. 58, 375-395· FRIEND, R.]. (1965). A study of sooty mould on lime trees (Tilia x vulgaris). Trans. Br. mycol. Soc. 48,367-37°. GXUMANN, E. A. (1928). Comparative morphology of the fungi, p. 265. (Translated by C. W. Dodge.) New York. HOGGAN, I. A. (1924). On Dematium pullulans de Bary. Trans. Br. mycol. Soc. 9, 100-107. LANJOUW,]. et al. (1956). International Code of Botanical Nomenclature, p. 43, Article 66. Utrecht, Netherlands. *MONTAGNE,]. P.F. C. (1849)' De Capnodio, nov.gen. Annls Sci. nat. Bot. (Ser. 3), 11,233. *NEGER, F. W. (1918). Experimentelle Untersuchungen tiber Russtaupilze. Flora, N.F. 10,67- 139. PERSOON, C. H. (1822). Mycologia europea. 1, p. 9. RABENHORST, L. (1887). Kryptogamen-Flora. 2, 74-77. Pilze, RABENHORST, C. H. (1910). Kryptogamen-Flora, 9, 266-269. Pilze. SACCARDO, P. A. (1886). Sylloge Fungorum. IV, p. 547. TULASNE, L. R. & TULASNE, C. (1863). Selecta fungorum carpologia. II, pp. 279-284. Paris: Imperial Press. (Translated by W. B. Grove, Oxford: Clarendon Press, 1931.) *WORONICHIN, N. N. (1926). Zur Kenntnis der Morphologie und Systematik der Russtaupilze Transkaukasiens. Annls mycol. 24, 231-265. (* Abstracts only available to the writer.)
(Accepted for publication 9 November 1964)