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A new species of Cylindrocarpon associated with pine galls in Honduras
Notes and brief articles the former apex, giving the impression of branching. The section figured, of IMI 7255, is of a very young conidiophore fascicle which has not yet
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produced conidia. The conidiophores are often slightly curved towards the apex. The stroma, such as it is. is similar in older conidiophore fascicles.
A NEW SPECIES OF CYLINDROCARPON ASSOCIATED WITH PINE GALLS IN HONDURAS BY C. BOOTH AND H. C. EVANS
Commonwealth Mycological Institute, Ferry Lan e, K ew, U.K. Cylindrocarpon carneum sp .nov . is described from galls on Pinus maximinoi in Honduras and from culture. Its biology is briefly discussed. In the course of a disease survey of Central American pine species, galls of unknown etiology were encountered on high altitude pines in Honduras. Amongst the fungi associated with these galls was an undescribed species of the genus Cylindrocarpon Wollenw. Although pathogenic involvement has not been proven, its distinctive morphology is considered to warrant a formal description. Cylindrocarpon carneum sp .nov. (F igs 1,2) (E t yrn , - referring to the bright pink colour of the
spore tendrils) In hospitis. Stromata pulvinata innata cavita in galla. Conidiophora ramosa, hyalina, phialidibus clavatis vel su bulatis terminantia, 10-16 (- 20) X 3-5Itm . Macroconi dia cylindrica vel fusoidea , curvula, utrinque rotundat a, 5-8 septata, 42-50 x 7-8 pm, in cirrhi. In cultura. Coloniis salmoneis, farinosis ; rever so aurant iaco. M ycelium aer ium hyalinum, f1occosum. M icroconidia vecta palrnata con idiophora, hyalina ovara ad obovoidea, 7-9 x 4-5 Itm. Macrocon idia variabilia, ovata vel cylindrica, leniter curvata utrinque rotundata, 1-8 septata, 12-50 x 4-8 Itm . Chlamydosporae absentes. Teleomorphosis incognita. In galla Pini maximinoi, Pela Nariz, Cordillera Opalaca, Intibuca, Honduras, H. C. Evans, 17 Nov. 1980, IMl 255248, holotypus; 9 Nov. 1981 , IMI 263816, paratypus. Culturae IMI 255248 et 263816 in collectione IMI depositae.
On host. Conidiomata stromatic, pustulate, on galls. Stromata deep-seated densely covered with obpyriform to subulate phialides, 10-16 (- 20) x 3-5 11m. Macroconidia forming pink mucilaginous tendrils ; fusoid, dorsiventrally curved, with rounded apex , frequently swollen in upper half, and tapering to a bluntly rounded base " (1-) 5-8 septate, (15-) 42-50 x (5-) 7-8 11m . In culture. Colonies salmon pink, powdery, orange reverse. Aerial my celium hyaline, floccose. M icroconidia abundant, aseptate, hyaline, oval to T rans. Br . my col. Soc. 82 (4) , (1984)
obovoid, 7-9 x 4-5 11m. Macroconidia on palmate conidiophores, each branch terminating in one or more phialides, highly variable, oval to cylindrical, curved with rounded ends, 1-8 septate, 1250 x 4-8 Ilm. Chlamydospores absent. Teleomorph unknown. Spe cimens examined: On gall on Pinus maximinoi, Pela Nariz, Cordillera Opalaca, Intibuca, Honduras, H . C. Evans, 17 N ov. 1980, IMI 255248 (holotype) ; 9 Nov. 1981 , IMI 263816 (parat yp e),
The conidiomata form within the fleshy, mucilaginous tissues of active, cerebriform galls (up to 24 em diam) on branches of Pinus maximinoi H. E. Moore, appearing initially as shiny, orange blisters or corrugations amongst the corky tissues or periderm. This resinous layer, composed of host cortical cells, ruptures under the pressure of the compact column of macroconidia which typically continues to develop into a long , bright pink (becom ing pale in dry specimens) tendril. The latter consists of tightly-adhering 6- to 8-septate conidia which are of uniform shape and size, with a characteristic rounded apex and tapering base, contrasting with the highly variable, 1- to 8-septate macroconidia, with smoothly-rounded ends usually encountered in culture isolates together with microconidia. Preliminary isolations, made from the internal gall tissues, produced cultures with well-organized sporodochia and compact macroconidial columns as on the host. Hence there was some doubt as to the true identity of the various isolates. This was resolved by preparing single spore cultures using host macroconidia. The latter germinate readily to produce hyphal primordia and within 2-3 day s hyphaI mats become visible on the agar surface. Microconidia develop rap idly and abundantly followed by macroconidia, although the delimitation between the spore types is not sharp, there being a gradation from 1- to 8-septate and from oval to cylindrical. The strictly dorsiventral
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Notes and brief articles
~J lb
B
Fig.
1 (A-C) Cylindrocarpon carneum on host. (A) Conidiogenous structures teased from compact stroma within gall; (B) Macroconidia from unopened conidioma; (C) Macroconidia from spore tendril.
conidia found on the host rarely develop in sub-culture. Futher examination of unopened pustules on the galls revealed that the macroconidial types produced in culture also occur during the early stages of conidiogenesis on the host. Conidial morphology would seem to be dependent upon nutritional or environmental factors. Trans. Br. my col. Soc. 82 (4), (1984)
Colonies on potato sucrose agar (PSA) cover a 9 ern Petri dish within 20 days at 20-24 "C. In old cultures the hyphae aggregate in brown mycelial ropes but chlamydospores were not observed. Chlamydospore-like swellings occasionally develop, however, within old macroconidia. Non-sporulating galls were commonly found on
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Notes and brief articles
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Fig. 2 . Cylindrocarpon carneum on host (A- C) and from culture (D- F) . (A) Spore tendrils on gall surface x 2; (B) T .S. through gall showing deep-seated stroma x 100 ; (C) Macroconidia from spore tendril ; (D) Macroconidia from ro-day-old PSA culture; (E) Microconidial production; (F ) Microconidial and initiation of macroconidial production. (C) and (D) mounted in water, (E) and (F) mounted in lactophenol and cotton blue, all x 700.
other P. maximinoi trees within the survey site, which lies in an area of remnant cloud forest (ea 1800 m a.s.1.) originally cleared for potato and cabbage cultivation. Morphologically similar galls were recorded also on P. maximinoi, and to a lesser extent on P. oocarpa Schiede, in the nea rby extensive forests between La Esperanza and Yamaranguila but all were sterile. Cylindrocarpon earneum was recollected the following year on old , necrotic galls together with several Nectria species, but single ascospore isolations did not reveal an anarnorph-teleomorph connexion. Trans . Br . mycol. Soc. 82 (4) , ( 19 84)
Cylindrocarpon earneum belongs to the Group 1 species of Cylindrocarpon as described by Booth (1966) and shows affinities with C. heteronema (Berk. & Br. ) Wollenw., the anamorph of Neetria galligena Bresadola, a tumour-inducing pathogen of various trees. This raises questions as to the role of C. carneum in the pine gall syndrome. The fact that C. carneum occurs in well-organized conidiomata on active galls, with no evidence of colonization by other micro-organisms, initially suggested that this fungus was the causal agent. However, pathogenicity tests conducted in an isolation
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Notes and brief articles
propagator at the C.M.I., using 6-month-old P. oocarpa seedlings, gave negative results over an 18-month observation period. This does not entirely exclude the possibility ofa direct pathogenic involvement although similar galls have since been collected from a P. "pseudostrobus ' ( = P. maximinoii provenance trial, and from other high altitude pine species, in Honduras, on which rust aecia assignable to the Cronartium quercuum (Berk.) Miyabe & Shirai complex (Burdsall & Snow, 1977 ) were present. Circumstantial evidence indicates, therefore , that P . maximinoi gall s are of rust origin and that C. cameum is functioning as a parasite of galled tissues rather than as a pathogen per se. Clearly, however, the question ofpathogenicity has not been resolved and it may be that there are distinct gall
types on P . maximinoi associated with different causal agents. We are indebted to Mr C. Hughes for rust gall specimens and to Mrs G. Butterfill and Mr D . Fry for technical and photographic assistance respectively . This work forms part of a project (R341o) funded by the U .K. Overseas Development Administration. REFERENCES
BOOTH,C. (1966). The genus Cylindrocarpon, Myc ological
Pap ers 104, 1-56 . BURDSALL ,
H. H. & SNOW, G . A. (19 77) . Taxonomy of
Cronartium quercuum and C. fusi form e. Mycologia 69, 503-508 .
VOLUTELLAR/A LAURINA TASSI, AN EARLIER NAME FOR WIESNERIOMYCES JAVAN/CUS KOORDERS BY P. M. KIRK
Commonwealth My cological In stitute, Ferry Lane, Kew, R ichmond, Surrey TW9 3AF, U.K.
!"
olutl!llaria laur ina is shown to be an earlier binomial for the ubiquitous species Wie sneriomyces ja uamcus. A study of the microfungi colonizing leaf litter of Cha etopelt is laurina (T assi) Sacco in Tassi, Boll. R . Laurus nobilis L. , the sweet bay , in the British Isles Orto bot. Siena 1: 14,1897 (1898). (K irk, 1981, 1982, 1983, 1984) has revealed Tassia laurina (T assi) H . & P. Sydow, Ann. mycol. Wiesner iomycesjavanicus Koorders (1907) to be one 17 : 44 (1919). of the most ubiquitous species recorded on this W. ja vanicus Koorders, Verh. K . ned. Akad. W et. substratum. 246 (1907)· Oudemans (1921) listed 51 species occurring on Tassi (1897) introduced Volutellaria laurina for the leaves of L. nobilis, with a further 74 on twigs, wood and bark. Considering these figures, it is a tuberculariaceous fungus he encountered on apparent that Saccardo and his contemporaries Laurus nobilis leaf litter collected in the botanical paid particular attention to this substratum during gardens, Siena . Saccardo (in Tassi, 1898) introduced their collecting activities. It was likely, therefore, the genus Chaetopeltis Sacco for V . laurina because that they collected W . ja vanicus and although this he considered it belonged in the Leptostromataceae, name dates from 1907 and was included by close to Leptothyrium but with the conidiomata Saccardo (1913), the probability that it had been bearing setae and producing scolecospores. Unfordescribed earlier under a different name is high. A tunately, Chaetopeltis Sacc., 1898, is a homonym of review of the 51 species listed by Oudemans has Chaetopeltis Berth., 1878, a genus of Algae, and Tassia H. & P. Sydow (1919) was, therefore, shown this to be the case. An examination of the holotype of Volutellaria introduced to replace it. Sutton (1977) included laurina Tassi (1897) has revealed that this provides Tassia in his compilation of generic names an earlier name for the taxon presently referred to proposed for Coelomycetes but made no comment W iesneriomyces javanicus (K oorders, 1907). The on its taxonomic status. The affinities of the type species of Volutellaria, follow ing new combination and synonym y is, V . acaroides (Sacc.) Sacc., are not clear. The species therefore, proposed. was not illustrated by Saccardo and it has not been Wiesneriomyces laurinus (T assi) P . M. Kirk possible to locate the holotype. From the descripcomb.nov. tion, however, it is clear that it differs from W . Volutellaria laurina Tassi, Atti R . Accad. Fisiocr. laurinus, especially in conidium morphology. Siena, ser. 4, 8: 551 (1897). Trans. Br. mycol. Soc. 82 (4) , (1984)
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745
produced conidia. The conidiophores are often slightly curved towards the apex. The stroma, such as it is. is similar in older conidiophore fascicles.
A NEW SPECIES OF CYLINDROCARPON ASSOCIATED WITH PINE GALLS IN HONDURAS BY C. BOOTH AND H. C. EVANS
Commonwealth Mycological Institute, Ferry Lan e, K ew, U.K. Cylindrocarpon carneum sp .nov . is described from galls on Pinus maximinoi in Honduras and from culture. Its biology is briefly discussed. In the course of a disease survey of Central American pine species, galls of unknown etiology were encountered on high altitude pines in Honduras. Amongst the fungi associated with these galls was an undescribed species of the genus Cylindrocarpon Wollenw. Although pathogenic involvement has not been proven, its distinctive morphology is considered to warrant a formal description. Cylindrocarpon carneum sp .nov. (F igs 1,2) (E t yrn , - referring to the bright pink colour of the
spore tendrils) In hospitis. Stromata pulvinata innata cavita in galla. Conidiophora ramosa, hyalina, phialidibus clavatis vel su bulatis terminantia, 10-16 (- 20) X 3-5Itm . Macroconi dia cylindrica vel fusoidea , curvula, utrinque rotundat a, 5-8 septata, 42-50 x 7-8 pm, in cirrhi. In cultura. Coloniis salmoneis, farinosis ; rever so aurant iaco. M ycelium aer ium hyalinum, f1occosum. M icroconidia vecta palrnata con idiophora, hyalina ovara ad obovoidea, 7-9 x 4-5 Itm. Macrocon idia variabilia, ovata vel cylindrica, leniter curvata utrinque rotundata, 1-8 septata, 12-50 x 4-8 Itm . Chlamydosporae absentes. Teleomorphosis incognita. In galla Pini maximinoi, Pela Nariz, Cordillera Opalaca, Intibuca, Honduras, H. C. Evans, 17 Nov. 1980, IMl 255248, holotypus; 9 Nov. 1981 , IMI 263816, paratypus. Culturae IMI 255248 et 263816 in collectione IMI depositae.
On host. Conidiomata stromatic, pustulate, on galls. Stromata deep-seated densely covered with obpyriform to subulate phialides, 10-16 (- 20) x 3-5 11m. Macroconidia forming pink mucilaginous tendrils ; fusoid, dorsiventrally curved, with rounded apex , frequently swollen in upper half, and tapering to a bluntly rounded base " (1-) 5-8 septate, (15-) 42-50 x (5-) 7-8 11m . In culture. Colonies salmon pink, powdery, orange reverse. Aerial my celium hyaline, floccose. M icroconidia abundant, aseptate, hyaline, oval to T rans. Br . my col. Soc. 82 (4) , (1984)
obovoid, 7-9 x 4-5 11m. Macroconidia on palmate conidiophores, each branch terminating in one or more phialides, highly variable, oval to cylindrical, curved with rounded ends, 1-8 septate, 1250 x 4-8 Ilm. Chlamydospores absent. Teleomorph unknown. Spe cimens examined: On gall on Pinus maximinoi, Pela Nariz, Cordillera Opalaca, Intibuca, Honduras, H . C. Evans, 17 N ov. 1980, IMI 255248 (holotype) ; 9 Nov. 1981 , IMI 263816 (parat yp e),
The conidiomata form within the fleshy, mucilaginous tissues of active, cerebriform galls (up to 24 em diam) on branches of Pinus maximinoi H. E. Moore, appearing initially as shiny, orange blisters or corrugations amongst the corky tissues or periderm. This resinous layer, composed of host cortical cells, ruptures under the pressure of the compact column of macroconidia which typically continues to develop into a long , bright pink (becom ing pale in dry specimens) tendril. The latter consists of tightly-adhering 6- to 8-septate conidia which are of uniform shape and size, with a characteristic rounded apex and tapering base, contrasting with the highly variable, 1- to 8-septate macroconidia, with smoothly-rounded ends usually encountered in culture isolates together with microconidia. Preliminary isolations, made from the internal gall tissues, produced cultures with well-organized sporodochia and compact macroconidial columns as on the host. Hence there was some doubt as to the true identity of the various isolates. This was resolved by preparing single spore cultures using host macroconidia. The latter germinate readily to produce hyphal primordia and within 2-3 day s hyphaI mats become visible on the agar surface. Microconidia develop rap idly and abundantly followed by macroconidia, although the delimitation between the spore types is not sharp, there being a gradation from 1- to 8-septate and from oval to cylindrical. The strictly dorsiventral
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746
Notes and brief articles
~J lb
B
Fig.
1 (A-C) Cylindrocarpon carneum on host. (A) Conidiogenous structures teased from compact stroma within gall; (B) Macroconidia from unopened conidioma; (C) Macroconidia from spore tendril.
conidia found on the host rarely develop in sub-culture. Futher examination of unopened pustules on the galls revealed that the macroconidial types produced in culture also occur during the early stages of conidiogenesis on the host. Conidial morphology would seem to be dependent upon nutritional or environmental factors. Trans. Br. my col. Soc. 82 (4), (1984)
Colonies on potato sucrose agar (PSA) cover a 9 ern Petri dish within 20 days at 20-24 "C. In old cultures the hyphae aggregate in brown mycelial ropes but chlamydospores were not observed. Chlamydospore-like swellings occasionally develop, however, within old macroconidia. Non-sporulating galls were commonly found on
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Notes and brief articles
747
Fig. 2 . Cylindrocarpon carneum on host (A- C) and from culture (D- F) . (A) Spore tendrils on gall surface x 2; (B) T .S. through gall showing deep-seated stroma x 100 ; (C) Macroconidia from spore tendril ; (D) Macroconidia from ro-day-old PSA culture; (E) Microconidial production; (F ) Microconidial and initiation of macroconidial production. (C) and (D) mounted in water, (E) and (F) mounted in lactophenol and cotton blue, all x 700.
other P. maximinoi trees within the survey site, which lies in an area of remnant cloud forest (ea 1800 m a.s.1.) originally cleared for potato and cabbage cultivation. Morphologically similar galls were recorded also on P. maximinoi, and to a lesser extent on P. oocarpa Schiede, in the nea rby extensive forests between La Esperanza and Yamaranguila but all were sterile. Cylindrocarpon earneum was recollected the following year on old , necrotic galls together with several Nectria species, but single ascospore isolations did not reveal an anarnorph-teleomorph connexion. Trans . Br . mycol. Soc. 82 (4) , ( 19 84)
Cylindrocarpon earneum belongs to the Group 1 species of Cylindrocarpon as described by Booth (1966) and shows affinities with C. heteronema (Berk. & Br. ) Wollenw., the anamorph of Neetria galligena Bresadola, a tumour-inducing pathogen of various trees. This raises questions as to the role of C. carneum in the pine gall syndrome. The fact that C. carneum occurs in well-organized conidiomata on active galls, with no evidence of colonization by other micro-organisms, initially suggested that this fungus was the causal agent. However, pathogenicity tests conducted in an isolation
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Notes and brief articles
propagator at the C.M.I., using 6-month-old P. oocarpa seedlings, gave negative results over an 18-month observation period. This does not entirely exclude the possibility ofa direct pathogenic involvement although similar galls have since been collected from a P. "pseudostrobus ' ( = P. maximinoii provenance trial, and from other high altitude pine species, in Honduras, on which rust aecia assignable to the Cronartium quercuum (Berk.) Miyabe & Shirai complex (Burdsall & Snow, 1977 ) were present. Circumstantial evidence indicates, therefore , that P . maximinoi gall s are of rust origin and that C. cameum is functioning as a parasite of galled tissues rather than as a pathogen per se. Clearly, however, the question ofpathogenicity has not been resolved and it may be that there are distinct gall
types on P . maximinoi associated with different causal agents. We are indebted to Mr C. Hughes for rust gall specimens and to Mrs G. Butterfill and Mr D . Fry for technical and photographic assistance respectively . This work forms part of a project (R341o) funded by the U .K. Overseas Development Administration. REFERENCES
BOOTH,C. (1966). The genus Cylindrocarpon, Myc ological
Pap ers 104, 1-56 . BURDSALL ,
H. H. & SNOW, G . A. (19 77) . Taxonomy of
Cronartium quercuum and C. fusi form e. Mycologia 69, 503-508 .
VOLUTELLAR/A LAURINA TASSI, AN EARLIER NAME FOR WIESNERIOMYCES JAVAN/CUS KOORDERS BY P. M. KIRK
Commonwealth My cological In stitute, Ferry Lane, Kew, R ichmond, Surrey TW9 3AF, U.K.
!"
olutl!llaria laur ina is shown to be an earlier binomial for the ubiquitous species Wie sneriomyces ja uamcus. A study of the microfungi colonizing leaf litter of Cha etopelt is laurina (T assi) Sacco in Tassi, Boll. R . Laurus nobilis L. , the sweet bay , in the British Isles Orto bot. Siena 1: 14,1897 (1898). (K irk, 1981, 1982, 1983, 1984) has revealed Tassia laurina (T assi) H . & P. Sydow, Ann. mycol. Wiesner iomycesjavanicus Koorders (1907) to be one 17 : 44 (1919). of the most ubiquitous species recorded on this W. ja vanicus Koorders, Verh. K . ned. Akad. W et. substratum. 246 (1907)· Oudemans (1921) listed 51 species occurring on Tassi (1897) introduced Volutellaria laurina for the leaves of L. nobilis, with a further 74 on twigs, wood and bark. Considering these figures, it is a tuberculariaceous fungus he encountered on apparent that Saccardo and his contemporaries Laurus nobilis leaf litter collected in the botanical paid particular attention to this substratum during gardens, Siena . Saccardo (in Tassi, 1898) introduced their collecting activities. It was likely, therefore, the genus Chaetopeltis Sacco for V . laurina because that they collected W . ja vanicus and although this he considered it belonged in the Leptostromataceae, name dates from 1907 and was included by close to Leptothyrium but with the conidiomata Saccardo (1913), the probability that it had been bearing setae and producing scolecospores. Unfordescribed earlier under a different name is high. A tunately, Chaetopeltis Sacc., 1898, is a homonym of review of the 51 species listed by Oudemans has Chaetopeltis Berth., 1878, a genus of Algae, and Tassia H. & P. Sydow (1919) was, therefore, shown this to be the case. An examination of the holotype of Volutellaria introduced to replace it. Sutton (1977) included laurina Tassi (1897) has revealed that this provides Tassia in his compilation of generic names an earlier name for the taxon presently referred to proposed for Coelomycetes but made no comment W iesneriomyces javanicus (K oorders, 1907). The on its taxonomic status. The affinities of the type species of Volutellaria, follow ing new combination and synonym y is, V . acaroides (Sacc.) Sacc., are not clear. The species therefore, proposed. was not illustrated by Saccardo and it has not been Wiesneriomyces laurinus (T assi) P . M. Kirk possible to locate the holotype. From the descripcomb.nov. tion, however, it is clear that it differs from W . Volutellaria laurina Tassi, Atti R . Accad. Fisiocr. laurinus, especially in conidium morphology. Siena, ser. 4, 8: 551 (1897). Trans. Br. mycol. Soc. 82 (4) , (1984)
Printed in Great Brita in