Phylogenetic tree resulting from a Bayesian analysis on the combined alignment of five genes (rpb2, ITS, actA, gapdh, tef1-α). Bayesian posterior probabilities (BPP), maximum likelihood bootstrap support values ( 80 %; MLBS) and parsimony bootstrap support values ( 80 %; PBS) are indicated at the nodes (BPP/MLBS/PBS). Values of BPP/MLBS/PBS equal to 1/100/100 were replaced with a hash (#). The scale bar represents the expected number of changes per site. Branches in a thicker stroke represent the branches present in the strict consensus parsimony tree. Species clades are delimited in coloured blocks, where strain numbers are written in black, host names in blue and country of origin in brown. The current species name and clade number are indicated to the right of the tree. Type strains are represented in bold. The tree was rooted to Zymoseptoria halophila (CBS 128854). 

Phylogenetic tree resulting from a Bayesian analysis on the combined alignment of five genes (rpb2, ITS, actA, gapdh, tef1-α). Bayesian posterior probabilities (BPP), maximum likelihood bootstrap support values ( 80 %; MLBS) and parsimony bootstrap support values ( 80 %; PBS) are indicated at the nodes (BPP/MLBS/PBS). Values of BPP/MLBS/PBS equal to 1/100/100 were replaced with a hash (#). The scale bar represents the expected number of changes per site. Branches in a thicker stroke represent the branches present in the strict consensus parsimony tree. Species clades are delimited in coloured blocks, where strain numbers are written in black, host names in blue and country of origin in brown. The current species name and clade number are indicated to the right of the tree. Type strains are represented in bold. The tree was rooted to Zymoseptoria halophila (CBS 128854). 

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Ramularia is a species-rich genus that harbours plant pathogens responsible for yield losses to many important crops, including barley, sugar beet and strawberry. Species of Ramularia are hyphomycetes with hyaline conidiophores and conidia with distinct, thickened, darkened, refractive conidiogenous loci and conidial hila, and Mycosphaerella sexual...

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... The species epithet of Ramularia hydrangeae-macro- phyllae reflects the name of the host on which it was first observed, Hydrangea macrophylla, from New Zealand (holotype in HAL). Within this clade the phylogenetic structure was not resolved consistently in all gene trees (data not shown; Fig. 2, clade 21) and, in accordance with the Genealogical Concor- dance Phylogenetic Species Recognition (GCPSR) concept, the transition from concordance to conflict determined the limit of these species ). Ramularia sparganii was described from Sparganium emersum from Sweden (holotype in C) and has not been reported from the Netherlands ). The species produces conidiophores that are sub- cylindrical to geniculate-sinuous, 5-40(-60) × 1-3 μm, catenate conidia, smooth, ellipsoid-fusoid, 0-1-septate, 8-30(-33) × 1.5-3 μm and with minute hila. The strain CBS 159.82 was possibly misidentified based on the host but was sterile in culture and morphological characters were not observed. Ramularia hellebori was described from Helleborus foetidus from Germany (lectotype in HAL), and was firstly re- ported from New Zealand on Helleborus orientalis (Braun & Hill 2002) and later on Helleborus niger (CBS 118408) ( ), but no ex-type culture was designated. This species description includes conidiophores that are subcylindrical to geniculate sinuous, 10-45 × 1.5-5 μm, conidia catenate, ellipsoid-ovoid to fusiform, verruculose, 0-1-septate, 6-20(-30) × 2-4 μm and minute hila. Ramularia rollandii was described from Iris pseudacorus from France (lectotype in PC) and the species was reported from New Zealand on an Iris × hollandica hybrid (CBS 122625) (Braun & Hill 2008), but no ex-type culture is known. This species produces conidiophores that are cylindrical to geniculate-sinuous, apically minutely sub- denticulate, 5-15(-20) × 2-3 μm, conidia solitary or in short chains, smooth to faintly verruculose, filiform to acicular, 15-40(-60) × 1-2 μm, 1-4-septate, with minute hila. Ramularia butomi is mycophylic and was originally described overgrowing ascomycetous stromata on dead leaves of Butomus umbellatus in Sweden (lectotype in B), but the strain CBS 114117 is not documented as hyperparasite in the database. This species produces conidiophores that are simple, subcylindrical to geniculate-sinuous, 8-60 × 1-4 μm, conidia catenate, narrowly ellipsoid-ovoid to subcylindrical-fusiform, (5-) 8-16(-24) × (1.5-)2-3(-4) μm, 0-1(-2)-septate, verruculose and with minute hila. Ramularia deusta var. alba is not reported from Ulex and the representative clade for this species has been designated in this study (Fig. 2, clade 62). All the species mentioned above have in common that the conidia are catenate and slightly verruculose to verruculose, with minute hila, but size and septation vary among them. It is necessary to collect fresh material from the type location and host for further observations. The only ex-type culture present in this clade is that of Ramularia hydrangeae-macrophyllae (CBS 122273) (Braun & Hill 2008), and in accordance with the GCPST concept, we accept that name for this clade. This species, now with a broad host range and wide geographical distribution, forms a highly supported clade (Fig. 2, clade 21, 1/100/100). Similar intraspecific variation, wide host range and geographical distribution have been observed before for Ramularia vizellae (Videira et al. 2015b; Fig. 2, clade 85). Strains CPC 25901 and CPC 25902 were isolated using the method developed for single ascospore isolation for Mycosphaerella ( Crous et al. 1991, Crous 1998, which means this species has a sexual morph (Fig. 51) Substrate and distribution: Only known from South ...
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... The species epithet of Ramularia hydrangeae-macro- phyllae reflects the name of the host on which it was first observed, Hydrangea macrophylla, from New Zealand (holotype in HAL). Within this clade the phylogenetic structure was not resolved consistently in all gene trees (data not shown; Fig. 2, clade 21) and, in accordance with the Genealogical Concor- dance Phylogenetic Species Recognition (GCPSR) concept, the transition from concordance to conflict determined the limit of these species ). Ramularia sparganii was described from Sparganium emersum from Sweden (holotype in C) and has not been reported from the Netherlands ). The species produces conidiophores that are sub- cylindrical to geniculate-sinuous, 5-40(-60) × 1-3 μm, catenate conidia, smooth, ellipsoid-fusoid, 0-1-septate, 8-30(-33) × 1.5-3 μm and with minute hila. The strain CBS 159.82 was possibly misidentified based on the host but was sterile in culture and morphological characters were not observed. Ramularia hellebori was described from Helleborus foetidus from Germany (lectotype in HAL), and was firstly re- ported from New Zealand on Helleborus orientalis (Braun & Hill 2002) and later on Helleborus niger (CBS 118408) ( ), but no ex-type culture was designated. This species description includes conidiophores that are subcylindrical to geniculate sinuous, 10-45 × 1.5-5 μm, conidia catenate, ellipsoid-ovoid to fusiform, verruculose, 0-1-septate, 6-20(-30) × 2-4 μm and minute hila. Ramularia rollandii was described from Iris pseudacorus from France (lectotype in PC) and the species was reported from New Zealand on an Iris × hollandica hybrid (CBS 122625) (Braun & Hill 2008), but no ex-type culture is known. This species produces conidiophores that are cylindrical to geniculate-sinuous, apically minutely sub- denticulate, 5-15(-20) × 2-3 μm, conidia solitary or in short chains, smooth to faintly verruculose, filiform to acicular, 15-40(-60) × 1-2 μm, 1-4-septate, with minute hila. Ramularia butomi is mycophylic and was originally described overgrowing ascomycetous stromata on dead leaves of Butomus umbellatus in Sweden (lectotype in B), but the strain CBS 114117 is not documented as hyperparasite in the database. This species produces conidiophores that are simple, subcylindrical to geniculate-sinuous, 8-60 × 1-4 μm, conidia catenate, narrowly ellipsoid-ovoid to subcylindrical-fusiform, (5-) 8-16(-24) × (1.5-)2-3(-4) μm, 0-1(-2)-septate, verruculose and with minute hila. Ramularia deusta var. alba is not reported from Ulex and the representative clade for this species has been designated in this study (Fig. 2, clade 62). All the species mentioned above have in common that the conidia are catenate and slightly verruculose to verruculose, with minute hila, but size and septation vary among them. It is necessary to collect fresh material from the type location and host for further observations. The only ex-type culture present in this clade is that of Ramularia hydrangeae-macrophyllae (CBS 122273) (Braun & Hill 2008), and in accordance with the GCPST concept, we accept that name for this clade. This species, now with a broad host range and wide geographical distribution, forms a highly supported clade (Fig. 2, clade 21, 1/100/100). Similar intraspecific variation, wide host range and geographical distribution have been observed before for Ramularia vizellae (Videira et al. 2015b; Fig. 2, clade 85). Strains CPC 25901 and CPC 25902 were isolated using the method developed for single ascospore isolation for Mycosphaerella ( Crous et al. 1991, Crous 1998, which means this species has a sexual morph (Fig. 51) Substrate and distribution: Only known from South ...
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... The species epithet of Ramularia hydrangeae-macro- phyllae reflects the name of the host on which it was first observed, Hydrangea macrophylla, from New Zealand (holotype in HAL). Within this clade the phylogenetic structure was not resolved consistently in all gene trees (data not shown; Fig. 2, clade 21) and, in accordance with the Genealogical Concor- dance Phylogenetic Species Recognition (GCPSR) concept, the transition from concordance to conflict determined the limit of these species ). Ramularia sparganii was described from Sparganium emersum from Sweden (holotype in C) and has not been reported from the Netherlands ). The species produces conidiophores that are sub- cylindrical to geniculate-sinuous, 5-40(-60) × 1-3 μm, catenate conidia, smooth, ellipsoid-fusoid, 0-1-septate, 8-30(-33) × 1.5-3 μm and with minute hila. The strain CBS 159.82 was possibly misidentified based on the host but was sterile in culture and morphological characters were not observed. Ramularia hellebori was described from Helleborus foetidus from Germany (lectotype in HAL), and was firstly re- ported from New Zealand on Helleborus orientalis (Braun & Hill 2002) and later on Helleborus niger (CBS 118408) ( ), but no ex-type culture was designated. This species description includes conidiophores that are subcylindrical to geniculate sinuous, 10-45 × 1.5-5 μm, conidia catenate, ellipsoid-ovoid to fusiform, verruculose, 0-1-septate, 6-20(-30) × 2-4 μm and minute hila. Ramularia rollandii was described from Iris pseudacorus from France (lectotype in PC) and the species was reported from New Zealand on an Iris × hollandica hybrid (CBS 122625) (Braun & Hill 2008), but no ex-type culture is known. This species produces conidiophores that are cylindrical to geniculate-sinuous, apically minutely sub- denticulate, 5-15(-20) × 2-3 μm, conidia solitary or in short chains, smooth to faintly verruculose, filiform to acicular, 15-40(-60) × 1-2 μm, 1-4-septate, with minute hila. Ramularia butomi is mycophylic and was originally described overgrowing ascomycetous stromata on dead leaves of Butomus umbellatus in Sweden (lectotype in B), but the strain CBS 114117 is not documented as hyperparasite in the database. This species produces conidiophores that are simple, subcylindrical to geniculate-sinuous, 8-60 × 1-4 μm, conidia catenate, narrowly ellipsoid-ovoid to subcylindrical-fusiform, (5-) 8-16(-24) × (1.5-)2-3(-4) μm, 0-1(-2)-septate, verruculose and with minute hila. Ramularia deusta var. alba is not reported from Ulex and the representative clade for this species has been designated in this study (Fig. 2, clade 62). All the species mentioned above have in common that the conidia are catenate and slightly verruculose to verruculose, with minute hila, but size and septation vary among them. It is necessary to collect fresh material from the type location and host for further observations. The only ex-type culture present in this clade is that of Ramularia hydrangeae-macrophyllae (CBS 122273) (Braun & Hill 2008), and in accordance with the GCPST concept, we accept that name for this clade. This species, now with a broad host range and wide geographical distribution, forms a highly supported clade (Fig. 2, clade 21, 1/100/100). Similar intraspecific variation, wide host range and geographical distribution have been observed before for Ramularia vizellae (Videira et al. 2015b; Fig. 2, clade 85). Strains CPC 25901 and CPC 25902 were isolated using the method developed for single ascospore isolation for Mycosphaerella ( Crous et al. 1991, Crous 1998, which means this species has a sexual morph (Fig. 51) Substrate and distribution: Only known from South ...
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... Previously identified as Ramularia inaequalis these strains in fact represent a separate species (Fig. 50), since they do not cluster together with the type of R. inaequalis (Fig. 2, clade 40). The morphology agrees well with R. hieracii-umbellati described on Hieracium umbellatum from Sweden. Cultures and sequences based on collections from Sweden are not available for comparison, but since the Korean material might belong to this species, we prefer to apply the latter name, at least tenta- tively. Hieracium umbellatum is a widespread circumpolar spe- cies. Strains of R. hieracii-umbellati form a highly supported clade (Fig. 2, clade 56, 1/100/100). Ramularia hieracii-umbellati formed a sister clade to R. rhabdospora, but the latter produces wider conidiophores [10-50(-115) × 2-8 μm] and larger cate- nate conidia [(10-)15-40(-50) × 3-7 μm], echinulate, ellipsoid- ovoid to cylindrical, and 0-3(-4)-septate Fig. ...
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... Previously identified as Ramularia inaequalis these strains in fact represent a separate species (Fig. 50), since they do not cluster together with the type of R. inaequalis (Fig. 2, clade 40). The morphology agrees well with R. hieracii-umbellati described on Hieracium umbellatum from Sweden. Cultures and sequences based on collections from Sweden are not available for comparison, but since the Korean material might belong to this species, we prefer to apply the latter name, at least tenta- tively. Hieracium umbellatum is a widespread circumpolar spe- cies. Strains of R. hieracii-umbellati form a highly supported clade (Fig. 2, clade 56, 1/100/100). Ramularia hieracii-umbellati formed a sister clade to R. rhabdospora, but the latter produces wider conidiophores [10-50(-115) × 2-8 μm] and larger cate- nate conidia [(10-)15-40(-50) × 3-7 μm], echinulate, ellipsoid- ovoid to cylindrical, and 0-3(-4)-septate Fig. ...
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... This species has been recently described (Park & Shin 2016) and is only known from South Korea. Until now, only two Ramularia species were known to infect Hydrangea hosts, R. hydrangeae Y.L. Guo & U. Braun (on Hydrangea bretsch- neideri, China, holotype in HMAS) and Ramularia hydrangeae- macrophyllae U. Braun & C.F. Hill (on Hydrangea-macrophylla, New Zealand, holotype in HAL). The isolates of Ramularia hydrangeicola cluster in a highly supported clade (Fig. 2, clade 70, 1/100/100) and are not conspecific with R. hydrangeae- macrophyllae (Fig. 2, clade 21 For additional synonyms see ...
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... Ramularia bosniaca was originally described on the host Scabiosa columbaria in Bosnia (holotype BPI 416442 [the neotype designated in , BPI 416443, is obsolete since holotype material has been traced in BPI]) and is the only species of Ramularia known to infect Scabiosa (Braun 1998). Ramularia bosniaca has been reported from several European countries but this is the first time it is reported from the Czech Republic. The description of these collections fit that of R. bosniaca (Braun 1998), except that conidia were shorter than 28 μm (Fig. 39). Additional collections from Montenegro may well reveal the strains from the Czech Republic to represent a new species. The phylogenetic analysis provides good support to this species clade (Fig. 2, clade 60, ...
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... Ramularia beticola (Fig. 38) is the causal organism of Ramularia leaf spot disease in sugar beet, table beet and fodder beet. The fungus forms pale brown leaf spots and affected leaves turn yellow, become necrotic and die. The impact of Ramularia leaf spot disease can vary significantly from season to season. Conditions of high humidity, moderate temperature (17-20 °C), high plant density and sulphur deficiency usually increase disease intensity and damage. It has been reported from North America (Oregon, Washington, California and Colo- rado), Europe (Ireland, UK, the Scandinavian countries, Belgium, France, Germany) and Russia ( Harveson et al. 2009). World- wide, yield losses in sugar beet due to plant pathogens and pests are estimated in general to be 26 % with, and more than 80 % without crop protection (Oerke & Dehne 2004). In the Netherlands, in spite of crop protection measures, the yield losses due to pests and diseases for top growers were 37.1 and 16.7 % on sandy and clay soils respectively ( Hanse et al. 2011). When treatments are applied timely, programmes of disease control in Denmark increased sugar yield by 10 %. Thus far, R. beticola has not shown signs of developing resistance to either strobilurin or triazol fungicides, but it remains important to apply fungicides efficiently by following monitoring programmes and respecting the recommended thresholds (www.FRAC.info) ( Thach et al. 2013). Ramularia beticola was described on Beta vulgaris from France in 1896. The strains used in this study clustered together in a single and highly supported clade (Fig. 2, clade 52, 1/100/100). Mycelium consisting of hyaline, septate, branched, smooth, 1 -3 μm diam hyphae. Conidiophores hyaline, thin-walled, smooth, erect, 1 -2(-4)-septate, straight to geniculate- sinuous, cylindrical-oblong, unbranched, (20-) 41 -60( -119) × 2 -2.5(-3) μm or reduced to conidiogenous cells. Conidiogenous cells integrated in mycelium or terminal in conidiophores, cylindrical-oblong and narrower at the top, geniculate-sinuous, (14-)19 -23(-35) × 2-2.5(-3) μm, with 1 -4 apical conidiogenous loci, almost flat or protuberant, thickened, darkened and refractive. Conidia hyaline, thin- walled, smooth, solitary or catenate, aseptate, with hila thickened, darkened and refractive. Ramoconidia sub- cylindrical to ovoid, (7.5-)10 -11( -14) × (3 -)4-4.5(-5) μm, with two apical hila. Intercalary conidia ovoid, 0 -1-septate, (8 -)9.5 -10.5(-13) × (3 -)4 -5(-7) μm, in branched chains of up to four conidia. Terminal conidia obovoid, (5 -) 7 -8(-10) × (2.5 -)4(-5) μm (on ...
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... Ramularia heraclei was originally described on Herac- leum sphondylium from the Netherlands (lectotype in L). Strains of Ramularia heraclei used in this study formed two sister clades that are both highly supported in the multigene phylogeny (Fig. 2, clade 78, 1/100/100). In literature , the description of this species is quite broad including conidiophores in fascicles or forming crustose-like layers, erect and simple, cylindrical to geniculate-sinuous and variable in length, 5-80(-110) × 2-6 μm and conidia which are catenate, 0-3-septate, smooth to verruculose, (8-)10-35(-45) × 2-6 μm. The herbarium mate- rial corresponding to strain CBS 108972 has short co- nidiophores, (5-)12-17(-20) × 2-3 μm and conidia which are catenate, verruculose, 0-1-septate, (6-)11-14(-25) × (2-) 3(-6) μm . The herbarium material corresponding to the strain CBS 108988 has longer conidiophores, (29-) 49-59(-82) × 2-3 μm and longer conidia, catenate, smooth to slightly verruculose, (0-)1-3-septate and (6.5-) 15.5-20(-36) × (2.5-)3.5-4(-6) μm. They both fit the morphological description in literature and strain CBS 108969, which was collected from the Netherlands and isolated from Heracleum sphondylium, the same location and host as the type species, and is herewith designated as epitype (Fig. 49). The morphology of Ramularia collections from hosts of Apiaceae that are preserved in herbaria are difficult to distinguish and several names were synonymised with R. heraclei ). The variation in morphology and the phylogeny indicate that this may be a species complex that needs further study and comparison with collections from other apiaceous hosts. In planta: Leaf spots rectangular, following the leaf nerves, yellowish to brown. Caespituli emerging through stomata, hyaline to buff. Conidiophores hyaline, thin-walled, smooth, erect, septate, cylindrical-oblong, straight to geniculate-sinuous, rarely branched, (14-)27-32(-41.5) × (2-)3(-4) μm. Conidiogenous cells terminal or intermediate in the conidiophore, cylindrical- oblong or geniculate-sinuous, (7-)10-13(-22) × (2-) 3(-4) μm, with multiple conidiogenous loci almost flat to protu- berant, thickened, darkened and refractive. Conidia hyaline, thin- walled, smooth, solitary or in short chains unbranched, cylindrical- oblong to obovate, (0-)1-2-septate, (8-)15-20(-25) × (2-) 2.5-3 μm with hila thickened, darkened and ...
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... pigmentosum Videira & Crous, sp. nov. Myco- Bank MB817152. Fig. ...
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... This species was described on Bunias orientalis collected in Sweden (lectotype in S). Ramularia buniadis (among other names) was synonymised with R. armoraciae, but since R. armoraciae clusters in clade 55 (Fig. 2), the name R. buniadis is again resurrected for this isolate (Fig. 2, clade 48). Although this isolate could be considered a good representative for epi- typification, it is sterile in culture and no herbarium material of the CBS isolate was ...
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... This species was described on Bunias orientalis collected in Sweden (lectotype in S). Ramularia buniadis (among other names) was synonymised with R. armoraciae, but since R. armoraciae clusters in clade 55 (Fig. 2), the name R. buniadis is again resurrected for this isolate (Fig. 2, clade 48). Although this isolate could be considered a good representative for epi- typification, it is sterile in culture and no herbarium material of the CBS isolate was ...
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... characteristics: On MEA, 15 mm diam, surface raised, folded, smooth mycelium, smoke-grey with olivaceous tinge, with small buff droplets, with margins lobate, convex, feathery, colony reverse olivaceous grey and ochraceous patches; on OA, 15 mm diam, surface low convex, smooth mycelium, white, with margins buff, naked, undulate, colony reverse rosy- buff; on PDA, 20 mm diam, surface low convex, smooth mycelium, white with greyish tinge with margins undulate, feathery, colony reverse rosy-buff and iron-grey patches. Notes: These strains were initially identified as R. inaequalis, but this species clusters in clade 40 (Fig. 2). Several names have been synonymised with R. inaequalis that refer to spe- cies isolated from different hosts and locations. These need to be recollected and re-examined since it appears that R. inaequalis is a species complex. Helminthotheca echioides is of Mediterranean origin, but now with a widespread, almost cosmopolitan neophytic distribution. The descriptions of both species named R. helminthiae are from the neophytic area of the host, but the origin of the species concerned is probably Mediterranean as well. A sporulating culture based on material collected on H. echioides in Turkey or adjacent countries should serve as epitype for this species, but is not yet avail- able. Therefore, the name R. helminthiae is only tentatively used for the present strains until appropriate cultures will be available. Ramularia helminthiae (Fig. 48) is supported as distinct from other included species by the phylogenetic ana- lyses (Fig. 2, Substrate and distribution: On Apium, Cicuta, Conium, Corian- drum, Hansenia, Heracleum, Levisticum, Malabaila, and Pasti- naca (Apiaceae); Asia, Africa, Caucasus, Europe, New Zealand, N. America and West ...
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... characteristics: On MEA, 15 mm diam, surface raised, folded, smooth mycelium, smoke-grey with olivaceous tinge, with small buff droplets, with margins lobate, convex, feathery, colony reverse olivaceous grey and ochraceous patches; on OA, 15 mm diam, surface low convex, smooth mycelium, white, with margins buff, naked, undulate, colony reverse rosy- buff; on PDA, 20 mm diam, surface low convex, smooth mycelium, white with greyish tinge with margins undulate, feathery, colony reverse rosy-buff and iron-grey patches. Notes: These strains were initially identified as R. inaequalis, but this species clusters in clade 40 (Fig. 2). Several names have been synonymised with R. inaequalis that refer to spe- cies isolated from different hosts and locations. These need to be recollected and re-examined since it appears that R. inaequalis is a species complex. Helminthotheca echioides is of Mediterranean origin, but now with a widespread, almost cosmopolitan neophytic distribution. The descriptions of both species named R. helminthiae are from the neophytic area of the host, but the origin of the species concerned is probably Mediterranean as well. A sporulating culture based on material collected on H. echioides in Turkey or adjacent countries should serve as epitype for this species, but is not yet avail- able. Therefore, the name R. helminthiae is only tentatively used for the present strains until appropriate cultures will be available. Ramularia helminthiae (Fig. 48) is supported as distinct from other included species by the phylogenetic ana- lyses (Fig. 2, Substrate and distribution: On Apium, Cicuta, Conium, Corian- drum, Hansenia, Heracleum, Levisticum, Malabaila, and Pasti- naca (Apiaceae); Asia, Africa, Caucasus, Europe, New Zealand, N. America and West ...
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... Ramularia calcea is a species with a wide distribution within Europe that was originally described on Symphytum officinale from Italy. Morphologically, the strains in this clade have conidiophores reduced to conidiogenous cells and nar- rower conidia (Fig. 40) than the in vivo description of R. calcea found in literature . Unfortunately, the her- barium specimen from which the culture was retrieved was not preserved which made it impossible to assess the morpho- logical characters of this species on host tissue. The clade formed by these two strains is highly supported by the phylogenetic analysis (Fig. 2, clade 42). Strain CBS 299.49 (Fig. 2, clade 36) is also under the name R. calcea in the CBS database, but since this is sterile it will be treated as Ramu- laria sp. C. Therefore, the correct phylogenetic placement of this species remains unresolved until material from the type host and location is recollected. Ramularia calcea has been reported on Symphytum officinale in both Germany and the Netherlands, among other countries. The morphological characteristics of CBS 114442 are identical to CBS 102612 and 102613, therefore, until more collections become avail- able, these strains will be treated as R. calcea here. Substrate and distribution: On Scrophularia (Scrophulariaceae); Asia, Caucasus, Armenia, Europe, N. ...
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... Ramularia calcea is a species with a wide distribution within Europe that was originally described on Symphytum officinale from Italy. Morphologically, the strains in this clade have conidiophores reduced to conidiogenous cells and nar- rower conidia (Fig. 40) than the in vivo description of R. calcea found in literature . Unfortunately, the her- barium specimen from which the culture was retrieved was not preserved which made it impossible to assess the morpho- logical characters of this species on host tissue. The clade formed by these two strains is highly supported by the phylogenetic analysis (Fig. 2, clade 42). Strain CBS 299.49 (Fig. 2, clade 36) is also under the name R. calcea in the CBS database, but since this is sterile it will be treated as Ramu- laria sp. C. Therefore, the correct phylogenetic placement of this species remains unresolved until material from the type host and location is recollected. Ramularia calcea has been reported on Symphytum officinale in both Germany and the Netherlands, among other countries. The morphological characteristics of CBS 114442 are identical to CBS 102612 and 102613, therefore, until more collections become avail- able, these strains will be treated as R. calcea here. Substrate and distribution: On Scrophularia (Scrophulariaceae); Asia, Caucasus, Armenia, Europe, N. ...
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... conidiophores and pigmented mycelium (Fig. 20), as well as 47 nucleotides in rpb2, and 5 in LSU. Etymology: composed of Terato-from Teratosphaeriaceae and ...
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... veronicae is known from several Veronica spp. worldwide with the exception of Australia and Antartica. Ramu- laria beccabungae has been described from several Veronica spp. in Europe and Asia. Some Ramularia species have been shown to be plurivorous while others can be seen as host specific. Phylogeny based on five partial gene sequences places sequences retrieved from Ramularia on Veronica chamaedrys (as ?R. chamaedryos), V. persica (as ?R. veronicae, but not the type host) and V. anagallis-aquatica (as ?R. beccabungae) in the same clade (Fig. 2, clade 61, 1/100/100), suggesting that a single species is involved. However, this assumption is still vague and unproven since the description of the sporulation in vitro is only based on a culture of R. chamaedryos on Veronica chamaedrys (Fig. 41). Sporulating cultures of R. veronicae and R. beccabungae based on isolations from the type hosts are necessary for comparison and to evaluate and explain possible differences in the conidial septation between in vivo and in vitro material. Therefore, a final taxonomic conclusion is not yet ...
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... consisting of hyaline, septate, branched, thin-walled, smooth, 1-2 μm diam hyphae. Conidiophores reduced to con- idiogenous cells. Conidiogenous cells hyaline, thin-walled, smooth, integrated in hyphae, cylindrical-oblong, (5-) 10-12.5(-16.5) × (1-)1.5(-2) μm, with 1 thickened and dark- ened apical locus, 1 μm diam. Conidia are catenate, forming ramoconidia, intercalary conidia and terminal conidia. Conidia (type I) hyaline, thin-walled, smooth, catenate, aseptate or occa- sionally 1-septate, with hila conspicuous, thickened and darkened, 1 μm diam; ramoconidia subcylindrical to fusiform, (8.5-) 12-15(-23) × (1.5-)2(-2.5) μm, with 2 apical hila; intercalary conidia subcylindrical, sometimes slightly curved, (6.5-) 10-13(-20) × (1.5-)2(-2.5) μm, in chains of up to five conidia; terminal conidia subcylindrical to obovoid, (3-) 5.5-6(-8) × (1.5-)2(-3) μm. Conidia (type II) brown, smooth, catenate, 1-4-septate, constricted at the septa, (5-) 11.5-14.5(-18.5) × (2-)2.5-3 μm, with hila thickened and darkened. Notes: A total of seven Ramularia species have been described from Rumex worldwide ) and two of these species form filiform, long conidia, i.e. R. pseudodecipiens and R. pratensis. Ramularia pseudodeci- piens is only known from the type collection in the USA (Wyoming), has larger conidia [(10-)25-45(-55) × 2-5 μm] that are consistently septate and, although sometimes con- stricted at the septa, they are always hyaline. Ramularia pra- tensis has a worldwide distribution and produces conidia of approximately the same size, (6-)8-25(-35) × (1.5-) 2-4(-5) μm, but they are never constricted at the septa or pigmented. In addition, the conidiophores in T. rumicicola (Fig. 23) were consistently reduced to conidiogenous cells while both R. pseudodecipiens and R. pratensis produce long conidiophores. The phylogenetic analysis supports this species clade (Fig. 1, apical locus, 1 μm diam. Conidia are catenate, forming ramoconidia, intercalary conidia and terminal conidia. Conidia (type I) hyaline, thin-walled, smooth, aseptate, hila thickened and darkened, 1 μm diam.; ramoconidia subcylindrical to fusiform, (7-)8.5-10(-15) × (1-)1.5-2 μm, with two apical hila; intercalary conidia, fusiform, (6-)8-10(-17) × (1-) 1.5-2 μm, in chains of up to five conidia; terminal conidia, fusiform to obovoid, (3.5-)5-6(-7) × (1-)1.5-2 μm. Conidia (type II) not observed. ...
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... Two Ramularia species (R. geranii var. geranii, R. pseudogeranii) and two ramularia-like species (Phacellium geranii and Pseudocercosporella magnusiana) have thus far been described from Geranium (Braun 1998). Ramularia geranii var. geranii produces conidia that are smooth to verruculose, ellipsoid-ovoid to fusiform, 0-3-septate and 10-40(-55) × (2-) 2.5-6(-7) μm. Ramularia pseudogeranii produces solitary obovoid conidia, 14-25 × 6-12 μm. The synnematous Pha- cellium geranii produces catenate conidia, ellipsoid-ovoid to fusoid, 12-28 × 4-7 μm. Pseudocercosporella magnusiana produces solitary conidia, subcylindrical-filiform to acicular (30-) 40-100(-120) μm, 2-8-septate, with hyaline, unthickened hi- lum. The morphological characters of R. geraniicola (Fig. 46) are also distinct from the closest species, R. variabilis that produces shorter conidiophores and narrower fusiform to obovoid conidia (Fig. 2, clade 50). Ramularia geraniicola has unique morpho- logical characters and forms a single lineage in the phylogenetic analysis (Fig. 2 For additional synonyms see or ...
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... Two Ramularia species (R. geranii var. geranii, R. pseudogeranii) and two ramularia-like species (Phacellium geranii and Pseudocercosporella magnusiana) have thus far been described from Geranium (Braun 1998). Ramularia geranii var. geranii produces conidia that are smooth to verruculose, ellipsoid-ovoid to fusiform, 0-3-septate and 10-40(-55) × (2-) 2.5-6(-7) μm. Ramularia pseudogeranii produces solitary obovoid conidia, 14-25 × 6-12 μm. The synnematous Pha- cellium geranii produces catenate conidia, ellipsoid-ovoid to fusoid, 12-28 × 4-7 μm. Pseudocercosporella magnusiana produces solitary conidia, subcylindrical-filiform to acicular (30-) 40-100(-120) μm, 2-8-septate, with hyaline, unthickened hi- lum. The morphological characters of R. geraniicola (Fig. 46) are also distinct from the closest species, R. variabilis that produces shorter conidiophores and narrower fusiform to obovoid conidia (Fig. 2, clade 50). Ramularia geraniicola has unique morpho- logical characters and forms a single lineage in the phylogenetic analysis (Fig. 2 For additional synonyms see or ...
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... species or a collective species composed of various races since on some hosts there are morphologically similar phytopathogenic species and in other hosts this species was morphologically distinguishable from other phytopathogenic spe- cies. Ramularia coleosporii was originally described parasitising Coleosporium melampyri on Melampyrum nemorosum from France (lectotype in HAL). All the strains of R. coleosporii used in this study cluster together in the same clade (Fig. 2, clade 66, 1/ 100/100) and are clearly separated from the other Ramularia spp., supporting the hypothesis that this is indeed a unique species. They were, however, all collected from South Korea, and a few isolates from other countries should be analysed to determine if it is a global species. It was the first time that this species was observed in association with the host Pileoaria shiraiana. To determine if R. coleosporii is truly mycophylic more studies need to be done to understand the biology and ecology of this species. For additional synonyms see ) or MycoBank. Description in vivo: See Braun (1998. Substrate and distribution: On Bromus, Festuca, Glyceria, Leu- copoa, Lolium, Phalaris, and Triticale (Poaceae) and Cannabis (Cannabaceae); Europe, N. America (Canada, Mexico, USA), S. America (Chile, Colombia), Asia (Japan, Russia), Australia and New ...
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... morphologically, T. persicariae (Fig. 22) produces longer conidia than T. rumicicola. Etymology: Named after the host genus Rumex, from which it was ...
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... Notes: The strain representing this species was originally iden- tified as Phacellium paspali. The characteristic Phacellium syn- nemata are sometimes also formed in culture. This species is represented by a single basal lineage in the Teratoramularia clade in the phylogenetic analysis (Fig. 1, clade XXX). Morphologically (Fig. 24), it is nearly impossible to distinguish it from the closest sister species T. ...
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... The valid publication of Cylindrosporium grevilleanum, the basionym of Ramularia grevilleana, dates back to Oudemans (1873). A detailed discussion of the complicated nomenclature of this species and its lectotypification has been published by Braun & Pennycook (2003). The phylogenetic analyses provide high support for this species clade (Fig. 2, clade 28, 1/100/100). Ramularia grevilleana causes Ramularia leaf spot disease of strawberry and other hosts of the Rosaceae. The most con- spicuous symptoms are leaf lesions but symptoms can also develop on fruits, calyxes, fruit trusses, petioles and stolons. It occurs worldwide on cultivated varieties as well as wild straw- berry species. In earlier years, the economic impact of the dis- ease was so great that Ramularia leaf spot was considered the most important strawberry disease. With increased emphasis on the development and use of resistant cultivars, Ramularia leaf spot disease, although still an important foliar disease is now of less concern (Maas 1984). The link between the sexual morph Mycosphaerella fragariae and the asexual morph R. grevilleana has been experimentally proven (Dudley 1889). Mycelium consisting of hyaline, septate, branched, smooth, 1-3 μm diam hyphae. Conidiophores hyaline, thin-walled, smooth, erect, 1-3-septate, straight to flexuous, cylindrical- oblong, unbranched (19-)41-53(-82) × (1.5-)2-2.5(-3) μm or reduced to conidiogenous cells. Conidiogenous cells inte- grated in mycelium or terminal on conidiophores, cylindrical- oblong, (15-)21.5-25.5(-36) × 2.0-2.5(-3) μm, with 1-2 apical conidiogenous loci almost flat to short cylindrical, thick- ened, darkened, refractive. Conidia hyaline, thin-walled, smooth to slightly verruculose, with hila thickened, darkened and refractive. Ramoconidia subcylindrical to clavate, (14-) 21-27(-44) × (2.5-)3-3.5(-4) μm, 0-3-septate, with 2-3 apical hila. Intercalary conidia hyaline, smooth, 0-3-septate, subcylindrical with apices rounded and broader, (14-) 19-25(-50) × (2.5-)3-4(-4.5) μm, in branched chains of up to four conidia. Terminal conidia aseptate, subcylindrical to obovoid, (5.5-)13-16.5(-25.5) × (2-)3-3.5(-4.5) ...
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... Ramularia gei was originally described on Geum urbanum from Sweden (holotype in S). The similarities between Acrotheca gei and Ramularia gei were discussed by Hughes (1953) and those between R. gei and R. submodesta were pointed out by von Höhnel (1904). The strains included here form a highly supported clade (Fig. 2, For additional synonyms see or ...
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... strain used in this study forms a single lineage (Fig. 2, clade 88), basal to R. endophylla, and is positioned on a very long branch, which supports this species as unique. No Ramularia species is currently known from Gaultheria. Unfortu- nately, the culture was sterile and the herbarium material is an old dried culture on which some conidiophores and con- idiogenous cells could be observed but were not sufficient to warrant a full description. (Fig. 44), so the molecular differences based on the sequence data are also provided. Substrate and distribution: On Geum (Rosaceae); Asia, Cau- casus, Europe, Iceland, N. ...
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... consisting of hyaline, septate, branched, smooth, 1.5-3 μm diam hyphae. Conidiophores hyaline, thin-walled, erect, septate, straight to geniculate-sinuous, cylindrical-oblong, unbranched, (65-)82-100(-150) × 2-2.5(-3) μm or reduced to conidiogenous cells, hyaline, smooth, integrated in mycelium or terminal in conidiophores, cylindrical-oblong, (14.5-) 19-22(-28) × 2-3 μm, with one apical conidiogenous locus, (Fig. 2, clade 4). The strains CBS 159.24 and CBS 160.24 produce very large conidia fitting with the original description and were isolated from the same host genus, but from a different species and country. Strain CBS 114566 was isolated from the same host as the type but from a different European country. Unfortunately it was sterile and morphological comparison with the original description was not possible. Strains CBS 159.24 and CBS 160.24 (Fig. 2, in clade 3, 1/100/100; Fig. 45) are considered good representatives of R. geranii both morphologically and phylogenetically, and CBS 160.24 is therefore chosen as ex-epitype. The strain CBS 114566 appears as R. geranii in the CBS database but it is not conspecific with the species in this clade and will be treated as a Ramularia sp. A for the time ...
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... consisting of hyaline, septate, branched, smooth, 1.5-3 μm diam hyphae. Conidiophores hyaline, thin-walled, erect, septate, straight to geniculate-sinuous, cylindrical-oblong, unbranched, (65-)82-100(-150) × 2-2.5(-3) μm or reduced to conidiogenous cells, hyaline, smooth, integrated in mycelium or terminal in conidiophores, cylindrical-oblong, (14.5-) 19-22(-28) × 2-3 μm, with one apical conidiogenous locus, (Fig. 2, clade 4). The strains CBS 159.24 and CBS 160.24 produce very large conidia fitting with the original description and were isolated from the same host genus, but from a different species and country. Strain CBS 114566 was isolated from the same host as the type but from a different European country. Unfortunately it was sterile and morphological comparison with the original description was not possible. Strains CBS 159.24 and CBS 160.24 (Fig. 2, in clade 3, 1/100/100; Fig. 45) are considered good representatives of R. geranii both morphologically and phylogenetically, and CBS 160.24 is therefore chosen as ex-epitype. The strain CBS 114566 appears as R. geranii in the CBS database but it is not conspecific with the species in this clade and will be treated as a Ramularia sp. A for the time ...
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... forming gall-like deformations. Mycelium con- sisting of hyaline, septate, sparsely branched, thin-walled hy- phae. Conidiophores reduced to the conidiogenous cells, erumpent, usually ampulliform but sometimes subcylindrical, aseptate, hyaline, thin-walled, mono-or polyblastic, sympodial, conidiogenous loci conspicuous, thickened and darkened. Con- idia formed singly, acrogenous, oblong-clavate to subcylindrical, hyaline, thin-walled, smooth, aseptate or 1-septate, hilum con- spicuous, thickened and darkened. Notes: Hawksworthiana is monotypic and was described based on H. peltigericola on a specimen of Peltigera polydactyla from the Isle of Mull in Scotland. It forms gall-like deformations on lichens of the genus Peltigera, and has been reported from Europe and North America. Hawksworthiana differs from Ramularia by its lichenicolous habit and morphological charac- ters such as the wide ampulliform conidiogenous cells, the conidiogenous loci and hila are not refractive, the absence of stroma-like structures and the symptoms caused on the host (Fig. 25). All attempts to culture this fungus from fresh collections have thus far proven ...
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... U. Braun, Int. J. Mycol. Lichenol. 3: 276. 1988. Fig. ...
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... While Ramularia has polyblastic, sympodial and cicatrised conidiogenous cells, Monodidymaria has monophialidic con- idiogenous cells (Fig. 26). This character excludes Monodidymaria from the Cercosporella/Ramularia complex, but due to their common taxonomical history, they are still studied together. Monodidymaria is in fact morphologically more similar to Cephalosporiopsis, but the latter genus comprises saprophytic soil hyphomycetes and its taxonomic status is not yet certain ). As a consequence, the genus is maintained until more data is available to clarify its taxonomic position ). Five species are known to belong in this genus and were isolated from several hosts (Chenopodium, Equisetum, Scirpus and Vitex) from Asia, Europe, North and South America , Seifert et al. ...
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... Ramularia diervilla was originally described on Diervilla lonicera from New York, USA (holotype in NYS), and is the only species of Ramularia known to infect this host. Although it has been previously reported from North America, this is the first report from Canada. This species formed a highly supported clade (Fig. 2, clade 41, 1/100/100). reported that when associated with its host, R. diervilla produces simple, straight to geniculate-sinuous conidiophores, 5-25 × 1.5-3.5 μm, and catenate, cylindrical-fusiform conidia, 5-25(-30) × 1.5-4 μm. The conidiophores described in culture are longer and the conidia are slightly narrower than what is described in vivo (Fig. 42). In the herbarium specimen corre- sponding to the isolate CPC 16859, the conidiophores are shorter than in culture [(25-)28-30(-33) × (1.5-)2(-3) μm] but more similar to the description provided by , while the conidial dimensions [(4-)8-9(-12) × (1.5-)2(-3) μm] are smaller than in culture, but still narrower than in . The cultures and specimens represented in this clade are considered here as representative material of the species until collections from the type location are examined. ...
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... Ramularia diervilla was originally described on Diervilla lonicera from New York, USA (holotype in NYS), and is the only species of Ramularia known to infect this host. Although it has been previously reported from North America, this is the first report from Canada. This species formed a highly supported clade (Fig. 2, clade 41, 1/100/100). reported that when associated with its host, R. diervilla produces simple, straight to geniculate-sinuous conidiophores, 5-25 × 1.5-3.5 μm, and catenate, cylindrical-fusiform conidia, 5-25(-30) × 1.5-4 μm. The conidiophores described in culture are longer and the conidia are slightly narrower than what is described in vivo (Fig. 42). In the herbarium specimen corre- sponding to the isolate CPC 16859, the conidiophores are shorter than in culture [(25-)28-30(-33) × (1.5-)2(-3) μm] but more similar to the description provided by , while the conidial dimensions [(4-)8-9(-12) × (1.5-)2(-3) μm] are smaller than in culture, but still narrower than in . The cultures and specimens represented in this clade are considered here as representative material of the species until collections from the type location are examined. ...
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... U. Braun, Nova Hedwigia 58: 195. 1994. Fig. ...
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... causing leaf spots. Caespituli amphigenous, whitish to pink or ochraceous. Mycelium consisting of hyaline to faintly pigmented, septate, branched, thin-walled hyphae forming well-developed stromata. Conidiophores arising from stromata, emerging through stomata or erumpent through the cuticle, often forming sporodochia, subcylindrical, subclavate, simple, thin- walled, smooth, hyaline or lightly pigmented, continuous or septate. Conidiogenous cells integrated, terminal, straight to moderately geniculate-sinuous, polyblastic and sympodial, con- idiogenous loci numerous, conspicuous, bulging, papilla-like, but not thickened and darkened, at most slightly refractive. Conidia formed singly, subglobose, obovoid, ellipsoid, aseptate, hyaline to faintly pigmented, thin-walled, smooth to verruculose; basal hilum not thickened or darkened; conidial secession schizolytic. Adapted from Notes: Neoovularia species are characterised by having unthickened but bulging and refractive conidiogenous loci, and by producing single, subglobose conidia with unthickened but refractive hila (Fig. 27). There are six species described in this genus that are phytopathogenic and cause distinct lesions on leaves and stems ). They have been observed from hosts belonging to four different families (Asteraceae, Fabaceae, Lamiaceae and Malvaceae) and located in Europe, Asia, Cau- casus and N. ...
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... U. Braun, Nova Hedwigia 54: 473. 1992. Fig. ...
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... latifolii does not appear as synonym of any of the other varieties and should, therefore, be considered as a synonym of R. deusta var. deusta. The strain used in this study (CBS 473,50; Fig. 2, clade 62) was previously identified as R. deusta f. latifolii and was deposited by Baker and Snyder in 1950, which makes it an authentic strain and a reliable representative of the species Ramularia deusta var. deusta until fresh material from the same location and host as the type material is recollected. Denmark, designated in Braun (1998), in C]. While R. didyma var. didyma conidiophores emerge through stromata and form catenate conidia, R. didyma var. exigua exhibits conidiophores erumpent through the cuticle, and R. didyma var. pulsatillae frequently forms solitary conidia. Ramularia didyma var. didyma has a wider distribution than the other two varieties . Ramularia didyma was identified as the causal agent of leaf spotting symptoms on Persian buttercups (Ranunculus asiaticus) in USA, California. These are colourful, cool-season perennials or annuals grown for the flowers and bulbs. Introduction of this pathogen into commercial production fields could cause signifi- cant economic loss (Blomquist & Warfield 2011). The ITS sequence (GenBank HQ442297) generated at that time is 100 % similar to the ITS sequence of the strains in this clade. Based on phylogenetic analyses in this study, this species forms a highly supported clade (Fig. 2, clade 72, 1/100/100). The morphology could not be observed since the cultures were sterile and no herbarium materials corresponding to the strains were pre- served. This clade is tentatively maintained as a representative of this species until fresh material from the type host and location is recollected. Mycelium consisting of hyaline, septate, branched, smooth, 1-2 μm diam hyphae. Conidiophores hyaline, thin-walled, smooth, erect, 1-2-septate, cylindrical-oblong, straight to sinuous, unbranched, (10-)26.5-35(-54) × (1-)1.5-2(-3) μm, or reduced to conidiogenous cells, terminal on conidiophores or intermediate in the mycelium, cylindrical-oblong, narrower at the top, (5.5-)14.5-19(-29) × 1.5-2(-3) μm, with one or two conidiogenous loci almost flat to protuberant; conidiogenous loci thickened, darkened and refractive. Conidia hyaline, thin-walled, smooth, catenate, with hila thickened, darkened and refractive. Ramoconidia cylindrical-oblong, sometimes sinuous or curved, (9.5-)14-17(-26.5) × (1.5-)2-2.5(-3) μm, 0-1-septate, with 2 apical hila. Intercalary conidia cylindrical-oblong, fusoid, some- times curved, aseptate, (8-)11-13(-19.5) × (1.5-)2(-3) μm, in branched chains of up to seven conidia. Terminal conidia cylindrical-oblong to obovoid, aseptate, (5-)7-8(-11) × (1.5-) 2(-2.5) μm. Sporulating on ...
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... latifolii does not appear as synonym of any of the other varieties and should, therefore, be considered as a synonym of R. deusta var. deusta. The strain used in this study (CBS 473,50; Fig. 2, clade 62) was previously identified as R. deusta f. latifolii and was deposited by Baker and Snyder in 1950, which makes it an authentic strain and a reliable representative of the species Ramularia deusta var. deusta until fresh material from the same location and host as the type material is recollected. Denmark, designated in Braun (1998), in C]. While R. didyma var. didyma conidiophores emerge through stromata and form catenate conidia, R. didyma var. exigua exhibits conidiophores erumpent through the cuticle, and R. didyma var. pulsatillae frequently forms solitary conidia. Ramularia didyma var. didyma has a wider distribution than the other two varieties . Ramularia didyma was identified as the causal agent of leaf spotting symptoms on Persian buttercups (Ranunculus asiaticus) in USA, California. These are colourful, cool-season perennials or annuals grown for the flowers and bulbs. Introduction of this pathogen into commercial production fields could cause signifi- cant economic loss (Blomquist & Warfield 2011). The ITS sequence (GenBank HQ442297) generated at that time is 100 % similar to the ITS sequence of the strains in this clade. Based on phylogenetic analyses in this study, this species forms a highly supported clade (Fig. 2, clade 72, 1/100/100). The morphology could not be observed since the cultures were sterile and no herbarium materials corresponding to the strains were pre- served. This clade is tentatively maintained as a representative of this species until fresh material from the type host and location is recollected. Mycelium consisting of hyaline, septate, branched, smooth, 1-2 μm diam hyphae. Conidiophores hyaline, thin-walled, smooth, erect, 1-2-septate, cylindrical-oblong, straight to sinuous, unbranched, (10-)26.5-35(-54) × (1-)1.5-2(-3) μm, or reduced to conidiogenous cells, terminal on conidiophores or intermediate in the mycelium, cylindrical-oblong, narrower at the top, (5.5-)14.5-19(-29) × 1.5-2(-3) μm, with one or two conidiogenous loci almost flat to protuberant; conidiogenous loci thickened, darkened and refractive. Conidia hyaline, thin-walled, smooth, catenate, with hila thickened, darkened and refractive. Ramoconidia cylindrical-oblong, sometimes sinuous or curved, (9.5-)14-17(-26.5) × (1.5-)2-2.5(-3) μm, 0-1-septate, with 2 apical hila. Intercalary conidia cylindrical-oblong, fusoid, some- times curved, aseptate, (8-)11-13(-19.5) × (1.5-)2(-3) μm, in branched chains of up to seven conidia. Terminal conidia cylindrical-oblong to obovoid, aseptate, (5-)7-8(-11) × (1.5-) 2(-2.5) μm. Sporulating on ...
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... causing leaf spots. Mycelium consisting of hyaline or subhyaline, septate, branched, thin-walled hyphae forming stromata or not. Conidiophores macronematous, usually in large fascicles, sometimes forming sporodochial and basistromatic conidiomata, emerging through stomata or erum- pent through the cuticle, straight, subcylindrical to geniculate- sinuous, simple, hyaline or faintly pigmented, continuous or septate, thin-walled, smooth or occasionally rough. Con- idiogenous cells integrated, terminal, polyblastic, percurrent and sympodial, conidiogenous loci inconspicuous, not thickened or darkened. Conidia solitary or catenate, ellipsoid-ovoid, sub- cylindrical or fusoid, hyaline or slightly pigmented, aseptate to 3- septate, thin-walled, smooth or almost so, hila unthickened and hyaline, conidial secession schizolytic. Notes: The genus Neoramularia was introduced by Braun (1991) to include species with inconspicuous, unthickened, hyaline conidiogenous loci and hila. The circumscription of Neoramularia was later modified to include species forming catenate conidia such as Neoramularia esfandiarii (Braun 1992). Ten species are currently known in this genus and have been isolated from different hosts in Asia, Europe and North America , Seifert et al. 2011). The type species is known from Kochia sp., Azerbaijan, and a photo- plate based on the holotype of R. eurotia, a synonym of Neoramularia kochiae is presented (Fig. 28). hyphae, forming well developed stromata. Conidiomata basi- stromatic and sporodochial. Conidiophores arranged in palisade- like fascicles, subcylindrical, subclavate, straight to flexuous, sinuous, rarely septate, hyaline to faintly pigmented, thin-walled, smooth, sometimes reduced to conidiogenous cells. Con- idiogenous cells integrated, terminal, polyblastic, sympodial, conidiogenous loci bulging, unthickened or with a thickened rim, not darkened but refractive. Conidia formed singly, ellipsoid- obovoid, subclavate, aseptate to 2-septate, base rounded to broadly truncate, hyaline to faintly pigmented, thin-walled, smooth to verruculose, hilum unthickened, not darkened but refractive, conidial secession schizolytic. Notes: Pseudodydimaria was established to accommodate Didymaria wyethiae, since it did not fit comfortably with the description of Ramularia, Pseudocercosporidium or Neoovularia. Pseudocercosporidium differs by having very long, branched conidiophores, formed singly or loosely grouped. Neoovularia differs by having aseptate, subglobose to ovoid conidia with narrow, darkened, refractive hila. Two species are currently known to belong to this genus, P. wyethiae (Fig. 29) and P. clematidis, reported from North America ...
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... causing leaf spots. Mycelium consisting of hyaline or subhyaline, septate, branched, thin-walled hyphae forming stromata or not. Conidiophores macronematous, usually in large fascicles, sometimes forming sporodochial and basistromatic conidiomata, emerging through stomata or erum- pent through the cuticle, straight, subcylindrical to geniculate- sinuous, simple, hyaline or faintly pigmented, continuous or septate, thin-walled, smooth or occasionally rough. Con- idiogenous cells integrated, terminal, polyblastic, percurrent and sympodial, conidiogenous loci inconspicuous, not thickened or darkened. Conidia solitary or catenate, ellipsoid-ovoid, sub- cylindrical or fusoid, hyaline or slightly pigmented, aseptate to 3- septate, thin-walled, smooth or almost so, hila unthickened and hyaline, conidial secession schizolytic. Notes: The genus Neoramularia was introduced by Braun (1991) to include species with inconspicuous, unthickened, hyaline conidiogenous loci and hila. The circumscription of Neoramularia was later modified to include species forming catenate conidia such as Neoramularia esfandiarii (Braun 1992). Ten species are currently known in this genus and have been isolated from different hosts in Asia, Europe and North America , Seifert et al. 2011). The type species is known from Kochia sp., Azerbaijan, and a photo- plate based on the holotype of R. eurotia, a synonym of Neoramularia kochiae is presented (Fig. 28). hyphae, forming well developed stromata. Conidiomata basi- stromatic and sporodochial. Conidiophores arranged in palisade- like fascicles, subcylindrical, subclavate, straight to flexuous, sinuous, rarely septate, hyaline to faintly pigmented, thin-walled, smooth, sometimes reduced to conidiogenous cells. Con- idiogenous cells integrated, terminal, polyblastic, sympodial, conidiogenous loci bulging, unthickened or with a thickened rim, not darkened but refractive. Conidia formed singly, ellipsoid- obovoid, subclavate, aseptate to 2-septate, base rounded to broadly truncate, hyaline to faintly pigmented, thin-walled, smooth to verruculose, hilum unthickened, not darkened but refractive, conidial secession schizolytic. Notes: Pseudodydimaria was established to accommodate Didymaria wyethiae, since it did not fit comfortably with the description of Ramularia, Pseudocercosporidium or Neoovularia. Pseudocercosporidium differs by having very long, branched conidiophores, formed singly or loosely grouped. Neoovularia differs by having aseptate, subglobose to ovoid conidia with narrow, darkened, refractive hila. Two species are currently known to belong to this genus, P. wyethiae (Fig. 29) and P. clematidis, reported from North America ...
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... U. Braun, Nova Hedwigia 53: 291. 1991. Fig. ...
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... phylogenetic trees based on the multigene dataset ( Fig. 2) that were generated with BA, ML and PA separated the strains into similar species clades. The phylogeny distributed the species into three main clades, and the position of single species clades varied with each gene and each phylogenetic method. The tree depicted a total of 86 clades, of which 30 are single lineages (clades 4, 6, 11, 12, 14, 18, 22, 25, 26, 33, 34, 36, 37, 39, 43, 44, 48, 51, 62-64, 69, 73, 75, 77, 80, 82-84, 88), 20 clades represent new species (clades 1, 5, 7, 15, 20, 24, 25, 44, 51, 56, 58, 70, 76, 77, 80, 83, 84, 88), and 12 clades contained good candidates for epitypification for existing species (clades 3, 16, 27, 38-40, 48, 50, 52, 67, 78, 79). These are discussed in further detail in the Taxonomy section ...
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... on the results of MrModelTest the Bayesian analysis was performed with the GTR+I+G substitution model, with inverse gamma rates and with dirichlet base frequencies for actA, gapdh and rpb2. The ITS partition was analysed with a SYM+I+G substitution model with fixed frequencies and with inverse gamma rates while the tef1-α partition was analysed with the HKY+I+G substitution model with inverse gamma rates and with dirichlet base frequencies. The alignment contained a total of 1 476 unique site patterns: 374 (rpb2), 178 (ITS), 191 (actA), 354 (gapdh), and 379 (tef1-α). The analysis generated 17 232 trees from which 12 924 were sampled and 4 308 were discarded (25 % burnin) and the final tree is depicted in Fig. 2. The Maximum Likelihood analysis using the GTRGAMMA model detected 1 415 distinct patterns and reached a final ML optimi- sation likelihood of -62205.001171. The bootstrap support values from the best-scoring tree were mapped on the Bayesian tree as the second value in the tree nodes ( Fig. 2; bootstrap values 80 %). The parsimony analysis generated the maximum of 1 000 equally most parsimonious trees. From the 2 499 char- acters analysed, 1 068 were constant, 182 were variable and parsimony-uninformative and 1 249 were parsimony-informative. The robustness of the trees obtained was evaluated by 1000 bootstrap replications. The bootstrap support values were mapped on the Bayesian tree as the third value in the tree nodes (Fig. 2, bootstrap values 80 %). A consensus parsimony tree was calculated from the equally most parsimonious trees and the branches were mapped with a thicker stroke on the Bayesian tree ( Fig. 2). The additional parameters calculated were TL = 14589, CI = 0.213, RI = 0.827 and RC = ...
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... on the results of MrModelTest the Bayesian analysis was performed with the GTR+I+G substitution model, with inverse gamma rates and with dirichlet base frequencies for actA, gapdh and rpb2. The ITS partition was analysed with a SYM+I+G substitution model with fixed frequencies and with inverse gamma rates while the tef1-α partition was analysed with the HKY+I+G substitution model with inverse gamma rates and with dirichlet base frequencies. The alignment contained a total of 1 476 unique site patterns: 374 (rpb2), 178 (ITS), 191 (actA), 354 (gapdh), and 379 (tef1-α). The analysis generated 17 232 trees from which 12 924 were sampled and 4 308 were discarded (25 % burnin) and the final tree is depicted in Fig. 2. The Maximum Likelihood analysis using the GTRGAMMA model detected 1 415 distinct patterns and reached a final ML optimi- sation likelihood of -62205.001171. The bootstrap support values from the best-scoring tree were mapped on the Bayesian tree as the second value in the tree nodes ( Fig. 2; bootstrap values 80 %). The parsimony analysis generated the maximum of 1 000 equally most parsimonious trees. From the 2 499 char- acters analysed, 1 068 were constant, 182 were variable and parsimony-uninformative and 1 249 were parsimony-informative. The robustness of the trees obtained was evaluated by 1000 bootstrap replications. The bootstrap support values were mapped on the Bayesian tree as the third value in the tree nodes (Fig. 2, bootstrap values 80 %). A consensus parsimony tree was calculated from the equally most parsimonious trees and the branches were mapped with a thicker stroke on the Bayesian tree ( Fig. 2). The additional parameters calculated were TL = 14589, CI = 0.213, RI = 0.827 and RC = ...
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... on the results of MrModelTest the Bayesian analysis was performed with the GTR+I+G substitution model, with inverse gamma rates and with dirichlet base frequencies for actA, gapdh and rpb2. The ITS partition was analysed with a SYM+I+G substitution model with fixed frequencies and with inverse gamma rates while the tef1-α partition was analysed with the HKY+I+G substitution model with inverse gamma rates and with dirichlet base frequencies. The alignment contained a total of 1 476 unique site patterns: 374 (rpb2), 178 (ITS), 191 (actA), 354 (gapdh), and 379 (tef1-α). The analysis generated 17 232 trees from which 12 924 were sampled and 4 308 were discarded (25 % burnin) and the final tree is depicted in Fig. 2. The Maximum Likelihood analysis using the GTRGAMMA model detected 1 415 distinct patterns and reached a final ML optimi- sation likelihood of -62205.001171. The bootstrap support values from the best-scoring tree were mapped on the Bayesian tree as the second value in the tree nodes ( Fig. 2; bootstrap values 80 %). The parsimony analysis generated the maximum of 1 000 equally most parsimonious trees. From the 2 499 char- acters analysed, 1 068 were constant, 182 were variable and parsimony-uninformative and 1 249 were parsimony-informative. The robustness of the trees obtained was evaluated by 1000 bootstrap replications. The bootstrap support values were mapped on the Bayesian tree as the third value in the tree nodes (Fig. 2, bootstrap values 80 %). A consensus parsimony tree was calculated from the equally most parsimonious trees and the branches were mapped with a thicker stroke on the Bayesian tree ( Fig. 2). The additional parameters calculated were TL = 14589, CI = 0.213, RI = 0.827 and RC = ...
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... on the results of MrModelTest the Bayesian analysis was performed with the GTR+I+G substitution model, with inverse gamma rates and with dirichlet base frequencies for actA, gapdh and rpb2. The ITS partition was analysed with a SYM+I+G substitution model with fixed frequencies and with inverse gamma rates while the tef1-α partition was analysed with the HKY+I+G substitution model with inverse gamma rates and with dirichlet base frequencies. The alignment contained a total of 1 476 unique site patterns: 374 (rpb2), 178 (ITS), 191 (actA), 354 (gapdh), and 379 (tef1-α). The analysis generated 17 232 trees from which 12 924 were sampled and 4 308 were discarded (25 % burnin) and the final tree is depicted in Fig. 2. The Maximum Likelihood analysis using the GTRGAMMA model detected 1 415 distinct patterns and reached a final ML optimi- sation likelihood of -62205.001171. The bootstrap support values from the best-scoring tree were mapped on the Bayesian tree as the second value in the tree nodes ( Fig. 2; bootstrap values 80 %). The parsimony analysis generated the maximum of 1 000 equally most parsimonious trees. From the 2 499 char- acters analysed, 1 068 were constant, 182 were variable and parsimony-uninformative and 1 249 were parsimony-informative. The robustness of the trees obtained was evaluated by 1000 bootstrap replications. The bootstrap support values were mapped on the Bayesian tree as the third value in the tree nodes (Fig. 2, bootstrap values 80 %). A consensus parsimony tree was calculated from the equally most parsimonious trees and the branches were mapped with a thicker stroke on the Bayesian tree ( Fig. 2). The additional parameters calculated were TL = 14589, CI = 0.213, RI = 0.827 and RC = ...
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... Ramularia collo-cygni was originally isolated from the host Hordeum vulgare collected in Italy, but the type specimen is pre- sumed missing ). The strains used in this study cluster together in a highly supported clade (Fig. 2, clade 38, 1/100/100) and a strain isolated from the same host that was collected in Austria is designated as neotype. Ramularia collo-cygni is the causal agent of Ramularia leaf spot disease on barley, a disease that has been known for more than 100 years but of which the importance has only been recognised in the last 30 years. The disease has been reported worldwide and on various cereals and grasses. On barley, the symptoms appear late in the season as reddish brown necrotic spots that lead to premature leaf senes- cence and subsequent grain yield loss. Environmental conditions such as temperature and humidity are key factors in activating the production of rubellin, a non-host specific toxin, by the fungus. The development of molecular diagnostic tools has improved the detection of the pathogen in plant tissue and seeds before symptom development. Vertical transmission of the fungus in barley has been confirmed ( Havis et al. 2015) and further evidence points to the existence of an endophytic life-style that shifts towards necrotrophy depending on plant health. Population studies using simple-sequence repeat markers and sequence analyses of housekeeping genes revealed a high genetic diversity in R. collo- cygni isolates ( Piotrowska 2014, Havis et al. 2015. A high level of genotypic diversity is usually indicative of sexual recombination, but the sexual morph of R. collo-cygni is yet to be identified. Control of the disease can be accomplished by timely fungicide application between the Zadoks growth stages (ZGS) 30 and ZGS 49, well before the symptoms develop that usually happens at stage ZGS 70. Ramularia collo-cygni has lost sensitivity to strobilurin-based fungicides due to the development of the G143A point mutation in the cytochrome b gene, which is now prevalent in most pop- ulations. The introduction of a new generation of SDHI fungicide has brought some leverage in disease control, but the rapid evolutionary potential displayed by this fungus suggests it can adapt to new control strategies quickly. Despite all the research performed so far several questions still need to be addressed to fully understand the biology of this species in order to develop appropriate control measures ( Havis et al. 2015). Ramularia collo- cygni also causes Tan Leaf Spot on turfgrass. Turfgrasses are used to control water and wind erosion of soil, and are used as ornamental plants and as ground cover of playing fields in many sports. The disease has been reported from Australia, Japan, New Zealand and North America ( Smiley et al. ...
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... Ramularia coryli was originally described on Coryllus avellana from France, and is currently the only Ramularia spe- cies known to infect this host ). The strain used in this study forms a single lineage (Fig. 2, clade 83), and is positioned on a very long branch, which supports this species as unique. Unfortunately, this strain proved to be sterile, and thus we could not compare it morphologically. This clade is for now maintained as representative of R. coryli until fresh material is collected and more information becomes available. Substrate and distribution: On Carpesium, Codonocephalum, Inula, and Pulicaria (Asteraceae); Asia, Caucasus, ...
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... Ramularia abscondita was originally described on Arctium lappa from France (lectotype in PC). It has a wide geographical distribution but has only been isolated from hosts belonging to the genus Arctium (Asteraceae), a plants genus commonly known as thistle burdock. Burdock root was used as bittering agent of beer before the introduction of hop and is very much used in Asian cuisine. Ramularia abscondita has been reported on Arctium tomentosum from Sweden (Braun 1998). The morphological description of this strain (Fig. 31) differs from the one in literature ) based on collections in vivo by having longer co- nidiophores and narrower conidiophores and conidia. These dif- ferences may be related to the fungus growing in culture and not being associated with its host. This strain forms a single lineage in the phylogenetic analysis (Fig. 2, clade 43) but will be tentatively considered as a good representative of this species until material from the type host and location is collected and cultured. Substrate and distribution: On Ranunculus (Ranunculaceae); Asia, Europe and N. ...
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... this study we applied the Consolidated Species Concept ( Quaedvlieg et al. 2014), a polyphasic approach combining the concordance of multiple gene genealogies with morphological and ecological information to improve fungal species delimitation. The genera mapped in Fig. 1 are discussed by clade order followed by a section describing and illustrating the allied genera of Ramularia for which only herbarium specimens were avail- able. The species of Ramularia resolved in Fig. 2 are discussed in alphabetical order in a third section to which a few important species not known from culture but of phytopathological impor- tance were ...
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... The strains in this clade were previously identified as R. didyma. Authentic strains of R. didyma cluster in clade 72 ( Fig. 2) and, besides R. didyma, R. acris is also common on Ranunculus acris. Since the strains were sterile in culture and no herbarium specimens were preserved, the morphological char- acters could not be compared with the description available in literature. The strains in this clade cluster in a highly supported clade (Fig. 2, Specimens examined: Greece, Macedonia region, Kozani, on leaves of Centaurea solstitialis, 28 Apr. 2004, D. Berner, culture CBS 120253 = BPI 844247. Turkey, Ankara, on Acroptilon repens, 14 Jul 1947 (lectotype W, No. 11177); near Isparta, on Acroptilon repens, 1 Sep. 1997, R. Sobhian (epitype designated here BPI 745883, MBT130827, culture ex-epitype CBS 120252). USA, California, on Cynara cardunculus, Oct. 2010, L. Davenport, cultures CPC 18723, CPC 18724. Substrate and distribution: On Acroptilon repens, Centaurea solstitiales and Cynara cardunculus (Asteraceae); Central Asia, Europe and N. ...
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... The strains in this clade were previously identified as R. didyma. Authentic strains of R. didyma cluster in clade 72 ( Fig. 2) and, besides R. didyma, R. acris is also common on Ranunculus acris. Since the strains were sterile in culture and no herbarium specimens were preserved, the morphological char- acters could not be compared with the description available in literature. The strains in this clade cluster in a highly supported clade (Fig. 2, Specimens examined: Greece, Macedonia region, Kozani, on leaves of Centaurea solstitialis, 28 Apr. 2004, D. Berner, culture CBS 120253 = BPI 844247. Turkey, Ankara, on Acroptilon repens, 14 Jul 1947 (lectotype W, No. 11177); near Isparta, on Acroptilon repens, 1 Sep. 1997, R. Sobhian (epitype designated here BPI 745883, MBT130827, culture ex-epitype CBS 120252). USA, California, on Cynara cardunculus, Oct. 2010, L. Davenport, cultures CPC 18723, CPC 18724. Substrate and distribution: On Acroptilon repens, Centaurea solstitiales and Cynara cardunculus (Asteraceae); Central Asia, Europe and N. ...
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... actinidiae Ablak., Bot. Mater. Otd. Sporov. Rast. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 13: 244. 1960. Fig. ...
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... The description of R. actinidiae available in literature in- cludes greyish caespituli, conidiophores, hyaline, simple, 20-40 μm long, conidia cylindrical, aseptate, 10.5-17 × 3 μm. The specimen observed has short conidiophores (12-) 19-23(-35) × (1.5)-2-2.5(-3) μm and conidial dimensions matching the ones in culture (5-)10-12(-21) × (1.5-)2(-3) μm. Ramularia actinidiae was originally described on Actinidia poly- gama from Russia. Braun (1998) commented on this species being insufficiently known, and that the type material was not available for study. The strain used in this study is from a different location to that of R. actinidiae but the description of the morphology is quite similar (Fig. 32). It forms a single lineage supported by the Bayesian multigene analysis (Fig. 2, clade 77) and is tentatively maintained as Ramularia actinidiae until fresh material from the same location and host as the type has been recollected (Actinidia polygama, ...
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... The description of R. actinidiae available in literature in- cludes greyish caespituli, conidiophores, hyaline, simple, 20-40 μm long, conidia cylindrical, aseptate, 10.5-17 × 3 μm. The specimen observed has short conidiophores (12-) 19-23(-35) × (1.5)-2-2.5(-3) μm and conidial dimensions matching the ones in culture (5-)10-12(-21) × (1.5-)2(-3) μm. Ramularia actinidiae was originally described on Actinidia poly- gama from Russia. Braun (1998) commented on this species being insufficiently known, and that the type material was not available for study. The strain used in this study is from a different location to that of R. actinidiae but the description of the morphology is quite similar (Fig. 32). It forms a single lineage supported by the Bayesian multigene analysis (Fig. 2, clade 77) and is tentatively maintained as Ramularia actinidiae until fresh material from the same location and host as the type has been recollected (Actinidia polygama, ...
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... Ramularia agastaches was originally described on Agastache rugosa from Japan and was synonymised with R. lamii var. lamii by Braun (1998) who was not able to examine the type specimen. The strains in this clade were previously identified as R. lamii, which is now restricted to species in clade 67 (Fig. 2). The strains in this clade form a highly supported clade by all three methods of phylogenetic analysis (Fig. 2, clade 46, 1/100/100). Morphologically (Fig. 33) the description does not match that of R. lamii available in literature ...
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... Ramularia agastaches was originally described on Agastache rugosa from Japan and was synonymised with R. lamii var. lamii by Braun (1998) who was not able to examine the type specimen. The strains in this clade were previously identified as R. lamii, which is now restricted to species in clade 67 (Fig. 2). The strains in this clade form a highly supported clade by all three methods of phylogenetic analysis (Fig. 2, clade 46, 1/100/100). Morphologically (Fig. 33) the description does not match that of R. lamii available in literature ...
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... Ramularia tovarae was originally described on Poly- gonum filiforme from Japan and its distribution was limited to the type location. The only strain available representative of this species formed a single lineage (Fig. 2, clade 37), but positioned on a long branch that supports this species as unique. Although the strain did not sporulate in culture, the morphology observed in vivo corresponded to that described in literature ). Therefore, this specimen is considered as a good representative (lectotype in HAL). Based on phylogenetic analyses in this study, strain CBS 113981 clustered within the Ramularia clade (Fig. 1, clade XIV) and formed a single lineage (Fig. 2, clade 64) in the multigene phylogeny. Since Phacellium is now considered a synonym of Ramularia, a new combination is proposed. Because the epithet "veronicae" is already occupied in Ramularia for a different species (Fig. 2, clade 64) the new epithet "veroniciola" is introduced. Ramularia veroniciola is the causative agent of leaf spot disease on Veronica species that are perennial plants used as ornamentals. The pathogen cau- ses brown roundish spots and develops conidiophores aggre- gated in synnemata. This species has been observed in several European countries and also in North America (Canada) ). It has recently been reported from China infecting V. sibirica and, although the disease incidence was low, it may become significant with the increase of the cultivated area ( Bai et al. 2013; ITS sequence GenBank HE995799). During recent field surveys in Hungary, the disease incidence affecting V. spicata and V. spuria varied between 90-100 %, and reached a severity between 30-60 % (Horv at et al. 2015; ITS sequences GenBank HQ690097 and JQ920427). The ITS sequence of the isolate CBS 113981 is identical to GenBank JQ920427, and differs from GenBank HQ690097 in 1 nucleo- tide and from GenBank HE995799 in 10 nucleotides. Unfortu- nately this strain did not sporulate in culture, and the corresponding herbarium specimen was not ...
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... Ramularia tovarae was originally described on Poly- gonum filiforme from Japan and its distribution was limited to the type location. The only strain available representative of this species formed a single lineage (Fig. 2, clade 37), but positioned on a long branch that supports this species as unique. Although the strain did not sporulate in culture, the morphology observed in vivo corresponded to that described in literature ). Therefore, this specimen is considered as a good representative (lectotype in HAL). Based on phylogenetic analyses in this study, strain CBS 113981 clustered within the Ramularia clade (Fig. 1, clade XIV) and formed a single lineage (Fig. 2, clade 64) in the multigene phylogeny. Since Phacellium is now considered a synonym of Ramularia, a new combination is proposed. Because the epithet "veronicae" is already occupied in Ramularia for a different species (Fig. 2, clade 64) the new epithet "veroniciola" is introduced. Ramularia veroniciola is the causative agent of leaf spot disease on Veronica species that are perennial plants used as ornamentals. The pathogen cau- ses brown roundish spots and develops conidiophores aggre- gated in synnemata. This species has been observed in several European countries and also in North America (Canada) ). It has recently been reported from China infecting V. sibirica and, although the disease incidence was low, it may become significant with the increase of the cultivated area ( Bai et al. 2013; ITS sequence GenBank HE995799). During recent field surveys in Hungary, the disease incidence affecting V. spicata and V. spuria varied between 90-100 %, and reached a severity between 30-60 % (Horv at et al. 2015; ITS sequences GenBank HQ690097 and JQ920427). The ITS sequence of the isolate CBS 113981 is identical to GenBank JQ920427, and differs from GenBank HQ690097 in 1 nucleo- tide and from GenBank HE995799 in 10 nucleotides. Unfortu- nately this strain did not sporulate in culture, and the corresponding herbarium specimen was not ...
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... Ramularia tovarae was originally described on Poly- gonum filiforme from Japan and its distribution was limited to the type location. The only strain available representative of this species formed a single lineage (Fig. 2, clade 37), but positioned on a long branch that supports this species as unique. Although the strain did not sporulate in culture, the morphology observed in vivo corresponded to that described in literature ). Therefore, this specimen is considered as a good representative (lectotype in HAL). Based on phylogenetic analyses in this study, strain CBS 113981 clustered within the Ramularia clade (Fig. 1, clade XIV) and formed a single lineage (Fig. 2, clade 64) in the multigene phylogeny. Since Phacellium is now considered a synonym of Ramularia, a new combination is proposed. Because the epithet "veronicae" is already occupied in Ramularia for a different species (Fig. 2, clade 64) the new epithet "veroniciola" is introduced. Ramularia veroniciola is the causative agent of leaf spot disease on Veronica species that are perennial plants used as ornamentals. The pathogen cau- ses brown roundish spots and develops conidiophores aggre- gated in synnemata. This species has been observed in several European countries and also in North America (Canada) ). It has recently been reported from China infecting V. sibirica and, although the disease incidence was low, it may become significant with the increase of the cultivated area ( Bai et al. 2013; ITS sequence GenBank HE995799). During recent field surveys in Hungary, the disease incidence affecting V. spicata and V. spuria varied between 90-100 %, and reached a severity between 30-60 % (Horv at et al. 2015; ITS sequences GenBank HQ690097 and JQ920427). The ITS sequence of the isolate CBS 113981 is identical to GenBank JQ920427, and differs from GenBank HQ690097 in 1 nucleo- tide and from GenBank HE995799 in 10 nucleotides. Unfortu- nately this strain did not sporulate in culture, and the corresponding herbarium specimen was not ...
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... Ramularia rubella was originally described on Rumex aquaticus from Germany, but it has a wide geographical distri- bution in association with the host Rumex, while it is very rarely observed infecting Polygonum s. lat. (Braun 1998). As a necrotroph, Ramularia rubella shows promise as a biological control agent against Rumex obtusifolius by causing severe defoliation, shoot and root weight loss (Huber-Meinicke et al. 1989). The available strains form a highly supported clade based on the employed phylogenetic methods (Fig. 2, clade 79, 1/100/100). The morphological description of the isolates (Fig. 61) in this clade is in agreement with the one presented in literature ), except the conidiophores were reduced to conidiogenous cells in culture. Because of the long, solitary conidia and sometimes broad conidiogenous loci and hila, some of the strains were initially confused with Cercosporella. ...
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... ≡ Ovularia rufibasis (Berk. & Broome) Massee, Brit. fung.-fl. 3: 322. 1893. ≡ Phacellium rufibasis (Berk. & Broome) U. Braun, Nova Hedwigia 54: 471. 1992. = Ramularia destructiva W. Phillips & Plowr., Grevillea 6(37): 22. 1877. = Ovularia monilioides Ellis & G. Martin, Amer. Naturalist 19: 76. 1885. Description in vivo: See Braun (1998 Substrate and distribution: On Comptonia and Myrica (Myr- icaceae); Asia, Canary Islands, Europe, N. America) Notes: Ramularia destructiva, described on Myrica gale from England (holotype in K), was reassigned to the genus Pha- cellium as Phacellium rufibasis (Braun 1992) due to the pro- duction of synnematous conidiophores. The genus Phacellium is now considered a synonym of Ramularia as the production of synnemata was deemed as an unreliable character to separate these two genera. The strain used in this study clusters within the genus Ramularia (Fig. 1, clade XIV), and formed a single lineage (Fig. 2, clade 82) basal to R. nyssicola (clade 81), but positioned on a very long branch, which sup- ports this species as unique. Unfortunately, the strain was sterile in culture and morphological data could not be evalu- ated. This lineage is for now maintained as a representative of R. rufibasis, until fresh material is collected and more infor- mation becomes available. This species causes the Ramularia dieback disease of Myrica faya in its natural habitat, affecting young shoots and causing leaf spots (Gardner & Hodges 1990). Myrica faya is considered an invasive plant in Hawaii and this pathogen represents a potentially good biocontrol agent, but no studies for field applications have been con- ducted thus ...
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... Ramularia agrimoniae was originally described on Agrimo- nia sp. from Siberia, Russia. Despite the reported distribution of this species across Europe and Asia, the available strains in this study all originate from South Korea (Fig. 2, Shin & J.D. Kim, Mycotaxon 81: 341. 2002, non Ramularia alangii Hasija, ...
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... was based on the phylogenetic position of the ex-type culture ( Videira et al. 2015b), which in this study is located in clade 63 (Fig. 2). (Fig. 2, Mycelium consisting of hyaline, septate, branched, smooth, 1-2 μm diam hyphae. Conidiophores reduced to conidiogenous cells, hyaline, thin-walled, smooth, integrated in mycelium or terminal in conidiophores, cylindrical-oblong and narrower at the top, (5.5-)12-15(-27) × (1.5-)2(-3) μm, with 1-2 apical conidiogenous loci almost flat to short cylindrical; conidiogenous loci thickened, darkened, refractive, 1 μm diam. Ramoconidia hyaline, thin-walled, smooth, subcylindrical to obclavate, (5-) 8.5-11(-19) × (2-)2.5-3(-4) μm, aseptate to 1-3-septate, with 2-3 apical hila. Intercalary conidia hyaline, smooth, aseptate or 1-3-septate, subcylindrical with apices rounded and broader, (5-)7-8(-11.5) × 2-2.5(-3) μm, in branched chains of up to seven conidia. Terminal conidia, hyaline, smooth, aseptate, obovoid, (3-)4.5-5(-6) × 2-2.5(-3) μm, hila thickened, dark- ened, refractive, 1 μm ...
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... was based on the phylogenetic position of the ex-type culture ( Videira et al. 2015b), which in this study is located in clade 63 (Fig. 2). (Fig. 2, Mycelium consisting of hyaline, septate, branched, smooth, 1-2 μm diam hyphae. Conidiophores reduced to conidiogenous cells, hyaline, thin-walled, smooth, integrated in mycelium or terminal in conidiophores, cylindrical-oblong and narrower at the top, (5.5-)12-15(-27) × (1.5-)2(-3) μm, with 1-2 apical conidiogenous loci almost flat to short cylindrical; conidiogenous loci thickened, darkened, refractive, 1 μm diam. Ramoconidia hyaline, thin-walled, smooth, subcylindrical to obclavate, (5-) 8.5-11(-19) × (2-)2.5-3(-4) μm, aseptate to 1-3-septate, with 2-3 apical hila. Intercalary conidia hyaline, smooth, aseptate or 1-3-septate, subcylindrical with apices rounded and broader, (5-)7-8(-11.5) × 2-2.5(-3) μm, in branched chains of up to seven conidia. Terminal conidia, hyaline, smooth, aseptate, obovoid, (3-)4.5-5(-6) × 2-2.5(-3) μm, hila thickened, dark- ened, refractive, 1 μm ...
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... Ramularia macrospora was described as a pathogen on Campanula pyramidalis from Germany (iconotype Pl. XI, figs 29-32). The strains used in this study cluster in the Ramularia clade (Fig. 1, clade XIV) but were not used in the multigene analysis because it was not possible to amplify and sequence the tef1-α partial gene. Although R. macrospora is usually associ- ated with members of the Campanulaceae (Braun 1998), it was recently observed infecting a host from the Aristolociaceae ( Mukhtar et al. 2012 Note: Ramularia malicola formed a single lineage (Fig. 2, clade 80) that is sister to R. rubella (Fig. 2, clade 79). This species is morphologically similar to R. rubella but differs by forming wider conidia (Fig. 57). Ramularia malicola was first isolated in a study related to sooty blotch and flyspeck on ...
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... Ramularia macrospora was described as a pathogen on Campanula pyramidalis from Germany (iconotype Pl. XI, figs 29-32). The strains used in this study cluster in the Ramularia clade (Fig. 1, clade XIV) but were not used in the multigene analysis because it was not possible to amplify and sequence the tef1-α partial gene. Although R. macrospora is usually associ- ated with members of the Campanulaceae (Braun 1998), it was recently observed infecting a host from the Aristolociaceae ( Mukhtar et al. 2012 Note: Ramularia malicola formed a single lineage (Fig. 2, clade 80) that is sister to R. rubella (Fig. 2, clade 79). This species is morphologically similar to R. rubella but differs by forming wider conidia (Fig. 57). Ramularia malicola was first isolated in a study related to sooty blotch and flyspeck on ...
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... Ramularia bellunensis was described in 1879 from Tanacetum parthenium, from Italy [lectotype, designated in Braun (1998), in PAD]. This species is represented in a single lineage (Fig. 2, clade 75). This is a new report for the Netherlands and New Zealand, as well as a first report on the host Argyr- anthemum frutescens . Although strain CBS 118417 is not in the phylogenetic trees, it is identical to CBS 116.43, except on one nucleotide in gapdh and one nucleotide in tef1-α. It was not included because at the time the trees were prepared we did not possess all the gene sequences. No ex-type strain of this species is available, and collections on Tanacetum from Italy are required to resolve its identity. Mycelium consisting of hyaline, septate, branched, smooth, 1-2 μm diam hyphae. Conidiophores hyaline, thin-walled, smooth, erect, 1-2(-4)-septate, straight to sinuous, cylindrical- oblong, unbranched (19.5-)43-58(-83) × 2-2.5(-3) μm or reduced to conidiogenous cells. Conidiogenous cells integrated in mycelium or terminal in conidiophores, cylindrical-oblong and narrower at the top, (7-)16.5-20(-30) × 2-2.5(-3) μm, with 1-4 apical conidiogenous loci, almost flat or protuberant, thickened, darkened and refractive. Conidia hyaline, thin-walled, smooth to slightly verruculose, catenate, with hila thickened, darkened and refractive. Ramoconidia subcylindrical to clavate, (8-)12-14.5(-22) × 3-4 μm, 0-1-septate, with two apical hila. Intercalary conidia subcylindrical, sometimes curved, ovoid, 0-1- septate, (8.5-)10.5-12.5(-20) × (2.5-)3(-4) μm, in branched ...
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... characterised by forming synnematous conidiomata that can be hyaline or slightly pigmented. The Phacellium strains in this study cluster within Ramularia (Fig. 1, clade XIV; Fig. 2, clade 64, clade 82) and a new Ramularia species that forms synnemata is described (Fig. 2, clade 76). These results support the hypothesis that, as in Pseudocercospora, synnematous conidiophores is a feature that is unreliable at generic level. Therefore, the genus Phacellium is tentatively synonymised with Ramularia until the exact phylogenetic position of its type species becomes known. Etymology: Named after its morphological similarity to the genus Ramularia, composed of xeno-(xenos, Greek for strange) and the latter genus ...
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... Ramularia armoraciae was first described on Armoracia rusticana from Germany (lectotype in HAL). Strain CBS 241.90 originates from the same country and was isolated from the same host as the holotype, and is therefore chosen as ex-epitype (Fig. 37). Phylogenetically, this species is highly supported by the BA and ML analysis (Fig. 2, clade 55). Substrate and distribution: On Aster, Galatella, Grindelia, Het- eropappus, Rudbeckia, Solidago (Asteraceae); Asia, Europe, N. ...
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... Ramularia ligustrina was described as a pathogen on Ligustrum vulgare in France, but was considered doubtful by Braun (1998) since type material or other collections agreeing with the description could not be traced. The species is insuffi- ciently known but the name is tentatively accepted here given its distinct phylogeny (Fig. 2 Substrate and distribution: On Adenophora, Asyneuma, Campanula, Gadellia, Legousia, and Phyteuma (Campanula- ceae), Aristolochia punjabensis (Aristolochiaceae); Asia, Cau- casus, Europe, N. America, ...
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... Braun (1998) used the name Ramularia plantaginis Ellis & G. Martin for Ramularia on Plantago major and other species characterised by verruculose conidia. This was based on the wrong assumption that R. plantaginis Peck was also published in 1882, which is, however, not correct since the latter name was published in 1880, which makes R. plantaginis Ellis & G. Martin an illegitimate homonym. Thus, R. kriegeriana is the oldest valid name for this fungus. This species has previously been reported from South Korea on Plantago asiatica, and is known from several Plantago species in Asia, Europe and N. America, including P. asiatica from China (Braun 1998). The strains in this study cluster together in a highly supported clade (Fig. 2, clade 65, 1/100/100) although a collection from Germany is required to fix the application of this name. Plantago asiatica is phyloge- netically close to P. major, the principal host of Ramularia krie- geriana. The two species belong in Plantago subgen. Plantago, in contrast to P. lanceolata, the principal host of R. rhabdospora, which belongs in subgen. Psyllium (Rønsted et al. 2002). For additional synonyms see ...

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... Filiella pastinacae was originally described on Pastinaca sativa L. from Finland, but it can infect various Apiaceous hosts worldwide. The Turkish specimen Filiella pastinacae morphologically fits to literature (Braun 1995, Bakalova and Borisova 2010, Videira et al. 2016), but conidia are slightly narrower and shorter. (Crous et al. 2013). ...
... The specimen was collected from the Yedigöller area morphologically fit to described by a previous study Crous 2005, Crous et al. 2013), but differed in having slightly narrower and fewer septa. It was originally described on Lotus uliginosus Schkuhr but later is able to infect Fabaceae members in the world (Videira et al. 2016). The Turkish specimen agrees with other species descriptions concerning the morphology of caespituli, conidiophores, and conidia, the only observable difference being in the size of conidia and conidiophores. ...
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Sixteen cercosporoid and ramularioid species belonging to nine genera are reported from Yedigöller National Park, Bolu Province, Türkiye. The species were identified based on conventional methods. Of these, Cercospora datiscicola, Cercospora mercurialis, Filiella pastinacae, Pseudocercospora sambucigena, Ramularia linaria, and Ramularia sphaeroidea are new records for Türkiye mycobiota. The description of the new recorded species are presented and discussed with their macro and micromorphological photographs. Özet Bolu ili Yedigöller Milli Parkı'ndan 9 cinse ait 16 cercosporoid ve ramularioid tür kaydedilmiştir. Tür teşhisleri geleneksel yöntemlere dayanarak yapılmıştır. Tespit edilen türlerden Cercospora datiscicola, Cercospora mercurialis, Filiella pastinacae, Pseudocercospora sambucigena, Ramularia linaria ve Ramularia sphaeroidea Türkiye mikobiyotası için yeni kayıttır. Yeni kaydedilen türlerin deskripsiyonları, makro ve mikromorfolojik fotoğrafları ile birlikte verilmiş ve tartışılmıştır.
... Ramularia is a rich genus that includes important plant pathogens such as R. collocygni and R. beticola, which caused severe economic losses to barley and sugar beet crops, respectively [16]. Ramularia leaf spot is an important disease on wheat, turfgrass, and perennial grasses and forms a potential threat to other agriculturally important crops [17]. ...
... The 28S rRNA gene (LSU), internal transcribed spacer regions (ITS), calmodulin (cmdA), translation elongation factor 1-α (tef1-α), histone H3 (his3), glyceraldehyde-3-phosphate dehydrogenase (gapdh), and RNA polymerase II second largest subunit (rpb2) genes were amplified and sequenced with the primers LR5/LSU1Fd [36,37], ITS4/V9G [38,39], CAL-228F/CAL-737R [40], EF1- 728F/TEF-1R [41,42], CylH3F/CylH3R [43,44], GPD1/GPD2 [45], and RPB2-5f2/RPB2-7 [46,47], respectively. PCR was performed as previously described by Videira [16]. ...
... All reference sequences were downloaded from GenBank and listed in Supplementary Table S1 [49][50][51][52][53], as described by Videira et al. [16]. Each sequence was aligned using MEGA 7.0.2. ...
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... The endophytic fungi genera Ramularia and Taphrina were also more abundant in grape samples with high levels of downy mildew. Ramularia, the white mold of plants, is a species-rich genus that harbors plant pathogens responsible for yield losses in many important crops, including barley, sugar beet, and strawberries [62]. It has been shown that barley plants with mlo resistance to downy mildew have an increased susceptibility to a new important disease-ramularia leaf spot [63]. ...
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... Global cotton yield has been affected by R. gossypii since 1883 7 . Historically, this species is most widespread and economically important in Brazil, although it has been mostly identified based upon morphological data 4,15 . Although conidiophore length is useful for separating Ramulariopsis species, taxonomic expertise is required. ...
... Although conidiophore length is useful for separating Ramulariopsis species, taxonomic expertise is required. Interestingly, several isolates previously putatively identified as R. gossypii were molecularly identified as either R. gossypii or R. pseudoglycines 15 . Comparison of ITS sequences of the Ramulariopsis isolates deposited in Gen-Bank (nucleotide matrices and phylogenetic tree available in TreeBASE; study number S28159) revealed that R. gossypii and R. pseudoglycines were described in previous studies 17,44 . ...
... The microscopical characteristics were analyzed by mounting asexual structures in clear lactoglycerol, and 50 measurements for each morphological parameter were carried out at a magnification of × 1,000 using a Leica DM2500 light microscope equipped with a Leica DFC 490 digital camera, coupled to a computer containing the Leica Qwin-Plus software. The morphological characteristics of the isolates were compared with the description of R. gossypii and R. pseudoglycines 14,15 . ...
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Whilst Brazil is the fourth largest cotton producer globally, incidence of ramularia leaf spot (RLS) has decreased yield. In 2017–18 and 2018–19, ca. 300 fungal samples were collected throughout Brazil. Hyphal tip cultures were obtained for amplification of the RNA polymerase II (RPB2), 28S rRNA, the ribosomal DNA internal transcribed spacers (ITS), actin (ACT), elongation factor (EF1-α) and histone H3 (HIS3) genomic regions. Additionally, sequences of the glyceraldehyde-3-phosphate dehydrogenase (GAPDH) were obtained by nanopore sequencing and the EF1-α region was selected as a marker for rapid recognition of Ramulariopsis species. Clade assignments based on the concatenated-sequence tree were identical to those in tree generated by RPB2-sequences, as well as in an RPB2 haplotype network and an ISSR (TGTC)⁴ dendrogram, in identification with species-specific primers and based on morphological comparisons. Out of 267 examined isolates, 252 were identified as Ramulariopsis pseudoglycines, indicating this species as the most widespread causal agent of cotton RLS in the Brazilian growing regions. Species-specific primers developed in the study that target the EF1-α gene provide an opportunity for extensive RLS sampling worldwide to study the distribution of Ramulariopsis species. Such data will aid breeders and plant pathologists in cotton disease resistance development and fungicide resistance avoidance.
... Hence, caution should be exercised when using small peptides as foliar fertilizers to prevent such risks [25,26]. Small peptides signi cantly inhibited the proliferation of Plectosphaerella and Coniothyrium compared to the control group. ...
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... Accordingly, numerous species have been transferred repeatedly from one of these genera to the other. The taxonomic problems related to this complex were extensively discussed in several studies (Hughes 1949;Sutton and Waller 1988;Braun 1990Braun , 1991aBraun , b, 1994aBraun , 1995Braun , 1998Verkley et al. 2004;Kirschner 2009;Videira et al. 2015Videira et al. , 2016Videira et al. , 2017. ...
... The ultrastructure of the conidiogenous loci are smooth and shaped as a truncated cone in Cercosporella, while a cladosporium-type circular rim with a central dome is seen in Ramularia (Kirschner 2009;Bensch et al. 2012). Cercosporella produces cup-shaped appressoria which are lacking in Ramularia (Kirschner 2009;Videira et al. 2016). Pseudocercosporella is characterised by unthickened and inconspicuous conidial loci and hila (Deighton 1973;Braun 1995;Frank et al. 2010). ...
... Pseudocercosporella is characterised by unthickened and inconspicuous conidial loci and hila (Deighton 1973;Braun 1995;Frank et al. 2010). (Videira et al. 2016). The phylogenetic placement of Ramularia and allied genera within the order Capnodiales was established by Videira et al. (2016Videira et al. ( , 2017 using polyphasic approaches based on multi-locus DNA sequences and morphological and cultural data. ...
Article
A colourless hyphomycetous fungus was found on living leaves of Peristrophe bicalyculata in India. Molecular phylogenetic analyses of several loci (LSU-RPB2-ITS) indicated a relationship with cercosporoid fungi (Mycosphaerellaceae). Since there is no other known lineage with similar morphology or DNA sequences, the new genus Neocercosporella is proposed with N. peristrophes comb. nov., based on Cercosporella peristrophes as type species. Pseudocercosporella andrographidis is recognized as new synonym. Similar ultrastructure of conidiogenous loci and hila confirm that N. peristrophes and P. andrographidis are conspecific. The conidiogenous loci are conical and have a small rim-like depression on the top encircling a small flat protuberant structure, which differs from other closely related members of the ramularioid complex. Neocercosporella and Cercosporella s. str. are paraphyletic as are several other genera in the Mycosphaerellaceae. Accordingly, we propose three new genera and four new combinations. The new genera are Neoacervuloseptoria gen. nov., Neocercosporella gen. nov. and Neoramulariopsis gen. nov. The new combinations are Neoacervuloseptoria fraxini comb. nov., Neocercosporella peristrophes comb. nov., Neoramulariopsis catenulata comb. nov. and Neoramulariopsis unguis-cati comb. nov.