Videira, S.I.R.; Groenewald, J.Z.; Nakashima, C.; Braun, U.; Barreto, R.W.; de Wit, P.J.G.M.; Crous, P.W. 2017: Mycosphaerellaceae – Chaos or clarity?. Studies in Mycology 87: 257-421.
![Fig. 31. Fulvia fulva (CPC 13652). A–H. Observations in vitro. A. Culture on V8. B, D. Conidiophore reduced to conidiogenous cell and catenate conidia. C. Conidiogenous cell and catenate conidia. E, F. Conidiophore and catenate conidia. G, H. Catenate conidia. Scale bars = 10 μm. © 2017 Westerdijk Fungal Biodiversity Institute. [Image: 4CJX]](https://data.landcareresearch.co.nz/Image/4CJX/SysPortalThumb)
![Fig. 31. Fulvia fulva (CPC 13652). A–H. Observations in vitro. A. Culture on V8. B, D. Conidiophore reduced to conidiogenous cell and catenate conidia. C. Conidiogenous cell and catenate conidia. E, F. Conidiophore and catenate conidia. G, H. Catenate conidia. Scale bars = 10 μm. © 2017 Westerdijk Fungal Biodiversity Institute. [Image: 4CJX]](https://data.landcareresearch.co.nz/Image/4CJX/SysPortalPreview)
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Videira, S.I.R.; Groenewald, J.Z.; Nakashima, C.; Braun, U.; Barreto, R.W.; de Wit, P.J.G.M.; Crous, P.W. 2017: Mycosphaerellaceae – Chaos or clarity?. Studies in Mycology 87: 257-421.
10.1016/j.simyco.2017.09.003
Article
Taxonomic concepts
Adelopus gaeumannii T. Rohde
Catenulocercospora fusimaculans (G.F. Atk.) C. Nakash., Videira & Crous 2017
Cercoseptoria handelii (Bubák) Deighton
Cercospora assamensis S. Chowdhury
Cercospora bougainvilleae Munt.-Cvetk.
Cercospora brachycarpa Syd.
Cercospora dioscoreae Ellis & G. Martin
Cercospora exitiosa Syd. & P. Syd.
Cercospora ferruginea Fuckel
Cercospora fusimaculans G.F. Atk.
Cercospora handelii Bubák
Cercospora henningsii Allesch.
Cercospora manihotis Henn.
Cercospora microsora Sacc.
Cercospora passaloroides G. Winter
Cercospora perfoliati Ellis & Everh.
Cercospora personata (Berk. & M.A. Curtis) Ellis & Everh.
Cercospora rosicola Pass.
Cercosporidium bougainvilleae (Munt.-Cvetk.) Sobers & C.P. Seym. 1969 [1968]
Cercosporidium graminis (Fuckel) Deighton 1967
Cercosporidium henningsii (Allesch.) Deighton 1976
Chuppomyces handelii (Bubak) U. Braun, C. Nakash., Videira & Crous 2017
Cladosporium personatum Berk. & M.A. Curtis
Cladosporium solanicola Viégas
Clarohilum henningsii (Allesch.) Videira & Crous
Distocercosporaster dioscoreae (Ellis & G. Martin) Videira, H.D. Shin, C. Nakash. & Crous
Distomycovellosiella brachycarpa (Syd.) U. Braun, C. Nakash., Videira & Crous 2017
Fulvia fulva (Cooke) Cif.
Fusarium maculans Berenger
Fusoidiella anethi (Pers.) Videira & Crous
Fusoidiella anethi (Pers.) Videira & Crous
Graminopassalora graminis (Fuckel) U. Braun, C. Nakash., Videira & Crous
Kirramyces hebes W.P. Wu, B. Sutton & Gange
Mycosphaerella berkeleyi W.A. Jenkins
Mycosphaerella handelii Crous & U. Braun
Mycosphaerella henningsii Sivan.
Mycosphaerella manihotis Ghesq. & Henrard
Mycosphaerella microsora Syd.
Mycosphaerella millegrana (Cooke) J. Schröt.
Mycosphaerella mori F.A. Wolf
Mycosphaerella rosicola B.H. Davis ex Deighton
Mycovellosiella brachycarpa (Syd.) Deighton
Mycovellosiella fulva (Cooke) Arx
Mycovellosiella solanicola (Viégas) Munt.-Cvetk.
Neophloeospora maculans (Berenger) Videira & Crous 2017
Nothopassalora personata (Berk. & M.A. Curtis) U. Braun, C. Nakash., Videira & Crous
Nothophaeocryptopus gaeumannii (T. Rohde) Videira, C. Nakash., U. Braun & Crous 2017
Paracercosporidium microsorum (Sacc.) U. Braun, C. Nakash., Videira & Crous 2017
Paramycovellosiella passaloroides (G. Winter) Videira, H.D. Shin & Crous 2017
Passalora assamensis (S. Chowdhury) U. Braun & Crous
Passalora bacilligera (Mont. & Fr.) Mont. & Fr.
Passalora bougainvilleae (Munt.-Cvetk.) R.F. Castañeda & U. Braun
Passalora brachycarpa (Syd.) U. Braun & Crous
Passalora dioscoreae (Ellis & G. Martin) U. Braun & Crous
Passalora ferruginea (Fuckel) U. Braun & Crous
Passalora fulva (Cooke) U. Braun & Crous
Passalora fusimaculans (G.F. Atk.) U. Braun & Crous
Passalora graminis (Fuckel) Höhn.
Passalora henningsii (Allesch.) R.F. Castañeda & U. Braun
Passalora microsora (Sacc.) U. Braun
Passalora passaloroides (G. Winter) U. Braun & Crous
Passalora perfoliati (Ellis & Everh.) U. Braun & Crous
Passalora personata (Berk. & M.A. Curtis) S.A. Khan & M. Kamal
Passalora rosicola (Pass.) U. Braun
Phaeocryptopus gaeumannii (T. Rohde) Petr.
Phaeoisariopsis personata (Berk. & M.A. Curtis) Arx
Phaeophleospora atkinsonii (Syd.) Pennycook & McKenzie
Phaeoramularia dioscoreae (Ellis & G. Martin) Deighton
Phaeoramularia fusimaculans (G.F. Atk.) X.J. Liu & Y.L. Guo
Phloeospora maculans (Berenger) Allesch.
Phloeospora mori (Lév.) Sacc.
Pluripassalora bougainvilleae (Munt.-Cvetk.) U. Braun, C. Nakash., Videira & Crous
Pseudocercospora handelii (Bubák) Deighton
Ragnhildiana ferruginea (Fuckel) U. Braun, C. Nakash., Videira & Crous
Ragnhildiana perfoliati (Ellis & Everh.) U. Braun, C. Nakash., Videira & Crous
Rosisphaerella rosicola (Pass.) U. Braun, C. Nakash., Videira & Crous 2017
Scoleciasis atkinsonii Syd.
Scolicotrichum graminis Fuckel
Descriptions
Notes: Distomycovellosiella is a monotypic genus that is morphologically similar to Mycovellosiella but is not congeneric with its type, Mycovellosiella cajani. Morphologically, Distomycovellosiella differs from Mycovellosiella by forming distoseptate conidia with slightly thickened and refractive loci (Fig. 24). Distomycovellosiella forms a clade well-supported by all three phylogenetic methods (Fig. 1, clade 47; Fig. 3, clade 9) and that is closely related to Pseudocercosporella as defined by its type, Pseudocercosporella bakeri.
Description in vitro (on SNA; CPC 13652): Mycelium composed of hyaline to pale brown hyphae, uniform in width, 2 μm diam. Conidiophores arising from hyphae, pale brown, smooth to rough, micro- or macronematous, multi-septate, simple or short branched, straight or sinuous, often strongly curved at the tip, 20–160 μm × 2.5–10 μm, variable in width, sometimes reduced to conidiogenous cell. Conidiogenous cells integrated, terminal and intercalary, proliferating sympodially, polyblastic, with rim-like conidiogenous loci that are darkened and thickened, 1–2.5 μm. Conidia catenate, often forming branched chains, ovoid, obovoid, ellipsoidal, sphaerical, cylindrical, straight or strongly curved, 10–30 × 5–10 μm, 1–4-septate, hila thickened and darkened, 1–2.5 μm diam.
The genus Fulvia is no longer considered a synonym of Passalora as a result of analysis of the type species, Fulvia fulva (≡ Cladosporium fulvum ≡ Passalora fulva), which was recollected and epitypified in this study. Fulvia fulva clusters close to Stromatoseptoria in the phylogenetic analyses (Fig. 1, clade 59; Fig. 3, clade 23). The single-gene trees indicate that both LSU and ITS are able to distinguish this species but rpb2 is more reliable. Fulvia fulva is the causal agent of tomato leaf mould, a disease that affects mostly the leaves of tomato but occasionally also stems, blossoms, petioles and fruit (Butler & Jones 1949, de Wit 1977, 1992, Jones et al. 1997). The interaction between Fulvia fulva and tomato is governed by a gene-for-gene relationship, a characteristic that made this organism an interesting model to study plant-pathogen interactions (de Wit 1981, 1992). The resistance of tomato against Fulvia fulva was genetically determined by the presence of Cf (Cladosporium fulvum) resistance genes of which now five have been cloned. Cf proteins mediate the recognition of effector proteins secreted by Fulvia fulva of which all encoding genes have been cloned (de Wit 2016). Fulvia fulva was once a devastating pathogen of tomato that required treatment with agrochemicals, but since various Cf genes from different wild Solanum species were introduced in commercial tomato cultivars by breeders the pathogen is now under control. Commercially grown tomato cultivars contain up to five different Cf genes (Cf-2, Cf-4, Cf-4E, Cf-5 or Cf-9) (Thomma et al. 2005).
epitype: CBS H-22950, MBT378581, culture ex-epitype CBS 142314
Notes: This genus is introduced to accommodate the species Phloeospora maculans that is not congeneric with the type of Phloeospora, Phloeospora ulmi, and clusters in a single strain lineage in the phylogenetic analyses performed in this study (Fig. 1, clade 55; Fig. 2, clade 19). Morphologically, Neophloeospora can be distinguished from Phloeospora by the subhyaline to pale brown conidia constricted at the septum.
Notes: The genus Nothophaeocryptopus is introduced to accommodate the species Phaeocryptopus gaeumannii which is not congeneric with the type of Phaeocryptopus, Phaeocryptopus nudus (Dothideales). The systematic position of Phaeocryptopus gaeumannii was originally determined based on a phylogeny of combined LSU and SSU sequences that placed it within the Mycosphaerellaceae (Capnodiales), followed by a phylogeny of ITS sequences that placed it in the Mycosphaerella heimii complex (Winton et al. 2007). In this study, the phylogenetic results agreed with the previous results of Winton et al. (2007), placing this species in a well-supported clade (Fig. 1, clade 23; Fig. 2, clade 29), closely related with Pallidocercospora
Description in vivo (CBS H-20777): Leaf spots absent or yellowish green, angular, 1–2 mm diam, delimited by leaf veins. Caespituli hypophyllous, olivaceous to pale brown. Mycelium internal, consisting of hyaline, branched, septate, 1–2 μm diam hyphae. Stromata absent or only formed as small aggregations of a few swollen substomatal hyphal cells. Conidiophores medium brown, arising from stomata, in fascicles of up to 12, unbranched or occasionally branched, straight to flexuous, usually up to 3-septate, occasionally with a single basal septum, 40–180 × 3–3.5 μm, geniculate at the apex. Conidiogenous cells integrated, terminal, somewhat swollen, 3–6.5 μm in width, polyblastic, proliferating sympodially, with conidiogenous loci flat, somewhat thickened and darkened, 1–2 μm diam. Conidia solitary, olivaceous to pale brown, thin-walled, smooth, straight or gently curved, basal cell ellipsoid-doliiform and obconical truncate without protruding, apical cell narrowly long-ellipsoid to subcylindrical, 21–68 × 4.5–8.5 μm, (0–)1(–3)-euseptate, constricted at basal septum, with hilum somewhat thickened, darkened and refractive, 1.5–2 μm diam.
Description in vitro (on V8; CBS 131547): Mycelium composed of hyaline to pale olivaceous brown, delicate hyphae, 2–2.5 μm width. Conidiophores macronematous, pale olivaceous brown to brown, simple or branched, straight to sinuous, smooth, paler towards the apex, 25–300 × 2.5–3.3 μm. Conidiogenous cells integrated, terminal, proliferating sympodially, polyblastic, conidiogenous loci located on the shoulders and the apex, slightly thickened and darkened, 2.5 μm diam. Conidia solitary, pale olivaceous brown to brown, cylindrical to obclavate, obconical truncate at the base, rounded or pointed at the apex, 13–37.5 × 2.5–5 μm, (0–)1-euseptate, constricted at the septum, hilum slightly thickened, darkened and refractive, 2.5 μm diam.
Passalora was the first genus introduced for cercosporoid hyphomycetes (Fries 1849) and a review of the taxonomical history of the genus has recently been published by Braun et al. (2013). In one of the most comprehensive examinations on this generic complex, Crous & Braun (2003) concluded that various genera (e.g. Mycovellosiella, Phaeoramularia, Fulvia) should be merged under the oldest name Passalora. After this revision Passalora included cercosporoid species with solitary, fasciculate to synnematous conidiophores and conidia formed singly or in chains, but in all cases with conspicuous (thickened and darkened) conidiogenous loci (scars) and mostly non-scolecosporous, pigmented conidia. This new concept was also supported by first molecular sequence analyses (Crous et al., 2000, Crous et al., 2001b). However, with the addition of more species and more phylogenetic markers, Passalora s. lat. has proven to be para- or polyphyletic (Thomma et al., 2005, Crous et al., 2009b, Crous et al., 2009d, Crous et al., 2013a). In addition, the type species has not been subjected to DNA sequence analyses before, and the passalora-like clades distributed throughout the Mycosphaerellaceae are not clearly connected with morphological groups within Passalora (e.g. mycovellosiella-like). In this study, we propose a good candidate for the epitypification of the type species of Passalora (CBS 131547). Phylogenetically, this strain forms a single species clade in all phylogenetic analyses performed (Fig. 1, clade 34; Fig. 2, clade 22), but without a strong link to other genera. With the additional epitypification of the type species of Fulvia (Fulvia fulva; Fig. 1, clade 59), Mycovellosiella (Mycovellosiella cajani; Fig. 1, clade 7) and Phaeoramularia (Phaeoramularia gomphrenicola; Fig. 1, clade 61), these names are resurrected and applied to different monophyletic clades and are no longer regarded as synonyms of Passalora s. str. The value of features such as mycelium internal and/or external, conidia solitary or in chains, remains doubtful and barely applicable for the discrimination of cercosporoid genera. Morphologically, Passalora s. str. is rather different from common passalora-like species (Fig. 15), in having sparsely septate, flexuous conidiophores, and predominantly smooth, olivaceous, 1–2-septate conidia constricted at the basal septum, with somewhat to distinctly thickened, darkened, and refractive loci. The placement of the hundreds of passalora-like species that are not known from their DNA is not yet possible, and these would for the interim have to be retained in Passalora s. lat. as a wide, morphologically circumscribed genus, pending cultures and results of DNA sequence analyses.
epitype: CBS H-20777, ex-epitype culture: CBS 131547
Hyphomycetous, phytopathogenic. Mycelium internal, consisting of hyaline, branched, septate hyphae. Stromata absent or small. Conidiophores emerging through stomata, in fascicles, unbranched or branched, straight to flexuous, at times with a single basal septum, usually up to 3-septate, medium brown, somewhat swollen in the conidiogenous region. Conidiogenous cells integrated, terminal, with flat, somewhat thickened and darkened loci. Conidia solitary, olivaceous to pale brown, thin-walled, smooth, straight or gently curved, mostly didymosporous, constricted at septum, with somewhat thickened, darkened and refractive hila.
Cited scientific names
- Adelopus gaeumannii T. Rohde 1936
- Catenulocercospora C. Nakash., Videira & Crous 2017
- Catenulocercospora fusimaculans (G.F. Atk.) C. Nakash., Videira & Crous 2017
- Cercoseptoria handelii (Bubák) Deighton 1976
- Cercospora assamensis S. Chowdhury 1957
- Cercospora bougainvilleae Munt.-Cvetk. 1957
- Cercospora brachycarpa Syd. 1930
- Cercospora dioscoreae Ellis & G. Martin 1882
- Cercospora exitiosa Syd. & P. Syd. 1907 [1906]
- Cercospora ferruginea Fuckel 1863
- Cercospora fusimaculans G.F. Atk. 1892
- Cercospora handelii Bubák 1909
- Cercospora henningsii Allesch. 1895
- Cercospora manihotis Henn. 1902
- Cercospora microsora Sacc. 1880
- Cercospora passaloroides G. Winter 1883
- Cercospora perfoliati Ellis & Everh. 1889
- Cercospora personata (Berk. & M.A. Curtis) Ellis & Everh. 1885
- Cercospora rosicola Pass. 1875
- Cercosporidium bougainvilleae (Munt.-Cvetk.) Sobers & C.P. Seym. 1969 [1968]
- Cercosporidium graminis (Fuckel) Deighton 1967
- Cercosporidium henningsii (Allesch.) Deighton 1976
- Chuppomyces handelii (Bubák) U. Braun, C. Nakash., Videira & Crous 2017
- Chuppomyces Videira & Crous 2017
- Cladosporium fulvum Cooke 1883
- Cladosporium personatum Berk. & M.A. Curtis 1875
- Cladosporium solanicola Viégas 1946
- Clarohilum henningsii (Allesch.) Videira & Crous 2017
- Clarohilum Videira & Crous 2017
- Distocercosporaster dioscoreae (Ellis & G. Martin) Videira, H.D. Shin, C. Nakash. & Crous 2017
- Distocercosporaster Videira, H.D. Shin, C. Nakash. & Crous 2017
- Distomycovellosiella brachycarpa (Syd.) U. Braun, C. Nakash., Videira & Crous 2017
- Distomycovellosiella U. Braun, C. Nakash., Videira & Crous 2017
- Fulvia Cif. 1954
- Fulvia fulva (Cooke) Cif. 1954
- Fusarium maculans Berenger 1844
- Fusoidiella anethi (Pers.) Videira & Crous 2017
- Graminopassalora graminis (Fuckel) U. Braun, C. Nakash., Videira & Crous 2017
- Graminopassalora U. Braun, C. Nakash., Videira & Crous
- Kirramyces hebes W.P. Wu, B. Sutton & Gange 1996
- Mycosphaerella berkeleyi W.A. Jenkins 1938
- Mycosphaerella handelii Crous & U. Braun 2003
- Mycosphaerella henningsii Sivan. 1985
- Mycosphaerella manihotis Ghesq. & Henrard 1924
- Mycosphaerella microsora Syd. 1940
- Mycosphaerella millegrana (Cooke) J. Schröt. 1894
- Mycosphaerella mori F.A. Wolf 1935
- Mycosphaerella rosicola B.H. Davis ex Deighton 1967
- Mycosphaerellaceae Lindau 1897
- Mycovellosiella brachycarpa (Syd.) Deighton 1974
- Mycovellosiella fulva (Cooke) Arx 1983
- Mycovellosiella solanicola (Viégas) Munt.-Cvetk. 1960
- Neophloeospora maculans (Berenger) Videira & Crous 2017
- Neophloeospora U. Braun, C. Nakash., Videira & Crous 2017
- Nothopassalora personata (Berk. & M.A. Curtis) U. Braun, C. Nakash., Videira & Crous 2017
- Nothopassalora U. Braun, C. Nakash., Videira & Crous 2017
- Nothophaeocryptopus gaeumannii (T. Rohde) Videira, C. Nakash., U. Braun & Crous 2017
- Nothophaeocryptopus Videira, C. Nakash. & Crous 2017
- Paracercosporidium microsorum (Sacc.) U. Braun, C. Nakash., Videira & Crous 2017
- Paracercosporidium Videira & Crous 2017
- Paramycovellosiella passaloroides (G. Winter) Videira, H.D. Shin & Crous 2017
- Paramycovellosiella Videira, H.D. Shin & Crous 2017
- Parapallidocercospora thailandica (Crous, Himaman & M.J. Wingf.) Videira & Crous 2017
- Parapallidocercospora Videira, Crous, U. Braun & C. Nakash. 2017
- Passalora assamensis (S. Chowdhury) U. Braun & Crous 2003
- Passalora bacilligera (Mont. & Fr.) Mont. & Fr. 1856
- Passalora bougainvilleae (Munt.-Cvetk.) R.F. Castañeda & U. Braun 1991
- Passalora brachycarpa (Syd.) U. Braun & Crous 2003
- Passalora dioscoreae (Ellis & G. Martin) U. Braun & Crous 2003
- Passalora ferruginea (Fuckel) U. Braun & Crous 2003
- Passalora Fr. 1849
- Passalora fulva (Cooke) U. Braun & Crous 2003
- Passalora fusimaculans (G.F. Atk.) U. Braun & Crous 2003
- Passalora graminis (Fuckel) Höhn. 1923
- Passalora henningsii (Allesch.) R.F. Castañeda & U. Braun 1989
- Passalora microsora (Sacc.) U. Braun 1995
- Passalora passaloroides (G. Winter) U. Braun & Crous 2003
- Passalora perfoliati (Ellis & Everh.) U. Braun & Crous
- Passalora personata (Berk. & M.A. Curtis) S.A. Khan & M. Kamal 1961
- Passalora rosicola (Pass.) U. Braun 1995
- Phaeocryptopus gaeumannii (T. Rohde) Petr. 1938
- Phaeoisariopsis personata (Berk. & M.A. Curtis) Arx 1983
- Phaeophleospora atkinsonii (Syd.) Pennycook & McKenzie 2002
- Phaeoramularia dioscoreae (Ellis & G. Martin) Deighton 1976
- Phaeoramularia fusimaculans (G.F. Atk.) X.J. Liu & Y.L. Guo 1982
- Phloeospora maculans (Berenger) Allesch. 1900 [1901]
- Phloeospora mori (Lév.) Sacc. 1884
- Pluripassalora bougainvilleae (Munt.-Cvetk.) U. Braun, C. Nakash., Videira & Crous 2017
- Pluripassalora Videira & Crous 2017
- Pseudocercospora handelii (Bubák) Deighton 1987
- Pseudocercospora platanigena Videira & Crous 2017
- Pseudophaeophleospora atkinsonii (Syd.) U. Braun, C. Nakash., Videira & Crous 2017
- Ragnhildiana ferruginea (Fuckel) U. Braun, C. Nakash., Videira & Crous 2017
- Ragnhildiana perfoliati (Ellis & Everh.) U. Braun, C. Nakash., Videira & Crous 2017
- Ragnhildiana Solheim 1931
- Rosenscheldiella brachyglottidis G.F. Laundon & Sivan. 1972 [1971]
- Rosenscheldiella korthalsellae A. Sultan, P.R. Johnst., D.-C. Park & A.W. Robertson 2011
- Rosenscheldiella Theiss. & Syd. 1915
- Rosisphaerella rosicola (Pass.) U. Braun, C. Nakash., Videira & Crous 2017
- Rosisphaerella Videira & Crous 2017
- Scoleciasis atkinsonii Syd. 1924
- Scolicotrichum graminis Fuckel 1870 [1869-70]
- Septoria mori Lév. 1846
- Zasmidium cerophilum (Tubaki) U. Braun, C. Nakash., Videira & Crous 2017
Metadata
971aba4b-0909-466a-a9e3-09390a1397d2
reference
Names_Fungi
20 April 2018
16 February 2023
