Circumscription of anamorph genera in Mycosphaerella based on ITS rDNA sequence and morphology

Uwe  Braun

The Mycosphaerellaceae represent thousands of fungal species that are associated with diseases on a wide range of plant hosts. Understanding and stabilising the taxonomy of genera and species of Mycosphaerellaceae is therefore of the utmost importance given their impact on agriculture, horticulture and forestry. Based on previous molecular studies, several phylogenetic and morphologically distinct genera within the Mycosphaerellaceae have been delimited. In this study a multigene phylogenetic analysis (LSU, ITS and rpb2) was performed based on 415 isolates representing 297 taxa and incorporating ex-type strains where available. The main aim of this study was to resolve the phylogenetic relationships among the genera currently recognised within the family, and to clarify the position of the cercosporoid fungi among them. Based on these results many well-known genera are shown to be paraphyletic, with several synapomorphic characters that have evolved more than once within the family....

A taxonomic and biogeographic overview of the genus Myrionora is provided. Two species are recognized, M. albidula (Willey) R. C. Harris and M. pseudocyphellariae (Etayo) S. Ekman & Palice comb. nov. The genus is characterized by polysporous asci, the presence of crystals in the hymenium and proper exciple that partly consist of lobaric acid, and a photobiont with large cells (mostly in the range 12–20 µm). Myrionora albidula is currently known from Germany, Norway, Sweden, the Russian Federation (Altayskiy Kray, Chelyabinskaya Oblast', Khabarovskiy Kray and Zabaykal'skiy Kray), and the United States (Alaska, Connecticut, Maine and Massachusetts). It inhabits bark of deciduous trees and shrubs and conifers over a wide range of latitudes. Myrionora pseudocyphellariae is known from Chile and Ecuador, where it has been encountered on lichens and decaying bark. Based on morphological characteristics, we conclude that Myrionora belongs in the Ramalinaceae.

hfycologza, 93(6), 2001, pp. 1081-1101. 0 2001 by The Mycological Society of America, Lawrence, KS 660448897 A phylogenetic redefinition of anamorph genera in Mycosphaerella based on ITS rDNA sequence and morphology Pedro W. Crousl Ji-Chuan Kang Dtpartmmt of Plant Pathology, University of Stellenbosch, I! Bag XI, Matieland 7602, South Afica Uwe Braun Martin-Luther-Universitat,FB. Biologze, Institut fur Geobotanik und Botanlscher Garten, Neuwerk 21, D06099 Halle (Saale), Germany Abstract: Anamorph genera associated with Mycosphaerella are separated on a combination of characters such as conidiomatal structure, the nature and arrangement of conidiophores, conidiogenesis, dehiscence scars and pigmentation. To date 27 genera have been reported as anamorphs of Mycosphaerella, 19 of which were included in this study. Based on the phylogeny of the ITS1, 5.8S, and ITS2 rDNA sequence data, Mycosphaerella proved to be monophyletic. The major group consisted of several small clusters correlating to specific anamorph genera, as well as other clusters containing an assemblage of anamorph genera. Although Mycosphaerella is monophyletic, it appears that many of the morphological characters defining different anamorph genera evolved more than once and for the most part do not represent true groups within Mycosphaerella. Nevertheless, the anamorph generic concepts contain valuable information of practical use to plant pathologists and mycologists working with these organisms. Based on these results, as well as a re-evaluation of the criteria upon which anamorph genera are distinguished in this complex, a reduced set of informative criteria and genera are proposed. The degree of scar thickening, darkening and refraction, as well as the presence or absence of pigmentation in conidiophores and conidia still appear to be useful features delimiting anamorph genera of Mycosphaerella. Key Words: Cercospora, Cladosporium, ITS phylogeny, Mycosphaerella, Mycovellosiella, Passalora, Pseudocercospora INTRODUCTION Bisby and Ainsworth (1942) stated that 'Nature may make species, but man has made the genera'. To this Accepted for publication April 19, 2001. Corresponding author, Email: pwc@maties.sun.ac.za ' end the genus Mycosphaerella Johanson (Dothideales, Mycosphaerellaceae) has been linked to at least 27 different coelomycete or hyphomycete anamorph genera (Kendrick and DiCosmo 1979), 23 of which were accepted by Crous et al (2000). The anamorphic genera are separated using a combination of characters including conidiomatal structure (pycnidia, acervuli, sporodochia, etc.), mycelium (presence or absence of superficial mycelium and texture thereof), conidiophores (arrangement, branching, pigmentation and ornamentation), conidiogenous cells (placement, proliferation and scar type) and conidia (formation, shape, septation, ornamentation, pigmentation and catenulation). One of the anamorph genera for which more than 3000 names have already been published is Cercospora Fresen. (Pollack 1987). Cercospora was last monographed by Chupp (1954), who proposed a broad concept for the genus, simply stating if hila were thickened or not, and if conidia were pigmented, single or in chains. Deighton, Sutton, Braun and several other workers divided the Cercospora-complex into smaller, more morphologically similar units based on different combinations of the characters discussed above (Braun 1995a, 1998). The abandonment of the "Chupp concept" has resulted in close to 50 genera being recognized in this complex (Braun 1995a). One of the reasons for this was the strict interpretation of the numerous conidiogenous events as defined by Sutton and Hennebert (1994), as well as the additional characters discussed above. In the ophiostomatoid fungi, however, Wingfield et al (1993) found that the number of anamorph genera attributed to Ceratoqstis Ellis & Halst. and Ophiostoma Syd. & P. Syd. could be reduced from sixteen to seven based on a re-evaluation of conidium ontogeny (Wingfield et al 1993). Several anamorph genera have been found to have species with conidiomata varying from mononematous, scattered conidiophores to sporodochia with a basal stroma, or from pycnidia to sporodochia to synnemata (Samuels and Seifert 1987, Seifert and Okada 1990). Based on similar observations Sutton (1980) and Nag Raj (1981) saw the need to abandon the distinction between hyphomycetes and coelomycetes, as acervuli were frequently found to form a continuum with more stromatic, sporodochial forms. If this plasticity is taken into consideration when examining the 23 anamorph genera accepted by Crous et a1 (2000), many appear superfluous. Crous (1998) was of the opinion that iVycosphaerella is polyphvletic, and that it could possibly be split along the lines as defined by its different anamorphs. However, in recent phylogenetic studies done on this group (Stewart et a1 1999, C:rous et a1 1999, 2000), ~Mjcosphnerellaproved to be mo~lophyletic,having several well-defined clades correlating with specific anamorph genera, and other clades corlsisting of a motley assemblage of species from diverse anamorph genera. The aims of the present study were thus to derive a molecular phylogeny of selected anamorph genera linked to ~Mjcosphner~lla, and to compare these data with morphology, re-evaluating the generic concepts of the respective genera studied. A further aim was to derive a new set of characters and concepts that would enable us to propose a simplified, but workable classificatiorl system, without the loss of too much taxonomic information. DLVAamfilijcatio?7 and sequencing.-Genomic DNA Iva, isolated from fungal mycelium (TABLEI) collected directly from malt extract agar (MEX) (Biolab, Midrand, South .Mrica) plates using a modification of the isolation protocol of Lee and Taylor (1990). DNA quantification was done by LT7 spectroscopy using a Beckman DU Series 7500 Spectrophotometer. Buffers and sequencing conditioris were as state d in Crous e t a1 (2000). The purified PCR products were sequenced using an M I Prism 377 DNA Sequencer (PerkillElmer, Norlvalk, Connecticut), lvith an AABIPRIShI1" Dye Terminator Cycle sequencing Ready Reaction Kit (Perkin Elmer, Thrrington, U.K.). Phjlogenetic a17a!ssis.-The nucleotide sequences of the 3.XS rRVA gene and the flanking internal transcribed spacers (ITS1 and ITS2) were assembled using Sequence NavigatorT\' version 1.0.1. (Perkin Elmer, Applied Biosysterns, Inc., Foster City California). Alignment of the sequence files was conducted using the CLUSTAL W software (Thompson et a1 1994). Adjustments for improvement were made by eye where necessary. .Uignment gaps were coded as a fifth base in the parsimony analysis. T h e parsimony analyses were performed with PAUP* version 4.0b2a (Swoffbrd 1999). The neighbor-joining distance lrlethod was also used to infer a gene tree. Clade stability waa assessed using 1000 neighbor-joining bootstrap replications. The parsimony tree scores including tree length, consistenc~index, retention index, rescaled consiste~icyindex and homoplasy index (TL, CI, RI, RC and HI) were also calcl~lated. Xlorphology-Isolates were either obtained from other culture collection^ or isolated fi-om infected plant material as explained in Crous (1998), and deposited in the culture collection of the Departrnerlt of Plant Patholo,?; Uni1-ersity of Stellenbosch. South ,Vi-ica (STE-U. T ~ B LIE) , and at the Centraalbureau voor Schimmelcultures, Utrecht, Nether- lands. Isolates were cultured on divided plates containing MEA and carnation-leaf agar ( C U ) (Fisher et a1 1982, Crous et a1 1992). (:ulture, were incubated at 25 C ullder near-llltraviolet light, and exarnined tceekll- for 1-2 mo. Mounts were prepared in lactophenol, examined usitig Nomarski interferewe phase contrast and hright-field phase contrasL ruicroscopy, and nieasure~ne~lts made at XI000 magnification. ~Molrcularplzy1ogenj.-Maximum parsimony analysis was c o n d ~ c t e dfor 383 parsimony-informative characters in the alignment using the heuristic search option with 1000 random sequence input orders for exact solution. The other 265 sites were excluded from the analysis to avoid fragmentary ambiguities observed in the ITS1 and ITS2 regions of the aligllment (data not shown). A total of 208 equally most parsi~nonioustrees (MPT) were obtained and compared fbr the best topolo'q using the Kishino-Hasegawa test (FIG. 1 ) . The 383 parsimony-informative characters in the alignment were also analyzed by means of the neighborjoining (NJ) rrlethod using u~lcorrected pair frequencies, and evaluated with 1000 bootstrap replications, which produced a similar tree t o p o l o g ~ (FIG. 2 ) , ~vithhigh bootstrap values for the relevant clades. Fusnriu?n prolzferntum (Matsush.) Nirenberg (GenBank X94l'il) was used as outgroup. Trvo species of Didj~nellnSacc. as well as Ran~z~lis$or.cr Miura were also included. The latter genus has been lirlkecl to teleornorphs in Taprsicc (Pers.) Fuckel (Crous et a1 2000). Sequence data of this stud)' were deposited at GenBank ('2F362048-3620'70), and the alignment in TreeBase (S.390, h1889). Based on the ITS rDNA phylogeny obtained for the taxa investigated in the present study (FIGS.1, 2 ) , five clusters were identified. These include a major morlophyletic ~Mjcosphaerrllacluster (NJ bootstrap = 84, FIG. 2) (with subclusters for cercosporoicl and coelomycete anamorphs), and two closely related clusters for Dissocorziu?n de Hoog, Oorschot & Hijrvegen (NJ bootstrap = 100, FIG. 2) and Clado.~$oriz~nz L,ink anamorphs (NJ bootstrap = 100, FIG. 2)).X fourth cluster represents Kamulzs;hora spp. rt.ith ;\.Iollzsin (Fr.) P. Karst. ( = 7apesin) teleomorphs (NJ bootstrap = 100, FIG.2). Species of Didymellci formed the fifth cluster (NJ bootstrap = 88, FIG. 2 ) . The present data support :l/Ijcosphn~rellcias monophyletic, with the I~issoc.oniurnand Cladosfio~iutnclusters representing sections or subgenera in 1L!~cosphnn-ella. ITS sequences of other related taxa in the Dothideccles, namely Botr;l;osphneria Ces. & De Not. (AF 195774),1)othidea Fr. (AF027764), Leptosphnrrin Ces. & De Not. (U04203), Phaeos$lzurrin I Mivake (U77359) and Guignardia Viala & Ravaz (AF312008AF312015) from GenBank were found to be nonalignable with those of Mycosphaerella. i2lorphology.-Several of the c~llturesobtained from other collections were sterile, thus making it impossible to confirm their identity (indicated as ? in TABLE I). Although cultures of only 19 anamorph genera were available for inclusion in the present study, numerous generic type and other specimens have been examined. The genera dealt with in this study are discussed below. DISCUSSION Cladosporium.-Cladospom'um Link (1816) is the oldest generic name for this genus. No monographic treatment of this important, but complicated genus has been attempted. Hughes (1958), who examined numerous types of hyphomycetes described in the lgth century, reassessed various old generic names and reduced the following genera to synonymy with Cladospom'um: Spar-ocladium Chev., 1826; Myxocladium Coi-da, 1837; Didymotrichum Bonord., 1851 and Heterospom'um Klotzsch ex Cooke, 1877. A review of the history of Cladosporium has been given by David (1997). The status of the genus Heterosponum has resulted in much controversy. De Vries (1952) reduced it to synonymy with Cladospom'um, a conclusion supported by Hughes (1958) and Ellis (1971, 1976).Arx (1981, 1983) re-introduced Heterospom'um, and McKemy and Morgan-Jones (1990) as well as Braun (1995,) agreed with this taxonomic concept. David (1997) examined Cladosponum and Heterospom'um by means of SEM and clearly demonstrated that the scars and hila in the two genera are similar, and of the 'Cladosporium-type', i.e., protuberant with a central dome surrounded by a raised rim. Based on these results, David (1997) placed Heterosporium in Cladospom'um and proposed the combination Cludospom'um subgen. Heterospom'um (Klotzsch ex Cooke) J.C. David, a treatment that should be followed. Arx (1983) considered Arroconidiella J.C. Lindq. & Alippi as synonym of Heterospom'um. This treatment is, however, not tenable since Acroconidiella is characterized by having tretic conidiogenous cells (Ellis 1971, David 1997). Furthermore, Arx (1983) also reduced Stenella Syd. to synonymy under Cladospo?ium. The scars in the former genus are, however, quite distinct from those of the latter genus (pileate, sensu David 1993), and these two genera should be retained as separate entities. Cladospom'um (incl. Heterospom'um) is characterized and distinguished from other Mycosphaerella anamorphs by its unique type of scars and conidial hila. FIG. 1. One of 208 equally most parsinlonious trees inferred from 383 parsimony-informative characters of aligned sequences of the 3.8s rRNA gene and flanking ITS1 and ITS2 regions using maximum parsimony a~lalysiswith heuristic search option and 1000 random sequence input orders (TL = 3873 steps, (:I = 0.312, RI = 0.348, RC = 0.171, HI = 0.688). The sequence of Fz~sariumprol@utum was used as outgroup to root the tree. Branch lengths are indicated. The peculiarity and separate position of Cladospo~iurn has also been confirmed in a previous molecular study (Crous et a1 2000). Descriptions and an expanded key to the species of Cladosporiz~rnavailable in culture have recently beer1 published by Ho et a1 (1999). The taxonomic concept applied in the latter paper is rather wide and not solely based on the structure of loci and conidial hila, so that a few species are included which belong in other genera, viz., .M~rouellosielln fulvn (Cooke) Arx [= Clado.sporiurrr I I 1 94.47 Phaeoisariopsis 1 Paracercospora I I I I 2552 462'75 336.33 Stigmina 1 P~eudocercospom I I I I I I 100 ,. , 1453 Mycovellosiella 221 Phaeoramularia 'efogloeopsis I I 1 346 Phaeophleospora I ., 31 3.76 Scolecosrigmina 1 Phaeophleospora C'webraunia ~ 9 4 1 7 1 Fusar~umprol?firafum outgroup I -. - 0.05 changes FIG. 2 . Neighbor-joining gene tree derived from 383 parsimony-informative characters in the alignment of ITS sequences using uncorrected pair frequencies. The bootstrap values representing 1000 bootstrap replicatiolls are indicated at the nodes of the tree. The sequence of Fusam'um prolzferatztm was used as outgroup to root the tree. fulvum Cooke, = Fuluza fulua (Cooke) Cif.], (,'ladorponunz malorum Ruehle [ = Pseudocladosponum k ~ l lermanzanum (Maraqas & I.H. Bredell) U. Braun] and Fuszcladzum @sum G. Winter [= CZ(~dosporzt~m carjzgenum (Ellis & Langl.)] Gottwald (the taxonomy of C. rnalorum and l? effuutum will be treated in separate monograph of Vtntuna Sacc. anamorphs by Braun and collaborators). Human pathogenic " Cladospomurn spp." must be excluded since they are anamorphs of ascomycetes belonging to the Herpotnrhzaceae, which has been confirmed by molecular studies (de Hoog et a1 1995, Masclaux et a1 1995, Takeo et a1 1995, Untereiner 2000). Some morphologically similar saprobic hyphomycetes are anamorphs of Caproventuria U. Braun [Wnturia, according to Untereiner (1997)], (Wnturiaceae) and have been placed in Pseudocladosporium U. Braun (1998). In the present study iZI. astmma (Fr.) Lindau was found to cluster in the Cludosporium clade (FIGS.1, 2). As the culture proved to be sterile, further research would be required to elucidate its possible relationship with C. polygonati M.B. Ellis, which also occurs on this host. Cladosporium (incl. Heterosporium) should be confined to saprobic and phytopathogenic (rarely hyperparasitic) hyphomycetes with cladosporium-like scars and hila and i%fycosphaerella teleomorphs (Mycosphaerellacene). I11 our analyses Clndosporium clustered adjacent to the main Mycosphaerella cluster, which suggests that it represents a separate section of iMycosphaerella. The issue of recognizing sections in iLfycosphaerella has been dealt with by several workers (Arx 1949, Barr 1972, David 1997), the correct nomenclature of which awaits further type and molecular studies. Ramularia and allied genera with colorless corzidiophore~ and conidia.-The genus Ramularia was described by Unger (1833). Ramularia and allied genera are characterized by having colorless conidiophores and conidia. A monograph of this complex of genera of colorless cercosporoid hyphomycetes ( Cercosporella Sacc., Ramularia, 1Veoramularia U. Braun, Neooz)ularin U. Braun, Phacellium Bonord., PseudocPrcosporella Deighton) has been published by Braun (1995a, 1998). Ouularia Sacc. (aseptate conidia) was reduced to synomymy with Ramularia sensu Saccardo (multiseptate conidia) (Hughes 1949, Sutton and W7aller 1988, Braun 1998). Ophiocladium Cavara was distinguished from this complex because of its curled conidiophores, and conidia that have somewhat eccentrically positioned scars. This genus was, howlever, reduced to synonymy with Ramularia (Braun 1988a, Sutton and M7aller 1988), as was supported by molecular data presented by Crous et a1 (2000). Conidium septation and unusual conidiophore shape with eccentric scars are thus characters to be considered at the species level. Passalora Fx, Cercospora, and allied genern with thickened, darkened conidiogenous loci and conidial hila.The genera in this complex are all very similar, and as more taxa were described, many intermediates were found that could not easily be allocated to any one of these. For the purpose of this study these genera are divided into five groups and discussed as such below. Asperisporium il/lnubl., Cercosporidium Earl; Dis- tocercospora Pons & R. Sutton, Passalora, Phaeoisariopsis Ferraris, Prathigada Subrum., Pseudocercosporidium Deighton, Quasiphloeospora B. Sutton, Crous & Shamoun. The separation of Passalora and Phaeoisariopsis is rather tenuous, and Braun (1995a) accepted Phaeoisnriopsis as the sjmnernatous counterpart of Pnssalom. Passalora was introduced by Fries (1849). Passalora bacilligera (Mont. & Fr.) Mont. & Fr. (- Cladosporium bacilligerum Mont. & Fr.), the type species, is characterized by having pigmented conidiophores and ellipsoid-fusiform, obclavate-subcylindric, (0-) 1(-3)septate, pigmented conidia, formed singly. Deighton (1967) emended the genus Passalora, confined it to three species occurrirlg on Alnus, and discussed the generic history in detail. Earle (1901) described the genus Cmosporidium [type species: Scolecotrichuv2 euphorbiar Tracy & Earle = Cercosporidium churtomium (Cooke) Deighton] . Earle (1902) reduced Cercosporidium to synonymy with Passalora. Deighton (1967) considered C;mcosporidium "a useful and definable genus, distinct from Passalorc~",but failed to give clear morphological differences between the two genera. Based on the descriptions in Deighton (1967), Cmosporidium was only distinguished from Passalora by forming welldeveloped stromata. The taxonomic value of stronlatal formation for the differentiation of genera in this complex is, however, very d o u b t f ~ ~Arx l . (1983) stated that even in Pu.ssalora b a r i l l i p , the type species of this genus, small substomatal stromata are often present. The size of the stromata is often influenced by the size and structure of the substomatal cavities, which are relatively small in hosts of the latter genus. rlrx (1983) followed the wider concept of P(~ssalora,which was generally used before Deighton (196'7) reintroduced Crrcosporidium, and maintained the latter genus in synonymy with Pas.saloru. Castaiieda and Braun (1989),Deighton ( I 990), and Braun (1995,) followed Arx's (1983) concept of Passalora (incl. Cercosporidium). Braun (1995b) discussed the differentiation of Pa.ssalora and allied genera in detail, provided a re-description of Passaloru ernend., and a key to the related genera. The following additional genera have been reduced to synonymy . . with Passalora (incl. Cercosfioridium) : Berteromjces Cif. (Deighton 1967) [type ipecies: B. aeneus Cif. = Passalhm cassiae (Henn.) U. Braun] and Orrophyllum Cif. (Braun et al 1999) [type species: 0. angelaemariae Cif. = Passalora gliricidiasis (Gonz. Frag. & Cif.) Castaiieda & U. Braun]. The genus Aspem's@orium,introduced by Maublanc (1913a, b), resembles Passaloru, but differs in having verrucose conidia (Ellis 1971, 1976, Arx 1983). Maublanc (1919a, h) considered Asperisporium cari- cae (Speg.) Maubl., the type species of this genus, to be the anamorph of Sphaerella caricae Maubl. ( = Mycosphaerella caricae Syd. & P. Syd.), but this connection has never been proven. The conidiogenous cells in Aspe-risporium spp. are usually only slightly geniculate and the scars, which agree well with those of Passalora spp., are usually more or less terminal and lateral, flat, and little protruding. Asperisporium spp. are usually easily distinguishable from Passalora spp., though there are a few intermediate taxa, e.g., Passalora scariolae Syd. (Deighton 1967) and l? milii (Syd.) de Vries (1952) with finely verruculose-rugose conidia. Although no molecular data are available to assess the validity of this genus, its separation seems doubtful. Pons and Sutton (1988) described Distocercospora to accommodate Cercos@orapachyderma Syd. & P. Syd. The slightly thickened conidial scars and pigmented conidia in Distocercospora resemble those of Passalora spp., and the frequently branched conidiophores in D. pachyderma are reminiscent of Mycovellosiella Rangel and some Phaeoramularia Munt.-Cvetk. species, but the conidia are predominantly distoseptate. As mentioned for taxa in the Pseudocercospora cluster, distoseptation appears to be important at species level, which questions the separation of Distocercospora from this complex. Pmthigada Subram. (Subramanian and Ramakrishnan 1956) is very close to Passalora (Braun 1995a), but differs, based on the present interpretation of this genus (Sutton 1994, Braun 1996, Braun and Melnik 1997), in having scolecosporous, pluriseptate, thick-walled, usually more or less rostrate conidia. Although molecular data are not yet available, this separation also appears uncertain. Quasiphloeospora (Sutton et a1 1996) was introduced to accommodate Cercospora saximontanensis Deighton, an unusual cercosporoid species characterized by having large subepidermal sporodochia (described as acerwli by Sutton et a1 1996), filiform conidiophores with percurrent as well as sympodial conidiogenous cells, and pale, but not quite colorless scolecospores. Sutton et a1 (1996) explained the differences between Quasiphloeospora and Cercospora as well as Pseudocercospora Speg., but failed to contrast it with Passalora, which is, indeed, quite similar. Quasiphloeospora differs from Passalora in having frequently percurrently proliferating conidiogenous cells and more or less cylindrical, rather cercosporalike conidia. It appears, therefore, that this genus may also eventually be reduced to synonymy with either the Passalora, or the Pseudocercospora complex. Given the mode of conidiogenesis, as well as the slightly thickened conidial scars, it may well prove a synonym of the latter. Pseudocercosporidium Deighton (1973) was described to accommodate Cercosporidium venezuelanu m Syd., a dematiaceous hyphomycete with long, erect, frequently branched conidiophores and cicatrized conidiogenous cells. This species is close to the Passalora/Mycovellosie22a/Phaeoramulariacomplex, but the structure of the conidiogenous loci, which are rather cercosporella-like, is quite distinct. Its status remains unclear. Phaeoisariopsis Ferraris (1909) was introduced to accommodate Isam'opsis gm'seola Sacc., the type species, which is a synnematous hyphomycete distinguished from Isariopsis Fresen. s.str. ( = Phacellium Bonord., see Braun 1998) by having pigmented conidiophores and conidia. Ferraris (1909) included three additional North American species, which were later excluded: Phaeoisariopsis grayiana (Ellis) Ferraris [E Fusicladium grayianum (Ellis) Deighton], P mexicana (Ellis & Everh.) Ferraris [ E Exosporium mexicanum (Ellis & Everh.) M.B. Ellis] and P pilosa (Earle) Ferraris [ = Morrisog-raphiurn pilosum (Schwein.) Deighton] (see Deighton 1990). Ellis (1971, 1976) and Arx (1983) recognized the genus Phaeoisan'opsis for synnematous hyphomycetes close to Cercospora, but included some species with large, dense, non-synnematous fascicles. Deighton (1990) re-examined and reassessed the genus Phaeoisariopsis. He considered the synnematous arrangement of conidiophores to be a feature unsuitable for generic differentiation. Phaeoisariopsis was therefore confined to a few species similar to l? griseola, characterized by non-geniculate conidiogenous cells with scars lying more or less flat against the conidiogenous cells. Species with conspicuously geniculate conidiogenous cells were placed in Passalora. Braun (1992, 1995a, b) disagreed and preferred to maintain Phaeoisariopsis as the synnematous counterpart to Passalora. The formation of svnnemata is, as the only feature for generic delimitation, very questionable. For instance, the genus Pseudocercospora also includes some synnematous species. Several species originally placed in Phaeoisariopsis, but with inconspicuous conidial scars, have been reallocated to Pseudocercospora (Deighton 1990). There are also some other genera of hyphomycetes with synnematous as well as nonsynnematous species, e.g., Spiropes Cif. (Ellis 1971). The differentiation between Ramularia and Phacellium (Braun 1998) is, however, not comparable since the two genera are also distinguished based on other features. Phaeoisariopsis is undoubtedly heterogenous and not tenable in the present sense as a synnematous counterpart to Passalora. The key for a future treatment of this genus is connected with the status of Phaeoisa~iopsis griseola, the type species. Recent molecular data (Crous et a1 2000), placed P gn'seola in the Pseudocmcospora complex, which is surprising at first sight. The close affinity of I? gmseola and Pseudocercospora spp. is, however, not totally unexpected, since the scars in this species are very thin, unthickened or almost so, and only very slightly darkened. In many collections, the conidiogenous loci range from being inconspicuous to subconspicuous, so that it is not surprising that Yen (Yen and Lim 1980), referred this species to Pseudocercosfiora. Yen introduced, however, the wrong combination [Pseudocercospora columnam's (Ellis & Everh.) J.M. Yen] and cited Isariopsis gn'seola, which is the oldest, valid name for this species, as synonym. However, he did not take into consideration that the genus Phaeoisum'opsis Ferraris (1909) antedates Pseudocerco.sporc~ Speg. (Spegazzini 1910), and that the inclusion of Plzaeoisariopsis gm'seola in the latter genus would reduce Psrudocmcosporn to synonymy with Phaeoisa?-iopsi.~,an action which was already indirectly done by Sawada (1922) who transferred Septonema vitis Lev., the type species of Pseudocercospora, to Phaeoisariopsis. Kecently, type material of Cercospora soli~naniSpeg. (LPS 918) has been re-examined. This species was reduced to synonymy with Phaeoisariopsis griseoln by Deighton (1990). The type material of C. .sol+mnni is also a collection with rather inconspicuous scars. The close affinity of Phaeoisam'opsis and Pseudocercospora was also recognized by Arx (1983). It appears, therefore, that I? griseolq the type species of Phaeoisam'opsis, is congeneric with Pseudocercospora. As in the case for Paracercospora Deighton (Stewart et a1 1999), it appears that genera with pigmented conidia and thin, unthickened or almost so, slightly darkened scars belong in Pseudocercospora. Species with consistent, conspicuously thickened and darkened scars should be assigned to Passalora. h formal merging of Plzaeoisariopsis and Pseudocercospora should, ho~vever,coincide with a proposal to conserve Pseudocercospora over Phaeoisariopsis. Cercospora. The genus Cercospora was introduced by Fresenius (in Fuckel 1863) for passalora-like fungi with pluriseptate conidia. The concept and circumscription of Cercospora was subsequently widened, and culminated in the treatment of the genus by Chupp (1954), which included almost all cercosporoid hyphomycetes. In other treatments the circumscription of this genus was redefined and narro\ved. Saccardo (1880) and Spegazzini (1910) considered Cercosporn ferruganea Fuckel with pigmented conidia to be typical for Cercospora, so that Spegazzini (1910) introduced Cercosporina Speg. for cercosporoid hyphomycetes with hpaline conidia. Deighton (1979) transferred C. ferruginea to ~M~covellosiella.Sutton and Pons (1980) dealt with Cercosf~orinain detail, and reduced this genus to synonymy with (,'rrcospora s.str. Other taxonomic treatments restricted the genus to hyaline-spored, scolecosporous species with conspicuous conidial scars, agreeing with Cercospo7-a apii, which was proposed by Clements and Shear (1931) to serve as lectotype (Muntaiiola 1960, Deighton 1976, Ellis 1976, Sutton and Pons 1980). This taxonoinic concept is now generally recognized and has been confirmed by molecular studies (Crous et al 2000), in which species of Cercospora s.str. formed a well-defined group. C~rrcosporapen icillata (Ces.) Fresen. [ = C. depazroides (Desm.) Sacc.] is, however, the true type species of this genus. The complicated history of its typification has been discussed in detail by Braun (1995a). Braun (1993) referred some species with colorless or subhyaline conidiophores to Cercospora, and pror ~ Braun for posed the subgenus H ~ a l o c r ~ c o s p oU. these taxa. The differentiation between almost or quite colorless C:ercospora spp. (subgeii. Hjc~locerc.osporn) and Cercosporella spp. has been discussed in detail by Braun (199.3a). Braun (1993) discussed the differentiation between C~rcospomand Passalom. C:rreospora s.str. has also been circumscribed by Pons and Sutton (1988), Braun (1993a, h) , and Braun and Melnik (1997). The structure of' the conspicuously thickened and darkened conidiogenous loci (scars) and conidial hila is very important for the identification of C;~rco.cporn spp. Pons et a1 (1985) provided ultrastructural details of these structures in C,'ercospora brticoln Sacc. David (1993) examined scar structures of cercosporoid hyphomycetes by means of SEM ant1 proposed the term " ~ h o s p o r a - t y p e " for taxa ~vith planate scars. I11 our analysis two strains with S r p t o ~ i nanamorphs (CBS 632.85, 183.97) clustered in the f~rrcosporn clade, apart from Septo7ia tm'tici (STE-U 658). In other analyses, however, where more Srjf~toriclisolates Jvere included (G. I'erkley, CBS, the Netherlands, pers cornm), these specific isolates grouped separate from the (~ercosporaclade, and have thus n o t been included in FIG. 3. Although not the focus of the present study, it appears that Septorin anainorphs have e\~olveclfrom several lineages within I\.lyco,~phaerella, and a separate study will be required to address this. Elletevera Lleighton, Eriocercospora Deighton, Fulvia C$, Mycovellosiella, Sirosporium Bubcik & Seraiwinn., Stenella (gen0-u with supe~jficialserondnr~l112'phne 07- Prert, extensiuelj branched conicliophore.r). IWJcovellosiella was established by Rarigel (1917) to replace I+losielln Range1 (1915) (nor1 Baillon 1887, .Scroph~ilnriacene).Muntariola (1960) and Deighton (1974) reintroduced and redefined Rangel's genus. ~ j c o u e ~ ~ o s i rhas l l a been generally recognized as a segregate of (,'ercospo?-a s.lat. for taxa with conspicuous conidial scars and assurgent to repent external secondary hyphae on which the conidiophores are borne terminally and as lateral branches. The conidia are formed singly or in chains. Arx (1974, 1981) reduced Mycovellosiella to synonymy with Cladosporium, but later (Arx 1983) maintained it as a separate genus. Scars of ~Uycovellosiellaare quite distinct from those of Cladospom'um, belonging to the Cercosporatype (David 1993). Ragnhildiana Solheim (1929) and Cercodeuterospora Curzi (1932) were reduced to synonymy with Mycovellosiella by Muntaiiola (1960) and Walkmmyces Thaung (1976) by Deighton (1979). The genus Ormathodium, described by Sydow (1928), was considered a synonym of Mycovellosiella by Muntaiiola (1960), but, unfortunately, type material of 0. stjracis Syd., the type species of this genus, is not preserved, so that this synonymy cannot be confirmed. Muntaiiola (1960) and Deighton (1974) separated Fulvia from Mycovellosiella by differences in habit (loosely fasciculate, without any tendency to form ropes or to ascend leaf hairs in Fulvia), although they emphasized the close resemblance of the two genera. Arx (1974, 1981) reduced Fulvia together with Mycovellosiella to synonymy with Cladosporium, but in 1983, he recognized Mycovellosiella as a separate genus and considered Fulvia a synonym of the latter, since the differences in habit, discussed by Deighton (1974), are not tenable. Several intermediate taxa exist, e.g., iUycovellosiella brachycarpa (Syd.) Deighton (1974) and M . trichostemmatis (Henn.) U . Braun (1999). Braun (1995a) followed Arx's (1983) concept and introduced the combination Mycovellosiella subgen. Fulvia (Cif.) U. Braun. David (1993) examined the structure of the scars in Fulvia and 1Vfjcoz1ellosiella and found some differences. The scars of Fulvia have been referred to the Stenella-type (pileate) and those of Mjcovellosiella to the Cercospora-type (planate). The number of species examined was, however, too small for a final conclusion. Further study of a wider range of taxa are necessary to resolve their scar types. Although Braun (1995a) recognized Fulvia as a subgenus of Mycovellosiella, it should for the interim be retained as synonym of Mycovellosiella. Deighton (1969) described Elletmera and Em'ocercospora for hyperparasitic ~Vycovellosiella(incl. fulvialike) hyphomycetes. Elletmera is characterized by having erect, fasciculate, frequently branched, pigmented conidiophores, slightly thickened and darkened conidial scars, and usually solitary (rarely catenate) conidia. Deighton (1969) compared Elletevera only with Fulvia and Mycovellosiella, and stated that it differs from the latter genus in the fasciculate arrangement of the conidiophores and from the former in the absence of regular catenation of the conidia. However, ~M~covellosiella and Fzclvia are connected by numerous intermediate taxa and have been united in one genus. Elletmera is only distinguished from Mycovellosiella by being hyperparasitic, but there is no clear basis for a morphological differentiation. The hyperparasitic habit alone, without any morphological basis for a differentiation, is not sufficient to introduce or maintain separate genera in the complex of cercosporoid hyphomycetes. Hyperparasitic species are, for instance, also known in Ramularia (Braun 1998) and Cladospom'um (Ellis 1976). Hence, Elletmera should not be maintained as a separate genus. Species of Eriocercospora possess superficial mycelium with solitary, lateral conidiophores, slightly thickened and darkened conidial scars, solitary mycovellosiella/passalora-like conidia, and are indistinguishable from ~Uycovellosiella.Deighton (1969) compared Eriocercospora with the genus Cercospora, and stated that it only differs from the latter genus in having only slightly thickened conidial scars and in its hyperparasitic habit. Therefore, Em'ocercospora cannot be maintained as a separate genus. Sirospom'um (BubBk 1912) is morphologically close to Mycovellosiella and only distinguished by having thick-walled, dictyosporous conidia. Siros~oriumis tentatively maintained as a separate genus until some molecular data become available to establish the value of oblique septa in this complex. Species with thin walls and without dictyospores should be excluded since they are undoubtedly congeneric with Mycovellosiella. Stenella was described by Sydow (1930) and recognized by Ellis (1971, 1976), who reduced Biharia Thirum. & Mishra (Thirumalachar and Mishra 1953) to synonymy with this genus. Deighton (1979) followed this concept of Stenella and differentiated it from il/lvcovellosiella based on the formation of verruculose superficial hyphae and usually roughwalled, catenate conidia. However, many species with conidia formed singly have been assigned to Stenella, so that the verruculose creeping hyphae remain the onlv reliable basis for the differentiation of the two genera. According to David (1993), the scars in Stenella are pileate and differ from the planate Cercospora-type scars. However, too few specimens have been examined to determine if this is a good distinguishing character. Molecular data also showed Stenella parkza Crous and Alfenas (anamorph of Mycosphaerella parkzz Crous & M.J. Wingf.), Stenella marasaszz (Crous & M.J. M'ingf.) B. Sutton & Crous (anamorph of Mycosphaerella marasaszz Crous & M.J. Wingf.) and Mycosphaerella markszz Carnegie & Keane (anamorph unknown) to cluster separately from the M~covelloszella/Passalora/Phaeoramulanaclades (Crous et al 2000, 2001). Stenella a r a p a t a Svd. clusi FIG.3. Clades of Mjcosphaerella spp. identified in the neighbor-joining gene tree derived in FIG. 2, indicating the overlap in conidiomata and conidiophore arrangement (A), conidium shape, septation and arrangement (B), proliferation of the conidiogenous cells (C) , and conidial hila and pigmentation (D) . Clade 1 = Pseudocercospora (incorporating Paracercospora, Phaeoisan'opsis, S t i p i n a , a passalora-like species with slightly thickened scars and Pseudophaeoramulan'a); clade 2 = Pseudocercospora; clade 3 = Passalora (incorporating Mjcovellosiella and Psc.udophaeoramula?-za);clade 4 = Cercospora (presently ters separately from other species of Stenella, however, suggesting that this genus is polyphyletic in Mycosphaerella. The relationship between Stenella and the Mycovellosiella/Passalora/Phaeoramularia complex deserves further investigation. Several intermediates between Stenella and Passalora exist, as there are some Stenella spp. with smooth conidia, etc. It appears therefore that Stenella should also be seen as part of the Passalora complex. The molecular data available in this study, however, are insufficient to resolve this issue. The relationship between Stenella and Stenellopsis Huguenin (1966) is also unclear. Stenellopsis is morphologically similar to Stenella. It has single, conspicuously verrucose conidia with scars that are somewhat thickened and darkened, but lacks verruculose superficial hyphae. However, the formation of superficial hyphae has now been recognized to be of lesser importance at the generic rank. For instance, it is not accepted at generic rank in Pseudocercospora (Deighton 1976) or Ramularia (Braun 1998), and must also be rejected in the iVlycovellosiella/Passalora/Phaeoramularia/Stenella complex. It appears, therefore, that Stenellopsis should also be seen as synonym. Verrucisporota, introduced by Shaw and Alcorn (1993),is similar to Stenellopsis. The scars are broad and of the Cercospora-type, and the conidia are rugose (David 1997). Parastenella J.C. David (David 1991, MorganJones 1998, = Stenellopsis Morgan-Jones 1980, non Huguenin 1966) resembles Stenella and Verrucisporota, but the conidial scars are unthickened, not darkened, and the conidiogenous cells are frequently intercalary and lateral. The fate of the latter two genera remains to be determined. Phaeoramularia and Tandonella S.S. Prasad & R.A.B. Vmma. Phaeoramularia, described by Muntaiiola (1960), was introduced for Ramulam'a-like dematiaceous hyphomycetes with fasciculate, simple or branched conidiophores and has been widely used since (Ellis 1971, 1976, Deighton 1979, Liu and Guo 1982, Pons and Sutton 1988, Hsieh and Goh 1990). Arx (1974) did not recognize the genus Phaeoramularia, and stated that he was not able to distinguish it from M~~covellosiella, Stenella, and other allied genera. In 1983 Arx treated Pizaeoramulam'a as a separate genus, but emphasized that this genus is morphologically not clearly separable from Mycovellosiella. Braun (in Braun and Melnik 1997) reallocated Fusicladium leviem' Magnus ( = E: kaki Hori & Yoshino) to Phaeoramularia and reduced Hormocladium Hohn. (type species: Fusicladium kaki; Hohnel 1919) to synonymy with Phaeoramularia. Phaeoramularia (1960), would be a later facultative synonym of Hormocladium, but monographic studies in Fusicladium, including SEM examinations, recently carried out by U. Braun and collaborators, showed that F: lmiem' is a true Fusicladium belonging to a group of species with catenate conidia. Hence, Hormocladium must be reduced to synonymy with Fusicladium and does not pertain to Phaeoramularia. Tandonella Prasad & Verma (1970) was introduced for a synnematous hyphomycete with thickened, darkened scars and hila and pigmented, catenate conidia. Sutton and Pascoe (1987) added a new species, compared Tandonella with Sclerographiopsis Deighton, and emended the circumscription of the two genera. Tandonella is a synnematous counterpart of Phaeoramularia, also having superficial mycelium. However, as stated earlier, both these features should not be employed at generic level, so that Tandonella is not tenable any longer as a separate genus. Reassessment of Mycovellosiella, Passalora, Phaeoramularia, and allied genera. Passalora and Phaeoramularia are only differentiated by the mode of conidial formation, either formed singly or in chains. Conidial formation is, however, a weak feature to employ at the generic level in this complex. Ramularia comprises, for instance, species with solitary as well as catenate conidia. Braun (1998) clearly demonstrated that the conidial formation (solitary vs catenate) is not applicable within Ramularin slat. for a separation of the genus into smaller taxonomic units. In Pseudocercospora, the conidia are usually formed singly, but a few species with catenate conidia are also known (Deighton 1976). Conidia in short chains are, for instance, known for Pseudocercospora nq'ctnnthis (A.K. Roy) U. Braun, Bagyan. & Jagad. (Braun et a1 1992), I? millettine Goh & W.H. Hsieh and l? noveboracensis Goh & W.H. Hsieh (Hsieh and Goh 1990). Pseudocercosporella Deighton is also a genus that usually forms solitary conidia, but a few taxa with catenate ones have also been included (Deighton 1973). Even in Cercospora s.str. the conidia may occasionally be formed in short chains [e.g., in C:Prcospora lartu- incorporating two septoria-like isolates that separate in other analyses, and thus do not belong in this clade, see text); clade 5 = Passalora (incorporating M~couellosiellaand Phaeoramularia, and one isolate of Thedgonia; clade 6 = Ramularia (incorclade 10 = Lecanosticta; porating Ouularia and Ophiocladium); clade 7 = unknown; clade 8 = Septona; clade 9 = Uwebraz~nia; clade I1 = Stenella; clade 12 = Phaeophleospora (incorporating Colletogloeopsisand Scokcostigmina) ; clade 13 = CT7uehaunia; clade 14 = Stenella; clade 15 = Di.rsoconium; clade 16 = Cladosporium (including Hetm-osporium). cue-satiuae Sawada (Hsieh and Goh 1990, as C. longissima Sacc.) and in C. ricinella Sacc. & Berl. (Braun et a1 1992)l. The formation of short conidial chains in Cercospora s.str. is often connected with the germination of conidia and microcyclic conidiation. This phenomenon is rather common and has been described and illustrated for many species, e.g., C. sesamigenaJ.M. Yen & Lim (1973), C. nurantia Heald & F.A. Wolf (Pons 1988) and Cercospora kikuchii T. Matsumoto & Tomoy. (Fernandez et a1 1991). Both 1W~covellosiellaand Stenella include species with solitary and catenate conidia. There are even several intermediate taxa in Pnssalora which occasionally form short conidial chains, e.g., Passcclo'ra janseana (Racib.) U. Brawl ( = C;ercospora orjzae Miyake) [see Constarltinescu 1975, under C;ercospora jun,seana (Racib.) Constant.], P heterospora (Hohn.) Hohn. (= Phaeoramularia hoehnelii S. Petzoldt, in Braun 1992) and R aratai (Speg.) U. Braun, Delhey & Kiehr (2000). Pons and Sutton (1996) described Cercosporidium dezghtonii, which was referred to Cerco.spom'dium ( = Passalora) although the conidia in this species are fomed in chains. In conclusion, it must be stated that among cercosporoid hyphomycetes the formati011 of single or catenate conidia is not tenable as a distinguishing character at generic rank. Phaeoramularia has to be reduced to synonymy with Passalora, which has also been confirmed by our molecular data, in which Passalora, Phaeoramularia and iVycovellosiella form mixed clusters. The differentiation between Passalora (incl. Phaeoramularia) and 1Wlycovellosiella (incl. Fulvia) , which comprises taxa with solitary as well as catenate conidia, is also difficult (Arx 1983, Pons and Sutton 1988). Various intermediate taxa exist, e.g., iVl~couellosiella Drachycarpa (Syd.) Deighton (Arx 1983), iM. solanitorvi (Gonz. Frag. & Cif.) Deighton (1974) and kf. tm'chostemmatis (Henn.) U . Braun (1999). Fuluia forms transitions from iMycovellosielln to Pussalora (incl. Phaeoramuluria) . The development of superficial secondary mycelium with solitary conidiophores is also a variable character. Deighton (in Ellis 1976) assigned, for instance, CPrcosp~ratithonine Baker & Dale to Phaeoramularia, but Hsieh and Goh (1990) found external hyphae with solitary conidiophores in this species. C:~rcospora eupatorii-odorati J.M. Yen has been placed in 1M~co7)ello.riella (Yen 1981) as well as Plzaeoramulam'a (Liu and Guo 1982). Threads with terminal, intercalary, and pleurogenous conidiogerlous cells may range in this group of genera from being decumbent, developing into creeping secondary hyphae, to erect, fasciculate, forming ropes or climbing leaf hairs, but all kinds of transitions mav exist. In culture, all differences in habit disappear. Therefore, the develop- ment of creeping superficial hyphae is, as in Pseudocercospora (Deighton 1976), Cercosporella (Braun 1995a), and Ramulnvia (Braun 1998), not tenable for the separation of cercosporoid genera. Even in Cercospora s.str., creeping hyphae with solitary conidiophores may be found [e.g., in Cercospora canescens Ellis & G. Martin (Yen and Lim 1969) and C. hjalofilispora J . M . Yen (1966)l. The main character that connects ~WjcovellosielLaand Fulvia is the formati011 of conidiogenous cells, which are integrated, terminal, intercalary pleurogenous, and often formed as short lateral branches or shoulders immediately below transverse septa. However, even in Phaeommz~laria g o m p h r ~ ~ z a(Speg.) e Munt.-Cvetk., the type species of Phaeoramula~ia[holotype (LPS 914), isotype (IMI 7706) examined], the fLll range of rnyco\~ellosiella/ fulvia-like conidiogenous threads have been found, with loosely fasciculate to decumbent, frequently branched conidiophores having terminal, intercalary as well as pleuroge~lousconidiogenous cells (Muntaiiola 1960, Deighton 1979, Pons and Sutton 1988). Hence, the type species of Phaeomrnularia is indistinguishable from LWl~cov~llosi~lla (incl. Fuluicc). Similar Phaeoramularia spp. with frequently branched conidiophores are rather common, e.g., Ph. acanthicoln (Hansf.) Deighton (1987), Plz. isotomae Deighton (1986), Ph. cz~curbiticola(Henn.) Deighton, Ph. lom ~ n s iD s eighton, E'h. sudanensis Deighton (Deighton 1979), Ph. parccdoxci Munt.-Cvetk. and I'h. iresines Munt.-Cvetk. (Muntaiiola 1960). Branched erect or decumbent threads are not useful as a character for taxoiiomic purposes on a generic level in cercosporoid hyphomycetes. Branched conidiophores are even present in Passalom baeillig~ra,the type species of Passalora, in P microsper~naFuckel and f? chaptomi u m (Cooke) , b x (Deighton 1967). There is not a true basis for the separation of Passalorn (incl. Plzaeommularia) and i~fycovellosiella(incl. F u k ~ i n,) so that the latter genus must also be included in Pu.rsalora slat., which is in agreement with our molecular data. Pa.rsalora emend. reveals the same range of morphological variation as observed in Ramularia and P.\eudocercospora. I-'assalo.ra may therefore be circumscribed as f o l l o ~ ~ s : Passalora Fr., Summa \reg Scand p. 500. 1849. emend. Cercosporidiun~Earle, Muhlenbergia 1: 16. 1901. lhllosi~llaRangel, Bolm Agric S Paulo, Ser 16 A, 2: 151. 1915, homonynl. ~~lycovellosiella Rangel, Arch Jard Bot Rio de Janeiro 2: 71. 1917. = C~rcodeuterospornCurzi, Boll Staz Patol \reg Roma, Ser 2. 12: 149. 1932. = R ~ g n l ~ l l d z c c nSolheirn, o hf~l\colog~a 23: 365. 1931. = = - = = Berteromyces Cif., Sydowia 8: 267. 1945. Fulvia Cif., Atti 1st Bot Univ Lab Critt Pavia, Ser 5, 10: 245-246. 1954. = Mycovellosiella subgen.Fulvia (Cif.)U. Braun, A monograph of Ccrcosporella, Rumularia and allied genera (phytopathogenic hyphomycetes),Vol. 1: 39. 1995. = Oreophylla Cif., Sydowia 8: 259. 1954. = Phaeoramularia Munt.-Cvetk.,Lilloa 30: 182. 1960. = Eriocercospora Deighton, Mycol Pap 118: 3. 1969. = Elleteueru Deighton, Mycol Pap 118: 17. 1969. = Tandonella Prasad and Verma, Indian Phytopathol 23: 111. 1970. = Phaeoisariopsis p.p. Anamorphs of Mycosphaerella spp., usually on living hosts, phytopathogenic, often causing leaf spots, occasionally hyperparasitic, rarely saprobic. Primary mycelium internal, secondary mycelium lacking to well-developed, external, superficial; hyphae branched, septate, smooth, hyaline to pigmented. Stromata absent to well-developed, substomatal to intraepidermal, rarely deeply immersed, subglobose to applanate, subhyaline to pigmented, composed of loosely to densely aggregated swollen hyphal cells. Conidiophores solitary, loosely to densely fasciculate or in sporodochial to synnematous conidiomata, arising from internal or superficial hyphae or substomatal to intraepidermal stromata, emerging through stomata, erumpent through the cuticle or arising from creeping hyphae, terminal or as lateral branches, conidiophores macronematous, unbranched or branched, continuous to pluriseptate, subhyaline to pigmented, smooth to finely verruculose; conidiogenous cells integrated, terminal, intercalary to pleurogenous or conidiophores reduced to conidiogenous cells, conidiogenous loci conspicuous, conidiogenous cells cicatrized, scars somewhat thickened and darkened-refractive, more or less applanate. Conidia solitary to catenate, in simple or branched chains, amerosporous to scolecosporous, aseptate to pluriseptate, euseptate, rarely with a few additional distosepta, pale to distinctly pigmented (if subhyaline, conidia non-scolecosporous, broad, 4-15 km diam., and with few septa, usually 0-4), smooth to finely verruculose, hila somewhat thickened and darkened-refractive, more or less truncate. Pseudocercospora and allied genera.-Pseudocercospora was introduced by Spegazzini (1910). Deighton (1976) re-introduced this forgotten name and widened the concept of this genus considerably to include a wide range of cercosporoids with inconspicuous scars. He reduced Helicominia L.S. Olive, Ancylospora Sawada and Cercocladospora G.P. Agarwal & S.M. Singh to synonymy with Pseudocercospora. Deighton (1976) distinguished Cercoseptom'a Petr. from Pseudocercospora by having narrow, acicular conidia, but both genera cannot be properly differentiated (Deighton 1987, Braun 198%). Arx (1983) merged Cercoseptom'a, which is characterized by having pigmented conidiophores, with the colorless genera Pseudocercosporella and Thedgonia B. Sutton. Presently Cercoseptoria is accepted as a synonym of Pseudocercospora (Hsieh and Goh 1990, Guo and Hsieh 1995, Crous and Braun 1996, Braun and Melnik 1997), which is also supported by the molecular data derived by Crous et a1 (2000) and in the present study. Pseudocercospora is morphologically highly variable, and accommodates a wide range of cercosporoid hyphomycetes with pigmented conidiophores and inconspicuous, unthickened, not darkened conidiogenous loci. Braun (1995a) demonstrated that all other characters are ~ariableand unsuitable for consideration at generic rank. Braun (1998) proposed a formal division of Pseudocercospora into five sections, based on morphological features of the conidiomata and conidia, but emphasised that these "units" should only be regarded as morphological groups, which are formally treated as sections, independent of their taxonomic value. The molecular data derived by Crous et a1 (2000) and in the present study show Pseudocercospora to be polyphyletic within Mycosphaerella, having evolved more than once from different Mq'cosphaerella holomorphs, and in several occasions having lost the teleomorph. This complex includes species with solitary or catenulate conidia, having euor distosepta, and scars which are inconspicuous to denticle-like, unthickened or almost so, not to slightly pigmented-refractive. Paracercospora Deighton (1979) was introduced for some cercosporoid hyphomycetes with subconspicuous conidial scars (thickened slightly along the rim). Paracercospora fijiensis (Morelet) Deighton was originally referred to as a species of Pseudocercospora (Deighton 1976). The differentiation of Pseudocercospora and Paracercospora is often difficult, and there are numerous intermediate taxa in Pseudocercospora that occasionally form paracercospora-like scars. In molecular analyses (Crous et a1 1999, Stewart et a1 1999, Crous et a1 2000), Paracercospora spp. grouped with Pseudocercospora spp. These results supported the synonymy of these two genera. Braun and Melnik (1997) included the description of the new genus Pseudophaeoramulam'a U. Braun, which contained cercosporoid hyphomycetes intermediate between Pseudocercospora and Phaeoramularia. The conidia are catenate, but the conidial scars are subconspicuous, flat, truncate, minutely thickened, not darkened to somewhat darkened and refractive. Pseudophaeoramulam'n geranii (M7.B. Cooke & C.G. Shaw) U. Braun, the type species of this genus, was originally placed in Pseudocercospora by Braun (1991). Based on our molecular data, Pseudopha~oramularia should be reduced to synonymy with Pseudocercospora. These results suggest, therefore, that the separation of taxa with unthickened scars, and minutely thickened scars, are not informative at the generic level. The "type" of scar thickening (inconspicuous, planate, pileate or protruding) could therefore be more important than previously accepted. These results suggest that the separation of taxa with unthickened, non-pigmented scars and almost unthickened, slightly pigmented-refractive scars are not informative at the generic level. Denticularia Deighton (1972) and Semipseudocercosporn J.M. Yen (1983) seem to be closely allied to Pseudocercospora, but the conidiogenous cells are distinctly denticulate and the conidia are non-scolecosporous. Because some species of Pseudocercospora also form denticles (Crous 1998), it may mean that these genera will eventually also be reduced to synonymy. Presently no cultures are available for study and hence this remains unresolved. Deighton (1976) recognized Pantospora Cif. (Ciferri 1938) as a separate genus close to Pseudocerco~porn, distinguished by synnematous conidiomata, percurrent as well as sympodial conidiogenous cells, and frequently dictyosporous conidia. These differences are, however, rather vague. The formation of dictyospores is the only clear character which distinguishes Pantosporn from Pse~~docercospo'ra, although conidia of Ps~udocercosporauitis (Ltv.) Speg., the type species of this genus, occasionally possess a few oblique or transverse septa. Molecular data on the species concerned are not available and the taxonomic value of dictyospores as a generic feature is not quite clear. As this feature can occur in the type of Ps~udocercosporn,however, it appears that Pant0.sporn may also represent a synonym of Pseudocercospora. Stigrninn plntani (Fuckel) Sacc., the type species of Stigminn, which has phragmo- and dictyospores, as well as eu- and distoseptate conidia, also clustered in the larger Pseudoem-cosporu clade in the present study. Furthermore, Ellis (1971) also lists several genera such as Annellophora S. Hughes, P ~ ~ z ~ d o s p i r o M.B. pes Ellis, Spiro@esCif. and S p o r i d ~ s m i u mLink, where taxa occur that have a mixtue of eu- and distoseptate conidia. Braun (1993) introduced Cerco.r.ticpinnfor some cercosporoid hyphomycetes that have been segregated from Stigrnina Sacc. Species of this genus are morphologically close to Pseudocercospora spp. (scars inconspicuous), but distinct by having consistently percurrently proliferating conidiogenous cells. However, the combination of sympodial and percurrent conidiogenous cells is not uncommon in P.~eudoc~rro.~po- ra. Some Cercostigmina spp. have ~Mycosplzaerellateleomorphs. One such an example is (=ercostigminn punctata (N7akef.) Crous (ibl. syzjp'i Crous) (Crous 1999). In a recent ~nolecularstudy comparing the taxa occurring on Eucaljptus, Crous et a1 (2001) included one isolate of this fungus (STE-U 1124), which only produces conidia on percurrently proliferating conidiogenous cells in culture, though also sy~npodiallyon the host (Sutton and Crous 1997). This species, belonging to Cercostigmina s.str., clustered with 97% bootstrap support to Pseudorercos~om, suggesting that C~rco.stigminnshould be reduced to synonymy M Jth ~ J-'Sez~do(~rcospom. Stigminn Sacc. (Saccardo 1880) was recently circumscribed by Sutton and Pascoe (1989) to represent foliicolous species allied to S. platani with pigmented structures, percurrently proliferating conidiogenous cells, and transversely, occasionally longitudinally distoseptate conidia. Genera that have since been segregated are Cercostigrnina ( = Pseudocrrco.sj)orr, see above), X e n o s t i p i n a Crous (19981, with euseptate conidia and sympodial and percurrently proliferating conidiogenous cells and Scoleco.stipinn LT. Braun. Scolerostigminn (Braun et a1 1999) was introduced for stigrnina-like species with thick-walled structures, and conidia having transverse eusepta, occasionally with a few longitudinal septa. One isolate of Stigmirta plrtani (CBS 336.33) that was included in this study clustered in the Pseudocerro.sporn clade. As with Cmostigmina, it supports the contention that the strictly percurrent proliferation of conidiogenous cells should be seen as a feature of value at species level. Furthermore, it also suggests that distoseptatiorl and oblique conidial septa are not evolutionarily informative in this group. Presently no species of S t i p i n a s.str. has a known L\f~cosphaerellu teleomorph. These findings suggest, however, that the teleornorphs, if found, ivould reside in lL1ycosphaerelln. One problem with placing Stigrnina in the Pseudocer.co.~porrgroup is that Stignintl (1880) predates Psrudoc~rco.sporu (l910), so this synonymy will have to be accompanied with a proposal to retain P.r.~zrdocercospomover Stignzina. Sutton and Hennebert (1994) commented that sporodochia are not fundamentally different from acer\uli and thus S l i p r ~ i n ais not that distinct from Leranosticta Syd. The fact that Lecnnosticta rricolc~ (Thiim.) Syd. clustered with other anamorphs of ~\{ycosphccrrella, casts doubt on the validity of Eruptio M.E. Barr as proposed by Barr (1996). More species will have to be included to resolve the fate of this genus, and to determine if it is homogenous. Sonderhenia H,]. Swart and J . Walker (1988) was established for pycnidial ananlorphs of ~\.Ijro~phaereI/n with pigmented structures, percurrently prolifer- ating conidiogenous cells and distoseptate conidia. Although similar to Phaeophleospora Rangel, it is chiefly distinguished by its distoseptate conidia. In a study comparing sequence data of Sonderhenia, it was shown to cluster between a Pseudocercospora and Mycovellosiella clade (Crous et a1 2001), further casting doubt on the value of distoseptation as character in this complex. Phaeophleospora was established by Rangel ( 1916) for pycnidial anamorphs of Mycosphaerella that have pigmented non-multi-euseptate, pigmented, smooth to rough conidia that form on pigmented, percurrently proliferating conidiogenous cells (Taylor and Crous 1999). Recently, however, collections were obtained of a species with conidiogenous cells that could also proliferate sympodially (Crous unpubl) . Colletogloeopsis Crous & M.J. Wingf. (Crous and M'ingfield 1997) represents acervular anamorphs of Mycosphaerella with brown, 0(-1)-septate, verruculose conidia that form on brown, percurrently or sympodially proliferating conidiogenous cells. The fact that species of these genera clustered together with high bootstrap support in the phylogenies derived from both ITS and LSU data (Crous et a1 2001) suggests that conidiomatal structure and conidial septation are of lesser importance in this complex than conidial pigmentation and conidiogenesis. Presently no cultures are available of Xenosti,pina Crous, thus its position cannot be clarified. However, given the flexibility of percurrent and sympodial proliferation occurring in some of the Mjcosphaer~llaanamorphs, the separation of this genus needs to be reconsidered. Crous and Corlett (1998) also recently described a Cercostipina synanamorph for Xenostigmzna wolJii Crous & Corlett. Given the variation in mode of proliferation, and the fact that oblique septa can also occur in Pseudocercospora, it appears probable that Xenostipina, like Cercostipina, could be a synonym of the Pseudocercospora/Stipina complex. A species that has to be placed in Scolecostipina U. Braun (sporodochia, pigmented structures, percurrently proliferating conidiogenous cells, conidia subcylindric-obclavate, transversely multi-euseptate) is S t i p i n a eucalypti Alcorn (CBS 313.76). This species, however, clustered with Colletogloeopsis (acervuli, pigmented structures, sympodial and percurrently proliferating conidiogenous cells, conidia subcylindrical, 0-1-euseptate) and Phaeophleospora (pycnidia, pigmented structures, sympodial and percurrent proliferating conidiogenous cells, conidia subcylindrical to obclavate, 0-multi-euseptate). If these genera were to be considered objectively the main distinguishing factor is their conidiomatal structure. To merge them into the oldest genus name, Phaeophleospora, would be to disregard conidial septation and conidiomatal structure. The abandonment of these features has already been argued elsewhere in this paper. These preliminary data also support this synonymy. How ever, to complicate matters, the type of Phaeophleospora, P eugeniae Range1 clustered apart from this group in a study by Crous et a1 (2001).It would seem, therefore, that although the synonymy proposed above could be feasible, it may have to await the inclusion of more species and other data sets to be finalized. In the interim, the following genera can be seen as part of the Pseudocercospora complex (with Phaeoisariopsis and Stigmina awaiting proposals to retain Pseudocercospora): Pseudocercospora Speg., Ann Mus Nac Hist B Aires 20: 438. 1910; emend Deighton (1976). Cercospo~iopszsMiura, Flora of Manchuria and East Mongolia, 3, Cryptogams 527. 1928 (nom. illegit.). = Cercoseptona Petr., Ann Mycol 23: 69. 1925. = Pantospora Cif., Ann Mycol 36: 242. 1938. = Anglospora Sawada, Rep Govt Res Inst Formosa 87: 78. 1944 (nom. illegit.). = Helicomzna L.S. Olive, Mycologia 40: 17. 1948. = Cercocladospora G.P. Agarwal & S.M. Singh, Proc Nat Acad Sci India, B, 42: 439. 1972 (1974), fide Deighton (1976). = Paracercospora Deighton, Mycol Pap 144: 47-48. 1979. = = = Cercost~grnznaU . Braun, Crjpt Bot 4: 107. 1993. Ps~udophaeoian~ulana U. Braun, Proc K o m a r o ~Bot Int 20: 18. 1997. The genera Dzssoconium de Hoog, van Oorschot & Hijwegen (1983) and Uwebraunia Crous & MJ. Wingf. (Crous and Wingfield 1996) are two morphologically indistinguishable anamorph genera linked to Mycosphaerella. Crous et a1 (1999) compared these taxa on a molecular basis, and concluded that they should be retained as separate entities, as they were phylogenetically distinct. It is interesting to note, however, that Dissoconium has a hyperparasitic habit, compared to the phytopathogenic one of Uwebmunia. As mentioned above for Elletevma and Mycovellosiella, several cercosporoid genera are now known that include saprobic, phytopathogenic, and hyperparasitic species. Furthermore, the two species of Uwebraunia also cluster separately (Crous et a1 1999, 2000), suggesting that this form has evolved more than once in Mycosphaerella. However, species of Dissoconium always cluster together, forming a clade adjacent to the main Mycosphaerella clade, and is an example of convergent evolution in the anamorph form. Conclusions.-Based on sequence data derived from the ITS and LSU regions in the present and previous studies (Stewart et a1 1999, Crous et a1 1999, 2000, 2001), the monophyly of Mycosphaerella is supported. Presently these data also support the separation of 1Mycosphaerella into three sections, namely the main Mjcosphaerella section, a section for species with Dissoconium anamorphs, and a section for taxa with Cladosporium anamorphs. Approximately 23 anamorph genera have thus far been linked to iVycosphaerella based on cultural data. Of the 19 anamorph genera investigated here, it appears that many are polyphyletic, and do not represent natural groups within iVycosphaerella (FIG. 3 ) . Furthermore, it should be renlembered that anamorphs are form genera. The only groups that were found to be clearCladospom'um ly defined were Cercospora, Ram~~lam'a, and llissoconium. As more species are added to the phylogeny, however, this could possibly also result in more polyphyletic groups, as is the case with the other anamorph genera. To revisit the genus-for-genus concept, one would assume that iVycosphaerella, being monophyletic, should have one anamorph genus. However, this approach would lead to a gross loss of valuable taxononlic inforrnation that will not serve mycology or plant pathology, and will have serious implications. The solution, therefore, is to propose a simplified but workable system, using the characters shown here to remain informative at the generic level. From our preliminary data we conclude that the following features are not useful for generic separation of 1Vjcosphaer~llaanamorphs: 1) Superficial mycelium with solitary conidiophores, and the surface texture thereof (it should be noted, however, that the separation of Stenella and Passalorn is still not clearly resolved); 2) The formation of stromata; 3) Conidiomatal structure (conidiophores solitary, fasciculate to synnematous, sporodochia to pycnidia and acermli) (FIG.3) ; 4) Conidial shape, size, and septation (even eusepta vs distosepta), as well as solitary vs catenate conidia; 5) Saprobic, hyperparasitic, and phytopathogenic habit. Characters of value at the generic level include: 1) Structure of conidiogenous loci (scars) and hila [unthickened or almost so, but slightly darkened appears to have the same value as being unthickened; note a Passalol-a sp. with such scars (STE-U 2554) in the Ps~udocercosporagroup]. Pigmentation and thickening of scars form a continuum between genera. The distinction should thus be between the degree of thickening, darkening, and refraction, building on the concepts developed by David (1997); 2) Presence or absence of pigmentation in conidiophores and conidia [some overlap exists here, and needs clarification, i.e., Passalora and C;wcosporu. Also note Thedgonia (STE-U 1661) in the Passalora group]. These conclusions are still preliminary, and as more taxa are sequenced and other genes also investigated, we hope that this will become clearer. The philosophy follo~iedhere thus supports a reduction in genera, and a widening of anamorph generic concepts within iWycosphaer~lla,which presently still appears to be monophyletic. The next steps would be to focus on specific anamorph genera of 1Vljcos~hnuella that could represent single groups, obtain more isolates of these taxa, and investigate other areas of their genomes. LITERATURE CITED h x Jhvon. 1949. Beitrage zur Kenntnis der Gattung Mycosphaerella. Sydowia 3:28-100. . 1974. The genera of fungi sporulating in pure culture. 2nd ed. Vaduz: Cramer. 315 p. . 1981. The genera of fungi sporulating in pure culture, 3rd ed. Vaduz: Cramer. 424 p. . 1983. i4fycosphaerella and its anamorphs. Proc K Ned Akad Wet Ser C Biol Med Sci 86(1):1.3-54. Barr ME. 1996. Planistromellaceae, a new family in the Dothideales. Mycotaxon 60:433-442. . 1972. Preliminary studies on the Dothideales i11 temperate North Anlerica. Contr Univ Mich Herb 9: 523-638. Bisby GR, iiinsworth GC. 1942. The rlulnbers of fungi. Trans Br Mycol Soc 26:16-19. Braun U. 1988a. Studies on Ra?nulam'a and allied genera (I). Int J Mycol Lichenol 3:271-285. . 1988b. Studies on Ra~nulariaand allied genera (11). Nova Hedwigia 45:333-349. . 1991. St~ldieson Ramularia and allied genera (K). Nova Hedwigia 53:291-305. . 1992. Taxonomic notes on some species of the CB-cospora-complex. Nova Hedwigia 55:211-221. . 1993. Kew genera of phytopathogenic Deuteromycetes. Cryptogam Bot 4:107-114. . 1995a. A nlonograph of Cm-cospo~~lla, Rnmz~laria and allied genera (phytopathogenic hyphomycetes). Vol. 1. Eching: IH\V Verlag. 333 p. . 1995b. Miscellaneous notes on phytopathogenic hyphomycetes (11). hlpcotaxon ,35223-241. . 1996. Taxonomic notes on some species of the Cur-cospora-complex (n'). Sydowia 48:205-2 17. . 1998. A monograph of Crr.cosporella, Ramularia and allied genera (phytopathogenic hyphomycetes). Vol. 2. Eching: IHW Verlag. 493 p. . 1999. Taxonomic notes on some species of the Curcospora complex (W). Cryptogam Mycol 20:155-177. , Bagayanarayana G, Jagadeeswar P. 1992. Notes on Indian Cercosporae and allied genera (11). Int J Mycol Lichenol 4:361-374. , Delhey R, Kiehr M.2000. Some new cercosporoid hyphomycetes from Argentina. Fungal Div 6: 18-33. , Melnik VA. 1997. Cercosporoid fungi from Russia and adjacent countries. Trudy Bot Inst Im VL Komarova (St. Petersburg) 20:l-130. , Mouchacca J, McKenzie EHC. 1999. Cercosporoid hyphomycetes from New Caledonia and some other South Pacific islands. N Z J Bot 37:297-327. BubLk F. 1912. Einige neue Pilze aus RuBIand. Hedwigia 52: 265-273. Castaiieda RF,Braun U. 1989. Cercospora and allied genera of Cuba (1). Cryptogam Bot 1:42-55. Chupp C. 1954. A monograph of the fungus genus Cercospora. Ithaca, New York: Published by the author. 667 p. Ciferri R. 1938. Mycoflora domingensis exsiccata. Ann Mycol 36:198-245. Clements FE, Shear, CL. 1931. Genera of Fungi. New York: Wilson. 496 p. Constantinescu, 0 . 1975. Studies on Cercospo'ra and similar fungi 11. New combinations in Cercospora and Mycouellosiella. Cryptogam Mycol 263-70. Crous PW. 1998. Mycosphaerella spp. and their anan~orphs associated with leaf spot diseases of Eucalyptus. Mycol Mem 21:l-170. . 1999. Species of n/Iycosphaerella (Ascomycetes) and related anamorphs occurring on Myrtaceae (excluding Eucalyptus). Mycol Res 103:607-621. , Aptroot A, Kang JC, Braun LT,Wingfield hfJ. 2000. The genus 1~cosphaerellaand its anamorphs. Stud Mycol 45: 107-121. , Braun U. 1996. Cercosporoid fungi from South Africa. Mycotaxon 57:233-321. , Corlett M. 1998. Reassessment of iMycosphaerella spp. and their anamorphs occurring on Platanus. Can J Rot 76:1523-1532. , Hong L, Wingfield BD, Wingfield MJ. 2001. ITS rDNA phylogeny of selected 1Mycosphaerella spp. and their anamorphs occurring on Myrtaceae. Mycol Res 105:425-431. --, Wingfield MJ, &ng JC. 1999. Uwebrnunia and Dissoconium, nvo morphologicaIly similar anamorph genera with distinct teleomorph affinity. Sydorvia 52:155-166. , Phillips AJL, Wingfield MJ. 1992. Effects of ctlltural conditions on vesicle and conidium morphology in species of Cylindrocladium and Cylindrocladiella. Mycologia 84:497-504. , Wingfield MJ. 1996. Species of Mycosphaerella and their anamorphs associated with leaf blotch disease of Eucalyptus in South Africa. Mycologia 88:441-458. --. 1997. Colletogloeopsis, a new coelomycete genus to accommodate anamorphs of two species of iMycosphaerella occurring on Eucahptus. Can J Bot 75: 667-674. Curzi M. 1932. De fungis et morbis africanis. I. De quibusdam hyphomycetibus parasitis Somalie. Bot R Staz Pato1 Veg N S 12:149-168. David JC. 1991. Parastenella, a new generic name for Heterosporium map~oliae.Mycol Res 95: 123-1 28. . 1993. A revision of taxa referred to ffetmosporium Wotzsche ex Cooke (Mitosporic Fungi) (PhD Thesis). UK: Universi? of Reading. 290 p. . 1997. contribution to the systematics of Cladosporium. Revision of the fungi preciously referred to Heterosporium. Mycol Pap 172:1-157. Deighton FC. 1967. Studies on Cercospora and allied genera. 11. Passalora, Cercosporidium and some species of Fusicladium on Euphorbia. Mycol Pap 112:l-80. . 1969. Microfungi. nr:some hyperparasitic hyphomycetes, and a note on Cercosporella uredinicola Sacc. Mycol Pap 188:l-41. . 1972. Four leaf-spotting hyphomycetes from Africa. Trans Br Myco Soc 59:419-427. . 1973. Studies on Cercospora and allied genera. n'. Cercosporella Sacc., Pseudocmosporella gen. no% and Pseudocercosporidium gen. nov. Mycol Pap 133:l-62. . 1974. Studies on Cercospora and allied genera. V. rMycouellosiella Rangel, and a new species of Ramulariopsis. Mycol Pap 137:l-75. . 1976. Studies on Cercospora and allied genera. W. Pseudocercospora Speg., Pantospora Cif. and Cercoseptoria Petr. Mycol Pap 140:l-168. . 1979. Sudies on Cercospora and allied genera. WI. New species and redispositions. Mycol Pap 144:l-56. . 1986. Misidentification of Cercospora effusa. Trans Br Mycol Soc 86:637-641. . 1987. New species of Psez~docercosporaand Mycoriellosiella, and new combinations into Pseudocercospora and Phaeornmularia. Trans Br Mycol Soc 88:365-391. . 1990. Observations on Phaeoisanopsis. Mycol Res 94:1096-1102. Earle FS. 1901. Some fungi from I'uerto Rico. Muhlenbergia 1:lO-17. . 1902. A much-named fungus. Torreya 2:159-160. Ellis MB. 1971. Dematiaceous hyphomycetes. Kew: CMI. 608 p. . 1976. More dematiaceous hyphomycetes. Kew: CMI. 507 p. Fernandez FA, Glawe DA, SinclairJB. 1991. Microcycle conidiation and nuclear beha~iorduring conidiogenesis in Cercospora kikuchii. Mycologia 83:752-757. Ferraris T. 1909. Osservazioni mycologische. Su specie del gruppo Hyphales (Hyphomycetae) . Ann Mycol 7:273286. Fisher NL, Burgess LW, Toussoun TA, Nelson PE. 1982. Carnation leaves as substrate and for preserving cultures of Fusarium species. Phytopathologv 72: 151-153. Fries EM. 1849. Summa vegetabilium Scandinaliae. Uppsala: Sectio posterior. 313 p. Fuckel KWGL. 1863. Fungi Rhenani exsiccati, Fasc. I-IV. Hedwigia 2:132-136. Guo YL, Hsieh WH. 1995. The genus Pseudocercospora in China. Mycosystema Monographicum 2:l-388. Ho MH, Castafieda RF, Dugan FM, Jong SC. 1999. Cladosporium and Cladophialophora in culture: descriptions and an expended key. Mycotaxon 72:113-157. Hohnel F \on. 1919. Fiinfte vorlaufige Mitteilung mycologischer Ergebnisse (Nr. 399-500). Ber Dt Bot Ges 37: 153-1 61. Hoog GS de, Gukho E, Masclaux F, Gerrits AHG van den, Kwon-Chung EJ, McGinnis MR. 1995. Nutritional phys- iology and taxonomy of human pathogenic Cladosporium-Xylohypha species. J Med Vet Mycol 33:339-347. , Oorschot CAN van, Hijwegen T. 1983. Taxonomy of the Dactylaria complex. 11. Dissoconium gen. nov. and Cordana Preuss. Proc K Ned Akad Wet Ser C Biol Med Sci 86(2):197-206. Hsieh N'H, Goh TK. 1990. Cwcospora species and similar fungi from Taiwan. Taiwan: Maw Chang Book Company. 376 p. Hughes SJ. 1949. Studies on some diseases of sainfoin (Onobrychis satiua) 11. The life history of Ramularia onobrychidis Allescher. Trans Br Mycol Soc 32:3439. . 1958. Revisiones hyphomycetum aliquot cum appendice de nominibus rejiciendis. Can J Bot 36:727836. Huguenin B. 1966. Micromycetes du Pacifique Sud (Troisieme contribution). (1) D6matii.e~de Nouvelle-Caledonie. Bull Soc Mycol Fr 81:686-698. Kendrick NB,DiCosmo F. 1979. Teleornorph-anamorph contlections in ascomycetes. In: Kendrick W'B, ed. The Whole Fungus. Ottawa: National bluseum of Natural Sciences. 1:283-310. Lee SB, Taylor<PV. 1990. Isolatiotl of DNA from fi~ngalmycelia and single spores. In: Innia MA, Gelfand DH, Sninsky JJ, MXite 'rJ, eds. PCR protocols: a guide to methods and applications. New York: Academic Press. p 282-287. Link HF. 1816. Observationes in ordines plantarum naturales. Dissertatio 11, sisteils nuperas de M u c e d i n ~ ~ et m Gastromycorurn ordinibus observationes. Mag Ges Naturf Freunde Berlin 7:25-45. Liu XJ, Guo 11. 1982. Studies on some species of the genus Pfzaeoramularia in China. Acta Phytopathol Sinica 12: 1-13, Masclaux F, Gukho E, Hoog GS de, Christen R. 1995. Phylogenetic relationships of human-pathogenic Cludosporium (Xylohypha) species inferred from partial LSL rRNA sequences. J Med Vet hfycol 33327-338. Maublanc A. 1913a. Sur une maladie de feuilles du papayer "C;c~rirapapaya". Lavoura 16:208-212. . 1913b. Sur une maladie de feuilles du papayer (Carica papaya). Bull Soc Mycol Fr 29353-338. McKemp JM, Morgan-Jones G. 1990. Studies in the genus Cladosporium sensu lato. 11. Collcerning fIetrrosporiurn gmcile, the causal agent organism of leaf spot disease of Iris species. Mycotaxon 39:425-440. Morgan-Jones G. 1980. Notes on hyphomycetes. XXXI'. Stenrllopsis gen. no\: Mycotaxon 10:403-408. . 1998. Notes on hyphoniycetes. LXXK. Concerning Parastenella magnoliae, its phylogenetic afffinic, and the relationship between it and its host, Magnolia p a n dz$!om. Mycotaxon 66:421-428. Muntaiiola M. 1960. rUgunos Hyphomycetes criticos. Notas y descripciones. 1,illoa 30: 183-21 3. Nag Raj TR. 1981. C;oelomycete systematics. In: Cole GT, Kendrick B, eds. The b i o l o ~of conidial fungi. New York: Academic Press. p 33-84. Pollack FG. 1987. An annotated cornpilation of Cmospora names. hlycol Mern 12:l-212. Pons N. 1988. Ocurrencia de Crrcospora aurantza sobre Citrus spp. en Venezuela. Fitopatol Venezol 1:8-13. Pons N, Sutton BC. 1988. Cercospora and similar fungi on yams (Dioscorea spp.). Mycol Pap 160:l-78. -. 1996. Cercospora and similar fungi on Heliotropium weeds. Mycol Res 100:813-820. --, Gay JL. 1985. Ultrastructure of conidiogenesis in Cwcospora beticola. Trans Br Mycol Soc 85:405416. Prasad SS, Verma RAB. 1970. A new genus of Moniliales from India. Indian Phytopathol 23: 111-1 13. Rangel E. 1915. Fungos parasitos do guando ( C a j a n u ~ijtdicus Spreng.). Bol Xgric S Paulo, Ser 16(2):145-136. . 1916. Fungos do Brasil, novo ou ma1 conhecidos por Ezigenia Rangel. Arch hIus Nac Rio de Janeiro 18: 1.55-164. . 1917. Algunos fungos rlobas d o Brasil. Xrch Jard Bot Rio de Ja~leiro259-74. Saccardo PA. 1880. Conspectus generum fungorum Italiae inferiorum, nempe ad Sphaeropsideas, Melanconicas et Hyphomycetas pertinentium, systemate sporologico dispositorum. Michelia 2:l-38. Samuels GJ, Seifert IL4. 1987. Taxonomic implications of variation arnong hypocrealeail anamorphs. In: Sugiya ma J , ed. Pleomorphic fungi: the diversity and its taxonon~icimplications. Tokyo, Japan: Kodansha Ltd. and Arnsterdan~:Elsevier. p 29-56. Sarvada K.1922. Descriptive catalogue of the Forrllosan fungi 11. Dep L4gricGoy Res Inst Taiwan Rep 2:139-164. Seifert KA, Okada G. 1990. Taxonomic implications of conidiomatal anatomy in synnernatous Hyphomycetes. Stud Mycol 32:29-40. Shaw DE, Alcorn JL. 1993. New names for Vwrztcispora and its species. Aust Syst Bot 6:273-276. Solheim LVG. 1929. Mol-phological studies of the genus Crr-cospom. I11 Biol Monogr 12:l-85. Spegazzini C. 1910. Mycetes Argentinenses, Ser. V. h r i hlus Nac Hist Nat B Ares 20329-467. Stewart EL, Liu Z, Crous PLY, Smb6 L. 1999. Phylogenetic relationships among some cercosporoid anamorphs of L~l~cosphaerelia based o n rDNA sequence analysis. Mycol Res 103:1491-1399. Subranlanian CV, Rarnakrishnan R. 1956. List of Indian fungi 1952-36. J Madras Univ Sect B 26:327-421. Sutton BC. 1980. The Coelomycetes. Fungi imperfecti with pycnidia, acervuli and stromata. England: CMI. 696 p. . 1994. Prathigadci twminaliae. IMI Descriptions of Fungi and Bacteria, No. 1181. Mycopathol 125:45-46. , Crous PM'. 1997. Lecano.rtictoj~sisg en. nov. and similar f~lngifrom Syzygiztm species. Mycol Res 101:21322.5. . Hennebert GL. 1994. Interconnections amongst anamorphs and their possible contribution to Ascomycete systematics. In: Hawksu-orth DL, ed. Ascornycete systematics. Problems and perspectives in the nineties. New York: Plenuni Press. p 77-98. , Pascoe I. 1987. Tandonella olenrine sp. nov. and generic separation of Tandonr~llaand Sclrro~graphiopsis. Aust J Bot 33:183-191. -. 1989. Reassessment of Pelto~oma,S t i p z l / n and Batcheloromyces and description of Hyphothyn'um gen. nov. Mycol Res 92:210-222. , Pons N. 1980. Notes on the original species of Cercosporina. Mycotaxon 12:201-218. , Shamoun SF, Crous PW. 1996. Two leaf pathogens of Ribes spp. in North America, Quasiphloeospora saximontanensis (Deighton) comb. nov. and Phloeosporella ribis (J.J. Davis) comb. nov. Mycol Res 100:979-983. , Waller JM. 1988. Taxonomy of Ophiocladium hordei, causing leaf lesions on Triticale and other Gramineae. Trans Br Mycol Soc 90:55-61. Swart HJ, Walker J. 1988. Australian leaf-inhabiting fungi XXVIII. Hendersonia on Eucalyptus. Trans Br Mycol SOC90:633-641. Swofford DL. 1999. PAUP* (Phylogenetic analysis using parsimony* and other methods), 4.0b2a. Sunderland, Massachusetts: Sinauer Associates. Sydow H. 1928. Novae fungorum species XIX. Ann Mycol 26:132-139. . 1930. Fungi venezuelani. Ann Mycol 28:29-224. Takeo K, Hoog GS de, Miyaji M, Nishimura K. 1995. Conidial surface ultrastructure of human-pathogenic and saprobic Cladosporium species. Antonie Leewenhoek 68: 51-55. Taylor JE, Crous PW. 1999. Phaeophleospora faureae comb. nov. associated with leaf spots on Faurea saligna (Proteaceae), with a key to known species of Phaeophleospora. Fungal Div 3:153-158. Thaung MM. 1976. New hyphomycetes from Burma. Trans Br Mycol Soc 66:211-215. Thirumalachar MJ, Mishra JN. 1953. Contribution to the study of fungi of Bihar, India-I. Sydowia 7:29-83. Thompson JD, Higgins DG, Gibson TJ. 1994. Clustal W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position spe- cific gap penalties and weight matrix choice. Nucl Acids Res 22:4673-4680. Unger F. 1933. Die Exantheme der Pflanzen. Wien. 423 p. Untereiner WA. 1997. Taxonomy of selected members of the ascomycete genus Capronia with notes on anamorph-teleomorph connections. Mycologia 89:120131. . 2000. Capronia and its anamorphs: exploring the value of morphological and molecular characters in the systematics of the Herpotrichiellaceae. Stud Mycol 45:141-149. Vries GA de. 1952. Contribution to the knowledge of the genus Cladosporium Link ex Fr. CBS, Baarn. 121 p. Wingfield MJ, Seifert KA, Webber JN, eds. 1993. Ceratoqstis and Ophiostoma: taxonomy, ecology and pathology. St. Paul, Minnesota: APS Press. 293 p. Yen JM. 1966. Etude sur les champignons parasites du SudEst asiatique. nr.Troisieme note sur quelques nouvelles especes de Cercospora de Singapour. Rev Mycol31: 109149. . 1981. ~ t u d esur les champignons parasites de SudEst asiatique, 42. Champignons parasites de Malaisie, 21. Bull Soc Mycol Fr 97:129-133. . 1983. Studies on parasitic fungi from South East Asia, 49. Parasitic fungi from malaysia, 25: Semipseudocercospora gen. nov. Mycotaxon 17:361-363. , Lim G. 1969. Etude sur les champignons parasites du Sud-Est asiatique. XIV. Huitieme note sur les Cercospora de Malaisie. Bull Soc Mycol Fr 85:459-474. -. 1973. Etude sur les champignons parasites du Sud-Est asiatique. XX. Dixikme note sur les Cercospora de Malaisie. Cah Pac 17:95-114. --. 1980. Cercospora and allied genera of Singapore and the Malay Peninsula. Gardens Bull Singapore 33:151-263.

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Circumscription of anamorph genera in Mycosphaerella based on ITS rDNA sequence and morphology

Circumscription of anamorph genera in Mycosphaerella based on ITS rDNA sequence and morphology

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