Verkley et al.S. rumicumPrimulaceaeSep<strong>to</strong>ria lysimachiaeRanunculaceaeSep<strong>to</strong>ria clematidisS. lycoc<strong>to</strong>niS. napelliRosaceaeSep<strong>to</strong>ria citri complexSphaerul<strong>in</strong>a geiSphaer. tirolensisSphaer. westendorpiiRubiaceaeSep<strong>to</strong>ria cruciataeS. coprosmaeRutaceaeSep<strong>to</strong>ria protearumSalicaceaeSphaerul<strong>in</strong>a frondicolaScrophulariaceaeSep<strong>to</strong>ria digitalisUrticaceaeSep<strong>to</strong>ria urticaeVerbenaceaeSep<strong>to</strong>ria verbenaeViolaceaeSep<strong>to</strong>ria paridisACKNOWLEDGEMENTSWe thank the technical staff of the <strong>CBS</strong> Collection for their support dur<strong>in</strong>g thisproject. Arien van Iperen (cultures) and Marjan Vermaas (pho<strong>to</strong>graphic plates)are acknowledged for their <strong>in</strong>valuable assistance. Huub van der Aa is thankedfor his help dur<strong>in</strong>g collect<strong>in</strong>g and identify<strong>in</strong>g material. Simeon Vanev is gratefullyacknowledged for his support with collect<strong>in</strong>g bibliographic data and translat<strong>in</strong>gRussian diagnoses. Part of the DNA barcode sequenc<strong>in</strong>g results used <strong>in</strong> this studywas f<strong>in</strong>ancially supported by the Fonds Economische Structuurversterk<strong>in</strong>g (FES,Dutch M<strong>in</strong>istery of Education, Culture and Science grant BEK/BPR-2009/137964-U,“Mak<strong>in</strong>g the Tree of Life Work”).REFERENCESAndrianova TV (1987). Problems of classification and phylogeny of Sep<strong>to</strong>ria<strong>species</strong>. Mikologiya i Fi<strong>to</strong>pa<strong>to</strong>logiya 21: 393–399.Aptroot A (2006). Mycosphaerella and its anamorphs 2. Conspectus ofMycosphaerella. <strong>CBS</strong> Biodiversity Series 5. <strong>CBS</strong>-<strong>KNAW</strong> Fungal BiodiversityCentre, Utrecht, The Netherlands.Arx JA von (1983). Mycosphaerella and its anamorphs. Proceed<strong>in</strong>gs Kon<strong>in</strong>klijkeNederlandse Akademie van Wetenschappen, Series C - Biological and MedicalSciences 86 (1): 15–54.Beach WS (1919). Biological spezialization <strong>in</strong> the genus Sep<strong>to</strong>ria. American Journalof Botany 6: 1–33.Braun U (1995). A monograph of Cercosporella, Ramularia and allied genera(Phy<strong>to</strong>pathogenic Hyphomycetes). Vol. 1. IHW-Verlag, Ech<strong>in</strong>g.Breeÿen A den, Groe<strong>new</strong>ald JZ, Verkley GJM, Crous PW (2006). Morphological andmolecular characterisation of Mycosphaerellaceae associated with the <strong>in</strong>vasiveweed, Chromolaena odorata. Fungal Diversity 23: 89–110.Cheewangkoon R, Crous PW, Hyde KD, Groe<strong>new</strong>ald JZ, To-anan C (2008). Speciesof Mycosphaerella and related anamorphs on Eucalyptus leaves from Thailand.Persoonia 21: 77–91.Cochran LC (1932). A study of two Sep<strong>to</strong>ria leaf spots of celery. Phy<strong>to</strong>pathology22: 791–812.Constant<strong>in</strong>escu O (1984). Taxonomic revision of Sep<strong>to</strong>ria-like fungi parasitic onBetulaceae. Transactions of the British Mycological Society 83: 383–398.Crous PW, Aptroot A, Kang J-C, Braun U, W<strong>in</strong>gfield MJ (2000). The genusMycosphaerella and its anamorphs. Studies <strong>in</strong> Mycology 45: 107–121.Crous PW, Groe<strong>new</strong>ald JZ (2005). Hosts, <strong>species</strong> and genotypes: op<strong>in</strong>ions versusdata. Australasian Plant Pathology 34: 463–470.Crous PW, Denman S, Taylor JE, Swart L, Palm, ME (2004a).Cultivation anddiseases of Proteaceae: Leucadendron, Leucospermum and Protea. <strong>CBS</strong>Biodiversity Series 2: 1–228.Crous PW, Groe<strong>new</strong>ald JZ, Pongpanich K, Himaman W, Arzanlou M, W<strong>in</strong>gfield MJ(2004b). Cryptic speciation and host specificity among Mycosphaerella spp.occurr<strong>in</strong>g on Australian Acacia <strong>species</strong> grown as exotics <strong>in</strong> the tropics. Studies<strong>in</strong> Mycology 50: 457–469.Crous PW, Kang J-C, Braun U (2001). A phylogenetic redef<strong>in</strong>ition of anamorphgenera <strong>in</strong> Mycosphaerella based on ITS rDNA sequences and morphology.Mycologia 93: 1081–1101.Crous PW, Slippers B, W<strong>in</strong>gfield MJ, Rheeder J, Marasas WFO, et al. (2006a).Phylogenetic l<strong>in</strong>eages <strong>in</strong> the Botryosphaeriaceae. Studies <strong>in</strong> Mycology 55:235–253.Crous PW, Verkley GJM, Groe<strong>new</strong>ald JZ, Samson RA (2009). <strong>CBS</strong> Labora<strong>to</strong>ryManual Series 1. <strong>CBS</strong>-<strong>KNAW</strong> Fungal Biodiversity Centre, Utrecht, TheNetherlands.Crous PW, W<strong>in</strong>gfield MJ, Mansilla JP, Alfenas AC, Groe<strong>new</strong>ald JZ (2006b).Phylogenetic reassessment of Mycosphaerella spp. and their anamorphsoccurr<strong>in</strong>g on Eucalyptus. II. Studies <strong>in</strong> Mycology 55: 99–131.Demaree JB, Wilcox MS (1943). The fungus caus<strong>in</strong>g the so-called “Sep<strong>to</strong>ria leafspotdisease” of raspberry. Phy<strong>to</strong>pathology 33: 986–1003.Diedicke H (1915). Pilze VII, Sphaeropsidaea, Melanconiae,. Kryp<strong>to</strong>gamenfl. MarkBrandenburg 9: 1–962.Farr DF (1991). Sep<strong>to</strong>ria <strong>species</strong> on Cornus. Mycologia 83: 611–623.Farr DF (1992). Species of Sep<strong>to</strong>ria on the Fabaceae, subfamily Faboidae, tribeGenistae. Sydowia 44: 13–31.Feau N, Hamel<strong>in</strong> RC, Bernier L (2006). Attributes and congruence of three moleculardata sets: <strong>in</strong>ferr<strong>in</strong>g phylogenies among Sep<strong>to</strong>ria-related <strong>species</strong> from woodyperennial plants. Molecular Phylogenetics and Evolution 40: 808–829.Gabrielson RL, Grogan RG (1964). The late blight organism Sep<strong>to</strong>ria apiicola.Phy<strong>to</strong>pathology 54: 1251–1257.Goodw<strong>in</strong> SB, Dunkle LD, Zismann VL (2001). Phylogenetic analysis of Cercosporaand Mycosphaerella based on the <strong>in</strong>ternal transcribed spacer region ofribosomal DNA. Phy<strong>to</strong>pathology 91: 648–658.Grove WB (1935). British stem and leaf fungi (Coelomycetes). Vol. I: Sphaeropsidales,Sphaerioideae with hyal<strong>in</strong>e conidia: 1–488.Hall TA (1999). BioEdit: A user-friendly biological sequence alignment edi<strong>to</strong>r andanalysis program for W<strong>in</strong>dows 95/98/NT. Nucleic Acids Symposium Series 41:95–98.Hebert PDN, Cyw<strong>in</strong>ska A, Ball SL, DeWaard JR (2003). Biological identificationsthrough DNA barcodes. Proceed<strong>in</strong>gs of the Royal Society of London SeriesB 270: 313–321.Huelsenbeck JP, Ronquist F (2001). MRBAYES: Bayesian <strong>in</strong>ference of phylogenetictrees. Bio<strong>in</strong>formatics 17: 754–755.Jørstad I (1965). Sep<strong>to</strong>ria and sep<strong>to</strong>roid fungi on dicotyledones <strong>in</strong> Norway. Skrifterutgitt av Det Norske Videnskaps-Akademi i Oslo, I: Mat-Naturv. Klasse 22:1-110.Ka<strong>to</strong>h K, Misawa K, Kuma K, Miyata T (2002). MAFFT: a novel method for rapidmultiple sequence alignment based on fast Fourier transform. Nucleic AcidsResearch 30: 3059–3066.Kuijpers AFA, Aptroot A (2004). A revision of Mycosphaerella section Longispora(Ascomycetes). Nova Hedwigia 75: 451–468.Lombard L, Crous PW, W<strong>in</strong>gfield BD, W<strong>in</strong>gfield MJ (2010). Species concepts <strong>in</strong>Calonectria (Cyl<strong>in</strong>drocladium). Studies <strong>in</strong> Mycology 66: 1–13.Markevičius V, Treigienė A (2003). Sphaeropsidales. Genus Sep<strong>to</strong>ria. MycotaLithuaniae 10, 3: 1–199.Mason-Gamer RJ, Kellogg EA (1996). Test<strong>in</strong>g for phylogenetic conflict amongmolecular data sets <strong>in</strong> the tribe Triticeae (Gram<strong>in</strong>eae). Systematic Biology 45:524–545.Muthumary J (1999). First contribution <strong>to</strong> a monograph of Sep<strong>to</strong>ria <strong>species</strong> <strong>in</strong> India.Madras: Centre for advanced studies <strong>in</strong> Botany.Nylander JAA (2004). MrModeltest v2. Program distributed by the author.Evolutionary Biology Centre Uppsala University.Petrak F (1925). Beiträge zur Pilzflora Südost-Galiziens und der Zentralkarpathen.Hedwigia 65: 179–330.Petrak F (1957). Die auf Aconiten vorkommenden Arten der Gattung Sep<strong>to</strong>ria.Sydowia 11: 375–379.Priest MJ (2006). Fungi of Australia. Sep<strong>to</strong>ria. ABRS, Canberra: CSIRO Publish<strong>in</strong>g,Melbourne.Punithal<strong>in</strong>gam E (1967a). Sep<strong>to</strong>ria chrysanthemella. CMI Descriptions of PathogenicFungi and Bacteria 137. Commonwealth Mycological Institute, Kew.Punithal<strong>in</strong>gam E (1967b). Sep<strong>to</strong>ria leucanthemi. CMI Descriptions of PathogenicFungi and Bacteria 138. Commonwealth Mycological Institute, KewPunithal<strong>in</strong>gam E (1967c). Sep<strong>to</strong>ria obesa. CMI Descriptions of Pathogenic Fungiand Bacteria 139. Commonwealth Mycological Institute, Kew.304
A <strong>new</strong> <strong>approach</strong> <strong>to</strong> <strong>species</strong> <strong>delimitation</strong> <strong>in</strong> Sep<strong>to</strong>riaPunithal<strong>in</strong>gam E (1967d). Sep<strong>to</strong>ria socia. CMI Descriptions of Pathogenic Fungi andBacteria 140. Commonwealth Mycological Institute, Kew.Punithal<strong>in</strong>gam E (1982). Sep<strong>to</strong>ria cucurbitacearum. CMI Descriptions of PathogenicFungi and Bacteria 740. Commonwealth Mycological Institute, Kew.Punithal<strong>in</strong>gam E, Holiday P (1972). Sep<strong>to</strong>ria lactucae. CMI Descriptions ofPathogenic Fungi and Bacteria 335. Commonwealth Mycological Institute,Kew.Punithal<strong>in</strong>gam E, Wheeler BEJ (1965). Sep<strong>to</strong>ria spp. occurr<strong>in</strong>g on <strong>species</strong> ofChrysanthemum. Transactions of the British Mycological Society 48: 423–439.Quaedvlieg W, Kema GHJ, Groe<strong>new</strong>ald JZ, Verkley GJM, Seifbarghi S, Razavi M,Gohari A, Mirzadi, Mehrabi R, Crous PW (2011). Zymosep<strong>to</strong>ria gen. nov.: a<strong>new</strong> genus <strong>to</strong> accommodate Sep<strong>to</strong>ria-like <strong>species</strong> occurr<strong>in</strong>g on gram<strong>in</strong>icoloushosts. Persoonia 26: 57–69.Quaedvlieg W, Groe<strong>new</strong>ald JZ, Jesús Yáñez-Morales M, Crous PW (2012). DNAbarcod<strong>in</strong>g of Mycosphaerella <strong>species</strong> of quarant<strong>in</strong>e importance <strong>to</strong> Europe.Persoonia 29: 101–115.Quaedvlieg W, Verkley GJM, Sh<strong>in</strong> H-D, Barre<strong>to</strong> RW, Algenas AC, Swart WJ,Groe<strong>new</strong>ald JZ, Crous PW (2013). Siz<strong>in</strong>g up Sep<strong>to</strong>ria. Studies <strong>in</strong> Mycology 75:307–390. (this volume).Radulescu E, Negru A, Docea E (1973). Sep<strong>to</strong>riozele d<strong>in</strong> România. Bucureşti:Editura Academiei Republicii Socialiste România.Rayner RW (1970). A mycological colour chart. CMI and British Mycological Society.Kew, UK.Saccardo PA (1884). Sylloge Fungorum: Sylloge Sphaeropsidearum etMelanconiearum. 3: 542. Padova, Italy.Saccardo PA (1895). Sylloge fungorum: Supplemental Universale, Pars III 11:1–753. Padova, Italy.Saccardo PA (1906). Sylloge fungorum: Supplemental Universale, Pars VII 18:1–838. Padova, Italy.Saccardo PA, Sydow P (1899). Sylloge Fungorum: Supplemental Universale, ParsIV 14: 1–1316. Padova, Italy.Sheridan JE (1968). Conditions for <strong>in</strong>fection of celery by Sep<strong>to</strong>ria apiicola. PlantDisease Report 52: 142–145.Sh<strong>in</strong> HD, Sameva EF (2004). Sep<strong>to</strong>ria <strong>in</strong> Korea (Plant Pathogens of Korea 11).National Institute of Agricultural Science and Technology, Republic of Korea.Simon UK, Groe<strong>new</strong>ald JZ, Stierhof Y-D, Crous PW, Bauer R (2009). A necrotrophicphy<strong>to</strong>pathogen form<strong>in</strong>g a special cellular <strong>in</strong>teraction with its host Aegopodiumpodagraria. Mycological Progress 9: 49–56.Smissen RD, Clement JC, Garnock-Jones PJ, Chambers JK (2002). Subfamilialrelationships with<strong>in</strong> Caryophyllaceae as <strong>in</strong>ferred from 5’ ndhF sequences.American Journal of Botany 89: 1336–1341.Stewart EL, Liu Z, Crous PW, Szabo LJ (1999). Phylogenetic relationships amongsome cercosporoid anamorphs of Mycosphaerella based on rDNA sequenceanalysis. Mycological Research 103: 1491–1499.Sut<strong>to</strong>n BC (1980). The Coelomycetes. Fungi imperfecti with Pycnidia, Acervuli andStromata. Commonwealth Mycological Institute, Kew, Surrey, England.Sut<strong>to</strong>n BC, Pascoe IG (1987). Sep<strong>to</strong>ria <strong>species</strong> on Acacia. Transactions of theBritish Mycological Society 89: 521–532.Sut<strong>to</strong>n BC, Pascoe IG (1989). Some Sep<strong>to</strong>ria <strong>species</strong> on native Australian plants.Studies <strong>in</strong> Mycology 31: 177–186.Sut<strong>to</strong>n BC, Hennebert GL (1994). Interconnections amongst anamorphs and theirpossible contribution <strong>to</strong> ascomycete systematics. In Ascomycete systematics:problems and perspectives <strong>in</strong> the n<strong>in</strong>eties (Hawksworth DL, ed.), pp. 77–98.Plenum Press, New York, USA.Sut<strong>to</strong>n BC, Waters<strong>to</strong>n JM (1966) Sep<strong>to</strong>ria apiicola. CMI Descriptions of PathogenicFungi and Bacteria 88. Commonwealth Mycological Institute, Kew.Sydow H von (1924). Beiträge zur Kenntnis der Pilzflora Neu-Seelands. I. AnnalesMycologici 22: 299–317.Tamura K, Dudley J, Nei M, Kumar S (2007). MEGA4: Molecular evolutionarygenetics analysis (MEGA) software version 4.0. Molecular Biology andEvolution 24: 1596–1599.Teterevnikova-Babayan DN (1987). Fungi of the genus Sep<strong>to</strong>ria <strong>in</strong> the U.S.S.R.Yerevan: Akademia Nauk Armyanskoi SSR.Vanev SG, Sameva EF, Bakalova GG (1997). Fungi Bulgaricae 3. OrdoSphaeropsidales. Sofia: Editio Academica Prof. Mar<strong>in</strong> Dr<strong>in</strong>ov.Verkley GJM (1998a). Ultrastructural evidence for two types of proliferation <strong>in</strong> as<strong>in</strong>gle conidiogenous cell of Sep<strong>to</strong>ria chrysanthemella. Mycological Research102: 368–372.Verkley GJM (1998b). Ultrastructure of conidiogenesis and conidia <strong>in</strong> two <strong>species</strong> ofSep<strong>to</strong>ria sensu la<strong>to</strong>. Mycologia 90: 189–198.Verkley GJM, Priest MJ (2000). Sep<strong>to</strong>ria and similar coelomyce<strong>to</strong>us anamorphs ofMycosphaerella. Studies <strong>in</strong> Mycology 45: 123–128.Verkley GJM, Star<strong>in</strong>k-Willemse M (2004a). A phylogenetic study of some Sep<strong>to</strong>ria<strong>species</strong> pathogenic <strong>to</strong> Asteraceae based on ITS ribosomal DNA sequences.Mycological Progress 3: 315–322.Verkley GJM, Star<strong>in</strong>k-Willemse M, Iperen A van, Abeln ECA (2004b). Phylogeneticanalyses of Sep<strong>to</strong>ria <strong>species</strong> based on the ITS and LSU-D2 regions of nuclearribosomal DNA. Mycologia 96: 558–571.Verkley GJM, Crous PW, Groe<strong>new</strong>ald JZ, Braun U, Aptroot A (2004c). Mycosphaerellapunctiformis revisited: morphology, phylogeny, and epitypification of the type<strong>species</strong> of the genus Mycosphaerella (Dothideales, Ascomycota). MycologicalResearch 108: 1271–1282.Waddell HT, Weber GF (1963). Physiology and pathology of Sep<strong>to</strong>ria <strong>species</strong> onChrysanthemum. Mycologia 55: 442–452.White TJ, Bruns T, Lee S, Taylor JW (1990). Amplification and direct sequenc<strong>in</strong>g offungal ribosomal RNA genes for phylogenetics. In: PCR pro<strong>to</strong>cols: a guide <strong>to</strong>methods and applications (Innis MA, Gelfland DH, S<strong>in</strong><strong>in</strong>sky JJ, White TJ, eds).Academic Press, San Diego, USA: 315–322.Zalar P, Hoog GS de, Schroers H-J (2007). Phylogeny and ecology of the ubiqui<strong>to</strong>ussaprobe Cladosporium sphaerospermum, with descriptions of seven <strong>new</strong><strong>species</strong> from hypersal<strong>in</strong>e environments. Studies <strong>in</strong> Mycology 58: 157–183.www.studies<strong>in</strong>mycology.org305