Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 A modern account of the genus Phyllosticta Wulandari NF1, 2*, Bhat DJ3 and To-anun C1* 1 Department of Entomology and Plant Pathology, Faculty of Agriculture, Chiang Mai University, Chiang Mai, Thailand. Microbiology Division, Research Centre for Biology, Indonesian Institute of Sciences (LIPI), Cibinong Science Centre, Cibinong, Indonesia. 3 Formerly, Department of Botany, Goa University, Goa-403 206, India 2 Wulandari NF, Bhat DJ, To-anun C 2013 – A modern account of the genus Phyllosticta. Plant Pathology & Quarantine 3(2), 145–159, doi 10.5943/ppq/3/2/4 Conidial states of Guignardia are in the genus Phyllosticta. In accordance to nomenclatural decisions of IBC Melbourne 2011, this paper validates species that were in Guignardia but are now accepted in Phyllosticta. The conclusions are arrived based on molecular analyses and morphological examination of holotypes of those species previously described in the genus Guignardia. Thirty-four species of Phyllosticta, viz. P. ampelicida (Engelm.) Aa, P. aristolochiicola R.G. Shivas, Y.P. Tan & Grice, P. bifrenariae O.L. Pereira, Glienke & Crous, P. braziliniae O.L. Pereira, Glienke & Crous, P. candeloflamma (J. Fröhlich & K.D. Hyde) Wulandari, comb. nov., P. capitalensis Henn., P. cavendishii M.H. Wong & Crous, P. citriasiana Wulandari, Gruyter & Crous, P. citribraziliensis C. Glienke & Crous, P. citricarpa (McAlpine) Aa, P. citrichinaensis X.H. Wang, K.D. Hyde & H.Y. Li, P. clematidis (Hsieh, Chen & Sivan.) Wulandari, comb. nov., P. cruenta (Fr.) J. Kickx f., P. cussoniae Cejp, P. ericarum Crous, P. garciniae (Hino & Katumoto) Motohashi, Tak. Kobay. & Yas. Ono., P. gaultheriae Aa, P. hostae Y.Y. Su & L. Cai, P. hubeiensis K. Zhang, Y.Y. Su & L. Cai, P. hymenocallidicola Crous, Summerell & Romberg, P. hypoglossi (Mont.) Allesch., Rabenh., P. ilicis-aquifolii Y.Y. Su & L. Cai, P. korthalsellae A. Sultan, P.R. Johnst, D.C. Park & A.W. Robertson, P. maculata Wong & Crous, P. morindae (Petr. & Syd.) Aa, P. musarum (Cooke) Aa, P. muscadinii (Luttr.) Wulandari, comb. nov., P. owaniana G. Winter, P. partenocissi K. Zhang, N. Zhang & L. Cai, P. philoprina (Berk. & M.A. Curtis) Aa, P. schimae Y.Y. Su & L. Cai, P. spinarum (Died.) Nag Raj & M. Morelet, P. styracicola Zhang, Y.Y. Su & L. Cai, and P. vaccinii Earle are accepted in this study based on examination of type material of “Guignardia” species, including three new combinations. It is anticipated that other species of Phyllosticta will be accepted, following future molecular studies. Key words – accepted species – Dothideomycetes – molecular phylogeny – monograph – plant diseases – saprobes – taxonomy Article Information Received 11 July 2013 Accepted 20 September 2013 Published online 13 October 2013 *Corresponding authors: Chaiwat To-anun – e-mail – chaiwat.toanun@gmail.com, Nilam Wulandari – e-mail – nilamwulandari@gmail.com 145 Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 Introduction Species of Guignardia Viala & Ravaz (Botryosphaeriales) produce simple brown, thin-walled ascomata, containing bitunicate and fissitunicate asci and variously shaped, hyaline, unicellular ascospores, with or without mucilaginous sheaths or appendages (Sivanesan 1984, Wulandari et al. 2010a,b, 2011, Glienke et al. 2011). The ascomata contain sparse, cellular pseudoparaphyses. Several species of Guignardia have been linked to Phyllosticta asexual states via cultural studies or observation of co-occurence of both states in close proximity on the host (van der Aa 1973, van der Aa & Vanev 2002, Motohashi et al. 2008a,b, 2010, 2011). Another asexual state, Leptodothiorella, is known in some species (Wulandari et al. 2010, Su & Cai 2012, Zhang et al. 2013b). There are 340 epithets listed under Guignardia in Index Fungorum (2013). Unfortunately, there is no monographic work on the genus Guignardia so far and one generally relies on publications containing described species on the hosts (Hyde 1995) or those introduced as new species (van der Aa 1973, Punithalingham 1974, Sivanesan 1984, Hyde 1995, Okane et al. 2001, 2003, van der Aa & Vanev 2002, Motohashi et al. 2008a,b, Wulandari et al. 2009, Motohashi et al. 2010, Wikee et al. 2011, Glienke et al. 2011, Wong et al. 2012, Su & Cai 2012, Zhang et al. 2013b). Phyllosticta is the name given to the asexual states of Guignardia (van der Aa 1973, Sivanesan 1984, Hyde 1995, Wulandari et al. 2010a,b), which are phytopathogens producing simple, brown, thin-walled conidiomata, containing round to oval, hyaline, unicellular conidia. The distinctive character of Phyllosticta is the thin mucilaginous sheath surrounding the spore and an apical appendage (van der Aa 1973, van der Aa & Vanev 2002). Species of Guignardia and Phyllosticta are found associated with leaf spots of numerous plants and even isolated as endophytes (Okane et al. 2008, Glienke et al. 2011). A few species are saprobic (Wang et al. 2008). Because of their phytopathogenic nature, Phyllosticta species have been studied more than the Guignardia states. van der Aa (1973) has provided a detailed account of the genus Phyllosticta and accepted 30 species in the 145 genus. van der Aa & Vanev (2002) updated the taxonomy of the genus and accepted 154 species. The species are differentiated largely on host specificity and conidial characters. The International Botanical Congress held in Melbourne, Australia, in 2011, made efforts to bring an end to dual nomenclature of fungi (McNeill et al 2012, Hawksworth et al. 2011). It was resolved that the connected sexual and asexual states will have only one name to represent the genus or species which previously had two (or more) names. Thus, it is required to make a choice of name between Guignardia or Phyllosticta as these genera have been unequivocally linked by cultural and molecular data (Glienke et al. 2011, Wikee et al. 2013, Zhang et al. 2013b). In this paper, we use Phyllosticta over Guignardia, as suggested in recent publications (Glienke et al. 2011, Su & Cai 2012, Wong et al. 2012, Wikee et al. 2013, Zhang et al. 2013a,b) and the latter is treated as a synonym of Phyllosticta. The genus Phyllosticta is placed in the family Phyllostictaceae in Botryosphaeriales (Liu et al. 2012). The reasons for using Phyllosticta over “Guignardia” are: 1. Phyllosticta was introduced by Persoon (1818) and is an earlier name than Guignardia introduced by Viala & Ravaz (1892). The earlier name takes priority (see International Code for Nomenclature of Algae, Fungi and Plants; http://www.iapttaxon.org/nomen/main. php?page=title). 2. There are more Phyllosticta names in various indices, i.e. Index Fungorum, Species Fungorum, MycoBank and GenBank, than Guignardia. 3. The records of Phyllosticta in search engines, such as Google Scholar are higher. 4. Phyllosticta is an important genus of plant pathogens and therefore important in quarantine. For example, P. citricarpa causes citrus black spot (CBS) and is an important quarantinable species in Europe. 5. Phyllosticta is often used in agricultural reference books (van der Aa & Vanev 2002). Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 In this paper, “Guignardia” species are described based on holotypes. In all, 127 specimens of “Guignardia” from 31 herbaria have been examined, including the type material of many taxa. Descriptions and illustrations of eight accepted species of Phyllosticta are documented based on holotypes and annotations of 26 accepted species of Phyllosticta are provided based on published literature. A key to all species of Phyllosticta is not provided at present because the identification is based partly on morphology and, for some species, on the phylogeny. Materials & Methods The holotypes of 127 species named “Guignardia” were obtained on loan from herbaria (B, BPI, BRIP, CBS, DAR, F, FH, G, K, KRA, LE, MA, NTU, NY, PAV, PDD, PH, PREM, SIENA, TNS, TUR, W, YAM, and ZT) (Holmgren & Holmgren 1998). Ascomata were rehydrated, carefully dissected under a microscope and mounted in lactic acid. Twenty ascospores and conidia were measured. Hand sections were made to determine and illustrate the wall of ascomata. An Olympus microscope with an attached camera lucida apparatus was used for drawing. Photomicrographs were made on a Nikon eclipse 80i with an EOS 450D Nikon camera. All line drawings were prepared by the first author. Morphological characters are recorded as in Wulandari et al. (2009). Results Thirty-four presently accepted species of Phyllosticta are annotated, based on recent publications and examination of type material of “Guignardia” species. It is anticipated that many other species of Phyllosticta will be accepted in the future following molecular studies. Taxonomy Phyllosticta Pers., Traité sur les Champignons Comestibles (Paris) 55: 147 (1818). Possible synonymys Caudophoma B.V. Patil & Thirum., Sydowia 20: 36 (1968) [1966] 146 Guignardia Viala & Ravaz, Bull. Soc. Mycol. Fr. 8: 63 (1892) Laestadiella Höhn., Ann. Mycol. 16: 50 (1918) Leptasteromella Petr., Sydowia 20: 235 (1968) [1966] Leptodothiorella Höhn., Hedwigia 60: 173, 175 (1918) Leptodothiorella Aa, Stud. Mycol. 5: 13 (1973) Leptophacidium Höhn., Sber. Akad. Wiss. Wien, Math. naturw. Kl., Abt. 1 127: 331 [3 repr.] (1918) Macrophyllosticta Sousa da Câmara, Anais Inst. Sup. Agron. Univ. Téc. Lisboa 3: 36 (1929) Montagnellina Höhn., Sber. Akad. Wiss. Wien, Math. naturw. Kl., Abt. 1 121: 387 [49 repr.] (1912) Myriocarpa Fuckel, Jb. Nassau. Ver. Naturk. 23–24: 116 (1870) [1869–70] Pampolysporium Magnus, Verh. Zool.Bot. Ges. Wien 50: 444 (1900) Phyllosphaera Dumort., Comment. Bot.: 86 (1822) Phyllostictina Syd. & P. Syd., Ann. Mycol. 14: 185 (1916) Polysporidium Syd. & P. Syd., Ann. Mycol. 6: 528 (1908) Phytopathogenic, causing leaf spots, endophytes or rarely saprobic on plants. Sexual states: Ascomata small, pseudothecia, solitary to aggregated, scattered, uniloculate, immersed in host tissue or becoming erumpent, globose to subglobose, dark brown, ostiolate, with or without an ostiolar papilla. Peridium thin, composed of thick-walled pseudoparenchymatous cells towards the exterior, with inner wall cells thin and hyaline. Pseudoparaphyses absent in mature pseudothecia, sometimes present as tubular cells in immature ascomata. Asci 8-spored, bitunicate, fissitunicate, clavate to cylindrical, with a short pedicel, with or without an ocular chamber. Ascospores seriate, hyaline, 1-celled, ovoid, ellipsoidal to rhomboidal, straight or curved, wide in the middle, guttulate, with distinct mucilaginous appendages at one or both ends which in some disappear or remain as remnants at spore ends. Asexual state: Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 Conidiomata epiphyllous, small, pycnidial, gregarious, immersed, reddish brown to dark brown, uniloculate, subglobose, papillate, ostiolate. Wall of conidioma thick in culture, continuous with epidermal or subepidermal stroma, composed of several layers, with cells of the outer layers thick-walled, dark brown, isodiametric or slightly flattened; inner layers with thin-walled cells. Conidiogenous cells holoblastic, hyaline, short, cylindrical. Conidia 1-celled, very rarely 2-celled, broadly ellipsoidal to subglobose or obovate, rounded at both ends, occasionally truncate at the base, slightly indented at the apex, hyaline, smoothwalled, coarsely granular, surrounded by a slime layer, usually with a hyaline, flexuous, narrowly cylindrical, mucilaginous apical appendage. Appendage often disappearing when mounted in water or other reagents. Spermogonia pycnidial, numerous, crowded on faded leaf spots with unbordered necrotic spots on the leaf, epi- or amphiphylous, uniloculate, subglobose, sometimes embedded in a multiloculate stroma, black, papillate, ostiolate. Wall of spermogonium, continuous with subepidermal stroma, consisting of several layers of cells; the cells of the outer layers thick-walled, dark brown, slightly flattened; inner layers thin-walled, hyaline, flattened. Spermatiogenous cells holoblastic, hyaline, cylindrical to narrowly flattened. Spermatia 1celled, hyaline, dumbbell-shaped to cylindrical, bluntly rounded at both ends. Accepted species in Phyllosticta Phyllosticta ampelicida (Engelm.) Aa, Stud. Mycol. 5: 28 (1973) ≡ Naemospora ampelicida Engelm., Trans. Acad. Sci., St Louis 2: 165 (1863) = Phyllosticta ampelopsidis Ellis & G. Martin, in G. Martin, J. Mycol 2: 14 (1886) = Phyllosticta labrusca Thüm., Die Pilze des Weinstockes: 189 (1878) = Phyllosticta vitea Sacc., in P. Sydow, Syll. Fung. 12: 564 (1896) = Phyllosticta viticola Thüm., Die Pilze des Weinstockes: 188 (1878) = Phyllosticta vulpinae Allesch., Rabenh. Kryptog.-Fl. 1, 6: 98 (1898); illeg. name (Art. 52) 147 Host – Vitis riparia (Vitaceae) Phyllosticta ampelicida is the causal agent of black rot in grapes and one of the destructive fungal pathogens in North American viticulture. Engelman (1861) described this species as Naemospora ampelicida without designating a type. van der Aa (1973) re-described and transferred the fungus to Phyllosticta, recording it as asexual state of Guignardia bidwellii (Ellis) Viala & Ravaz (van der Aa & Vanev 2002). Neotypifying this species using a strain from ATCC, Zhang et al. (2013b) concluded that P. ampelicida is not only the asexual state of G. bidwellii but also host specific on Vitis sp. Zhang et al. (2013b) observed that P. ampelicida (ATCC 200578) is morphologically similar to the protologue of P. ampelicida as described by Engelman (1863) but distinct from P. bidwellii (CBS 111645) in host, morphology and distribution. Phyllosticta parthenocissi, which occurs on Parthenocissus sp. (Vitaceae) differs from P. ampelicida (Zhang et al. 2013b) in size of conidia. Cultural characteristics, aerial mycelium and multi-gene phylogenies (ITS, EF, actin and GPDH) also show that P. ampelicida is distinct from P. parthenocisii (Zhang et al. 2013b). Phyllosticta aristolochiicola R.G. Shivas, Y.P. Tan & Grice, Persoonia 29: 154 (2012) Host – on leaves of Aristolochia acuminata (Aristolochiaceae) Phyllosticta aristolochiicola is associated with leaf spots and shot-hole disease of Aristolochia clematitis and A. sempervirens in Queensland, Australia (Shivas & Alcorn 1996). Phyllosticta ariestolochia Tassi is a replacement name for P. tassiana Saccardo & Traverso occurring on Aristolochia (Aristolochiaceae), a homonym and illegitimate name (van der Aa & Vanev 2002). A GenBank nucleotide blast result for ITS, showed that P. aristolochiicola is most similar to P. cordilinophylla (97% similarity), while LSU sequence data showing this species to be close to P. abietis and P. minima (99% similarity). This species forms a distinct clade from P. abietis and P. minima (Crous et al. 2012a). Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 Phyllosticta bifrenariae O.L. Pereira, Glienke & Crous, Persoonia 26: 52 (2011) Host – Bifrenaria harrisoniae (Orchidaceae) Previously reported as a pathogen on Bifrenaria harrisoniae (Silva et al. 2008), this species is also recorded from healthy leaves of orchids in Brazil (Glienke et al. 2011). Phyllosticta bifrenariae is distinct from P. capitalensis and P. aplectri (other species of Phyllosticta recorded on orchids) in that the conidia of P. bifrenariae are larger (10–16 × 7–9 μm) than those of P. capitalensis [(10– )11–12(–14) × (5–)6–7 μm] and P. aplectri (5– 8 × 4–6 μm). Glienke et al. (2011) reported that P. bifrenariae is phylogenetically distinct from other Phyllosticta species, forming a separate clade with 97% bootstrap support. Phyllosticta braziliniae O.L. Pereira, Glienke & Crous, Persoonia 26: 53 (2011) Host – Mangiferae indica (Anacardiaceae) Several species of Phyllosticta have been recorded from Mangifera indica (Glienke et al. 2011, van der Aa 1973, van der Aa & Vanev 2002). Phyllosticta anacardiacearum is a replacement name of Phyllostictina mangiferae (van der Aa 1973). van der Aa & Vanev (2002) reported P. mortonii and P. mangiferae as excluded species of Phyllosticta. The conidia of P. braziliniae (8–10–11(12.5) x (5–)6(–7) μm) differ in size from P. capitalensis [(10–)11–12(–14) × (5–)6–7 μm)]. Without causing any leaf spot disease on mango, P. braziliniae is ecologically distinct from P. anacardiacearum. Phylogenetically P. braziliniae is distinct from other Phyllosticta species with 97% bootstrap value (Glienke et al. 2011). Phyllosticta candeloflamma (J. Fröhlich & K.D. Hyde) Wulandari, comb. nov. Fig. 1–2 MycoBank: MB 805537 = Guignardia candeloflamma J. Fröhlich & K.D. Hyde, Mycol. Res. 99: 110 (1995) MycoBank: MB 363083 Leaf spots 1–2.7 × 0.6–3 mm, ellipsoidal, pale brown, with thin, darker brown border, zonate in the centre, round to ellipsoidal, medium brown, containing numerous ascomata. Ascomata 50–130 μm 148 diam., 50–80 μm high, amphigenous, sometimes hyphogenous, black, globose to subglobose, immersed in leaf tissues, coriaceous, solitary to clustered, ostiolate, with ostiole as black dots at the centre. Peridium 17.5–31.25 μm wide, composed of 2–3 layers of cells textura angularis and pigmented outwardly and around the ostiole, paler in the innermost layer. Pseudoparaphyses not observed. Asci 50–90 × 19–25 μm ( x = 68 × 22 μm, n = 20), 8-spored, bitunicate, with an ocular chamber 3–7 μm high, clavate to broadly clavate, rounded at the apex, where the diam. is 14–25 μm, tapering gradually to a 6–7 × 3–10 μm long pedicel attached to the basal peridium. Ascospores 17–22 × 8–11 μm ( x = 20 × 9 μm, n = 20), biseriate to triseriate, ellipsoidal, hyaline to greenish, 1-celled, coarse-guttulate, smooth-walled, with polar mucilagenous sheath at both side ends, with sheath 3.3–13.2 μm wide extended at the base up to 15 μm long to candle-flame-shape. Fig. 1 – Phyllosticta candeloflamma (BRIP 20472, holotype) a. Leaf spots. b. Ascomata on the host surface. c. Section of ascoma. d. Peridium comprising one strata of textura angularis cells; note cells with thickened brown cell walls at the ostiolar region. e–f. Asci. g–i. Ascospores. – Scale bars a = 5 mm, b = 100 μm, c–d = 15 μm, e–i = 10 μm. Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 Host – On living leaves of Pinanga sp. (Arecaceae) Known distribution – Australia, Indonesia. Material examined – Australia, Queensland, Smithfield, Australia, on leaves of Pinanga sp., 12 February 1992, K.D. Hyde, (BRIP 20472, holotype); Indonesia, Irian Jaya, on leaves of Pinanga sp., March 1992, K.D. Hyde (BRIP 20398, isotype). Phyllosticta candeloflamma can form zonate spots on Pinanga leaves. This species is distinct from other species of Phyllosticta in having a polar mucilaginous sheath at each end of the ascospores with the basal appendage extending to a candle-flame-shaped appendage. Fig. 2 – Phyllosticta candeloflamma (BRIP 20472, holotype) line drawing. j. Section of ascoma (darkened area are fungal cells, in the arrowed region). k. Mature ascus. l. Immature asci. m. Ascospores. – Scale bars j = 50 μm, k– l = 25 μm, n = 20 μm. Phyllosticta capitalensis Henn., Hedwigia 48: 13 (1908) Fig. 3–4 = Guignardia aleuritis (Vassiljevsky) Aa (1973), Stud. Mycol. 5: 88 (1973) 149 ≡ Melanops aleuritis Vassiljevsky, Notulae Systematicae Section Cryptogamica Instituti Botanici Academiae Scientiarum U.S.S.R. 5: 9 (1940) = Guignardia allamandae A.K. Pande & V.G. Rao, J. Econ. Taxon. Bot. 14: 158 (1990) = Phyllosticta concinna (Syd.) Aa, Stud. Mycol. 5: 49 (1973) = Guignardia concinna (Syd.) Aa, Stud. Mycol. 5: 49 (1973) = Melanops concinna Syd., Annls Mycol. 24: 372–373 (1926) = Guignardia endophyllicola Okane, Nakagiri & Tad. Ito, Can. J. Bot. 79: 103 (2001) = Guignardia eucalyptorum Crous, South Africa J. Bot. 59: 605 (1993) = Phyllosticta eucalyptorum Crous, South Africa J. Bot. 59: 605 (1993) = Guignardia smilacis A.J. Roy, Indian Phytopath. 20: 348 (1968) = Guignardia graminea Lobik, Material on flora and fauna observations of Terek region 25 = Guignardia pipericola F. Stevens, Trans. Illinois State Acad. Sci. 10: 183 (1917) = Guignardia smilacinae (Dearn. & House) Dearn. & House, New York State Museum Bull..266: 73 (1925). = Guignardia bumeliae J.H. Mill. & G.E. Thomps., Mycologia 32: 4 (1940) = Guignardia helicteres F. Stevens, Trans. Illinois State Acad. Sci. 10: 183 (1917) = Guignardia festiva (Syd.) Sacc., Sylloge Fung. 24(2): 783 (1928) = Guignardia foeniculata (Mont.) Arx & E. Mull, Beitr. Kryptfl. Schweiz 11(no. 1): 48 (1954) = Guignardia cocogena (Cook) Punith., Mycol. Pap. 136: 21 (1974) = Guignardia capsici Punith., Mycol. Pap. 136: 13 (1974) = Guignardia castanopsidis W.H. Hsieh, Chi Y. Chen & Sivan., Mycol. Res. 101: 901 (1997) = Guignardia cocoicola Punith., Mycol. Pap. 136: 15 (1974) = Guignardia codiae Thaung, Trans. British Mycol. Soc. 66: 107 (1976) = Guignardia coffeae Punith. & B.S. Lee, in Punithalingam, Mycol. Pap. 149: 19 (1981) = Guignardia dyerae Punith. & P.H. Wong, in Punithalingam, Mycol. Pap. 149: 28 (1981) = Guignardia perseae Punith., Mycol. Pap. 136: 43 (1974) = Guignardia pini Sivan., Trans. British Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 Mycol. Soc. 73: 169 (1979) = Guignardia heveae Syd. & P. Syd., Ann. Mycol. 14: 360 (1916) = Guignardia miconia Seixas & R.W. Barreto, in Seixas, Barreto & Killgore, Mycologia 99: 103 (2007) = Guignardia musicola Wulandari, L. Cai & K.D. Hyde, Crypto. Mycol. 31: 412 (2010) = Guignardia musae F. Stevens, Bull. Bernice P. Bishop Museum, Honolulu, Hawaii 19: 101 (1925) = Guignardia stephensii Wulandari & K.D. Hyde, in Wulandari, To-anun, Cai, AbdElsalam & Hyde, Crypto. Mycol. 31: 406 (2010) = Guignardia musae F. Stevens, Bull. Bernice P. Bishop Museum, Honolulu, Hawaii 19: 101 (1925) Ascomata 93–125 μm diam., 100–125 μm high, present on the upper surface of the leaves, black, globose to subglobose, immersed to semi-immersed in leaf tissues or growth media. Peridium 13–25 μm wide, with one strata of 2–3 layers of cells textura angularis with thickened brown walls around the ostiole. Pseudoparaphyses not observed. Asci 31–50 × 8–13 μm ( x = 41 × 10 μm, n = 20), 8-spored, bitunicate, cylindrical to cylindro-clavate, rounded at the apex, where the diameter is 5– 11 μm, tapering gradually to a 4–6 × 3–5 μm long pedicel attached to the basal peridium. Ascospores 11–13 × 3–4 μm ( x = 13 × 3 μm, n = 20), uniseriate to biseriate, ellipsoidal wide in the middle, obtrullate, hyaline to greenish, 1celled, coarse-guttulate, smooth-walled, with or without mucilaginous pads at the ends. Hosts – As saprobe on dead leaves of Aleurites fordii, A. moluccana (= A. triloba), A. cordata, A. montana (Euphorbiaceae), on fallen leaf of Smilax aspera (Smilacaceae), on dead stems of Atropis distans (Poaceae), on fallen leaves of Bumelia lycioides (Sapotaceae), on dead leaves of Vagnera stellate (Convallariceae, formerly Alliaceae); as endophyte of Rhododendron pulchrum (Ericaceae); leaf spots on living leaves of Allamanda cathartica (Apocynaceae), Cordia ferruginae (Boraginaceae), Eucalyptus dives (Myrtaceae), Helicteres (Sterculiaceae), Heptapleurum venulosum (Schefflera bengalensis) (Araliaceae), Piper sp. (Piperaceae), and Sumbaviopsis albicans (Euphorbiaceae) (Table 1). 150 Known distribution – Brazil, Brunei Darussalam, Carribean, Costa Rica, Dominican Republic, Eastern Cape, Georgia, Germany, Hong Kong, India, Indonesia, Italy, Jamaica, Japan, Malaysia, Myanmar, Netherlands, New Zealand, Panama, Pakistan, Philippines, Puerto Rico, South Africa, Sri Lanka, Taiwan, Thailand, Tasmania, China, UK, USA, Venezuela, Virgin Islands. Material examined – Georgia, Batumi, on leaves of Aleurites fordii, 24 September 1935, V.L. Vassiljevsky, (LE 34833 lectotype of G. aleuritis, designated here), ibid. (LE 34834, LE 34835, paratypes of G. aleuritis). ibid., Island Sheldobai on stems of Atropis distans, 15 July 1925, D.K. Volgunov, V.L. Komarov Botanical Institute. Prof. Popov Street 2. Saint Petersburg 197376, Russia (LE 122701, from the holotype of G. graminea). India, Dhobighat, Raniket, Almora UP, on leaves of Smilax aspera, 3 July 1963, A.J. Roy, Eidgenossische Technische Hochschule, Zurich (ETH, holotype of G. smilacis); ibid., Pune, Maharashtra, on leaves of Allamanda cathartica, February 1978, B.R.D. Yadav, Herbarium Cryptogamie Ajrekar Mycological Herbarium (AMH 4049, holotype of G. allamandae). Costa Rica (La Caja bei), 6 January 1925, H. Sydow (BPI 1110937, holotype of G. concinna). Japan, on living leaves of Rhododendron pulchrum cv. “Ohmurasaki” as endophytic cluture (IFO–H 12230, holotype of G. endophyllicola). South Africa, Tzaneen, Northern Transvaal, on leaves of Eucalyptus dives, 27 March 1991, P.W. Crous, (PREM 51291, holotype of G. eucalyptorum), USA, New York, Karner, on leaves of Vagnera stellata, April, C.H. Peck, (DAOM 4710, holotype of G. smilacinae); Georgia, Milledgeville, on fallen leaves of Bumelia lycioides, 20 March 1939, George E. Thompson & Julian H. Miller, Julian H. Miller Mycological Herbarium, Georgia Museum of Natural History, University of Georgia (GA 8357, holotype of G. bumelliae, NY, isotype of G. bumelliae). Pakistan, Lahore, Botanical garden on leaves of Heptapleurum venulosum, 25 July 1960, S. Ahmed (IMI 83132, holotype of G. foeniculata). Puerto Rico, on leaves of Piper sp., 22 September 1913, F.L. Stevens Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 Table 1 Synopsis of species synonymized under Phyllosticta capitalensis Species Guignardia aleuritis G. allamandae G. bumeliae G. cephalotaxi G. concinna G. endophyllicola G. eucalyptorum G. festiva G. foeniculata G. graminea G. helicteres G. pipericola G. smilacinae G. smilacis G. sterculiae Host plants (and mode of life) Aleurites fordii (dead leaves) Allamanda cathartica (leaves pathogen) Bumelia lycioides (dead leaves) Cephalotaxi drupaceae (living leaves) Cordia verruginae (living leaves) Rhododendron pulchrum (healthy leaves) Ascomata (μm) Peridium (μm) Ascospore shape* Asci (μm) Ascospores (μm) 93–125 × 100–125 13–25 A 90–150 × 94–100 12.5–15 A 70–100 × 70–90 20–25 B 70–160 × 100–160 17–20 C to B 70–100 × 94–100 12.5–15 C 118–156 × 8–139 23–39 A 31–50 × 8–13 41–67.5 × 12.5–15 44–76 × 10–24 45–80 × 13–17 50–80 × 18–24 38–54 × 14–15 11–13 × 3–4 15–20 × 6.6 15–19 × 7–8 11–18 × 5–9 14–20 × 7.5–10 9–16 × 4–5 Eucalyptus dives (weak leaf pathogen) Sumbaviopsis albicans (living leaves) Heptapleurum venulosum (living leaves) Atropis distans (dead leaves) Helicteres sp. (living leaves) Piper sp. (dead leaves) Vagnera stellata (dead leaves) Smilax aspera (dead leaves) Sterculia foetidae (dead leaves) 60–175 × 58–213 25–30 A 150–200 × 170–200 15–20 C to B 80–130 × 70–120 15–20 A 88–138 × 88–138 19–25 B 70–100 × 94–100 12.5–15 B 100–150 × 150–200 15–20 A 150–200 × 88–100 30–55 B 56–94 × 38–56 7.5–12.5 A 113–162 × 108–155 13–25 C 45–112 × 11–17 35–75 × 15–17 50–94 × 8–20 33–60 × 6–15 50–94 × 8–20 73–95 × 16–18 39–75 × 10–16 41–94 × 11–18 56–106 × 19–25 11–16 × 3–6 13–18 × 6–8 11–16 × 4–6 9–15 × 3–6 14–18 × 5–9 15–20 × 6–13 10–19 × 5–13 11–17 × 4–6 15–21 × 8–13 A = ellipsoidal swollen in the middle and obtrullate, B = obtrullate, C = clavate 151 Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 No. 4998 (ILL 9747, isotype of G. pipericola); ibid., Barcelona, on leaves of Helicteres jamaicensis, 10 August 1915, F.L. Stevens, Herbarium C.E. Chardon No. 1366, University Illinios Herbarum 9260 (BPI 1366, isotype of G. helicteres); ibid., on leaves of Helicteres, 7 July 1924, F.L. Stevens, Cornell University, Insular Experiment Station (NY 2615, holotype of G. helicteres). Philippines, Palawan Island, Palawan Province, Brooks Point (Addison Peak), on living leaves of Sumbaviopsis albicans, February 1911, A.D.E. Palmer, Philippine Island Plants No. 12637 (F 10766, from the holotype of G. festiva, BPI 598372, in poor condition). Phyllosticta capitalensis is described here based on the holotype specimen of G. aleuritis from dead leaves of Aleurites fordii, and other collections, viz. G. allamandae from Allamanda cathartica, G. endophyllicola from healthy leaves of Rhododendron sp., G. eucalyptorum as a weak pathogen on leaves of Eucalyptus dives, G. graminea from dead leaves of Atropis distans, G. smilacinae from dead leaves of Vagnera stellata, G. smilacis from dead leaves of Smilax aspera, and other species are synonymized under P. capitalensis as ascospores are ellipsoidal in shape and swollen in the middle. A synopsis of the synonyms is given in Table 1. Five authentic specimens of Guignardia aleuritis (= “Melanops aleuritica” (sic) Vassil.) have been examined and none of these handwritten labels read epithet as “aleuritidis” and designated as “sp. nov.”. Nevertheless, curator of LE (Dr Vadim Melnik, pers. comm.) is convinced that this material is what Vassiljevsky based his description of the new taxon (Vassiljevsky 1940). We designate the best of these specimens, LE 34833 as the lectotype to represent the taxon. Fresh collections from Thailand also have shown that P. capitalensis is generally found on Euphorbiaceae. Phyllosticta capitalensis occurs on many tropical plants as an endophyte (Bayen et al 2002, Glienke et al. 2011, Motohashi et al. 2011, Okane et al. 1998, 2001, 2003, Photita et al 2001, Wikee et al. 2011, 2013, Wulandari et al. 2011), as a weak pathogen (Crous et al. 1993, Glienke et al. 2011, Thongkantha et al. 2008) or as a saprobe following senescence of 145 host tissue as in Ficus leaves (Wang et al. 2008), Castanopsis acuminatissima, Smilax aspera (Roy 1967) and Aleurites fordii (Vassiljevsky 1940). There have been several molecular studies on Phyllosticta capitalensis (Baayen et al. 2002, Everett & Rees-George 2006, Glienke et al. 2011, Motohashi et al. 2008a,b, 2009, 2010, Pandey et al. 2004, Peres et al. 2007, Okane et al. 2001, 2003, Rodrigues et al. 2004, Sultan et al. 2011, Wang et al. 2012, Wickert et al. 2012, Wikee et al. 2011, 2013, Wong et al. 2012, Wulandari et al. 2009, 2012), the latest being by Wikee et al. (2013) which all show the wide host range of this endophytic species and its synonyms. Fig. 3 Phyllosticta capitalensis (PREM 51291, holotype of Guignardia eucalyptorum) on CLA media. a. Ascomata (arrowed). b. Ascomata on weed tissue in the media. c–d. Peridium comprising one strata of 2–3 layers of cells textura angularis without thickened brown cells. e–g. Asci. h–j. Ascospores – Scale bars: a–b = 3 mm, c–d = 10 μm, e–g = 15 μm, h–j = 5 μm. Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 Citrus spp.: P. citricarpa, P. citriasiana, P. citrichinaensis and P. capitalensis (Kiely 1948, Wulandari et al. 2009, Zhang et al. 2011). P. citriasiana differs from these other species in morphology and phylogeny. P. citriasiana found in Thailand occurs on Citrus maxima. Phyllosticta citricarpa forms a yellow zone in oatmeal agar whereas P. citriasiana does not. The conidia of P. citriasiana are larger (10– )12–14(–16) μm × (5–)6–7(–8) μm than those of P. citricarpa (9–10 × 6–7 μm). Although the species is pathogenic to Citrus maxima, this has not been included in EPPO most wanted species list (EPPO/CABI 1997). Phylogenetically, this species clusters in a separate clade to other Phyllosticta species with 93% bootstrap support (Wulandari et al. 2009). Fig. 4 Phyllosticta capitalensis (PREM 51291, holotype of Guignardia eucalyptorum) line drawing. k. Section of ascoma (arkened area fungal cells in arrowed region). l. Asci. m. Ascospores – Scale bars: k = 25 μm, l = 50 μm, m = 10 μm. Phyllosticta cavendishii M.H. Wong & Crous, Fungal Diversity 56: 177 (2012) Host – Musa sp. Wong et al. (2011) stated that the morphology of P. cavendishii is different from P. musarum, P. stevenshii, P. maculata and P. capitalensis also occurring on Musaceae. Conidia of P. cavendishii are smaller (12–)13– 16(–17) μm × 8–9(–10) μm than P. maculata (15–)16–19(–21) μm × (9–)10–12(–13) μm. Further, a maximum parsimonious tree showed P. cavendishii in a separate clade from other Phyllosticta species with 88% bootstrap support (Wong et al. 2011). P. cavendishii occurs on Musa cv. Cavendish and other cultivars. Phyllosticta citriasiana Wulandari, Gruyter & Crous, Fungal Diversity 34: 31 (2009) Host – Citrus maxima (Rutaceae) Four species of Phyllosticta occur on 146 Phyllosticta citribraziliensis C. Glienke & Crous, Persoonia 26: 54 (2011) Host – Citrus sp. (Rutaceae) P. citribraziliensis occurs on healthy leaves of Citrus sp. in Brazil (Glienke et al. 2011). This species is distinct in having larger conidia (8–)10–12(–13) μm × 6–7(–8) μm than the type of P. spinarum (8–)9.8(–12) μm × (6– )6.6(–7) μm. Phyllogenetically, this species forms a separate clade from P. spinarum with 68% bootstrap support (Glienke et al. 2011). Phyllosticta citricarpa (McAlpine) Aa, Stud. Mycol. 5: 40 (1973) ≡ Phoma citricarpa McAlpine, Fungus Diseases of Citrus trees in Australia: 21 (1899) = Guignardia citricarpa Kiely, Proc. Linn. Soc. N.S.W. 73: 259 (1948) Host – Citrus sinensis (Rutaceae) So far known from Australia, morphologically and phylogenetically P. citricarpa is different from P. citribraziliensis also described from Citrus (Glienke et al. 2011). Unfortunately, the culture that we examined was in poor condition. P. citricarpa has larger conidia than P. capitalensis, 9–10 × 6–7 μm vs. 8–10 × 4–5 μm. P. citricarpa forms a yellow hallow in oatmeal agar medium. P. citricarpa is closely related to P. citriasiana with 93% bootstrap support. P. citricarpa is an important quarantine plant pathogen (EPPO/CABI 1997). Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 Phyllosticta citrichinaensis X.H. Wang, K.D. Hyde & H.Y. Li, Fungal Diversity 52: 215 (2011) Host – Citrus maxima (Rutaceae) Wang et al. (2011) stated that P. citrichinaensis differs from other Phyllosticta species described from Citrus spp. based on conidia dimension, cultural characteristics and phylogeny. P. citrichinaensis (8–12 × 6–9 μm) has larger conidia than P. citribraziliensis (8– 12 × 6–7 μm). P. citrichinaesis has a paler appearance in culture as compared to P. citriasiana. The maximum growth temperature of P. citrichinaensis (30–35ºC) is somewhat intermediate between P. citricarpa (30–36ºC) and P. citriasiana (30–33ºC). The phylogenetic tree for combined three genes (ITS, actin and TEF1) showed a distinctive clade from P. citribraziliensis with 68% bootstrap support. Phyllosticta clematidis (W.H. Hsieh, Chi Y. Chen & Sivan.) Wulandari, comb. nov. Fig. 5 MycoBank: MB 805538 = Guignardia clematidis W.H. Hsieh, Chi Y. Chen & Sivan., Mycol. Res. 101: 901 (1997) MycoBank: MB 437714 Pathogen causing discoloration on tip of the leaf and the leaf becomes tan coloured: the area of infection wider in the vein and in the middle of leaf; paler coloured region and around the infection with numerous ascomata. Ascomata 160–180 μm diam., 130–140 μm high, superficial on the leaves, black, globose to subglobose, sometimes immersed in plant tissues, coriaceous, solitary to clustered, ostiole inconspicuous. Peridium 15–20 μm wide, comprising one strata of 2–3 layers of cells textura angularis with thickened brown walls. Pseudoparaphyses not observed. Asci 60–112 × 19–29 μm ( x = 91 × 25 μm, n = 20), 8-spored, bitunicate, ocular chamber 1–5 μm high, broadly cylindrical to cylindro-clavate, rounded at the apex, where the diam. is 12.5– 22.5 μm, tapering gradually to a 4–6 × 2.5– 37.5 μm long pedicel attached to the basal peridium. Ascospores 12.5–25 × 5–11 μm ( x = 19 × 8 μm, n = 20), biseriate or occasionally overlapping triseriate, obtrullate, hyaline to greenish, 1-celled, coarse-guttulate, smoothwalled, without a mucilaginous sheath. Host – Causing leaf spot of Clematis hayatai (Ranunculaceae) 147 Known distribution – Taiwan. Material examined – Taiwan, Piluchi, on leaves of Clematis hayatai, 28 April 1995, W.H. Hsieh (IMI 369599, holotype) The main character that distinguishes this species from all other Phyllosticta sexual states is the ascospore appendage (Fig. 6, arrowed). The ascospores inside the asci are diamond shaped and when released they are broadly fusiform. P. clematidis has larger conidia (12.5–25 × 5–11 μm) than P. capitalensis (15–17 × 5–6 μm) (Glienke et al. 2011). Fig. 5 – Phyllosticta clematidis (IMI 369599, holotype) a. Leaf spot. b. Ascomata on the host surface. c–d. Peridium comprising one strata of 2–3 layers of cells textura angularis with thickened brown walls. e–f. Asci. g. Ascospores with an appendage (arrowed). – Scale bars a = 3 mm, b = 100 μm, c–d = 16 μm, e–f = 29 μm, g = 11 μm. Phyllosticta cruenta (Fr.) J. Kickx f., Mém. Acad. R. Sci. Lett. Arts. Brux. 23: 22 (1849) Fig. 6 ≡ Sphaeria cruenta Fr., Syst. Mycol. 2: 581 (1823) Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 arising from the cells lining the pycnidial locule. Conidia 10–15 × 4–6 μm ( x = 12 × 5 μm, n = 20), hyaline to greenish, 1-celled, coarse-guttulate, smooth-walled, globose, ellipsoidal, clavate or obclavate, with an obtuse apex, sometimes truncate at the base, surrounded by 1–2 μm thick mucilaginous sheath which persists at maturity and in some conidia with a single, 2–6 μm long, hyaline, curved or straight apical appendage. Host – Causing leaf spot of Polygonatum multiflorums (Liliaceae). Known distribution – Romania. Material examined – Romania, CaraşSeverin, on leaves of Polygonatum multiflorum, 8 June 1976, G. Negrean, Herbarium Mycologicum Romanicum No. 2728, Herbarium Horticulture Botany Matrit (MA 8727, holotype). The conidial dimensions of the holotype (10–15 × 4–6 μm) differ from that of CBS 858.71 (12–21 × 5–10 μm) examined by van der Aa. van der Aa (1973) described two species occurring on Polygonatum spp., i.e P. subeffusa and P. cruenta. Fig. 6 – Phyllosticta cruenta (MA 8727, holotype). a–b. Leaf spot. c. Pycnidia. d. Peridium comprising one strata of 2–3 layers of cells textura angularis with thick brown walls. e. Conidiogenous cells. f. Conidia. – Scale bars – b = 200 µm, c = 100 µm, d = 30 µm, e = 15 µ m, f = 6 µm. = Guignardia reticulata (DC.) Aa, Stud. Mycol. 5: 23 (1973) = Phyllosticta convallariae Pers., Traité sur les Champignons Comestibles (Paris): 148 (1819); inval. name (Art. 32. 1) Pycnidia 143–170 μm diam., 88–136 μm high, on the surface of the leaf, black, globose to subglobose, partly immersed in leaf tissues, coriaceous, solitary to clustered, ostiolate, with ostioles as black dots in the centre, often growing together with ascomata. Peridium 11–23 μm wide composed of 2–3 layers of cells textura angularis pigmented in the outmost layers around ostiole, with inner wall layer cells paler. Conidiogenous cells 6– 14 × 2–3 μm ( x = 9 × 2 μm, n = 20), holoblastic, determinate, discrete, rarely integrated, hyaline, cylindrical to doliiform, 148 Phyllosticta cussoniae Cejp, Bothalia 10: 342 (1971) Fig. 7 = Guignardia cussoniae Crous, South Africa J. Bot. 59: 606 (1993) Pycnidia 160–180 μm diameter, 100– 160 μm high, on the surface of the leaf, black, globose to subglobose, partly immersed in leaf tissues, coriaceous, solitary to clustered, ostiolate, with ostioles as black dots in the centre, often growing together with ascomata. Peridium 15–31 μm wide composed of two to three layers of cells, arranged in a textura angularis and pigmented in the outer layers and around ostiole, paler inner layer. Conidiogenous cells 6–14 × 2–3 μm ( x = 9 × 2 μm, n = 20), holoblastic, determinate, discrete, rarely integrated, hyaline, cylindrical to doliiform, arising from the cells lining the pycnidial locule. Conidia 8–15 × 4–6 μm ( x = 11 × 5 μm, n = 20), hyaline to greenish, 1celled, coarse-guttulate, smooth-walled, globose, ellipsoidal, clavate or obclavate, with an obtuse apex, sometimes truncate at the base, surrounded by 2–5 μm ( x = 3 μm, n = 20) thick mucilaginous sheath which persists at maturity and in some conidia with a single, 6– Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 12 μm ( x = 10 μm, n = 20) long, hyaline, curved or straight apical appendage. Host – Causing leaf spot of Cussonia umbellifera (Araliaceae). Known distribution – South Africa. Material examined – South Africa, Eastern Transvaal, Schagen, Nelspruit district, on living leaf of Cussonia umbellifera, 25 December 1933, L.C.C. Liebenberg (PREM32821, holotype). Conidia of P. cussoniae (8–15 × 4–6 μm) are smaller than those of P. capitalensis (8–15(–18) × 6–10 μm). Phylogenetically, P. cussoniae segregates into a distinct clade from other species of Phyllosticta, such as P. bifrenariae, P. citricarpa, P. citriasiana, P. citribraziliensis, P. spinarum and P. hypoglossi with 98% bootstrap support (Glienke et al. 2011). Fig. 7 – Phyllosticta cussoniae (PREM 32821, holotype). a, b. Pycnidia. c–d. Peridium comprising one strata of 2–3 layers of cells textura angularis with an apex of thickened brown walls. e. Conidiogenous cells. f–g. 149 Conidia. – Scale bars b = 200 µm, c = 100 µm, d = 30 µm, e–f = 15 µm, f = 6 µm. Phyllosticta ericarum Crous, Persoonia 28: 212 (2011) Host – Erica gracilis (Ericaceae) Phyllosticta ericae, P. ericarum and P. gaultheriae were described from Ericaceae (van der Aa 1973, van der Aa & Vanev 2002, Crous 2011). P. ericae is identical to P. pyrolae based on morphological characters. P. pyrolae is similar to P. ericae described from Erica carnea (Ericaceae). Okane et al. (2001) showed that P. pyrolae is distinct from 14 strains from Ericaceae based on ITS sequence data. Conidia of P. ericarum ((10–)11–12(–14) × (5–)6–7 μm)) are slightly longer than those of P. pyrolae (4.5–7.5 × 4–9 μm). This species also has smaller conidia than P. capitalensis. Blast results data revealed that P. ericarum is closely related to Guignardia philoprina in ITS sequence data and P. hymenocallidicola in LSU sequence data (Crous et al. 2012). Phyllosticta garciniae (I. Hino & Katum.) Motohashi, Tak. Kobay. & Yas. Ono, Mycoscience 51: 94 (2010) Fig. 8 = Guignardia garciniae I. Hino & Katum., Bull. Fac. Agric. Yamaguti Univ. 16: 607 Ascomata 175–250 μm diam., 250–375 μm high, on the upper surface of the leaves, black, slightly brown to black, globose to subglobose, semi-immersed to semi immersed in leaf tissues. Peridium 55–125 μm wide, one strata of 2–4 layers of cells textura angularis with thickened brown walls around ostiole. Pseudoparaphyses not observed. Asci 50–95 × 16–21 μm ( x = 16 × 21 μm, n = 20), 8-spored, bitunicate, cylindrical to cylindro clavate, rounded at the apex, where the diam. is 10–16 μm, tapering gradually to a 6–19 × 6–8 μm long pedicel attached to the basal peridium. Ascospores 13–21 × 6–10 μm ( x = 17 × 8 μm, n = 20), uniseriate to biseriate, ellipsoidal, widest 2/5 near the apex (obtrullate), hyaline to greenish, 1-celled, coarse-guttulate, smoothwalled, with mucilaginous sheath at one end. Host – On living leaves of Garcinia spicata (Clusiaceae). Known distribution – Japan. Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 Material examined – Japan, Ryukyu Island on leaves of Garcinia spicata, 24 June 1961, Hino & Katumoto, Herbarium laboratory of Plant Pathology Faculty of Agriculture Yamaguti University Number 21744, Japan (TFM 21744, holotype of Guignardia garciniae). P. garciniae is distinct from other Phyllosticta species in that ascomata are black with beak whitest on the top, a peridium more than 50 μm thick, and a polar pad at one side of the ascospores. Motohashi et al. (2010) found the asexual stages of this fungus. This is a good species in the genus because of its unique ascomata, peridium and ascospore features. Phyllosticta gaultheriae Aa, Stud. Mycol. 5: 56 (1973) = Guignardia gaultheriae Aa, Stud. Mycol. 5: 56 (1973) Host – Gaultheria procumbens (Ericaceae) van der Aa (1973) stated that P. gaultheriae is comparable to P. pyrolae in regard to conidial size (4–9 × 4–7 μm in P. gaultheriae and 4.5–7.5 × 4–9 μm in P. pyrolae). They differ only in apical appendage that reduces into an inconspicuous sheath. Su & Cai (2012) also mentioned the variable and distinct conidial dimension of P. gaultheriae and P. ilicis-aquifolii. Phylogenetically P. gaultheriae is distinct from P. bidwellii (CBS 447.70 strain) and P. ilicis-aquifolii (Sultan et al. 2011, Su & Cai 2012, Zhang et al. 2013b). Phyllosticta hostae Y.Y. Su & L. Cai, Persoonia 28: 79 (2012) Host – Hosta plantaginia (Liliaceae) P. hostae is the only Phyllosticta species on Hosta plantaginia. Conidia of P. hostae (8–15(–18) × 6–10 μm) are smaller than those of P. hypoglossi (8–15 × 5–9 μm). Phylogenetic analyses by maximum parsimony tree and combined four genes (ITS, EF, actin and GPDH) showed that this species is distinct and in a separate subclade from P. cussoniae, P. hypoglossi, P. citribraziliensis and P. spinarum with 99% bootstrap value (Su & Cai 2012). Fig. 8 – Phyllosticta garciniae (TFM 21744, from the holotype of Guignardia garciniae). a– b. Appearance of ascomata on the host surface with withest on the top (arrowed). c–d. Peridium comprising one strata of 2–3 cells textura angularis with thickened brown walls. e–g. Asci. h–i. Ascospores with mucilaginous sheath (arrowed). – Scale bars b = 250 μm, c, e–i = 10 μm, d = 20 μm. 150 Phyllosticta hubeiensis K. Zhang, Y.Y. Su & L. Cai, Mycol. Progress 12 (On Line article) (2013) Host – Viburnum odoratissimum (Adoxaceae) van der Aa (1973) stated that the conidial dimension of P. hubeiensis (10–14.5 × 6–9 μm) is smaller than P. concentrica (8–20 × 4–14 or 10–15 × 7–10 μm). The conidiogenous cells and spermatia also differ in size (van der Aa 1973). This species is special in that its conidia fail to detach from the conidiogenous cells. Phylogenetically P. hubeiensis is closely related to P. concentrica (previously described as the asexual state of Guignardia philoprina) and P. gaultheriae in ITS sequences data (Zhang et al. 2013a). Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 Phyllosticta hymenocallidicola Crous, Summerell & Romberg, Persoonia 27: 139 (2011) Host – Hymenocallis littoralis (Amaryllidaceae) P. hymenocallidicola is the only species that occurs on Hymenocallis littoralis as P. hymenocallidis was found to be Phoma narcissi (van der Aa 1973). Conidia of P. hymenocallidicola [(8–)9–10(–11) × (6–)6.5–7 μm] are smaller than those of P. capitalensis [8–15(–18) × 6–10 μm)]. P. hymenocallidicola is phylogenetically distinct from other Phyllosticta species that have been submitted to GenBank (Crous et al. 2011, Glienke et al. 2011). Crous et al. (2011) showed that P. hymenocallidicola is close to P. owaniana based on mega BLAST using ITS sequences. Phyllosticta hypoglossi (Mont.) Allesch., Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1(6): 163 (1898) = Macrophoma hypoglossi (Mont.) Berl. & Voglino, Atti Soc. Veneto-Trent. Sci. Nat. 10(1): 192 (1886) = Phoma hypoglossi (Mont.) Sacc., Syll. Fung. 3: 162 (1884) = Phyllostictina hypoglossi (Mont.) Petr. & Syd., Beih. Reprium nov. Spec. Regni veg.: 203 (1927) ≡ Sphaeropsis hypoglossi Mont., Annls Sci. Nat., Bot., sér. 3, 12: 307 (1849) Host – Ruscus hypoglossum (Liliaceae) The conidial dimensions and other characters show differences from other Phyllosticta species occurring on Liliaceae. P. hypoglossi has bigger conidia and a longer appendage sheath compared to P. yuccae, i.e., 7.5–15.4 × 6–9.5 μm vs. 8–15(–18) × 6–10 μm and 4–15 μm vs. 10–35 μm for P. hypoglossi. This species forms a distinct clade from other Phyllosticta species (Glienke et al. 2011, Su & Cai 2012). Phyllosticta ilicis-aquifolii Y.Y. Su & L. Cai, Persoonia 28: 81 (2012) Host – Ilex aquifolium (Aquifoliaceae) P. ilicis-aquifolii is characterized by its large conidia and long mucoid sheath or appendage. The conidial dimensions of P. ilicis–aquifolii (4–9 × 4–7 μm) are smaller than P. gaultheriae (10–18 × 6–9 μm). The 151 spermatial state and conidiogenous cells also differ in shape and dimensions. P. ilicisaquifolia has grey to leaden grey growth in culture with an irregular margin, whereas P. gaultheriae has greenish to black cultures with a lobed margin (van der Aa 1973). P. ilicisaquifolii is phylogenetically close to P. gaultheriae based on ITS sequences with 94% bootstrap support. Phylogenetic tree generated from DNA combined sequences alignment showed that this species is closely related to P. gaultheriae and P. yuccae with 100% bootstrap support (Su & Cai 2012). Phyllosticta korthalsellae A. Sultan, P.R. Johnst., D.C. Park & A.W. Robertson, Stud. Mycol. 68: 241 (2011) Host – Korthalsella lindsayi (Viscaceae) This is first report of a Phyllosticta from Korthalsella from New Zealand. Based on ITS sequence data, it was inferred that P. korthalsella is related to P. eugeniae and Guignardia bidwellii with 76% bootstrap support (Sultan et al. 2011). Phyllosticta maculata Wong & Crous, Fungal Diversity 56: 180 (2012) Fig. 9 = Guignardia musae Racib., Bull. Int. Acad. Sci. Lett. Cracovie, Cl. sci.math. nat. Sér. B, sci. nat. 3: 388 (1909) Ascomata 100–125 μm high, 75–150 μm diam., on upper and lower surface of leaf and on banana fruit skin, globose to subglobose, black, semi-immersed in plant tissues, coriaceous, solitary to clustered, ostiolate, with ostiole as black centrally located dots. Peridium 12.5–20 μm wide, upper part composed of compressed brownish, thinwalled cells, 1–4 cells thick, lower part hyaline, composed of flattened, dark brown cells, darkest around the ostiole. Asci 49–105 × 16– 28 μm ( x = 74 × 21 μm, n = 20), 8-spored, bitunicate, broadly cylindro-clavate, rounded at the apex, where the diam. is 8–21 μm, tapering gradually to a 5–10 × 5–10 μm long pedicel attached to the basal peridium, ocular chamber 3–8 μm high. Ascospores 20–25 × 8–13 μm ( x = 22 × 10 μm, n = 20), uniseriate or occasionally overlapping biseriate, clavate to oblong, hyaline to greenish, 1-celled, coarseguttulate, smooth-walled, without a Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 mucilaginous appendages. Host – on living leaves causing freckle leaves of Musa spp. (Musaceae) Known distribution – American Samoa, Australia, Indonesia, Malaysia, Papua New Guinea. Material examined – Indonesia, Bogor, on leaves of Musa acuminata, Raciborski, (KRA 063561, from holotype of Guignardia musae). This species was recently reidentified by Wong et al. (2012). The new name “maculata” was derived from the commonly caused freckle diseases worldwide that occurs on Musa spp. Four other species of Phyllosticta are known on Musaceae, P. musae Racib. (Raciborski 1909), P. musarum, P. capitalensis and P. cavendishii (Wong et al. 2012). Phylogenetically, P. maculata forms a distinct clade, from other Phyllosticta occuring on Musaceae, and is close to P. vaccinii and P. cavendishii, with 73% based on LSU sequence and 88% on ITS sequence bootstrap support (Wong et al. 2012). Fig. 9 – Phyllosticta maculata (KRA 063561, from holotype of Guignardia musae) a. Ascomata on the host surface. b–c. Peridium comprising sterile tissue of plant cells. d–e. Asci. f–g. Ascospores. – Scale bars: a = 100 μm, b–c = 20 μm, d = 30 μm, e–g = 10 μm. 152 Phyllosticta morindae (Petr. & Syd.) Aa, Stud. Mycol. 5: 69 (1973) = Guignardia morindae (Koord.) Aa, Stud. Mycol. 5: 69 (1973) = Phyllostictina morindae Petr. & Syd., Feddes Repert., Beih. 42: 200 (1927) = Physalospora morindae Koord., Verh. K. Akad. Wet., tweede sect. 13(4): 190 (1907) Host – Morinda citrifolia (Rubiaceae) The type is not available in BO (Mien A. Rifa’i pers. comm.). Phyllosticta morindae is distinct from other species in having smaller conidia than P. capitalensis and P. maculata. Phyllosticta musarum (Cooke) Aa, Stud. Mycol. 5: 72 (1973) = Macrophoma musae (Sacc.) Berl. & Voglino, Atti Soc. Veneto–Trent. Sci. Nat. 10(1): 187 (1886) = Phoma musae Sacc., Syll. Fung. 3: 163 (1884) = Sphaeropsis musarum Cooke, Grevillea 8: 93 (1880) Host – Musa sp. (Musaceae) P. musarum is morphologically distinct from other species of Phyllosticta occurring on banana (Musaceae). It occurs only on leaves of banana types AAB and ABB. P. musarum differs from P. cavendishii by its conidial dimensions and phylogenetic affinities. P. musarum has bigger conidia [(12–)13–16(–20) × (7–)9–10(–11) µm] than P. cavendishii, [(12–)13–16(–17) × 8–9(–10) µm]. Phylogenetic data showed that P. musarum is closely related to P. cavendhishii with 63% bootstrap support. Phyllosticta muscadinii (Luttr.) Wulandari, comb. nov. et stat. nov. Fig. 10–11 MycoBank: MB 805539 = Guignardia bidwellii f. musacadinii Luttr., Phytopathology 36: 913 (1946) MycoBank: MB 346168 Pathogenic, leaf spots, irregular to regular with a red brown border and cream centre with numerous ascomata and pycnidia. Ascomata visible on the surface of the leaf as black dot, 70–100 μm diam., 94–100 μm high, black, globose to subglobose, immersed in plant tissue, coriaceous, solitary to clustered, Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 ostiolate, with ostioles as black dots in the centre. Peridium 12.5–15 μm wide, comprising one strata of 2–6 layers of cells textura angularis with thickened brown walls. Pseudoparaphyses tubular. Asci 50–80 × 18–35 μm ( x = 77 × 15 μm, n = 20), 8-spored, bitunicate, with an ocular chamber 1–2 μm high, broadly cylindrical to cylindric-clavate, rounded at the apex, where the diam. is 13–18 μm, tapering gradually to a 7.5–16 × 2–6 μm very long pedicel attached to the basal peridium. Ascospores 14–20 × 7.5–10 μm ( x = 15 × 7 μm, n = 20), biseriate or occasionally overlapping uniseriate, ovate to oblong, hyaline to greenish, 1-celled, coarse guttulate, smoothwalled, without a hyaline mucilaginous sheath at the base. Pycnidia 50–100 μm diam., 70–100 μm high, black, globose to subglobose, immersed in plant tissues, coriaceous, solitary to clustered, ostiolate, ostioles as a black dots in the centre. Peridium 10–15 μm wide, comprising one strata of 1–2 layers of cells textura angularis with thickened brown walls. Conidiogenous cells short cylindrical 16–41 × 11–15 μm ( x = 24 × 13 μm, n = 20). Conidia 7–10 × 4–7 μm ( x = 8 × 6 μm, n = 20), oblong, 1-celled, containing coarse guttules, smoothwalled, 1–2 μm wide mucilaginous sheath and 3–6 (4) μm long appendage. Host – Causing leaf spots of Muscadinia rotundifolia (Vitaceae) Known distribution – USA. Material examined – USA, on overwintered leaves of Muscadinia rotundifolia, Experiment Spalding Co., 15 April 1945, E.S. Luttrell, Georgia Cryptogams (BPI 598288, isotype). Phylogenetically there is a species complex of Phyllosticta in Vitaceae. Zhang et al. (2013b) mentioned that P. ampelicida, P. bidwellii and P. parthenocissi form three distinct clades. This finding is similar to Witch et al. (2012) who also showed that more than two species occur on Vitaceae (Witch et al. 2012, Zhang et al. 2013b). P. muscadinii is a good species based on pathogenicity tests (Luttrell 1946). Furthermore P. muscadinii has smaller conidia than P. ampelicida and P. parthenocissi. 153 Fig. 10 – Phyllosticta muscadinii (BPI 598288, isotype). a, h. Leaf spot. b, i. Ascomata on the host surface of fruits. c–j. Peridium comprising one strata of 2–4 layers of cells of textura angularis with thickened brown walls. d–e. Asci with pseudoparaphyses. f–g. Ascospores. i. Pycnidia on the host surface. j–k. Section of pycnidium. l–m. Conidiogenous cells. n–p. Conidia. – Scale bars: c = 50 μm, d–e = 25 μm, f–g = 10 μm, j = 100 μm, k–m = 15 μm, n–p = 7 μm. Phyllosticta owaniana G. Winter, Hedwigia 24: 31 (1885) Host – Brabejum stellatifolium (Proteaceae) Phyllosticta owaniana is a distinct species and forms a separate clade in the genus (Wulandari et al. 2009, Glienke et al. 2012, Zhang et al. 2013). It differs from P. capitalensis by its shorter and wider conidia (10–14 × 7–8.5 μm for P. owaniana and 15–17 × 5–6 μm for P. capitalensis) (van der Aa 1973, van der Aa & Vanev 2002, Glienke et al. 2011). Phyllosticta partenocissi K. Zhang, N. Zhang & L. Cai, Mycologia 105(4): 1030 (2013) Host – Parthenocissus quinquefolia (Vitaceae) Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 This species is phylogenetically distinct from other species (Wulandari et al. 2009, Wikee et al. 2011, Sultan et al. 2011, Zhang et al. 2013b). Phyllosticta philoprina is distinct from P. concentrica (Motohashi et al. 2009), which was previously synonymised as the asexual state of Guignardia philoprina (van der Aa 1973). Phyllosticta schimae Y.Y. Su & L. Cai, Persoonia 28: 80 (2012) Host – Schima superba (Theaceae) Phyllosticta schimae is distinct from P. parthenocissi based on morphology and phylogeny (Su & Cai 2012). Phyllosticta schimae has longer but narrower conidia (7–13 × 4–7 μm) than those of P. parthenocissi (7.5– 10 × 6–9 μm). Maximum parsimony derived from four sequences (ITS, EF, GDPH and actin) showed that P. shimae is closely related to P. parthenocissi (strain CBS 111645) with 51% bootstrap support (Zhang et al. 2013b). Fig. 11 – Phyllosticta muscadinii (BPI 598288, isotype) line drawing. a. Section of ascoma on the leaf. b. Asci. c. Ascospores. – Scale bars = 25 μm. Morphologically this fungus is distinct from other species of Phyllosticta occurring on Vitaceae. The dimensions of conidia differ from P. ampelicida (Zhang et al. 2013). Phyllosticta parthenocissi was described as a new species and a replacement name for strain CBS 111645, previously described as Guignardia bidwellii. The maximum parsimony tree showed that P. parthenocissi associated with Parthenocissus is a distinct clade from that of seven strains from Japan associated with Vitis sp. with 55% bootstrap support (Zhang et al. 2013b). Phyllosticta philoprina (Berk. & M.A. Curtis) Aa, Stud. Mycol. 5: 44 (1973) = Guignardia philoprina (Berk. & M.A. Curtis) Aa, Stud. Mycol. 5: 44 (1973) = Guignardia ilicis (Jacz.) Schrantz, Bull. Trimmest. Soc. Mycol. Fr. 76: 327 (1961) = Guignardia rhodorae (Cook) B.H. Davis, Mycologia 38: 48 (1946) Host – Taxus baccata (Taxaceae) 154 Phyllosticta spinarum (Died.) Nag Raj & M. Morelet, Bull. Soc. Sci. nat. Arch. Toulon et du Var 34(219): 12 (1978) ≡ Phoma spinarum Died., Krypt.-Fl. Brandenburg (Leipzig) 9: 148 (1912) Host – Juniperus sp. (Cupressaceae) Phyllosticta spinarum has smaller conidia lacking a mucoid sheath as compared to P. citribraziliensis. Phyllosticta spinarum is phylogenetically distinct from other species in maximum parsimony tree when using analysis of ITS, EF, actin and GDPH gene sequences (Glienke et al. 2011). It forms a separate clade from P. citribraziliensis with 68% bootstrap support (Glienke et al. 2011). Phyllosticta styracicola K. Zhang & L. Cai, Mycological Progress 12: 551 (2013) Host – Styrax grandiflorus (Styracaceae) Phyllosticta styracicola is a distinct species with conidia 9.5–13 × 6.5–9 μm, which are smaller than P. schimae (7–13 × 4–7 μm) and longer than P. parthenocissi (7.5–10 × 6–9 μm). Phyllosticta styracicola clustered in a separate clade from P. schimae and P. parthenocissi with 87% bootstrap support (Zhang et al. 2013a). Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 Phyllosticta vaccinii Earle, Bull. Torrey Bot. Club 24: 31 (1897) ≡ Phyllosticta vaccinii (Earle) Shear, Mycologia 15: 131 (1923) = Phyllosticta sparsa Bonar, Mycologia 20: 296 (1928) = Phyllosticta vaccinii Demaree & Wilcox, Phytopathology 37: 292 (1947) Host – Vaccinium macrocarpon (Ericaceae) Polymorphism exists in P. vaccinii and this species has already been epitypified by Zhang et al. (2013b). The parsimony tree presented in Zhang et al. (2013b) indicates that Guignardia vaccinii Shear is not the same biological species as Phyllosticta vaccinii as they fall into different clades in sequence analysis (Zhang et al. 2013b). Guignardia vaccinii Shear is morphologically and phylogenetically distinct from Phyllosticta vaccinii, which was previously recorded as the asexual state of Guignardia vaccinii. This anamorph and teleomorph connection was proposed by Shear in 1907 (Zhang et al. 2013b). Weidemann et al. (1982) regarded P. elongata as the anamorph state of G. vaccinii, this connection need to be clarified. The clade E in the paper published by Zhang et al. (2013b), known as the ex-type of Guignardia vaccinii (CBS 126.22) forms a separate clade from ATCC 46255 epitypified as P. vaccinii by Zhang et al. (2013b). Discussion Species of Phyllosticta were named mainly on morphological characters and molecular phylogeny based on recent publications. Some species are distinguished because of their unique appendages, as in P. candeloflamma and P. clematidis. Phyllosticta garciniae has an unusual thick peridium in the ascomata. Both sexual and asexual states were present on the P. muscadinii specimen examined. The material of P. candeloflamma had only the sexual state whereas only asexual state was present on P. cussoniae. Phyllosticta capitalensis was recorded on numerous tropical plants. Guignardia mangiferae is a genetically and morphologically distinct species from previously recorded Phyllosticta species on Mangiferae indica. The name P. mangiferae has already been used and it is necessary to 155 choose another name as and when more isolates become available; thus the name of G. mangiferae is still available as a valid name (Glienke et al. 2011). Pure cultures from single spore isolations of several Phyllosticta species are not available and therefore gene sequence data and analyses have not been possible for many species. It is important to recollect and restudy the taxonomy and phylogeny of important old species. There are several problems, which inhibited taxonomic work during the course of this study, and are noted below: 1. Low numbers of ascomata in the voucher holotype specimens. 2. The low quality of holotype specimens; some specimens were in poor condition. 3. Need for epitypifying many species. 4. The need for a polyphasic approach and multi-gene phylogenies to accurately resolve species (Wang et al. 2012, Zhang et al. 2013b). A single gene that can resolve species well is needed. 5. Need for cultures of each species for obtaining sequences for phylogenetic analyses. Single spore isolation of many Phyllosticta speies is not possible (Crous et al. 2009) and attempts to isolate from ascomata or pycnidia may be necessary (Zhang et al. 2009c). Acknowledgements This study was funded in part by the Thailand Research Fund (grant DBG5380011). The directors and curators of herbaria B, BPI, BRIP, DAR, F, FH, G, K, KRA, LE, MA, NTU, NY, PAV, PDD, PH, PREM, SIENA, TNS, TUR, W, YAM, and ZT are profoundly thanked, for loaning specimens including holotypes. The Mushroom Research Foundation is thanked for a PhD scholarship to Nilam F. Wulandari during 2007–2011. The University of Hong Kong, Hong Kong and the School of Science, Mae Fah Luang University, Thailand and CBS, The Netherlands, are thanked for laboratory facilities. Dr Pedro W Crous is thanked for partial funding of this research. Dr Kevin D. Hyde is acknowledged for valuable comment and discussion on this manuscript. Dr Richard T. Hanlin is thanked for supplying valuable references. Dr Vadim Mel’nik is thanked for translating some articles Plant Pathology & Quarantine — Doi 10.5943/ppq/3/2/4 from Rusian. Dr Keichi Motohashi is thanked for Japanese references on Phyllosticta and “Guignardia”. Dr Okane Izumi is thanked for cultures of G. endophyllicola. References Aa HA van der, Vanev S. 2002 – A Revision of the species described in Phyllosticta. Centraalbureau voor Schimmelcultures, Utrecht, The Netherlands. 1–49. Aa HA van der. 1973 – Studies in Phyllosticta 1. 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