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Mycosphaerellaceous Fungi and New Species of Venustosynnema and Zasmidium on Ferns and Fern Allies in Taiwan

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Mycosphaerellaceous Fungi and New Species of Venustosynnema and Zasmidium on Ferns and Fern Allies in Taiwan Phytotaxa 176 (1): 309–323 ISSN 1179-3155 (print edition) www.mapress.com/phytotaxa/ Article PHYTOTAXA Copyright © 2014 Magnolia Press ISSN 1179-3163 (online edition) http://dx.doi.org/10.11646/phytotaxa.176.1.29 Mycosphaerellaceous fungi and new species of Venustosynnema and Zasmidium on ferns and fern allies in Taiwan ROLAND KIRSCHNER & LI-CHIA LIU 1 Department of Life Sciences, National Central University, Jhongli City, 320 Taoyuan County, Taiwan email: [email protected] Abstract Selected fungi on ferns in Taiwan are described, illustrated, annotated and complemented with first DNA sequences. The ranges of distribution, host species, morphology, and the taxonomy of some species were revised. ITS rDNA sequences proved to be useful for distinguishing between species on the same host. A new species of Venustosynnema found on dead leaves of Selaginella moellendorfii differs from the other species of the genus by its small conidiomata with smooth central seta and reniform conidia. A new species of Zasmidium described on living fronds of Dicranopteris linearis is particular by its hyaline, smooth conidia. New data are derived from new collections of Pseudocercospora athyrii, Ps. cyatheae, and Ps. pteridophytophila hitherto only known from the type collections and of Mycosphaerella gleicheniae. Ps. christellae is considered a synonym of Ps. pteridophytophila. M. gleicheniae and Ps. cyatheae are new records for Taiwan. Key words: Ascomycota, Capnodiales, Cyclosorus, Deparia, Dicranopteris, plant pathogens, Sphaeropteris Introduction Stevenson (1945) was probably the pioneer researcher who published an overview of fungi parasitic on ferns. The most recent overview of saprobic, parasitic and symbiotic fungi associated with ferns was published by Mehltreter (2010). The relationship between rust fungi and ferns was first to attract the attention of phylogenetic mycologists (see Sjamsuridzal et al. 1999 and references therein). Molecular analyses indicated that fern rusts were not basal to the rusts on seed plants (Sjamsuridzal et al. 1999). In contrast to the rust fungi, which are biotrophs on their host plants, evidence for parasitism by other fungi on ferns is often not clear or is only concluded from field observations. These studies mainly focused on the systematics of sexual and asexual Ascomycota including saprobic as well as presumably parasitic species (e.g. Bubák 1916, Cannon 1997, Castañeda Ruiz & Heredia 2000, Dingley 1972, Medel & Lorea-Herrnández 2008, Samuels & Rogerson 1990). The study of Ascomycota on ferns is still in the pioneer stage with biodiversity being recorded and the study of saprobic fungi on fern leaves is expected to result in new discoveries (Mehltreter 2010). Among the Ascomycota, mycosphaerellaceous fungi, including sexual species of Mycosphaerella and related genera, as well as their corresponding cercosporoid anamorphs, are particularly species-rich and in most cases are considered to be highly specific to host genera and families of vascular plants (Crous & Braun 2003). Taiwan has one of the highest density of ferns and is, therefore, particularly suitable for studying the fungi on ferns. Given approximately 700 species of ferns known each for Taiwan, Japan, and Thailand, the species density is 0.019, 0.003, and 0.001 taxa/km2, respectively (calculated from Lu & Yang 2005). Because of the high population and species density of ferns, fungi preferring ferns as substrate are also likely to be found frequently in Taiwan. Approximately 30 species of Pseudocercospora are known from ferns (Braun et al. 2013), and six species were described from and are only known from Taiwan (Hsieh & Goh 1990, Kirschner & Chen 2007). A worldwide study on species of the cercosporoid genus Periconiella on ferns was published by Braun (2004); two of the species were later recorded from Taiwan (Kirschner & Chen 2007, 2010). Braun et al. (2013) compiled information concerning cercosporoid hyphomycetes on ferns worldwide based on the literature and herbaria. Cercosporoid fungi are considered comparatively host-specific and are identified to species mainly based on morphological Accepted by Kevin Hyde: 13 Mar. 2014; published: 20 Aug. 2014 309 Licensed under a Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0 comparison of congeneric species known from members of the same host genus or family. The conspectus of Mycosphaerella by Aptroot (2006) contains more than 30 taxa on ferns and fern-allies, although several species on different ferns were morphologically identical with M. filicum (Desm.) Starbäck (Aptroot 2006). Classification of ferns, however, has been unsettled over the last decades including frequent changes to concepts of families and genera (Christenhusz et al. 2011, Frey et al. 2009, Kramer et al. 1990, followed in Knapp 2011, Rothfels et al. 2012, Smith et al. 2006). In contrast to fungi parasitic on specific seed plants, which can often be conveniently identified based on the knowledge of the host genera and families, this approach might be misleading for the fungi on ferns. Several species of fern-inhabiting cercosporoid fungi are known only from the type collection and lack cultures and DNA data. It is, therefore, difficult to rely only on the original morphological descriptions for identification. In particular, conidiophore lengths can vary considerably between different specimens of the same species (Kirschner 2013). The aim of the study was to increase our knowledge of the fungal diversity of fungi on ferns from Taiwan by providing collections of species previously being unknown or only known from the type, revising their morphology, and depositing cultures and DNA sequences as far as possible. TABLE 1. Collection data and GenBank accession numbers of species of Mycosphaerella, Pseudocercospora, and Zasmidium from Taiwan with ITS rDNA sequences newly generated in this study. Species Host Collection no., Collection data in Taiwan GenBank strain no. 1 no. M. gleicheniae T.S. Ramakr. Dicranopteris linearis RoKi 3613 New Taipei City, Yingge, trail to KJ201929 & K. Ramakr. (Burm. f.) Underw. var. Yingge Rock, 100–250 m, 11 April linearis (Gleicheniaceae) 2012 M. gleicheniae Dicranopteris linearis var. RoKi 3945 Taoyuan County, Dasi (Daxi) KJ201930 linearis (Gleicheniaceae) Township, Weiliao Old Trail, ca. 150 m, 29 September 2013 Ps. athyrii Goh & W.H. Hsieh Deparia longipes (Ching) RoKi 3943 Taoyuan County, Dasi (Daxi) KJ201932 Shinohara Township, Weiliao, ca. 150 m, 29 (Dryopteridaceae) September 2013 Ps. camelliicola U. Braun & Camellia japonica cultivar RoKi 3774, Hsinchu County, Naluo, 12 October KJ201933 C.F. Hill (Theaceae) BCRC 2012 FU30031 Ps. chrysanthemicola (J.M. Chrysanthemum sp. RoKi 3708, Taoyuan County, Kuanyin Township, KJ201934 Yen) Deighton (Asteraceae) BCRC Lotus Garden Leisure Farm, 4 August FU30006 2012 Ps. cyatheae C. Nakash. & S. Sphaeropteris lepifera (J. RoKi 3808, Taipei City, Yangmingshan, ca. 450 m, KJ201935 Inaba Sm. ex Hook.) R.M. Tryon BCRC 7 December 2012 (Cyatheaceae) FU30106 Ps. ebulicola (W. Yamam.) Sambucus javanica Blume RoKi 3933 Taoyuan County, Shihmen Dam, 10 KJ201936 Deighton (Adoxaceae) September 2013 Ps. jussiaeae-repentis Ludwigia octovalvis (Jacq.) RoKi 3793, New Taipei City, Yingge, ca. 50 m, 29 KJ201937 (Sawada) Goh & W.H. Hsieh P.H. Raven (Onagraceae) FU30038 October 2012 Ps. lindericola (W. Yamam.) Lindera megaphylla Hemsl. RoKi 3605, Taoyuan County, Shihmen Dam, ca. KJ201931 Goh & W.H. Hsieh (Lauraceae) BCRC 34946 300 m, 1 April 2012 Ps. pteridophytophila Goh & Cyclosorus parasiticus (L.) RoKi 3602 New Taipei City, Yingge, trail to Yingge KJ201938 W.H. Hsieh Farw. (Thelypteridaceae) Rock, 100–250 m, 25 March 2012 Ps. puerariicola (W. Yamam.) Pueraria montana (Lour.) RoKi 3769, Hsinchu County, Smangus Bridge, ca. KJ201939 Deighton Merr. (Fabaceae) BCRC 900 m, 12 October 2012 FU30030 Ps. wedeliae (A.K. Kar & M. Sphagneticola trilobata (L.) RoKi 3588, Yilan County, Beiguan Tidal Park at KJ201940 Mandal) Deighton2 Pruski (Asteraceae) BCRC 34945 the coast, 12 March 2012 Z. dicranopteridis R. Dicranopteris linearis RoKi 3953 Taipei City, Wenshan District, KJ201941 Kirschner (Burm. f.) Underw. var. Maokong, ca. 300 m, 20 October 2013 linearis (Gleicheniaceae) 1 RoKi=R. Kirschner, dried specimen deposited in TNM, BCRC=Bioresource Collection and Research Center, Hsinchu, Taiwan, strain number; 2 Described in detail in Kirschner (2013). 310 • Phytotaxa 176 (1) © 2014 Magnolia Press KIRSCHNER & LIU Materials and methods Specimens of ferns, fern-allies, and seed plants were collected in northern Taiwan during the years 2012–2013. The host plants were identified using the Flora of Taiwan (Huang & Editorial Committee of the Flora of Taiwan 1993–2003). The monographs by Knapp (2011) and Chang et al. (2012) were also consulted. The type specimen of Pseudocercospora pteridophytophila was loaned from the herbarium of the Plant Pathology Department, National Chung Hsing University, Taichung, Taiwan (NCHUPP). Fern leaf spots and dead leaves were examined under a dissecting microscope for the presence of fungi. Fungi were identified using Hsieh & Goh (1990) and original publications of the species. Morphology, cultivation and DNA techniques were applied as described in Kirschner & Okuda (2013). Leaf fragments with mature ascomata of a Mycosphaerella species were fixed beneath the lid of a Petri dish containing 2% malt extract agar. Ascospores shot from the ascomata onto the agar were allowed for germination for 1 day and then investigated microscopically.
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