In Press at Mycologia, preliminary version published on March 23, 2015 as doi:10.3852/14-122

Short title: Systematics of the Cosmospora viliuscula Systematics of the Cosmospora viliuscula species complex Cesar S. Herrera1 Department of Plant Science and Landscape Architecture, University of Maryland, 2112 Plant Sciences Building, College Park, Maryland 20742 Amy Y. Rossman Gary J. Samuels Systematic Mycology and Microbiology Laboratory, USDA, B-010A, 10300 Baltimore Ave, Beltsville, Maryland 20705 Olinto Liparini Pereira Departamento de Fitopatologia, Universidade Federal de Viçosa, 36570-000 Viçosa, Minas Gerais, Brasil Priscila Chaverri Department of Plant Science and Landscape Architecture, University of Maryland, 2112 Plant Sciences Building, College Park, Maryland 20742 and Escuela de Biología, Universidad de Costa Rica, Apartado 11501-2060, San 15 Pedro, San José, Costa Rica Abstract: The genus Cosmospora includes nectrioid fungi that grow on polypores and xylariaceous fungi. The collections growing on xylariaceous fungi have been identified recently as Cosmospora viliuscula. In this paper the phylogeny and taxonomy of C. viliuscula are investigated. A phylogeny was generated with maximum likelihood and Bayesian inference methods applied to a three-partition dataset (ITS, 28S, MCM7-RPB1-TUB2). Based on these results, we demonstrate that Cosmospora viliuscula represents a diverse species complex comprising more than 10 species. Seven new species are described, including three single-strain lineages, and the sexual states of C. arxii and C. khandalensis are described for the first time. The

Copyright 2015 by The Mycological Society of America.

sexual states of these fungi tend to have a high degree of morphological homoplasy, making it difficult to differentiate among them based on morphological characters alone. However, the apparent host specificity of species in this complex aide in the diagnosis of these fungi. In addition, the RPB1 marker provides sufficient resolution to distinguish these fungi. Key words: Ascomycota, fungal systematics, Hypocreales, mycoparasite, Nectria INTRODUCTION Cosmospora Rabenh. sensu Rossman (Nectriaceae, Hypocreales, Sordariomycetes, Ascomycota) included nectrioid species with small, reddish, non- or weakly stromatic perithecioid ascomata that are KOH+, smooth and thin-walled and collapse laterally when dry (Rossman et al. 1999). Asexual states of Cosmospora encompassed Chaetopsina Rambelli, Stilbella Lindau, Volutella Fr. and asexual states described as acremonium- and fusarium-like (Rossman et al. 1999). Recently Cosmospora was restricted to species growing on polypores (Hymenochaetaceae, Fomitopsidaceae, Polyporaceae; Basidiomycota) and xylariaceous fungi (Xylariaceae, Xylariales, Sordariomycetes, Ascomycota) and having acremonium-like asexual states. To accommodate cosmospora-like fungi excluded from Cosmospora sensu stricto, these genera were erected or revived: Chaetopsina, Cylindrocladiella Boesew., Fusicolla Bonord., Macroconia (Wollenw.) Gräfenhan et al., Microcera Desm., Pseudocosmospora C. Herrera & P. Chaverri, Stylonectria Höhn. and Volutella (Schoch et al. 2000; Luo and Zhang 2010, 2012; Gräfenhan et al. 2011; Herrera et al. 2013a). Collections growing on stromata of Xylariaceae that produce green colonies in vitro have been identified as Cosmospora vilior (Starbäck) Rossman & Samuels (see Samuels et al. 1991). However it was recognized early on that the type specimen of this species did not occur on a xylariaceous fungus (Samuels et al. 1990), and for this reason C. vilior was transferred to the newly described genus Pseudocosmospora C. Herrera & P. Chaverri, which includes species that

occur mainly on Eutypa Tul. & C. Tul. and Eutypella (Nitschke) Sacc. (Diatrypaceae, Xylariales; Herrera et al. 2013a). Cosmospora occurring on xylariaceous fungi were referred to as C. viliuscula (Samuels, Yoshim. Doi & Rogerson) Rossman & Samuels because its type specimen is on a xylariaceous fungus (Herrera et al. 2013a). In the present study we demonstrate that C. viliuscula, thought previously to be a single species, represents a species complex. We recognize seven new species in this complex based on multilocus phylogenetic analyses and host specificity. Our results demonstrate the high degree of homoplasy within the Hypocreales and the need to apply cultural and molecular methods for defining species. MATERIALS AND METHODS Herbarium specimens and cultures.—Fresh specimens were collected in Argentina, Brazil, Costa Rica, Peru and USA. Pure cultures were obtained by isolating single ascospores from the freshly collected samples on cornmeal dextrose agar (CMD; Difco™ cornmeal agar + 2% w/v dextrose [Fisher Scientific Co., Fair Lawn, New Jersey] supplemented with 1% [v/v] penicillin-streptomycin-neomycin antibiotic solution [Sigma-Aldrich St Louis, Missouri]). Dried specimens are deposited at the U.S. National Fungus Collections (BPI), Beltsville, Maryland. Cultures derived from fresh specimens were deposited at the CBS-KNAW Fungal Biodiversity Centre (CBS), Utrecht, the Netherlands. Accession numbers are provided (SUPPLEMENTARY TABLE I). Specimens were borrowed from the U.S. National Fungus Collections (BPI); Royal Botanic Gardens Kew (K); William and Lynda Steere Herbarium, New York Botanical Garden (NY); New Zealand National Fungal Herbarium (PDD); and Herbarium of the Botany Department, Swedish Museum of National History (S). Cultures corresponding to these specimens were obtained from the culture collection at USDA, ARS, Systematic Mycology and Microbiology Laboratory (personal isolates of Amy Rossman [A.R.], Clark T. Rogerson [C.T.R.] and Gary J. Samuels [G.J.S.]) and CABI Bioscience Fungus Collection, Egham, Surrey, UK (formerly International Mycological Institute, IMI). Additional cultures were obtained from CBS; Keith A. Seifert (K.A.S.), Agriculture and Agri-Food Canada, Ottawa, Ontario, Canada, and the Ministry of Agriculture, Forestry and Fisheries (MAFF), Tsukuba, Japan (see SUPPLEMENTARY TABLE I). SUPPLEMENTARY Non-authentic material examined is provided (APPENDIX I.

Morphological characterization.—Features of the ascomata (e.g. color, shape, size, ornamentation, habit) were characterized with a stereoscope (Olympus SZX12 Olympus, Tokyo, Japan). Characteristics of the asci and ascospores were obtained by rehydrating the ascomata in 3% KOH, isolating the centrum on a glass slide and observing these structures with a compound microscope (Olympus BX50; Olympus, Tokyo, Japan). Colony morphology was characterized on CMD and Difco™ potato dextrose agar (PDA). Each strain was grown two independent times with two duplicates (n = 4) on each medium at 25 C in an incubator with alternating periods of fluorescent light and darkness (12h/12h). Colony growth rates were measured weekly for 2 wk. Colony color is described with the terminology of Rayner (1970). Asexual states were characterized from cultures grown on synthetic nutrient-poor agar (SNA; Nirenberg 1976) following the system of Hirooka et al. (2012). Measurements of continuous characters were made with Scion Image software beta 4.0.2 (Scion Corp., Frederick, Maryland) and summarized by descriptive statistics (e.g. minimum, maximum, mean and standard deviation). Because the data were not normally distributed, the nonparametric Kruskal–Wallis test (stats package in R [R Core Team 2013]) was used to examine whether there were significant morphological differences among species followed by multiple pairwise comparisons using the nparcomp package (type = Tukey). DNA extraction, PCR, and sequencing.—DNA extraction was performed as described in Hirooka et al. (2010). For P.C. 1285 and P.C. 1306 (personal collections of Priscila Chaverri), DNA was amplified directly from the centra of a few ascomata, as described in Herrera et al. (2013b), because living cultures could not be obtained. Six partial loci comprising two ribosomal DNA regions (nuc rDNA region encompassing the internal transcribed spacers 1 and 2, along with the 5.8S rDNA [ITS] and nuc 28S rDNA D1-D2 domains [28S]) and four protein-coding regions (the minichromosome maintenance complex component 7 gene (MCM7); the RNA polymerase II largest subunit gene (RPB1); the translation elongation factor 1-α gene (TEF1); and the β-Tubulin gene (TUB2) were amplified and sequenced. The primers used to amplify these gene regions are listed (TABLE I). PCR mixtures were prepared as described in Herrera et al. (2013a). PCR amplifications were carried out in an Eppendorf Mastercycler thermo-cycler (Eppendorf, Westbury, New York) under the specified cycle conditions (TABLE I). PCR products were cleaned and sequenced as described in Herrera et al. (2013a). Sequences were assembled and edited with Sequencher 4.9 (Gene Codes Corp., Madison, Wisconsin). Sequences were deposited in GenBank (accession numbers are listed in SUPPLEMENTARY TABLE I).

Phylogenetic analyses.—Multiple sequence alignment for each locus was performed via the MAFFT 6 web service (mafft.cbrc.jp/alignment/server/; Katoh et al. 2002, 2013) with the E-INS-i alignment strategy and the 1PAM /κ = 2 scoring matrix for nucleotide sequences. Alignments were manually edited in Mesquite 2.75 (Maddison and Maddison 2011). Alignments were deposited in TreeBASE under accession number 16192. PartitionFinder (Lanfear et al. 2012) was used with default settings to find the best-fit partitioning scheme among the loci. CONCATEPILLAR 1.4 (Leigh et al. 2008) was used to determine which partitions could be concatenated and analyzed to estimate a phylogeny following Herrera et al. (2013a) except that the α-level was corrected to 0.025. Nucleotide substitution models were inferred for each partition using JModeltest 2.1.2 (Guindon and Gascuel 2003, Darriba et al. 2012) with the settings described in Herrera et al. (2013a). The phylogeny for Cosmospora was estimated with maximum likelihood (ML) and Bayesian inference (BI) following Herrera et al. (2013a). Maximum likelihood analyses were performed with GARLI 2.0 (genetic algorithm for rapid likelihood Inference; Zwickl 2006) via the GARLI web service (www.molecularevolution.org; Cummings and Huskamp 2005; Bazinet and Cummings 2008, 2011). Maximum likelihood bootstrap proportions (ML BP) are reported. Bayesian analyses were performed in MrBayes 3.2.2 (Ronquist et al. 2012). Bayesian inference posterior probabilities (BI PP) are reported. Tracer 1.5 (Rambaut and Drummond 2007) was used to confirm whether the negative log likelihoods had reached convergence. Species Recognition.—Species were recognized in Cosmospora based on the modified criterion used by Pringle et al. (2005) of the genealogical concordance phylogenetic species recognition (GCPSR; Taylor et al. 2000) under which a putative species is recognized if a clade is well supported in the majority of single gene trees. Single gene trees were generated with MrBayes 3.2.2 as described above, except that only 5 000 000 Markov chain Monte Carlo generations were run. Morphological and ecological species recognition were used to support the species inferences made when applying GCPSR. Thus a clade, or putative species, may be associated with unique morphological and/or ecological traits that set it apart from other closely related species.

RESULTS Phylogenetic analyses.—PartitionFinder determined four partitions: ITS, 28S, MCM7+RPB1+TUB2 and tef1. The null hypothesis that these partitions were congruent was tested in CONCATEPILLAR, which rejected this null hypothesis (P < 0.025). These loci were

concatenated because a separate analysis failed to reject the null hypothesis that the partitions ITS, 28S and MCM7+RPB1+TUB2 were congruent (P = 0.043). The resulting matrix comprised 3323 characters; 680 of these were parsimony informative, 255 characters were parsimony uninformative and 1997 characters were invariable. Tef1 was analyzed separately (SUPPLEMENTARY FIG. 5). Individual loci are characterized (TABLE I). The topologies of the phylogenetic trees generated with ML (FIG. 1; −lnL 18351.215) and BI (−lnL 18094.710) were incongruent and differed primarily in relationships within Cosmospora (node A in FIG. 1; 74% ML BP; 100% BI PP). Support values at most deep nodes were low (< 70% ML BP; < 90% BI PP). Well-supported groups (> 70% ML BP; > 90% BI PP) in both phylogenies included C. coccinea, C. viridescens and a large subclade (node B) encompassing the C. viliuscula species complex. Cosmospora coccinea and C. fomiticola, a single-strain lineage recognized below, occur on polypores. Cosmospora viridescens sensu stricto occurs on ascomycetous and other substrates. All lineages within the C. viliuscula species complex corresponding to recognized species received high ML BP and BI PP support (FIG. 1). Ten of these lineages, each distinguished by specific fungal hosts belonging to the Xylariaceae, were recognized as species. The Bayesian phylogeny placed C. lavitskiae within the C. viliuscula species complex, but the fungal host of this species has not been identified. The taxonomic identity of another five isolates in paraphyletic sister-group relation to C. scruposae (Cosmospora sp. 3–7) could not be resolved. At least two loci supported the recognized species (SUPPLEMENTARY FIGS 1–6; TABLE II). Morphological studies.—Little morphological variation exists among members of the Cosmospora viliuscula complex, with some exceptions. For example, the ascospores of C. novazelandica (described below) are longer and wider compared to other species. The conidia of

C. arxii (W. Gams) Gräfenhan & Schroers and C. stilbohypoxyli (described below), differ from each other in conidial width and are the longest in the complex. TAXONOMY Cosmospora Rabenh., Hedwigia 2:59. 1862. Generic type: Cosmospora coccinea Rabenh., Fungi europ. exsicc.: No. 459. 1862. Sexual state: Ascomata nonstromatic, superficial, collapsing laterally, scarlet, smooth, less than 300 μm high. Ascomatal walls 20–35 μm wide, of a single region of intertwined hyphae. Asci cylindrical, eight-spored, ascospores uniseriately arranged. Ascospores ellipsoidal, one-septate, equally two-celled, slightly constricted at septum, minutely verrucose, verrucose, or tuberculate, yellow-brown at maturity. Culture and asexual state: Acremonium-like or verticillium-like. Colonies on PDA velvety, slightly floccose, olivaceous. Colonies on CMD dark-green, flat. Habitat: On polypores (e.g. species of Fomes [Fr.] Fr., Fomitopsis P. Karst., Inonotus P. Karst., and Stereum Hill ex Pers.), xylariaceous fungi (e.g. Annulohypoxylon Y.M. Ju, J.D. Rogers & H.M. Hsieh, Hypoxylon Bull., Kretzschmaria Fr., Stilbohypoxylon Henn., and Xylaria Hill ex Schrank); often isolated from soil. Distribution: Cosmopolitan. Descriptions and illustrations: Gräfenhan et al. (2011), Rossman et al. (1999). Notes: The present paper follows the concept of Cosmospora provided by Gräfenhan et al. (2011) who restricted the genus to cosmospora-like fungi that tend to occur on other fungi, particularly on polypores and xylariaceous fungi, and which possess acremonium-like or verticillium-like asexual states. Cosmospora are suspected to be mycoparasitic based on the description of the attack of a Cosmospora on the ascomata of Annulohypoxylon truncatum (Schwein.) Y.M. Ju, J.D. Rogers & H.M. Hsieh (see Tsuneda 1982). We attempted to obtain the

DNA from the associated xylariaceous fungus (for identification purpuses) by amplifying DNA directly from ascomata centra, but most of the generated sequences were those of the associated Cosmospora. KEY TO SPECIES OF COSMOSPORA SENSU STRICTO 1. Colonies on CMD olivaceous or olivaceous black at the center………………………………..2 1. Colonies on CMD citrine, dark green, greenish yellow, honey or white………………………3

2. Colonies on CMD olivaceous at the center, citrine or citrine-green toward the margin, 35–40 mm diam.…………………………………….………………………………………...C. lavitskiae 2. Colonies on CMD olivaceous black at the center, becoming citrine or olivaceous or grayish yellow-green toward the margin, 30–42 mm diam.……………………….…………C. viridescens

3. On Polyporaceae………………………………………………………………………………...4 3. On Xylariaceae (excluding Biscogniauxia Kuntze)…………………………………………….5

4. On Inonotus spp.; ascospores tuberculate………………………………………...….C. coccinea 4. On Fomes fomentarius; ascospores verrucose………………………….…………..C. fomiticola

5. On Hypoxylon fragiforme and H. howeanum………………………..…………………..C. arxii 5. On other xylariaceous fungi………………………………………………….…………………6

6. On Annulohypoxylon spp…………………………………………….………………….………7 6. On other xylariaceous fungi………………………………………………….………………....9

7. On Annulohypoxylon bovei; ascospore mean length 10 μm.....………………..C. novazelandica 7. On Annulohypoxylon spp.; ascospore mean length < 10 μm…………………………………….8 8. On Annulohypoxylon cohaerens and A. multiforme; ascospores 6.5–9.0 μm long (mean = 7.5 μm); Colonies on CMD amber or white…………….………………………….C. annulohypoxyli 8. On an Annulohypoxylon sp.; ascospores 5.5–8.0 μm long (mean = 6.8 μm); colonies on CMD dark-green………………………………………….………………………………C. khandalensis 9. On Kretzschmaria spp…………………………………………………………………………10 9. On other xylariaceous fungi……………………………………….…………………………..14 10. Ascospore mean length > 10 μm; on Kretzschmaria cetrarioides …………………….C. rickii 10. Ascospore mean length < 10 μm…………………………….……………………….…..…...11 11. Conidiophores unbranched or sparingly branched …………………………….…………….12 11. Conidiophores branched. ……………………………………………….……………………13 12. Conidia 3.0–6.0 × 1.0–2.5 μm (mean 7 × 1.8 μm); colonies on CMD dark-green, pale-luteous at the margin (48–63 mm diam); on Hypoxylon cyclopicum Speg. or K. micropus………………...……….………...……….………...……….………...……C. micropedis 12. Conidia 3.5–6.5 × 1.5–3.0 μm (mean 4.7 × 2.2 μm); colonies on CMD dark green or greenish yellow, hyaline at the margin (44–52 mm diam); on K. deusta ....……….….C. ustulinae 13. Conidiophores unbranched or once dichotomously branched; conidia 3.5–6.5 μm long (mean 4.4 μm); colonies on CMD dark green, greenish yellow at the margin; on K. clavus or K. cf. pavimentosa.………………………………………….…………………….……………...C. clavi

13. Conidiophores unbranched or penicillate with primary, secondary, rarely tertiary branches, terminating in phialides; conidia 2.0–6.5 μm long (mean = 4.0 μm); colonies on CMD white, 27– 55 mm diam; on Kretzschmaria cf. deusta…………….………………….……...……C. viliuscula 14. On Stilbohypoxylon quisquiliarum; conidia 4.5–7.5 × 2.0–3.5 (mean 5.9 × 2.8 μm); colonies on CMD 8–11.5 mm diam (mean = 9.5 mm)………………………………..…..C. stilbohypoxyli 14. On Xylaria scruposa; conidia 3.5–6.5 × 1.5–3.0 μm (mean = 4.8 × 2.0 μm); colonies on CMD 33–55 mm diam (mean = 49.5 mm)…………………………………………………..C. scruposae Cosmospora annulohypoxyli C. Herrera & P. Chaverri, sp. nov. FIG. 2. MycoBank MB 808499 Typification: USA. Louisiana: East Baton Rouge Parish, Port Hudson, Port Hudson State Commemorative Area, on Annulohypoxylon cf. cohaerens, on bark of an unidentified tree, 19 Aug 1996, G.J. Samuels, M. Blackwell & M. Camara (holotype BPI 744521). Ex-holotype culture G.J.S. 96-186 = CBS 137823. Etymology: In reference to the host Annulohypoxylon species. Sexual state: Ascomata solitary, sometimes in groups of a few (< five), subglobose with a blunt apex, 207–274 × 181–232 μm (mean = 232.6 × 198.4; SD 21.2, 15.2; n = 10). Asci cylindrical, 50–68 × 4.0–5.5 μm (mean = 59.0 × 5.0; SD 4.9, 0.6; n = 20). Ascospores ellipsoidal, 6.5–9.0 × 3.0–4.0 μm (mean = 7.5 × 3.6; SD 0.5, 0.2; n = 60), minutely verrucose. Culture and asexual state: Colonies on PDA 30–39 mm diam (mean = 35.1; SD 2.8; n = 7), greenish olivaceous, sometimes centrally grayish yellow-green, velvet-like, sometimes slightly floccose at center, radially furrowed, reverse amber. Colonies on CMD 40–46 mm diam (mean = 42.7; SD 2.4; n = 7), flat, amber, or white, reverse concolorous. Asexual state acremonium-like; conidiophores simple, unbranched, or dichotomously branched. Phialides

monophialidic, cylindrical, (26–)36–68 μm long (mean = 50.4; SD 8.5; n = 20), 2.0–2.5 μm wide at base (mean = 2.2; SD 0.3; n = 20), 1.0–1.5 μm wide at apex (mean = 1.1; SD 0.2; n = 20), collarette flared, hyaline. Conidia ovoid to ellipsoidal, 3.0–5.5 × 1.5–2.5 μm (mean = 4.0 × 2.0; SD 0.5, 0.2; n = 60), unicellular, smooth, hyaline. Habitat: On Annulohypoxylon cohaerens (Pers.) Y.M. Ju et al. and A. multiforme (Fr.) Y.M. Ju et al. (Xylariaceae), on bark of unidentified trees. Distribution: United States (Indiana, Louisiana). Notes: This species is known to grow only on Annulohypoxylon cohaerens and A. multiforme. Cosmospora novazelandica and C. khandalensis also occur on Annulohypoxylon, but the former has longer ascospores (mean = 10 μm) and the latter possesses shorter ascospores (mean = 6.8 μm). Cosmospora arxii (W. Gams) Gräfenhan & Schroers, Stud Mycol 68: 95. 2011. FIG. 3. ≡ Acremonium arxii W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart) p 123. 1971.

Sexual state: Ascomata solitary, globose with a blunt apex, 172–226 × 162–209 μm (mean = 206.2 × 186.8; SD 19.4, 18.7; n = 8). Asci cylindrical, 56–68 × 4.5–6 μm (mean = 63.1 × 5.2; SD 3.7, 0.4; n = 17). Ascospores ellipsoidal, 6.5–8.5(–10.0) × 3.0–4.0 μm (mean = 7.6 × 3.5; SD 0.7, 0.3; n = 40), minutely verrucose. Culture and asexual state: Colonies on PDA 16–23 mm diam (mean = 20.1; SD 2.1; n = 8), salmon to saffron, buff to white at margin, with zonate luteous rings, velvet-like, reverse concolorous. Colonies on CMD 23–31 mm diam (mean = 27.1; SD 3.0; n = 8), flat, hyaline, reverse concolorous. Asexual state acremonium-like; conidiophores simple, unbranched or dichotomously branched. Phialides monophialidic, cylindrical, 42–68 μm long (mean = 52.1; SD 7.7; n = 20), 1.5–2.5 μm wide at base (mean = 2.0; SD 0.2; n = 20), 1.0–1.5 μm wide at apex

(mean = 1.2; SD 0.1; n = 20), collarette flared, hyaline. Conidia ellipsoidal, 4.0–8.5 × 1.5–2.5 μm (mean = 5.4 × 2.0; SD 1.0, 0.2; n = 60), unicellular, smooth, hyaline. Habitat: On Hypoxylon fragiforme (Pers.) J. Kickx f. and H. howeanum Peck (Xylariaceae). Distribution: France, Germany, USA (Kentucky, New York). Additional descriptions and illustrations: Gams (1971), Gräfenhan et al. (2011). Notes: Identity of these isolates was confirmed by sequencing an additional locus, RNA polymerase II (RPB2; A.R. 4521: JQ014128; G.J.S. 10-247: JQ014118) and comparing these sequences to the RPB2 sequence of the ex-type culture (CBS 748.69: HQ897725; retrieved from GenBank). CBS 748.69 was 99% similar to G.J.S. 10-247 and 97% similar to A.R. 4521. Unique to this species is its fungal hosts (Hypoxylon fragiforme or H. howeanum) and its salmon-pink colonies on PDA. Cosmospora clavi C. Herrera & P. Chaverri, sp. nov. FIG. 4. MycoBank MB808506 Typification: COSTA RICA. Heredia: Puerto Viejo de Sarapiquí, La Selva Biological Station, Sendero Tres Rios and Camino Experimental Norte, 10°26′7.3″N, 84°00′31.4″W, 600 m, on Kretzschmaria clavus (Fr.) Sacc., 17 Mar 2010, P. Chaverri (P.C. 1167), G.J. Samuels, A.Y. Rossman, C. Salgado & C. Herrera (holotype BPI 892896). Ex-holotype culture G.J.S. 10-112 = CBS 137812. Etymology: Referring to the fungal host of the type specimen, Kretzschmaria clavus. Sexual state: Ascomata solitary, globose with blunt apex to obpyriform, 181–258 × 135– 245μm (mean = 214.3 × 179.1; SD 27.8, 30.5; n = 15). Asci cylindrical, 49–67 × 5–6 μm (mean = 56.3 × 5.2; SD 3.9, 0.4; n = 30). Ascospores ellipsoidal, 6.0–9.0 × 3.0–4.5 μm (mean = 7.5 ×

3.9; SD 0.6, 0.3; n = 90), verrucose. Culture and asexual state: Colonies on PDA (32–)37–53 mm diam (mean = 43.7; SD 6.5; n = 15), greenish-olivaceous, with black droplets forming at center and a sienna pigment diffusing in medium, or olivaceous buff towards center, sulfur yellow toward margin, velvet-like, floccose, reverse umber. Colonies on CMD 50–61 mm diam (mean = 54.4; SD 3.4; n = 16), flat, dark green, becoming greenish yellow at margin, reverse concolorous. Asexual state acremonium-like; conidiophores simple, unbranched, or dichotomously branched, rarely with three phialides. Phialides monophialidic, cylindrical, 21–68 μm long (mean = 47; SD 11.2; n = 49), 1.5–2.5 μm wide at base (mean = 2.1; SD 0.3; n = 49), 1.0–1.5 μm wide at apex (mean = 1.1; SD 0.1; n = 39), collarette flared, hyaline. Conidia rarely ovoid, ellipsoidal to reniform, 3.5–6.5 × 1.5–2.5 μm (mean = 4.4 × 2.1; SD 0.5, 0.2; n = 150), unicellular, smooth, hyaline. Habitat: On Kretzschmaria clavus and K. cf. pavimentosa (Xylariaceae), on bark of unidentified trees. Distribution: Known from Brazil, Costa Rica, Puerto Rico and Venezuela, possibly panNeotropical. Notes: This species grows on Kretzschmaria clavus and on ustulinoid Kretzschmaria species. Cosmospora clavi has longer and wider ascopores compared to C. micropodis and C. ustulinae, and has longer conidia compared to C. viliuscula. Cosmospora coccinea Rabenh., Fungi europ. exsicc.: no. 459. 1862. FIG.5. = Nectria cosmariospora Ces. & de Not., Comm Soc Crittog Ital 1(4): 195. 1863. ≡ Dialonectria cosmariospora (Ces. & De Not.) Cooke, Grevillea 12: 110. 1884. = Verticillium olivaceum W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart) p 123. 1971.

Asexual state: Acremonium-like. Habitat: On Inonotus spp. (Hymenochaetaceae) on Fagus and Alnus.

Distribution: Europe. Additional descriptions and illustrations: Gams (1971; as V. olivaceum), Samuels et al. (1991; as N. cosmariospora), Rossman et. al (1999), Gräfenhan et al. (2011). Notes: Cosmospora coccinea was lectotypified by Rossman et al. (1999) and reported to grow on pores of Inonotus. Its ascospores are tuberculate. Cosmospora cymosa is also known to occur on Inonotus and is presumed to be sister to C. coccinea (Gräfenhan et al. 2011). However the sexual state of C. cymosa is unknown. It is possible that C. cymosa is a morphological variant of C. coccinea. Cosmospora fomiticola C. Herrera & P. Chaverri, sp. nov. FIG. 6. MycoBank MB808500 Typification: NEW ZEALAND. Westland: Mount Aspiring National Park, Haast Pass, on Fomes fomentarius (L.) Fr., 12 Apr 1983, G.J. Samuels, R.E. Beever & R.H. Petersen (holotype PDD 46398). Ex-holotype culture G.J.S. 83-194 = CBS 137813. Etymology: From fomitis (Latin) = tinder and -cola (Latin) = growing on; referring to the name of the host, Fomes fomentarius (the tinder fungus). Sexual state: Ascomata solitary, rarely in groups of a few, globose to ovoid with an acute apex, collapsing laterally, orange, 212–304 × 176–272 μm (mean = 240.4 × 201.3; SD 33.2, 35.4; n = 6). Asci cylindrical, 59–85 × 5–8 μm (mean = 73.9 × 6.0; SD 7.7, 1.1; n = 10). Ascospores ellipsoidal, 7.0–9.5 × 3.0–4.0 μm (mean = 7.9 × 3.6; SD 0.6, 0.3; n = 50), minutely verrucose. Culture and asexual state: Colonies on PDA 24–27.5 mm diam (mean = 25.8; n = 2), white, with a sienna pigment diffusing into medium, velvet-like, slightly floccose, reverse umber. Colonies on CMD 12–17 mm diam (mean = 14.5; n = 2), flat, honey, with white aerial mycelium toward center, with an olivaceous buff pigment diffusing into medium, reverse concolorous.

Asexual state acremonium-like; conidiophores branched with 3–4 phialides. Phialides monophialidic, cylindrical, 19–42 μm long (mean = 35; SD 6.3; n = 10), 1.5–3.0 μm wide at base (mean = 1.9; SD 0.4; n = 10), 1.0–1.5 μm wide at apex (mean = 1.1; SD 0.1; n = 10), collarette flared, hyaline. Conidia ovoid to ellipsoidal, 3.0–5.0 × 1.5–2.5 μm (mean = 3.7 × 2.0; SD 0.5, 0.2; n = 30), unicellular, smooth, hyaline. Habitat: Fomes fomentarius (Polyporaceae). Distribution: New Zealand. Notes: Cosmospora fomiticola is the only species of the genus known to occur on Fomes fomentarius. Apart from host differences, this species differs from C. coccinea in the ornamentation of the ascospores. The ascospores of C. coccinea are tuberculate, while the ascospores of C. fomiticola are verrucose. Cosmospora khandalensis (Thirum. & Sukapure) Gräfenhan & Seifert, Stud Mycol 68: 96. 2011. FIG. 7. ≡ Cephalosporium khandalense Thirum. & Sukapure, Mycologia 58: 359. 1966.

Sexual state: Ascomata solitary, globose with a blunt apex, 163–228 × 143–208 μm (mean = 188.9 × 164.9; SD 20.3, 19.5; n = 11). Asci cylindrical, 44–61 × 4–7 μm (mean = 52.7 × 5.2; SD 4.5, 0.7; n = 15). Ascospores ellipsoidal, 5.5–8.0 × 2.5–4.0 μm (mean = 6.8 × 3.3; SD 0.5, 0.3; n = 80), verrucose. Culture and asexual state: Colonies on PDA (23–)31.5–43 mm diam (mean = 36.5; SD 5.8; n = 12), citrine to citrine green, becoming buff to white toward margin, sometimes with a sienna pigment diffusing in medium, floccose, reverse sienna. Colonies on CMD 40–51 mm diam (mean = 45.6; SD 3.4; n = 11), flat, dark green, pale luteous at margin, with sparsely spread white aerial mycelium, reverse concolorous. Asexual state acremonium-like; conidiophores simple, unbranched or dichotomously branched. Phialides monophialidic, cylindrical, 34–64 μm long

(mean = 50.5; SD 9.2; n = 30), 1.5–2.5 μm wide at base (mean = 2.0; SD 0.2; n = 30), 1.0–1.5 μm wide at apex (mean = 1.0; SD 0.1; n = 30), collarette flared, hyaline. Conidia ovoid to ellipsoidal, 3.0–5.0 × 1.5–2.5 μm (mean = 3.8 × 2.0; SD 0.4, 0.2; n = 90), unicellular, smooth, hyaline. Holotype: INDIA. Maharashtra: Khandala, on decaying stem and stump of Bambusa, Aug 1964, M. J. Thirumalachar, holotype HACC 148 (not seen), isotypes CBS H-15076 (not seen) = K(M) 169348, ex-type cultures ATCC 16091 (not seen) = CBS 356.65 (not seen) = IMI 112790 = MUCL 7974 (not seen). Habitat: Annulohypoxyon sp. (Xylariaceae); asexual state on decaying stem and stump of Bambusa. Distribution: Argentina, Brazil, India, Japan. Additional descriptions and illustrations: Sukapure & Thirumalachar (1966), Hirooka et al. (2008; as Cosmospora triqua), Gräfenhan et al. (2011). Notes: The sexual state of Cosmospora khandalensis is reported for the first time. Isolates derived from single ascospores (A.R. 4798, A.R. 4799, P.C. 1306) formed a well supported clade with the ex-type culture of C. khandalensis (IMI 112790). Compared to other Cosmospora species growing on Annuloxypoxylon species, C. khandalensis has shorter ascospores. Cosmospora lavitskiae (Zhdanova) Gräfenhan, Seifert & Schroers, Stud Mycol 68: 96. 2011. FIG. 8. ≡.Gliomastix lavitskiae Zhdanova, Mikrobiol. Zhurn. 28: 37. 1966.

Typification: UKRAINE. Poltowa region: on plant debris from rhizosphere soil of Zea mays, 15 Jun 1968 (holotype, D.K. Zabolotny Institute of Microbiology and Virology of the National Academy of Sciences of Ukraine, not seen). Ex-type cultures ATCC 18666 (not seen) = CBS 530.68 (not seen) = IMI 133984 = VKM F-1324 (not seen). Sexual state: Unknown.

Culture and asexual state: Colonies on PDA 31–36 mm diam (mean = 34.3; SD 1.6; n = 10), centrally white, becoming sulfur yellow toward margin, or citrine green, with a buff to ochraceous pigment diffusing in medium, floccose, reverse umber. Colonies on CMD 35–40 mm diam (mean = 37.5; SD 1.7; n = 10), flat, olivaceous at center, citrine or citrine-green toward margin, reverse concolorous. Asexual state acremonium-like; conidiophores simple, unbranched, or dichotomously branched, rarely with three phialides. Phialides monophialidic, cylindrical, 31– 50 μm long (mean = 38.3; SD 4.4; n = 30), 2.0–3.0 μm wide at base (mean = 2.5; SD 0.2; n = 30), 1.0–1.5 μm wide at apex (mean = 1.1; SD 0.1; n = 30). Conidia ellipsoidal to reniform, unicellular, smooth, hyaline, 3.5–6.0 × 1.0–3.0 μm (mean = 4.4 × 1.9; SD 0.5, 0.3; n = 90). Habitat: Isolated from soil rhizosphere of Zea mays. Distribution: Canada, Ukraine, USA (Illinois). Descriptions and illustrations: Zhdanova (1966), Gams (1971). Notes: The collection information associated with one isolate of C. lavitskiae (G.J.S. 96251) suggests that the sexual state occurs on unidentified pyrenomycete, but this information could not be confirmed because the specimen is lost. Bayesian analysis placed C. lavitskiae within the C. viliuscula species complex, which suggests that the host could be a xylariaceous fungus. Unlike other Cosmospora species, colonies of C. lavitskiae on CMD are flat, olivaceous at the center, citrine or citrine-green toward the margin. Cosmospora micropedis C. Herrera & P. Chaverri, sp. nov. FIG. 9. MycoBank MB808501 Typification: FRENCH GUIANA. Saül: ca. 20 km SW of Saül (03°60′N, 53°20′W) toward Mont Galbao (03°50′N, 53°20′W), 650 m, on Kretzschmaria micropus (Fr.) Sacc, 22 Jan 1986, G.J. Samuels (3182) & J.R. Boise (holotype NY). Ex-holotype culture G.J.S. 86-108 =

CBS 137814. Etymology: Referring to the fungal host of the type specimen, Kretzschmaria micropus. Sexual state: Ascomata solitary, superficial, subglobose with a blunt apex, 186–227 × 167–233 μm (mean = 208.3 × 196.8; SD 14.7, 23.1; n = 6). Asci cylindrical, 50–61 × 4–7 μm (mean = 55.5 × 5.3; SD 3.5, 0.6; n = 9). Ascospores ellipsoidal, 5.0–8.5 × 3.0–4.5 μm (mean = 6.9 × 3.7; SD 0.6, 0.3; n = 84), verrucose. Culture and asexual state: Colonies on PDA 41–54 mm diam (mean = 47.6; SD 4.3; n = 12), greenish glaucous, or centrally grayish yellow-green, becoming sulfur yellow toward margin, with an amber pigment diffusing into medium, velvet-like, reverse amber. Colonies on CMD 48– 63 mm diam (mean = 57.6; SD 5.2; n = 12), flat, dark green, becoming pale luteous toward margin, with white aerial mycelium sparsely spread, reverse concolorous. Asexual state acremonium-like; conidiophores simple, unbranched. Phialides monophialidic, cylindrical, 22–53 μm long (mean = 41.2; SD 7.9; n = 29), 1.5–2.5 μm wide at base (mean = 2.1; SD 0.2; n = 29), 1.0–1.5 μm wide at apex (mean = 1.1; SD 0.1; n = 29), collarette flared, hyaline. Conidia ovoid to ellipsoidal, 3.0–6.0 × 1.0–2.5 μm (mean = 3.7 × 1.8; SD 0.5, 0.2; n = 90), unicellular, smooth, hyaline Habitat: On Kretzschmaria micropus and Hypoxylon cyclopicum Speg. (Xylariaceae). Distribution: Brazil, Costa Rica, French Guiana and Guadeloupe, possibly panNeotropical. Notes: This Cosmospora species grows on Kretzschmaria micropus and Hypoxylon cyclopicum, which are considered to be morphological variants of the same species (Rogers and Ju 1998). The conidia of C. micropedis are generally narrower compared to those of other Cosmospora growing on Kretzschmaria species.

Cosmospora novazelandica C. Herrera & P. Chaverri, sp. nov. FIG. 10. MycoBank MB808502 Typification: NEW ZEALAND. South Island: Buller, 21 Km S. of Murchison, on Annulohypoxylon bovei (Speg.) Y.M. Ju, J.D. Rogers & H.M. Hsieh, 17 Apr 1983, G.J. Samuels, P.R. Johnson, R.E. Beever & R.H. Petersen (holotype PDD 46401). Ex-holotype culture G.J.S. 83-197 = CBS 124032. Etymology: In reference to the geographical origin of this species, New Zealand. Sexual state: Ascomata solitary, superficial, obpyriform, with cells protruding around apex (appearing roughened), 292– 330 × 211– 330 μm (mean = 312.1 × 289.9; SD 19.1, 54.1; n = 4). Asci cylindrical to clavate, 70–90 × (5–)6–8 μm (mean = 80.5 × 7.0; SD 5.8, 0.8; n = 18). Ascospores ellipsoidal, 8.0–11.5 × 4.0–6.0 μm (mean = 10.0 × 4.8; SD 0.6, 0.4; n = 60), minutely verrucose. Culture and asexual state: Colonies on PDA 16–20 mm diam (mean = 18.3; SD 1.7; n = 4), centrally olivaceous buff, becoming primrose towards margin, zonate, with a citrine pigment slightly diffusing into medium, velvety, radially furrowed, reverse olivaceous. Colonies on CMD 18–19 mm diam (mean = 18.5; SD 0.6; n = 4), flat, citrine, with an citrine pigment diffusing into medium, reverse greyish yellow-green to citrine green. Asexual state acremonium-like; conidiophores simple, unbranched. Phialides monophialidic, cylindrical, 38–51 μm long (mean = 42.7; SD 4.2; n = 10), 2.0–3.0 μm wide at base (mean = 2.2; SD 0.4; n = 10), 1.0 μm wide at apex (mean = 1.1; SD 0.1; n = 10), collarette flared, hyaline. Conidia ovoid to ellipsoidal, 2.5–5.5 × 1.5–2.5 μm (mean = 3.8 × 1.8; SD 0.5, 0.2; n = 50), unicellular, smooth, hyaline. Habitat: On Annulohypoxylon bovei (Xylariaceae) on unidentified bark. Distribution: New Zealand.

Notes: Cosmospora novazelandica is presumably host specific to Annulohypoxylon bovei. This species is unique in the genus with respect to the ornamentation at the apex of the ascomata with the protruding cells giving the apex a roughened appearance. Ascoma ornamentation is generally absent in Cosmospora. In addition, this species has the largest ascospores among species of the Cosmospora viliuscula species complex. Cosmospora rickii (Rehm) Rossman & Samuels, Stud Mycol 42: 124. 1999. ≡ Nectria rickii Rehm, Hedwigia 44: 2. 1905. = Nectria episphaeria var. kretzschmariae Henn., Bot. Jb. 14(4): 364. 1891. ≡ Nectria kretzschmariae (Henn.) Weese, Akad. Wiss. Wien Sitzungsber., Math.-Naturwiss. Kl., Abt. 1, 125: 506. 1916. = Nectria stigme Rehm, Hedwigia 44: 2. 1905.

Typification: BRAZIL. Estado Do Rio Grande Do Sul: Sao Leopoldo, on disintegrated stroma of Kretzschmaria cetrarioides (as K. lichenoides), 1903, S.J. Rick, F10125 (holotype of N. rickii, S). Asexual state: Unknown. Habitat: Fungicolous on Kretzschmaria cetrarioides (Welw. & Curr.) Sacc. Distribution: Brazil, Republic of the Congo. Additional specimens examined: BRAZIL. Estado Do Rio Grande Do Sul: Sao Leopoldo, on Kretzschmaria cetrarioides (as K. lichenoides), 1903, S.J. Rick, F10189 (holotype of N. stigma, S). REPUBLIC OF THE CONGO (as Loango): on Kretzschmaria cetrarioides (as K. pechuelii), 1876, Pechuel-Lösche (holotype of Nectria episphaeria var. kretzschmariae, B, not seen; destroyed by fire in 1943). Notes: Weese (1916) was the last researcher to have examined all three holotype specimens. He concluded that N. episphaeria var. kretzschmariae, N. rickii and N. stigme represented the same species. He raised N. episphaeria var. kretzschmariae to species rank in

1916. The principle of priority applies only to names within the same rank, thus N. rickii or N. stigme, published in the same article in 1905, have equal priority and provide older names at the species rank. Rossman et al. (1999) selected N. rickii to represent this species and transferred the name to Cosmospora. The holotype of N. rickii consists of the mature form (ascospores ellipsoidal, 14.0–18.5 × 6.0–9.0 μm, mean = 16.0 × 7.5 μm, minutely verrucose, yellow-brown), while the holotype of N. stigme represents an immature form of the same species (ascospores ellipsoidal, 7.0–9.5(–13.0) × 4.0–5.0(–7.5) μm, mean = 8.0 × 4.1 μm, minutely verrucose, yellow-brown). All three type specimens are reported to occur on the same host, Kretzschmaria cetrarioides, which supports the idea that they represent the same species. A living culture from a specimen is needed to determine the phylogenetic placement of this species. Cosmospora scruposae C. Herrera & P. Chaverri, sp. nov. FIG. 11. MycoBank MB808503 Typification: FRENCH GUIANA. Cayenne: Montagne de Kaw, Route de l'est, km 50, on Xylaria scruposa (Fr.) Fr., 25 Mar. 1986, G.J. Samuels (4487) & C. Feuillet (holotype NY). Ex-holotype culture G.J.S. 86-331 = CBS 137816. Etymology: In reference to its fungal host, Xylaria scruposa. Sexual state: Ascomata solitary, superficial, subglobose with an acute apex, 177–239 × 159–224 μm (mean = 212.6 × 189.2; SD 17.0, 18.0; n = 18). Asci cylindrical, eight-spored, 45– 71 × 4–6 μm (mean = 56.4 × 4.8; SD 6.4, 0.5; n = 20). Ascospores ellipsoidal, 6.0–9.0 × 3.0–5.0 μm (mean = 7.3 × 3.7; SD 0.6, 0.3; n = 130), verrucose. Culture and asexual state: Colonies on PDA 28–41.5 mm diam (mean = 33.3; SD 3.8; n = 16), grayish yellow-green, with a sienna to umber pigment diffusing into medium, or primrose, with white aerial mycelium densely floccose in center, margin white, velvet-like, reverse sienna

to umber. Colonies on CMD 33–55 mm diam (mean = 49.5; SD 6.6; n = 17), flat, white, sometimes with an amber pigment, reverse concolorous. Asexual state acremonium-like; conidiophores simple, unbranched, or dichotomously branched, rarely terminating in three phialides. Phialides monophialidic, cylindrical, 36–66 μm long (mean = 59.6; SD 9.0; n = 38), 1.5–3.0 μm wide at base (mean = 2.1; SD 0.4; n = 38), 1.0–1.5 μm wide at apex (mean = 1.1; SD 0.2; n = 38), collarette flared, hyaline. Conidia ovoid to ellipsoidal, 3.5–6.5 × 1.5–3.0 μm (mean = 4.8 × 2.0; SD 0.8, 0.3; n = 90), unicellular, smooth, hyaline. Habitat: On Xylaria cf. scruposa (Xylariaceae). Distribution: French Guiana, Guyana, Puerto Rico and Venezuela, possibly pan Neotopical. Notes: Cosmospora species growing on Xylaria tend to have ascomata with an acute apex. Although fresh cultures were apparently dark green as indicated by the dried culture specimens in herbarium packets, subcultured isolates of C. scruposae produced a white colony on CMD, thus the white colony cannot be considered to be a unique character of this species. It suggests that the isolates have degraded in storage such that they no longer produce the pigments. Cosmospora stilbohypoxyli C. Herrera & P. Chaverri, sp. nov. FIG. 12. MycoBank MB808504 Typification: ARGENTINA. Tucuman Province: San Javier, on Stilbohypoxylon quisquiliarum (Mont.) J.D. Rogers & Y.M. Ju, on decorticated wood, 20 Apr 2011, C. Salgado (holotype BPI 892897). Ex-holotype culture A.R. 4783 = CBS 137834. Etymology: In reference to its fungal host, Stilbohypoxylon quisquiliarum. Sexual state: Ascomata solitary, superficial, subglobose with blunt apex, 152–188 × 145– 185 μm (mean = 169 × 158; SD 11.2, 12.8; n = 8). Asci narrowly clavate, 45–61 × 4– 6 μm

(mean = 51.6 × 4.9; SD 4.5, 0.5; n = 10). Ascospores ellipsoidal, 6.5–8.0(–10.0) × 3.0–4.0 μm (mean = 6.9 × 3.5; SD 0.4, 0.2; n = 40), minutely verrucose. Culture and asexual state: Colonies on PDA 5–6 mm diam (mean = 5.3; SD 0.6; n = 4) after 14 d at 25 C, velvet-like, slightly floccose, herbage green, with a slight sulfur yellow pigment diffusing into medium, reverse concolorous. Colonies on CMD 8–11.5 mm diam (mean = 9.5; SD 1.5; n = 4) after 14 d at 25 C, flat, greenish yellow at center, becoming white toward margin, reverse concolorous. Asexual state acremonium-like; conidiophores simple, unbranched. Phialides monophialidic, cylindrical, 34–44 μm long (mean = 40.1; SD 3.2; n = 10), 2.0–3.0 μm wide at base (mean = 2.3; SD 0.3; n = 10), 1.0–1.5 μm wide at apex (mean = 1.3; SD 0.2; n = 10), collarette flared, hyaline. Conidia ovoid to ellipsoidal, 4.5–7.5 × 2.0–3.5 μm (mean = 5.9 × 2.8; SD 0.6, 0.3; n = 30), unicellular, smooth, hyaline. Habitat: Stilbohypoxylon quisquiliarum (Xylariaceae) on unidentified bark. Distribution: Argentina, Guyana, Venezuela, possibly pan Neotopical. Notes: Cosmospora stilbohypoxyli is known only on Stilbohypoxylon quisquiliarum. This species is slow growing on PDA and CMD compared to other species. In additon, C. stilbohypoxyli possesses wider conidia compared to other species. Cosmospora ustulinae (Teng) C. Herrera & P. Chaverri, comb. nov. FIG. 13. MycoBank MB808505 ≡ Nectria ustulinae Teng, Sinensia, Shanghai 4: 275. 1934 (Basionym).

Typificaction: CHINA. Kiangsu: Pao-hua Shan, on Kretzschmaria deusta (as Hypoxylon ustulatum), [date unknown], S.C. Teng No. 2027 (holotype of Nectria ustulinae, BPI 553261). USA. Tennessee: Blount County, Great Smoky Mountains National Park, 25 mi W Gatlinburg, Cades Cove, Gum Swamp, 1800 feet, 35°35′13.7″N, 83°50′19.2″W, on stroma of

Kretzschmaria deusta, 06 Sep 2005, S. Huhndorf (Epitype designated here, BPI 871089). Exepitype culture A.R. 4215 = CBS 137835. Sexual state: Ascomata solitary, in clusters (< 10), rarely densely aggregated, superficial, subglobose with a blunt apex, 185–225 × 146–195 μm (mean = 205.6 × 169.9; SD 15.9, 17.7; n = 6). Asci narrowly clavate, 41–61 × 4–5 μm (mean = 50.3 × 4.8; SD 6.5, 0.4; n = 12). Ascospores ellipsoidal, 6.0–8.5 × 2.5–5.0 μm (mean = 7.0 × 3.6; SD 0.6, 0.4; n = 62), verrucose. Culture and asexual state: Colonies on PDA 35–51 mm diam (mean = 42.9; SD 5.3; n = 14), sulfur yellow or centrally dark herbage green, becoming grayish yellow-green at margin, with a slight greenish yellow or sienna pigment diffusing into medium, velvet-like, sometimes with black droplets forming at center of colony, reverse concolorous. Colonies on CMD 44–52 mm diam (mean = 48.4; SD 2.8; n = 14), flat, dark-green, or centrally greenish yellow, becoming hyaline toward margin, reverse concolorous. Asexual state acremonium-like; conidiophores simple, unbranched, or dichotomously branched. Phialides monophialidic, cylindrical, 35–57 μm long (mean = 45.9; SD 5.8; n = 28), 1.5–3.0 μm wide at base (mean = 2.3; SD 0.3; n = 28), 1.0– 1.5 μm wide at apex (mean = 1.1; SD 0.1; n = 28), collarette flared, hyaline. Conidia ovoid to ellipsoidal, 3.5–6.5 × 1.5–3.0 μm (mean = 4.7 × 2.2; SD 0.7, 0.3; n = 120), unicellular, smooth, hyaline. Habitat: On Kretzschmaria deusta (Hoffm.) P.M.D. Martin (Xylariaceae). Distribution: China, Japan, Portugal, USA (AL, FL, MI, NY, TN). Additional description: Teng (1934; no illustrations). Notes: This species grows on Kretzschmaria deusta in temperate forests. In contrast to the asexual state of C. viliuscula, with highly branched, penicillate conidiophores, the conidiophores of C. ustulinae are simple, unbranched or branch into two phialides. Although the Japanese

collection (BPI 892905) might make a better epitype on the basis of geographical proximity to the type locality, we opted against this because this collection is poor with few immature ascomata remaining. In addition, the ex-epitype isolate is the most phylogenetically similar to the isolate of the Japanese collection. Cosmospora viliuscula (Samuels) Rossman & Samuels, Stud Mycol 42: 126 (1999). FIG. 14. ≡ Nectria viliuscula Samuels, Mem. New York Bot Gard 59: 44. 1990.

Typification: INDONESIA. North Sulawsi: Eastern Dumoga-Bone National Park, Gn. Muajat, Danau Alia, 1400 m, 00°45′N, 124°25′E, on Kretzschmaria cf. deusta (as Hypoxylon cf. deustum), 26 Oct 1985, G.J. Samuels 2385 (Holotype of Nectria viliuscula, BO, not seen. Isotype NY 01013285). AUSTRALIA. Queensland: Lake Barrine, 750 m, 17.0°14.0′43.0″S, 145°38.0′, 21.0″E, on Kretzschmaria cf. deusta (as Ustulina deusta), 20 Feb 2009, P. Chaverri (P.C. 858) & A.Y. Rossman (epitype designated here, BPI 878994). Ex-epitype culture G.J.S. 09-411 = CBS 137821. Sexual state: Ascomata solitary or in clusters (< 10), superficial, subglobose with a blunt apex, 198–255 × 159–236 μm (mean = 224.9 × 236; SD 20.7, 29.4; n = 10). Asci cylindrical, 46– 63 × 4–6(–7) μm (mean = 54.2 × 5.1; SD 4.9, 0.7; n = 12). Ascospores ellipsoidal, 6.0–10.0 × 2.0–5.0 μm (mean = 7.8 × 3.8; SD 0.8, 0.4; n = 54), verrucose. Culture and asexual state: Colonies on PDA 39–56 mm diam (mean = 46.4; SD 6.7; n = 11), centrally sulfur yellow, becoming white toward margin, with white aerial mycelium sparsely or densely spread, or greenish olivaceous, with a sienna pigment diffusing into medium, velvetlike, reverse sienna. Colonies on CMD 27–55 mm diam (mean = 45.8; SD 10.2; n = 12), flat, white, reverse concolorous. Asexual state acremonium-like to penicillate; conidiophores simple, unbranched, or dichotomously branched; forming primary, secondary and rarely tertiary

branches, each terminal branch producing 2–4 phialides. Phialides monophialidic, cylindrical, 28–57 μm long (mean = 43.6; SD 6.3; n = 30), 1.4–2.5 μm wide at base (mean = 1.8; SD 0.3; n = 30), ~ 1.0 μm wide at apex (mean = 1.0; SD 0.1; n = 30), collarette flared, hyaline. Conidia ovoid to ellipsoidal, 2.0–5.0 × 1.5–3.0 μm (mean = 4.0 × 2.1; SD 0.6, 0.3; n = 90), unicellular, smooth, hyaline. Habitat: Fungicolous on Kretzschmaria cf. deusta (Xylariaceae). Distribution: Australia, Costa Rica, Indonesia, New Zealand. Additional descriptions and illustrations: Samuels et al. (1990, 1991), Rossman et al. (1999). Notes: This species grows on Kretzschmaria cf. deusta in tropical forests and is unique in the genus in producing penicillate conidiophores. Cosmospora viridescens (C. Booth) Gräfenhan & Seifert, Stud Mycol 68: 96. 2011. ≡ Nectria viridescens C. Booth, Mycol Pap 73: 89. 1959.

Typification: UK, England: Yorkshire, Sawley Woods, on black pyrenomycete on branches of Salix, 22 Apr 1954, C. Booth (holotype of Nectria viridescens, IMI 56376, not seen. Isotype BPI 553304 = DAOM 83074, not seen). Wales: Llanrwst, Gwydyr Forest, on Ruzenia spermoides, on Betula, May 1958, C. Booth (paratype, K(M) 169349.) Ex-paratype culture IMI 73377a. Habitat: On bone and fungicolous on Ruzenia spermoides (Lasiosphaeriaceae). Distribution: Czech Republic, Denmark and United Kingdom. Descriptions and illustrations: Booth (1959), Gräfenhan et al. (2011). Notes: The isolate IMI 73377a represents the only living culture derived from specimens cited by Booth (1959) in his original description of Nectria viridescens. By definition K(M) 169349 represents a paratype of C. viridescens, and phylogenetic placement of this species in the strict sense was possible with this isolate. It clustered with other isolates previously considered to

be C. viridescens (see Gräfenhan et al. 2011), but the clade may consist of a species complex. The ascospores are ellipsoidal, 7.5–12.5 × 3.5–4.5 μm (mean = 9.4 × 4.3 μm), slightly constricted at septum, minutely verrucose and yellow-brown. Colonies on CMD are flat, olivaceous black at the center, becoming citrine or olivaceous and grayish yellow-green toward the margin; and 30– 42 mm diam. ADDITIONAL ACCEPTED SPECIES NOT TREATED IN THIS PAPER Cosmospora berkeleyana (P. Karst.) Gräfenhan, Seifert & Schroers, Stud Mycol 68: 95. 2011. ≡ Verticillium berkeleyanum P. Karst., Meddeland. Soc Fauna Fl Fenn 18: 64. 1891. ≡1Acremonium berkeleyanum (P. Karst.) W. Gams, Netherlands J Pl Patho 88: 76. 1982.

Asexual state: Acremonium-like. Habitat: On Inonotus radiatus and Stereum hirsutum. Distribution: Canada, Finland, Germany, the Netherlands. Descriptions and illustrations: Karsten (1891), Gams (1971) and Gams & Zaayen (1982). Note: The sexual state of this species is unknown. The identity of the isolate CBS 258.70 reported to be growing on Inonotus radiatus is questionable. Given the host, the isolate could be a morphological variant of Cosmospora coccinea. This isolate has not been sequenced. Cosmospora butyri (J.F.H. Beyma) Gräfenhan, Seifert & Schroers, Stud Mycol 68: 96. 2011. ≡ Tilachlidium butyri J.F.H. Beyma, Zentralbl. Bakteriol., 2 Abt. 99: 388. 1938. ≡2Acremonium butyri (J.F.H. Beyma) W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart) p. 126. 1971.

Asexual state: Acremonium-like. Habitat: Isolated from butter. Distribution: Denmark (only known from the type). Descriptions and illustrations: Beyma (1938) and Gams (1971).

Notes: The sexual state of this species is unknown. In Gräfenhan et al. (2011), C. butyri (CBS 301.38) is sister to the clade comprising C. coccinea and C. cymosa. An isolate with the same accession number fell outside the Hypocreales in Summerbell et al. (2011), which suggested that more than one fungus might be preserved under the same collection number (Gräfenhan et al. 2011). Cosmospora cymosa (W. Gams) Gräfenhan, Seifert & Schroers, Stud. Mycol. 68: 96. 2011. ≡ Acremonium cymosum W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart) p. 131. 1971.

Asexual state: Acremonium-like. Habitat: On Inonotus radiatus. Distribution: Germany. Descriptions and illustrations: Gams (1971). Notes: The sexual state of this species is unknown. The phylogeny in Gräfenhan et al. (2011) includes C. coccinea and C. cymosa as sister taxa. It is possible that C. cymosa is a morphological variant of C. coccinea, given that they are 97% similar based on ITS sequences. However protein-coding loci showed them to be more divergent. Differences are found primarily at third codon position sites of the protein-coding genes. DISCUSSION Cosmospora.—Cosmospora is based on its type, C. coccinea Rabenh., which was lectotypified by Rossman et al. (1999). Its asexual state was described as Verticillium olivaceum W. Gams, but it is unrelated to Verticillium Nees, which is typified by V. dahliae Kleb. (Plectosphaeriales). Morphologically the asexual state of C. coccinea varies from acremonium-like (conidiophores unbranched, single phialides) to verticillium-like (branching into 1–3 phialides). The asexual state is the character that distinguishes species in Cosmospora sensu stricto from other cosmospora-like genera (Gräfenhan et al. 2011), except Pseudocosmospora. Species of

Pseudocosmospora also have acremonium-like asexual states but are unrelated to Cosmospora (Herrera et al. 2013a). All Pseudocosmospora occur on Diatrypaceae (with the exception of P. joca [Samuels] C. Herrera & P. Chaverri, which occurs on Biscogniauxia capnodes [Berk.] Y.M. Ju & J.D. Rogers, a member of the Xylariaceae), whereas Cosmospora grow on Xylariaceae, Hymenochaetaceae and Polyporaceae. Cosmospora sensu stricto is monophyletic and well supported (node labeled A in FIG. 1; 74% ML BP; 100% BI PP). However deep nodes in the Cosmospora phylogeny remain poorly resolved. One possible explanation for this lack of resolution is that Cosmospora could have undergone a fast early radiation. The relatively short inner branches of the Cosmospora phylogeny further support this idea. Similar phylogenetic patterns (i.e. poor support and short branches) have been observed in other organisms that have undergone rapid radiations (see Chen et al. 2003, Fishben and Soltis 2004, Pons et al. 2005, Kelly et al. 2009). Reconstructing deep node relationships in systems undergoing early radiations is difficult because phylogenetic signal is often masked by multiple substitutions at the same site, which increase with time (reviewed in Donoghue and Sanderson 1992). Phylogenetic resolution can be improved by increasing the number of characters (i.e. sampling more loci) and/or by sampling more taxa (reviewed in Hillis and Wiens 2000). Species recognition.—Species were recognized as described in Herrera et al. (2013a), except that we follow the modified GCPSR criterion used by Pringle et al. (2005). All clades (groups with two or more taxa) with high support (> 95% BI PP) in the majority of single gene trees were recognized as species (SUPPLEMENTARY FIGS 1–6). RPB1 is the only locus that supported all the recognized species (TABLE II), and thus it may be considered to the most reliable locus to diagnose Cosmospora species. In most cases we observed a high degree of morphological homoplasy, particularly in the sexual state, which makes it difficult to rely solely on

morphological characters to diagnose Cosmospora species. Host specificity and, in some instances, morphological characters were used as further support for recognizing clades as species. We consider the host to be one of the most useful characters for diagnosing Cosmospora species, particularly members of the Cosmospora viliuscula species complex. Cosmospora strains or specimens clustering together in a clade tend to have the same host species or hosts with similar morphology and several species specific to a particular host species, received strong support in the analyses (eg. C. arxii occurring on H. fragiforme/H. howeanum, C. clavi on K. clavus/ Kretzschmaria sp., C. micropedis on K. micropus/H. cyclopicum, C. scruposae on X. scruposa). Whereas C. viliuscula, producing penicillate conidiophores, grows on K. cf. deusta, C. ustulinae, producing simple conidiophores, occurs on K. deusta. The former host occurs in tropical regions, while the native range of K. deusta is in temperate regions (Rogers and Ju 1998). The host of C. khandalensis could not be identified to species. A more holistic approach that involves the study of both the parasite and host and the identification of host species based on the comparison of DNA sequences will be required to clarify the degree of host specificity of many Cosmospora species. We also recognize three single-strain lineages as species based on the rule of rarity (reviewed in Lim et al. 2012) given the apparent high fidelity of Cosmospora species to their hosts. Cosmospora fomiticola, C. novazelandica, and C. stilbohypoxyli occur on Fomes fomentarius, Annulohypoxylon bovei and Stilbohypoxylon quisquiliarum respectively. With extensive searching in herbaria, we were able to identify additional specimens with the same hosts and similar morphology as observed in these species. However no extant cultures are associated with these specimens, and thus DNA could not be extracted. These species have been poorly sampled, most likely due to lack of collectors in their regions of origin. Apart from distinct hosts, these species bear morphological differences that set them apart from other

recognized species. For example, cells that protrude from the apex of the ascomata give C. novazelandica a roughened appearance, and C. stilbohypoxyli grows extremely slowly on PDA. In addition, these species are characterized by relatively long branches on the phylogeny, which suggest that multiple substitutions per site have occurred since their speciation. ACKNOWLEDGMENTS This study was supported by the United States National Science Foundation (NSF) PEET grant DEB-0731510 Monographic studies in the Nectriaceae, Hypocreales: Nectria, Cosmospora and Neonectria’ to P. Chaverri, A.Y. Rossman and G.J. Samuels. OLP acknowledges financial support from FAPEMIG and CNPq. We appreciate the fresh specimens sent to us from Canada by Keith A. Seifert. We thank the many people that helped us with collecting trips: A. Romero and R. Sanchez in Argentina; C. Mendez in Costa Rica; P. Johnston in New Zealand; and T. Iturriaga in Venezuela. We gratefully acknowledge the assistance of the curators and their staff of the herbaria from which specimens were generously loaned. These herbaria include: U.S. National Fungus Collection (BPI); Royal Botanic Gardens, Kew (K); William and Lynda Steere Herbarium, New York Botanical Garden (NY); the New Zealand Fungal and Plant Disease Collection (PDD); and Herbarium of the Botany Department, Swedish Museum of National History (S).

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LEGENDS FIG. 1. ML phylogeny (best tree; −lnL = 18351.215) of C. viliuscula species complex based on a three partition (ITS, 28S, MCM7-RPB1- TUB2) dataset. Label A indicates the Cosmospora node. Label B indicates the Cosmospora viliuscula species complex. Black bold branches indicate > 70% ML BP and > 90% BI PP. Gray bold branches indicate > 90% BI PP. FIG. 2. Cosmospora annulohypoxyli BPI 737773 (A, E), BPI 744521(holotype; B–D, F), G.J.S 96-186 (ex-type; G–I, K) and G.J.S 95-199 (J). A. Habit. B. Habit. C. Ascomata on natural substrata. D. Ascoma in 3% KOH. E. Asci. F. Ascospore. G. Phialides. H. Conidia. I–J. Cultures after 3 wk at 25 C on PDA. K. Culture after 3 wk at 25 C on CMD. Bars: A = 3 mm, B = 4 mm, C = 200 μm, D = 100 μm, E−H = 10 μm, I−K = 10 mm. FIG. 3. Cosmospora arxii BPI 892899 (A, C, E), BPI 879925 (B, D), G.J.S. 10-247 (G–H) and A.R. 4521 (I–J). A. Habit. B. Ascomata on natural substrata. C. Ascoma in 3% KOH. D. Asci. E. Ascospore. F. Conidia. G. Phialides. H–I. Cultures after 3 wk at 25 C on PDA. J. Culture after 3 wk at 25 C on CMD. Bars: A = 4 mm, B = 200 μm, C = 100 μm, D–G = 10 μm, H–J = 10 mm. FIG. 4. Cosmospora clavi BPI 892896 (holotype; A, C–F), BPI 745249 (B), G.J.S. 10-112, (ex-type; G–J) and G.J.S. 84-290 (K–L). A−B. Habit. C. Ascomata on natural substrata. D. Ascoma in 3% KOH. E. Asci. F. Ascospores. G–H. Phialides. I. Conidia. J–K. Cultures after 3 wk at 25 C on PDA. L. Culture after 3 wk at 25 C on CMD. Bars: A, B = 4 mm; C = 200 μm; D = 100 μm; E–I = 10 μm; J–L = 10 mm. FIG. 5. Cosmospora coccinea BPI 802729 (A–D), A.R. 2741(E–H, J) and A.R. 2743 (I). A. Habit. B. Ascomata on natural substrata. C. Ascoma in 3% KOH. D. Ascospore. E–F. Phialides. G. Conidia. H–I. Cultures after 3 wk at 25 C on PDA. J. Culture after 3 wk at 25 C on CMD. Bars: A = 4 mm, B = 200 μm, C = 100 μm, D–G = 10 μm, H–J = 10 mm. FIG. 6. Cosmospora fomiticola PDD 46398 (holotype; A–D) and G.J.S. 83-194 (ex-type; F–I). A. Habit. B. Ascomata on natural substrata. C. Ascoma in 3% KOH. D. Ascus. E. Ascospore. F. Phialides. G. Conidia. H. Culture after 3 wk at 25 C on PDA. I. Culture after 3 wk at 25 C on CMD. Bars: A = 3 mm, B = 200 μm, C = 100 μm, D–G = 10 μm, H–I = 10 mm. FIG. 7. Cosmospora khandalensis BPI 892901(A–E), A.R. 4798 (F–H), A.R. 4799 (I, K) and IMI 112790 (ex-type; J). A. Habit. B. Ascomata on natural substrata. C. Ascoma in 3% KOH. D. Asci. E. Ascospores. F–G. Phialides. H. Conidia. I–J. Cultures after 3 wk at 25 C on PDA. K. Culture after 3 wk at 25 C on CMD. Bars: A = 5 mm, B = 200

μm, C = 100 μm, D–H = 10 μm, I–K= 10 mm. FIG. 8. Cosmospora lavitskiae IMI 133984 (ex-type; A–D). A. Culture after 3 wk at 25 C on PDA. B. Culture after 3 wk at 25 C on CMD. C. Phialide. D. Conidia. Bars: A, B = 10 mm; C, D = 10 μm. FIG. 9. Cosmospora micropedis NY GJS 3182 (holotype; A–B, E), BPI 892904 (C–D), G.J.S. 10-121 (F), G.J.S. 86108 (ex-type; G, I) and G.J.S. 10-113 (H). A. Habit. B. Ascomata on natural substrata. C. Ascoma in 3% KOH. D. Asci. E. Ascospore. F. Phialides and conidia. G–H. Cultures after 3 wk at 25 C on PDA. I. Culture after 3 wk at 25 C on CMD. Bars: A = 1 mm, B = 200 μm, C = 100 μm, D–F = 10 μm, G–I = 10 mm. FIG. 10. Cosmospora novazelandica PDD 46401 (holotype; A–E) and G.J.S. 83-197 (ex-type; F–I). A. Ascoma on natural substrata. B. Ascoma in 3% KOH. C. Cells protruding around the Ascomatal apex. D. Asci. E. Ascospore. F. Phialides and conidia. G. Culture after 3 wk at 25 C on PDA. H. Culture after 3 wk at 25 C on CMD. Bars: A = 200 μm; B = 100 μm; C–F = 10 μm; G, H = 10 mm. FIG. 11. Cosmospora scruposae BPI 745150 (A, C), NY GJS 4487 (holotype; B), BPI 744778 (D–E), G.J.S. 86331(ex-type; F–H) and G.J.S. 86-315 (I–J). A. Habit. B. Ascomata on natural substrata. C. Ascoma in 3% KOH. D. Asci. E. Ascospores. F. Phialide. G. Conidia. H–I. Cultures after 3 wk at 25 C on PDA. J. Culture after 3 wk at 25 C on CMD. Bars: A = 2 mm, B= 200 μm, C = 100 μm, D–G = 10 μm, H–J = 10 mm. FIG. 12. Cosmospora stilbohypoxyli BPI 892897 (holotype; A–E) and A.R. 4783 (ex-type; F–I). A. Habit. B. Ascomata on natural substrata. C. Ascoma in 3% KOH. D. Asci. E. Ascospores. F. Phialides. G. Conidia. H. Culture after 3 wk at 25 C on PDA. I. Culture after 3 wk at 25 C on CMD. Bars: A = 1 mm; B = 200 μm; C = 100 μm; D–G = 10 μm; H, I = 10 mm. FIG. 13. Cosmospora ustulinae BPI 871089 (epitype; A–E), A.R. 4215 (ex-epitype; F–H) and G.J.S. 92-95 (I−J). A. Habit. B. Ascomata on natural substrata. C. Ascoma in 3% KOH. D. Asci. E. Ascospore. F. Phialides. G. Conidia. H, I. Cultures after 3 wks at 25 C on PDA. J. Culture after 3 wk at 25 C on CMD. Bars: A = 4 mm, B = 200 μm, C = 100 μm, D–G = 10 μm, H–J = 10 mm. FIG. 14. Cosmospora viliuscula BPI 878994 (epitype; A–E), G.J.S. 09-411 (ex-epitype; F–G, I–J) and G.J.S. 73-2 (H). A. Habit. B. Ascomata on natural substrata. C. Ascoma in 3% KOH. D. Ascus. E. Ascospore. F. Phialides. G. Conidia. H−I. Cultures after 3 wk at 25 C on PDA. J. Culture after 3 wk at 25 C on CMD. Bars: A = 4 mm, B = 200 μm, C = 100 μm, D–G = 10 μm, H–J = 10 mm.

FOOTNOTES Submitted 12 May 2014; accepted for publication 2 Feb 2015 1

Corresponding author. E-mail: [email protected]

TABLE I. Genes/loci used in the phylogenetic analyses. Information on the characteristics of individual loci, models of nucleotide substitution, primers and PCR protocols are indicated Locus ITS 28S mcm7 + rpb1 + tub2 tef1 Nucleotide substitution GTR+I+G HKY+I GTR+I+G HKY+I+G models Number of 574 783 1966 351 characters Phylogenetically 28 28 624 201 informative sites Uninformative polymorphic 46 21 188 71 sites Invariable sites 354 723 920 27 LR5, btub-TI, tef1-728, ITS5, mcm7-709for, LROR crpb1a, rpb1c btub-T2 tef1-986 Primers used ITS4 mcm7-1348rev (Vilgalys (Castlebury et (O’Donnell & (Carbone (reference) (White et (Schmitt et al. & Hester al. 2004) Cigelnik & Kohn al. 1990) 2009) 1990) 1997) 1999) PCR protocol: annealing temp and cycles

53 C, 1 min, 40×

56 C, 50 s, 38×

50 C, 2 min, 40×

55 C, 30 s, 35×

66 C, 55 s, 9× 56 C, 55 s, 35×

TABLE II. Support (BI PP) received by each recognized species in each locus and the combined dataset. Posterior probabilities higher than 90% are reported. Species not supported are represented by a minus symbol Species Locus Combined ITS 28S mcm7 rpb1 tef1 tub2 − − 100 100 94 100 100 C. annulohypoxili 100 100 100 100 98 100 100 C. arxii − − 98 100 − − 100 C. clavi C. coccinea 100 100 N/A 100 99 100 100 98 − 100 97 94 100 100 C. khandalensis − − 100 100 − 100 100 C. lavitskiae − − 100 100 − 100 100 C. micropodis − − − 100 − 100 100 C. scruposae − − 99 100 97 100 100 C. ustulinae 100 − 100 100 93 100 100 C. viliuscula 95 − 100 94 − 100 100 C. viridescens

A

B

G.J.S. 83-194 / Fomes fomentarius / New Zealand Cosmospora fomiticola IMI 362240 / Vitis stem / Unknown Cosmospora sp. 1 A.R. 2743 / Inonotus nodulosus / Germany A.R. 2741 / Inonotus nodulosus / Germany C. coccinea CBS 102433 / Tilia dead tree / Czech Republic IMI 73377a / Ruzenia spermoides / Wales C. viridescens A.R. 2783 / Bone / Denmark CBS 102430 / Picea abies dead standing trunk / Czech Republic G.J.S. 96-7 / Kretzchmaria sp. / Puerto Rico Cosmospora cf. clavi G.J.S. 84-290 / Kretzschmaria clavus / Brazil CBS 251.78 / Kretzchmaria sp. / Brazil C. clavi CBS 123941 / Kretzchmaria sp. / Brazil G.J.S. 10-112 / Kretzschmaria cf. clavus / Costa Rica CBS 448.96 / Kretzchmaria sp. / Puerto Rico G.J.S. 01-301 / Xylaria sp. / Thailand Cosmospora sp. 3 A.R. 4783 / Stilbohypoxylon quisquiliarum / Argentina C. stilbohypoxyli K.A.S. 3751 / Xylaria cf. polymorpha / Canada Cosmospora sp. 7 G.J.S. 90-224 / Xylaria scruposa / Venezuela G.J.S. 90-217 / Xylaria scruposa / Venezuela C.T.R. 71-62 / Xylaria scruposa / Jamaica G.J.S. 86-278 / Xylaria scruposa / French Guiana G.J.S. 86-315 / Xylaria scruposa / French Guiana C. scruposae G.J.S. 86-331 / Xylaria scruposa / French Guiana G.J.S. 96-6 / Xylaria scruposa / Puerto Rico G.J.S. 86-320 / Xylaria scruposa / French Guiana G.J.S. 85-200 / Xylaria sp. / Indonesia Cosmospora sp. 6 G.J.S. 95-142 / Xylaria sp. / Uganda Cosmospora sp. 5 IMI 318025 / Xylaria cf. polymorpha / England Cosmospora sp. 4 IMI 389101 / Clethra arborea stump / Portugal G.J.S. 92-95 / Kretzschmaria deusta / USA MAFF 241532 / Kretzschmaria cf. deusta / Japan C. ustulinae A.R. 4215 / Kretzschmaria deusta G.J.S. 73-2 / Kretzschmaria cf. deusta / New Zealand C. viliuscula G.J.S. 09-411 / Kretzschmaria cf. deusta / Australia G.J.S. 10-114 / Kretzschmaria cf. deusta / Costa Rica G.J.S. 86-108 / Kretzschmaria micropus / French Guiana G.J.S. 10-113 / Hypoxylon cyclopicum / Costa Rica C. micropedis G.J.S. 10-121 / Hypoxylon cyclopicum / Costa Rica P.C. 1285 / Hypoxylon cf. cyclopicum / Brazil A.R. 4770 / Annulohypoxylon sp. / Argentina A.R. 4799 / Annulohypoxylon sp. / Argentina IMI 112790 / Bambusa decaying stem and stump / India C. khandalensis P.C. 1306 / Annulohypoxylon sp. / Brazil A.R. 4798 / Annulohypoxylon sp. / Argentina MAFF 241500 / unidentified pyronomycete / Japan G.J.S. 95-199 / Annulohypoxylon cf. multiforme / USA C. annulohypoxyli G.J.S. 96-186 / Annulohypoxylon cf. cohaerens / USA G.J.S. 83-197 / Annulohypoxylon bovei / New Zealand C. novazelandica G.J.S. 82-275 / Annulohypoxylon sp. / New Zealand Cosmospora sp. 2 A.R. 4521 / Hypoxylon cf. howeanum / France C. arxii G.J.S. 10-247 / Hypoxylon fragiforme / USA G.J.S. 96-251 / unidentified black pyrenomycete / USA K.A.S. 1105 / on woodpile / Canada C. lavitskiae IMI 133984 / plant debris from rhizosphere soil of Zea mays / Ukraine G.J.S 10-193 - Dialonectria episphaeria A.R. 4562 - Pseudocosmospora eutypellae A.R. 4547 - Corallomycetella elegans // A.R. 4580 - Microcera larvarum // 0.04