Xylaria species associated with fallen leaves and petioles

Ju, Yu-Ming; Hsieh, Huei-Mei

doi:10.1186/s40529-023-00377-w

Original Article
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Published: 14 July 2023

Xylaria species associated with fallen leaves and petioles

Botanical Studies volume 64, Article number: 19 (2023) Cite this article

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Abstract

Background

Xylaria species growing on fallen leaves and petioles have not been treated systematically. One source of confusion in this group of Xylaria species has stemmed from X. filiformis, which is an ancient name published in 1805 as Sphaeria filiformis and has commonly labeled on specimen packets that contain leaf- and petiole-inhabiting Xylaria species. Here we clarified the identity of X. filiformis and distinguish it from the species that are easily confused with it, notably X. simplicissima, to which most specimens labeled as X. filiformis are referred. Our research also led us to encounter many other leaf- and petiole-inhabiting Xylaria species, prompting a comprehensive study of this group of fungi.

Results

Forty-five foliicolous and caulicolous species of Xylaria were studied, including nine newly described species—X. allima, X. appendiculatoides, X. hispidipes, X. minuscula, X. neblinensis, X. spiculaticlavata, X. vermiformis, X. vittatipiliformis, and X. vittiformis; three unnamed species—X. sp. AR1741, X. sp. GS7461A, and X. sp. GS7461B; X. simplicissima, a name newly combined with Xylaria from Rhizomorpha simplicissima; and X. noduliformis and X. imminuta, which are two new replacement names, respectively, for X. maitlandii var. nuda and X. hypsipoda var. microspora. The 45 taxa can be classified into three groups by stromatal shape and conspicuousness of perithecial mounds on the stromatal surface: (i) the X. filiformis group contains 10 species, (ii) the X. phyllocharis group contains 19 species, and (iii) the X. heloidea group contains 16 species. One of the newly described or unnamed species belongs to the X. filiformis group—X. vermiformis; nine of them belong to the X. phyllocharis group—X. allima, X. appendiculatoides, Xylaria minuscula, X. neblinensis, X. sp. AR1741, X. sp. GS7461B, X. spiculaticlavata, X. vittatipiliformis, and X. vittiformis; and three of them belong to the X. heloidea group—X. hispidipes, X. imminuta, and X. sp. GS7461A.

Conclusion

The 45 species of Xylaria associated with fallen leaves and petioles can be identified by using the dichotomous identification key that we provided herein. It is important to note that most of the studied species are represented by only one or several specimens and many have not been recollected and cultured.

Background

Fallen leaves and petioles represent one of the four general substrate types, with which Xylaria species are associated. Most Xylaria species produce stromata on dead wood and branches, while fewer species are associated with fallen fruits and seeds (Ju et al. 2018), termite nests and soil (Hsieh et al. 2022; Ju and Hsieh 2007; Ju et al. 2022; Rogers et al. 2005; Wangsawat et al. 2021), and fallen leaves and petioles. One issue in identifying Xylaria species inhabiting fallen leaves and petioles stemmed from the name X. filiformis (Albertini & Schwein.) Fr., which has been commonly applied to Xylaria collections from fallen leaves, petioles or even other substrates worldwide. One particular case is the confusion between X. filiformis, an uncommonly collected species, and X. simplicissima (Pers.) Y.-M. Ju & H.-M. Hsieh, a more commonly encountered species but often immature, because of their overlapping distributions in Europe and likely in North America. Specimens containing X. simplicissima were labeled as X. filiformis in most cases. Similar confusion can occur with other species that have filiform or cylindrical stromata with evident perithecial mounds on the surface.

Recent molecular phylogenetic studies (Hsieh et al. 2010; U’Ren et al. 2016) have shown that leaf- and petiole-inhabiting Xylaria species are not necessarily closely related, even though they share the same growth substrate type. These species formed more than one clade among Xylaria species associated with other substrate types.

Xylaria species growing on fallen leaves and petioles have not been treated systematically, posing an obstacle in identifying these fungi. Rogers et al. (1987), while commenting on page 170 on several small leaf-inhabiting Xylaria species collected from rain forests of North Sulawesi, Indonesia, stated that these Xylaria species “are in urgent need of study on a worldwide basis.” Xylaria species that inhabit leaves and petioles are often overlooked because they produce delicate and often minute stromata compared to those growing on other substrate types. In many cases, their stromata are only produced scarcely, being represented by one or several stromata only. It is not uncommon that stromata of different species are growing together even on the same leaf; examples can be found in the isotype specimens of X. phyllophila Ces. (K ex Cooke herb.), where it is mixed with X. minuscula Y.-M. Ju & H.-M. Hsieh, and the isotype of X. phyllocharis Mont. (PC ex Leprieur herb.), where it is mixed with X. aristata Mont. var. aristata and X. spiculaticlavata Y.-M. Ju & H.-M. Hsieh.

In this study, we aimed to resolve the issues revolving around the name X. filiformis, particularly in answering what the species really is and those species that have been confused with it. We went through published Xylaria names from fallen leaves and petioles and managed to study their type and/or authentic materials. Xylaria species growing on fallen leaves and petioles are separated into three groups herein, i.e., the X. filiformis group, the X. phyllocharis group, and the X. heloidea Penz. & Sacc. group, mainly by stromatal shape, compactness of perithecial aggregation, and conspicuousness of perithecial mounds on the stromatal surface. It is important to note that these groups are not natural assemblages and that the grouping is by no means an attempt to reflect close relatedness within each group. Instead, the grouping is simply a convenient way to convey the general morphology of the taxa included within each group.

The X. filiformis group contains X. filiformis and nine other species—X. castilloi San Martín & J. D. Rogers, X. diminuta San Martín & J. D. Rogers, X. duranii San Martín & Vanoye, X. eugeniae San Martín, Vanoye & P. Lavín, X. meliacearum Læssøe, X. noduliformis Y.-M. Ju & H.-M. Hsieh, X. simplicissima, X. vagans Petch, and X. vermiformis Y.-M. Ju, H.-M. Hsieh, which are characterized by filiform stromata and half-exposed to fully exposed perithecial mounds. Xylaria noduliformis, although having half-exposed perithecial mounds, is like species of the X. phyllocharis group in possessing stouter stromata. The X. phyllocharis group contains 19 taxa—X. allima Y.-M. Ju & H.-M. Hsieh, X. appendiculata Ferd. & Winge, X. appendiculatoides Y.-M. Ju & H.-M. Hsieh, X. asperata J. D. Rogers, Callan, Rossman & Samuels, X. foliicola G. Huang & L. Guo, X. kamatii Pande, X. lima Höhn., X. luxurians (Rehm) C. G. Lloyd, X. maitlandii (Dennis) D. Hawksworth, X. minuscula, X. neblinensis Y.-M. Ju & H.-M. Hsieh, X. phyllocharis, X. phyllophila, X. polysporicola H.-X. Ma & X.-Y. Pan, X. sp. AR1741, X. sp. GS7461B, X. spiculaticlavata, X. vittatipiliformis Y.-M. Ju, H.-M. Hsieh & Fournier, and X. vittiformis Y.-M. Ju & H.-M. Hsieh. Their perithecial mounds are unexposed, being inconspicuous in most species but undulate in X. appendiculatoides, X. luxurians, X. minuscula, X. neblinensis, and X. phyllophila. The 16 taxa of the X. heloidea group—X. amphithele San Martín & J. D. Rogers, X. aristata var. aristata, X. aristata var. hirsuta Theiss., X. axifera Mont., X. clusiae K. F. Rodrigues, J. D. Rogers & Samuels, X. delicatula Starb., X. heloidea, X. hispidipes Y.-M. Ju & H.-M. Hsieh, X. hypsipoda Massee, X. imminuta Y.-M. Ju & H.-M. Hsieh, X. memecyli Pande, X. nainitalensis Dargan, X. petchii C. G. Lloyd, X. pisoniae D. Scott, J. D. Rogers & Y.-M. Ju, X. sicula Pass. & Beltr., and X. sp. GS7461A—are characterized by having a capitate or obconical fertile part of stromata. Their perithecia are clumped together in stromata, with X. petchii being peculiar in having perithecia clumped together near the top of a stroma but scattered below. The apex of stromata is fertile in certain taxa but sterile in others.

Methods

Materials of leaf- and petiole-inhabiting Xylaria taxa were studied from various herbaria, including BO, BPI, C, HAST, HBG, K, L, NY, PC, PH, S, UPS, and WSP. Herbarium abbreviations are in accordance with Index Herbariorum (http://sweetgum.nybg.org/science/ih/). Cultures were obtained by placing tissue or ascospores from freshly collected stromata on SME medium (Kenerley and Rogers 1976). Resulting colonies were transferred to 9-cm plastic Petri dishes containing 2% Difco oatmeal agar (OA), from which the culture descriptions were made, and were incubated at 20 °C under 12 h fluorescent light. Asci and ascospores were mounted in water and Melzer’s reagent and examined using differential interference microscopy and bright-field microscopy. Images of stromata, perithecia, asci and ascospores were taken and measured with the software Piximetre 5.10, which is available at http://ach.log.free.fr/Piximetre/.

Methods for obtaining sequences of nuc rDNA internal transcribed spacers (ITS1-5.8 S-ITS2 = ITS) followed Hsieh et al. (2009).

Taxonomy

Accepted taxa

Xylaria allima Y.-M. Ju & H.-M. Hsieh, sp. nov. Fig. 1A–G.

MycoBank MB848538.

Typification PANAMA. Canal Zone, Fort Sherman area, on fallen leaves, 5 Aug 1945, Martin, G. W. 6154, as X. appendiculata Ferd. & Winge by Dennis, R. W. G. (holotype of X. allima K[M] 236751).

Etymology Near X. lima in having a tomentose stromatal surface and a non-cellular appendage on each end of the ascospores.

Stromata cylindrical at fertile part, unbranched, with an acute apex, on a long, hirsute stipe, 8.6–16.5 mm in total length, 2–6 mm long × 0.6–1.1 mm broad at fertile part; surface dark vinaceous brown, with slight perithecial mounds, overlain entirely with brown tomentum, underlain with a thin, soft layer ca. 10 μm thick or less, concolorous with the surface; interior white, homogeneous, soft. Perithecia spherical, 250–350 μm broad. Ostioles coarsely conic-papillate, tilting upwards, 100–120 μm high × 110–130 μm broad at base. Asci not intact, with an apical ring staining blue in Melzer’s iodine reagent, urn-shaped, 3.5–4 μm high × 3.5–4 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid-inequilateral, with narrowly rounded to broadly rounded ends, smooth, (14.5–)15–16.5(–17) × (8–)8.5–9.5(–10) µm (15.7 ± 0.7 × 9.0 ± 0.4 μm, N = 40), with a straight germ slit slightly less than spore-length on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages; epispore smooth.

Notes Xylaria allima features a dark vinaceous brown, tomentose stromatal surface, beneath which a black layer is lacking, and conic-papillate ostioles tilting upwards. These features are also shared by X. lima, which can be readily separated from X. allima by smaller ascospores (10–)10.5–12(–14) × (5–)6–7(–7.5) µm.

Xylaria appendiculata sensu Dennis (1956) is based on the collection Martin 6154 (K[M] 236751), which, however, differs from X. appendiculata that we define herein by the tomentose stromatal surface and larger ascospores. The ascospore size range given by Dennis (1956) is 12–15 × 7–8 μm, somewhat smaller than our measurement. It should be noted that a duplicate of the collection Martin 6154 in BPI is not the same as the current species but is instead X. lima, which has smaller ascospores.

Xylaria amphithele San Martín & J. D. Rogers, Mycotaxon 34: 304. 1989. Figs. 1H–K, 12H.

Stromata conical to globose at fertile part, unbranched, with or without a sterile apex, on a long, glabrous stipe, up to 50 mm in total length, 1.5–2.5 mm long × 1–2.5 mm broad at fertile part; surface blackish, with conspicuous to half-exposed perithecial mounds, lacking an outer layer, underlain with a thin, soft layer ca. 10 μm thick or less, concolorous with the surface; interior white, homogeneous, soft. Perithecia spherical, 300–550 μm broad. Ostioles slightly papillate, ca. 50 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 130–220 μm total length, the spore-bearing part 80–120 μm long × 7–9 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, inverted hat-shaped, 3.5–4.5 μm high × 2–3 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid-inequilateral, with narrowly rounded ends, smooth, (12–)12.5–15.5(–17) × (5–)6–7.5(–8) µm (14.0 ± 1.5 × 6.7 ± 0.8 μm, N = 60), with a straight germ slit nearly spore-length to spore-length on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages, sometimes retaining a cellular appendage within a non-cellular appendage; epispore smooth.

Cultures and anamorph. Colonies reaching the edge of 9-cm Petri dish at 1.5 wk, whitish to pinkish, becoming blackish from center outwards, appressed, azonate, with diffuse margins. Reverse uncolored. Stromata arising copiously from the entire colony surface, cylindrical, tapering upwards, unbranched, up to 2.5 cm long × ca. 0.6 mm diam, becoming black from base upwards, white on the surface of upper part. Anamorph not produced.

Specimens examined FRENCH WEST INDIES. Guadeloupe, ravine Blondeau, on dead leaves, 4 Sep 2005, Lechat, C. L. 5352 (cultured) (HAST 145965), GenBank: ITS = GU300083. MEXICO. Tamaulipas state, Gómez Farias, on dried fallen leaves in median subdeciduous rain forest, Jul 1987, San Martín, F. 207 (isotype of X. amphithele WSP ex Rogers herb.).

Notes Xylaria amphithele is known thus far from Mexico and Guadeloupe, both located in Central America. It is characterized primarily by having the fertile part of stromata conical to globose and the ascospore enclosed within a hyaline sheath conspicuously swelling at both ends. Xylaria amphithele clustered with two other foliicolous species, X. vagans and X. aristata, in the phylogenetic study of Hsieh et al. (2010), where these two species were labeled as X. sp. 6 and X. sicula f. major, respectively.

Xylaria appendiculata Ferd. & Winge, Bot. Tidsskr. 29: 17. 1908. Figs. 1L–P, 2A, B.

≡ Xylosphaera appendiculata (Ferd. & Winge) Dennis, Kew Bull. 13: 102. 1958.

≡ Podosordaria appendiculata (Perd. & Winge) P. M. D. Martin, J. S. African Bot. 36: 131. 1970, nom. inval. ICN Art. 41.1 (Shenzhen Code); J. S. African Bot. 42: 79. 1976.

? = Xylaria phyllocharis Mont. var. hirtella Theiss., Ann. Mycol. 6: 343. 1908.

Stromata short-cylindrical to long-cylindrical at fertile part, unbranched, with a blunt or acuminate apex, on a hirsute stipe or subsessile to astipitate, (1 –)8–20(–28) mm in total length, 1–13 mm long × 0.7–1.5 mm broad at fertile part; surface brown to dark vinaceous brown except for black perithecial mounds, lacking perithecial mounds when young, becoming slightly undulate, finely longitudinally striated at maturity, lacking an outer layer, underlain with a thin, soft layer ca. 10 μm thick or less, concolorous with the surface; interior white, homogeneous, soft. Perithecia spherical, 200–250 μm broad. Ostioles coarsely conic-papillate, tilting upwards, 110–130 μm high × 100–130 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 100–145 μm total length, the spore-bearing part 80–95 μm long × 9–13 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, urn-shaped, 3–4 μm high × 3–3.5 μm broad. Ascospores dark brown, unicellular, ellipsoid-inequilateral, with narrowly rounded ends, smooth, (11.5–)12.5–14(–15) × (6–)6.5–7.5(–9) µm (13.1 ± 0.8 × 7.2 ± 0.5 μm, N = 60), with a straight germ slit nearly spore-length to spore-length on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages; epispore smooth.

Specimens examined FRENCH GUIANA. ca. 10 km SW of Saul, track from Saul leading SW toward Mt. Galbao, elev. ca. 200 m, on dead leaves, 7–9 Jan 1986, Samuels, G. J. & Boise, J. GS2643, as X. appendiculata (HAST 145966 ex NY); ca. 20 km SW of Saul, Mt. Galbao, Upland moist forest with relatively few epiphytes and mosses on trees, alt. 650 m, on dead leaves, 22 Jan 1986, Samuels, G. J. & Boise, J. GS3164, as X. appendiculata (HAST 145967 ex NY). FRENCH WEST INDIES. Guadeloupe, Basse Terre, on fallen leaves of Clusia venosa, Nov 1992, Vivant, J. GUAD60, as X. appendiculata (WSP 74830). GUYANA. Cuyuni-Mazaruni Region, Mazaruni Subregion, Foothills immediately S of Mt. Ayanganna, alt. 550–650 m, on leaves, 26 Feb 1987, Samuels, G. J. et al. GS4843, as X. phyllocharis var. hirtella (BPI 881476 ex NY), immature; Cuyuni-Mazaruni Region, VII. Mazaruni Subregion, VII-2, Headwaters of Kangu R, W branch ca. 4 km NW of eastern peak of Mt. Ayanganna, first talus slope on sandstone, alt. 700 m, on dead leaves, 5–7 Mar 1987, Samuels, G. J., Pipoly, J., Gharbarran, G. & Chin, J. GS5034, as X. appendiculata (HAST 145968); Cuyuni-Mazeruni, Region VII, Mazeruni Subregion VII-2 and Potaro-Siparuni Region No. VIII, Potaro Subregion VIII, eastern side of Mt. Ayanganna, on decaying dicot leaves, 11–12 Mar 1987, Samuels, G. J., Pipoly, J., Gharbarran, G. & Chin, J. GS5123, as X. appendiculata (BPI 883911 ex NY). US VIRGIN ISLANDS. St. Croix, Caledonia Valley, on leaves of Crescentia, 2 Feb 1906, Raunkiaer, C., Plantae ex Ind. Occid. 1780 (holotype of X. appendiculata C). VENEZUELA. Edo. Merida, above Lagunillas, La Trampa, Loma El Burro, in remnant forest, alt. 2100 m. 8°33′0″N 71°27′0″W, on decaying Clusia leaves, 12 Nov 1990, Samuels, G. J., Hein, B., Huhndorf, S. M., Holmquist, O., Iturriaga, T. & Hererra, M. GS7018, as X. appendiculata (BPI 882429); Edo. Miranda, Parque Nacional Guatopo, trail between Agua Blanca and La Cruceta, alt. 500–600 m, on dicot leaves, 27–30 Nov 1990, Samuels, G. J., Hein, B. & Huhndorf, S. M. GS7636, as X. appendiculata (BPI 88201).

Notes Xylaria appendiculata was named and illustrated with emphasis on the conspicuous non-cellular appendages on the ascospore ends (Ferdinandsen and Winge 1908). Unlike most examined specimens where stromata are cylindrical and stipitate, the holotype specimen of X. appendiculata has stunted, sessile to short-stipitate stromata 1–2 mm high × 0.7–1 mm broad. The current species belongs to the X. phyllocharis group and differs from other species in the group with a similar ascospore size range mainly by having a glabrous stromatal surface, lacking an outer stromatal layer, and possessing non-cellular ascospore appendages.

Xylaria appendiculata was likely included within X. phyllocharis in Fournier et al. (2020), who noted a wide range of variations in ascospore morphology. Xylaria appendiculata differs from X. phyllocharis by having more conspicuous ostioles, larger ascospores, and a prominent hyaline sheath swelling at ascospore ends.

BPI 881476 is immature and has whitish perpendicular pegs arising from the young stromatal surface (Fig. 2A, B). These pegs, presumably synnematal remnants, seem to largely wither off during the maturation process and leave little trace of them on mature stromata. The original material of X. phyllocharis var. hirtella Theiss. appears to be missing. Theissen (1908) described the stromata being short-apiculate on top and having stipes covered with dark hairs. The stromatal surface had half-liberated perithecial mounds with hemispherical to conical ostioles. The ascospores were surrounded by a delicate hyaline sheath that hardly becomes appendiculate on the ends. The ascospore size range was not originally given but was later reported as 12–14 × 6.5–8.5 μm (Theissen 1909). Theissen’s descriptions of X. phyllocharis var. hirtella (Theissen 1908, 1909) seem to suggest the current species except for the hyaline sheath not being conspicuously detached at the ascospore ends.

The interpretation of X. appendiculata in (Dennis 1956) based on Martin, G. W. 6154 from Panama (K[M] 236751) is X. allima, which has a tomentose stromatal surface.

Xylaria appendiculatoides Y.-M. Ju & H.-M. Hsieh, sp. nov. Fig.2C–H.

MycoBank MB848539.

Typification BRAZIL. on fallen leaves, Rick, J. 42, as X. filiformis (holotype of X. appendiculatoides BPI 713892 ex Lloyd herb. 11867).

Etymology Referring to its similarity to X. appendiculata, from which it differs in having larger ascospores and protuberant perithecial mounds.

Stromata cylindrical at fertile part, unbranched, with the apex broken off, probably acute, on a glabrous stipe, 34–59 mm in total length, 8–11 mm long × 0.7–1 mm broad at fertile part, up to 35 mm long at stipe; surface blackish brown to black, slightly polished, with conspicuous perithecial mounds and fine longitudinal striations, lacking an outer layer, underlain with a thin, soft layer ca. 10 μm thick or less, concolorous with the surface; interior white, homogeneous, soft. Perithecia spherical, 250–350 μm broad. Ostioles conic-papillate, tilting upwards, 110–130 μm high × 120–150 μm broad at base. Asci not intact, with an apical ring staining deep blue in Melzer’s iodine reagent, urn-shaped, 4–4.5 μm high × 3.5–4 μm broad. Ascospores dark brown, unicellular, ellipsoid-inequilateral, with acute to narrowly to broadly rounded ends, smooth, (14–)15–16(–17) × (6.5–)7.0–7.5(–8) µm (15.3 ± 0.9 × 7.3 ± 0.3 μm, N = 40), with a straight germ slit spore-length on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages; epispore smooth.

Additional specimen examined BRAZIL. on fallen leaves, Rick, J. 21, as X. cf. filiformis (BPI 713890 ex Lloyd herb. 11868).

Notes Xylaria appendiculatoides can easily be confused with X. appendiculata due to their similar stromatal morphology, coarsely conic-papillate ostioles, and prominent secondary appendages on the ascospores. However, they can be separated by the former species having protuberant perithecial mounds and longer ascospores (13.9–)14.4–16.2(–17.1) µm compared to (11.4–)12.3–13.9(–15.1) µm for X. appendiculata. The current species is known only from two specimens identified as X. filiformis in the Lloyd herbarium (BPI).

Xylaria aristata Mont. var. aristata, Ann. Sci. Nat., Bot., sér. IV, 3: 106. 1855. Figs. 2I–O and 12I.

≡ Xylosphaera aristata (Mont.) Dennis, Kew Bull. 13: 102. 1958.

≡ Podosordaria aristata (Mont.) P. M. D. Martin, J. S. African Bot. 36: 91. 1970, nom. inval. ICN Art. 41.1 (Shenzhen Code); J. S. African Bot. 42: 79. 1976.

= Xylaria oocephala Penz. & Sacc., Malpighia 11: 500. 1897.

? = Xylaria setocephala Yates, Philipp. J. Sci. 12: 379. 1917.

? = Xylaria bogoriensis C. G. Lloyd, Mycol. Writings 7: 1309. 1924.

= Xylaria hainanensis Y. F. Zhu & L. Guo, Mycosystema 30: 527. 2011.

= Xylaria sphaerica G. Huang & L. Guo, in G. Huang, R. S. Wang, L. Guo & N. Liu, Mycotaxon 130: 300. 2015.

Stromata capitate at fertile part, unbranched, with a mucronate to acicular apex, on a long, glabrous or hirsute stipe, 12–35 mm in total length, 1.5–2.5 mm broad at fertile part; surface grayish brown, dark brown to blackish, lacking perithecial mounds or with slight perithecial mounds, overlain with a thin grayish pellicle cracked reticulately into plaques 100–200 μm broad but gradually worn off when overmature, underlain with a carbonized layer ca. 50 μm thick; interior white, homogeneous, fragile. Perithecia spherical, 300–400 μm broad. Ostioles slightly papillate, ca. 100 μm broad at base. Asci with eight ascospores arranged in uniseriate to partially biseriate manner, cylindrical, 115–165 μm total length, the spore-bearing part 65–80 μm long × 6–10 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, urn-shaped, 2.5–3.5 μm high × 2.5–3 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid-inequilateral, with narrowly rounded ends frequently bearing a cellular appendage on one end, smooth, (10–)10.5–12.5(–14) × (5.5–)6–7(–7.5) µm (11.5 ± 1.1 × 6.5 ± 0.5 μm, N = 80), with a straight germ slit nearly spore-length to spore-length on the ventral side, lacking a hyaline sheath; epispore smooth.

Cultures and anamorph. Colonies reaching the edge of 9-cm Petri dish in 4 wk, whitish, immediately becoming blackish in patches from center outwards, appressed, azonate, with diffuse margins. Reverse pale pink. Stromata arising copiously from the entire colony surface, cylindrical, tapering upwards, unbranched, up to 2.5 cm long × 0.3–0.5 mm diam, becoming black from base upwards, white on the surface of upper part. Anamorph not produced.

Specimens examined BRAZIL. Pyrenopolis, on dead leaves, 1912, Brockes, F. A., as X. oocephala (S F43698 ex Sydow herb.). FRENCH GUIANA. on fallen leaves, Leprieur, F. R. 1209 (holotype of X. aristata var. aristata [in 2 packets] PC 0096736 & 0096737 ex Montagne herb. 10093 & 10094, respectively); Cayenne City, on dead dicot leaves, Mar 1986, Samuels, G. J. GS4508, as X. cf. aristata (BPI 881467 ex NY). GUYANA. Kopinang Village, Mt. Wokomung, alt. 1540–1570 m, on dead leaves, 12 Jul 1989, Samuels, G. J., Boom, B. M. & Bacchus, G. GS6692, as X. aristata (HAST 145969). INDIA. Kerala State, Kannur District, Andaloorkavu, on petiole of a fallen leaf, 5 Aug 2011, Deepna Latha, K. P. DKP20, as X. aristata (HAST 145970). INDONESIA. Java, Hortus Bogoriensis, in ramulis dejectis, 28 Dec 1896, Penzig, O. (holotype of X. oocephala BO 3310, isotype PAD); Java, Hortus Bogoriensis, on dead leaves, Jan 1950, van Steenis, C. G. G. J., as X. oocephala (L ex Boedijn herb.); Java, Hortus Bogoriensis, on fallen leaves of Palaquium sp., Feb 1921, van Overeem, C. 37a (holotype of X. bogoriensis BPI 713657 ex Lloyd herb. 10316, isotype L ex U 120499), immature; Java, Hortus Bogoriensis, on petioles, 17 Nov 2017, Ju, Y.-M. 1823 (cultured), as X. aristata (HAST 145971), GenBank: ITS = OQ883719; Java, Hortus Bogoriensis, on petioles, Apr 1923, van Overeem, C., as X. oocephala (BO 521). MEXICO. Oaxaca State, Temazcal Town, in median subdeciduous tropical rain forest, on lamina and midribs of fallen leaves of “zapote chico”, 7 Oct 1988, San Martín, F. 1051B, as X. delicatula (WSP ex Rogers herb.); Oaxaca State, Temazcal Town, in median subdeciduous tropical rain forest, on petioles and lamina of fallen leaves, 8 Oct 1988, San Martín, F. 1121, as X. delicatula (WSP ex Rogers herb.); Quintana Roo State, Othón P. Blanco Municipality, Ejido La Unión, in median subdeciduous tropical rain forest, on lamina and veins of fallen leaves, 12 Nov 1988, San Martín, F. 1271, as X. delicatula (WSP ex Rogers herb.); Quintana Roo State, Othón P. Blanco Municipality, Ejido La Unión, in median subdeciduous tropical rain forest, on lamina and veins of fallen leaves, 15 Nov 1988, San Martín, F. 1323, as X. delicatula (WSP ex Rogers herb.); Quintana Roo State, Othón P. Blanco Municipality, Ejido La Unión, km 12 Cafetal-Mahahual Rd. in low subdeciduous tropical rain forest, on petioles and lamina of fallen leaves, 16 Nov 1988, San Martín, F. 1300, 1367, 1427 & 1450, as X. delicatula (WSP ex Rogers herb.). PHILIPPINES. Luzon, Province of Tayabas, Basiad, on fallen leaves of Garcinia, Dec 1916, Yates, H. S. 25647 (holotype of X. setocephala BPI 584093, isotypes BPI 584094 ex MO, BPI 713652 ex Lloyd herb. 10092, K[M]), immature. TAIWAN. Ping-tung Co., Heng-chun, Ken-ting, on dead leaves, 13 Jul 2001, Ju, Y.-M. & Hsieh, H.-M. 90071613, as X. oocephala (HAST 145972), GenBank: ITS = GU300081.

Notes Stromata of X. aristata var. aristata are overlain with a grayish, reticulately cracked pellicle that gradually wears off at maturity. The stromatal apex is usually long and aciculate in its young stage and becomes short and apiculate when mature. Xylaria delicatula is similar in stromatal features but has darker, nearly semicircular to broadly ellipsoid-inequilateral ascospores. The cultures that we obtained produce copious stromata but lack an anamorph, much as reported by San Martín et al. (1997) (as X. delicatula).

The type materials of X. setocephala from the Philippines and X. bogoriensis from Indonesia are immature, but their stromatal features agree with those of X. aristata var. aristata. The protologues of X. hainanensis and X. sphaerica from China indicate that these two names are in synonymy with X. aristata var. aristata.

Xylaria aristata Mont. var. hirsuta Theiss., Ann. Mycol. 6: 344. 1909. Fig. 3A–C.

Typification BRAZIL. on fallen leaves, Theissen, F. (lectotype [designated here, MycoBank Typification No. 10013065] of X. aristata var. hirsuta S F5664 ex Bresadola herb.).

Stromata capitate at fertile part, unbranched, with a rounded top, on a long, hirsute stipe, 10 mm in total length, 1.8 mm high × 1.6 mm broad at fertile part; surface grayish brown dark brown, with inconspicuous perithecial mounds, overlain with a thin grayish pellicle cracked reticulately into plaques 100–200 μm broad, underlain with a carbonized layer ca. 30 μm thick; interior white, homogeneous, fragile. Perithecia spherical, 400–500 μm broad. Ostioles coarsely conic-papillate, ca. 130 μm high × ca. 150 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 140–170 μm total length, the spore-bearing part 100–130 μm long × 7–9 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, urn-shaped, 2.5–3 μm high × 2.5–3 μm broad. Ascospores dark brown, unicellular, ellipsoid-inequilateral, with narrowly rounded to acute ends frequently bearing a cellular appendage on one end, smooth, (13.5–)14–16(–17) × (5.5–)6–7(–7.5) μm (15.2 ± 1.0 × 6.3 ± 0.4 μm, N = 40), with a straight germ slit nearly spore-length to spore-length on the ventral side, lacking a hyaline sheath; epispore smooth.

Notes Xylaria aristata var. hirsuta is known only from its type specimen, which is highly similar to var. aristata except for its longer ascospores. Some collections of var. aristata have an ascospore length range approaching that of var. hirsuta. Further collections may bridge the gap between the two varieties.

Xylaria asperata J. D. Rogers, Callan, Rossman & Samuels, in Rogers, Callan, Rossman & Samuels, Mycotaxon 31: 120. 1988. Fig. 3D–K.

Stromata ellipsoid, cylindrical to cylindric-fusoid at the continuous or discrete fertile parts, unbranched to occasionally branched once, with an acicular apex, on a long, hirsute stipe, 16–27 mm in total length, 1.2–3.8(–8.5) mm high × 0.8–1.4(–2.4) mm broad at fertile part; surface dark brown to blackish, with inconspicuous to conspicuous perithecial mounds, overlain with a dark brown striped layer on fertile part and dark long spikes entirely, underlain with a thin, black layer ca. 10 μm thick; interior white, homogeneous, soft. Perithecia spherical, 200–300 μm broad. Ostioles coarsely conic-papillate, ca. 150 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 120–160 μm total length, the spore-bearing part 80–95 μm long × 10–11 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, cylindrical to urn-shaped, 2.5–3 μm high × 3–3.5 μm broad. Ascospores dark brown, unicellular, ellipsoid, strongly inequilateral, with narrowly rounded ends sometimes pinched on one end, smooth, (10.5–)11–12.5(–14.5) × (7–)7.5–8.5(–9) µm (11.9 ± 0.8 × 7.9 ± 0.5 μm, N = 40), with a straight germ slit spore-length or nearly so on the ventral side, lacking a hyaline sheath; epispore smooth.

Anamorph was reported by Rogers et al. (1988) as synnemata on the surface of the stromata collected from the field.

Specimens examined VENEZUELA. Cerro de la Neblina, along Rio Mawarinuma, just outside Canon Grande, vicinity of Neblina Base Camp, on fallen leaves of Ficus cf. tonduzii, Apr–May 1984, Samuels, G. J. GS1410 (paratype of X. asperata NY); Territorio Federal Amazonas, Cerro de la Neblina, 5.1 km NE Pico Phelps, alt. 1730–1850 m, on dead leaves, 3 Feb 1985, Rossman, A. Y. AR2224 (isotype of X. asperata BPI 584098); Territorio Federal Amazonas, Neblina Base Camp on Rio Mawarinuma, alt. 140 m, on thin leaves, 27 Jan 1985, Rossman, A. Y. AR2394 (paratype of X. asperata BPI 584099).

Notes Xylaria asperata is unique in having dark spikes covering the entire stromatal surface. The dark long spikes appear to be the synnematal remnants persisting to the maturity. The outer stromatal layer is split into fine stripes, which are somewhat obscured by the spikes. The ascal apical rings are slightly broader than high, a feature not commonly found among Xylaria species.

Xylaria axifera Mont., Ann. Sci. Nat., Bot., sér. IV, 3: 107. 1855. Fig. 3L–O.

≡ Xylosphaera axifera (Mont.) Dennis, Kew Bull. 13: 102. 1958.

≡ Podosordaria axifera (Mont.) P. M. D. Martin, J. S. African Bot. 36: 131. 1970, nom. inval. ICN Art. 41.1 (Shenzhen Code); J. S. African Bot. 42: 79. 1976.

Stromata capitate at fertile part, unbranched, with an acicular apex, on a long hirsute stipe, 6–25 mm in total length, 2–2.8 mm high × 2–3 mm broad at fertile part, up to 18 mm long × 0.5–1 mm at stipe; surface dark brown at the fertile part, yellowish brown at the sterile apex and stipe, with inconspicuous to conspicuous perithecial mounds, overlain with a dull reddish brown layer cracked reticulately into plaques 100–300 μm broad, underlain with a carbonized layer 50–80 μm thick; interior white, homogeneous, fragile. Perithecia spherical, 500–800 μm broad. Ostioles slightly papillate, ca. 100 μm broad at base. Asci with eight ascospores arranged in uniseriate to partially biseriate manner, cylindrical, 250–300 μm total length, the spore-bearing part 175–195 μm long × 7–10 μm broad, with an apical ring staining deep blue in Melzer’s iodine reagent, coffin-shaped to urn-shaped with an anchor-like upper rim, 7.5–9 μm high × 4.5–5.5 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid-inequilateral, with broadly rounded ends, sometimes with one or both ends slightly pinched, smooth, (21.5–)22.5–24.5(–26) × (6.5–)7–8(–9) µm (23.6 ± 1.0 × 7.4 ± 0.6 μm, N = 40), with a straight to slightly oblique germ slit spore-length on the dorsal side, surrounded with a hyaline sheath swelling at both ends to form long tubular non-cellular appendages up to 40 μm; epispore smooth.

Cultures and anamorph were reported by Læssøe and Lodge (1994). The anamorph was not produced in culture but on the stromata collected from the field.

Specimens examined CUBA. on petioles, Wright, C. (K[M]); on petioles, Wright, C., Fungi Cubenses Wrightiani 798 (6 parts), as X. axifera (K[M]). DOMINICAN REPUBLIC. Jarabacoa, coffee plantation #24, 650 elev., on petioles of Schefflera, 27 Mar 1993, Lodge, D. J. 2, as X. axifera (K[M] 25655). GUYANA. on petioles, Leprieur, F. R. 1192 (holotype of X. axifera PC 0086061 ex Montagne herb. 10095, isotypes K[M] 130076 ex Berkeley herb., K[M] ex Currey herb., PC ex Leprieur herb.). PERU. Dep. Huanuco, Tingo María, grounds of Universidad Agraria, on petiole of Araliaceae, 6 Jul 1987, Læssøe, T. TL-P-197, as X. axifera (K[M] 20529). PUERTO RICO. Bosque Nacional del Caribe, Bisley, on fallen petioles of Schefflera, 3 Dec 1997, Legon, N. W. PR317, as X. axifera (K[M] 57023); Luquillo Exp. Forest, El Verde Research Area, on petioles of Schefflera morototoni, 15 May 1988, Lodge, D. J. PR605, as X. axifera (K[M] 25564); Luquillo Exp. Forest, El Verde Research Area, on petioles of Schefflera morototoni, 21 Jan 1991, Lodge, D. J. PR614, as X. axifera (K[M] 25563); Luquillo Exp. Forest, El Verde Research Area, on petioles of Schefflera morototoni, Jan 1991, Lodge, D. J. PR682, as X. axifera (K[M] 25576); Luquillo Exp. Forest, El Verde Research Area, on petioles of Schefflera morototoni, 14 Sep 1992, Lodge, D. J. PR860, as X. axifera (K[M] 25575); Luquillo Exp. Forest, El Verde Research Area, on petioles of Schefflera morototoni, 12 Dec 1992, Lodge, D. J. PR957, as X. axifera (K[M] 25561); Sierra de Cayey, Guavate, on petioles of Schefflera morototoni, 8 Dec 1990, Lodge, D. J. et al. PR613, as X. axifera (K[M] 25565); Sierra de Cayey, nr. Jájome Alto, on petioles of Schefflera morototoni, 18 Jun 1988, Lodge, D. J. PR607, as X. axifera (K[M] 25562). TRINIDAD AND TOBAGO. Port of Spain, Emperor Valley, on petioles of Panax, Thaxter, R. 8709, as X. axifera (K[M] ex NY).

Notes Xylaria axifera differs from other leaf- and petiole-inhabiting species by having a reddish brown outer stromatal layer cracked into fine scales and ascospores longer than 20 μm, possessing a straight to slightly oblique germ slit on the dorsal side and a long tubular non-cellular appendages at each end, and a preference for petioles. Læssøe and Lodge (1994) considered X. axifera to be host-specific to Araliaceae.

Xylaria axifera Mont. var. perexigua Penz. & Sacc. [INDONESIA. Java, Hortus Bogoriensis, in fragmentis ligneis et foliis coriaceis dejectis, 21 Dec 1896, Penzig, O. (holotype BO 3317)] has prominent perithecial mounds on the stromatal surface but is immature as already documented in Penzig and Saccardo (1897). The stromata in the holotype packet bear no resemblance to those of X. axifera. In general, they are like those of X. sicula, X. pisoniae, and their likes.

Xylaria castilloi San Martín & J. D. Rogers, in San Martín, Rogers & Lavín, Revta Mex. Micol. 13: 60. 1997.

For descriptions of the teleomorph, see San Martín et al. (1997) where illustrations of stromata, ascus, and ascospores are also provided. Xylaria castilloi is characterized by the following features: stromata cylindrical at fertile part, unbranched, with an obtuse sterile apex, on a hirsute stipe, 15–20 mm in total length × 2–3 mm broad at fertile part, with a blackish surface where perithecial mounds are half-exposed to fully exposed, overlain with stiff hairs and remnants of outer stromatal layer, with a white, fairly hard interior; perithecia spherical, 400–500 μm broad, with an inconspicuous to slightly papillate ostiole; ascospores brown, ellipsoid-inequilateral to crescentic, with narrowly rounded ends, (11.5–)12–14(–15) × (4.5–)5 μm, with a straight germ slit spore-length or nearly so on the ventral side, lacking a hyaline sheath.

Cultures and anamorph were reported by San Martín et al. (1997) with obovoid to ellipsoid conidia (3.5–)4–4.5(–5) × 2–2.5 μm.

Notes Xylaria castilloi is known only from its type material collected from Mexico (San Martín et al. 1997). Among the species with the stromatal surface overlain with hairs or spikes, X. castilloi can easily be separated from X. allima, X. asperata, X. lima, and X. maitlandii mainly by its half-exposed to fully exposed perithecial mounds.

Xylaria clusiae K. F. Rodrigues, J. D. Rogers & Samuels in Samuels & Rogerson, Mem. N. Y. Bot. Gard. 64: 176. 1990.

= Xylaria lindericola H.-X. Ma & X.-Y. Pan, in X.-Y. Pan, Z.-K. Song, Z. Qu, T.-D. Liu & H.-X. Ma, MycoKeys 86: 55. 2022.

See Samuels and Rogerson (1990) for a description and Ju et al. (2018) for notes and illustrations. The ascospores are dark brown to blackish brown, ellipsoid-inequilateral, (12.5–)13–15(–16) × (7.5–)8–9(–10) µm (13.9 ± 0.8 × 8.6 ± 0.5 μm, N = 40), with a spore-length germ slit, lacking a hyaline sheath. Xylaria clusiae grows on fallen leaves and occasionally fallen fruits of Clusia species (Samuels and Rogerson 1990). The protologue of X. lindericola indicates that this name is in synonymy with X. clusiae.

Xylaria delicatula Starb., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 27, 3: 18. 1901. Fig. 4A–H.

Stromata capitate at fertile part, unbranched, with a mucronate to acicular apex, on a long, glabrous stipe, 7–24 mm in total length, 1.1–2.2 mm long × 1.0–1.6 mm broad at fertile part; surface dark brown to blackish, lacking perithecial mounds or with slight perithecial mounds, overlain with a whitish pellicle when young, cracked reticulately into plaques 100–300 μm and gradually worn off at maturity, underlain with a carbonized layer ca. 50 μm thick; interior white, homogeneous, soft. Perithecia spherical, 300–400 μm broad. Ostioles papillate, ca. 100 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 140–170 μm total length, the spore-bearing part 70–80 μm long × 7–8 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, urn-shaped, 3–4.5 μm high × 3–3.5 μm broad. Ascospores dark brown, unicellular, nearly semicircular to broadly ellipsoid-inequilateral, with broadly rounded ends to, less frequently, narrowly rounded ends sometimes bearing a cellular appendage on one end, smooth, (10–)10.5–12.5(–14) × (5.5–)6–7(–7.5) µm (11.3 ± 1.0 × 6.6 ± 0.5 μm, N = 70), with a straight germ slit spore-length or nearly so on the ventral side, lacking a hyaline sheath; epispore smooth.

Specimens examined BRAZIL. Matto Grosso, Cuyabá, on decayed leaves, 22 Mar 1894, Malme, G. A. 577 (holotype of X. delicatula S F43701). FRENCH GUIANA. between SW of Saul, track between Saul and Mt. Galbao, alt. 450–500 m, on decaying leaves, 11 Jan 1986, Samuels, G. J. & Boise, J. GS2775, as X. delicatula (HAST 145973), GenBank: ITS = OQ883720; km 16 on road between Sinnamary and St. Elie, Orstom Research Area “Ecerex”, primary forest, on decaying dicot leaves, 20 Feb–1 Mar 1986, Samuels, G. J. GS4039, as X. aristata (BPI 881417 ex NY). GUYANA. Demerara-Mahaica Region, IV, Mahaica Subregion, IV-1, Linden Highway between Georgetown and Yarowcabra, Yarowcabra Forestry Station, elev. 50–100 m, on decaying leaf, 26, 27 Apr 1987, Samuels, G. J. & Pipoly, J. GS5555, as X. delicatula (BPI 883855); Mt. Wokomung, on decaying leaves, no date, Samuels, G. J. & Pipoly, J. GS6369, as X. aristata (NY).

Notes Xylaria delicatula resembles X. aristata var. aristata and var. hirsuta in stromatal morphology. It differs from these two taxa by frequently having darker ascospores and strongly inequilateral ascospores with broadly rounded ends.

Xylaria diminuta San Martín & J. D. Rogers, in San Martín, Rogers & Lavín, Revta Mex. Micol. 13: 63. 1997.

For a description of the teleomorph, see San Martín et al. (1997) where illustrations of stromata, ascus, and ascospores are also provided. Xylaria diminuta is characterized by the following features: stromata cylindrical at fertile part, unbranched, with an acute apex, on a glabrous stipe, 5–13 mm total length × 0.5–1.5 mm broad, with a blackish surface where the perithecial mounds are half-exposed to fully exposed and frequently clumped together along the axis, with a white, hard interior; perithecia spherical, 200–300 μm broad, with a papillate ostiole; ascospores light brown to brown, ellipsoid-inequilateral, with rounded to narrowly rounded ends, (5.5–)6–7 × 3–3.5(–4) µm, lacking a hyaline sheath, with a straight germ slit nearly spore-length on the ventral side.

Notes Xylaria diminuta belongs to the X. filiformis group and is unique among the foliicolous Xylaria species in having the smallest ascospore size range.

Xylaria duranii San Martín & Vanoye, in San Martín, Lavín & Rogers, Mycotaxon 79: 342. 2001.

? = Xylaria betulicola H.-X. Ma, Lar.N. Vassiljeva & Y. Li, in H.-X. Ma & Y. Li, Sydowia 70: 39. 2018.

For a description of the teleomorph, see San Martín et al. (2001) where illustrations of stromata, ascus, and ascospores are also provided. Xylaria duranii is characterized by the following features: stromata filiform at fertile part, unbranched, with an acute apex, on a villose stipe, 10–20 mm in total length × 1–1.5 mm broad, with a blackish surface where perithecial mounds are half-exposed to fully exposed, with a white, soft interior; perithecia spherical, 300–400 μm broad, with a slightly papillate to papillate-hemispherical ostiole; ascospores light brown to brown, ellipsoid-inequilateral to somewhat crescentic, with narrowly rounded ends sometimes pinched at one end, 12–14.5(–16) × 4–4.5(–5) µm, with a straight to slightly sigmoid germ slit spore-length on the ventral side, lacking a hyaline sheath.

Notes This Mexican species grows on leaves and fruit remains of Quercus polymorpha and is known only from its type material. It belongs to the X. filiformis group and can be separated from the species in this group with similar ascospore size ranges by having a glabrous stromatal fertile part and lacking a hyaline sheath on the ascospores. The protologue of X. betulicola in general suggests its resemblance to X. duranii except for the stromata terminating into a long acicular stromatal apex.

Xylaria eugeniae San Martín, Vanoye & P. Lavín, in San Martín, Rogers & Lavín, Revta Mex. Micol. 13: 64. 1997.

For a description of the teleomorph, see San Martín et al. (1997) where illustrations of stromata, ascus, and ascospores are also provided. Xylaria eugeniae is characterized by the following features: stromata filiform at fertile part, unbranched, with an acute apex, on a glabrous stipe, 15–20 mm in total length × 1–1.2 mm broad, with a blackish surface where perithecial mounds are half-exposed to fully exposed, with a white, soft interior; perithecia spherical, 200–400 μm broad, with a papillate to hemispherical ostiole; ascospores brown to dark brown, ellipsoid-inequilateral, with narrowly rounded ends sometimes slightly pinched, 12–13.5 × 4–5 μm with a straight germ slit spore-length on the ventral side, surrounded with a hyaline sheath not swelling at ends.

Notes Xylaria eugeniae is known only from Mexico and grows on the fallen leaves of Eugenia capuli. It belongs to the X. filiformis group and resembles X. filiformis and X. vagans but differs by the hyaline sheath surrounding the ascospores not forming a non-cellular appendage at each end.

Xylaria filiformis (Albertini & Schwein.) Fr., Summa Veg. Scand. II, p. 382. 1849. Fig. 4I–N.

≡ Sphaeria filiformis Albertini & Schwein, Consp. Fung. (Leipzig), p. 2. 1805; Albertini & Schwein.: Fr., Syst. Mycol. II, p. 329. 1823.

≡ Hypoxylon filiforme (Albertini & Schwein.) Rabenh., Deutsch. Krypt. Flor. I, p. 223. 1844.

≡ Xylosphaera filiformis (Albertini & Schwein.) Dennis, Kew Bull. 13: 103. 1958.

≡ Podosordaria filiformis (Albertini & Schwein.) P. M. D. Martin, J. S. African Bot. 36: 131. 1970, nom. inval. ICN Art. 41.1 (Shenzhen Code); J. S. African Bot. 42: 79. 1976.

Stromata filiform at fertile part, unbranched, with a long acicular apex, on a glabrous stipe, 14–50 mm in total length, 5–8 mm long × ca. 0.7 mm broad at the broadest of fertile part, 6–20 mm long at stipe; surface black, slightly polished, with half-exposed perithecial mounds and fine longitudinal striations, lacking an outer layer, underlain with a thin, soft layer ca. 20 μm thick, concolorous with the surface; interior white, homogeneous, soft. Perithecia depressed-spherical to spherical, 300–400 μm broad × 200–350 μm high. Ostioles papillate to conic-papillate, ca. 70 μm high × 100–130 μm broad at base. Asci not intact, with an apical ring staining blue in Melzer’s iodine reagent, inverted hat-shaped, 2 μm high × 2–2.5 μm broad. Ascospores light brown, unicellular, short fusoid-inequilateral, with narrowly rounded ends frequently pinched on one or both ends, smooth, (9.5–)11.5–13.5(–14.5) × (4–)4.5–5.5(–6) µm (12.5 ± 1.1 × 5.0 ± 0.5 μm, N = 70), with a straight germ slit spore-length on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages; epispore smooth.

Specimens examined FRANCE. on fallen leaves and petioles of Acer platanoides?, 1853–1860, Roberge, M., Desmazières’ Plantes Cryptogames de France sér. II 377, as X. filiformis (K[M] 169694, K[M] 169692). GERMANY. Niesky, on leaves, Schweinitz, L. D. V. (lectotype of Sphaeria filiformis PH ex Schweinitz herb., isolectotypes BPI 799790 ex Schweinitz herb., BPI 799791 ex Schweinitz herb.), immature; Lichterfelde b. Berlin, auf alten Blättern, Aug 1888, Sydow, P., Sydow’s Mycotheca Marchica 2242, as X. filiformis (HBG, S F131054), immature, with developing perithecia. UK. England, Cumbria County, Meathop Wood, on decaying leaves, 28 Jan 1970, Howard, P. J., comm. Frankland, J., as X. filiformis (K[M] 163347). USA. Georgia, Rabun Co., on fallen leaves of Ilex paca, 19 Aug 1932, Miller, J. H. 4804, as X. filiformis (S F131037).

Notes Despite that X. filiformis has been commonly reported worldwide on fallen leaves, it is likely that this species is largely restricted to Europe. Xylaria filiformis as defined herein has uncommonly been collected, and only two of the examined specimens listed above are mature: K(M) 169694 and K(M) 163347, collected from France and the UK, respectively. The former has all the features as described above, while the latter is overmature, with asci and hyaline sheath no longer available. These two specimens formed the basis for X. filiformis in Dennis (1971) despite of his belief that this species is cosmopolitan.

The specimen S F131037 from Georgia, USA has ascospores (9.5–)10.5–11.5(–12.5) × (3.5–)4–5(–5.5) µm, slightly smaller than those from Europe but otherwise fits X. filiformis and is tentatively included here. Rogers and Samuels (1986) reported three species resembling X. filiformis from New Zealand: X. cf. filiformis, Xylaria taxonomic species 1, and Xylaria taxonomic species 2, which have ascospores (12–)13–16(–21) × 5–7(–8) µm, (14.5–)16–20(–22) × (6.5–)7–8.5(–9) µm, and 16–17(–19) × 7–8(–8.5) µm, respectively. These three New Zealand species differ from X. filiformis by having larger ascospores and lacking a hyaline sheath surrounding their ascospores and likely represent undescribed species. Xylaria cf. filiformis that San Martín et al. (1997) reported differs from typical X. filiformis mainly by having larger ascospores (12.5–)13–17.5 × 5.5–6 μm.

We studied many American and European specimens labeled as X. filiformis, most of which are in fact X. simplicissima from herbaceous stems and only occasionally from fallen leaves of various plants. Xylaria simplicissima can easily be separated from X. filiformis by its larger ascospores (15–)16.5–19(–21.5) × (5–)5.5–6.5(–7.5) µm that lack a hyaline sheath. Xylaria vagans highly resembles X. filiformis but differs primarily by its darker, ellipsoid-inequilateral ascospores and its geographic distribution in the tropics and subtropics.

Albertini and Schweinitz (1805) clearly described and illustrated X. filiformis as a fungus emerging from fallen leaves in Niesky, Germany. The color drawings indicate that the stromata are sterile, with one of them bearing immature perithecia arranged in close set, half-buried with the upper conical portion exposed. Schweinitz’s specimen in PH and BPI collected from Niesky also confirms its leaf-inhabiting habit.

Desmazières was probably the first person to record ascospores of X. filiformis, which were given as “0^mm,015” (= 15 μm) long on the label of his exsiccata Plantes Cryptogames de France sér. II 377 issued between 1853 and 1860. He also provided a detailed description for the species on this label and mentioned that the exsiccata contained material collected from leaves of Acer platanoides, Carpinus, Castanea, Fagus, Populus, Quercus, and Salix. It is thus possible that the Desmazières exsiccata is composed of stromata from multiple gatherings. The two duplicates of the Desmazières exsiccata in K, upon which Dennis (1971) based his interpretation of X. filiformis, contain light brown to brown ascospores with a prominent non-cellular appendage on each end. Nonetheless, Nitschke (1867) did not find ascospores in the duplicate of this exsiccata that he studied nor in Rabenhorst’s Fungi Europaei 57 or his own collections and some Kunze specimens of X. filiformis. According to Nitschke (1867), the Kunze specimens are probably the original material from Albertini and Schweinitz and correspond well to their description and illustration. Fuckel (1869) reported that no ascospores were found in his specimens collected from leaves of various plants around Oestrich, Germany. Ascospores of X. filiformis were given as 13–14 × 5–6 μm in Karsten (1879), 13–14 × 5–6 μm in Winter (1887), 12–15 × 5 μm in Saccardo (1882), and 12–15 × 4–5 μm in Traverso (1906).

Xylaria foliicola G. Huang & L. Guo, in G. Huang, L. Guo & N. Liu, Mycotaxon 129: 150. 2014.

For a description of the teleomorph, see Huang et al. (2014) where illustrations of stromata, asci, and ascospores are also provided. Xylaria foliicola is characterized by the following features: stromata cylindrical at fertile part, unbranched or branched occasionally, with a long acicular apex, on a glabrous stipe, 23–35 mm in total length, 7–18 mm long × 1–2 mm broad at fertile part, with a black surface lacking perithecial mounds or with slight perithecial mounds, overlain with a dull grayish brown peeling layer split into narrow or thread-like stripes, with a white interior; perithecia subspherical to ellipsoid, 400–650 μm broad, with a papillate ostiole; ascospores brown, ellipsoid-inequilateral, with narrowly rounded ends, smooth, (8.5–)9–11 × 4–6 μm, with a straight germ slit spore-length on the ventral side, lacking a hyaline sheath.

Notes Xylaria foliicola is known only from its type material collected from Yunnan, China (Huang et al. 2014). Xylaria minuscula is similar to X. foliicola in stromata overlain with a narrowly striped outer peeling layer but differs from it mainly by larger ascospores.

Xylaria heloidea Penz. & Sacc., Malpighia 11: 498. 1897.

See Ju et al. (2018) for a description and illustrations. Xylaria heloidea is not exclusively associated with fallen leaves and does not appear to have a specificity to certain hosts.

Xylaria hispidipes Y.-M. Ju & H.-M. Hsieh, sp. nov. Fig. 5A–D.

MycoBank MB848540.

Typification INDONESIA. North Sulawesi, Eastern Dumoga-Bone National Park, at confluence of Toraut & Tumpah Rivers, Project Wallace Base Camp, on leaves, Sep–Nov 1985, Samuels, G. J. GS1981, as X. hypsipoda (holotype of X. hispidipes NY 01089626, isotype BO 19978).

Etymology Referring to the coarse rigid erect hairs covering the stromatal stipes.

Stromata capitate at fertile part, unbranched, with a mucronate apex, on a long hirsute stipe, 15–40 mm in total length, 1–3.5 mm high × 2–3 mm broad at fertile part, 14–38 mm long at stipe; surface dull grayish brown, lacking perithecial mounds, overlain with a thin pellicle cracked reticulately into plaques 100–200 μm broad, underlain with a carbonized layer 300–400 μm thick; interior white, homogeneous, soft. Perithecia spherical, 300–600 μm broad. Ostioles papillate, ca. 100 μm high × 110–140 μm broad at base. Asci with eight ascospores arranged in uniseriate to partially biseriate manner, cylindrical to clavate, at least 160 μm total length, the spore-bearing part 70–100 μm long × 12–20 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, urn-shaped, 4.5–6 μm high × 3–5 μm broad. Ascospores dark brown, unicellular, ellipsoid-inequilateral, with narrowly rounded ends, smooth, (15–)15.5–17(–18) × (6.5–)7.5–9(–9.5) µm (16.3 ± 0.8 × 8.2 ± 0.6 μm, N = 40), with a straight germ slit spore-length or nearly so on the ventral side, lacking a hyaline sheath; epispore smooth.

Cultures obtained by Rogers et al. (1987) (as X. hypsipoda) with stromata produced on OA but without an observed anamorph.

Notes Xylaria hispidipes is similar to X. hypsipoda in having a capitate stromatal fertile part overlain with a thin pellicle that is cracked reticulately into plaques on a long stipe. In fact, the type material of X. hispidipes was previously identified as X. hypsipoda by Rogers et al. (1987). Xylaria hypsipoda can be separated from the current species by its stromata with a rounded fertile apex and a glabrous stipe, less conspicuous ostioles, and smaller ascospores surrounded with a hyaline sheath that swells at both ends. The asci from the type material are now fragmentary; the measurement is thus adopted from Rogers et al. (1987), where colonies on oatmeal agar were described but no anamorph was produced.

Xylaria hypsipoda Massee, Bull. Misc. Inform. Kew nos. 153–154: 174. 1899. Fig. 5E–I.

Stromata capitate at fertile part, unbranched, with a rounded top, on a long glabrous stipe slightly roughened with scars of fallen conidial pegs, (20–)45–70 mm in total length, 2.6–3.4 mm high × 2.7–4 mm broad at fertile part; surface dull grayish brown, lacking perithecial mounds, overlain with a thin pellicle cracked reticulately into plaques 100–200 μm broad, underlain with a carbonized layer 90–100 μm thick; interior white, homogeneous, soft. Perithecia spherical, 500–600 μm broad. Ostioles minutely papillate, ca. 70 μm broad at base. Asci not intact, with an apical ring staining blue in Melzer’s iodine reagent, urn-shaped, 3–3.5 μm high × 3.5 μm broad. Ascospores dark brown, unicellular, ellipsoid-inequilateral, with narrowly to broadly rounded ends, smooth, (13–)13.5–15(–16.5) × (6–)6.5–7.5(–8) µm (14.1 ± 0.8 × 7.0 ± 0.5 μm, N = 40), with a straight germ slit spore-length on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages; epispore smooth.

Specimens examined SINGAPORE. Bukit Mardi, on fallen leaves, 1897, Ridley, H. N. 34 (holotype of X. hypsipoda K[M] 156834). PHILIPPINES. Luzon, Makiling, on dead sheath of Arenga saccharifera, 27 Sep 1920, Sanchez A. 9570, as X. hypsipoda (BPI 713971 ex Lloyd herb. 10074, BPI 585083 ex Reinking herb.).

Notes Xylaria hypsipoda is unique among foliicolous Xylaria species in stromata having a large stature topped by a capitate fertile part. The collection reported as X. hypsipoda by Rogers et al. (1987) from North Sulawesi, Indonesia is newly described herein as X. hispidipes. Lloyd (1923) reported a collection of X. hypsipoda from the Philippines, which was sent to him by O. A. Reinking. He presented a photograph in Fig. 2363 showing two stromata characterized by a globose head on a long slender stipe. However, he did not find spores in the specimen but quoted the spore size as 12 × 6–7 μm, likely provided by Reinking. The specimen (BPI 713971) from the Lloyd herbarium contains only long slender stipes arising from the sheath of Arenga saccharifera, without any trace of globose heads. A duplicate of the same collection (BPI 585083) no longer contains any Xylaria material.

Xylaria imminuta Y.-M. Ju & H.-M. Hsieh, nom. nov. Fig. 5J, K.

MycoBank MB848541.

≡ Xylaria hypsipoda Massee var. microspora J. D. Rogers, Callan & Samuels, Mycotaxon 29: 162. 1987.

Typification INDONESIA. North Sulawesi, Eastern Dumoga-Bone National Park, at confluence of Toraut & Tumpah Rivers, Project Wallace Base Camp, on leaves, Sep–Nov 1985, Samuels, G. J. GS1954 (holotype of X. hypsipoda var. microspora BO 19982).

Etymology Referring to its miniature stromata.

Stromata capitate at fertile part, unbranched, with a rounded top, on a long hirsute stipe, 60–85 mm long in total length, 0.6–1.2 mm high × 0.6–0.8 mm broad at fertile part; surface blackish brown, lacking perithecial mounds, overlain with a thin pellicle cracked reticulately into plaques 70–120 μm broad, underlain with a thin black layer ca. 20 μm thick. Perithecia not measured. Ostioles minutely papillate, ca. 50 μm broad at base. Asci not measured. Ascospores as in X. hispidipes but smaller, 8–9(–9.5) × 4–4.5(–6.6) µm.

Notes Xylaria imminuta was originally described as X. hypsipoda var. microspora by Rogers et al. (1987) who noted that it is a diminutive form of X. hypsipoda. The collection identified as X. hypsipoda by Rogers et al. (1987) is described herein as a new species X. hispidipes, which differs from X. imminuta mainly by its larger stromata with a minute sterile apex and larger ascospores (15–)15.5–17(–18) × (6.5–)7.5–9(–9.5) µm.

The holotype at BO is now rather scanty, consisting of less than ten tiny stromata, only three of which still bear a fertile part. As such, we did not exploit the specimen further but instead refer to the ascospore morphology described in Rogers et al. (1987).

Xylaria kamatii Pande, Nova Hedwigia 24: 13. 1973.

For a description of the teleomorph, see Pande (1973) where illustrations of stroma, perithecia, ascus, and ascospores are also provided. Xylaria kamatii is characterized by the following features: stromata cylindrical at fertile part, unbranched, with a blunt apex, on a glabrous stipe, 10 mm in total length, 4 mm long × 0.8 mm broad at fertile part, with a black surface where perithecial mounds are inconspicuous, not shrinking upon drying, with a white interior; perithecia spherical to obovoid, 320–480 μm broad × 400–480 μm high, with a slightly papillate ostiole; ascospores brown, ellipsoid-inequilateral, 8–10 × 4–6 μm, with a straight germ slit much less than spore-length, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages.

Notes Xylaria kamatii is known only from its type collection, which was made from fallen leaves of Memecylon umbellatum (Melastomataceae) in India (Pande 1973). We made a request for loaning the type specimen preserved at AMH but did not receive a response. The protologue indicates that X. kamatii possesses a short germ slit on its ascospores, which are enclosed by a hyaline sheath. The ascospores in Fig. 2D of Pande (1973) were depicted as having a non-cellular appendage at each end.

Xylaria lima Höhn., Denkschr. Kaiserl. Akad. Wiss. Wien, Math.-Naturwiss. Kl. 83: 27. 1907. Fig. 6A–F.

= Xylaria mexicana San Martín, J. D. Rogers & P. Lavín, Revta Mex. Micol. 13: 66. 1997.

Stromata cylindrical at fertile part, unbranched, with an acuminate apex, on a hirsute stipe, 11–21 mm in total length, 3.1–7.5 mm long × 0.8–1.1 mm broad at fertile part; surface dark vinaceous brown, with slight perithecial mounds, overlain with a dark brown tomentum, underlain with a thin, soft layer ca. 10 μm thick or less, concolorous with the surface; interior white, homogeneous, soft. Perithecia spherical, 150–250 μm broad. Ostioles conic-papillate, tilting upwards, 90–110 μm high × 90–110 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 105–145 μm total length, the spore-bearing part 75–105 μm long × 9.5–10.5 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, urn-shaped, 3.5–5 μm high × 3.5–4.5 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid-inequilateral, with narrowly rounded ends, smooth, (10–)10.5–12(–14) × (5–)6–7(–7.5) µm (11.3 ± 0.8 × 6.3 ± 0.5 μm, N = 70), with a straight germ slit spore-length or nearly so on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages; epispore smooth.

Anamorph was reported by San Martín et al. (1997) (as X. mexicana) on the surface of immature stromata collected from the field.

Specimens examined BRAZIL. São Paulo, Raiz de Serra, on dead leaf, Jun 1901, von Höhnel, F. 4371 (holotype of X. lima FH ex von Höhnel herb.). FRENCH GUIANA. primary forest at km 16 on road between Sinnamary and St. Elie, Orstom Research Area, “Ecerex”, on decaying dicot leaf, 20 Feb–1 Mar 1986, Samuels, G. J. GS3964, as X. appendiculata (BPI 881412). PANAMA. Canal Zone, Fort Sherman area, on fallen leaves, 5 Aug 1945, Martin, G. W. 6154, as X. appendiculata (BPI 583953 ex MO 186287).

Notes Xylaria lima features a tomentose stromatal surface, conic-papillate ostioles tilting upwards, and a non-cellular appendages at each end of the ascospores. It is similar to X. allima, from which X. lima differs primarily by its smaller ascospores.

The protologue of X. mexicana suggests that it is in synonym with X. lima. San Martín et al. (1997) reported the anamorph from immature stromata in nature, where hyaline, smooth, ellipsoid conidia (white in mass) are produced, with a size range (4–)4.5–5 × (1.5–)2(–2.5) µm.

Xylaria luxurians (Rehm) C. G. Lloyd, Mycol. Writings 5: Xylaria notes p. 29. 1918. Fig. 6G–K.

Basionym: Xylaria carpophila (Pers.) Fr. var. luxurians Rehm, Hedwigia 40: 147. 1901.

≡ Xylosphaera luxurians (Rehm) Dennis, Kew Bull. 13: 104. 1958.

Stromata cylindrical at fertile part, dichotomously or trichotomously branched once to thrice at stipe, bearing 1–3 clavae on each terminal branch, with a mucronate apex on each clava, on a long, glabrous or hirsute stipe, up to 7 cm in total length, 3–5 mm long × 1.1–2 mm broad at fertile part; surface black, with conspicuous perithecial mounds, overlain with a striped peeling layer, underlain with a thin black layer ca. 20 μm thick; interior white, homogeneous, soft. Perithecia spherical, 450–550 μm broad. Ostioles coarsely papillate, 150–200 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 130–185 μm total length, the spore-bearing part 95–140 μm long × 9.5–13 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, inverted hat-shaped, 3.5–4.5 μm high × 3–3.5 μm broad. Ascospores light brown to brown, unicellular, ellipsoid-inequilateral to crescentic, with narrowly rounded ends, smooth, (20.5–)21.5–23.5(–24.5) × (6–)6.5–8(–9) µm (22.6 ± 1.1 × 7.4 ± 0.6 μm, N = 40), with a straight to slightly oblique germ slit spore-length or nearly so on the ventral side, lacking a hyaline sheath; epispore smooth.

Specimens examined BRAZIL. St. Catharina, Blumenau, probably on fallen petioles, Jul 1888, Ule, E. 786 (holotype of X. carpophila var. luxurians S-F62514 ex Rehm herb., isotypes HBG, K[M] [in 2 packets]).

Notes Xylaria luxurians has stromata with a branched, long stipe that bears one to three clavae on each terminal branch. The photographs shown in Fig. 6G–K were taken from the isotype at HBG, while photographs from the holotype were presented in Ju et al. (2016). One of the two K(M) packets was illustrated by Dennis (1956).

Xylaria luxurians was originally published as a variety of X. carpophila, a fructicolous species quite different from X. luxurians (Ju et al. 2018). According to the protologue of X. carpophila var. luxurians (Rehm 1901), the type material was collected “in petiolis foliorum?”. Further collections are needed to determine the substrate type and the host range. Xylaria luxurians is probably only represented by the type collection thus far. Dennis (1956) cited two specimens for X. luxurians: one from the type collection and one from Maggs, D. H. L.C.T.A. 569 from Trinidad (K[M]), which is actually X. meliacearum. Dennis (1956) also noted that X. luxurians may be represented by two other specimens: a Venezuelan specimen reported in Miller (1934) and Bertero 1723 from Chile. We have not examined Miller’s Venezuelan specimen, but Miller (1934) considered it seemingly the same as X. gracillima sensu Lloyd (1918), which is based on Rick’s Brazilian material. Three specimens identified by Lloyd as X. gracillima [BRAZIL. Rick, J. (BPI 714177 ex Lloyd herb. 10439; BPI 714178 ex Lloyd herb. 182; BPI 714179 ex Lloyd herb. 12636)] were located in his herbarium and redetermined as X. luxurians by Miller. They have an oblique to slightly sigmoid germ slit on the dorsal side of the ascospores and fit X. chordaeformis C. G. Lloyd (Ju et al. 2016). It is thus plausible that Miller’s Venezuelan specimen is not X. luxurians but X. chordaeformis. The specimen Bertero 1723 [CHILE. Juan Fernández, ad corticem Urticae excelsae, 1830, as X. hypoxylon var. cupressiformis (K[M] 236761), mistaken as X. hypoxylon var. uniformis Mont. in Dennis (1956)] is X. zealandica Cooke.

Two specimens identified as X. luxurians by C. G. Lloyd [BRAZIL. Rick, J. (BPI 714175 ex Lloyd herb. 11859; BPI 714176 ex Lloyd herb. 10462)] are X. chordaeformis.

Xylaria maitlandii (Dennis) D. Hawksworth, Trans. Br. Mycol. Soc. 61: 199. 1973. Fig. 6L–P.

≡ Xylosphaera maitlandii Dennis, Revista Biol. (Lisboa) 1: 181. 1958.

Stromata cylindrical at fertile part, unbranched, with a mucronate apex, on a hirsute stipe, 40–50 mm long in total length, 15–17 mm long × 1.6–1.7 mm broad at fertile part; surface dark brown, lacking perithecial mounds or with slight perithecial mounds, overlain with a striped hirsute peeling layer, underlain with a carbonized layer ca. 40 μm thick; interior white, homogeneous, coriaceous. Perithecia spherical, 300–400 μm broad. Ostioles slightly papillate, 60–80 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 125–145 μm total length, the spore-bearing part 65–75 μm long × 7–8.5 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, inverted hat-shaped, 2.5–3 μm high × 2.5–3 μm broad. Ascospores dark brown, unicellular, ellipsoid-inequilateral, with broadly rounded ends, smooth, (10–)10.5–11.5(–12) × (5.5–)6–6.5(–7) µm (10.9 ± 0.5 × 6.4 ± 0.3 μm, N = 40), with a straight germ slit spore-length on the ventral side, lacking a hyaline sheath; epispore smooth.

Specimen examined UGANDA. Entebbe, Kitubilu Forest, on twigs lying on ground amongst leaf mould, July 1919, Maitland, T. D. (holotype of Xylosphaera maitlandii K[M] 236752).

Notes Xylaria maitlandii resembles X. arbuscula Sacc. and its affinities in having stromata with a mucronate apex and striped outer peeling layer but is unique in having hairs on the outer peeling layer. It remains to be verified if X. maitlandii is associated with petioles because the original paper slip enclosed in the packet indicates that the material was collected from twigs. Dennis (1958), however, changed the substrate to fallen petioles in the protologue.

Xylaria meliacearum Læssøe, in Læssøe & Lodge, Mycologia 86: 441. 1994. Fig. 7A–H, I, L, M.

Stromata filiform at fertile part, unbranched, with an acute apex, on a glabrous stipe, 17–65 mm long in total length, 3–17 mm long × 1.6–2.5 mm broad at fertile part; surface sulfur yellow at axis but blackish brown at perithecial mounds, with 2/3-exposed to fully exposed perithecial mounds, overlain with a sulfur-yellow outer layer attached with synnematal remnants at places, gradually ruptured by perithecial mounds into flaky remnants and sloughing off afterwards, underlain with a thin black layer ca. 20 μm thick; interior white, homogeneous, soft. Perithecia spherical, 500–800 μm broad. Ostioles papillate, 120–140 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 180–270 μm total length, the spore-bearing part 120–170 μm long × 7–8 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, urn-shaped to cylindrical with 1–2 constrictions, 5.5–7 μm high × 3.5–4.5 μm broad. Ascospores brown, unicellular, ellipsoid-inequilateral, with narrowly rounded ends, smooth, (19–)21.5–27.5(–31.5) × (5–)5.5–7(–8) µm (24.5 ± 3.1 × 6.2 ± 0.7 μm, N = 108), with a spiral germ slit spore-length or slightly less than spore-length on the ventral side, lacking a hyaline sheath; epispore smooth.

Cultures and anamorph were reported by Læssøe and Lodge (1994). Stromata were produced on OA with the anamorph formed on stromatal initials. The anamorph formed on the stromata collected from the field is located on the stromatal surface as synnemata or as a continuous layer.

Specimens examined FRENCH GUIANA. Paul Isnard Area, Mt. Decou Decou, on twig, 11–12 Mar 1986, Samuels, G. J. & Searwar, P. GS4251, as X. meliacearum (HAST 145976 ex NY); ca. 10 km SW of Saul, track from Saul leading SW toward Mt. Galbao, elev. ca. 200 m, on twig, 7–9 Jan 1986, Samuels, G. J. & Boise, J. GS2600, as X. meliacearum (BPI 881642); ca. 15 km SW of Saul, along trail from Saul to Mt. Galbao, alt. 600 m, on dead twigs, 14–29 Jan 1986, Samuels, G. J. & Boise, J. GS2879, as X. meliacearum (HAST 145975 ex NY); ca. 17.5 km SW of Saul, Mt. Galbao, Camp 3, alt. 350 m, on leaf rachis, 24–26 Jan 1986, Samuels, G. J. & Boise, J. GS3203, as X. meliacearum (BPI 881654). PUERTO RICO. Luquillo Mts., El Verde Research Area, on petioles of Guarea guidonia, 16 Oct 1992, Lodge, D. J. PR894 (paratype of X. meliacearum WSP ex Rogers herb.). TRINIDAD AND TOBAGO. Trinidad, Heights of Aripo, on dead wood, 11 Jun 1945, Maggs, D. H. L.C.T.A. 569, as X. luxurians (K[M]). VENEZUELA. Edo. Trujillo, Parque Nacional Guaramacal, ca. 10 km SW of Batatal, La Defensa, along Río Saguás, Campamiento Granja Bocono, in forest along trail to water source, alt. 2000 m, on petiole of Cecropia (Urticaceae)?, 20, 23 Nov 1990, Samuels, G. J., Hein, B. & Huhndorf, S. M. GS7461, as X. meliacearum (HAST 145977 ex BPI).

Notes Xylaria meliacearum belongs to the X. filiformis group and differs from its species mainly by larger ascospores that have a spiral germ slit. It is noteworthy that the anamorph is born on immature stromata as upright, sulfur-yellow synnematal pegs and becomes synnematal remnants that can still be observed adhering to the outer stromatal layer. The ascospore size range of X. meliacearum is fairly broad, much as given by Læssøe and Lodge (1994) where the ascospores were measured “(18.5–)19.1–30.0(–33.0) × (4.0–)4.6–6.6(–7.9) µm (means from various collections 20.1–29.7 × 4.7–7.5 µm)”.

The Venezuelan collection GS7461 is mixed with two other species, X. sp. GS7461A and X. sp. GS7461B, on the same petiole (Fig. 7H). All of these three species have a spiral germ slit on their ascospores but differ by the combinations of stromatal morphology and ascospore size range.

Xylaria memecyli Pande, Nova Hedwigia 24: 14. 1973, as “memecylonii.”

For a description of the teleomorph, see Pande (1973) where illustrations of stromata, perithecia, asci, and ascospores are also provided. Xylaria memecyli is characterized by the following features: stromata peltate at fertile part, concave underneath, unbranched, convex on top, on a long, glabrous stipe, 10–15 mm in total length, 1 mm high × 2 mm broad at fertile part, with a dark brown to blackish surface lacking perithecial mounds, with a whitish to pale yellow interior; perithecia spherical to obovoid, 400–480 μm broad × 512–590 μm high; ascospores brown, ellipsoid-inequilateral, 12–14 × 6–8 μm, with a straight germ slit spore-length, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages.

Notes Xylaria memecyli is known only from the type collection, which was made from fallen leaves of Memecylon umbellatum (Melastomataceae) in India (Pande 1973).We made a request for a loan of the type preserved at AMH but did not receive a reply. Although certain important features, such as stromatal surface, perithecial aggregation in stromata, ostioles, and certain ascospore traits, were not described in the protologue, the stromatal shape and ascospore morphology suggest that X. memecyli is a distinct species. The stromatal size was not given by Pande (1973); the measurement provided above was made from the two stromata on a leaf presented in Fig. 2 by Pande (1973), which were sketched in natural habit as mentioned in the figure legend.

Xylaria minuscula Y.-M. Ju & H.-M. Hsieh, sp. nov. Figs. 8A–M, 10E, 12 K, L.

MycoBank MB848542.

Typification TAIWAN. I-lan Co., Yuan-shan, Fu-shan, on dead leaves of Castanopsis carlesii var. sessilis, 27 Oct 2001, Ju, Y.-M. & Hsieh, H.-M. 90102701 (cultured) (holotype of X. minuscula HAST 145978), GenBank: ITS = OQ883721.

Etymology Referring to its minute stromata.

Stromata cylindrical at fertile part, unbranched, with a long acicular apex, on a glabrous stipe, 3–14 mm in total length, 1–5 mm long × 0.6–1.1 mm broad at fertile part; surface black, with conspicuous to half-exposed perithecial mounds, overlain with a dull grayish brown peeling layer split into narrow or thread-like stripes, underlain with a thin black layer ca. 30 μm thick; interior white, homogeneous, soft. Perithecia spherical, 300–500 μm broad. Ostioles coarsely papillate, 110–120 μm high × 180–190 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 130–180 μm total length, the spore-bearing part 80–95 μm long × 7–8 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, inverted hat-shaped, 2.5–3 μm high × 2.5–3 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid-inequilateral, with narrowly rounded ends bearing a tiny hyaline cellular appendage on one end, smooth, (13.5–)14–15(–17) × (4.5–)5–6(–7) µm (14.5 ± 0.8 × 5.7 ± 0.5 μm, N = 56), with a straight germ slit spore-length on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages; epispore smooth.

Cultures and anamorph. Colonies reaching the edge of 9-cm Petri dish in 3 wk, white, velvety, appressed, azonate, with diffuse margins. Reverse uncolored. Stromata arising at periphery, conical, unbranched, 1–2 mm high × 1–2 mm diam at base, white, immediately becoming black from base upwards. Anamorph not produced.

Additional specimens examined MALAYSIA. Sarawak, on dead leaves, Beccari, O., as X. phyllocharis (K[M] 236749 ex Cooke herb.), mixed with the lectotype of X. phyllophila. TAIWAN. Nan-tou County, Yu-chee, Lien-hwa-chee, on fallen leaves, 20 Aug 1991, Ju, Y.-M. 80082005, as X. sp. (WSP).

Notes Xylaria minuscula was reported as an unnamed Xylaria sp. (80082005) from Taiwan by Ju and Rogers (1999) who suspected that X. phyllophila might be a tenable name for it. However, X. phyllophila differs from X. minuscula mainly by having larger ascospores (15–)16.5–18(–19) × (7.5–)8–9(–9.5) µm. Xylaria minuscula resembles X. foliicola in stromatal morphology but differs by its ascospores having a larger size range, i.e., (13.5–)14–15(–17) × (4.5–)5–6(–7) µm vs. (8.5–)9–11 × 4–6 μm, and being enclosed within a hyaline sheath.

Xylaria nainitalensis Dargan, J. Bio. Res. 3: 43. 1983. Fig. 8N–R.

Stromata capitate at fertile part, unbranched, with a long acicular apex, on a glabrous stipe, 91–147 mm in total length, 1.3–2.9 mm long × 1–2.7 mm broad at fertile part; surface black, with conspicuous perithecial mounds, lacking an outer layer, underlain with a thin, black layer ca. 20 μm thick; interior white, homogeneous, soft. Perithecia spherical, 350–450 μm broad. Ostioles coarsely papillate, 90–130 μm high × 150–180 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 150–180 μm total length, the spore-bearing part 80–95 μm long × 7–8.5 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, inverted hat-shaped, 2.5–3 μm high × 2.5–3 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid-inequilateral, with narrowly rounded ends sometimes bearing a tiny hyaline cellular appendage on one end, smooth, (10.5–)11.5–13.5(–15) × (5–)5.5–6.5(–7.5) µm (12.3 ± 1.0 × 6.2 ± 0.5 μm, N = 50), with a straight germ slit spore-length on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages; epispore smooth.

Specimens examined INDIA. on twig, Narula, A. M. 313, as X. nainitalensis (WSP ex Rogers herb.); Uttar Pradesh, Nainital, Sat Tal, on dead leaves of Quercus incana, 17 Aug 1973, Dargan, J. S. 13117 (isotype of X. nainitalensis K(M) 236748).

Notes Xylaria nainitalensis features a capitate stromatal fertile part with conspicuous perithecial mounds on the surface and terminating into a long acicular apex. Its ascospores enclosed within a hyaline sheath that forms a non-cellular appendage at each end. A tiny cellular appendage can sometimes be seen on one end of the ascospores.

A specimen [USA. Florida, San Felasco Reserve, on fallen leaves and petioles, 10 Aug 1985, Rogers, J. D., as X. cf. filiformis (WSP ex Rogers herb.)], bearing a capitate stromatal fertile part that terminates into a long thin apex, looks much like X. nainitalensis but is immature.

Xylaria neblinensis Y.-M. Ju & H.-M. Hsieh, sp. nov. Fig. 8S–W.

MycoBank MB848544.

Typification VENEZUELA. Territorio Federal Amazonas, Cerro de la Neblina, 6.2 km NE Pico Phelps, alt. 1390–1515 m, on fallen leaves of Clusia sp., 22 Feb 1985, Rossman, A. Y. AR2233, as X. cf. brachiata (holotype of X. neblinensis BPI 583728, isotype K[M]).

Etymology Referring to the collecting site Cerro de la Neblina, Venezuela.

Stromata cylindrical at fertile part, unbranched, with an acute apex, on a glabrous stipe, 15–30 mm in total length, 8–12 mm high × 0.6–0.8 mm broad at fertile part; surface blackish brown, with conspicuous perithecial mounds, lacking an outer layer, with the layer immediately beneath the surface extremely thin, less than 10 μm thick; interior white, homogeneous, soft. Perithecia spherical, 200–300 μm broad. Ostioles papillate, 60–80 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 80–95 μm total length, the spore-bearing part 55–65 μm long × 7.5–8.5 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, inverted hat-shaped, 2–2.5 μm high × 2–2.5 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid-inequilateral, with narrowly rounded to broadly rounded ends, smooth, (9–)9.5–10.5(–11.5) × (5–)5.5–6(–6.5) µm (10.2 ± 0.5 × 5.6 ± 0.3 μm, N = 40), with a straight germ slit spore-length on the ventral side, lacking a hyaline sheath; epispore smooth.

Notes Xylaria neblinensis is in general similar to X. appendiculatoides in stromatal morphology, having a blackish brown surface with conspicuous perithecial mounds and a pointed apex. The ascospores of X. neblinensis differ from those of X. appendiculatoides by having a smaller size range and lacking a hyaline sheath.

Rogers et al. (1988) tentatively referred to this fungus as X. cf. brachiata Sacc. but noted that this fungus was not known to them. A larger ascospore size of 14.5–16 × 6.5–7.5 μm was given in Rogers et al. (1988), where specimens AR2232 and AR2284 were also cited for this fungus, in addition to AR2233. It is possible that larger ascospores can be found in these two collections. The notes enclosed in the packet of AR2233 indicate that the ascospore size is much like our measurement. Although the stromatal apex is currently broken off, an acute stromatal apex was nicely depicted by A. Rossman on the notes.

Xylaria noduliformis Y.-M. Ju & H.-M. Hsieh, nom. nov. Fig. 9A–E.

MycoBank MB848545.

≡ Xylosphaera maitlandii Dennis var. nuda Dennis, Revista Biol. (Lisboa) 1: 183. 1958.

≡ Xylaria maitlandii (Dennis) D. Hawksworth var. nuda (Dennis) D. Hawksworth, Trans. Br. Mycol. Soc. 61: 200. 1973.

Typification UGANDA. Entebbe, Kitubilu Forest, on twigs lying on ground amongst leaf mold, Jun 1919, Maitland, T. D. 484A (holotype of Xylosphaera maitlandii var. nuda K[M] 236727).

Etymology Referring to the stromatal surface with fully exposed perithecial mounds.

Stromata cylindrical at fertile part, unbranched or branched once at clavae, with an acuminate or mucronate apex, on a glabrous stipe, 29–53 mm in total length, 9–22 mm long × 1.2–2.7 mm broad at fertile part; surface black, with fully exposed perithecial mounds, overlain with a brown striped peeling layer obscured with the perithecial mounds fully exposed, underlain with a thin, black layer ca. 30 μm thick; interior white, homogeneous, soft. Perithecia spherical, 400–550 μm broad. Ostioles slightly papillate, often encircled with a paler area, 50–80 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 180–210 μm total length, the spore-bearing part 70–80 μm long × 7.5–8.5 μm broad, with an apical ring staining deep blue in Melzer’s iodine reagent, inverted hat-shaped, 2–2.5 μm high × 2–2.5 μm broad. Ascospores dark brown to blackish brown, unicellular, ellipsoid-inequilateral, with narrowly to broadly rounded ends, smooth, (9–)9.5–10.5(–11) × (5.5–)6–6.5(–7) µm (10.0 ± 0.5 × 6.0 ± 0.2 μm, N = 40), with a straight germ slit spore-length on the ventral side, lacking a hyaline sheath; epispore smooth.

Notes Xylaria noduliformis is only known from the type material and is characterized by half-exposed, highly crowded perithecial mounds. Dennis (1958) recorded X. noduliformis (as Xylosphaera maitlandii var. nuda) from the same kind of fallen petioles where the original material of X. maitlandii was collected. However, like X. maitlandii, X. noduliformis may not be associated with petioles because the original paper slip enclosed in the type packet of X. maitlandii indicates twigs rather than petioles as the substrate.

Xylaria petchii C. G. Lloyd, Mycol. Writings 7: 1310. 1924. Fig. 9F–L.

= Xylaria filiformoidea Hladki & A. I. Romero, Fungal Diversity 42: 83. 2010.

Stromata capitate to short-cylindrical at fertile part, unbranched or branched occasionally at stipe, with an apiculate apex, on a glabrous stipe, 7.5–20 mm in total length, 2–5 mm high × 1–2 mm broad at fertile part; surface black, somewhat polished, with conspicuous to half-exposed perithecial mounds clumped together on upper part, frequently with several loose, fully exposed perithecial mounds scattered below, lacking an outer layer, underlain with a thin, soft layer ca. 10 μm thick or less, concolorous with the surface; interior white, homogeneous, soft. Perithecia spherical, 400–500 μm broad. Ostioles conic-papillate, 110–150 μm high × 150–190 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 95–125 μm total length, the spore-bearing part 55–70 μm long × 6–7 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, inverted hat-shaped, 2–2.5 μm high × 2–2.5 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid-inequilateral, with narrowly to, less frequently, broadly rounded ends, smooth, (7.5–)8.5–9.5(–10) × (3.5–)4–4.5(–5) µm (8.9 ± 0.5 × 4.4 ± 0.2 μm, N = 40), with a straight germ slit spore-length on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages; epispore smooth.

Specimens examined SRI LANKA. Peradeniya, on fallen leaves, 29 Jan 1919, Petch, T. 6126 (holotype of X. petchii BPI 714305 ex Lloyd herb. 10407, isotype K[M] 169689).

Notes Xylaria petchii is unique in having most perithecia compacted together near the apex of stromata with several loosely scattered below. Petch collected this fungus and sent two stromata to Lloyd, as shown in Fig. 2960 in Lloyd (1924), upon which Lloyd’s description of X. petchii was based. The holotype packet now contains only one stroma, while the isotype packet at K contains seven stromata. The stromatal habit of X. petchii is also shown in X. filiformoidea, which is described from Argentina with ascospores 8–9 × 4–5 μm and enclosed within a hyaline sheath (Hladki and Romero 2010). Xylaria filiformoidea is inseparable from X. petchii, and these two names are thus considered in synonymy. Xylaria petchii is briefly noted in Ju et al. (2016).

Xylaria phyllocharis Mont., Ann. Sci. Nat., Bot., sér. IV, 3: 108. 1855. Fig. 9M–T.

≡ Xylosphaera phyllocharis (Mont.) Dennis, Kew Bull. 13: 105. 1958.

Stromata cylindrical at fertile part, unbranched, with an acuminate to mucronate apex, on a glabrous stipe, 11–40 mm in total length, 5–16 mm long × 0.7–1.3 mm broad at fertile part; surface dark vinaceous brown, with slight perithecial mounds, lacking an outer layer, underlain with a thin, soft layer ca. 30 μm thick, concolorous with the surface; interior white, homogeneous, soft. Perithecia spherical, 100–250 μm broad. Ostioles papillate, 80–120 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 95–130 μm total length, the spore-bearing part 65–85 μm long × 7–9 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, short-cylindrical to urn-shaped, 2.5–3.5 μm high × 2.5–3.5 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid, strongly inequilateral, with broadly rounded ends sometimes bearing a tiny hyaline cellular appendage on one end, smooth, (9–)10–11.5(–12.5) × (4.5–)5.5–6.5(–7) µm (10.6 ± 0.7 × 6.0 ± 0.6 μm, N = 100), with a straight germ slit spore-length on the ventral side, lacking a hyaline sheath; epispore smooth.

Cultures and anamorph were reported by San Martín et al. (1997) with obovoid to ellipsoid conidia 4–5 × 2.5–3 μm.

Specimens examined FRENCH GUIANA. on fallen leaves, Leprieur, F. R. 1208 (holotype of X. phyllocharis PC 0086048 ex Montagne herb., isotype PC 0086071 ex Leprieur herb.); ca. 15 km SW of Saul, along trail from Saul to Mt. Galbao, wet forest with moss and epiphyte covered trees, alt. 600–650 m, on dead leaves, 14–29 Jan 1986, Samuels, G. J. & Boise, J. GS2962 (HAST 145979 ex NY); Saul, Monts La Fumee, dry primary forest, alt. ca. 400 m, on dead leaves, 4–6 Feb 1986, Samuels, G. J. & Boise, J. GS3519 (HAST 145980 ex NY). FRENCH WEST INDIES. Guadeloupe, Trace de Sofaïa, on dead leaves, 1 Sep 2005, Lechat, C. L. 5302 (HAST 145981 ex LIP), GenBank: ITS = GU322445; Martinique, Morne Rouge, La Propreté forest track, on dead leaves in rainforest, 6 Jun 2014, Fournier, J. MJF 14080 (HAST 145152 ex LIP).

Notes Xylaria phyllocharis can easily be confused with X. appendiculata due to their similar stromatal morphologies and ascospore size ranges. However, X. appendiculata can be distinguished from X. phyllocharis by its more conspicuous, conic-papillate ostioles and its ascospores that are not strongly inequilateral and are surrounded by a hyaline sheath that swells on both ends. Fournier et al. (2020) reported X. phyllocharis from French West Indies and acknowledged variation in ascospore size and shape as well as in the presence or absence of a hyaline sheath among examined specimens; it is likely that some of these specimens are assignable to X. appendiculata. Like the stromatal surfaces of X. phyllocharis and X. appendiculata, those of X. lima and X. allima also have a vinaceous tinge; however, those of the latter two species are entirely tomentose.

The holotype in the Montagne herbarium (PC) is not fully mature, with the stromatal surface still overlain with some powdery remnants, but the isotype from the Leprieur herbarium (PC) is fully mature. The isotype packet from the Leprieur herbarium contains three different Xylaria species, each glued onto a separate paper card as shown in Fig. 9O. Xylaria aristata is on the upper left, X. phyllocharis is on the lower left, and the newly described X. spiculaticlavata is on the right.

Xylaria phyllophila Ces., Atti Accad. Sci. Fis. 8: 15. 1879. Fig. 10A–G.

Stroma cylindrical at fertile part, unbranched, with an acute apex, on a glabrous stipe, 22 mm in total length, 9 mm high × 1 mm broad at fertile part; surface blackish brown, with conspicuous perithecial mounds, lacking an outer layer, underlain with a thin, soft layer ca. 10 μm thick or less, concolorous with the surface; interior white, homogeneous, soft. Perithecia spherical, 350–500 μm broad. Ostioles papillate, 50–80 μm broad at base. Asci not intact, with an apical ring staining deep blue in Melzer’s iodine reagent, inverted hat-shaped, 3.5–4 μm high × 3–4 μm broad. Ascospores dark brown, unicellular, ellipsoid-inequilateral, with narrowly rounded ends, smooth, (15–)16.5–18(–19) × (7.5–)8–9(–9.5) µm (17.3 ± 0.9 × 8.7 ± 0.5 μm, N = 40), with a straight germ slit spore-length or nearly so on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages; epispore smooth.

Specimen examined MALAYSIA. Sarawak, on dead leaves, Beccari, O., as X. phyllocharis (lectotype [designated here, MycoBank Typification No. 10013067] of X. phyllophila K[M] 236749 ex Cooke herb.).

Notes Xylaria phyllophila can be readily distinguished from species in the X. phyllocharis group with long cylindrical stromata and conspicuous perithecial mounds by having ascospores longer than 16 μm and enclosed within a hyaline sheath.

Type material of X. phyllophila was not located at RO. A part of the type material sent to Cooke by Cesati was kept in two packets at K; both were labeled as X. phyllocharis, reflecting the doubt raised in the protologue that X. phyllophila could be X. phyllocharis. There are five stromata attached to the leaf enclosed within K(M) 236749, but only two of them are fertile. One stroma has a long cylindrical fertile part and larger ascospores (15–)16–18.5(–19) × 8–9.5 μm, while the other has an ellipsoid fertile part and smaller ascospores 13.5–15 × 5.5–6.5 μm. Cesati (1879) originally described X. phyllophila as having a cylindrical fertile part, 0.5 to 1 cm long, umber in color, torulose, apiculate or indeed aristate but did not give the ascospore measurement. Cooke (1883) drew a long cylindrical stroma in accordance with Cesati’s description and gave the ascospore measurement as 20 × 10 μm. Thus, the stroma with a long cylindrical fertile part is lectotypified for X. phyllophila, and the description herein is based on one stroma only; it should be noted that the stromatal surface has been partially ruined by insects. The other fertile stroma with an ellipsoid fertile part belongs to X. minuscula (Fig. 10E). The other K packet contains a leaf attached with two stromata with an ellipsoid fertile part. We have not studied the material enclosed in this packet microscopically but suspect that it is X. minuscula.

Xylaria pisoniae D. Scott, J. D. Rogers & Y.-M. Ju, in Rogers, Scott & Ju, Harvard Pap. Bot. 6: 189. 2001. Fig. 10H–O.

Stromata capitate to conic at fertile part, unbranched or infrequently branched, with an acute apex, on a glabrous stipe, up to 3–15 mm in total length, 1–3 mm high × 1–2 mm broad at fertile part; surface black, with conspicuous perithecial mounds, lacking an outer layer, underlain with a thin, black layer ca. 20 μm thick; interior white, homogeneous, soft. Perithecia spherical, 200–400 μm broad. Ostioles slightly papillate, sometimes encircled with a paler area, ca. 50 μm broad at base. Asci with eight ascospores arranged in uniseriate to partially biseriate manner, cylindrical, 120–155 μm total length, the spore-bearing part 65–95 μm long × 7–9 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, inverted hat-shaped to cylindrical, 2.5–3 μm high × 2–2.5 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid-inequilateral, with narrowly to broadly rounded ends, smooth, (10–)10.5–12(–12.5) × (5–)5.5–6(–6.5) µm (11.3 ± 0.6 × 5.7 ± 0.3 μm, N = 40), with a straight germ slit spore-length or nearly so on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages; epispore smooth.

Cultures and anamorph were reported by Rogers et al. (2001). The anamorph was found on the stromata produced in culture and collected from the field. The teleomorph was produced on OA in 6 wk.

Specimen examined USA. Hawaiian Islands, Hawaii, Volcanoes National Park, Pu’u Puaulu (Bird Park), on dead leaves of Pisonia brunoniana, 15 Jan 2000, Rogers, J. D. (isotype of X. pisoniae WSP).

Notes Xylaria pisoniae has fairly minute stromata characterized by an acute apex and a capitate or conic fertile part roughened by conspicuous perithecial mounds. It is known only from the type specimen collected from the Hawaiian Islands. Xylaria nainitalensis and X. sicula are morphologically similar to X. pisoniae, from which they can be readily distinguished by their acicular stromatal apex that is much longer than the fertile part.

Xylaria polysporicola H.-X. Ma & X.-Y. Pan, in X.-Y. Pan, Z.-K. Song, Z. Qu, T.-D. Liu & H.-X. Ma, MycoKeys 86: 57. 2022.

For a description of the teleomorph, see Pan et al. (2022) where illustrations of stromata, asci, and ascospores are also provided. Xylaria polysporicola is characterized by the following features: stromata cylindrical at fertile part, unbranched or occasionally branched once at clavae, with an acute apex, on a glabrous stipe, 1–4 cm in total length, 2–15 mm long × 0.5–1.6 mm broad at fertile part; stromatal surface black, slightly polished, with conspicuous perithecial mounds and fine longitudinal striations, lacking an outer layer, underlain with a thin, soft layer ca. 70 μm thick, concolorous with the surface, with a white, soft interior; perithecia subspherical, 450–600 μm diam, with a papillate ostiole ca. 70 μm broad at base; ascospores brown to dark brown, ellipsoid-inequilateral, with broadly to narrowly rounded ends, sometimes with one end slightly pinched, smooth, (11.5–)12.5–14.5(–15) × 5.5–8 μm, with a straight germ slit slightly less than spore-length on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages.

Notes Xylaria polysporicola is keyed out in the X. phyllocharis group herein owing to the cylindrical stromata and undulate perithecial mounds. It resembles X. appendiculata and X. appendiculatoides but can be distinguished by having larger perithecia and less conspicuous ostioles, which are papillate and approximately 70 μm broad at base as opposed to coarsely conic-papillate and more than 100 μm broad at base. Nonetheless, the presence of fine longitudinal striations on the slightly polished black stromatal surface suggests that X. polysporicola may be related to X. filiformis and X. vagans, where the perithecial mounds are exposed and ascospores are slightly smaller.

Xylaria sicula Pass. & Beltr., Atti della R. Accademia dei Lincei, Transunti, ser. III, 7: 36. 1882. Fig. 10P–W.

≡ Thamnomyces siculus (Pass. & Beltrani) Maire, Mycotheca Boreali-Africana, fasc. 4, no. 100. 1913.

≡ Podosordaria sicula (Pass. & Beltrani) P. M. D. Martin, J. S. African Bot. 36: 134. 1970.

= Xylaria sicula Pass. & Beltr. f. major Ciccarone, Nuovo G. Bot. Ital. 53: 357. 1947.

Stromata capitate, depressed-capitate or obconical at fertile part, unbranched, with a long acicular apex, on a glabrous stipe, up to 3–20 mm in total length, 0.7–1.5 mm high × 1–1.5 mm broad at fertile part; surface black, with conspicuous to half-exposed perithecial mounds, lacking an outer layer, underlain with a thin, black layer ca. 20 μm thick; interior white, homogeneous, soft. Perithecia spherical, 400–550 μm broad. Ostioles slightly papillate, sometimes encircled with a paler area, ca. 60 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 115–165 μm total length, the spore-bearing part 70–90 μm long × 6.5–8 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, inverted hat-shaped, 2.5–3 μm high × 2.5–3 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid-inequilateral, with narrowly to broadly rounded ends, smooth, (8.5–)9.5–11(–12) × (4–)4.5–6(–6.5) µm (10.3 ± 0.7 × 5.3 ± 0.6 μm, N = 40), with a straight germ slit slightly less than spore-length to nearly spore-length on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages; epispore smooth.

Specimens examined ALGERIA: on dead olive leaves, Maire, R., as X. sicula (BPI 714381 ex Lloyd herb. 12667, MPU), immature; on dead olive leaves, Nov 1920, Maire, R. 7337, as X. sicula (BPI 714379 ex Lloyd herb. 10408), immature; L’Alma, sur les feuilles tombées d’Olea europaea L., 19 Oct 1912, Maire, R., Maire’s Mycotheca Boreali-Africana 100, as X. sicula (S F43691 ex Sydow herb., BPI 586419), immature. GREECE: Corfu, Stefanos, near Agios, on fallen leaves of Olea europaea, 12 May 2001, Spooner, B. M., as X. sicula (K[M] 136350). SPAIN: Balearic Islands, Menorca, on decaying Olea leaf litter and debris, 21 May 2002, Spooner, B. M., as X. sicula (K[M] 99656).

Notes Xylaria sicula is mainly distributed in the Mediterranean region, fruiting on fallen olive leaves. The holotype, collected from Sicily, Italy, could not be located by Graniti (1959) who designated the type of X. sicula f. major from Kenya as the neotype of X. sicula. For notes on the literature of X. sicula, see Fournier (2014). Xylaria sicula f. major mentioned in Hsieh et al. (2010) is not this species but rather X. aristata var. aristata.

Martin (1970) recombined the epithet with Podosordaria but cited a reference by Passerini and Beltrani (1883) instead of the protologue in Passerini and Beltrani (1882), where the basionym was published. According to ICN 41.8 (Shenzhen Code), Martin’s error is to be corrected (Martin 1976) and does not affect the valid publication of Podosordaria sicula in 1970.

A collection from BRAZIL [São Leopoldo, on fallen leaves, Theissen, F. 10, as X. aristata (S F72249 ex Bresadola herb.)], which has ascospores (10.0–)10.4–11.4(–11.7) × (4.5–)4.9–5.5(–5.8) µm and evident perithecial mounds, is similar to X. sicula. It remains to be determined if this collection is truly X. sicula, a host-specific species growing on olive leaves in the Mediterranean region, far away from Brazil.

Xylaria simplicissima (Pers.) Y.-M. Ju & H.-M. Hsieh, comb. nov. Figs. 11A–G, 12J.

MycoBank MB848546.

Basionym: Rhizomorpha simplicissima Pers., Mycol. Eur. (Erlanga) I, p. 59. 1822.

≡ Xylaria subularis Fr., Summa Veg. Scand. II, p. 382. 1849.

Typification FRANCE. on herbaceous stems, Chaillet (lectotype [designated here, MycoBank Typification No. 10013068] of Rhizomorpha simplicissima UPS F-004396 ex Fries herb.), immature. FINLAND. North Karelia, Outokumpu, Rikkaranta, Eskola, N 62.7709, E 28.757, on herbaceous stems, Oct 2011, Pennanen, M. MP111004 (cultured), comm. Fournier, J. (epitype [designated here, MycoBank Typification No. 10013069] of Rhizomorpha simplicissima HAST 145982), GenBank: ITS = OQ883722.

= Sphaeria stipiticola Swartz, Summa Veg. Scand. (Swartz), p. 55. 1814.

= Xylaria filiformis (Albertini & Schwein.) Fr. f. caulincola Rehm, Ann. Mycol. 10: 57. 1912.

= Xylaria jiangsuensis R. S. Wang & L. Guo, in G. Huang, R. S. Wang, L. Guo & N. Liu, Mycotaxon 130: 302. 2015.

= Xylaria crinalis H.-X. Ma, Lar.N. Vassiljeva & Y. Li,, in H.-X. Ma & Y. Li, Sydowia 70: 39. 2018.

Stromata filiform at fertile part, unbranched or rarely branched at clavae, with a long acicular apex, on a glabrous glossy stipe, (10–)25–55(–65) mm in total length, 3–15 mm long × 1–2 mm broad at fertile part; surface blackish, with half-exposed to fully exposed perithecial mounds, lacking an outer layer, underlain with a thin, black layer ca. 20 μm thick; interior white, homogeneous, fragile. Perithecia spherical, 550–750 μm broad. Ostioles slightly papillate, ca. 70 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 175–220 μm total length, the spore-bearing part 110–130 μm long × 7–8 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, cylindrical to urn-shaped, 3–3.5 μm high × 2.5–3 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid to shortly fusoid, inequilateral, with narrowly rounded ends, smooth, (15–)16.5–19(–21.5) × (5–)5.5–6.5(–7.5) µm (17.7 ± 1.2 × 6.1 ± 0.5 μm, N = 100), with a straight germ slit spore-length on the ventral side, lacking a hyaline sheath; epispore with a lumpy appearance along the germ slit.

Cultures and anamorph. Colonies reaching the edge of 9-cm Petri dish in 2 wk, whitish, mostly submerged, azonate, with diffuse margins. Reverse uncolored. Stromata arising in 2–3 concentric zones, cylindrical, tapering upwards, unbranched, up to 2.6 cm × 0.5–1 mm diam, becoming black from base upwards, white on the surface of upper part and orangish at tip, with abundant clear droplets on the surface. Anamorph not produced.

Additional specimens examined that are mature AUSTRIA. Österreich, Steiermark, Schladminger Tauern, Kleinsölktal S von Gröbming, Aufstieg von der Putzentalalm zur Oberen Alm, ca. 1500 m alt., on stems of Cicerbita alpina, 18 Sep 1991, Scheuer, C., as X. filiformis (WSP 70041 ex GZU). CANADA. Ontario, London, on dead herbaceous stems, Aug–Oct 1911, Dearness, J., Bartholomew’s Fungi Columbiani 3600, as X. filiformis (HBG, S F131049 ex Sydow herb., WSP 182159); Ontario, London, on old Archangelica stems, Jul–Aug 1911, Dearness, J., Rehm’s Ascomyceten 1969 (holotype of X. filiformis var. caulincola S F131060 ex Rehm herb., isotypes HBG, S F131063 ex Sydow herb.); Ontario, London, on old stems of Archangelica sp. and other plants, 22, 29 Jul 1911, Dearness, J. 2138, as X. filiformis var. caulincola (S F131064 ex Rehm herb.); Ontario, near London, on various dead herbaceous stems, 20 Sep 1911, Dearness, J., as X. filiformis var. caulincola (HBG); Ontario, pr. London, in caulibus herbarium, praecipue Solidaginis, Sep 1911–1913, Dearness, J., Sydow’s Fungi Exotici Exsiccati 258, as X. filiformis f. caulincola (HBG, L 1038608 ex U 034503, S F131051 ex Sydow herb., S F131053 ex Sydow herb., S F131057 ex Sydow herb.). GERMANY. Oberammergau, Kapellenwand, on stem of Lunaria rediviva, Sep 1889, Allescher, as X. filiformis (S F271420 ex Bresadola herb.). JAPAN. Honshu, Tochigi Pref., Okka-Nikko, Koto-Ku, Bakujo, on Sasa sp., 26 Aug 1983, Samuels, G. J. GS83-342, as X. cf. filiformis (WSP ex Rogers herb.). RUSSIA. Berdicino, prov. Jaroslawl, in caulibus emortuis Epilobii angustifolii L., Aug 1911, Serebrianikow, Tranzschel & Serebrianikow’s Mycotheca Rossica 328, as X. filiformis (HBG); Russland, St. Petersburger Forst-Institut, in petiolis Fraxini excelsioris, Sep 1895, collectors unknown, as X. filiformis (S F271421 ex Rehm herb.). USA. New Hampshire, Chocorua, on dead stems and leaves, 4 Aug 1909, Farlow, W. G., Thaxter’s Reliquiae Farlowianae 97, as X. filiformis (S F131048); North Dakota, Kulm, on herbaceous stems of Iva xanthiifolia, 16 Aug 1923, Brenckle, J. F., Brenckle’s Fungi Dakotenses 575, as X. filiformis f. caulincola (WSP 23494).

Additional specimens examined that are immature AUSTRIA. Niederösterreich, on petioles of Rubus sp., Jun 1905, Strasser, P., as X. filiformis (S F271422), immature; Österreich, Steiermark, Rottenmanner Tauern, am Pölsbach S von St. Johann a. Tauern, ca. 1015 m alt., on stems of Scirpus sylvaticus, 12 Aug 1984, Scheuer, C., as X. filiformis (WSP 70040 ex GZU), immature. CZECH REPUBLIC. Reichenberg in Bohemia, ad petiolos arborum frondosarum, praesertim Aceris, Siegmund Jun., W., Rabenhorst’s Fungi Europaei 917, as X. filiformis (BPI 739338 ex Nitschke herb., HBG [in 2 packets], S F131047 ex Rehm herb.), immature; Reichenberg in Bohemia, on used carpet knitted from plant fibers, Siegmund Jun., W., Rabenhorst’s Fungi Europaei 57, as X. filiformis (HBG, HBG ex Magnus herb., S F131040 ex Rehm herb., S F131041 ex Sydow herb., S F131046 ex Sydow herb., S F131050 ex Sydow herb.), immature. EUROPE. no detailed locality, on herbaceous stem, Mougeot, J. B. 989, as Rhizomorpha simplicissima (L 0117822 ex Persoon herb.), immature. FINLAND. Mustiala, På Gråbo, on herbaceous stems, 17 Dec 1865, Karsten, P. A., Fungi Fennici Exsiccati 559, as X. subularis (K[M]), immature. HUNGARIA. in pratis ”Nagymezö” montium ”Bükk hegyseg”, on stems of Aconitum variegatum ssp. gracile, Oct 1957, Tóth, S., Poelt & Scheuer’s Reliquiae Petrakianae 1697, as X. filiformis (S F131039), immature. ITALY. Andalo, on stems of Senecio cordatus ssp. gracile, Aug 1896, Bresadola, G., as X. filiformis (S F131045 ex Bresadola herb.), immature; Bei Paneveggio in Süd-Tyrol, auf faulenden Stengeln von Aconitum u. Cirsium spinossimum, Aug 1882, Arnold, as X. filiformis (S F258794 ex Rehm herb.), immature; on leaves, May 1882, Bresadola, G., as X. filiformis (S F131052 ex Bresadola herb.), immature. NORWAY. Norvegia, Tr., Lyngen, Övre Karnes, kbl. 1634 III, UTM DC 61 NE, in Filipenduleto ulmariae, 3 Aug 1977, Sivertsen, S., as X. filiformis (S F131013), immature. SLOVAKIA. Prencow, in foliis emortuis, 28 Oct 1889, Kmet, A., Fungi Schemnitzienses, as X. filiformis (HBG, S F271423 ex Sydow herb.), immature; Prencow, Mt. Sytno, in caulibus emortius Ononidis spinosae L., 27 Jun 1888, Kmet, A., Fungi Schemnitzienses, as X. filiformis (HBG), immature; Prencow, na háj, in diversis foliis emortuis, 8 Oct 1888, Kmet, A., Fungi Schemnitzienses, as X. filiformis (S F271424), immature. SWEDEN. Bohuslän: Rödbo parish, Ellesbo, on decaying stems, culms and leaves of various plants under Scirpus silvaticus at the edge of a pond, 25 Jul 1942, Nathorst-Windahl, T., Lundell & Nannfeldt’s Fungi Exsiccati Suecici 1459, as X. filiformis (S F130992), immature; Gästrikland, Gävle, Lövudden, on decaying stems of Aegopodium podagraria, 16 Jul 1945, Nannfeldt, J. A. 7747, Lundell & Nannfeldt’s Fungi Exsiccati Suecici 1460, as X. filiformis (S F131023), immature; Jämtland, Åre s:n, Åre, on herbaceous stems of Aconitum septentrionale, 22 Aug 1931, Eliasson, A. G., as X. filiformis (S F131004), immature; Medelpad, Borgsjö par., Granbodåsen chalet, on dead stalks of Urtica dioeca in mixed forest, 11 Sep 1995, Lundqvist, N. 20263, as X. filiformis (S F130991), immature; on dead stem, Swartz, O. (lectotype of Sphaeria stipiticola S F131030 ex Swartz herb.), immature; Skåne, on herbaceous stems, collectors unknown, as Sphaeria filiformis (UPS F-012070 ex Fries herb.), immature; Stockholm, on herbaceous stems, collectors unknown, as X. stipiticola (UPS F-133310 ex Fries herb.), immature; Uppland, Fasterna sn, Grindstugan i Granby, på gamla hallonpinnar, Oct 1988, Anderberg, A., as X. filiformis (S F131009), immature; Uppland, Funbo parish, Storudden, on decaying petioles of Fraxinus excelsior, 5 Jul 1942, Åberg, E., Lundell & Nannfeldt’s Fungi Exsiccati Suecici 1461b, as X. filiformis (S F131027), immature; Uppland, Funbo parish, Storudden, on decaying stems of Urtica dioica, 5 Jul 1942, Åberg, E., Lundell & Nannfeldt’s Fungi Exsiccati Suecici 1461a, as X. filiformis (S F131025), immature; Uppland, Lidingö, Hersbyvägen, på c 10 cm bred stambas av Lonicera tatarica (rosentry), Apr 1985, Bremer 2254, as X. filiformis (S F131019), immature; Uppland, Stocksund stn till gamla Stocksundsbron, på multnade stjälkar, 17 Oct 1952, Berggren, G., as X. filiformis (S F131018), immature; Uppland, Uppsala par., Uppsala, Hort. Bot., on wood, Sep 1853, Fries, E., as X. subularis (UPS F-176043 ex Fries herb.), immature.

Notes Xylaria simplicissima is distributed in temperate regions of Eurasia and North America, where it is commonly associated with dead herbaceous stems and less commonly with fallen leaves. Most herbarium specimens labeled as X. filiformis that were collected from these regions are in fact X. simplicissima. Xylaria filiformis differs from X. simplicissima mainly by having a more delicate stromatal axis and smaller ascospores that possess a hyaline sheath. Xylaria simplicissima is represented by numerous specimens, but immature stromata are more commonly encountered.

An authentic specimen labeled as Rhizomorpha simplicissima (L 0117822) was located in the Persoon herbarium (L), but it has been relabeled as Sphaeria filiformis. This specimen is immature and lacks collecting data. Chaillet collected the original material of R. simplicissima, as indicated in the protologue (Persoon 1822). A specimen (F-004396) labeled as R. simplicissima from the Fries herbarium (UPS) was identified as a Chaillet specimen from “Frankrike” (France in Swedish) on a paper slip enclosed within the packet; it is herein designated as the lectotype. However, this specimen is immature, and a fully mature Finnish collection, from which a culture was obtained, is designated as the epitype herein.

The specimens labeled as S. stipiticola and X. subularis that we studied are immature. The only specimen labeled as X. subularis from the Fries herbarium (UPS) was collected in 1853, after the name was published, and contains material of X. simplicissima. Specimen S F131030 from the Swartz herbarium (S), originally labeled as S. stipiticola and relabeled as X. filiformis, contains stromata with developing perithecia and is considered the holotype of S. stipiticola. The stromatal surface has been ruined by insects at places. A Fries specimen labeled as both X. stipiticola and X. filiformis was collected from Stockholm and has a similar insect-gnawing pattern to that of S F131030, of which it may be a duplicate.

Xylaria simplicissima has a long, entangling history with X. filiformis. Fries (1816) tentatively identified a fungus collected from herbaceous stems in Skåne, Sweden as S. filiformis but was certain that it was the same as S. stipiticola Swartz (Swartz 1814). Later, Fries (1823) divided S. filiformis into two varieties based on substrate types: var. filiformis grew on fallen leaves and petioles, known to him probably only from Albertini and Schweinitz (1805), while var. b, equivalent to S. stipiticola, grew on herbaceous stems. It should be noted that S. stipiticola is a nomen nudum, being published without a description/diagnosis. Var. b was later equated by Fries (1828, 1830) to Rhizomorpha simplicissima Pers. Fries (1849) eventually recognized var. b as a distinct species from X. filiformis [≡ S. filiformis] and named it X. subularis Fr., with R. simplicissima (as “simplex P.”) in synonymy. Xylaria subularis is thus a superfluous name of R. simplicissima.

In North America, X. simplicissima was described as X. filiformis f. caulincola (Rehm 1912), which is based on a Canadian collection made by J. Dearness. Lloyd (1918b) identified a Canadian specimen from Dearness as X. filiformis, with an ascospore size of 16 × 5 μm; this is likely X. simplicissima. The description of X. filiformis in Ellis and Everhart (1887) also indicates X. simplicissima. The protologues of X. jiangsuensis and X. crinalis leave little doubts that they are in synonymy with X. simplicissima.

Xylaria sp. AR1741 Fig. 11H–J.

Stroma cylindrical at fertile part, unbranched, with an acute apex, on a glabrous stipe, 15 mm in total length, 7 mm long × 0.5 mm broad at fertile part; surface blackish brown, slightly polished, with slight perithecial mounds and fine longitudinal striations, lacking an outer layer, underlain with a thin, soft layer ca. 20 μm thick, concolorous with the surface; interior white, homogeneous, soft. Perithecia spherical, 250–350 μm broad. Ostioles conic-papillate, tilting upwards, 100–120 μm high × 100–120 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 90–130 μm total length, the spore-bearing part 85–100 μm long × 9–11 μm broad, with an apical ring staining deep blue in Melzer’s iodine reagent, urn-shaped, 4.5–5.5 μm high × 4–5 μm broad. Ascospores dark brown, unicellular, ellipsoid-inequilateral, with broadly to, less frequently, narrowly rounded ends, smooth, (14–)14.5–16(–16.5) × (6.5–)7–8(–8.5) µm (15.2 ± 0.6 × 7.6 ± 0.6 μm, N = 40), with a straight germ slit spore-length on the ventral side, surrounded with a hyaline sheath swelling at both ends to form cone-shaped non-cellular appendages; epispore smooth.

Specimen examined VENEZUELA. Territorio Federal Amazonas, Cerro de la Neblina, San Carlos de Rio Negro, near airport, on IVIC plot, on dead leaves, 5 Jan 1985, Rossman, A. AR1741-434 (WSP ex Rogers herb.).

Notes This small Venezuelan collection represents an undescribed species but contains only a fragmented stroma. It closely resembles X. appendiculatoides but has cone-shaped secondary appendages at the ascospore ends. Rogers et al. (1988) suspected that this fungus resembles X. lima, which differs by having a tomentose stromatal fertile part and smaller ascospores with a cone-shaped secondary appendage at each end.

Xylaria sp. GS7461A Fig. 7H, J, N, O.

Stromata depressed-capitate at fertile part, unbranched, with an acute apex, on a glabrous stipe, 18–19.5 mm in total length, 1.2–1.4 mm long × 1.6–2.2 mm broad at fertile part; surface dull grayish brown, with conspicuous to half-exposed perithecial mounds, overlain with a dull grayish brown thin layer cracked reticulately into fine plaques 100–200 μm broad, underlain with a black layer ca. 30 μm thick; interior white, homogeneous, soft. Perithecia spherical, 500–600 μm broad. Ostioles papillate, 160–190 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 200–240 μm total length, the spore-bearing part 140–180 μm long × 10.5–13.5 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, urn-shaped, 5.5–6.5 μm high × 4–5 μm broad. Ascospores brown, unicellular, ellipsoid-inequilateral, with narrowly rounded ends, smooth, (24.5–)26–29(–31) × (7.5–)8–9(–9.5) µm (27.7 ± 1.5 × 8.7 ± 0.5 μm, N = 40), with a spiral germ slit spore-length or slightly less than spore-length on the ventral side, lacking a hyaline sheath; epispore smooth.

Specimen examined VENEZUELA. Edo. Trujillo, Parque Nacional Guaramacal, ca. 10 km SW of Batatal, La Defensa, along Río Saguás, Campamiento Granja Bocono, in forest along trail to water source, alt. 2000 m, on petiole of Cecropia (Urticaceae)?, 20, 23 Nov 1990, Samuels, G. J., Hein, B. & Huhndorf, S. M. GS7461A (HAST 145983 ex BPI).

Notes Xylaria sp. GS7461A is characterized by a depressed-capitate stromatal fertile part born on a long glabrous stipe and topped with an acute apex and by a spiral germ slit on the ascospores. Interestingly, X. sp. GS7461A grows on the same petiole as X. meliacearum and X. sp. GS7461B (Fig. 7H), and all three species have a spiral germ slit on their ascospores. This may give the false impression that they are different manifestations of the same species. However, they are significantly different by their combinations of stromatal morphology and ascospore size range. The stromatal fertile part and ascospores of X. meliacearum, X. sp. GS7461A, and X. sp. GS7461B are torulose/(19–)20–23(–27) × (4.5–)5–6 (–6.5) µm, depressed-capitate/(24.5–)26–29(–31) × (7.5–)8–9(–9.5) µm, and cylindrical/(22.5–)23.5–25(–26) × (6–)6.5–7.5(–8) µm, respectively. Xylaria sp. GS7461A, and X. sp. GS7461B remain undescribed due to limited materials, with only two and one stromata available for study, respectively.

Xylaria sp. GS7461B Fig. 7H, K, P, Q.

Stroma cylindrical at fertile part, unbranched, with a mucronate apex, on a glabrous stipe, 5 mm in total length, 3.4 mm long × 1.6 mm broad at fertile part; surface dull brown, with inconspicuous perithecial mounds, overlain with a dull brown thin layer splitting into vague stripes, underlain with a carbonized layer ca. 70 μm thick; interior whitish, homogeneous, soft. Perithecia spherical, 550–650 μm broad. Ostioles papillate, 120–140 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 185–230 μm total length, the spore-bearing part 140–175 μm long × 8–10 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, urn-shaped, 4–6 μm high × 3.5–4.5 μm broad. Ascospores brown, unicellular, ellipsoid-inequilateral to crescentic, with narrowly rounded ends sometimes bearing a tiny hyaline cellular appendage on one end, smooth, (22.5–)23.5–25(–26) × (6–)6.5–7.5(–8) µm (24.2 ± 0.8 × 7.1 ± 0.5 μm, N = 40), with a spiral germ slit spore-length or slightly less than spore-length on the ventral side, lacking a hyaline sheath; epispore smooth.

Specimen examined VENEZUELA. Edo. Trujillo, Parque Nacional Guaramacal, ca. 10 km SW of Batatal, La Defensa, along Río Saguás, Campamiento Granja Bocono, in forest along trail to water source, alt. 2000 m, on petiole of Cecropia (Urticaceae)?, 20, 23 Nov 1990, Samuels, G. J., Hein, B. & Huhndorf, S. M. GS7461B (HAST 145984 ex BPI).

Notes Xylaria sp. GS7461B is reminiscent of species in the X. arbuscula group, where the point-topped stromata are cylindrical and overlain with an outer layer that splits into stripes, but can be separated from the species in the group by its larger ascospores with a spiral germ slit. It grows on a petiole where X. meliacearum and X. sp. GS7461A are also present (Fig. 7H). Also see the notes on X. sp. GS7461A.

Xylaria spiculaticlavata Y.-M. Ju & H.-M. Hsieh, sp. nov. Fig. 9O, U–Y.

MycoBank MB848547.

Typification FRENCH GUIANA. on a fallen leaf, Leprieur, F. R. 1208? (holotype of X. spiculaticlavata PC 0086071 ex Leprieur herb.), mixed with X. phyllocharis and X. aristata.

Etymology Referring to the cylindric-clavate stromatal fertile part studded with conical ostioles, resembling a medieval spiked battle club.

Stroma cylindric-clavate at fertile part, unbranched, with a blunt apex, on a glabrous stipe, 76.6 mm in total length, 10.8 mm long × 2.3 mm broad at fertile part; surface brown, lacking perithecial mounds, lacking an outer layer, underlain with a thin, soft layer ca. 30 μm thick, concolorous with the surface; interior white, homogeneous, soft. Perithecia spherical, 600–800 μm broad. Ostioles coarsely conical, slightly tilting upwards, 240–270 μm high × 270–310 μm broad at base. Asci not intact, with an apical ring staining deep blue in Melzer’s iodine reagent, cylindrical, 7.5–8 μm high × 4–4.5 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid-inequilateral, with broadly to narrowly rounded ends, sometimes with one end slightly pinched, smooth, (22–)23.5–27(–28) × (8.5–)9–10.5(–11) µm (25.1 ± 1.8 × 9.7 ± 0.7 μm, N = 10), with a straight germ slit spore-length or nearly so on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages; epispore smooth.

Notes Xylaria spiculaticlavata is peculiar among foliicolous Xylaria species in having a plane stromatal fertile part studded with conspicuous conical ostioles, perithecia broader than 600 μm, and ascospores longer than 20 μm and broader than 9 μm. Material of X. spiculaticlavata is mixed in the isotype packet of X. phyllocharis from the Leprieur collection (PC), where materials of X. phyllocharis and X. aristata are also enclosed. The materials of these three species are separately glued on three paper cards as shown in Fig. 9O, where the stroma of X. spiculaticlavata is on the right-hand side bearing the pencil writing “Herb. Leprieur 1208?”. Although the material is represented by one stroma only, we formally describe it as new because it possesses a suite of highly distinctive features among the foliicolous Xylaria species and appears to be an extremely rare species having not been recollected since Leprieur made the collection during 1835–1849 (Montagne 1855).

Xylaria vagans Petch, Ann. Roy. Bot. Gard. (Peradeniya) 6: 68. 1915. Fig. 11K–Q.

= Xylaria hedyosmicola H.-X. Ma & X.-Y. Pan, in X.-Y. Pan, Z.-K. Song, Z. Qu, T.-D. Liu & H.-X. Ma, MycoKeys 86: 52. 2022.

Stromata filiform at fertile part, unbranched, with a long acicular apex, on a glabrous stipe, 20–53 mm in total length, 4–11.5 mm long × 0.7–1.1 mm broad at fertile part; surface black, slightly polished, with half-exposed perithecial mounds and fine longitudinal striations, lacking an outer layer, underlain with a thin, soft layer ca. 20 μm thick, concolorous with the surface; interior white, homogeneous, soft. Perithecia spherical, 300–550 μm broad. Ostioles conic-papillate, 90–120 μm high × 120–160 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 125–150 μm total length, the spore-bearing part 75–90 μm long × 6.5–8 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, inverted hat-shaped, 2.5–3 μm high × 2–2.5 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid-inequilateral, with narrowly rounded ends sometimes bearing a tiny hyaline cellular appendage on one end, smooth, (11–)11.5–13(–13.5) × (4.5–)5–6(–6.5) µm (12.1 ± 0.7 × 5.5 ± 0.4 μm, N = 80), with a straight germ slit spore-length on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages; epispore smooth.

Specimens examined SRI LANKA. Central Province, Nuwara Eliya District, Hakgala, elev. 5400 ft, on fallen leaves, Jan 1914, Petch, T. 4093 (holotype of X. vagans K[M] 144168, isotype BPI 713891 ex Lloyd herb. 10383). USA. Hawaiian Islands, Hawaii, Hilo, on dead leaves of Melonchia umbellata, 26 Aug 2000, Hemmes, D. DEH-1977 (WSP ex Rogers herb.); Hawaiian Islands, Hawaii, Leilani Estates, on dead leaves of mango and Melastoma sp., Apr 1996, Hemmes, D. DEH-1091 (WSP ex Rogers herb.); Hawaiian Islands, Hawaii, Leilaris Estates, on dead leaves of Tibouchina semidecandra or soil, 2 Jan 1996, Hemmes, D. DEH-1052 (WSP ex Rogers herb.).

Notes Xylaria vegans is quite similar to X. filiformis in stromatal morphology and ascospore size range. Xylaria filiformis appears to be a temperate counterpart of X. vegans, from which it differs by having light brown, short fusoid-inequilateral ascospores with narrowly rounded ends frequently pinched on one or both ends.

A specimen [SRI LANKA. Central Province, Nuwara Eliya District, Hakgala, elev. 5400 ft, on fallen leaves, Petch, T. 4094, as X. vagans (BPI 713889 ex Lloyd herb. 10386)] that Lloyd received from Petch contains rhizomorphs. However, Petch (1915), while discussing horse-hair blights, mentioned that X. vagans was common in Hakgala on dead leaves but “not necessarily in direct connection with the black rhizomorphs.” An Indian specimen [INDIA. on fallen leaves, Feb 1980, Narula, A. M. H-13 (WSP ex Rogers herb.)] labeled as X. filiformis is similar to X. vagans but has slightly larger ascospores (14–)14.5–15.5(–16.5) × (5.5–)6–6.5(–7) µm (14.9 ± 0.6 × 6.2 ± 0.4 μm, N = 20).

Xylaria vagans was included in Hsieh et al. (2010) as X. sp. 6 from Hawaiian Islands, which was suspected by Rogers and Ju (2012) to be an undescribed species. Xylaria hedyosmicola described by Pan et al. (2022) possesses features of X. vagans, of which we consider a synonym. The synonymy is further corroborated by the high sequence identities at three loci between X. sp. 6 (DEH-1052) and X. hedyosmicola (FCATAS856) from Hainan, China, shown by BLAST searches: 98.99% (GQ478217 vs. MZ221183) at β-tubulin, 99.92% (GQ844795 vs. MZ683407) at RPB2, and 98.66% (GU300082 vs. MZ227121) at ITS.

Xylaria vermiformis Y.-M. Ju, H.-M. Hsieh, sp. nov. Fig. 12A–G.

MycoBank MB848548.

Typification INDIA. Simla, on way to Taradevi Temple, on fallen angiospermous leaves, 9 Sep 1982, Singh, M. 381, as X. cf. sicula (holotype of X. vermiformis WSP ex Rogers herb.).

Etymology Referring to the wavy, worm-shaped stromata.

Stromata filiform at fertile part, unbranched, with a long acicular apex, on a glabrous stipe, 35–83 mm in total length, 2.4–6.3 mm long × 0.5–0.9 mm broad at fertile part; surface black, with half-exposed perithecial mounds, lacking an outer layer, underlain with a thin, soft layer ca. 20 μm thick, concolorous with the surface; interior white, homogeneous, soft. Perithecia depressed-spherical, 250–400 μm broad × 150–250 μm high. Ostioles conic-papillate, 80–120 μm high × 110–140 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 95–125 μm total length, the spore-bearing part 55–70 μm long × 5.5–6.5 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, inverted hat-shaped, 1.5–2 μm high × 1.5–2 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid to shortly fusoid, inequilateral, with narrowly rounded ends, smooth, (9–)9.5–10.5(–11) × (3.5–)4–4.5(–5) µm (10.1 ± 0.5 × 4.4 ± 0.3 μm, N = 40), with a straight germ slit spore-length or nearly so on the ventral side, lacking a hyaline sheath; epispore smooth.

Notes Xylaria vermiformis has half-exposed perithecial mounds arranged on the filiform stromata that terminate into an acicular stromatal apex and thus in general resembles X. filiformis, X. simplicissima, and X. vagans. Xylaria filiformis and X. vagans can be readily separated from X. vermiformis by having a hyaline sheath swelling at both ends to form papillate non-cellular appendages, and X. simplicissima can be separated mainly by having larger ascospores.

Xylaria vittatipiliformis Y.-M. Ju, H.-M. Hsieh & Fournier, sp. nov. Figs. 12M, 13A–G.

MycoBank MB848549.

Typification FRENCH WEST INDIES. Guadeloupe, Basse-Terre, Petit-Bourg, trail to La Mamelle, hygrophilic rainforest, on a dead leave in the litter, 14 Aug 2010, Lechat, C. CLLGUAD029 (cultured) (holotype of X. vittatipiliformis LIP, isotype HAST 145985), GenBank: ITS = OQ883723.

Etymology Referring to the stromata overlain with an outer layer that splits into band-like stripes and terminates into a long hair-like apex.

Stromata cylindrical at fertile part, unbranched, with a long acicular apex, on a glabrous stipe, 61–78 mm in total length, 3–6 mm long × 0.8–1 mm broad at fertile part; surface black, with inconspicuous to conspicuous perithecial mounds, overlain with a grayish brown outer peeling layer split into band-like stripes, underlain with a black layer 40–50 μm thick; interior white, homogeneous, soft. Perithecia spherical, 400–500 μm broad. Ostioles slightly papillate, ca. 70 μm high × 100–150 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, 110–160 μm total length, the spore-bearing part 70–85 μm long × 7–8.5 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, inverted hat-shaped, 2.5 μm high × 2–2.5 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid-inequilateral, with narrowly to broadly rounded ends frequently bearing a tiny cellular appendage on one end, smooth, (10–)11–12(–12.5) × (5.5–)6–7(–7.5) µm (11.4 ± 0.6 × 6.5 ± 0.5 μm, N = 40), with a straight germ slit spore-length or nearly so on the ventral side, lacking a hyaline sheath; epispore smooth.

Cultures and anamorph. Colonies reaching the edge of 9-cm Petri dish in 3 wk, white, velvety, appressed, azonate, with diffuse margins. Reverse uncolored. Stromata and anamorph not produced.

Notes Xylaria vittatipiliformis is characterized by cylindrical stromata topped with a long acicular apex and overlain with an outer peeling layer splitting into band-like stripes. Its stromatal surface is much like that of X. vittiformis, from which X. vittatipiliformis can be readily separated by having larger ascospores that lack a surrounding hyaline sheath. The type material of X. vittatipiliformis was described and illustration as Xylaria sp. CLLGUAD 029 by Fournier et al. (2020).

Xylaria vittiformis Y.-M. Ju & H.-M. Hsieh, sp. nov. Fig. 13H–N.

MycoBank MB848550.

Typification BRAZIL. Rio Grande do Sul, São Leopoldo, in foliis aridis, 1907, Theissen, F., Rick’s Fungi Austro-Americani 353, as X. phyllocharis Mont. (holotype of X. vittiformis HBG, isotype HBG ex Magnus herb.).

Etymology Referring to the band-shaped outer peeling layer on the stromatal surface.

Stromata cylindrical to cylindric-fusoid at fertile part, unbranched, with an acuminate apex, on a glabrous stipe, 7.7–14 cm in total length, 2.3–4.1 cm long × 0.6–0.9 mm broad at fertile part; surface black, with inconspicuous to conspicuous perithecial mounds, overlain with a grayish brown outer peeling layer split into band-like stripes, underlain with a thin, black layer ca. 10 μm thick; interior white, homogeneous, soft. Perithecia spherical, 150–200 μm broad. Ostioles slightly papillate, ca. 30 μm high × 30–50 μm broad at base. Asci with eight ascospores arranged in uniseriate manner, cylindrical, immeasurable due to lacking intact stipes, the spore-bearing part 55–65 μm long × 6–7 μm broad, with an apical ring staining blue in Melzer’s iodine reagent, inverted hat-shaped, 1.5 μm high × 2–2.5 μm broad. Ascospores brown to dark brown, unicellular, ellipsoid-inequilateral, with narrowly rounded ends, smooth, (8–)8.5–10(–11) × (4–)4.5–5(–5.5) µm (9.3 ± 0.6 × 4.8 ± 0.3 μm, N = 40), with a straight germ slit spore-length on the ventral side, surrounded with a hyaline sheath swelling at both ends to form papillate non-cellular appendages; epispore smooth.

Additional specimen examined BRAZIL. Rio Grande do Sul, São Leopoldo, on fallen leaves, Theissen, F., Theissen’s Decades Fungorum Brasiliensium 105, as X. phyllocharis (HBG).

Notes Xylaria vittiformis resembles X. vittatipiliformis in having the outer peeling layer of stromata splitting into band-like stripes but differs from the latter by having smaller ascospores enclosed within a hyaline sheath which forms a non-cellular appendage at each end and the sterile stromatal apex not being acicular or long hair-like. Known material of X. vittiformis was collected by F. Theissen from Brazil and distributed among the exsiccata Fungi Austro-Americani 353 by J. Rick and Decades Fungorum Brasiliensium 105 by F. Theissen as X. phyllocharis. Xylaria phyllocharis lacks an outer peeling layer on stromata and has a vinaceous brown tinge on the stromatal surface.

Identification key to accepted taxa

Species that have a newly proposed name herein or that are likely undescribed are in boldface. This key is constructed mainly for identification purpose and is not intended to suggest the phylogenetic relationships between the treated species.

1.
Perithecia contained in capitate or obconical head or aggregated into clumps along stromata...3 (X. heloidea group).
1′.
Perithecia distributed along stromata, not aggregated into clumps...2.
2.
Stromata filiform; perithecial mounds half-exposed to fully exposed, discrete or loosely packed... 18 (X. filiformis group).
2′.
Stromata cylindrical; perithecial mounds inconspicuous to conspicuous or half-exposed in rare cases, often closely packed...27 (X. phyllocharis).
3.
Ascospores longer than 20 μm long...4.
3′.
Ascospores shorter than 20 μm long...5.
4.
Ascospores (21.5–)22.5–24.5(–26) × (6.5–)7–8(–9) µm, with a straight germ slit, surrounded with a hyaline sheath forming a long, tubular non-cellular appendage at each end...X. axifera.
4′.
Ascospores (24.5–)26–29(–31) × (7.5–)8–9(–9.5) µm, with a spiral germ slit, lacking a hyaline sheath...X. sp. Samuels GS7461A.
5.
Stromatal fertile parts with plane to inconspicuous perithecial mounds...6.
5′.
Stromatal fertile parts with conspicuous to half-exposed perithecial mounds...13.
6.
Ascospores 8–9(–9.5) × 4–4.5(–6.6) µm....X. imminuta.
6′.
Ascospores longer than 10 μm...7.
7.
Perithecia distributed on the top of stromatal fertile parts only...8.
7′.
Perithecia distributed on the entire stromatal fertile parts...9.
8.
Stromatal fertile parts capitate; ascospores (12.5–)13–15(–16) × (7.5–)8–9(–10) µm...X. clusiae.
8′.
Stromatal fertile parts peltate; ascospores 12–14 × 6–8 μm ...X. memecyli.
9.
Stromatal fertile parts globose; ascospores (13–)13.5–15(–16.5) × (6–)6.5–7.5(–8) µm, surrounded with a hyaline sheath swelling at both ends to form non-cellular appendages...X. hypsipoda.
9′.
Ascospores lacking a hyaline sheath...10.
10.
Ascospores (15–)15.5–17(–18) × (6.5–)7.5–9(–9.5) µm...X. hispidipes.
10′.
Ascospores smaller...11.
11.
Ascospores dark brown, nearly semicircular to broadly ellipsoid-inequilateral, with broadly rounded ends to, less frequently, narrowly rounded ends, (10–)10.5–12.5(–14) × (5.5–)6–7(–7.5) µm...X. delicatula.
11′.
Ascospores brown to dark brown, ellipsoid-inequilateral, with narrowly rounded ends...12.
12.
Ascospores (10–)10.5–12.5(–14) × (5.5–)6–7(–7.5) µm...X. aristata var. aristata.
12′.
Ascospores (13.5–)14–16(–17) × (5.5–)6–7(–7.5) µm...X. aristata var. hirsuta.
13.
Ascospores (14.5–)15.5–18(–19) × (5–)5.5–6.5(–7) µm...X. heloidea.
13′.
Ascospores shorter than 15 μm...14.
14.
Stromata with a long acicular apex, much longer than the fertile part...15.
14′.
Stromata with an acute apex, shorter than the fertile part...16.
15.
Ascospores (10.5–)11.5–13.5(–15) × (5–)5.5–6.5(–7.5) µm...X. nainitalensis.
15′.
Ascospores (8.5–)9.5–11(–12) × (4–)4.5–6(–6.5) µm...X. sicula.
16.
Ascospores (12–)12.5–15.5(–17) × (5–)6–7.5(–8) µm...X. amphithele.
16′.
Ascospores shorter...17.
17.
Ascospores (10–)10.5–12(–12.5) × (5–)5.5–6(–6.5) µm...X. pisoniae.
17′.
Ascospores (7.5–)8.5–9.5(–10) × (3.5–)4–4.5(–5) µm...X. petchii.
18.
Ascospores encircled with a hyaline sheath...19.
18′.
Ascospores lacking a hyaline sheath...21.
19.
Ascospores brown to dark brown, 12–13.5 × 4–5 μm, with the hyaline sheath not swelling at ends... X. eugeniae.
19′.
Ascospores with the hyaline sheath swelling at both ends to form non-cellular appendages...20.
20.
Ascospores short fusoid-inequilateral, light brown, (9.5–)11.5–13.5(–14.5) × (4–)4.5–5.5(–6) µm; mainly distributed in northern temperate regions...X. filiformis.
20′.
Ascospores ellipsoid-inequilateral, brown to dark brown, (11–)11.5–13(–13.5) × (4.5–)5–6(–6.5) µm; mainly distributed in the tropics and subtropics...X. vagans.
21.
Stromata hairy on the entire surface; ascospores (11.5–)12–14(–15) × (4.5–)5 μm...X. castilloi.
21′.
Stromata glabrous on the fertile part...22.
22.
Ascospores mostly longer than 16 μm...23.
22′.
Ascospores mostly shorter than 16 μm...24.
23.
Ascospore germ slit straight, (15–)16.5–19(–21.5) × (5–)5.5–6.5(–7.5) µm...X. simplicissima.
23′.
Ascospore germ slit spiral, (19–)21.5–27.5(–31.5) × (5–)5.5–7(–8) µm...X. meliacearum.
24.
Ascospores longer than 11 μm long Ascospores 12–14.5(–16) × 4–4.5(–5) µm...X. duranii.
24′.
Ascospores shorter than 11 μm long...25.
25.
Ascospores (5.5–)6–7 × 3–3.5(–4) µm...X. diminuta.
25′.
Ascospores larger...26.
26.
Stromata slender, with a long acicular apex; ascospores (9–)9.5–10.5(–11) × (3.5–)4–4.5(–5) µm...X. vermiformis.
26′.
Stromata more slender, with an acuminate or mucronate apex; ascospores (9–)9.5–10.5(–11) × (5.5–)6–6.5(–7) µm...X. noduliformis.
27.
Stromatal fertile parts overlain with spikes or a tomentum...28.
27′.
Stromatal fertile parts not overlain with spikes or a tomentum...31.
28.
Stromata overlain with stiff spikes on the surface; ascospores (10.5–)11–12.5(–14.5) × (7–)7.5–8.5(–9) µm, lacking a hyaline sheath...X. asperata.
28′.
Stromata overlain with a tomentum... 29.
29.
Mature stromata overlain with a striped peeling layer; ascospores (10–)10.5–11.5(–12) × (5.5–)6–6.5(–7) µm, lacking a hyaline sheath...X. maitlandii.
29′.
Mature stromata not overlain with an outer layer; ascospores surrounded a hyaline sheath swelling at both ends to form non-cellular appendages...30.
30.
Ascospores (14.5–)15–16.5(–17) × (8.0–)8.5–9.5(–10) µm...X. allima.
30′.
Ascospores (10–)10.5–12(–14) × (5–)6–7(–7.5) µm...X. lima.
31.
Ascospores (22.5–)23.5–25(–26) × (6–)6.5–7.5(–8) µm, with a sigmoid to spiral germ slit...X. sp. Samuels GS7461B.
31′.
Ascospores with a straight to slightly oblique germ slit...32.
32.
Ascospores longer than 16 μm long...33.
32′.
Ascospores shorter than 16 μm long...35.
33.
Ascospores (15–)16.5–18(–19) × (7.5–)8–9(–9.5) µm...X. phyllophila.
33′.
Ascospores longer than 20 μm...34.
34.
Perithecial mounds conspicuous; ascospores light brown to brown, (20.5–)21.5–23.5(–24.5) × (6–)6.5–8(–9) µm...X. luxurians.
34′.
Perithecial mounds plane to inconspicuous; ascospores (22–)23.5–27(–28) × (8.5–)9–10.5(–11) µm...X. spiculaticlavata.
35.
Ascospores 8–10 × 4–6 μm, with a much less than spore-length germ slit ...X. kamatii.
35′.
Ascospores with a spore-length germ slit or nearly so...36.
36.
Mature stromata overlain with a longitudinally split peeling layer...37.
36′.
Mature stromata lacking an outer layer...40.
37.
Stromatal outer peeling layer split into band-like stripes...38.
37′.
Stromatal outer peeling layer split into narrow or thread-like stripes...39.
38.
Stromata terminating into an acuminate sterile apex; ascospores (8–)8.5–10(–11) × (4–)4.5–5 (–5.5) µm, surrounded with a hyaline sheath swelling at both ends to form non-cellular appendages...X. vittiformis.
38′.
Stromata terminating into a long acicular apex; ascospores (10–)11–12(–12.5) × (5.5–)6–7(–7.5) µm, lacking a hyaline sheath, commonly bearing a cellular appendage...X. vittatipiliformis.
39.
Ascospores (8.5–)9–11 × 4–6 μm ...X. foliicola.
39′.
Ascospores (13.5–)14–15(–17) × (4.5–)5–6(–7) µm...X. minuscula.
40.
Ascospores lacking a hyaline sheath...41.
40′.
Ascospores surrounded with a hyaline sheath swelling at both ends...42.
41.
Stromatal surface slightly undulate; ascospores (9–)10–11.5(–12.5) × (4.5–)5.5–6.5(–7) µm...X. phyllocharis.
41′.
Stromatal surface tuberculate; ascospores (9–)9.5–10.5(–11.5) × (5–)5.5–6(–6.5) µm...X. neblinensis.
42.
Ascospores (14–)14.5–16(–16.5) × (6.5–)7–8(–8.5) µm, with cone-shaped non-cellular appendages...X. sp. AR1741.
42′.
Ascospores with papillate non-cellular appendages...43.
43.
Stromatal surface black; ostioles papillate, not tilting upwards; ascospores (11.5–)12.5–14.5(–15) × 5.5–8 μm...X. polysporicola.
43′.
Stromatal surface brown, dark brown to blackish brown; ostioles coarsely conic-papillate, tilting upwards...44.
44.
Stromatal surface tinged vinaceous, with slight perithecial mounds; ascospores (11.5–)12.5–14(–15) × (6–)6.5–7.5(–9) µm...X. appendiculata.
44′.
Stromatal surface not tinged vinaceous, with conspicuous perithecial mounds; ascospores (14–)15–16(–17) × (6.5–)7.0–7.5(–8) µm...X. appendiculatoides.

Discussion

The diversity of leaf- and petiole-inhabiting Xylaria species is mainly found in tropical and subtropical regions except for X. filiformis and X. simplicissima, which are prevalent in the northern temperate regions. Host specificity is unclear for most of these species due to the scarcity of available collections. Xylaria sicula specifically grows on fallen olive leaves primarily in the Mediterranean region. Læssøe and Lodge (1994) considered X. axifera and X. meliacearum collected from Puerto Rico to be specific to the petioles of the plant families Araliaceae and Meliaceae, respectively. However, collections of X. meliacearum from French Guiana and Venezuela seem to indicate a broader host range for the species. Xylaria clusiae and X. pisoniae were named after their hosts Clusia sp. (Samuels and Rogerson 1990) and Eugenia capuli (Rogers et al. 2001), respectively, but it is not yet clear if they are genuinely host-specific. Similar cases are found in the species epithets of X. lindericola, X. hedyosmicola, and X. polysporicola, which reflect the assumption that they prefer fallen leaves of Lindera robusta, Hedyosmum orientale, and Polyspora hainanensis, respectively (Pan et al. 2022).

As outlined in the dichotomous key above, the 45 species are separated into three groups based on stromatal shape, compactness of perithecial aggregation, and conspicuousness of perithecial mounds. Other diagnostic stromatal features include whether the stromatal apex is fertile or sterile, whether an outer stromatal layer persists into maturity as seen in X. foliicola, X. minuscula, X. noduliformis, X. vittatipiliformis, and X. vittiformis, and whether there are overlying spikes as in X. asperata or hairs as seen in X. allima, X. castilloi, X. lima, and X. maitlandii. Long and thread-like stromatal apices are noticeable and can be found in X. filiformis, X. foliicola, X. minuscula, X. nainitalensis, X. sicula, X. simplicissima, X. vagans, X. vermiformis, and X. vittatipiliformis. However, caution should be taken because the long stromatal apices can easily break off as seen in some herbarium specimens. For instance, the type material of X. vagans was described and illustrated as having a long, thin stromatal apex (Petch 1915) but is no longer as such. Fortunately, this feature has seldom been overlooked and can be retrieved by consulting the original descriptions and/or illustrations. Finely longitudinally striated stromatal surfaces that are often slightly polished can be found in X. appendiculata, X. appendiculatoides, X. filiformis, X. polysporicola, X. sp. AR1741, and X. vagans. This feature has diagnostic value and may eventually prove valuable in demonstrating the close relatedness of these species.

Ostioles can be slightly papillate, papillate, conic-papillate or coarsely conic-papillate as seen in species such as X. appendiculata and X. allima. The most salient ostioles is seen in X. spiculaticlavata, where the ostiolar bases are 270–310 μm broad, much broader than those of other species, which are less than 200 μm broad. Ostioles that tilt upwards conspicuously can be found in X. appendiculatoides, X. allima, X. lima, X. sp. AR1741, and X. spiculaticlavata and are useful for identification.

Ascospores are smooth-walled except for X. simplicissima, which has a lumpy appearance on the epispore along the germ slit. The hyaline sheath of ascospores is present in a half of the species, swelling at ascospore ends to form non-cellular appendages that are papillate in most species but tubular in X. axifera and cone-shaped in X. sp. AR1741. The ascospore germ slits are spore-length or nearly so in most species but much less than spore-length in X. kamatii (Pande 1973).

Cultures and/or anamorphs are known from approximately one-quarter of the leaf- and petiole-inhabiting Xylaria species. In the present study, cultures were obtained from five species: X. amphithele, X. aristata var. aristata, X. minuscula, X. simplicissima, and X. vittatipiliformis, but none of them produced anamorphs culture. Cultures have also been obtained from X. axifera (Læssøe and Lodge 1994), X. castilloi (San Martín et al. 1997), X. hispidipes (Rogers et al. 1987) (as X. hypsipoda), X. meliacearum (Læssøe and Lodge 1994), X. phyllocharis (San Martín et al. 1997)d pisoniae (Rogers et al. 2001). Among these species, X. axifera and X. hispidipes did not produce anamorphs in culture, but the anamorph of X. axifera was found on stromata collected from the field. The anamorphs of X. castilloi, X. meliacearum, and X. pisoniae were produced on the stromata produced in culture as well as collected from nature. Cultures have not been obtained from X. asperata, but its anamorph was reported as synnemata on the stromatal surface (Rogers et al. 1988). The anamorph of X. lima was reported by San Martín et al. (1997) (as X. mexicana) on the surface of immature stromata collected from the field.

Conclusion

Most of the known leaf- and petiole-inhabiting Xylaria species are only represented by one or several dried herbarium specimens, indicating the necessity of recollecting them through further intensive collecting activities. DNA extractions from these herbarium specimens are normally unpermitted due to their status as type, historical or authentic specimens, or unsuitable because the specimens are scanty or moldy. Although teleomorphic characteristics are sufficient for recognizing 45 species, additional species may be revealed with data from cultures and/or anamorphs, which are currently quite limited and only available from approximately one-quarter of the species. Molecular phylogenetic studies, such as Hsieh et al. (2010) and Ren et al. (2016), where several leaf- and petiole-inhabiting Xylaria species were included, suggested the incoherence of these species, signifying the possibility of their multiple origins. Many more species are required to form a comprehensive molecular phylogenetic study on leaf- and petiole-inhabiting Xylaria species to reflect their phylogenetic relationships in taxonomy.

Availability of data and materials

Specimens collected by us are deposited at the herbarium HAST. Cultures are available at BCRC. ITS sequences are deposited at GenBank.

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Acknowledgements

This study is dedicated to the late Prof. Jack D. Rogers of Washington State University, Pullman, who was a pioneer in modernizing the taxonomy of the genus Xylaria. He inspired Y-MJ to study Xylaria species on a worldwide basis and collaborated with Y-MJ and H-MH in revising the genus until his passing in 2021. We greatly appreciate Gary Samuels and Jacques Fournier for providing their collections for this study and curators of BO, BPI, C, FH, HBG, K, L, NY, PC, PH, S, UPS, and WSP for locating and loaning specimens.

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This work was supported by Grant MOST 111-2311-B-001-038-MY2 from National Science and Technology Council of Taiwan to Y-MJ.

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Institute of Plant and Microbial Biology, Academia Sinica, Nankang, Taipei, 11529, Taiwan
Yu-Ming Ju & Huei-Mei Hsieh

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Yu-Ming Ju
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Y-MJ and H-MH were responsible for forming the species concepts. Y-MJ recorded morphological traits, obtained cultures, wrote the manuscript, and prepared figure plates. H-MH obtained ITS sequences and deposited them at GenBank. Y-MJ served as the project leader. Both authors read and approved the final manuscript.

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Correspondence to Yu-Ming Ju.

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Ju, YM., Hsieh, HM. Xylaria species associated with fallen leaves and petioles. Bot Stud 64, 19 (2023). https://doi.org/10.1186/s40529-023-00377-w

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Received: 20 March 2023
Accepted: 10 May 2023
Published: 14 July 2023
DOI: https://doi.org/10.1186/s40529-023-00377-w

Xylaria species associated with fallen leaves and petioles

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