Introduction
Trypetheliaceae is an almost entirely tropical, corticolous lichenized family (Aptroot & Lücking Reference Aptroot and Lücking2016). Members of this family were first described by, for example, Zenker (Reference Zenker1829) early in the 19th century from pieces of medicinal bark (mainly Cinchona for quinine) that had been collected in South American forests. However, until recently only a small number of species was known. At the start of the present century, less than 200 species were known in this family (Aptroot & Lücking Reference Aptroot and Lücking2016).
Aptroot & Lücking (Reference Aptroot and Lücking2016) published a revisionary conspectus of the whole family, accepting 418 species, almost a third of which were published simultaneously as new to science (Aptroot & Cáceres Reference Aptroot and Cáceres2016; Aptroot et al. 2016 Reference Aptroot, Ertz, Etayo Salazar, Gueidan, Mercado Diaz, Schumm and Weerakoona , Reference Aptroot, Mendonça, Andrade, Silva, Martins, Gumboski, Fraga Júnior and Cáceresb ; Flakus et al. Reference Flakus, Kukwa and Aptroot2016; Luangsuphabool et al. Reference Luangsuphabool, Lumbsch, Aptroot, Piapukiew and Sangvichien2016; Lücking et al. 2016 Reference Lücking, Nelsen, Aptroot, Benatti, Binh, Gueidan, Gutiérrez, Jungbluth, Lumbsch and Marcellia ). Since then, 14 additional species have been published in the family by Cáceres & Aptroot (Reference Cáceres and Aptroot2016, Reference Cáceres and Aptroot2017), Diederich et al. (Reference Diederich, Lücking, Aptroot, Sipman, Braun, Ahti and Ertz2017), Etayo & Aptroot (Reference Etayo and Aptroot2017) and Soto Medina et al. (Reference Soto Medina, Aptroot and Lücking2017), bringing the total number of species now accepted in the Trypetheliaceae to 432.
Aptroot et al. (2016 Reference Aptroot, Cáceres, Johnston and Lückingc ) predicted the existence of many more species of Trypetheliaceae than considered by Aptroot & Lücking (Reference Aptroot and Lücking2016). The present paper describes a further ten new species from various parts of the world, collected by the authors. Some were gathered subsequent to the revisionary synopsis. Others date from further back and came to light during a revision of identified specimens in the herbarium of the Botanischer Garten und Botanisches Museum in Berlin (B), using the new monograph (Aptroot & Lücking Reference Aptroot and Lücking2016). This revision also revealed many new country records and one new continental record. The countries of origin of the new species are mostly situated in northern South America. This reflects in part a geographical bias in the collections at B but is also in line with the observation that this area is the centre of diversity for this family (Aptroot & Cáceres Reference Aptroot and Cáceres2016).
The generic concept applied here follows the phylogenetic studies by Nelsen et al. (Reference Nelsen, Lücking, Aptroot, Andrew, Cáceres, Rivas Plata, Gueidan, Cañez, Knight and Ludwig2014) and Lücking et al. (2016 Reference Lücking, Nelsen, Aptroot, Barillas de Klee, Bawingan, Benatti, Bungartz, Cáceres, Canêz and Chavesb ). The characters found to be useful for the delimitation of species include: thallus aspect, thickness, colour, smoothness and gall induction as well as the presence/absence, width and colour of prothallus; ascospore colour, septation, shape, size and number per ascus; hamathecium inspersion and colour of inspersion; presence of lichexanthone and variously coloured anthraquinones in or on ascospores and/or ascomata and/or the hamathecium and/or pseudostromata and/or ostioles. Aptroot & Lücking (Reference Aptroot and Lücking2016) provide a more complete treatment and illustration of the characters of these species.
Material and Methods
Identification and descriptive work was carried out in Soest, The Netherlands with an Olympus SZX7 stereomicroscope and an Olympus BX50 compound microscope with interference contrast, connected to a Nikon Coolpix digital camera and, in Berlin, Germany with a Wild M7 stereomicroscope and an Olympus CX41 compound microscope. Sections were mounted in tap water, in which all measurements were also taken.
The chemistry of all specimens was investigated under UV light, and often tested with 10% KOH, generally on sections. The chemistry was further investigated by thin-layer chromatography (TLC) using solvent A (Orange et al. Reference Orange, James and White2001).
Taxonomy
Astrothelium bullatothallinum Aptroot & Sipman sp. nov.
MycoBank No.: MB 827202
Astrothelium which is close to A. aeneum but differs in having a bullate thallus with thick cortex in a mosaic with a grey prothallus.
Type: Venezuela, Bolivar, Cerro Guaiquinima, along Rio Carapo, alt. 800 m, 11 February 1990, H. J. M. Sipman 26957 (B—holotype).
Fig. 1 New Astrothelium species, all from holotypes. A–C, A. bullatothallinum, habitus; D–F, A. cayennense, habitus. Note the two discharged, and now dark, ascospores on the ostiole in Fig. 1F. Scales: A, B, D & E=1 mm; C & F=0·5 mm. In colour online.
Thallus corticate, smooth, bullate, somewhat shiny, discontinuous in a reticulate pattern, covering areas up to 15 cm diam., c. 0·2 mm thick, yellowish orange, with a thick cortex, intermixed with patches of a greyish prothallus, not inducing gall formation in the host bark.
Ascomata globose, 0·3–0·5 mm diam., in groups of 2–15 in lines or in irregular groups, with a brown surface with orange pruina, different from the thallus, distinctly raised above the thallus. Wall dark brown all around, up to c. 70 μm thick. Ostioles apical, not fused, flat to concave, brown. Hamathecium not inspersed with oil globules. Asci with 8 ascospores. Ascospores hyaline, 3-septate, 20–25×6–9 μm, fusiform, ends rounded, IKI−, lumina diamond-shaped, not surrounded by a gelatinous layer.
Pycnidia not observed.
Chemistry. Thallus UV−, ascomata UV+ red, yellow pigment on ascomata KOH+ blood red. TLC: an anthraquinone, probably parietin.
Etymology. The name refers to the bullate thallus.
Distribution and ecology. On tree bark in tropical rainforest. So far known only from Venezuela.
Discussion. This species is close to A. flavostromatum Aptroot & M. Cáceres (Aptroot & Cáceres Reference Aptroot and Cáceres2016) and A. kunzei (Fée) Aptroot & Lücking (Aptroot & Lücking Reference Aptroot and Lücking2016), which differ by their essentially green thalli without anthraquinones, and also to A. aeneum (Eschw.) Aptroot & Lücking (Aptroot & Lücking Reference Aptroot and Lücking2016), which differs mainly by the continuous, non-bullate thallus that is not in a mosaic with a grey prothallus and the ascomata that are not in distinct groups.
Astrothelium cayennense Aptroot & Sipman sp. nov.
MycoBank No.: MB 827203
Astrothelium similar to A. flavomegaspermum but with yellow pigment in the pseudostroma near the ostioles.
Type: French Guiana, Montsinery, c. 20 km W of Cayenne, “Risque tout” forest track, alt. 50 m, March 1985, A. Aptroot 15125 (B—holotype; ABL—isotype).
Thallus corticate, smooth, shiny, covering areas up to 5 cm diam., c. 0·2 mm thick, ochraceous green, not surrounded by prothallus, inducing gall formation in the host bark as numerous c. 3–5 mm wide hemispherical warts.
Ascomata pyriform, 0·8–1·3 mm diam., solitary, completely immersed in pseudostromata which are c. 1·5–2·5 mm wide, mostly covered by the thallus cortex except for an area around the ostiole c. 0·5 mm wide. Pseudostromata with dark yellow pigment beneath the cortex, especially near the ostiole. Wall carbonized all around, up to c. 80 μm thick. Ostioles apical, simple, flat, dark brown, surrounded by a non-corticated pale area c. 0·4 mm wide. Hamathecium inspersed with yellow oil globules. Asci with 4 ascospores. Ascospores hyaline, muriform, 295–330×35–40 µm, ellipsoid, IKI−, without distinctly thickened median septum, not surrounded by a gelatinous layer.
Pycnidia not observed with certainty, although some of the many black dots around the ostioles might represent young pycnidia.
Chemistry. Thallus and ascoma UV−, yellow pigment in pseudostroma KOH+ violet, yellow pigment in hamathecium unchanged in KOH.
Etymology. Named after the collecting locality of the type, Cayenne.
Distribution and ecology. On tree bark in tropical rainforest. So far known only from French Guiana.
Discussion. This species is most similar to A. flavomegaspermum Aptroot & Etayo (Etayo & Aptroot Reference Etayo and Aptroot2017) but has a yellow pigment in the pseudostromata near the ostioles. It also resembles A. meristosporum (Mont. & Bosch) Aptroot & Lücking by the large perithecia and the large, muriform ascospores, and the type specimen of A. cayennense was erroneously considered as a neotropical representative of A. meristosporum, a palaeotropical species (Aptroot & Lücking Reference Aptroot and Lücking2016). Astrothelium cayennense differs by the absence of lichexanthone in the thallus, the presence of a yellow pigment in the pseudostromata, and the larger ascospores without a thickened median septum or median constriction.
Astrothelium diaphanocorticatum Aptroot & Sipman sp. nov.
MycoBank No.: MB 827205
Astrothelium with a bullate thallus with thick hyaline cortex and 3-septate ascospores, 25–28×10–12 μm.
Type: Papua New Guinea, Simbu, Mount Wilhelm, along track from Keglsugl to Pindaunde Valley, alt. 2900 m, 3–8 August 1992, H. J. M. Sipman 35615 (B—holotype).
(Fig. 2)
Fig. 2 Astrothelium diaphanocorticatum, holotype. A–D, habitus of thallus warts; E & F, sections through a thallus wart. Scales: A=1 mm; B–D=0·5 mm; E & F=0·1 mm. In colour online.
Thallus olive-green to yellowish green, bullate, partly made up of almost spherical globules of c. 1 mm diam., with a thick, vitreous cortex where the algal cells can be clearly seen as groups of green dots from above, not surrounded by a prothallus, not inducing gall formation in the bark.
Ascomata globose, 0·7–1·3 mm diam., superficial, solitary or a few fused laterally, dark brown to black, not covered by the thallus, not in pseudostromata or pseudostromata black and not well developed. Wall carbonized, up to c. 90 μm thick. Ostioles apical, black, flat. Hamathecium not inspersed. Asci with 8 ascospores, c. 120–150×16–20 μm. Ascospores hyaline, 3-septate, 25–28×10–12 μm, ellipsoid, IKI−, lumina diamond-shaped, not surrounded by a gelatinous layer.
Pycnidia not observed.
Chemistry. Thallus and ascoma UV−, KOH−. TLC: no secondary substances detected.
Etymology. The name refers to the translucent cortex.
Distribution and ecology. On tree bark in montane forest. So far known only from Papua New Guinea.
Discussion. Astrothelium diaphanocorticatum is rather unique within the genus in that the thallus consists of almost spherical globules of c. 1 mm diam., with a thick hyaline cortex so transparent that the algal cells can be clearly seen under the stereomicroscope as groups of green dots. The unusually thick, vitreous cortical layer is at variance with what is usually found in lichens where the photobionts are sheltered from direct insolation by an opaque fungal cortex. In habitats with higher levels of UV-radiation (see e.g. Bjerke et al. Reference Bjerke, Lerfall and Elvebakk2002), such as high mountains and high latitudes, these lichens tend to contain higher concentrations of cortical pigments. Astrothelium diaphanocorticatum, however, grows in montane forest in Papua New Guinea at 2900 m altitude, where the photobionts apparently have no need of this protection and they are found under the dense canopy in a usually misty environment. The curved, glassy cortex of the globules may serve to focus the scarce light onto the photobionts. The function of a glassy cortical layer to focus light onto the photobionts of lichens is better known in the so-called window lichens (e.g. Kilias Reference Kilias1984; Timdal Reference Timdal2017).
Somewhat similar, bullate thalli occur in several Astrothelium species of very humid habitats, such as A. bullatum Flakus & Aptroot, A. komposchii Aptroot, A. megacrypticum Lücking et al., A. papillosum (P. M. McCarthy) Aptroot & Lücking, A. puiggarii (Müll. Arg.) Aptroot & Lücking, A. rimosum Aptroot, A. simplex Aptroot & S. M. Martins and A. tetrasporum Aptroot & M. Cáceres. All of these species differ by having much larger, >50 μm long, pluriseptate or muriform ascospores, and a cortex that is not as thick or clear.
Additional specimen examined. Papua New Guinea: Morobe, Saruwaged Range, near Honzeukngon Village S of Derim in Timbe Valley, alt. 2500 m, 1987, A. Aptroot 18070 (ABL).
Astrothelium macroeustomum Aptroot & Sipman sp. nov.
MycoBank No.: MB 827207
Astrothelium with joint lateral ostioles, a UV+ yellow ostiolar region and 5-septate ascospores of 50–55 ×12–17 μm.
Type: French Guiana, Saül, along piste to Crique Limonade, alt. 300 m, 13 January 1988, H. J. M. Sipman 31769 (B—holotype).
Fig. 3 New Astrothelium species, all from holotypes. A & B, A. macroeustomum, habitus. C & D, A. palaeoexostemmatis; C, habitus; D, ascus, the spores stained blue by IKI. E, A. quasimamillanum, habitus. Scales: A & C=1 mm; B & E=0·5 mm; D=50 μm. In colour online.
Thallus corticate, mostly smooth, somewhat shiny, continuous, covering areas up to 9 cm diam., under 0·1 mm thick, pale ochraceous brown, surrounded by a dark brown prothallus ≤ 0·5 mm wide, not inducing gall formation in the host bark.
Ascomata pyriform, c. 0·6–1·0 mm diam., mostly 2–5 aggregated, mostly immersed in the bark tissue below the thallus. Wall carbonized, up to c. 80 μm thick. Ostioles eccentric, fused, strongly convex, black, surrounded by a yellowish white pruinose ring of 0·3–0·5 mm diam. Hamathecium not inspersed with oil globules. Asci with 8 ascospores. Ascospores hyaline, 5-septate, 50–55×12–17 μm, fusiform, ends pointed, IKI− or very pale blue, not surrounded by a gelatinous layer.
Pycnidia not observed.
Chemistry. Thallus and ascoma UV−, KOH−, ostiole UV+ yellow. TLC: lichexanthone.
Etymology. The name refers to the large ascospores and the resemblance to A. eustomum (Mont.) Müll. Arg.
Distribution and ecology. On tree bark in tropical rainforest. So far known only from French Guiana.
Discussion. This is yet another member of the A. eustomum group which has whitish, UV+ yellow ostiolar regions strongly contrasting with the thallus. Within this group, A. macroeustomum is characterized by large, 5-septate ascospores. Thus, it is closest to A. eumultiseptatum Aptroot & M. Cáceres (Aptroot & Cáceres Reference Aptroot and Cáceres2016), which differs by having longer (>65 μm) ascospores with more (9–11) septa.
Additional specimen examined. French Guiana: same data, H. J. M. Sipman 31757 (B).
Astrothelium minicecidiogenum Aptroot & Sipman sp. nov.
MycoBank No.: MB 827209
Astrothelium with muriform ascospores 70–90×20–25 μm, without pseudostromata, with solitary ascomata, lateral ostioles and an inspersed hamathecium.
Type: Costa Rica, Alajuela, Volcán Tenorio National Park, surroundings of Pilón Biological Station, alt. 700 m, 16 March 2004, A. Aptroot 60498 (INB—holotype).
(Fig.: see Aptroot et al. (Reference Aptroot, Lücking, Sipman, Umaña and Chaves2008), page 56, Fig. 7C & D)
Thallus corticate, smooth, shiny, bullate, covering areas ≤7 cm diam., c. 0·2 mm thick, olive-green, not surrounded by prothallus, not inducing gall formation in the host bark.
Ascomata pyriform, 0·7–1·3 mm diam., solitary, fully immersed in the bark, below the thallus cortex, not in pseudostromata. Wall carbonized all around, up to c. 80 μm thick. Ostioles lateral, simple, flat, black. Hamathecium inspersed with oil globules. Asci with 8 ascospores. Ascospores hyaline to somewhat ochraceous, muriform, 70–90×20–25 μm, fusiform, IKI−, without a distinctly thickened median septum, not surrounded by a gelatinous layer.
Pycnidia not observed.
Chemistry. Thallus and ascoma UV−, KOH−. TLC: no secondary substances detected.
Etymology. The name refers to the small ascospores and the resemblance to A. cecidiogenum (Aptroot & Lücking) Aptroot & Lücking.
Distribution and ecology. On tree bark in tropical rainforest. So far known only from Costa Rica.
Discussion. This material keys out as A. puiggarii in group 8, couplet 5 and is described under that name on page 881 in Aptroot & Lücking (Reference Aptroot and Lücking2016). It was described previously in Aptroot et al. (Reference Aptroot, Lücking, Sipman, Umaña and Chaves2008) on page 53 and the material from Costa Rica was illustrated in Fig. 7C & D on page 56. However, in the original description (Müller Reference Müller1883) and in some additional material examined from Brazil and French Guiana, A. puiggarii has two ascospores per ascus, 170–340×30–60 µm. Therefore the 8-spored specimens are described here as a separate species.
Astrothelium palaeoexostemmatis Sipman & Aptroot sp. nov.
MycoBank No.: MB 827211
Astrothelium similar to A. exostemmatis but with larger, I+ blue ascospores.
Type: Thailand, Chiang Mai, Doi Suthep, Kings Palace, 18°49'N, 99°53'E, alt. 1550 m, oak/chestnut forest, 19 December 1991, P. A. Wolseley & B. Aguirre-Hudson 5771 (B 60 0173986—holotype; BM—isotype).
Thallus corticate, smooth, shiny, covering areas over 5 cm diam., endophloeodic, c. 200 μm thick, ochraceous brown (25 years in herbarium), with c. 50 μm thick cortex, prothallus unknown (thallus margin not present), not inducing gall formation in the host bark.
Ascomata subspherical, 0·4–0·6 mm diam., simple, solitary, dense and in part contiguous but not forming distinct pseudostromata, covered by the thallus and with only the black ostiole visible, sometimes a larger part of or the complete perithecium is visible as a black spot through the transparent thallus cortex. Wall dark brown all around, KOH+ greenish black, c. 40 μm thick, the inner 20 μm darkest. Ostioles apical, black, surrounded by an area of pale thallus c. 0·4 mm wide. Hamathecium clear. Asci with 8 ascospores, c. 300×50 µm. Ascospores hyaline, muriform, 85–100×20–24 µm, composed of c. 26×4 loculi, ellipsoid, IKI+ blue, with rather thin septa, without a distinctly thickened median septum, not surrounded by a gelatinous layer.
Pycnidia not observed.
Chemistry. Thallus and ascoma UV−, no pigment visible.
Etymology. The name refers to the occurrence in the Palaeotropics and the similarity with the neotropical species A. exostemmatis (Müll. Arg.) Aptroot & Lücking.
Distribution and ecology. On tree bark in montane tropical forest. So far known only from northern Thailand.
Discussion. This species superficially resembles a Pyrenula sp. but the hamathecium and ascus structure, and the hyaline ascospores, place it clearly in the Trypetheliaceae. It is most similar to A. exostemmatis in its immersed, single, simple perithecia and 8 per ascus muriform ascospores without a thickened central septum. However, it has larger, I+ blue ascospores and occurs in the Palaeotropics rather than the Neotropics.
Muriform, I+ blue or violet ascospores are an uncommon feature in Astrothelium but Aptroot & Lücking (Reference Aptroot and Lücking2016) mention five species: A. amylosporum Flakus & Aptroot, A. auratum (R. C. Harris) Aptroot & Lücking, A. bullatum Flakus & Aptroot, A. sanguinarium (Malme) Aptroot & Lücking and A. sanguineoxanthum Aptroot with this characteristic. Observations of specimens in the lichen herbarium at B show that A. subdisjunctum (Müll. Arg.) Aptroot & Lücking also has I+ blue ascospores. All these species share simple, not astrothelioid or cryptothelioid, perithecia which may be aggregated in pseudostromata or not, and ascospores without a more strongly thickened median septum. However, A. auratum, A. sanguinarium and A. sanguineoxanthum differ by their orange or red pigmentations; A. amylosporum is distinguished by its large perithecia which are not covered by the thallus; A. bullatum differs by its bullate thallus and large ascospores; A. subdisjunctum generally has 4-spored asci.
Astrothelium quasimamillanum Aptroot & C. Mendonça sp. nov.
MycoBank No.: MB 827213
Astrothelium with muriform ascospores of 30–33×9·5–10·5 μm, without pseudostromata, with solitary ascomata, lateral ostioles and an inspersed hamathecium.
Type: Brazil, Rondônia, Porto Velho, Parque Natural Municipal, alt. 100 m, 2015, C. Mendonça ISE 23813 (ISE—holotype; ABL—isotype).
Fig. 4 New species of Astrothelium and Pseudopyrenula, all from holotypes. A, A. quasimamillanum, habitus; B & C, A. studerae, habitus; D & E, A. tanianum, habitus; F, P. miniflavida, habitus. Scales: A, C, E & F=0·5 mm; B & D=1 mm. In colour online.
Thallus corticate, smooth, shiny, covering areas ≤7 cm diam., less than c. 0·1 mm thick, dark brown, surrounded by a black line denoting the prothallus c. 0·4 mm wide, not inducing gall formation in the host bark.
Ascomata pyriform, 0·6–0·9 mm diam., solitary, mostly immersed in the bark, but just discernible as black structures below the thallus cortex, not in pseudostromata. Wall carbonized all around, up to c. 80 μm thick. Ostioles lateral, simple, flat, black. Hamathecium inspersed with oil globules. Asci with a very wide ocular chamber, of more than half the width of the ascus, with 8 ascospores. Ascospores hyaline, muriform, 10–12×1–2-septate, 30–33×9·5–10·5 μm, ellipsoid, IKI−, without a distinctly thickened median septum, not surrounded by a gelatinous layer.
Pycnidia not observed.
Chemistry. Thallus and ascoma UV−, KOH−. TLC: no secondary substances detected.
Etymology. The name refers to the superficial resemblance to Pyrenula mamillana (Ach.) Trevis.
Distribution and ecology. On tree bark in tropical rainforest. Known only from Brazil at present.
Discussion. This species looks externally very much like a Pyrenula of the mamillana group but has hyaline ascospores and asci of the Trypetheliaceae type with a very wide ocular chamber. It is furthermore characterized by the combination of the following characters: muriform ascospores of 30–33×9·5–10·5 μm, lacking a pseudostroma, a lateral ostiole and an inspersed hamathecium. There are no similar species in Trypetheliaceae.
Astrothelium studerae Aptroot & M. Cáceres sp. nov.
MycoBank No.: MB 827214
Astrothelium with astrothelioid ascomata, lichexanthone only in the pseudostromata, and 3-septate ascospores of 21·5–23·0×6·5–7·5 μm.
Type: Brazil, Alagoas, Quebrangulo, Pedra Talhada private area, alt. 500–700 m, 21–23 October 2017, M. E. S. Cáceres & A. Aptroot ISE 42813 (ISE—holotype; ABL—isotype).
Thallus olive-green to yellowish green, bullate, surrounded by a black line denoting the prothallus c. 0·3 mm wide, not inducing gall formation in the host bark.
Ascomata pyriform, 0·7–1·1 mm diam., 2–5 aggregated with a common ostiole, black, in often somewhat unclear, thallus-like but paler to whitish, slightly raised pseudostromata. Wall carbonized, ≤c. 70 μm thick. Ostioles apical, brown, flat, c. 0·1 mm wide, often surrounded by a wide area where the pseudostromatal tissue is abraded and the black ascoma wall exposed. Hamathecium inspersed with oil globules. Asci with 8 ascospores. Ascospores hyaline, 3-septate, 21·5–23·0×6·5–7·5 μm, ellipsoid, IKI−, lumina diamond-shaped, surrounded by a gelatinous layer up to 7 μm thick.
Pycnidia not observed.
Chemistry. Thallus UV−, KOH−, pseudostromata UV+ yellow, KOH−. TLC: lichexanthone.
Etymology. Named in honour of Anita Studer, the protector of the isolated patch of Atlantic rainforest in Alagoas where the new species was collected.
Distribution and ecology. On tree bark in low montane forest. So far known only from Brazil.
Discussion. This species is similar to A. interjectum R. C. Harris (Aptroot & Lücking Reference Aptroot and Lücking2016) but differs by the inspersed hamathecium.
Astrothelium tanianum Aptroot & Sipman sp. nov.
MycoBank No.: MB 827215
Astrothelium with a bullate thallus, solitary ascomata, covered by thallus, and (9–)11(–15)-septate ascospores, 75–100×20–22 μm.
Type: Malaysia, Johor, Gunung Pulai Forest Reserve, alt. 150 m, 26 November 2000, H. J. M. Sipman, B. Tan & Farida 46414 (B—holotype).
Thallus corticate, bullate, shiny, covering areas of ≤3 cm diam., c. 0·2 mm thick, pale olive-green, not surrounded by a prothallus, not inducing gall formation in the host bark.
Ascomata globose, 0·7–1·3 mm diam., solitary, not in pseudostromata, but immersed in thallus verrucae, which can have a rather irregular surface and outline. Wall black all around, up to c. 60 μm thick. Ostioles flat, apical to often somewhat eccentric, 0·1 mm diam., black. Hamathecium inspersed with oil globules. Asci with 8 ascospores. Ascospores hyaline, (9–)11(–15)-septate, 75–100×20–22 μm, fusiform, ends pointed, IKI−, lumina diamond-shaped, surrounded by a 1 μm thick gelatinous layer.
Pycnidia not observed.
Chemistry. Thallus and ascoma UV−, KOH−. TLC: no secondary substances detected.
Etymology. Named in honour of one of the collectors, the late Philippine bryologist Benito Tan.
Distribution and ecology. On tree bark in tropical lowland rainforest. So far known only from Malaysia.
Discussion. This species is characterized by the bullate thallus, which covers the solitary ascomata, and (9–)11(–15)-septate, 75–100×20–22 μm ascospores. It resembles species of the A. annulare group, such as A. macrosporum (Makhija & Patw.) Aptroot & Lücking (Aptroot & Lücking Reference Aptroot and Lücking2016), but lacks the annulus surrounding the ostiole.
Pseudopyrenula miniflavida Aptroot & A. D. Nunes sp. nov.
MycoBank No.: MB 827216
Pseudopyrenula with a yellow inspersed hamathecium, inspersion dissolving in KOH (not changing in colour) and 3-septate ascospores, 15–17×5·5–6·5 μm.
Type: Brazil, Sergipe, Quissamãa, Mata do IFS, alt. 50 m, 5 September 2016, A. D. Nunes ISE 41583 (ISE—holotype).
Thallus whitish grey, not corticate.
Ascomata hemispherical, solitary, erumpent, black but often partly white pruinose. Ostioles apical. Hamathecium inspersed with yellow oil droplets, which dissolve but do not change colour in KOH. Ascospores 3-septate, 15–17×5·5–6·5 μm.
Pycnidia not observed.
Chemistry. Thallus and ascoma UV−, KOH−, yellow pigment in hamathecium dissolving in KOH. TLC: no secondary substances detected.
Etymology. The name refers to the small ascospores and the yellow oil droplets in the hamathecium.
Distribution and ecology. On tree bark in Atlantic rainforest remnant. So far known only from Brazil.
Discussion. This species is characterized by ascospores of 15–17×5·5–6·5 μm and a hamathecium with yellow oil droplets dissolving, but not changing colour, in KOH. It does not closely resemble any known species in Pseudopyrenula as it has the smallest ascospores in the genus and an apparently unique pigment in the hamathecium.
New Country Records
For each accepted species in the Trypetheliaceae, Aptroot & Lücking (Reference Aptroot and Lücking2016) listed the countries from which those species had been reported in the literature, and gave specimen citations from additional countries. Here we report additions to these lists. Some of these are in species groups where the species concept in Aptroot & Lücking (Reference Aptroot and Lücking2016) is much narrower than that previously used. In cases where the collective species had already been reported from a country, the record given here is marked with #, signifying that the species has been reported from that country in a much wider sense under a different name. All pertinent material would need to be checked to ascertain if it belongs to the currently accepted species reported herein.
Astrothelium aenascens Aptroot # (previously included within the concept of A. aeneum s.l.). Colombia: Caqueta, Araracuara, along road to airstrip, 0°36'S, 72°26'W, alt. 300 m, epiphyte on small tree in secondary vegetation and gardens, 1988, Sipman 27825 & Duivenvoorden (B).
Astrothelium amazonum (R. C. Harris) Aptroot & Lücking. Ecuador: Prov. Zamora-Chinchipe, Reserva Biológica San Francisco, 30 km E of Loja on road to Zamora, transect 2, 3°58·3'S, 79°4·7'W, alt. 2000 m, montane tropical rainforest, on bark, 2001, Nöske & Sipman 126 (B).
Astrothelium chrysoglyphum (Vain.) Aptroot & Lücking. Dominica: Pointe Round, Parish of St. Peter, alt. c. 30 m, dry scrub woodland, 1963, Imshaug 33485 (B).
Astrothelium degenerans (Vain.) Aptroot & Lücking. Guyana: Rupununi Distr., Marudi Mts, c. 1 km along trail from NorMan Mines camp to Aishalton, 2°15'N, 59°10'W, alt. 300–400 m, in disturbed vegetation on lateritic clay, growing on bark of Jacaranda copaia, 1982, Stoffers, Görts-van Rijn, ter Welle & Bonsen 251e (B).
Astrothelium heterophorum Nyl. Vanuatu: Espiritu Santo, logging area near Lavatmas (N of Sara), 48 km NNW of Luganville, 15°7'S, 167°1'E, alt. 300 m, poor lowland forest on Endospermum medullosum, Antiaris toxicaria & Pometia pinnata-dominated flats, upper branches of large felled tree (Endospermum medullosum), 1998, Streimann 62787 & Ala (B).
Astrothelium inspersaeneum E. L. Lima et al. # (previously included within the concept of A. aeneum s.l.). Colombia: Santander, municipality of Charala, corregimiento Virolín, Cañaverales, near finca St. Helena, 6°05'N, 73°12'W, alt. 1900 m, on little trees in pasture fields around farm in mountain valley, 1988, Sipman 27555 & Aguirre (B).—Ecuador: Prov. Zamora-Chinchipe, Cordillera Numbala, Reserva Biologica San Francisco, S of road Loja-Zamora, transecto 1, near Refugio, 3°58'S, 79°04'W, alt. 2450 m, on shrub branches in shrubby forest on mountain ridge, 2003, Sipman 51458 (B).—Malaysia: State of Sabah, Ranau Distr., Kinabalu Park, S-slope of Mount Kinabalu, surroundings of Headquarters, 6°05'N, 116°35'E, alt. c. 1650 m, tall forest on slope along Kiau View trail, near Ranau highway, from trunk and crown branches of felled tree, 1989, Sipman 31459 & Tan (B). New to the Asian continent.
Astrothelium leioplacum (Müll. Arg.) Aptroot & Lücking. Guyana: Upper Takutu District, c. 35 km S of Aishalton, c. 4 km N of Kuyuwini Landing, along track to Karaudanawa, c. 2°08'N, 59°15'W, alt. c. 250 m, epiphyte on scattered shrubs and trees both along the edge of and within small savannah, 1992, Sipman 57079 (B).
Astrothelium meiophorum (Nyl.) Aptroot & Lücking # (previously included within the concept of A. nitidiusculum s.l.). Singapore: Campus of National University of Singapore, around Kent Ridge, 1°18'N, 103°45·5'E, alt. c. 50 m, parkland with scattered buildings and roads, on c. 20–30 cm diam. trunk of small Cassia fistula tree, 2000, Sipman 45532 (B), 45538 (B); ibid., Botanic Gardens, 1°18'N, 103°48'E, alt. c. 50 m, parkland with scattered trees and shrubs, on c. 30 cm diam. tree trunk of Podocarpus gracilior (no. 5389, planted 1932), 2000, Sipman 45710 (B); ibid., Botanic Gardens, Plumeria Garden, 1°18'N, 103°48'E, alt. c. 50 m, parkland with scattered trees and shrubs, on trunks and branches of Plumeria, 2000, Sipman 45747 (B).
Astrothelium meristosporum (Mont. & Bosch) Aptroot & Lücking. Vanuatu: Espiritu Santo, Big Bay-Luganville Road, 26 km NW of Luganville, 15°19'S, 167°01'E, alt. 380 m, regrowth on limestone, Alphitonia, Macaranga-dominated, on semi-exposed Macaranga stem, 1998, Streimann 62268 & Ala (B).
Astrothelium nicaraguense Lücking et al. Costa Rica: Puntarenas, near Las Cruzes Garden (district of Coto Brus), c. 4 km SSE of San Vito, c. 8°43'N, 82°57'W, alt. c. 1300 m, premontane rainforest zone, on trunk of isolated tree on exposed hill ridge, 1978, Sipman 11962 (B); ibid., Parque Internacional La Amistad (AC Amistad Pacifico), Cerro Biolley, 30 km NNW of San Vito near Biolley, lower trail from road to Sabanas Esperanza, 9°04'N, 83°03'W, alt. 1300–1400 m, upland savannah zone, disturbed savannah vegetation with abundant shrubs and trees, on bark (lower trunk), 2002, Sipman 48111 (B).
Astrothelium nigratum (Müll. Arg.) Aptroot & Lücking # (previously included within the concept of A. nitidiusculum s. lat.). Colombia: Caqueta, 2·5 km NE of Araracuara, 0°37'S, 72°23'W, alt. 250 m, c. 30 m tall, hardly disturbed forest on low terrace of Caquetá River (parcelas de Marcela Torres), on canopy branches of Brosimum, 1988, Sipman 27907& Duivenvoorden (B).
Astrothelium nigrum Aptroot & M. Cáceres. Venezuela: Estado Bolivar, Cerro Guaiquinima, in central part of upper plateau (near camp 4), 5°40'N, 63°34'W, alt. c. 950 m, low, mossy forest on rocky slope towards stream, epiphytic, 1990, Sipman 26617 (B, VEN).
Astrothelium nitidiusculum (Nyl.) Aptroot & Lücking. Philippines: Luzon Island, Prov. Sorsogon, Irosin, alt. c. 250 m, on number 14650 (Gymnartocarpus woodii Merr.), 1915, Elmer 14659 (B).
Astrothelium norisianum Lücking et al. Mexico: Chiapas, municipality of Ocozocoautla, c. 5 km antes del aéropuerto San Juan en la carretera 190, Tuxtla Gutiérrez-México, 16°45·0'N, 93°16·5'W, alt. 1200 m, región Depresion Central, bosque caducifolio bajo seco sobre roca calcárea, epífita, alt. 0–2 m, 1994, Wolf & Sipman 2255 (B).—Costa Rica: Limón, 40 km S of Limón, Reserva Biológica Hitoy Cerere, near biological station, pasture fields along entrance road, 9°40'N, 83°02'W, alt. 130 m, submontane rainforest zone, on trunk of scattered trees in pastureland, 2004, Sipman 51612 (B); ibid., Puntarenas, San Vito de Coto Bruz, Estación Biológico Las Cruzes, 8°47·1'N, 82°57·6'W, alt. c. 1200 m, trail to Rio Java, on branches of fallen Ficus pertusa on fallen trunk, 2004, Sipman 53235 (B).
Astrothelium perspersum Aptroot & Ertz. Venezuela: Edo. Merida, Distr. Sucre, finca Los Topos (San Juanito), Chiguará, 8°31·5'N, 71°20'W, alt. c. 1200 m, on bark of fence post between meadow and road (former montane mossy forest area), 1996, Sipman 38061 & Morales (B).
Astrothelium phlyctaena (Fée) Aptroot & Lücking # (previously included within the concept of Trypethelium ochroleucum (Eschw.) Nyl. s. lat.). Venezuela: Estado Bolivar, Cerro Guaiquinima, in central part of upper plateau, along Rio Carapo (near camp 3-nuevo), 5°49'N, 63°32'W, alt. c. 800 m, on trees on rocky slope with low forest in deep clefts and along the river, 1990, Sipman 26971 (B).
Astrothelium robustum Müll. Arg. Mexico: Chiapas, municipality of La Trinitaria, Parque Nacional Lagunas de Montebello, Paso del Soldado, 16°07·1'N, 91°43·1'W, alt. 1500 m, Pinus maximinoi and Quercus sapotifolia forest, N exposure , epiphyte, alt. 0–2 m, 1994, Wolf & Sipman 2083 (B).—French Guiana: Saül, tropical primary forest along piste to Crique Limonade, about halfway, 3°35'N, 53°13'W, alt. c. 300 m, on trunk and branches of fallen tree, 1988, Sipman 31764 (B).
Astrothelium rufescens (Müll. Arg.) Aptroot & Lücking # (previously included within the concept of A. nitidiusculum s. l.). Costa Rica: Puntarenas, Fila Cedro (AC Amistad Pacifico), Las Alturas station, 25 km NE of San Vito near Alturas, surroundings of station and trail into forest, 8°57'N, 82°50'W, alt. 1600 m, montane rainforest zone, pasture with group of trees bordering forest, on bark (lower trunk), 2002, Sipman 47817a (B).
Astrothelium solitarium Aptroot & M. Cáceres. Venezuela: Estado Bolivar, Cerro Guaiquinima, in central part of upper plateau, along Rio Carapo (near camp 4), 5°40'N, 63°34'W, alt. c. 1000 m, sandstone flats with boggy Stegolepis vegetation and scrub, 1990, Sipman 26683 (B, VEN).
Astrothelium straminicolor (Nyl.) Aptroot & Lücking. Singapore: SE side of MacRitchie Reservoir, 1°21'N, 103°50'E, alt. c. 50 m, secondary forest with primary forest remnants, on tree trunk within reach of the ground, 2000, Sipman 45888 & Tan (B).
Astrothelium subdisjunctum (Müll. Arg.) Aptroot & Lücking. Colombia: Caqueta, 2·5 km NE of Araracuara, 0°37'S, 72°23'W, alt. 250 m, c. 30 m tall, hardly disturbed forest on low terrace of Caquetá River (parcelas de Marcela Torres), on canopy branches of Brosimum, 1988, Sipman 27909 & Duivenvoorden (B).
Astrothelium subendochryseum Lücking et al. Costa Rica: Prov. Cartago, SW-slope of Volcán Turrialba, N of Santa Cruz, Fila Bonilla, along Camino El Trazado, 10°01'N, 83°43'W, alt. 2100 m, pastureland with relic trees of cloud forest, 1985, Sipman 20355 (B).
Astrothelium subfuscum Kremp. Singapore: Sembawan Park, on N-coast, 1°28'N, 103°51'E, alt. c. 2 m, grassland with scattered trees and shrubs, on tree trunk within reach of the ground, 2000, Sipman 46404 & Tan (B).
Astrothelium subscoria Flakus & Aptroot # (previously included within the concept of A. nitidiusculum s. lat.). Guyana: Upper Takutu District, c. 45 km S of Aishalton, c. 3 km S of Kuyuwini Landing, along trail to Kassikaityu River, 1°55'N, 59°15'W, alt. c. 230 m, epiphyte in light scrub with Clusia around tiny savannah, 1992, Sipman 57022 (B).
Astrothelium vulcanum Aptroot # (previously included within the concept of Trypethelium ochroleucum s. lat.). El Salvador: Santa Ana, municipality of Metapán, Parque Nacional Montecristo, Hacienda San José Ingenio, Caja de Agua de Metapán, 14°22'N, 89°23'W, alt. c. 900 m, on tree in Pinus-Quercus forest on lateritic soil on slope, 1993, Sipman, Berendsohn & Ladino 37410 (B).
Bathelium madreporiforme (Eschw.) Trevis. China: Yunnan Prov., Xishuangbanna, Xishuangbanna Tropical Botanical Garden in Menglun, 21°55·6'N, 101°15·4'E, alt. 550 m, on Anacardium occidentale trunks scattered in parkland, 2002, Sipman 50476 (B, KUN).
Bathelium nigroporum (Makhija & Patw.) Aptroot & Lücking. Papua New Guinea: Madang Prov., S side of Ramu Valley, 8–10 km W of Brahman Mission, 5°44·9'S, 145°19·7'E, alt. 100 m, logging site in lowland forest remnant, 1995, Sipman 38820 (B).
Bathelium porinosporum Lücking et al. China: Yunnan Prov., Xishuangbanna, Xishuangbanna Tropical Botanical Garden in Menglun, 21°55·6'N, 101°15·4'E, alt. 550 m, on trunks scattered in parkland, 2002, Sipman 50458 (B, KUN).
Pseudopyrenula flavoreagens Aptroot & M. Cáceres. French Guiana: Saül, 2 km S of the village, “sentier lemonade”, 3°32'N, 53°12'W, alt. 180–210 m, lowland moist forest, in upper canopy of Hura crepitans, 1986, Montfoort & Ek 8 (B).
Trypethelium ornatum Müll. Arg. Bolivia: Alto Beni, San Antonio, 15°39·770'S, 67°10·399'W, alt. 410 m, 23 year-old plantation, owned by Angel Flores, on Theobroma cacao bark, 1999, Derakshani 53 (B).
Updates to Aptroot & Lücking (Reference Aptroot and Lücking2016)
The Conselho Nacional de Desenvolvimento Científico e Tecnológico (Processos 401186/2014-8 and 459155/2014-8) is thanked for funding the collecting in Brazil, and for a research grant to MESC (207282/2015-3). AA thanks the Stichting Hugo de Vries-Fonds for travel grants.
