Research Article |
Corresponding author: Sonja Kistenich ( sonjadk@nhm.uio.no ) Academic editor: Imke Schmitt
© 2018 Sonja Kistenich, Jouko K. Rikkinen, Holger Thüs, Charles S. Vairappan, Patricia A. Wolseley, Einar Timdal.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Kistenich S, Rikkinen JK, Thüs H, Vairappan CS, Wolseley PA, Timdal E (2018) Three new species of Krogia (Ramalinaceae, lichenised Ascomycota) from the Paleotropics. MycoKeys 40: 69-88. https://doi.org/10.3897/mycokeys.40.26025
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Krogia borneensis Kistenich & Timdal, K. isidiata Kistenich & Timdal and K. macrophylla Kistenich & Timdal are described as new species, the first from Borneo and the two latter from New Caledonia. The new species are supported by morphology, secondary chemistry and DNA sequence data. Krogia borneensis and K. isidiata contain sekikaic and homosekikaic acid, both compounds reported here for the first time from the genus. Krogia macrophylla contains an unknown compound apparently related to boninic acid as the major compound. DNA sequences (mtSSU and nrITS) are provided for the first time for Krogia and a phylogeny of the genus based on 15 accessions of five of the six accepted species is presented. Krogia antillarum is reported as new to Brazil, Guatemala and Mexico.
Borneo, New Caledonia, lichens, Phyllopsora , phylogeny, rainforest, TLC
Krogia Timdal is a corticolous genus occurring in tropical humid forests and rainforests. It closely resembles the much more common genus Phyllopsora Müll. Arg. in thallus morphology, but differs mainly in having a weak or absent amyloid reaction in the tholus of the asci and filiform, curved ascospores that are spirally arranged in the ascus (
Three species of Krogia are known: K. antillarum Timdal (the West Indies;
During revision of material of Phyllopsora from Southeast Asia and Oceania, we have come across material of three apparently undescribed Krogia species. There are no published sequences of Krogia, but we have provided sequences of the mitochondrial small subunit (mtSSU) and of the nuclear ribosomal transcribed spacer region (ITS) from the three putative new species and from two of the three previously described species. The sequences, some of which were taken from an unpublished paper on the phylogeny of the Ramalinaceae (
The specimens of the three new species were discovered during ongoing global studies of Phyllopsora by Kistenich and Timdal in material provided by Rikkinen (New Caledonia) and Thüs, Vairappan and Wolseley (Borneo), with additional specimens provided by A. Elvebakk (New Caledonia) and A. Paukov (Borneo). The specimens are deposited in B, BM, BORH, H, O and PC. DNA sequences of the two previously described Krogia species were generated from specimens in B and O and from a specimen provided by P. Diederich (hb Diederich). Additionally, we included 14 mtSSU and 12 ITS sequences (Table
Specimens used in this study with voucher information, major lichen substances and GenBank accession numbers. New sequences are indicated by accession numbers in bold.
Species and sequence ID | Voucher | Major lichen substances | mtSSU | ITS |
---|---|---|---|---|
Aciculopsora salmonea | Costa Rica, 2004, Lücking 17543 (BR), isotype | – | MG925842 | MG925948 |
Bacidia rosella | Sweden, 1997, Ekman 3117 (BG) | – | AY300877 | AF282086 |
Bacidia rubella | Switzerland, van den Boom 41103 (LG DNA 578) | – | JQ796830 | JQ796852 |
Bacidia sipmanii | Tenerife, Sérusiaux s.n. (LG DNA 361) | – | JQ796832 | JQ796853 |
Bacidina brittoniana | USA, 1999, Ekman 3657 (BG) | – | – | MG925954 |
Bacidina delicata | France, Sérusiaux s. n. (LG DNA 369) | – | JQ796834 | JQ796854 |
Bacidina neosquamulosa | Netherlands, van den Boom 41056 (LG DNA 490) | – | JQ796837 | JQ796855 |
Bacidina phacodes | Sweden, 1998, Ekman 3414 (UPS) | – | AY567725 | AF282100 |
Eschatogonia prolifera I | Peru, 2006, Timdal 10207 (O) | didymic acid | MG925870 | MG925969 |
Eschatogonia prolifera II | Peru, 2006, Timdal 10429 (O) | didymic acid | MG925871 | MG925970 |
Krogia antillarum I | Trinidad And Tobago, 2008, Rui & Timdal 10844 (O), paratype | 4-O-methylcrypto-chlorophaeic acid | MH174271 | MH174281 |
Krogia antillarum II | Guatemala, 2002, Andersohn s.n. (B) | 4-O-methylcrypto-chlorophaeic acid | MH174272 | – |
Krogia antillarum III | Mexico, 1994, Wolf & Sipman 2052 (B) | 4-O-methylcrypto-chlorophaeic acid | MH174273 | MH174282 |
Krogia antillarum IV | Brazil, 2015, Dahl, Kistenich, Timdal & Toreskaas AM-39 (O) | 4-O-methylcrypto-chlorophaeic acid | MH174274 | MH174283 |
Krogia borneensis I | Malaysia, 2013, Vairappan & Thüs L291 (BORH), holotype | sekikaic acid, homosekikaic acid | MH174275 | – |
Krogia borneensis II | Malaysia, 2012, Wolseley, Thüs & Vairappan D-3-10-2 (BM) | sekikaic acid, homosekikaic acid | MH174276 | – |
Krogia borneensis III | Malaysia, 2014, Paukov 2234 (B) | sekikaic acid, homosekikaic acid | MH174277 | – |
Krogia borneensis | Malaysia, 1997, Wolseley Q21 p.p. (BM) | sekikaic acid, homosekikaic acid | – | – |
Krogia borneensis | Malaysia, 2013, Vairappan & Thüs L229 (BM) | sekikaic acid, homosekikaic acid | – | – |
Krogia coralloides I | Mauritius, 1991, Krog & Timdal MAU51/83 (O), holotype | boninic acid, unknown | MG925875 | MG925977 |
Krogia coralloides II | Mauritius, 2016, Diederich 18455 (hb. Diederich) | boninic acid, unknown | MH174278 | MH174284 |
Krogia isidiata I | New Caledonia, 2005, Elvebakk 05:633 (O), holotype | sekikaic acid, homosekikaic acid | – | MH174285 |
Krogia isidiata II | New Caledonia, 2016, Rikkinen 34385 (H) | sekikaic acid, homosekikaic acid | MH174279 | MH174286 |
Krogia isidiata | New Caledonia, 2016, Rikkinen 35034 (H) | sekikaic acid, homosekikaic acid | – | – |
Krogia isidiata | New Caledonia, 2016, Rikkinen 35688 (H) | sekikaic acid, homosekikaic acid | – | – |
Krogia macrophylla I | New Caledonia, 2016, Rikkinen 36047 (H) | unknown | – | MH174287 |
Krogia macrophylla II | New Caledonia, 2016, Rikkinen 36077 (H), holotype | unknown | – | MH174288 |
Krogia macrophylla III | New Caledonia, 2016, Rikkinen 35037 (H) | unknown | – | MH174289 |
Krogia macrophylla IV | New Caledonia, 2011, Rikkinen 38565 (H) | unknown | MH174280 | MH174290 |
Physcidia wrightii I | Mauritius, 1991, Krog & Timdal MAU14/14 (O) | sekikaic acid, divaricatic acid | MG925911 | – |
Physcidia wrightii II | Mauritius, 1991, Krog & Timdal MAU13/10 (O) | sekikaic acid, divaricatic acid | MG925912 | – |
Toninia cinereovirens | Norway, 1994, Haugan & Timdal 7953 (O) | – | AY567724 | AF282104 |
Waynea californica | USA, 1995, Ekman L1486 (UPS) | – | MG925947 | – |
Microscope sections were cut using a freezing microtome and mounted in water, 10% KOH (K), lactophenol cotton blue and a modified Lugol’s solution, in which water was replaced by 50% lactic acid. Amyloid reactions were observed in the modified Lugol’s solution after pretreatment in K and crystals of lichen substances were observed using polarised light.
Thin-layer chromatography (TLC) was performed in accordance with the methods of
We extracted DNA from apothecia and/or thallus tissue of 14 Krogia specimens. The DNA extraction followed the protocol described by
We assembled and edited the resulting sequences using the software Geneious R9 (
We used PartitionFinder2 (
Three Bacidia De Not. species, B. rosella (Pers.) De Not., B. rubella (Hoffm.) A. Massal. and B. sipmanii M. Brand et al., were used as outgroup in all phylogenetic analyses based on the molecular phylogeny of the Ramalinaceae (
The alignment was subjected to maximum likelihood analyses using Garli 2.01 (
We analysed the alignment phylogenetically using MrBayes 3.2.6 (
The results of the TLC analyses are shown in Table
We successfully generated DNA sequences for 14 Krogia specimens, including 10 mtSSU and 10 ITS sequences (Table
Initial RAxML analyses produced congruent gene trees of the mtSSU and ITS datasets; only unsupported (< 0.7) topological differences between the consensus trees were observed. We therefore continued with the subsequent phylogenetic analyses. PartitionFinder2 suggested three subsets and two different substitution models, the GTR+G model for (1) mtSSU, (2) ITS1 and ITS2 and the K80+I model for (3) 5.8S. The likelihood score of the best tree found by Garli was –8023.487881. The Bayesian analysis halted automatically after 3 million generations, when the ASDSF in the last 50% of each run had fallen below 0.01. We used 6004 trees for constructing the final majority-rule consensus tree. The phylogenetic results generated by Garli and MrBayes showed no incongruences. The extended majority-rule consensus tree of our alignment (Fig.
Hypothesis of the phylogenetic relationships and placement of the 15 Krogia accessions. It shows the extended majority-rule consensus tree resulting from the Bayesian MCMC analysis with Bayesian PP ≥ 0.7 (above branch) and/or Garli maximum likelihood BS ≥ 50 (below branch) and branch lengths. Strongly supported branches (PP ≥ 0.95 and BS ≥ 95) are marked in bold; branches with PP ≥ 0.95 and BS ≥ 70 are marked in bold grey; branches only supported by PP ≥ 0.7 are marked with an asterisk above the branch. Bacidia rosella, B. rubella and B. sipmanii were used as outgroup. Scale bar indicates 0.05 changes per site.
The genus Krogia was first described by Timdal in 2002 and only few reports of the genus have been published since (
All accessions of Krogia included in our molecular phylogeny (Table
Our specimens of the genus Krogia were typically found amongst collections of undetermined tropical rainforest lichens, particularly amongst those tentatively named Phyllopsora.
On detailed microscopic examination of specimens of the new species K. borneensis, we discovered a thin, unicellular cortex on the upper and lower side of the thallus. This type of cortex, with a single layer of rounded or cuboid cells and a thick cell wall, is characteristic for the tropical genus Eschatogonia Trevis. (
Krogia is resolved as the phylogenetic sister to a clade consisting of the type species of Bacidina, B. phacodes (Körb.) Vězda and B. brittoniana (Riddle) LaGreca & Ekman (Fig.
In recent years, lichenologists have increasingly focused on tropical regions and many new species have been described each year (e.g.
The species differs from K. isidiata in forming lacinules as vegetative dispersal units, not isidia, and from the other species in the genus in producing sekikaic and homosekikaic acid.
Malaysia, Borneo, Sabah, Maliau conservation area, trail between Nepenthes Camp and waterfall Takob Akob, 4°43.4'N, 116°52.2'E, 900–1000 m alt., in low (few metres) and open pristine montane "Kerangas" (heath) forest with higher trees mostly along a small stream, on smooth barked tree in the vicinity of the stream, 2013-02-23, C. Vairappan & H. Thüs L291 (BORH, holotype) [TLC: sekikaic and homosekikaic acid; GenBank: MH174275 (mtSSU)].
Thallus effuse, squamulose; squamules up to 1 mm wide, deeply divided into 0.1–0.2 wide lobes, ascending, imbricate, flattened, elongated to partly linear, often slightly laterally constricted, greyish-green with patches of red (K+ purple) spots, epruinose, glabrous; margin concolorous with upper side, not fibrillose; lower side white; lacinules formed by tips of the lobes. Upper cortex composed of a single layer of thick-walled cells with angular to shortly cylindrical lumina (resembling Eschatogonia-type), not containing crystals (polarised light!); algal layer 30–40 µm thick, filled with crystals dissolving in K; medulla composed of loosely interwoven hyphae, not containing crystals dissolving in K; lower cortex resembling upper cortex, both continuing over the edge of the squamule; prothallus brownish-black, often well developed. Apothecia (present in the holotype only) up to 0.6 mm diam. when simple, forming aggregates up to 1.5 mm diam., medium brown with red patches or entirely reddish-brown, more or less plane, with an indistinct, slightly paler, often flexuose margin; excipulum pale brown to colourless, composed of radiating, closely conglutinated hyphae, in inner part containing colourless crystals dissolving in K; hypothecium partly to entirely stained by a blood red pigment which dissolves in K with a purple effusion; epithecium colourless, not containing crystals. Ascospores filiform, curved, non-septate, spirally arranged in ascus, 20–31 × ca. 1.0 µm (n=10, from holotype). Conidiomata not seen.
Sekikaic acid (major), homosekikaic acid (major). Spot tests: all negative, except for red patches being K+ purple.
The species is known from five localities in Borneo.
Ecology. The species occurred in rather low "Kerangas" (heath) forest vegetation or on transition vegetation between the heath and oak/conifer (particularly Agathis) forest at higher elevations (ca. 1000 m) on very poor soils on sandstone (Fig.
The specific epithet refers to its occurrence in Borneo.
The medium-sized, flattened squamules make the species morphologically most similar to the neotropical K. antillarum. The squamules are more elongated, often linear and with more lateral constrictions in K. borneensis than in K. antillarum, which has more fan-shaped squamules. The former species has a thin, unicellular cortex on both upper and lower side, whereas the latter has a multicellular (20–30 µm thick) upper cortex and lacks a lower cortex (
Krogia isidiata shares the secondary chemistry (sekikaic and homosekikaic acid) with K. borneensis, but they differ in their vegetative dispersal units, the former producing cylindrical isidia, the latter flat lacinules (fragmenting squamules). The upper cortex of K. isidiata is multicellular (15–30 µm thick) and the lower cortex is absent.
Malaysia, Borneo. Sabah: Danum, plot 88, dipterocarp forest logged in 1988, 4°58'N, 117°50'E, 131 m alt., 1997-04-30, P. Wolseley Q21 p.p. (BM 001104020); Danum valley, pristine lowland dipterocarp forest 4°57.96'N, 117°47.32'E, 200–400 m alt., 2012, P. Wolseley, H. Thüs & C. Vairappan D-3-10-2 (BORH); Maliau conservation area, trail between Nepenthes Camp and waterfall Takob Akob, transition between pristine montane "Kerangas" (heath forest) and montane oak-conifer (Agathis) forest, 4°42.6'N, 116°52.5'E, 900–1000 m alt., 2013, C. Vairappan & H. Thüs L229 (BM); Ranau district, Kinabalu park, Musang camp on the Tambuyukon trail (loc. T98), 6°12.720'N, 116°40.891' E, 1429 m alt., epiphytic, 2014-12-09, A. Paukov 2234 (B).
The species differs from K. borneensis in forming isidia as vegetative dispersal units, not lacinules, and from the other species in the genus in producing sekikaic and homosekikaic acid.
New Caledonia, Province Sud, 20 km NNE of Nouméa, along dirt mountain road to Mt Dzumac, 3–400 m S of Seismic Station, ca. 22°03'S, 166°25'E, 830 m alt., on unidentified tree trunk in forest near the road, 2005-12-06, A. Elvebakk 05:633 (O L-186393, holotype; CANB, isotype [not seen]) [TLC: sekikaic and homosekikaic acid; GenBank: MH174285 (ITS)].
Thallus effuse, squamulose; squamules up to 0.4 mm wide, rounded and adnate when young, later becoming somewhat elongated with a crenulate and slightly ascending margin, flattened, green, with scattered patches of red (K+ purple) spots, epruinose, glabrous; margin concolorous with upper side, not fibrillose; lower side white; isidia attached marginally to the squamules, simple or sparingly branched, up to 1.8 mm long and 0.1 mm wide. Upper cortex composed of a few layers of thick-walled, irregularly or mainly periclinally orientated hyphae with angular to shortly cylindrical lumina, 15–30 µm thick, lacking an epinecral layer, not containing crystals (polarised light!); algal layer 30–40 µm thick, filled with crystals dissolving in K; medulla composed of loosely interwoven hyphae, containing crystals in the upper part; lower cortex lacking; prothallus brownish-black, well developed. Apothecia up to 0.8 mm diam. when simple, often forming aggregates up to 1.6 mm diam., dark reddish-brown to brownish-black, more or less plane, with a rather distinct, concolorous or slightly darker, flexuose margin; excipulum dark reddish-brown throughout, composed of radiating, closely conglutinated, thick-walled hyphae with narrowly cylindrical lumina, inner part containing crystals dissolving in K; hypothecium dark reddish-brown, composed of closely conglutinated, thick-walled hyphae with narrowly cylindrical lumina, containing crystals dissolving in K; epithecium colourless, not containing crystals (but crystals present in hymenium below). Ascospores filiform, curved, simple, spirally arranged in ascus, ca. 20–30 × ca. 1.0 µm (estimate of curved spores). Conidiomata not seen.
Sekikaic acid (major), homosekikaic acid (major). Spot tests: all negative, except for red patches being K+ purple.
The species is known from four collections at three localities in New Caledonia.
Ecology. The species grows on tree trunks in moist or mesic tropical forests and woodlands (Fig.
The specific epithet refers to its vegetative dispersal units, isidia.
This species and K. macrophylla are the only isidiate species of Krogia. They differ morphologically mainly in the size and shape of the squamules. In K. isidiata, they are small (up to 0.4 mm wide), rounded and adnate to somewhat elongated and with a slightly ascending margin and, in K. macrophylla, large (up to 3 mm wide), elongated and ascending even when young. In K. isidiata, the squamules are attached to a prothallus, whereas in the latter species, a prothallus has not been observed. The former species contains sekikaic and homosekikaic acid, the latter an unknown compound resembling boninic acid.
Krogia isidiata shares the secondary chemistry with K. borneensis; see that species for discussion.
New Caledonia. Province Sud: Yaté, dense forests along road RP 3 about 5 km west of Yaté, on tree trunk, 22°10'03.63"S, 166°54'10.15"E, 410 m alt., 2016-09-20, J. Rikkinen 34385 (H); Blue River Provincial Park, dense riparian forest near camp site on river bank, on tree trunk, 22°05'54.79"S, 166°38'20.24"E, 200 m alt. 2016-09-22, J. Rikkinen 35034 (H); Blue River Provincial Park, dense forest between camp site and road GR NC1, on tree trunk, 22°05'47.63"S, 166°38'22.54"E, 220 m alt., 2016-09-24, J. Rikkinen 35688 (H, PC).
The species differs from all other species of the genus in forming larger (up to 3 mm wide, vs. up to 0.3–1.5 mm wide in the other species) squamules and, except for K. coralloides, in producing an unknown compound resembling boninic acid.
New Caledonia, Province Sud, Mont Mou Nature Reserve, in low dense mist forest along foot path to the mountain summit, on tree trunk, 22°03'39.66"S, 166°20'53.54"E, 1162 m alt., 2016-09-26, J. Rikkinen 36077 (H, holotype [TLC: unknown compound resembling boninic acid; GenBank: MH174288 (ITS)]; PC, isotype).
Thallus effuse, squamulose; squamules up to 3 mm wide, at first rounded, later becoming incised and deeply divided into up to 1 mm wide lobes, ascending even when young, often imbricate, flattened or with an up-turned tip, greyish-green, with patches of purple (K+ bluish-black) spots, epruinose, glabrous; margin concolorous with upper side, not fibrillose; lower side white; isidia (present in one specimen) attached marginally to the squamules, simple or sparingly branched, up to 1.6 mm long and 0.2 mm wide. Upper cortex composed of thick-walled, irregularly orientated hyphae with angular to cylindrical lumina, 50–80 µm thick, lacking an epinecral layer, not containing crystals (polarised light!); algal layer 25–35 µm thick, filled with crystals dissolving in K; medulla composed of loosely interwoven hyphae, upper part containing crystals dissolving in K; lower cortex lacking; prothallus lacking. Apothecia up to 1 mm diam. when simple, often forming aggregates up to 6 mm diam., pale to medium brown, with purple patches, plane to weakly convex, with an indistinct, slightly paler, often flexuose margin; excipulum pale brown to colourless, composed of radiating, closely conglutinated, thick-walled hyphae with narrowly cylindrical lumina, not containing crystals; hypothecium pale brown to colourless, composed of closely conglutinated, thick-walled hyphae with narrowly cylindrical lumina, not containing crystals; epithecium colourless, not containing crystals; purple pigment occurring patchily in exciple, hypothecium and hymenium. Ascospores filiform, curved, simple, spirally arranged in ascus, ca. 20–30 × ca. 1.0 µm (estimate of curved spores). Conidiomata not seen.
Krogia macrophylla A field photograph of JR36047B field photograph of holotype C herbarium photograph of holotype. Scale bar: 1 mm. Photo: J. Rikkinen (A, B), E. Timdal (C).
An unknown compound resembling boninic acid (major) and traces of additional compounds. Spot tests: all negative, except for purple patches being K+ deeper purple to bluish-black.
The species is known from three localities in New Caledonia.
The species grows on tree trunks in moist or wet tropical forests (Figs
The specific epithet refers to the large squamules.
In the examined material, one specimen (Rikkinen 38565) is isidiate, whereas the others are not. Our first assumption, that two species were involved, was not confirmed by the phylogeny (Fig.
Krogia macrophylla has a similar secondary chemistry to K. coralloides (an unknown substance resembling boninic acid as the major constituent) but differs in lacking the boninic acid that co-occurs as the major constituent in K. coralloides (
New Caledonia. Province Sud: Blue River Provincial Park, dense riparian forest near camp site on river bank, on tree trunk, 22°05'54.79"S, 166°38'20.24"E, 200 m alt., 2016-09- 22, J. Rikkinen 35037 (H); locality data as for holotype, J. Rikkinen 36047 (H); Mont Humboldt Nature Reserve, close to Mont Humboldt refuge, in low dense mist forest along foot path from shelter towards the mountain summit, on tree trunk, 21°52'46.79"S, 166°24'49.17"E, 1320 m alt., 2011-11-09, J. Rikkinen 38565 (H).
1 | Squamules large, up to 3 mm wide and with up to 1 mm wide lobes; containing an unknown compound resembling boninic acid | K. macrophylla |
– | Squamules smaller, up to 1.5 mm wide and with up to 0.4 mm wide lobes; chemistry various | 2 |
2 | Thallus with isidia; containing sekikaic and homosekikaic acid | K. isidiata |
– | Thallus without isidia; chemistry various | 3 |
3 | Squamules minute, up to 0.3 wide and with up to 0.1 mm, simple lobes, forming a microphyllinous crust; not containing lichen substances | K. microphylla |
– | Squamules medium sized, up to 1.5 mm wide and with up to 0.4 mm wide, coralloid elongated lobes; containing lichen substances | 4 |
4 | Thallus with brownish black hypothallus; containing sekikaic and homosekikaic acid | K. borneensis |
– | Thallus without distinct hypothallus; chemistry different | 5 |
5 | Squamules mainly flattened; lobes up to 0.4 mm wide; containing 4-O-methylcryptochlorophaeic acid | K. antillarum |
– | Squamules mainly convex; lobes up to 0.1 mm wide; containing boninic acid and an unknown, similar compound | K. coralloides |
We would like to thank Arve Elvebakk and Alexander Paukov for providing us with their field collections; the specimen of the latter (A. Paukov 2234 (B)) was collected during a sampling trip to Sabah. We are grateful to Mika Bendiksby for assistance in the DNA lab and for valuable discussions. We acknowledge the Sabah Biodiversity Council, the administration units at Danum Valley and Maliau Conservation Areas for granting permits to access and work in the studied sites in the framework of the NHM “Danum – Maliau Quantitative Inventory” (JKM/MBS.1000-2/2 (77), MBMC 2012/15) and for the provision of logistic support. J.R. would like to thank Jérôme Munzinger (Montpellier) for help in the preparation of the fieldwork in New Caledonia and Alexander R. Schmidt and Leyla Seyfullah (University of Göttingen) for planning and organising the field trips. Fieldwork and collection in southern New Caledonia were kindly permitted by the Direction de l’Environnement (Province Sud), permit 17778/DENV/SCB delivered in November 2011 and permit 20112388-2016/ARR/DENV delivered in October 2016.
Krogia antillarum is reported here as new to Brazil, Guatemala and Mexico from the following examined specimens: Brazil. Rio de Janeiro: Parque Nacional do Itatiaia, along trail to Três Picos, 22.4358°S, 44.6118°W, 1090 m alt., on tree trunk in Atlantic rainforest, 2015-11-27, M.S. Dahl, S. Kistenich, E. Timdal & A.K. Toreskaas AM-39 (O L-202829). Guatemala. Depto. Alta Verapaz: NE of Cobán-Aragon, at the borders of Rio Cahabon (tierra fría), 1700 m alt., cloud forest, on Liquidambar tyraciflua, 2002-09-13, C. Andersohn s.n. (B 60-127330. Mexico. Chiapas: Municipio La Trinitaria, Parque Nacional Lagunas de Montebello, Paso del Soldado, 16°07'07"N, 91°43'09"W, 1500 m alt., bosque de Pinus maximinoi y Quercus sapotifolia, exposición N, epífita, 1994-11-29, J. Wolf & H. Sipman 2052 (B 60-110597).