Fungal Diversity

, Volume 58, Issue 1, pp 13–32

High diversity of Graphidaceae (lichenized Ascomycota: Ostropales) in Amazonian Perú

Authors

    • Department of Biological SciencesUniversity of Illinois-Chicago
    • Department of BotanyThe Field Museum
  • Robert Lücking
    • Department of BotanyThe Field Museum
Article

DOI: 10.1007/s13225-012-0172-y

Cite this article as:
Plata, E.R. & Lücking, R. Fungal Diversity (2013) 58: 13. doi:10.1007/s13225-012-0172-y

Abstract

A survey of crustose microlichens at Los Amigos Biological Station in Amazonian Peru revealed 116 species of Graphidaceae at this site. This is the second highest number of Graphidaceae ever reported for a single site world-wide, after the Surumoni crane station in Venezuela, with 131 species, and followed by Fakahatchee Strand Park Preserve in Florida, with 111 species. Based on the number of Graphidaceae found at Los Amigos, we predict the total lichen species richness at this site to be approximately 700 species. Of the 116 species encountered at Los Amigos, 59 were graphidoid species (former Graphidaceae s.str.) and 67 thelotremoid species (former Thelotremataceae). The following 18 species are described as new: Ampliotrema sorediatum Rivas Plata & Lücking, spec. nova, Chapsa hypoconstictica Rivas Plata & Lücking, spec. nova, Chapsa scabiocarpa Rivas Plata & Lücking, spec. nova, Chapsa subsorediata Rivas Plata & Lücking, spec. nova, Diorygma nigricans Rivas Plata & Lücking, spec. nova, Fissurina flavomedullosa Rivas Plata & Lücking, spec. nova, Fissurina platythecioides Rivas Plata & Lücking, spec. nova, Graphis apertoinspersa Rivas Plata & Lücking, spec. nova, Graphis pitmanii Rivas Plata & Lücking, spec. nova, Leucodecton inspersum Rivas Plata & Lücking, spec. nova, Ocellularia cicra Rivas Plata & Lücking, spec. nova, Ocellularia fenestrata Rivas Plata & Lücking, spec. nova, Ocellularia microsorediata Rivas Plata & Lücking, spec. nova, Ocellularia natashae Rivas Plata & Lücking, spec. nova, Ocellularia plicata Rivas Plata & Lücking, spec. nova, Ocellularia protoinspersa Rivas Plata & Lücking, spec. nova, Ocellularia pustulata Rivas Plata & Lücking, spec. nova, and Thelotrema amazonicum Rivas Plata & Lücking, spec. nova.

Keywords

LichensThelotremoidTropicalRainforestCorticolousMadre de Dios

Introduction

With the recent inclusion of Thelotremataceae, Gomphillaceae, and Asterothyriaceae, Graphidaceae is the largest family of tropical lichens, with around 2300 species and, considering the amount of undescribed ones, possibly the largest family of lichenized fungi (Rivas Plata et al. 2012a). The family is especially rich in tropical lowland rain forests, but detailed surveys of particular sites are rare, mainly due to the lack of taxonomic resources in the past. This has changed with several large revisions published recently (Staiger 2002; Frisch et al. 2006; Lücking et al. 2009a; Rivas Plata et al. 2010a), which provide keys to identify many of the species in large genera such as Chapsa, Graphis, and Thelotrema.

As part of an ecological study testing the use of Graphidaceae as bioindicators of ecological continuity or forest health (Rivas Plata et al. 2012b), following an earlier study in Costa Rica (Rivas Plata et al. 2008), we surveyed the species richness of Graphidaceae at a single site in Amazonian Peru, the Centro de Investigación y Capacitación Río Los Amigos (CICRA) or Los Amigos Research and Training Center. The area of Los Amigos is located in the southwestern part of the Amazon basin at the base of the Andes, about a day trip away from Puerto Maldonado (Department of Madre de Dios), between Manú National Park in the northwest and Bahuaja-Sonene National Park in the southeast (Fig. 1). The average altitude is 270 m above sea level, with the station situated on a high terrace at the confluence of the Madre de Dios and Los Amigos rivers. The station is contiguous to the Los Amigos Conservation Concession (LACC), which protects a diversity of upland and lowland forest types and aquatic habitats with a total area of about in 1,450 km². The concession is 145,735 hectares dominated by lowland humid forest, where altitude varies from 200 to 350 meters above sea level and annual rainfall is between 2,000 and 3,000 mm (Ascorra et al. 1999; Pitman 2008).
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Fig. 1

Map showing geographical location of study site

The area around the station is sparsely populated, with approximately two persons per square kilometer, mostly gold miners. Due to previous more extensive mining activities, smaller parts of the surrounding forest are slightly to strongly disturbed or represent regrowth in various stages of succession, whereas most of the area represents more or less undisturbed rain forest with minor impact from the activities of indigenous tribes that inhabit(ed) the area. The station was established in 2000 by the Peruvian NGO Asociación para la Conservación de la Cuenca Amazónica (ACCA) and the US-based Amazon Conservation Association (ACA). The station is administered by ACCA, in partnership with ACA and the Amazon Center for Environmental Education and Research (ACEER; Pitman 2010).

The objective of the study was to identify all species of Graphidaceae encountered by means of systematic transect sampling, covering an area of primary forest, another of disturbed and secondary forest, and a third representing planted trees around the station. While the ecological results are presented in a separate paper (Rivas Plata et al. 2012b), the taxonomic results are given here, with a complete list of species including 18 new to science. The number of species found is discussed in the framework of tropical lichen diversity and species concepts.

Material and methods

The collections treated here were made by the authors in August 2008 at Los Amigos Biological Station. The exact locality data are: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest; Aug 2008, leg. E. Rivas Plata & R. Lücking.

The vegetation at the station corresponds to lowland tropical rainforest. Variation encountered within the collection area was entirely due to different forest succession and management, including patches of primary forest, forest regrowth at different stages between pioneer vegetation and oldgrowth secondary forest, and trees planted around the station (Pitman 2008). The collections studied here originated from all succession and management types, although most species were found in primary and oldgrowth secondary forest and on exposed, planted trees. We employed two methods of sampling: (1) Quantitative sampling along two 900 m long transects including 100 trees, to compare different successional stages in their diversity and composition of Graphidaceae; this method and the results are described in a separate paper (Rivas Plata et al. 2012b). (2) Opportunisting sampling following the methods suggested by Gradstein et al. (1996), in areas and on trees not forming part of the quantitative sampling effort; in opportunisting sampling we also included three recently fallen trees to obtain a selection of canopy species.

Identification work was done at the Field Museum of Natural History in Chicago. Specimens were examined using LEICA MS5 and OLYMPUS SZX12 dissecting microscopes and ZEISS Axioscop 2 and OLYMPUS BH-2 compound microscopes, in part connected to JENOPTIC ProgRes C3 and C5 digital microscope cameras. Images were also made with NIKON Coolpix 5400 and NIKON Coolpix 8400 digital cameras. All specimens are kept at F. Anatomical measurements refer to specimens mounted in water; for iodine reactions, we used FLUKA 62650 Lugol solution. Spot tests with P (for psoromic and protocetraric acids) were done on small, separated pieces of thalli with the cortex scraped off, using para-phenyldiamine crystals freshly dissolved in alcohol. Spot tests with K (for norstictic and stictic acids) were done under the compound microscope using sections of thalli and ascomata. For most specimens, we also employed thin-layer chromatography (TLC) using mostly solvent C (Orange et al. 2001; Lumbsch 2002). In addition, selected specimens were sequenced targeting mtSSU, nuLSU, and RPB2 genes (Rivas Plata and Lumbsch 2011).

Nomenclature follows Staiger (2002), Frisch et al. (2006), Lücking et al. (2009a), and Rivas Plata et al. (2010a, 2012a). Authentic type material was studied for about 90 % of the species of Graphidaceae and their accepted synonyms, to ensure that no previous names were available for the newly described material.

Results and discussion

A total of 116 species of Graphidaceae was identified from Los Amigos Biological Station (Table 1). This is the second highest number reported world-wide, following 131 taxa reported from the Surumoni crane station in Venezuela (Komposch and Hafellner 1999) and surpassing the 111 species recently reported from Fakahatchee Strand Park Preserve in Florida (Lücking et al. 2011). Although these numbers are not exactly comparable, since the sampling techniques were partially different (quantitative with canopy access in Venezuela; opportunistic only in Florida), they give a good approximation of the species richness to be expected in certain forest types, as all three sites more or less correspond to lowland rainforest in its wide sense.
Table 1

Species of Graphidaceae identified at Los Amigos Biological Station, new species in bold. Collection numbers are given except for new species described in detail below

Acanthotrema brasilianum (Hale) Frisch (Rivas Plata 1-07-A, 8-02-C, 8-07-C, 8-08-D, 8-09-B, 8-09-C, 8-09-D)

Ampliotrema amplius (Nyl.) Kalb ex Kalb (Rivas Plata FM-02a, FM-extra)

Ampliotrema lepadinoides (Leight.) Kalb (Rivas Plata 8-02-A, 8-extra, FM-extra)

Ampliotrema sorediatum Rivas Plata & Lücking, spec. nova (see below)

Carbacanthographis stictica Staiger & Kalb (Rivas Plata 8-02-A, 8-03-A, 8-05-A, 8-07-A, 8-09-A, 8-09-E, 8-10-B)

Chapsa albomaculata (Sipman) Sipman & Lücking (Rivas Plata 8-04-A, 8-06-C, 8-09-A)

Chapsa alborosella (Nyl.) Frisch (Rivas Plata 1-07-C)

Chapsa cinchonarum (Fée) Frisch (Rivas Plata 1-06-C, 8-10-E)

Chapsa crispata (Müll. Arg.) Mangold (Rivas Plata 1-07-C, 1-canopy, 1-extra, 8-01-D, 8-09-A, 8-extra)

Chapsa dissuta (Hale) Mangold (Rivas Plata 8-05-D)

Chapsa hypoconstictica Rivas Plata & Lücking, spec. nova (see below)

Chapsa scabiocarpa Rivas Plata & Lücking, spec. nova (see below)

Chapsa subsorediata Rivas Plata & Lücking, spec. nova (see below)

Chapsa thallotrema Lücking & N. Salazar (Rivas Plata 1-07-E)

Chapsa velata (Müll. Arg.) Cáceres & Lücking (Rivas Plata FM-04a, FM-05c)

Chapsa spec. (sorediate 1; see below)

Chapsa spec. (sorediate 2; see below)

Chapsa spec. (sorediate 3; see below)

Clandestinotrema pauperius (Nyl.) Rivas Plata, Lumbsch & Lücking (Rivas Plata FM-01a)

Diorygma confluens (Fée) Kalb, Staiger & Elix (Rivas Plata FM-02b, FM-07 g, FM-08e)

Diorygma epiglaucum (Müll. Arg.) Kalb, Staiger & Elix (Rivas Plata FM-10b)

Diorygma nigricans Rivas Plata & Lücking, spec. nova (see below)

Diorygma poitaei (Fée) Kalb, Staiger & Elix (Rivas Plata 1-01-C)

Diorygma tibellii Kalb, Staiger & Elix (Rivas Plata 1-04-C, FM-07e)

Dyplolabia afzelii (Ach.) A. Massal. (Rivas Plata 1-01-A, FM-02c, FM-04b, FM-05f, FM-09a)

Fissurina aggregatula Common & Lücking (Rivas Plata 1-07-C)

Fissurina cingalina (Nyl.) Staiger (Rivas Plata 1-07-B, 1-08-E, 1-09-A, 1-09-C, 1-canopy, 1-extra, 8-03-A, 8-03-B, 8-04-B, 8-04-D, 8-05-A, 8-05-D, 8-05-E, 8-06-A, 8-07-D, 8-08-A, 8-09-A, 8-10-B, 8-10-C, 8-10-E, 8-extra)

Fissurina dumastii Fée (Rivas Plata 1-08-C, 1-canopy, 8-01-E, 8-03-A, 8-03-B, 8-03-E, 8-04-B, 8-04-D, 8-06-A, 8-08-A, 8-08-C, 8-09-A, 8-09-C, 8-09-E, 8-10-B, 8-10-D)

Fissurina dumastioides (Fink) Staiger (Rivas Plata 1-06-E)

Fissurina flavomedullosa Rivas Plata & Lücking, spec. nova (see below)

Fissurina furfuracea (Leight.) A. W. Archer (Rivas Plata 1-01-A)

Fissurina insculpta Mont. (Rivas Plata 8-09-canopy, 8-extra)

Fissurina platythecioides Rivas Plata & Lücking, spec. nova (see below)

Fissurina radiata Mont. (Rivas Plata FM-01c)

Fissurina spec. (Rivas Plata FM-07j, FM-08d)

Glyphis atrofusca (Müll. Arg.) Lücking (Rivas Plata FM-05a)

Glyphis cicatricosa Ach. (Rivas Plata FM-07a)

Graphis acharii Fée (Rivas Plata FM-07 h)

Graphis apertoinspersa Rivas Plata & Lücking, spec. nova (see below)

Graphis chrysocarpa (Raddi) Spreng. (Rivas Plata 1-01-A, 1-01-C, 1-01-G)

Graphis caesiella Vain. (Rivas Plata FM-06b, FM-09c)

Graphis conferta Zenk. (Rivas Plata 1-01-C, FM-06c)

Graphis duplicata Ach. (Rivas Plata 1-01-G)

Graphis furcata Fée (Rivas Plata 1-01-C, 1-01-G, FM-01e)

Graphis glaucescens Fée (Rivas Plata 1-03-D, 1-07-C, 8-05-D)

Graphis illinata Eschw. (Rivas Plata FM-07b)

Graphis implicata Fée (Rivas Plata 1-02-E, 1-03-A)

Graphis pitmanii Rivas Plata & Lücking, spec. nova (see below)

Graphis sitiana Vain. (Rivas Plata FM-01 g)

Graphis tenella Ach. (Rivas Plata 1-01-C, FM-07c)

Graphis vestitoides (Fink) Staiger (Rivas Plata 1-01-G)

Gyrotrema album Kalb (Rivas Plata 8-03-B, 8-05-D)

Leucodecton expallescens (Nyl.) Rivas Plata & Lücking (Rivas Plata 8-extra)

Leucodecton inspersum Rivas Plata & Lücking, spec. nova (see below)

Malmographina plicosa (Meissn.) Cáceres, Rivas Plata & Lücking (Rivas Plata FM-03b, FM-09d)

Melanotrema platystomum (Mont.) Frisch (Rivas Plata 1-06-C, 1-06-D, 1-06-E)

Myriotrema clandestinum (Fée) Hale (Rivas Plata 8-01-A, 8-01-B, 8-04-B, 8-07-A, 8-08-D, 8-09-D)

Myriotrema erodens R. C. Harris (Rivas Plata 1-03-D, 1-06-A, 1-07-C, 1-07-E, 1-08-B, 1-08-C, 1-10-A, 1-10-C, 8-01-C, 8-04-E, 8-06-A, 8-06-E, 8-09-D)

Myriotrema foliicola (Hale) Hale (Rivas Plata 1-08-A, 1-09-C, 8-03-D, 8-05-C, 8-07-A, 8-08-B, 8-08-D, 8-08-E)

Myriotrema microporum (Mont.) Hale (Rivas Plata 8-09-A)

Myriotrema myrioporum (Tuck.) Hale (Rivas Plata 8-05-D, 8-10-D)

Myriotrema pulverulentum (Hale) Hale (Rivas Plata 8-01-A)

Myriotrema viride Nagarkar & Hale (Rivas Plata 8-01-A, 8-03-E)\

Myriotrema spec. (isidiate-sorediate; see below)

Ocellularia albula (Nyl.) Zahlbr. (Rivas Plata 1-canopy)

Ocellularia calvescens (Fée) Müll. Arg. (Rivas Plata 1-08-A, 8-09-C)

Ocellularia cavata (Ach.) Müll. Arg. (Rivas Plata 1-01-A)

Ocellularia cicra Rivas Plata & Lücking, spec. nova (see below)

Ocellularia exigua Müll. Arg. (Rivas Plata 8-05-D)

Ocellularia fenestrata Rivas Plata & Lücking, spec. nova (see below)

Ocellularia fumosa (Ach.) Müll. Arg. (Rivas Plata 8-09-canopy)

Ocellularia garoana Patw. & Nagarkar (Rivas Plata 8-01-A, 8-07-B)

Ocellularia gerardii Sipman (Rivas Plata 1-canopy, 1-extra, 8-03-D)

Ocellularia gymnocarpa (Nyl.) Zahlbr. (Rivas Plata 1-07-C, 1-08-B, 8-01-D, 8-03-B, 8-03-E, 8-04-D, 8-04-E, 8-05-E, 8-07-C, 8-07-E, 8-08-D, 8-09-D, 8-10-E)

Ocellularia laeviusculoides Sipman & Lücking (Rivas Plata 8-01-B, 8-08-B)

Ocellularia mauritiana Hale (Rivas Plata 8-03-D, 8-05-A)

Ocellularia microascidium (Vain.) Zahlbr. (Rivas Plata 8-09-canopy)

Ocellularia microsorediata Rivas Plata & Lücking, spec. nova (see below)

Ocellularia natashae Rivas Plata & Lücking, spec. nova (see below)

Ocellularia papillata (Leight.) Zahlbr. (Rivas Plata 8-01-B, 8-01-C, 8-01-D, 8-01-E, 8-03-B, 8-06-C, 8-09-B, 8-09-C, 8-09-D, 8-09-E, 8-10-D, 8-extra)

Ocellularia perforata (Leight.) Müll. Arg. (Rivas Plata 1-08-D, 8-01-B, 8-01-E, 8-09-C)

Ocellularia plicata Rivas Plata & Lücking, spec. nova (see below)

Ocellularia pluripora Hale (Rivas Plata 8-05-E, 8-07-B)

Ocellularia protoinspersa Rivas Plata & Lücking, spec. nova (see below)

Ocellularia psorbarroensis Sipman (Rivas Plata 1-08-A, 1-08-B, 1-09-A, 1-canopy, 8-01-D, 8-02-A, 8-02-D, 8-03-A, 8-08-B, 8-08-D, 8-08-E, 8-09-A, 8-09-E, 8-10-B, 1-extra, 8-extra)

Ocellularia pustulata Rivas Plata & Lücking, spec. nova (see below)

Ocellularia terebrata (Ach.) Müll. Arg. (Rivas Plata 1-08-C, 8-07-B, 8-10-B, 8-10-C, 8-10-E)

Ocellularia thryptica Hale (Rivas Plata 1-03-D, 8-extra)

Ocellularia vezdana Frisch (Rivas Plata 1-09-A, 8-01-E, 8-05-A)

Ocellularia violacea Räsänen (Rivas Plata 8-04-B)

Ocellularia viridipallens Müll. Arg. (Rivas Plata 8-03-D, 8-04-E, 8-08-B)

Ocellularia spec. (papillose; see below)

Ocellularia spec. (sorediate 1; see below)

Ocellularia spec. (sorediate 2; see below)

Ocellularia spec. (sorediate 3; see below)

Phaeographis brasiliensis (A. Massal.) Kalb & Matthes-Leicht (Rivas Plata 1-01-A)

Phaeographis decipiens Müll. Arg. (Rivas Plata 1-01-A)

Phaeographis extrusa (Stirt.) Zahlbr. (Rivas Plata FM-07d)

Phaeographis flavescens Dal Forno & Eliasaro (Rivas Plata FM-06d, FM-07i)

Phaeographis haematites (Fée) Müll. Arg. (Rivas Plata FM-01f, FM-03a, FM-04d, FM-06f, FM-09e)

Phaeographis heterochroides Zahlbr. (Rivas Plata FM-05e)

Phaeographis inconspicua (Fée) Müll. Arg. (Rivas Plata FM-05b, FM-08c)

Phaeographis intricans (Nyl.) Vain. (Rivas Plata 1-01-C)

Phaeographis leprieurii (Mont.) Staiger (Rivas Plata FM-01 h, FM-03c, FM-04e, FM-08b, FM-09b)

Phaeographis neotricosa Redinger (Rivas Plata 1-01-A, 1-01-C, FM-03 h)

Phaeographis scalpturata (Ach.) Staiger (Rivas Plata 1-01-A, 1-01-C, 1-06-A, FM-01i, FM-01 l, FM-03d, FM-04c, FM-07f, FM-08a, FM-10a)

Phaeographis subfulgurata (Nyl.) Zahlbr. (Rivas Plata FM-01 k)

Phaeographis subtigrina (Vain.) Zahlbr. (Rivas Plata FM-05d, FM-09f)

Phaeographis aff. subtigrina (Vain.) Zahlbr. (Rivas Plata FM-01j, FM-03e, FM-06e)

Platygramme aff. caesiopruinosa (Fée) Fée (Rivas Plata FM-05 g)

Redingeria glaucoglyphica (Sipman) Frisch (Rivas Plata 8-02-A, 8-10-B, 8-10-extra)

Sarcographa labyrinthica (Ach.) Müll. Arg. (Rivas Plata FM-03f)

Stegobolus anamorphus (Nyl.) Frisch (Rivas Plata 8-04-E, 8-07-B, 8-07-D)

Stegobolus subwrightii (Hale) Lücking (Rivas Plata 8-07-C, 8-10-A)

Stegobolus wrightii (Tuck.) Frisch (Rivas Plata 8-07-C)

Thelotrema amazonicum Rivas Plata & Lücking, spec. nova (see below)

The total number of corticolous lichens reported for the Florida site was approximately 400; therefore, using the richness of Graphidaceae as an indicator, Los Amigos could harbour around 400 or more corticolous lichen species. Indeed, other groups not treated in this paper, such as Trypetheliaceae, are also extremely rich at Los Amigos (Nelsen et al., unpubl. data). In addition, foliicolous taxa, which were not collected during this study, can reach nearly 300 species in tropical lowland rain forest (Lücking 2008), suggesting that the total number of lichen species at Los Amigos could be around 700. This estimate is consistent with other predictions (Lücking et al. 2009b) and might even be conservative considering that the lichen-rich canopy (Komposch and Hafellner 1999, 2000, 2003; Rivas Plata et al. 2008) was not sampled. We can assume that the number of Graphidaceae for Los Amigos has not yet reached its saturation point, as many of these species are canopy dwellers (Hale 1974, 1978, 1981; Komposch and Hafellner 2000).

Ampliotrema sorediatum Rivas Plata & Lücking, spec. nova

(Fig. 2A–B)
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Fig. 2

A–B. Ampliotrema sorediatum (holotype). C–D. Chapsa hypoconstictica (holotype). E–F. Chapsa scabiocarpa (holotype). G–H. Chapsa subsorediata (holotype)

MycoBank: MB800195

Etymology: The epithet refers to the sorediate thallus, an unusual feature in the genus.

Differing from Ampliotrema lepadinoides in the sorediate thallus and yellow pruina on the apothecial disc.

Thallus corticolous, grey-olive, up to 5 cm diam. and 50–100 μm thick, continuous; surface smooth to uneven, with dense, proso- to paraplectenchymatous cortex, abundantly sorediate; soralia 0.1 mm diam., punctiform to maculate, white, granular; photobiont Trentepohlia, with angular-rounded to elongate cells 7–11 × 5–7 μm large; photobiont layer and medulla incrusted with numerous clusters of calcium oxalate crystals; medulla white. Apothecia rounded, prominent to sessile, 0.6–1 mm diam.; disc partially covered by 0.2–0.3 mm wide pore, brown-black, yellow-pruinose; margin entire, forming a brown rim around the pore, covered by thalline layer forming shallow verrucae. Columella absent. Excipulum paraplectenchymatous, upper half carbonized; periphysoids absent; hypothecium prosoplectenchymatous, 10–15 μm high, colorless. Hymenium 150–200 μm high, strongly and densely inspersed; paraphyses unbranched except in uppermost part of hymenium; asci fusiform, 140–180 × 30–40 μm. Ascospores 8/ascus, oblong to cylindrical, 13–19-septate, 60–80 × 10–12 μm, 6–8 times as long as wide, with thick septa and lens-shaped lumina, colorless, I + violet-blue. Secondary chemistry: protocetraric and virensic acids (exposed medulla P + orange-red; thallus section K–), apothecial disc with (pale) yellow anthraquinone (K + red).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata 8-08-A (F, holotype; USM, isotype), 1-10-E, 8-02-B, 8-07-A, 8-07-E, 8-08-D, 8-08-E, 8-10-A, 8-10-B (F, USM, paratypes).

Notes: Ampliotrema sorediatum is the first sorediate species in the genus (Kalb 2004; Frisch et al. 2006). Most similar is A. dactylizum (Hale) Sipman, Lücking & Grube (Sipman et al. 2012), which forms coarse isidioid structures that break off at the base which then can resemble a soralium. The latter species is also anatomically similar to A. sorediatum but forms an orange rather than yellow pruina on the apothecial disc.

Chapsa hypoconstictica Rivas Plata & Lücking, spec. nova

(Fig. 2C–D)

MycoBank: MB800196

Etymology: The epithet refers to the unusual chemistry.

Differing from Chapsa albomaculata in containing hypostictic acid as only substance.

Thallus corticolous, light grey-green, up to 3 cm diam. and 40–70 μm thick, continuous; surface smooth to uneven, compact but with loose cortex or partially ecorticate; photobiont Trentepohlia, with angular-rounded to elongate cells 7–11 × 5–7 μm large; photobiont layer and medulla incrusted with numerous clusters of calcium oxalate crystals; medulla white. Apothecia angular-rounded, erumpent, 0.3–0.6 mm diam., usually aggregate into groups of 3–7; disc partially covered by excipulum, pale brown, white-pruinose; margin lobulate, fused (but excipulum sometimes layered), yellow-white to pale brown, white-pruinose. Columella absent. Excipulum paraplectenchymatous, colorless; periphysoids present; hypothecium prosoplectenchymatous, 10–15 μm high, colorless. Hymenium 60–80 μm high, clear; paraphyses unbranched, thick, apically moniliform; asci fusiform, 60–80 × 15–20 μm. Ascospores 8/ascus, ellipsoid, 5–7-septate, 15–20 × 5–6 μm, 3–4 times as long as wide, with thick septa and lens-shaped lumina, colorless, I + violet-blue. Secondary chemistry: hypoconstictic acid (exposed medulla P–; thallus section K–).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata 1-08-B (F, holotype), 8-06-A, 8-06-C (F, paratypes).

Notes: This species is morphologically and anatomically very similar to Chapsa albomaculata (Sipman) Sipman & Lücking (Rivas Plata et al. 2010a; Sipman et al. 2012) in the pale green thallus, small, aggregate apothecia, and amyloid ascospores with lens-shaped lumina. The only difference is in the chemistry, with C. albomaculata (which was also found at this locality) having stictic and constictic acid (K + yellow) and the new species having the rare chemistry of hypoconstictic acid (K–) as the only detectable substance. A similar chemistry is thus far only known from the unrelated Topeliopsis elixii Kalb & Frisch (Frisch and Kalb 2006). Both C. albomaculata and C. hypoconstictica resemble C. phlyctidioides (Müll. Arg.) Mangold (Rivas Plata et al. 2010a), which produces stictic acid ad major and several satellite substances but lacks hypoconstictic acid and its ascospores are I– (non-amyloid).

Chapsa scabiocarpa Rivas Plata & Lücking, spec. nova

(Fig. 2E–F)

MycoBank: MB800197

Etymology: The epithet refers to the layered apothecial margin.

Differing from Chapsa laceratula in the smaller, transversely septate ascospores.

Thallus corticolous, olive-green, up to 3 cm diam. and 30–50 μm thick, continuous, mostly endoperidermal; surface smooth to uneven, with loose, irregular cortex; photobiont Trentepohlia, with angular-rounded to elongate cells 7–11 × 6–8 μm large; photobiont layer endoperidermal, lacking crystals; medulla absent. Apothecia angular-rounded, immersed-erumpent, 0.3–0.5 mm diam.; often aggregate or fused into groups of 2–3; disc partially covered by excipulum, pale grey-brown, white-pruinose; margin fissured-lobulate, fused but excipulum distinctly layered, yellow-white to pale brown. Columella absent. Excipulum paraplectenchymatous, brown to apically dark brown; periphysoids present; hypothecium prosoplectenchymatous, 5–10 μm high, colorless. Hymenium 90–110 μm high, clear; paraphyses unbranched; asci fusiform, 90–100 × 15–20 μm. Ascospores 8/ascus, ellipsoid, 5–7-septate, 20–25 × 7–8 μm, 2.5–3.5 times as long as wide, with thick septa and lens-shaped lumina, colorless, I + violet-blue. Secondary chemistry: no substances detected by TLC (thallus P–; thallus section K–).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata 1-07-E (F, holotype).

Notes: This new species exhibits a morphotype intermediate between Chapsa, Thelotrema, and Topeliopsis, also found in species such as C. laceratula (Müll. Arg.) Rivas Plata & Lücking and C. scabiomarginata (Hale) Rivas Plata & Lücking (Rivas Plata et al. 2010a). The mostly endophloedic, corticate thallus resembles that of C. platycarpella (Vain.) Frisch, whereas the apothecia are akin to Topeliopsis, with a main split between the innermost excipulum and outer layers suggesting a double margin as in Thelotrema. Chapsa scabiocarpa is distinguished from the morphologically similar C. laceratula and C. scabiomarginata by its ascospores (large and muriform in the other two species). It should not be confused with either C. albomaculata, C. hypoconstictica, and Thelotrema amazonicum occurring at the same locality. The first two have a thicker, epiperidermal thallus with a less dense cortex and chapsoid apothecia and also differ in their chemistry, whereas the latter has a farinose thallus, apothecia with a double margin, and larger ascospores.

Chapsa subsorediata Rivas Plata & Lücking, spec. nova

(Fig. 2G–H)

MycoBank: MB800198

Etymology: The epithet refers to the unusual chemistry.

Differing from Chapsa sorediata in the absence of lichen substances.

Thallus corticolous, light grey-green, up to 5 cm diam. and 40–70 μm thick, continuous; surface smooth to uneven, with indistinct, loose cortex, in part endoperidermal, abundantly sorediate; soralia 0.2–0.5 mm diam., maculate, white, granular; photobiont Trentepohlia, with angular-rounded to elongate cells 7–10 × 5–7 μm large; photobiont layer and medulla incrusted with clusters of calcium oxalate crystals; medulla indistinct, white. Apothecia angular-rounded, immersed-erumpent, 0.5–1 mm diam.; disc partially covered by crumbling excipulum, pale brown, thickly white-pruinose; margin lobulate with lobules erect to recurved, fused (but excipulum sometimes layered), yellow-white to pale brown, strongly white-pruinose. Columella absent. Excipulum paraplectenchymatous, colorless; periphysoids present; hypothecium prosoplectenchymatous, 10–15 μm high, colorless. Hymenium 70–80 μm high, clear; paraphyses unbranched, thick, apically moniliform; asci fusiform, 60–80 × 15–20 μm. Ascospores 8/ascus, elliopsoid, 5–9-septate, 20–25 × 6–8 μm, with thick septa and lens-shaped lumina, colorless, I + violet-blue. Secondary chemistry: no substances detected by TLC (thallus P–; thallus section K–).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata 1-07-A (F, holotype), 1-07-D (F, paratype).

Notes: Thus far the genus Chapsa contains three species with sorediate thallus, C. sorediata Kalb, C. defectosorediata Lücking, and C. thallotrema Lücking & N. Salazar (Rivas Plata et al. 2010a; Lumbsch et al. 2011). The latter two have a distinctly corticate thallus and large ascospores and are not closely related to C. subsorediata. Chapsa sorediata is very close to the present material but differs in the larger apothecia and soralia and the presence of stictic and constictic acids (Kalb 2009). Except for the soralia, C. subsorediata is most similar to C. crispata, but the latter has a more distinct cortex and a thinner pruina on the apothecial discs.

Diorygma nigricans Rivas Plata & Lücking, spec. nova

(Fig. 3A–B)
https://static-content.springer.com/image/art%3A10.1007%2Fs13225-012-0172-y/MediaObjects/13225_2012_172_Fig3_HTML.gif
Fig. 3

A–B. Diorygma nigricans (holotype). C–D. Fissurina flavomedullosa (holotype). E–F. Fissurina platythecioides (holotype). G–H. Graphis apertoinspersa (holotype)

MycoBank: MB800199

Etymology: The epithet refers to the brown-black, non-pruinose lirellae.

Differing from Diorygma confluens in the epruinose apothecial disc.

Thallus corticolous, light green-grey, up to 5 cm diam. and 80–150 μm thick, continuous but partially flaking off; surface uneven, ecorticate, compact to minutely farinose; photobiont Trentepohlia, with angular-rounded to elongate cells 8–12 × 6–8 μm large; photobiont layer and medulla incrusted with numerous clusters of calcium oxalate crystals; medulla white; below with a massively carbonized basal layer about 30–70 μm thick. Lirellae flexuose, densely branched and often in stellate clusters, immersed, with complete thalline margin, 0.5–3 mm long, 0.2–0.4 mm wide; disc exposed, brown-black, non-pruinose; proper margin indistinct, labia inconspicuous, visible only in young lirellae, entire; thalline margin thick, white. Excipulum entire, completely carbonized, 30–50 μm wide laterally and up to 100 μm thick at the base including the basal carbonized thallus layer; laterally up to the top covered by corticate algiferous thallus including clusters of crystals; hypothecium prosoplectenchymatous, 10–15 μm high, colorless. Hymenium 100–130 μm high, colorless, clear; epithecium granulose, 10–15 μm high, brownish; paraphyses unbranched except in uppermost part of hymenium; asci fusiform, 100–130 × 30–40 μm. Ascospores single, ellipsoid, richly muriform, 80–120 × 20–30 μm, 3.5–4.5 times as long as wide, colorless, I + violet-blue. Secondary chemistry: lichexanthone (thallus surface UV + yellow), stictic and constictic acids (major), cryptostictic, hypostictic, and hypoconstictic acids (minor or traces).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata FM-12 (F, holotype).

Notes: This new species is described with some hesitation. It is very similar to the widespread Diorygma confluens in all aspects (carbonization, chemistry), except for the more delicate, abundantly branched lirellae with inconspicuous labia and completely lacking a pruina, their exposed, brown-black discs forming a strong contrast with the surrounding thallus. Almost all species of Diorygma have strongly pruinose lirellae, except for D. erythrellum (Mont.) Kalb, Staiger & Elix which resembles a Platythecium (Kalb et al. 2004). The type material of the new species is well-developed and does not show any evidence being an aberrant form of D. confluens. We have examined many collections of the latter and the lirellae are always thickly pruinose and feature well-developed labia.

Fissurina flavomedullosa Rivas Plata & Lücking, spec. nova

(Fig. 3C–D)

MycoBank: MB800200

Etymology: The epithet refers to the yellow medulla.

Differing from Fissurina dumastii in the yellow medulla.

Thallus corticolous, green-grey, up to 10 cm diam. and 60–120 μm thick, continuous; surface smooth to uneven, with dense, prosoplectenchymatous cortex; photobiont Trentepohlia, with angular-rounded to elongate cells 7–10 × 5–7 μm large; photobiont layer with scattered clusters of calcium oxalate crystals; medulla yellow; below with a massively carbonized basal layer about 20–30 μm thick. Lirellae flexuose, irregularly branched, immersed-erumpent, fissurine, with complete thalline margin, 1–2 mm long, 0.07–0.1 mm wide; disc concealed; proper margin indistinct, labia entire, thin, covered by algiferous thalline layer except uppermost part which is yellow-white, non-pruinose; thalline margin thick, green-grey. Excipulum thin, non-carbonized, 20–30 μm wide and thick at the base, yellow-brown; laterally covered by corticate algiferous thallus including clusters of crystals; hypothecium prosoplectenchymatous, 5–10 μm high, colorless. Hymenium 80–100 μm high, colorless, clear; epithecium indistinct, 3–7 μm high, yellowish; paraphyses unbranched; asci fusiform, 70–90 × 15–18 μm. Ascospores 8/ascus, ellipsoid, transversely 3-septate, 10–13 × 5–6 μm, with up to 2 μm thick outer wall, 1.8–2.3 times as long as wide, with thick septa and lens-shaped lumina, colorless, I + violet-blue. Secondary chemistry: medulla with yellow anthraquinone (exposed medulla K + orange-red).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata 8-10-A (F, holotype; USM, isotype).

Notes: This new species is highly unusual in the genus Fissurina as it is the only one known with either a pigmented medulla or a carbonized basal hypothallus. A pigmented medulla is very common in Ocellularia s.lat. and has also been described from Myriotrema (Lumbsch et al. 2011), but to our knowledge this is the first case of a pigmented medulla in a lirellate Graphidaceae. A carbonized basal hypothallus is characteristic of several species of Diorygma (Kalb et al. 2004) but does not seem to have been reported from other genera. Except for these characters, the new species is superficially most similar to Fissurina dumastii (Staiger 2002) which, however, also differs in the non-amyloid ascospores. Within Fissurina, strongly amyloid ascospores with 3-septate septa appear to be restricted to species with either thick labia or excipulum carbonization (Lücking et al. 2011). The thallus with pigmented medulla of the new species very much resembles that of Myriotrema endoflavescens Hale ex Lücking, except that the latter lacks a carbonized hypothallus (Lumbsch et al. 2011).

Fissurina platythecioides Rivas Plata & Lücking, spec. nova

(Fig. 3E–F)

MycoBank: MB800201

Etymology: The epithet refers to the resemblance of the species with Platythecium.

Differing from Fissurina incrustans in the larger ascospores.

Thallus corticolous, light green, up to 5 cm diam. and 70–120 μm thick, continuous; surface smooth to uneven, with dense, prosoplectenchymatous cortex; photobiont Trentepohlia, with angular-rounded to elongate cells 7–12 × 5–7 μm large; photobiont layer lacking crystals except near the lirellae; medulla white. Lirellae straight to curved, mostly unbranched, immersed, fissurine but gaping, with complete thalline margin, 0.5–2 mm long, 0.2–0.3 mm wide; disc exposed, light orange-brown; proper margin indistinct, labia entire, thin, covered by algiferous thalline layer except uppermost part which is yellow-white, non-pruinose; thalline margin thick, green-grey. Excipulum thin, non-carbonized, 20–30 μm wide, orange-brown; laterally covered by corticate algiferous thallus including large clusters of calcium oxalate crystals; hypothecium prosoplectenchymatous, 5–10 μm high, colorless. Hymenium 120–150 μm high, colorless, clear; epithecium indistinct, 3–7 μm high, colorless; paraphyses unbranched; asci fusiform to clavate, 100–130 × 20–25 μm. Ascospores 8/ascus, ellipsoid to almost fusiform, muriform, 30–50 × 15–20 μm, with up to 2 μm thick outer wall, with thin septa and rectangular lumina, 2–2.5 times as long as wide, colorless, I–. Secondary chemistry: no substances detected by TLC (thallus surface or exposed medulla P–; thallus section K–).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata 8-01-C (F, holotype).

Notes: Fissurina platythecioides at first glance resembles a Platythecium due to the gaping lirellae exposing the orange-brown disc. However, the excipulum and ascospore type place this taxon into the genus Fissurina (Staiger 2002). Several species of Fissurina have somewhat gaping lirellae, including the widespread F. incrustans Fée, but the disc is usually more immersed. Fissurina incrustans differs from the new species otherwise in the smaller, weakly amyloid ascospores. No other species of Fissurina appears to have similarly sized ascospores as those found in F. platythecioides.

Graphis apertoinspersa Rivas Plata & Lücking, spec. nova

(Fig. 3G–H)

MycoBank: MB800202

Etymology: The epithet refers to the partially exposed disc and inspersed hymenium, an unusual combination in the genus.

Differing from Graphis anfractuosa in the lirellae with exposed apothecial disc and lateral thalline margin.

Thallus corticolous, white, up to 3 cm diam. and 40–70 μm thick, continuous; surface uneven, with thin, prosoplectenchymatous cortex; photobiont Trentepohlia, with angular-rounded to elongate cells 6–12 × 5–8 μm large; photobiont layer incrusted with numerous clusters of calcium oxalate crystals; medulla white. Lirellae straight to curved, unbranched to sparsely branched, prominent, with basal to thin lateral thalline margin, 1–5 mm long, 0.15–0.2 mm wide; disc exposed at maturity but remaining narrow, dark grey-brown, non-pruinose; proper margin distinct, labia entire, black, non-pruinose; thalline margin basal to lateral but remaining thin and not reaching top of labia. Excipulum entire, completely carbonized, 40–70 μm wide and thick at the base, black; basally to laterally covered by thin, corticate algiferous thallus including clusters of crystals; hypothecium prosoplectenchymatous, 5–10 μm high, colorless. Hymenium 80–100 μm high, colorless, strongly inspersed, inspersion partially dissolving in K (type B according to Lücking 2009); epithecium granulose, 5–10 μm high, brownish; paraphyses unbranched; asci fusiform, 70–100 × 20–25 μm. Ascospores 8/ascus, oblong-ellipsoid, transversely 5–9-septate, 20–30 × 6–8 μm, 3–4 times as long as wide, colorless, I + violet-blue. Secondary chemistry: no substances detected by TLC (thallus surface or exposed medulla P–; thallus section K–).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata FM-1b (F, holotype), FM-06a (F, paratype).

Notes: This new species is similar to Graphis anfractuosa in the delicate lirellae with completely carbonized excipulum, inspersed hymenium, small transversely septate ascospores and lack of secondary substances (Lücking 2009; Lücking et al. 2009a). It differs however, consistently in the exposed disc, the less prominent lirellae with basal to lateral thalline margin, and the more strongly inspersed hymenium.

Graphis pitmanii Rivas Plata & Lücking, spec. nova

(Fig. 4A–B)
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Fig. 4

A–B. Graphis pitmanii (holotype). C–D. Leucodecton inspersum (holotype). E–F. Ocellularia cicra (E, holotype; F, paratype). G–H. Ocellularia fenestrata (holotype)

MycoBank: MB800203

Etymology: Named in honour of Dr. Nigel C. A. Pitman, for his contributions to tropical ecology and directorship of Los Amigos Research and Training Center in Amazonian Peru.

Differing from Graphis tetralocularis in the basally carbonized excipulum and lack of lichen substances.

Thallus corticolous, white-grey, up to 5 cm diam. and 30–70 μm thick, continuous; surface uneven, with thin, prosoplectenchymatous cortex; photobiont Trentepohlia, with angular-rounded to elongate cells 7–12 × 5–7 μm large; photobiont layer incrusted with numerous clusters of calcium oxalate crystals; medulla white. Lirellae flexuose, irregularly branched, erumpent, with apically thin complete thalline margin, 0.5–2 mm long, 0.1–0.15 mm wide; disc concealed; proper margin distinct, labia entire, appearing dark grey due to corticate, non-algiferous thalline cover, non-pruinose; thalline margin laterally thick, pale grey. Excipulum entire, completely carbonized, 30–50 μm wide and thick at the base, black; laterally covered by corticate algiferous thallus including clusters of crystals; hypothecium prosoplectenchymatous, 5–10 μm high, colorless. Hymenium 70–100 μm high, colorless, clear; epithecium granulose, 5–10 μm high, brownish; paraphyses unbranched; asci fusiform, 70–100 × 15–20 μm. Ascospores 8/ascus, ellipsoid, transversely 3-septate, 15–20 × 5–6 μm, 3–3.5 times as long as wide, colorless, I + violet-blue. Secondary chemistry: no substances detected by TLC (thallus surface or exposed medulla P–; thallus section K–).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata 8-09-canopy (F, holotype; USM, isotype), 01-01-D, FM-01d, FM-03 g (F, USM paratypes).

Etymology: This new species is dedicated to Dr. Nigel Pitman, former director of the Centro de Investigación y Capacitación Río Los Amigos (CICRA) or Los Amigos Research and Training Center, for his invaluable contributions to the study of tropical ecosystems.

Notes: This new species is unusual in the genus Graphis in having very small, 3-septate ascospores (Lücking 2009; Lücking et al. 2009a). The only other species in the genus with such small ascospores is G. tetralocularis C. Bock & Hauck from tropical Africa, but that species has a laterally carbonized excipulum and supposedly contains traces of atranorin (Bock and Hauck 2005). Another similar species is G. sitiana Vain., which has 3–5-septate ascospores, white-pruinose labia and a more or less ecorticate thallus (Lücking et al. 2009a).

Leucodecton inspersum Rivas Plata & Lücking, spec. nova

(Fig. 4C–D)

MycoBank: MB800204

Etymology: The epithet refers to the inspersed hymenium.

Differing from Leucodecton expallescens in the inspersed hymenium and larger ascospores.

Thallus corticolous, pale green-grey, up to 5 cm diam. and 80–120 μm thick, continuous; surface verrucose, with dense, prosoplectenchymatous cortex; photobiont Trentepohlia, with angular-rounded to elongate cells 7–10 × 5–7 μm large; photobiont layer and medulla incrusted with large clusters of calcium oxalate crystals; medulla white. Apothecia rounded, immersed-erumpent, 0.15–0.25 mm diam.; disc partially covered by 0.1–0.2 mm wide pore, pale brown to flesh-colored, very thinly white-pruinose; margin entire, white, proper excipulum distinctly free (double margin). Columella absent. Excipulum paraplectenchymatous, light brown to brown; periphysoids absent; hypothecium prosoplectenchymatous, 5–10 μm high, colorless. Hymenium 100–130 μm high, inspersed; paraphyses unbranched; asci fusiform, 100–120 × 15–22 μm. Ascospores 4–8/ascus, ellipsoid, muriform, 25–40 × 12–15 μm, with slightly thickened septa and rectangular lumina, colorless, I– (non-amyloid). Secondary chemistry: stictic, constictic, cryptostictic, and hypostictic acids (exposed medulla P + orange; thallus section with K + yellow efflux).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata 8-extra (F, holotype), 8-03-A (F, paratype).

Notes: Leucodecton inspersum is most similar to L. expallescens (Nyl.) Rivas Plata & Lücking, which also has small, colorless, muriform ascospores and stictic acid, but the ascospores in that species are much smaller (12–20 × 6–8 μm), the hymenium is not inspersed, and the thallus lacks distinct verrucae and features a loose rather than dense cortex (Rivas Plata et al. 2010a). The thallus morphology of L. inspersum is indeed unusual in the genus and only L. dactyliferum (Hale) Lücking has a verrucose thallus and L. compunctum (Ach.) A. Massal. a prosoplectenchymatous cortex. The only other species with inspersed hymenium are L. bisporum (Nyl.) Sipman & Lücking and L. oxysporum (Redinger) Rivas Plata & Lücking, but both have brown ascospores and differ further in thallus morphology.

Ocellularia cicra Rivas Plata & Lücking, spec. nova

(Fig. 4E–F)

MycoBank: MB800205

Etymology: The epithet honors the official name of Los Amigos Biological Station: Centro De Investigación Y Capacitación Río Los Amigos (CICRA).

Differing from Ocellularia thryptica in the inspersed hymenium.

Thallus corticolous, light olive-green, up to 5 cm diam. and 50–80 μm thick, continuous; surface smooth to uneven, with dense, prosoplectenchymatous cortex; photobiont Trentepohlia, with angular-rounded to elongate cells 7–10 × 5–8 μm large; photobiont layer and medulla with scattered clusters of calcium oxalate crystals; medulla white. Apothecia rounded to angular in outline, erumpent, 0.4–0.6 mm diam.; disc covered by 0.2–0.3 mm wide pore and columella; margin entire, covered by thalline layer up to the pore. Columella present, simple, apically to upper half carbonized. Excipulum prosoplectenchymatous, apically to upper half carbonized; periphysoids absent; hypothecium prosoplectenchymatous, 10–15 μm high, colorless. Hymenium 80–100 μm high, thinly but distinctly inspersed; paraphyses unbranched; asci fusiform, 80–100 × 15–20 μm. Ascospores 8/ascus, ellipsoid, 5-septate, 15–20 × 6–8 μm, 2–3 times as long as wide, with thick septa and lens-shaped lumina, colorless, I + violet-blue. Secondary chemistry: protocetraric and virensic acids (exposed medulla P + orange-red; thallus section K–).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata 1-09-A (F, holotype), 1-08-B, 1-08-E, 1-canopy, 8-04-B, 8-04-D, 8-05-A, 8-05-C, 8-05-D, 8-06-A, 8-08-A, 8-10-B (F, USM, paratypes).

Notes: Ocellularia cicra belongs in a complex of species with small, transversely septate, hyaline ascospores and protocetraric acid as major secondary compound, most of these being present in the collections studied here. So far, none of the species in this group is known to have an inspersed hymenium. Except for hymenial inspersion, the most similar species is O. thryptica, which agrees in the carbonization of excipulum and columella. Most other species in this group, including the common O. perforata, have a colorless to brown excipulum and columella or, such as O. violacea, a non-carbonized excipulum combined with a carbonized columella. Ocellularia gymnocarpa has both the excipulum and columella carbonized but the latter is broad-stump-shaped. Ocellularia fumosa agrees with O. cicra in apothecial anatomy and hymenial inspersion but has a pigmented medulla and otherwise lacks lichen substances.

Ocellularia fenestrata Rivas Plata & Lücking, spec. nova

(Fig. 4G–H)

MycoBank: MB800206

Etymology: The epithet refers to fenestrate columella, an unusual feature in the genus.

Differing from Ocellularia psorbarroensis in the smaller, erumpent apothecia.

Thallus corticolous, olive-green, up to 5 cm diam. and 60–100 μm thick, continuous; surface smooth to uneven, with dense, prosoplectenchymatous cortex; photobiont Trentepohlia, with angular-rounded to elongate cells 7–12 × 5–8 μm large; photobiont layer lacking crystals except near the apothecia; medulla indistinct, white. Apothecia rounded to angular in outline, immersed-erumpent, 0.4–0.6 mm diam.; disc covered by 0.2–0.3 mm wide pore and simple to fenestrate columella; margin entire, covered by thalline layer except around the pore, appearing as pale to brown rim. Columella present, simple to irregularly fenestrate and connecting to the lateral excipulum, non-carbonized, colorless to brown in upper parts. Excipulum prosoplectenchymatous, light brown to brown; periphysoids absent; hypothecium prosoplectenchymatous, 10–15 μm high, colorless. Hymenium 90–100 μm high, clear; paraphyses unbranched; asci fusiform, 90–100 × 15–20 μm. Ascospores 8/ascus, ellipsoid, 5-septate, 15–20 × 6–8 μm, 2–3 times as long as wide, with thick septa and lens-shaped lumina, colorless, I + violet-blue. Secondary chemistry: psoromic, subpsoromic and 2′-O-demethylpsoromic acids (exposed medulla P + yellow; thallus section K–).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata 8-extra (F, holotype; USM, isotype).

Notes: This new species belongs in a complex of species characterized by olive-green thalli, immersed-erumpent to prominent apothecia, small, transversely septate, hyaline ascospores and variously columellate and carbonized apothecia. Within this group, several species produce psoromic acid: Ocellularia terebrata and several other species feature a distinctly carbonized excipulum and columella; O. minutula Hale, O. pluripora Hale, and O. pluriporoides Homchantara & Coppins have a carbonized columella contrasting with a pale, non-carbonized excipulum. The only species with uncarbonized excipulum and columella producing psoromic acid are the isidiate O. isidioalbula Mangold and O. psorbarroensis. The latter agrees with the new species in most aspects but its apothecia are much larger and prominent and the columella, while becoming irregular, is not distinctly fenestrate. Molecular data (Rivas Plata et al. 2012c) confirm that O. psorbarroensis and O. fenestrata are different taxa. Morphologically most similar, including an often fenestrate columella, is O. mauritiana, which differs in producing protocetraric acid. The new species should not be confused with Myriotrema glaucophaenum (Kremp.) Hale and M. costaricense (Müll. Arg.) Hale, which have prominent apothecia with colorless excipulum and irregular pseudocolumella and are genetically not closely related to Ocellularia (Rivas Plata et al. 2012c).

Ocellularia microsorediata Rivas Plata & Lücking, spec. nova

(Fig. 5A–B)
https://static-content.springer.com/image/art%3A10.1007%2Fs13225-012-0172-y/MediaObjects/13225_2012_172_Fig5_HTML.gif
Fig. 5

A–B. Ocellularia microsorediata (holotype). C. Ocellularia plicata (holotype). D. Ocellularia protoinspersa (holotype). E–F. Ocellularia pustulata (holotype). G. Ocellularia natashae (paratype). H. Thelotrema amazonicum (holotype)

MycoBank: MB800207

Etymology: The epithet refers to the small soredia produced on the thallus surface.

Differing from Ocellularia calvescens in the sorediate thallus and partially carbonized excipulum.

Thallus corticolous, light green, up to 3 cm diam. and 50–80 μm thick, continuous; surface smooth to uneven, with dense, prosoplectenchymatous cortex, abundantly sorediate; soralia 0.1–0.2 mm diam., at first maculate but soon becoming confluent to form irregular to reticulate patches (sometimes starting out as elongate soralia along thallus cracks), white, granular; photobiont Trentepohlia, with angular-rounded to elongate cells 7–11 × 5–7 μm large; photobiont layer and medulla incrusted with clusters of calcium oxalate crystals; indistinct, medulla white. Apothecia rounded to irregular in outline, erumpent, 0.4–0.8 mm diam.; disc covered by narrow, 0.05–0.1 mm wide pore; margin entire, covered by thalline layer but often irregularly sorediate. Columella absent. Excipulum prosoplectenchymatous, dark brown to carbonized in outermost layers; periphysoids absent; hypothecium prosoplectenchymatous, 10–15 μm high, colorless. Hymenium 120–160 μm high, clear; paraphyses unbranched; asci fusiform, 120–150 × 15–20 μm. Ascospores 8/ascus, ellipsoid, 5–7-septate, 20–25 × 6–8 μm, 3–4 times as long as wide, with thick septa and lens-shaped lumina, colorless, I + violet-blue. Secondary chemistry: psoromic, subpsoromic and 2′-O-demethylpsoromic acids (exposed medulla P + yellow; thallus section K–).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata 1-07-C (F, holotype), 1-03-A, 1-07-E, 1-08-C, 1-10-A, 1-10-E, 8-04-E, 8-10-B (F, USM, paratypes).

Notes: The new species is characterized by the sorediate thallus in combination with ecolumellate apothecia, small and transversely septate, colorless ascospores, and psoromic acid as major secondary compound. Only few species in Ocellularia are known to produce soralia, viz. O. africana Frisch, O. discoidea (Ach.) Müll. Arg., O. flavosorediata Frisch, and O. sorediata Hale. All four produce columellate apothecia and differ in further details: O. africana produces protocetraric acid and has prominent apothecia with wider pore; O. discoidea, while having the same chemistry as O. microsorediata, has larger apothecia with wide pore and fully carbonized excipulum and (broad-stump-shaped) columella, as well as larger ascospores; O. flavosorediata Frisch differs in producing hypoprotocetraric acid plus a yellow medullary pigment and has fully carbonized excipulum and columella, in addition to larger ascospores; and O. sorediata lacks secondary substances and has smaller apothecia and smaller, 3-septate ascospores only (the statement in Frisch 2006: 201 for O. sorediata to produce protocetraric acid and having a reticulate columella is incorrect). Apart from the soralia, O. microspora is most similar to O. calvescens (Fée) Müll. Arg. in morphology, anatomy, and chemistry, but that species also differs in having a fully carbonized excipulum. Another similar sorediate species with psoromic acid, Myriotrema pulverulentum (Hale) Hale, has apothecia with larger pore, unpigmented excipulum, and irregular pseudocolumella.

Ocellularia natashae Rivas Plata & Lücking, spec. nova

(Fig. 5G)

MycoBank: MB800211

Etymology: With this new species we honor our daughter Natasha.

Differing from Ocellularia xantholeuca in the white medulla and larger ascospores.

Thallus corticolous, olive-green, up to 3 cm diam. and 60–90 μm thick, continuous; surface uneven to coarsely verrucose, with dense, prosoplectenchymatous cortex; photobiont Trentepohlia, with angular-rounded to elongate cells 8–12 × 5–8 μm large; photobiont layer and medulla incrusted with clusters of calcium oxalate crystals; medulla white. Apothecia rounded to irregular in outline, erumpent, 0.4–0.6 mm diam.; disc covered by 0.15–0.25 mm wide pore and white-tipped columella; margin entire, forming a dark brown rim around the pore, covered by thalline layer, irregularly bumpy to verrucose. Columella present, finger-like, upper half carbonized. Excipulum prosoplectenchymatous, upper half carbonized; periphysoids absent; hypothecium prosoplectenchymatous, 10–15 μm high, colorless. Hymenium 120–160 μm high, strongly inspersed; paraphyses unbranched; asci fusiform, 100–140 × 15–20 μm. Ascospores 8/ascus, oblong, 11–15-septate, 30–50 × 8–10 μm, with thick septa and lens-shaped lumina, colorless, I + violet-blue. Secondary chemistry: hirtifructic and conhirtifructic acid and low (main) cinchonarum unknown (exposed medulla P–; thallus section K–).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata 1-extra (F, holotype), 1-08-C, 8-01-C (F, paratypes).

Notes: Ocellularia natashae is characterized by a strongly inspersed hymenium, in combination with the rare chemistry of hirtifructic and and conhirtifructic acid and the main cinchonarum unknown. Hymenial inspersion is otherwise only known from O. circa, newly described above, with protocetraric acid and smaller ascospores, O. fumosa (Ach.) Müll. Arg., which has a pale yellow-orange medulla but otherwise lacks secondary compounds, O. khuntanense (Homchantara & Coppins) Lumbsch & Papong, which differs in having a pale excipulum and columella and producing psoromic acid, O. rimosa Hale, which lacks a columella and secondary compounds altogether, and O. xantholeuca Müll. Arg., which has a pale yellow-orange medulla and in addition the same chemical compounds as the new species, O. natashae. Ocellularia xantholeuca was considered a chemical strain (II) of O. fumosa by Mangold et al. (2009), but the presence of such rare substances as hirtifructic and conhirtifructic acid with cinchonarum (versus no substances in O. fumosa) warrants the recognition as separate species. Our new species, O. natashae, is chemically very similar to O. xantholeuca but lacks the medullary pigment; in addition, the apothecia in O. xantholeuca are more regularly rounded; and the ascospores are smaller (25–40 μm) and have less septa (5–9).

Ocellularia plicata Rivas Plata & Lücking, spec. nova

(Fig. 5C)

MycoBank: MB800208

Etymology: The epithet refers to the ridged thallus appearing folded.

Differing from Ocellularia wirthii in the folded thallus.

Thallus corticolous, yellow-white, up to 5 cm diam. and 70–120 μm thick, continuous; surface stongly uneven to irregularly ridged, with thin, prosoplectenchymatous cortex; photobiont Trentepohlia, with angular-rounded to elongate cells 7–10 × 5–7 μm large; photobiont layer lacking crystals except near the apothecia; medulla white. Apothecia rounded to angular in outline, erumpent, 0.4–0.7 mm diam.; disc covered by narrow, 0.1–0.2 mm wide pore and simple columella; margin entire, covered by thalline layer except around the pore, appearing as pale to brown rim. Columella present, simple, carbonized in upper half. Excipulum prosoplectenchymatous, carbonized in upper half; periphysoids absent; hypothecium prosoplectenchymatous, 10–15 μm high, colorless. Hymenium 100–120 μm high, clear; paraphyses unbranched; asci fusiform, 100–120 × 20–25 μm. Ascospores 8/ascus, oblong-ellipsoid, 7-septate, 25–35 × 7–9 μm, 3–4 times as long as wide, with thick septa and lens-shaped lumina, colorless, I + violet-blue. Secondary chemistry: psoromic, subpsoromic and 2′-O-demethylpsoromic acids (exposed medulla P + yellow; thallus section K–).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata 8-09-canopy (F, holotype).

Notes: Ocellularia plicata is a further new species in a morphotype group formed by taxa with with small, transversely septate, hyaline ascospores and psoromic acid. With several other species in the group it shares the yellow-white rather than olive-green thallus with strongly sculptured surface: Ocellularia antillensis Hale and O. wirthii Mangold, Elix & Hale (Mangold et al. 2008) have a verrucose thallus, whereas in O. albobullata Lücking, Sipman & Grube the thallus is strongly bullate and the excipulum and columella lack carbonization. The new species appears most similar to O. wirthii but is genetically distinct (Rivas Plata et al. 2012c); apart from the ridged rather than verrucose thallus, another feature separating it are the comparatively larger ascospores which are unusually large for a species in this group.

Ocellularia protoinspersa Rivas Plata & Lücking, spec. nova

(Fig. 5D)

MycoBank: MB800209

Etymology: The epithet refers to the inspersed hymenium in combination with protocetraric acid as secondary substance.

Differing from Ocellularia cicra in the carbonized columella and excipulum.

Thallus corticolous, light olive-green, up to 5 cm diam. and 60–80 μm thick, continuous; surface uneven to finely verrucose, with dense, prosoplectenchymatous cortex; photobiont Trentepohlia, with angular-rounded to elongate cells 8–12 × 5–8 μm large; photobiont layer and medulla with clusters of calcium oxalate crystals; medulla white. Apothecia rounded, prominent, 0.5–0.8 mm diam.; disc covered by 0.2–0.3 mm wide pore and columella; margin entire, covered by thalline layer up to the pore. Columella present, simple to broad-stump-shaped, completely carbonized. Excipulum prosoplectenchymatous, upper half carbonized; periphysoids absent; hypothecium prosoplectenchymatous, 10–15 μm high, colorless. Hymenium 120–150 μm high, strongly and densely inspersed; paraphyses unbranched; asci fusiform, 100–120 × 12–15 μm. Ascospores 8/ascus, oblong, 7-9-septate, 30–40 × 6–8 μm, 4.5–5.5 times as long as wide, with thick septa and lens-shaped lumina, colorless, I + violet-blue. Secondary chemistry: protocetraric and virensic acids (exposed medulla P + orange-red; thallus section K–).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata 8-10-E (F, holotype).

Notes: This new species also produces an inspersed hymenium, a rare feature within Ocellularia and only known from a handful of species (see discussion under O. cicra above and O. natashae below). The only other inspersed species with protocetraric acid is O. cicra; however, O. protoinspersa differs from that taxon by the carbonized excipulum and carbonized, broad columella, as well as the smaller ascospores. The new species is very similar to O. gymnocarpa in apothecial morphology and chemistry, including the broad columella, but the latter has a clear hymenium and smaller ascospores. Ocellularia protoinspersa resembles a species of Ampliotrema, but the latter genus never produces a columella.

Ocellularia pustulata Rivas Plata & Lücking, spec. nova

(Fig. 5E–F)

MycoBank: MB800210

Etymology: The epithet refers to the pustulate thallus surface.

Differing from Ocellularia microsorediata in the pustulate thallus.

Thallus corticolous, light green-grey to olive-grey, up to 5 cm diam. and 50–100 μm thick, continuous; surface uneven to irregularly verrucose, with dense, prosoplectenchymatous cortex, abundantly pustulate-sorediate; pustules 0.2–0.3 mm diam. and up to 0.5 mm high, resembling dactyls, irregular to sometimes grouped into 2–3 or branched, eventually breaking up at the tip and becoming sorediate, soredia of the same color as the thallus (not white); photobiont Trentepohlia, with angular-rounded to elongate cells 7–12 × 5–7 μm large; photobiont layer and medulla incrusted with clusters of calcium oxalate crystals; indistinct, medulla white. Apothecia rounded to irregular in outline, prominent, 0.5–1 mm diam.; disc covered by 0.1–0.3 mm wide pore; margin entire, covered by thalline layer up to the pore, often irregularly verrucose or pustulate. Columella absent. Excipulum prosoplectenchymatous, brown to dark brown; periphysoids absent; hypothecium prosoplectenchymatous, 10–15 μm high, colorless. Hymenium 140–170 μm high, clear; paraphyses unbranched; asci fusiform, 120–150 × 15–18 μm. Ascospores 8/ascus, elliopsoid, 5–7-septate, 20–25 × 7–10 μm, with thick septa and lens-shaped lumina, colorless, I + violet-blue. Secondary chemistry: psoromic, subpsoromic and 2′-O-demethylpsoromic acids (C–, K–, P + yellow).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 70 ° 6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata 1-08-C (F, holotype).

Notes: This is another new species of Ocellularia with psoromic acid and vegetative propagules. In many aspects, including the ascospores, it is similar to O. microsorediata, but differs clearly in the pustulate-dactyliferous thallus with the pustules partially breaking up to form soredia at the tips, whereas in O. microsorediata the thallus is sorediate from the beginning; also, the color of the soredia is different and O. pustulata has more prominent apothecia with a less carbonized, lighter brown excipulum. To our knowledge, there is only one other thelotremoid species in Graphidaceae with pustulate-dactyliferous thallus, Wirthiotrema santessonii (Hale) Rivas Plata & Frisch. Apart from having smaller, more immersed apothecia and a different ascospore type, that species differs chemically in producing stictic acid (Frisch 2006; Rivas Plata et al. 2010b).

Thelotrema amazonicum Rivas Plata & Lücking, spec. nova

(Fig. 5H)

MycoBank: MB800212

Etymology: The epithet refers to the unusual ecogeography for a Thelotrema species.

Differing from Thelotrema pseudosubtile in the completely ecorticate, farinose thallus.

Thallus corticolous, light grey-green, up to 10 cm diam. and 30–50 μm thick, continuous; surface farinose, ecorticate; photobiont Trentepohlia, with angular-rounded to elongate cells 7–10 × 5–7 μm large; photobiont layer incrusted with numerous clusters of calcium oxalate crystals; medulla indistinct, white. Apothecia angular-rounded, immersed, 0.25–0.4 mm diam.; disc partially covered by excipulum, pale brown, white-pruinose; proper margin separated from thalline margin by distinct split (double margin), fissured, brown; thalline margin irregular, brown, white-pruinose. Columella absent. Excipulum paraplectenchymatous, colorless; periphysoids present; hypothecium prosoplectenchymatous, 5–10 μm high, colorless. Hymenium 100–120 μm high, clear; paraphyses unbranched; asci fusiform, 100–120 × 15–20 μm. Ascospores 8/ascus, ellipsoid, 9–13-septate, 40–50 × 7–8 μm, 5–6 times as long as wide, with thick septa and lens-shaped lumina, colorless, I + violet-blue. Secondary chemistry: no substances detected by TLC (thallus P–; thallus section K–).

Material examined: Peru: Madre de Dios: Los Amigos Research and Training Center, Centro de Investigación y Capacitación Río Los Amigos (CICRA), 90 km W of Puerto Maldonado; 12 ° 35′ S, 7n6′ W, 270 m; tropical lowland rain forest, on bark of tree in secondary forest; Aug 2008, Rivas Plata 8-10-B (F, holotype).

Notes: This species shares the ascospore type and lack of secondary substances with a number of species in Thelotrema: T. defossum (Müll. Arg.) Mangold, T. lathraeum Tuck., T. pseudosubtile Mangold, T. suecicum (H. Magn.) P. James, and T. subtile Tuck. (Rivas Plata et al. 2010a; Sipman et al. 2012). All these have a distinct, loose to dense (T. lathreum) cortex and the thallus surface appears always compact and never farinose. The apothecia in T. suecicum are prominent and larger, and the apothecia of T. pseudosubtile and T. subtile are also larger. Except for T. pseudosubtile, all species have smaller ascospores, and in T. defossum, which otherwise agrees in the small, immersed apothecia, the ascospore have thinner septa and are only weakly amyloid. All species except T. pseudosubtile are also tropical-montane to extratropical and are not expected to occur in lowland rain forest. Thelotrema amazonicum superficially resembles Chapsa albomaculata and C. hypostictica found at the same locality but these have larger apothecia with more exposed disc, lack a double margin, and the apothecial margin is erect to recurved; also the chemistry is different.

Sterile species of Graphidaceae

Seven distinct species of Graphidaceae were found in the sterile condition only, producing either soralia or isidoid papillae. Their phylogenetic placement was determined using molecular methods where possible, and all sequenced material clustered within the Ocellularia-clade (Rivas Plata et al. 2012c). However, sequences were not obtained from Chapsa spec. 1, 2, and 3, which on account of their chemistry and thallus morphology might belong in the genus Chapsa. The taxa can be distinguished by the nature of their propagules, thallus morphology, and secondary chemistry:

Chapsa sp. sorediate 1: thallus with excavate soralia, ecorticate, yellow-green, hypoconstictic acid (Rivas Plata 8-03-C, 8-03-D, 8-10-B).

Chapsa sp. sorediate 2: thallus with white soralia, ecorticate, brown, cryptostictic acid (Rivas Plata 1-06-D, 1-07-E, 1-08-A, 1-08-B, 1-09-A).

Chapsa sp. sorediate 3: thallus with green-white soralia, corticate, light green, cryptostictic acid (Rivas Plata 1-08-B).

Myriotrema sp. isidiate-sorediate: thallus with white, excavate soralia producing papilliform isidia, corticate, light green, no substances (Rivas Plata 8-09-canopy).

Ocellularia sp. papillose: thallus with isidioid, white papillae, green-grey, loose cortex, protocetraric acid (Rivas Plata 1-07-E, 1-08-A, 8-08-D, 8-08-E).

Ocellularia sp. sorediate 1: thallus with yellow-white soralia, ecorticate, brown, protocetraric acid (Rivas Plata 1-03-C, 8-08-D).

Ocellularia sp. sorediate 2: thallus with white soralia, corticate, olive-brown, two unknown substances (Rivas Plata 1-04-C, 1-08-C).

Ocellularia sp. sorediate 3: thallus with linear, green-white soralia, corticate, light green, protocetraric acid (Rivas Plata 1-03-B, 1-06-D, 8-09-canopy).

Acknowledgements

This study was made possible by several grants provided by the United States National Science Foundation (NSF) to The Field Museum: “Phylogeny and Taxonomy of Ostropalean Fungi” (DEB 0516116; PI Lumbsch, Co-PI Lücking); “Neotropical Epiphytic Microlichens” (DEB 0715660; PI Lücking), and “ATM – Assembling a taxonomic monograph: The lichen family Graphidaceae” (DEB 1025861; PI Lumbsch, Co-PI Lücking). Fieldwork was possible thanks to the doctoral thesis grant: “Estructura de la comunidad de liquenes crustosos, con enfasis en la familia Thelotremataceae, basada en especificidad de forofitos, microclima y nivel de disturbio forestal en la Concesión de Conservación Los Amigos (sureste peruano)” (MOORE Y7 10330; PI Rivas Plata), awarded by the Asociación para la Conservación de la Cuenca Amazónica (ACCA). The first author is also grateful to a stipend from the Lester Armour Fund through the Field Museum which enabled part of the laboratory and data analysis work. Curators of the herbaria in B, BM, CANB, FH, G, H, M, NY, S, TUR, UPS, US, and W provided loans of or access to valuable type material and other significant collections.

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© Mushroom Research Foundation 2012