Ten new species of lichenized Basidiomycota in the genera Dictyonema and Cora (Agaricales: Hygrophoraceae), with a key to all accepted genera and species in the Dictyonema clade

Abstract

As part of a larger systematic and taxonomic revision, including molecular phylogenetic analysis, of lichenized Basidiomycota in the Dictyonema clade, ten species are described as new from tropical America, seven in the foliose genus Cora and three in the filamentous genus Dictyonema: Cora arachnoidea J. E. Hern. & Lücking, sp. nov., C. aspera Wilk, Lücking & E. Morales, sp. nov., C. byssoidea Lücking & Moncada, sp. nov., C. cyphellifera Dal-Forno, Bungartz & Lücking, sp. nov., C. inversa Lücking & Moncada, sp. nov., C. squamiformis Wilk, Lücking & Yánez-Ayabaca, sp. nov., C. strigosa Lücking, E. Paz & L. Salcedo, sp. nov., Dictyonema aeruginosulum Lücking, Nelsen & Will-Wolf, sp. nov., D. diducens Nyl. ex Lücking, sp. nov., D. metallicum Lücking, Dal-Forno & Lawrey, sp. nov., and D. obscuratum Lücking, Spielmann & Marcelli, sp. nov. We discuss the taxonomic status of the six names historically established for species belonging in the genus Cora and reinstate the names C. gyrolophia Fr., C. pavonia (Sw.) Fr., and C. reticulifera Vain., providing diagnostic features for these, whereas the status of C. glabrata (Spreng.) Fr. and C. bovei Speg. remains uncertain. The following new combinations are introduced: Cora hirsuta (Moncada & Lücking) Moncada & Lücking, comb. nov., C. minor (Lücking, E. Navarro & Sipman) Lücking, comb. nov., Corella melvinii (Chaves, Lücking & Umaña) Lücking, Dal-Forno & Lawrey, comb. nov., Cyphellostereum phyllogenum (Müll. Arg.) Lücking, Dal-Forno & Lawrey, comb. nov., Dictyonema caespitosum (Johow) Lücking, comb. nov., D. irrigatum (Berk. & M. A. Curtis).
Lücking, comb. nov., D. phyllophilum (Parmasto) Lücking, Dal-Forno & Lawrey, comb. et stat. nov., and D. scabridum
(Vain.) Lücking,
comb. et stat. nov. Keys are presented to the five currently accepted genera and 40 currently recognized
species in the genera Cyphellostereum, Dictyonema, Cora, and Corella.

Full text

Accepted by Mohammad Sohrabi: 15 Sept. 2013; published: 24 Oct. 2013 1
PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Copyright © 2013 Magnolia Press
Phytotaxa 139 (1): 1–38 (2013)
www.mapress.com/phytotaxa/Article
http://dx.doi.org/10.11646/phytotaxa.139.1.1
Ten new species of lichenized Basidiomycota in the genera Dictyonema and Cora
(Agaricales: Hygrophoraceae), with a key to all accepted genera and species in
the Dictyonema clade
ROBERT LÜCKING1, MANUELA DAL-FORNO2, JAMES D. LAWREY2, FRANK BUNGARTZ3,
MARÍA E. HOLGADO ROJAS4, JESÚS E. HERNÁNDEZ M.5, MARCELO P. MARCELLI6, BIBIANA
MONCADA7, EDUARDO A. MORALES8, MATTHEW P. NELSEN1,9, ELIAS PAZ10, LUIS SALCEDO11,
ADRIANO A. SPIELMANN12, KARINA WILK13, SUSAN WILL-WOLF14 & ALBA YÁNEZ-AYABACA3
1Integrative Science and Collections, The Field Museum, 1400 South Lake Shore Drive, Chicago, Illinois 60605, U.S.A.;
email: rlucking@fieldmuseum.org
2Department of Environmental Science and Policy, George Mason University, 4400 University Drive, Fairfax, VA 22030-4444, U.S.A.
3Biodiversity Assessment, Charles Darwin Foundation (AISBL), Puerto Ayora, Santa Cruz, Galápagos, Ecuador
4Facultad de Ciencias Biologicas, Universidad Nacional de San Antonio Abad del Cusco, Av. de la Cultura, Nro. 733, Cusco, Perú
5Fundación Instituto Botánico de Venezuela, División de Plantas no Vasculares, Sección Hongos y Líquenes, Ave. Salvador Allende,
Jardín Botánico de Caracas, Universidad Central de Venezuela, Apartado # 2156, Caracas 1010-A, Venezuela
6Instituto de Botânica, Núcleo de Pesquisa em Micologia, Caixa Postal 68041, São Paulo/SP, CEP 04045-972, Brazil
7Licenciatura en Biología, Universidad Distrital Francisco José de Caldas, Cra. 4 No. 26B-54, Torre de Laboratorios, Herbario,
Bogotá, Colombia
8Herbario Criptogámico, Universidad Católica Boliviana San Pablo, Calle M. Márquez esq. Plaza Jorge Trigo s/n, P.O. Box 5381,
Cochabamba, Bolivia
9Committee on Evolutionary Biology, University of Chicago, 1025 E. 57th Street, Chicago, Illinois 60637, U.S.A.
10Facultad de Ciencias Biologicas, Universidad Nacional de San Antonio Abad del Cusco, Av. de la Cultura, Nro. 733, Cusco, Perú
11Laboratorio de Ecofisiología Vegetal, Facultad de Ciencias Naturales y Matemática, Universidad Nacional Federico Villarreal,
Lima, Perú
12Laboratório de Botânica, Departamento de Biologia, Centro de Ciências Biológicas e da Saúde, Universidade Federal de Mato
Grosso do Sul, Cidade Universitária, Caixa Postal 549, CEP 79070-900, Campo Grande, Mato Grosso do Sul, Brazil
13Laboratory of Lichenology, W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, PL-31-512 Kraków, Poland
14Department of Botany, University of Wisconsin-Madison, 430 Lincoln Dr., Madison, Wisconsin 53706, U.S.A.
Abstract
As part of a larger systematic and taxonomic revision, including molecular phylogenetic analysis, of lichenized
Basidiomycota in the Dictyonema clade, ten species are described as new from tropical America, seven in the foliose
genus Cora and three in the filamentous genus Dictyonema: Cora arachnoidea J. E. Hern. & Lücking, sp. nov., C. aspera
Wilk, Lücking & E. Morales, sp. nov., C. byssoidea Lücking & Moncada, sp. nov., C. cyphellifera Dal-Forno, Bungartz
& Lücking, sp. nov., C. inversa Lücking & Moncada, sp. nov., C. squamiformis Wilk, Lücking & Yánez-Ayabaca, sp.
nov., C. strigosa Lücking, E. Paz & L. Salcedo, sp. nov., Dictyonema aeruginosulum Lücking, Nelsen & Will-Wolf, sp.
nov., D. diducens Nyl. ex Lücking, sp. nov., D. metallicum Lücking, Dal-Forno & Lawrey, sp. nov., and D. obscuratum
Lücking, Spielmann & Marcelli, sp. nov. We discuss the taxonomic status of the six names historically established for
species belonging in the genus Cora and reinstate the names C. gyrolophia Fr., C. pavonia (Sw.) Fr., and C. reticulifera
Vain., providing diagnostic features for these, whereas the status of C. glabrata (Spreng.) Fr. and C. bovei Speg. remains
uncertain. The following new combinations are introduced: Cora hirsuta (Moncada & Lücking) Moncada & Lücking,
comb. nov., C. minor (Lücking, E. Navarro & Sipman) Lücking, comb. nov., Corella melvinii (Chaves, Lücking &
Umaña) Lücking, Dal-Forno & Lawrey, comb. nov., Cyphellostereum phyllogenum (Müll. Arg.) Lücking, Dal-Forno &
Lawrey, comb. nov., Dictyonema caespitosum (Johow) Lücking, comb. nov., D. irrigatum (Berk. & M. A. Curtis)
Lücking, comb. nov., D. phyllophilum (Parmasto) Lücking, Dal-Forno & Lawrey, comb. et stat. nov., and D. scabridum
(Vain.) Lücking, comb. et stat. nov. Keys are presented to the five currently accepted genera and 40 currently recognized
species in the genera Cyphellostereum, Dictyonema, Cora, and Corella.

LÜCKING ET AL.2 • Phytotaxa 139 (1) © 2013 Magnolia Press
Key words: Acantholichen, Bolivia, Brazil, Colombia, Corella, Costa Rica, Cyphellostereum, Ecuador, Mexico,
Panama, paramo, Peru, puna
Introduction
Most lichenized fungi belong in Ascomycota, while few Basidiomycota are lichenized, being mainly found in
the orders Agaricales, Cantharellales, Hymenochaetales, and Lepidostromatales (Oberwinkler 1970, 2012;
Redhead et al. 2002; Hibbett et al. 2007; Lawrey et al. 2009; Hodkinson & Lücking 2013). The family
Hygrophoraceae (Agaricales) contains two diverse, lichenized clades: Lichenomphalia Redhead, Lutzoni,
Moncalvo & Vilgalys (Redhead et al. 2002: 38) s.lat. and Dictyonema C. Agardh ex Kunth (Kunth 1822: 1)
s.lat. (Lawrey et al. 2009), forming either agaricoid-omphalinoid mushrooms associated with green algae or
cyphelloid to stereoid-corticioid basidiomata lichenized with cyanobacteria (Oberwinkler 1970, 2012;
Parmasto 1978; Redhead et al. 2002; Chaves et al. 2004; Lawrey et al. 2009). Some authors considered
Dictyonema s.lat. to represent several different genera in different families, based on features related to
growth form, presence of clamp connections, and nature of the photobiont (Hariot 1891, 1892; Metzner
1934). Foliose forms were usually treated as Cora Fr. (Fries 1825: 300) and Corella Vain. (Vainio 1890: 242),
whereas filamentous forms were assigned to either Laudatea Johow (Johow 1884: 398), Dictyonema, or
Rhipidonema Mattir. (Mattirolo 1881: 265), depending on appressed or shelf-like growth and the absence or
presence of clamp connections. Other workers included such forms, at least partially, as ontogenetic and
ecological variations within a single species (Möller 1893; Oberwinkler 1970; Parmasto 1978; Larcher &
Vareschi 1988). Oberwinkler (1970, 1980, 1984, 2001, 2012) provided excellent morphological and
anatomical treatments of Dictyonema and related groups, discussing the taxonomic value of certain
characters, and Parmasto (1978) eventually accepted five species in a single genus, Dictyonema.
Until recently, Dictyonema s.lat. was best known by the two supposedly common and widespread tropical
montane species, the foliose D. glabratum (Spreng.) D. Hawksw. (Hawksworth 1988: 101) [= Cora pavonia
(Sw.) Fr. (Fries 1825: 300, 1838: 556)] and the filamentous D. sericeum (Sw.) Berk. (Berkeley 1843: 639) (see
Mitidieri et al. 1964; Feige 1969; Oberwinkler 1970, 1984, 2001; Parmasto 1978; Coxson 1987a–c; Fritz-
Sheridan & Portecop 1987; Iacomini et al. 1987; Fritz-Sheridan 1988; Larcher & Vareschi 1988; Wolf 1993;
Lange et al. 1994; Piovano et al. 1995; Thomas et al. 1997; Azenha et al. 1998; Trembley et al. 2002a, b;
Carbonero et al. 2002; Elifio et al. 2002). Taxonomic and phylogenetic studies, however, suggest that both
names comprise a fairly large number of species, many undescribed (Chaves et al. 2004; Lücking 2008;
Lawrey et al. 2009; Yánez et al. 2012; Dal-Forno et al. 2013). This also has implications on the correct use of
the epithets glabratum and pavonia for foliose forms, for which Hawksworth (1988) pointed out that, in case
of synonymy, glabratum would be the correct usage, being sanctioned through its adoption by Fries (1821).
Molecular analyses suggested that Dictyonema s.lat. can be divided into five genera, Cyphellostereum D.
A. Reid (Reid 1965: 336), Dictyonema s.str., Acantholichen P. M. Jørg. (Jørgensen 1998: 444), Cora, and
Corella (Lawrey et al. 2009; Dal-Forno et al. 2013), and this concept was applied to a treatment of Galápagos
basidiolichens (Yánez et al. 2012). These genera are well-distinguished morphologically and anatomically
(Oberwinkler 1984, 2012): Cyphellostereum and Dictyonema have filamentous thalli with the mycobiont
forming a hyphal sheath around the cyanobacterial filaments, the sheath consisting of irregular hyphae
(lacking haustoria) in Cyphellostereum and of jigsaw-puzzle-shaped cells (forming haustoria) in Dictyonema
s.str. Cyphellostereum has cyphelloid basidiomata emerging from the undifferentiated lichenized thallus,
whereas Dictyonema s.str. has stereoid-corticioid basidiomata that develop from the underside of the
lichenized thallus. Acantholichen forms a microsquamulose thallus, while species of Cora and Corella have
foliose-macrosquamulose thalli forming distinct layers (cortex, photobiont layer and medulla) and producing
corticioid basidiomata on the lobe underside. Corella brasiliensis Vain. (Vainio 1890: 243) was regarded by
Parmasto (1978) as a juvenile or underdeveloped form of Dictyonema pavonia (= glabratum) of no taxonomic
value. While morphologically similar to Cora (Vainio 1890; Metzner 1934; Oberwinkler 1970; Xavier Filho
& Vicente 1979), this taxon in reality forms a separate genus sister to Acantholichen (Dal-Forno et al. 2013),
supported by its different cortex structure.

Phytotaxa 139 (1) © 2013 Magnolia Press • 3
DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
As a result of our ongoing phylogenetic studies in the Dictyonema clade, we provide here a first revision
of the taxonomy of foliose and appressed-filamentous forms, clarifying the status of several historic names
available for this group, and describing ten species as new in the genera Cora and Dictyonema.
Material and Methods
Fresh material for this study was collected during field work in Bolivia, Brazil, Colombia, Costa Rica,
Ecuador, and Mexico, mostly in the framework of a Neotropical workshop project by RL and a phylogenetic
revision of Dictyonema s.lat. by JDL, RL and MDF. In addition, we revised a large number of herbarium
specimens housed at the Instituto de Biodiversidad in Costa Rica (INB), the Universidad Distrital Francisco
José Caldas in Colombia (UDBC), and collections from several parts of the Neotropics at the Field Museum
of Natural History in Chicago (F) and the US National Herbarium at the Smithsonian Museum of Natural
History in Washington (US). We also examined historical type material from BM, F, G, H, PC, TUR, UPS, and
W. Most specimens were studied at The Field Museum using standard techniques of light microscopy and
thin-layer chromatography (Orange et al. 2001; Lumbsch 2002). We used a standardized protocol for
morphological, anatomical, and chemical characters to describe each specimen.
For each species, we also cited the ITS barcode sequence, following recommendations by Kõljalg et al.
(2013).
Taxonomic Treatment
Parmasto (1978) recognized only a single species of Dictyonema with a foliose, corticate thallus, viz. D.
glabratum (Spreng.) D. Hawksw. The discovery of two further species, D. minus Lücking, E. Navarro &
Sipman (Chaves et al. 2004: 247) and D. hirsutum Moncada & Lücking (Lumbsch et al. 2011: 48), together
with the results of molecular phylogenetic analyses (Lawrey et al. 2009; Dal-Forno et al. 2013), suggested
that this taxon actually contains many different species. It therefore seemed prudent to revise historically
published names in this group.
Notably, among the over 50 species described in Dictyonema s.lat. (Parmasto 1978), only six represent
the genus Cora in being foliose with upper cortex of the Cora type (Dal-Forno et al. 2013). Thelephora
pavonia Sw. (Swartz 1806: 1930) is a replacement name for Ulva montana Sw. from Jamaica (Swartz 1788:
148), an illegitimate name antedated by U. montana Lightfoot (Lightfoot 1777). This name is also cited as T.
pavonia Weber & D. Mohr (Weber & Mohr 1805; Fries 1838, Saccardo 1888b), but this is an error since
Weber & Mohr (1805) did not publish a separate name or species, but referred to Swartz's (1806) upcoming
publication on the replacement name: "... Thaelaephora pavonia, oder ehemaligen Ulva montana Swartz, ..."
(Weber & Mohr 1805: 326). Thelephora glabrata Spreng. was described from Guadeloupe (Sprengel 1820:
51) and Cora gyrolophia Fr. from Mauritius (Fries 1838: 556). The latter name is based on material invalidly
published as Gyrolophium elegans (G. 'mauritianum') by Kunze in Von Krombholz (1831: 76, tab 5, fig. 16).
Cora bovei Speg. was established on material from Tierra del Fuego in Argentina (Spegazzini 1888: 169), C.
reticulifera Vain. was described from Brazil (Vainio 1890: 241), and Wainiocora ciferrii Tomas. from Panama
(Tomaselli 1950: 106); the latter in a separate genus supposedly due to a different photobiont, which turned
out to be an incorrect observation (Oberwinkler 1970; Parmasto 1978).
Thelephora pavonia (≡ Ulva montana) and Wainiocora ciferrii appear to represent the same taxon, as far
as can be judged from the type specimens, here accepted as Cora pavonia (Sw.) Fr. [≡ C. pavonia (Weber &
D. Mohr) Fr., nom. illeg.]. Cora pavonia is a relatively large species growing on soil between mosses with a
concentrically undulate, grey-brown surface and a hymenophore forming narrow, elongate, concentric lines
with slightly involute margins, common and widespread in Neotropical paramos (Fig. 1). The name Cora
pavonia is commonly attributed to Fries (1825), but was validly published later (Fries 1838), since in 1825,
Fries did not associate the epithet pavonia with the name Cora (ICN Art. 35.2). In 1838, he cited the wrong

LÜCKING ET AL.4 • Phytotaxa 139 (1) © 2013 Magnolia Press
authorship and page number for the basionym, as "Weber et Mohr, Beytr. 1, p. 236" (Fries 1838: 556), an error
repeated by Saccardo (1888b), but this does not preclude valid publication of the combination (ICN Art.
41.3), especially since the name mentioned in Weber & Mohr (1805), as mentioned above, is an indirect
reference to Swartz (1806).
FIGURE 1. Cora pavonia. A. Specimen in the field (Colombia, Lücking s.n.). B. Lobe underside enlarged showing hymenophore
with involute margins (Venezuela, Hernández 1778). C. Part of the lectotype of Ulva montana (≡ Thelephora pavonia) in BM showing
undulate lobe surface and underside with hymenophore. Scale in A = 10 mm, in B–C = 1 mm.

Phytotaxa 139 (1) © 2013 Magnolia Press • 5
DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
FIGURE 2. Cora spp. A–B. Cora glabrata (lectotype of Thelephora glabrata in UPS). C. Cora bovei (isotype in NY). D–F. C.
reticulifera. D. Lobe underside with hymenophore (isotype in BM). E. Lobe showing upper side (Ecuador, Cole 123). F. Lobe
underside with hymenophore (Ecuador, Lücking 26201). Scale in A–C = 5 mm, in D, F = 1 mm, in E = 10 mm.
The type of Thelephora glabrata is not well-developed (Fig. 2A–B), but represents a probably rare or
locally confined species in the Caribbean growing on more or less bare soil that we have not currently
recollected with certainty. It is composed of 5–10 semicircular lobes per thallus, with the lobes 1–2 cm wide
and 1–3 cm long, unbranched or once branched, but lacking radial branching sutures, white to pale yellowish
to greyish brown in the herbarium. The upper surface is glabrous except for scattered, unbranched, up to 0.3

LÜCKING ET AL.6 • Phytotaxa 139 (1) © 2013 Magnolia Press
mm long trichomes near the lobe margins; the involute margin is very minutely arachnoid to almost glabrous.
The lower surface is finely felty-arachnoid, becoming yellowish white in the herbarium. The thallus is 200–
300 µm thick in section, with the upper cortex formed by a 25–50 µm thick layer of rather loosely packed,
irregular hyphae supported by a 20–30 µm high 'medullary' layer of spaced groups of densely packed,
anticlinal hyphae. The photobiont layer is 100–150 µm thick; clamp connections were not observed. No
hymenophore is developed in the type.
Cora gyrolophia is a larger, epiphytic species on palm trunks similar to C. pavonia, but with grey rather
than brown color (Kunze in Von Krombholz 1831: 76, plate 5: 16). The species was originally and invalidly
published as Gyrolophium elegans, and the Index Fungorum also carries the name G. 'mauritianum' [IF
439913], referring to the same original publication, but the epithet mauritianum is not mentioned there.
Because of its ecology and distribution, we consider Cora gyrolophia a distinct species, but fresh material is
required to clarify its status, if the species still exists. Cora bovei is a rather small species growing on soil in
southern South America, but the type material is not well-developed (Fig. 2C) and fresh collections are
required to elucidate its status; its cortex is similar to the new species C. squamiformis described below. Cora
reticulifera also grows on soil, but is a species of montane forest rather than paramo; the hymenophore
consists of numerous minute, flat patches connected in reticulate fashion (Fig. 2D–F). This species has been
recollected and sequenced (Dal-Forno et al. 2013); it was found that the peculiar morphology of the
hymenophore is a good character. This means that currently we accept C. gyrolophia, C. pavonia, and C.
reticulifera as good species with distinctive character and tentatively accept C. bovei and C. glabrata pending
further studies.
Considering the taxonomic changes necessary in the recognition of species in the genus Cora, we are now
in the curious situation that this taxon is one of the best studied tropical lichens in terms of ecomorphology,
ecophysiology, and biochemistry (Mitidieri et al. 1964; Feige 1969; Oberwinkler 1970, 1984, 2001; Parmasto
1978; Coxson 1987a–c; Fritz-Sheridan & Portecop 1987; Iacomini et al. 1987; Fritz-Sheridan 1988;
Hawksworth 1988; Larcher & Vareschi 1988; Wolf 1993; Lange et al. 1994; Piovano et al. 1995; Thomas et
al. 1997; Azenha et al. 1998; Trembley et al. 2002a, b; Carbonero et al. 2002; Elifio et al. 2002). It is even
cited (as Dictyonema glabratum) in the "Listing of Interesting Plants of the World" [http://
www.newcrops.uq.edu.au/listing/species_pages_D/Dictyonema_glabratum.htm]. However, without revising
the material used in these studies, it is impossible to ascertain which species were actually investigated.
In order to facilitate identification of currently recognized taxa, we have added keys to the five accepted
genera and to species of Cyphellostereum, Dictyonema, Cora, and Corella, following the new species
descriptions.
Cora arachnoidea J. E. Hern. & Lücking, sp. nov. (Fig. 3)
Mycobank #805376
Genbank ITS barcoding sequence: KF443233
Differing from the morphologically similar and closely related Cora hirsuta in the larger thallus and lobes with brown
color when fresh and the shorter, arachnoid tomentum on the upper surface.
Holotype:—VENEZUELA. Mérida: Parque Nacional Sierra Nevada, surroundings of Laguna de Mucubají;
8º 47' N, 70º 49' W, 3626 m; 6 December 2009, Hernández 1780 (VEN).
Thallus terricolous between bryophytes, rarely epiphytic on bryophyte-laden branches, foliose, up to 10
cm across, composed of 5–10 semicircular lobes per thallus; lobes 1–3(–5) cm wide and 1–5(–7) cm long,
unbranched or once branched but lacking radial branching sutures, greyish brown to brown with slight
concentric color zonation when fresh, with thickened, involute, white margins, becoming pale yellowish grey
in the herbarium. Upper surface densely and shortly arachnoid-hirsute over entire surface (barely visible when
fresh); trichomes densely interwoven basally but apically free and irregularly arranged, 0.2–0.3 mm long and

Phytotaxa 139 (1) © 2013 Magnolia Press • 7
DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
25–50 µm thick at the base, composed of loosely agglutinated hyphae; involute margin with underside very
minutely arachnoid; lower surface ecorticate, finely felty-arachnoid (representing the exposed medulla), white
when fresh and becoming yellowish white in the herbarium. Thallus in section 250–350 µm thick, with upper
cortex, photobiont layer, and medulla; upper cortex formed by a 25–50 µm thick layer of rather loosely
FIGURE 3. Cora arachnoidea. A. Specimen in the field (Colombia, Lücking 32700). B. Lobe enlarged showing tomentose upper
surface (holotype). C. Lobe underside showing hymenophore with involute, byssoid margins (Venezuela, Hernández 1782). Scale in
A, C = 5 mm, in B = 1 mm.

LÜCKING ET AL.8 • Phytotaxa 139 (1) © 2013 Magnolia Press
packed, periclinal, 4–5 µm thick hyphae supported by a 20–30 µm high 'medullary' layer of spaced groups of
densely packed, anticlinal, 3–5 µm thick hyphae; photobiont layer 50–150 µm thick, irregular, composed of
clusters of short, coiled cyanobacterial filaments wrapped in a dense, paraplectenchymatous hyphal sheath
formed by jigsaw puzzle-shaped cells, clusters 20–30 µm diam., individual photobiont cells 10–12 µm broad
and 6–8 µm long, dark blue-green to yellow-green in upper portions, penetrated by tubular fungal hyphae;
heterocytes sparse, hyaline to pale yellow, 8–10 µm wide and 5–6 µm long; cells of hyphal sheath wavy in
lateral outline, 3–4 µm thick; medulla 50–100 µm thick, composed of loosely woven, irregularly arranged to
more or less periclinal hyphae 4–5 µm thick; clamp connections not observed.
Hymenophore developed as irregular to angular or elongate, resupinate patches dispersed on the
underside, patches 3–10 mm diam., with pale yellow, smooth surface and byssoid margins; hymenophore in
section 50–100 µm thick, composed of a paraplectenchymatous layer resting on loose, 4–6 µm thick,
generative medullary hyphae and supporting the hymenium; hymenium composed of numerous, palisade-like
basidioles and scattered basidia; basidioles 20–30 × 5–6 µm; basidia 25–35 × 5–7 µm, 4-sterigmate;
basidiospores (few seen) ellipsoid, non-septate, hyaline, 7–8 × 2.5–3.5 µm.
Chemistry: no substances detected by TLC.
Distribution and Ecology:—This species is known from several collections from Costa Rica, Colombia,
Venezuela, and Bolivia; it is probably widespread in the northern Andes and the Costa Rican Cordilleras. It is
a typical paramo species, mostly growing on soil between bryophytes in exposed situations.
Etymology:—The epithet refers to the arachnoid tomentum on the upper surface.
Remarks:—Cora arachnoidea is the second species known with a tomentose surface, after Cora hirsuta
(Moncada & Lücking) Moncada & Lücking, comb. nov. [Mycobank #805388; bas.: Dictyonema hirsutum
Moncada & Lücking in Lumbsch et al., Phytotaxa 18: 48 (2011); holotype: Colombia, Lücking 25900
(UDBC; isotype: F!)]. The latter differs from C. arachnoidea in the smaller thallus and lobes furnished with a
much thicker tomentum easily visible even when hydrated, and a zonate margin with an olive-green, glabrous,
submarginal zone and a white, tomentose margin (Lumbsch et al. 2011). Cora arachnoidea is a good example
how markedly specimens can differ in the living, hydrated stage compared to rather non-descript herbarium
material, a possible explanation why this genus has been a stumbling block for lichenologists and mycologists
in the past and only a single species has been recognized by most authors (Parmasto 1978; Hawksworth 1988;
Oberwinkler 2001). Field images are practically indispensable for correct identifications in this genus.
Additional specimens examined:—MEXICO. Jalisco: Parque Nacional Volcán Nevado de Colima,
beyond entrance station in La Joya area near the campground; 19° 35' N, 103° 36' W, 3415 m; 26 December
2006, Egan 17538 (OMA). COSTA RICA. Sán José: Los Santos Forest Reserve, Cerro de la Muerte
(Pacífico Central Conservation Area), Talamanca Ridge, km 90 on road (ruta 2) from Cartago to San Isidro,
access road to towers on summit; 83° 45' W, 9° 34' N, 3400–3500 m; upper montane cloud forest and
subalpine paramo zone, disturbed low paramo shrub with Chusquea, on bryophyte, exposed; September 2007,
Lücking R18 (F). COLOMBIA. Cundinamarca: Páramo de Sumapaz, Laguna de Chizacá; 4° 17' N, 74° 12'
W, 3700–3750 m; wet paramo with Espeletia; August 2010, Lücking 32700 (F, UDBC). VENEZUELA.
Venezuela. Mérida: Parque Nacional Sierra Nevada, surroundings of Laguna de Mucubají; 8º 47' N, 70º 49'
W, 3626 m; 6 December 2009, Hernández 1779, 1782 (VEN). BOLIVIA. Santa Cruz: Caballero, Siberia
region near La Palma; 17° 49' S, 64° 40' W, 2582 m; Yungas cloud forest, epiphytic on bark; 12 December
2004, Wilk 2780a (KRAM).
Cora aspera Wilk, Lücking & E. Morales, sp. nov. (Fig. 4)
Mycobank #805377
Genbank ITS barcoding sequence: KF443231
Differing from the superficially similar Cora arachnoidea in the absence of a dense, distinct upper tomentum and in the
epiphytic growth habit, and from the closely related C. pavonia in the plane lobe surface with scattered, indistinct
upper tomentum, and in the epiphytic growth habit.

Phytotaxa 139 (1) © 2013 Magnolia Press • 9
DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
Holotype:—BOLIVIA. Santa Cruz: Caballero, Siberia region near La Palma; 17° 49' S, 64° 40' W, 2582 m;
Yungas cloud forest, epiphytic on bark; 12 December 2004, Wilk 2780b (KRAM; isotype: LPB).
FIGURE 4. Cora aspera. A. Specimen in the field (Colombia, Lücking 33332). B. Lobe enlarged showing rough upper surface
(holotype). C. Lobe underside showing hymenophore with slightly involute, smooth margins (holotype). Scale in A–B = 10 mm, in C
= 1 mm.
Thallus epiphytic on twigs and branches of trees, foliose, up to 7 cm across, composed of 1–5 semicircular
lobes per thallus; lobes 1–5 cm wide and 1–5 cm long, often branched and with short radial branching sutures,
light greenish grey with slight concentric color zonation when fresh, with thin but distinct, involute, white to
light grey margins, becoming white to (dark) grey in the herbarium. Upper surface rough and thinly scabrose

LÜCKING ET AL.10 • Phytotaxa 139 (1) © 2013 Magnolia Press
in thin, concentric lines but lacking a continuous, distinct tomentum; trichomes where present in concentric
lines, free and irregularly arranged, 0.1–0.15 mm long and 5–10 µm thick at the base, composed of
agglutinated hyphae; involute margin usually shortly pilose; lower surface ecorticate, finely felty-arachnoid
(representing the exposed medulla) to almost glabrous, light grey when fresh and becoming white in the
herbarium. Thallus in section 200–300 µm thick, with upper cortex, photobiont layer, and medulla; upper
cortex formed by a 25–50 µm thick layer of rather loosely packed, irregularly arranged to nearly periclinal, 4–
5 µm thick hyphae supported by an indistinct, 20–30 µm high 'medullary' layer of spaced groups of densely
packed, anticlinal, 3–5 µm thick hyphae; photobiont layer 70–120 µm thick, composed of clusters of short,
coiled cyanobacterial filaments wrapped in a dense, paraplectenchymatous hyphal sheath formed by jigsaw
puzzle-shaped cells, clusters 20–30 µm diam., individual photobiont cells 10–13 µm broad and 5–8 µm long,
dark blue-green to lighter green in upper portions, penetrated by tubular fungal hyphae; heterocytes sparse,
hyaline to pale yellow, 9–12 µm wide and 5–6 µm long; cells of hyphal sheath wavy in lateral outline, 3–4 µm
thick; medulla 50–100 µm thick, composed of loosely woven, irregularly arranged to more or less periclinal
hyphae 4–5 µm thick; clamp connections not observed.
Hymenophore developed as elongate, resupinate patches forming more or less concentric ridges on the
underside, patches 1–10 mm long and 0.5–1 mm broad, with pale yellow, smooth surface and smooth,
involute margins; hymenophore in section 50–100 µm thick, composed of a paraplectenchymatous layer
resting on loose, 4–6 µm thick, generative medullary hyphae and supporting the hymenium; hymenium
composed of numerous, palisade-like basidioles and scattered basidia; basidioles 20–35 × 5–6 µm; basidia
25–40 × 6–7 µm, 4-sterigmate; basidiospores not observed.
Chemistry: no substances detected by TLC.
Distribution and Ecology:—This species is known from several collections from Costa Rica, Colombia,
Ecuador, Bolivia, and Peru. It appears to be a primarily epiphytic species, growing on twigs and branches of
trees and shrubs in (upper) montane rain forest and paramo vegetation, where it competes with other foliose
macrolichens such as Leptogium spp., Lobariella spp., and Sticta spp.
Etymology:—The epithet refers to the rough appearance of the surface especially when dry.
Remarks:—Parmasto (1978) and other authors (Mitidieri et al. 1964; Feige 1969; Oberwinkler 1970,
1984, 2001; Parmasto 1978; Coxson 1987a–c; Fritz-Sheridan & Portecop 1987; Iacomini et al. 1987; Fritz-
Sheridan 1988; Hawksworth 1988; Larcher & Vareschi 1988; Wolf 1993; Lange et al. 1994; Piovano et al.
1995; Thomas et al. 1997; Azenha et al. 1998; Trembley et al. 2002a, b; Carbonero et al. 2002; Elifio et al.
2002) considered Dictyonema glabratum (including Cora pavonia) to be a species with wide distribution and
broad ecological amplitude, being found on a wide range of substrata. The data now available indicate that
this is not the case. The many species recognized phylogenetically and morphologically also have distinct
substrate preferences, growing either on bare soil, among bryophytes, on rock, or epiphytic on branches,
rarely on tree trunks. Cora aspera is one of a few species growing typically as an epiphyte and it is thus far the
largest and most common epiphytic species known in the genus. It resembles the distantly related C.
arachnoidea in dry condition but can be distinguished by the lack of a dense tomentum covering the entire
upper surface and by the much finer, almost reticulate hymenophore. The latter is similar to that found in the
more closely related C. pavonia, but that species differs by its terrestrial growth in bryophyte mats and its
distinctly brownish color when fresh, as well as its coarsely undulate surface.
Additional specimens examined:—COSTA RICA. Puntarenas: Coto Brus, San Vito, Las Cruces
Biological Station and Botanical Garden; September 2007, Lücking 21016 (F). BOLIVIA. La Paz: Murillo,
Valle de Zongo, Laguna de Viscachani, a las orillas de la laguna; 16º 12' S, 68º 08' W, 3805 m; piso altoandino
con pajonales y vegetación baja, 13 November 2007, Lücking 23564 (F, LPB). Cochabamba: Chapare,
Incachaca; 17° 13' S, 65° 50' W, 2018 m; 7 July 2009, Lücking 29128 (F, HCUCB). Cochabamba: Chapare,
Corani; 17° 16' S, 65° 54' W, 3262 m; 7 July 2009, Lücking 29356, 29364 (F, HCUCB). PERU. Cuzco: Aguas
Calientes, near Machu Picchu; August 2009, Ve ra s .n . (F).

Phytotaxa 139 (1) © 2013 Magnolia Press • 11
DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
Cora byssoidea Lücking & Moncada, sp. nov. (Fig. 5)
Mycobank #805378
Genbank ITS barcoding sequence: KF443234
Differing from the morphologically similar Cora hirsuta in the only marginally present, arachnoid tomentum and the
epiphytic growth, and from the closely related C. inversa in the distinct upper tomentum and the absence of soredia.
FIGURE 5. Cora byssoidea. A. Aspect of typical habitat in the Colombian paramo near Bogotá. B. Lobe enlarged showing arachnoid-
byssoid upper surface (holotype). C. Lobe underside showing hymenophore with strongly involute margins (holotype). Scale in B–C =
1 mm.

LÜCKING ET AL.12 • Phytotaxa 139 (1) © 2013 Magnolia Press
Holotype:—COLOMBIA. Cundinamarca: Choachí, Páramo El Verjón; 4º 33' N, 74º 00' E; 3200 m; 18
August 2008, Lücking 25901 (F).
Thallus epiphytic on thin branches and twigs of paramo shrubs, foliose, up to 3 cm across, composed of
1–3 semicircular lobes per thallus; lobes 1–2 cm wide and 1–2 cm long, unbranched, light grey when fresh,
with thickened, involute, grey margins, becoming white to pale yellowish grey in the herbarium. Upper
surface glabrous except for a broad submarginal zone with appressed, arachnoid-byssoid tomentum;
trichomes densely interwoven and irregularly arranged, 0.1–0.2 mm long and 5–6 µm thick at the base,
composed of single hyphae; involute margin with underside minutely arachnoid; lower surface ecorticate,
finely felty-arachnoid (representing the exposed medulla), white when fresh and becoming yellowish white in
the herbarium. Thallus in section 250–400 µm thick, with upper cortex, photobiont layer, and medulla; upper
cortex formed by a 50–100 µm thick layer of rather loosely woven, irregularly arranged, 4–6 µm thick hyphae
supported by a 30–50 µm high 'medullary' layer of irregularly arranged to anticlinal, 4–6 µm thick hyphae,
towards the margin no such distinction visible and the upper cortex entirely formed by loosely woven,
irregularly arranged hyphae causing the tomentose appearance; photobiont layer 100–200 µm thick, irregular,
composed of clusters of short, coiled cyanobacterial filaments wrapped in a dense, paraplectenchymatous
hyphal sheath formed by jigsaw puzzle-shaped cells, clusters 30–50 µm diam., individual photobiont cells 10–
12 µm broad and 6–8 µm long, dark blue-green to yellow-orange in upper portions, penetrated by tubular
fungal hyphae; heterocytes sparse, hyaline to pale yellow, 8–10 µm wide and 5–6 µm long; cells of hyphal
sheath wavy in lateral outline, 3–4 µm thick; medulla 30–50 µm thick, composed of loosely woven,
irregularly arranged to more or less periclinal hyphae 4–5 µm thick; clamp connections not observed.
Hymenophore developed as irregular to elongate, resupinate patches dispersed on the underside, patches
1–3 mm long and 0.5–1 mm broad, with pale yellow, smooth surface and strongly involute, smooth margins;
hymenophore in section 50–100 µm thick, composed of a paraplectenchymatous layer resting on loose, 4–6
µm thick, generative medullary hyphae and supporting the hymenium; hymenium composed of numerous,
palisade-like basidioles and scattered basidia; basidioles 25–30 × 5–7 µm; basidia 25–35 × 5–8 µm, 4-
sterigmate; basidiospores ellipsoid, non-septate, hyaline, 7–9 × 3–4 µm.
Chemistry: no substances detected by TLC.
Distribution and Ecology:—This species is known from a single collection growing on a shrub in the
Colombian paramo regions. Due to its small size, it is certainly overlooked.
Etymology:—The epithet refers to the arachnoid-byssoid submarginal tomentum on the upper surface.
Remarks:—This is another new species with partially tomentose upper surface. It is most similar to Cora
hirsuta (Lumbsch et al. 2011), which was found at the same locality, but differs in the nature of the tomentum,
which is formed by erect trichomes of agglutinated hyphae in C. hirsuta and by an irregularly dissolved
cortical layer of single hyphae in C. byssoidea. Also, whereas C. hirsuta has a glabrous submarginal zone,
with the tomentum developed towards the center of the lobes, in C. byssoidea the tomentum is only seen close
to the margin. The two species are actually not closely related and fall in two different clades within the genus
(Dal-Forno et al. 2013). The sister species of C. byssoidea is C. inversa (see below), which differs markedly
in its upper surface being glabrous and in the irregular lobe margins producing dark soredia.
Cora cyphellifera Dal-Forno, Bungartz & Lücking, sp. nov. (Fig. 6)
Mycobank #805379
Genbank ITS barcoding sequence: KF443242
Differing from Cora pavonia in the light aeruginous color and pitted surface, the stereoid-cyphelloid hymenophore, and
the epiphytic growth habit, and from the closely related C. arachnoidea in the pitted, undulate, otherwise glabrous
surface, the stereoid-cyphelloid hymenophore, and the epiphytic growth habit.
Holotype:—ECUADOR. Imbabura: Andes, Cantón Cotacachi; 22º 29.8' N, 78º 27' 24.6'' W; 2053 m; small

Phytotaxa 139 (1) © 2013 Magnolia Press • 13
DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
entrance driveway towards the Reserva Alto Chocó near Intag, just before the small bridge over the river; 26
June 2012, Dal-Forno 1808 (GMUF).
FIGURE 6. Cora cyphellifera. A. Specimen in the field (holotype). B. Lobe underside showing cyphelloid hymenophores (holotype).
C. Aspect of habitat at type locality in Ecuador. Scale in A–B = 10 mm.
Thallus epiphytic on twigs and branches of trees, foliose, up to 15 cm across, composed of 20–30
semicircular lobes per thallus; lobes 3–5 cm wide and 2–3 cm long, lacking branching sutures, light

LÜCKING ET AL.14 • Phytotaxa 139 (1) © 2013 Magnolia Press
aeruginous with slight concentric color zonation when fresh, with shallow concentric ridges (8–11 per cm lobe
length) and shallowly but distinctly pitted, with thin but distinct, involute, white to light grey margins,
becoming light yellowish grey to dark grey in the herbarium. Upper surface glabrous; involute margin finely
arachnoid; lower surface ecorticate, glabrous, light aeruginous when fresh and becoming light yellowish grey
in the herbarium. Thallus in section 285–400 µm thick, with upper cortex, photobiont layer, and medulla;
upper cortex formed by a 25–35 µm thick layer of rather densely packed, periclinal, 4–5 µm thick hyphae
supported by an indistinct, 80–120 µm high 'medullary' layer of spaced groups of densely packed, anticlinal,
3–5 µm thick hyphae; photobiont layer 60–80 µm thick, composed of clusters of short, coiled cyanobacterial
filaments wrapped in a dense, paraplectenchymatous hyphal sheath formed by jigsaw puzzle-shaped cells,
clusters 40–70 µm diam., individual photobiont cells 8–11 µm broad and 6–8 µm long, bluish green to
orange-yellow in upper portions, penetrated by tubular fungal hyphae; heterocytes sparse, hyaline to pale
yellow, 9–12 µm wide and 5–6 µm long; cells of hyphal sheath wavy in lateral outline, 3–4 µm thick; medulla
100–200 µm thick, composed of loosely woven, irregularly arranged to more or less periclinal hyphae 4–5 µm
thick; clamp connections not observed.
Hymenophore developed as stereoid to cyphelloid structures irregularly dispersed along the margins on
the underside, 5–10 mm long and 10–15 mm broad, with white, smooth surface and smooth margins;
hymenophore in section 70–100 µm thick, composed of a paraplectenchymatous layer resting on loose, 4–6
µm thick, generative medullary hyphae and supporting the hymenium; hymenium composed of numerous,
palisade-like basidioles and scattered basidia; basidioles 20–35 × 5–8 µm; basidia 18–25 × 7–9 µm, 4-
sterigmate; basidiospores ellipsoid to lacrymoid, non-septate, hyaline, 7–8 × 2.5–3.5 µm.
Chemistry: no substances detected by TLC.
Distribution and Ecology:—This species is known from the type collection in a montane rain forest in
northern Ecuador, where it was found growing on small trees in open, disturbed forest patches.
Etymology:—The epithet refers to the unusual type of hymenophore.
Remarks:—This remarkable new species is characterized by its distinctly aeruginous color, the pitted
surface, and particularly the hymenophore becoming cyphelloid, differing markedly from all other species of
the genus, including the closely related C. arachnoidea (see above). Cora pavonia (see above) also has an
undulate lobe surface, but is brownish in the field, lacks pits, has a corticioid hymenophore, and always grows
terrestrial between bryophytes. In contrast to other species of Cora where the hymenophore is corticioid and
evenly distributed on the lobe underside, in C. cyphellifera it almost looks like the lichenized thallus is
parasitized by a non-lichenized, cyphelloid mushroom. The hymenophore is very similar to the basidiomata
found in the related genus Cyphellostereum (Lawrey et al. 2009). In his lengthy account on what he
considered ecomorphological variation of a single species, Möller (1893) reported Cora lichens with bluish
thalli that produced cyphelloid basidiomata, concluding that supposedly 'free-living' basidiomata and those
that are lichenized and form Cora thalli represent the same fungal species. It is very likely that he had
observed the same species as described here and did not consider the possibility that different fungal species
can form very similar fruiting bodies.
Cora inversa Lücking & Moncada, sp. nov. (Fig. 7)
Mycobank #805380
Genbank ITS barcoding sequence: KF443237
Differing from Cora hirsuta and the closely related C. byssoidea in the lobes with glabrous upper surface, tomentose-
strigose lower surface, and submarginally produced soredia, and from C. minor in the larger lobes and submarginally
formed soredia.
Holotype:—COLOMBIA. Cundinamarca: Choachí, Páramo El Verjón; 4º 33' N, 74º 00' E; 3200 m; 18
August 2008, Lücking 25902 (F).

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DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
FIGURE 7. Cora inversa. A. Specimen in the field showing irregular-fuzzy lobe margins and clusters of trichomes projecting from
lower surface (Colombia, Lücking 33340). B. Lobes enlarged showing marginal soredia (Colombia, Lücking 33308). C. Lobe
underside showing arachnoid surface (Colombia, Lücking 25902). Scale in A–B = 10 mm, in C = 1 mm.
Thallus epiphytic between bryophytes on thin branches and twigs of paramo shrubs or at the base on
mossy soil, foliose, up to 5 cm across, composed of 1–5(–10) semicircular lobes per thallus; lobes 1–3 cm
wide and 1–3 cm long, unbranched, white when fresh, with thickened, involute, irregular to fuzzy, white
margins and a narrow, dark, submarginal zone forming granular soredia, white in the herbarium. Upper
surface glabrous; involute margin with underside arachnoid-strigose; dark olive-brown submarginal zone

LÜCKING ET AL.16 • Phytotaxa 139 (1) © 2013 Magnolia Press
forming soredia composed of cyanobacterial granules 30–50 µm in diam. embedded in a
paraplectenchymatous hyphal sheath; lower surface ecorticate, finely arachnoid (representing the exposed
medulla) to distinctly hirsute-strigose in parts caused by the formation of clusters of longer trichomes
composed of agglutinated hyphae, white when fresh and becoming yellowish white in the herbarium. Thallus
in section 200–300 µm thick, with upper cortex, photobiont layer, and medulla; upper cortex formed by a 50–
100 µm thick layer of loosely woven, irregularly arranged, 4–6 µm thick hyphae covered by a thin layer of
distinctly periclinal, compacted hyphae and supported by a 30–50 µm high 'medullary' layer of irregularly
arranged to anticlinal, 4–6 µm thick hyphae; photobiont layer 50–100 µm thick, irregular, composed of
clusters of short, coiled cyanobacterial filaments wrapped in a dense, paraplectenchymatous hyphal sheath
formed by jigsaw puzzle-shaped cells, clusters 30–50 µm diam., individual photobiont cells 9–12 µm broad
and 5–6 µm long, yellow-orange to olive-yellow in upper portions, penetrated by tubular fungal hyphae;
heterocytes sparse, hyaline to pale yellow, 8–10 µm wide and 4–5 µm long; cells of hyphal sheath wavy in
lateral outline, 3–4 µm thick; medulla 30–50 µm thick, composed of loosely woven, irregularly arranged to
more or less periclinal hyphae 4–5 µm thick; clamp connections not observed.
Hymenophore not observed.
Chemistry: no substances detected by TLC.
Distribution and Ecology:—This species known from several collections growing on shrubs,
particularly at their base, in the Colombian paramo.
Etymology:—The epithet refers to the partially strigose underside as opposed to a similar tomentum
formed on the upper side by Cora hirsuta.
Remarks:—Cora inversa is one of several species producing soredia in the genus, most of which being
undescribed. Sorediate margins are also known from Cora minor (Lücking, E. Navarro & Sipman)
Lücking, comb. nov. [Mycobank #805389; bas.: Dictyonema minus Lücking, E. Navarro & Sipman in
Chaves et al., Bryologist 107: 247 (2004); holotype: Costa Rica, Navarro 1688 (INB-3789873; isotypes, CR,
F)], but in that species they are formed directly on the involute margin, whereas in C. inversa, they are formed
in a thin submarginal zone on the upper side. In contrast to most other species of Cora, the lobes in C. inversa
are not perfectly round but slightly irregular, together with the white surface and dark submarginal zone
giving the species a very characteristic appearance. Cora hirsuta and the closely related C. byssoidea (see
above) differ in the tomentose upper surface and the regularly rounded lobe margins lacking soredia;
herbarium material can easily be confused if soredia are indistinct and the upper and lower surface are not
properly recognized.
Additional specimens examined:—COLOMBIA. Cundinamarca: Choachí, Páramo El Verjón; 4º 33'
N, 74º 00' E; 3200 m; 18 August 2008, Lücking 25903 (F).
Cora squamiformis Wilk, Lücking & Yánez-Ayabaca, sp. nov. (Fig. 8)
Mycobank #805382
Genbank ITS barcoding sequence: KF443240
Differing from the morphologically similar Cora bovei in the smaller lobes with plane surface, and from the closely
related C. pavonia in the much smaller, often irregularly bent lobes giving the thallus a squamulose appearance.
Holotype:—BOLIVIA. La Paz: Franz Tamayo, Madidi National Park, Sanchez Pass between Pelechuco and
Keara; 14° 43' S, 69° 08' W, 4677 m; high mountain vegetation, on ground between mosses; 13 October 2007,
Wilk 7577 (KRAM; isotypes: F, LPB).
Thallus on soil between bryophytes, macrosquamulose, up to 3 cm across, composed of 3–5(–10)
semicircular lobes per thallus; lobes 0.5–1 cm wide and 0.5–1 cm long, unbranched or sparsely branched,
olive-grey to grey with indistinct color zonation when fresh, with thickened, involute, white margins, darker
grey to brownish-grey in the herbarium; lobes ascending and typically with much bent, sinous margins

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DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
('salad'-like). Upper surface glabrous but appearing rough; involute margin with underside finely arachnoid;
lower surface ecorticate, arachnoid (representing the exposed medulla), white when fresh and becoming grey-
white in the herbarium. Thallus in section 200–300 µm thick, with upper cortex, photobiont layer, and
medulla; upper cortex much reduced, formed by a 20–50 µm thick layer of loosely woven, irregularly
arranged, 4–6 µm thick hyphae and a 15–25 µm thick layer of strongly compacted, periclinal hyphae 3–5 µm
FIGURE 8. Cora squamiformis. A. Specimen in the field (Colombia, Lücking s.n.). B. Specimen with strongly bent and sinuose
margins (Colombia, Lücking s.n.). C. Specimen in the herbarium (holotype). Scale in A = 10 mm, in B–C = 5 mm.

LÜCKING ET AL.18 • Phytotaxa 139 (1) © 2013 Magnolia Press
thick and with brownish color, 'medullary' layer absent; photobiont layer 50–100 µm thick, irregular,
composed of clusters of short, coiled cyanobacterial filaments wrapped in a dense, paraplectenchymatous
hyphal sheath formed by jigsaw puzzle-shaped cells, clusters 30–50 µm diam., individual photobiont cells 9–
13 µm broad and 5–6 µm long, green to yellow-orange in upper portions, penetrated by tubular fungal hyphae;
heterocytes sparse, hyaline to pale yellow, 8–10 µm wide and 4–5 µm long; cells of hyphal sheath wavy in
lateral outline, 3–4 µm thick; medulla 30–80 µm thick, composed of loosely woven, irregularly arranged to
more or less periclinal hyphae 4–5 µm thick; clamp connections not observed.
Hymenophore not observed.
Chemistry: no substances detected by TLC.
Distribution and Ecology:—This species is known from several collections growing on soil between
bryophytes in the Ecuadorian and Bolivian high Andes.
Etymology:—The epithet refers to the thallus appearing squamulose rather than foliose.
Remarks:—Cora squamiformis is phylogenetically closely related to C. pavonia (Dal-Forno et al. 2013)
and was found with the latter in the same habitat in Ecuador. The two species differ markedly in thallus and
lobe size and lobe configuration, with C. pavonia having much larger thalli and lobes not growing close to the
ground and lacking sinuouse margins, but having a coarsely undulate surface instead. A particular feature of
C. squamiformis appears to be the compacted instead of 'medullary' upper cortex, which at first glance is
similar to the cortex of Corella species, but in the latter the cortex is distinctly paraplectenchymatous and
lacks free hyphae. Cora bovei from southern Argentina (Spegazzini 1888) is similar to C. squamiformis in
general appearance and cortex structure but, as far as can be judged from the depauperate type material, forms
larger lobes with concentrically undulate surface.
Additional specimens examined:—ECUADOR. Napo: Papallacta; 3300 m; disturbed wet paramo
vegetation; 4 July 2010, Lücking 32300 (F). BOLIVIA. La Paz: Franz Tamayo, Madidi National Park,
Sanchez Pass between Pelechuco and Keara; 14° 43' S, 69° 08' W, 4602 m; high mountain vegetation (puna),
on ground between mosses; 13 October 2007, Wilk 7587 (F, KRAM). Eliodoro Camacho, Pumasane,
crossroads to Pelechuco and Charazani; 15° 15' S, 69° 03' W, 4536 m; high mountain vegetation (puna), on
ground between mosses; 12 October 2007, Wilk 7446 (F, KRAM).
Cora strigosa Lücking, E. Paz & L. Salcedo, sp. nov. (Fig. 9)
Mycobank #805383
Genbank ITS barcoding sequence: KF443241
Differing from the morphologically similar Cora hirsuta and the closely related C. byssoidea in the strigose tomentum
developed mostly submarginally.
Holotype:—PERU. Cuzco: Piscacucho; 13° 10' S, 72° 21' W, 2700–3800 m; disturbed montane rainforest
and pasture; 4 August 2009, Paz & Salcedo 3 (F).
Thallus on rocks associated with other lichens (Hypotrachyna and Rimelia), foliose, up to 10 cm across,
composed of 1–3 semicircular lobes per thallus; lobes 1–3 cm wide and 1–2 cm long, unbranched, greenish
grey when fresh, with thin, involute, grey margins, white-grey in the herbarium. Upper surface densely
hirsute-strigose or sometimes glabrous towards the base, with the trichomes arranged in broad, concentric
zones; trichomes free, more or less projecting radially towards the margin (as if combed), 1–1.5 mm long and
25–50 µm thick at the base, composed of agglutinated hyphae; involute margin with underside very minutely
arachnoid; lower surface ecorticate, finely felty-arachnoid (representing the exposed medulla), white-grey.
Thallus in section 250–350 µm thick, with upper cortex, photobiont layer, and medulla; upper cortex formed
by a 25–50 µm thick layer of rather loosely packed to indistinctly periclinal, 4–5 µm thick hyphae supported
by a 25–50 µm high 'medullary' layer of spaced groups of densely packed, anticlinal, 3–5 µm thick hyphae;
photobiont layer 50–150 µm thick, irregular, composed of clusters of short, coiled cyanobacterial filaments
wrapped in a dense, paraplectenchymatous hyphal sheath formed by jigsaw puzzle-shaped cells, clusters 20–

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DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
30 µm diam., individual photobiont cells 10–13 µm broad and 5–7 µm long, dark blue-green to orange-yellow
in upper portions, penetrated by tubular fungal hyphae; heterocytes sparse, hyaline to pale yellow, 8–10 µm
wide and 5–6 µm long; cells of hyphal sheath wavy in lateral outline, 3–4 µm thick; medulla 50–80 µm thick,
composed of loosely woven, irregularly arranged to more or less periclinal hyphae 4–5 µm thick; clamp
connections not observed.
FIGURE 9. Cora strigosa. A. Aspect of typical habitat in the Peruvian Andes near Machu Picchu. B. Lobe enlarged showing strigose
upper surface (holotype). C. Lobe underside showing hymenophore with finely arachnoid surface (holotype). Scale in B = 5 mm, in C
= 1 mm.

LÜCKING ET AL.20 • Phytotaxa 139 (1) © 2013 Magnolia Press
Hymenophore developed as irregular to elongate, resupinate patches arranged in reticulate pattern or more
or less concentric zones on the underside, patches 1–3 mm long and 0.5–1 mm broad, with white to pale
yellowish, finely arachnoid surface and slightly involute, finely byssoid margins; hymenophore in section 50–
100 µm thick, composed of a paraplectenchymatous layer resting on loose, 4–6 µm thick, generative
medullary hyphae and supporting the hymenium; hymenium composed of numerous, palisade-like basidioles
and scattered basidia, as well as numerous projecting hairs formed by single, cylindrical hyphae 20–50 µm
long and 4–5 µm thick; basidioles 25–35 × 5–6 µm; basidia 30–40 × 5–7 µm, 4-sterigmate; basidiospores not
observed.
Chemistry: no substances detected by TLC.
Distribution and Ecology:—This species is known from a single locality in a heavily disturbed montane
rain forest in Peru, near Machu Picchu.
Etymology:—The epithet refers to the radially projecting, conspicuous trichomes.
Remarks:—This species at first glance resembles Cora hirsuta (Lumbsch et al. 2011) in forming rather
large, free trichomes, but is not closely related to the latter, as it falls into another clade (Dal-Forno et al.
2013). Morphologically it can be distinguished by the trichomes developing up to and especially near the
margin, whereas C. hirsuta features a thin, glabrous, submarginal zone of different color (Lumbsch et al.
2011). Also, the finely arachnoid surface of the hymenophore, caused by numerous hyphae projecting from
the hymenium surface, is unique within the genus. More closely related is C. byssoidea (see above), which
differs in the more irregular, interwoven tomentum and the glabrous hymenophore surface.
Additional specimens examined:—PERU. Cuzco: Piscacucho; 13° 10' S, 72° 21' W, 2700–3800 m;
disturbed montane rainforest and pasture; 4 August 2009, Díaz & Jihuallanco s.n. (F).
Dictyonema aeruginosulum Lücking, Nelsen & Will-Wolf, sp. nov. (Fig. 10)
Mycobank #805384
Genbank ITS barcoding sequence: EU825955
Differing from the morphologically similar Dictyonema phyllophilum and D. schenckianum in the abundant, coarse,
irregular, finger-like projections formed by the vegetative thallus, and from D. irpicinum in the appressed-
filamentous growth habit and the lack of clamp connections.
Holotype:—COSTA RICA. Alajuela: Volcán Tenorio National Park, Pilón Biological Station, Arenal-
Tempisque Conservation Area, Tilarán Ridge, 140 km NW of San José, 25 km NNW of Tilarán, near Bijagua,
access road to station and river; 84° 59' W, 10° 43' N, 700 m; lower montane cloud forest zone, exposed trees
and fence posts along pasture, on bark (lower stem), exposed; 16 March 2004, Nelsen 3754 (INB; isotypes: F,
WIS).
Thallus epiphytic on tree trunks, appressed filamentous, covering large areas of the substrate, forming a
compressed mat of irregularly arranged to more or less horizontal, densely interwoven, dark aeruginous fibrils
resting on a white, byssoid hypothallus; thallus densely furnished with irregular, finger-like projections
laterally covered with fibrils, the projections appearing stiff but softening when moistened, becoming
branched and confluent, up to 10 mm high and 2 mm broad. Thallus in section 300–800 µm thick (excluding
the projections), composed of an upper photobiont layer 200–400 µm thick and a lower medulla (forming the
hypothallus) 100–400 µm thick; photobiont layer composed of numerous cyanobacterial filaments wrapped in
a closed hyphal sheath formed by jigsaw puzzle-shaped cells, connected to loose hyphae towards the medulla;
medulla composed of a loose network of interwoven hyphae sparsely intermingled with cyanobacterial
filaments; cyanobacterial filaments composed of 8–12 µm wide and 4–5 µm high, blue-green cells penetrated
by tubular fungal hyphae; heterocytes sparse, pale yellow, 7–11 µm wide and 3–4 µm high; cells of hyphal
sheath wavy in lateral outline, 3–4 µm thick; medullary hyphae and those associated with hyphal sheath
straight, 4–6 µm thick, lacking clamp connections. Projections in section formed by a network of medullary

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DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
hyphae 4–6 µm thick, lacking clamp connections, loosely intermingled with cyanobacterial filaments
internally and with a denser layer of filaments formed on the outside except the apical regions.
FIGURE 10. Dictyonema aeruginosulum (holotype). A. Thallus with projections. B. Thallus surface enlarged showing filaments. C.
Hymenophore. Scale = 1 mm.
Hymenophore developed as irregular, resupinate patches on the thallus surface or on the underside of the
projections and then soon becoming inverted and exposed, with pale yellow, smooth surface; hymenophore in
section 50–100 µm thick, composed of a paraplectenchymatous layer resting on loose medullary hyphae and

LÜCKING ET AL.22 • Phytotaxa 139 (1) © 2013 Magnolia Press
supporting the hymenium; hymenium composed of numerous, palisade-like basidioles and scattered basidia;
basidioles 10–20 × 5–7 µm; basidia 15–25 × 5–8 µm, 4-sterigmate; basidiospores (few seen) ellipsoid to
narrowly drop-shaped, non-septate, hyaline, 7–9 × 3–4 µm.
Chemistry: no substances detected by TLC.
Distribution and Ecology:—This species is thus far known from montane rain forest in the northern
Cordillera de Tilarán in Costa Rica, forming extensive mats on the trunks of semi-exposed trees of Syzygium
jambos in a pasture along the road in an area with abundant precipitation. Unfortunately, a few years after
collecting the material, the trees in this spot were completely logged, so the holotype population is likely
extirpated.
Etymology:—The epithet refers to the characteristic blue-green color of this species, while most other
species are either more bluish or greenish.
Remarks:—Dictyonema aeruginosulum is one of several species now segregated from D. sericeum,
disentangling the broad concept of that species laid out by Parmasto (1978). While that author focused on
mycological features of the basidiomata and regarded variation in thallus morphology as of no taxonomic
value, molecular phylogenetic data clearly show that D. sericeum sensu Parmasto contains a large number of
different species and even the shelf-like forms representing D. sericeum in a narrow sense are more than one
species (Dal-Forno et al. 2013). Due to the distinct white hypothallus formed by a well-developed, laterally
projecting medullary layer, D. aeruginosulum is most similar to D. phyllophilum (Parmasto) Lücking, Dal-
Forno & Lawrey, comb. et stat. nov. [Mycobank #805390; bas.: D. sericeum f. phyllophilum Parmasto, Nova
Hedwigia 29: 113 (1978); holotype: Malaysia (Borneo: Sarawak), Beccari 222 (B; isotype: W!)]. It differs
from the latter chiefly in the conspicuous finger-like projections. Also, all known collections of D.
phyllophilum are sterile. Phylogenetically, the two species do not appear to be closely related. Finger-like
projections, though smaller, are also known from D. scabridum (Vain.) Lücking, comb. et stat. nov.
[Mycobank #805391; bas.: Rhipidonema irpicinum f. scabridum Vain., Ann. Acad. Sci. Fenn., Ser. A, 19(15):
29 (1923); syn.: Dictyonema ligulatum f. scabridum (Vain.) Parmasto, Nova Hedwigia 29: 120 (1978);
lectotype (Parmasto 1978: 120): Philippines, Weber 1391 (TUR-Vainio 32883!; isotype: W!)] and from D.
irpicinum Mont. (Montagne 1848: 119; holotype in PC checked), which both differ in the shelf-like growth
and the presence of clamp connections, and D. scabridum also in the densely arranged fibrils forming an
almost compact surface (Parmasto 1978).
Additional specimens examined:—COSTA RICA. Alajuela: Volcán Tenorio National Park, Pilón
Biological Station, Arenal-Tempisque Conservation Area, Tilarán Ridge, 140 km NW of San José, 25 km
NNW of Tilarán, near Bijagua, access road to station and river; 84° 59' W, 10° 43' N, 700 m; lower montane
cloud forest zone, exposed trees and fence posts along pasture, on bark (lower stem), exposed; 15 March
2004, Will-Wolf 12733 (F, INB, USJ, WIS).
Dictyonema metallicum Lücking, Dal-Forno & Lawrey, sp. nov. (Fig. 11)
Mycobank #805385
Genbank ITS barcoding sequence: KF443222
Differing from the morphologically similar and related Dictyonema hernandezii in the thin, completely appressed thallus
and the dark blue color, with a metallic shimmer when dry.
Holotype:—ECUADOR. Pichincha: Río Guajalito Protected Forest; 0°09’S, 78°39’W, 1800 m; montane
rainforest, on tree trunk; September 2008, Lücking 26255 (QCNE; isotype: F).
Thallus epiphytic on tree trunks and overgrowing nearby bryophytes, appressed filamentous, in irregular,
dispersed to confluent patches, each 1–5 cm across and entire thallus eventually covering larger areas of the
substrate, forming a strongly compressed mat of horizontal, loosely interwoven, dark blue fibrils completely
embedded in a gelatinous, silvery prothallus with strongly metallic shimmer. Thallus in section 25–50 µm
thick, composed of an irregular photobiont layer but lacking a discernible medulla; photobiont layer

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DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
composed of numerous cyanobacterial filaments wrapped in a closed hyphal sheath formed by jigsaw puzzle-
shaped cells; cyanobacterial filaments composed of 10–13 µm wide and 4–6 µm high, dark aeruginous blue
cells penetrated by tubular fungal hyphae; heterocytes sparse, hyaline, 8–12 µm wide and 4–6 µm high; cells
of hyphal sheath wavy in lateral outline, 3–4 µm thick; hyphae associated with hyphal sheath straight, 4–6 µm
thick, lacking clamp connections; compacted prothallus mostly found by densely arranged empty hyphal
sheaths admixed with straight hyphae.
FIGURE 11. Dictyonema metallicum (Ecuador, Lücking 26203). A–B. Specimen in the field. C. Thallus surface enlarged showing
filaments. Scale = 1 mm.

LÜCKING ET AL.24 • Phytotaxa 139 (1) © 2013 Magnolia Press
Hymenophore not observed.
Chemistry: no substances detected by TLC.
Distribution and Ecology:—This species is thus far known from montane rain forest in Ecuador,
forming dispersed mats on the trunks of shaded trees in the rain forest understory, also overgrowing nearby
epiphytic bryophytes.
Etymology:—The epithet refers to the metallic shimmer of the thallus when dry.
Remarks:—Dictyonema metallicum is similar to the recently described D. hernandezii Lücking, Lawrey
& Dal-Forno (Lumbsch et al. 2011: 46) in having the fibrils embedded in a gelatinous matrix formed by the
fungal prothallus and hypothallus. The latter differs in the much thicker thallus which forms a thick, bulging,
zonate marginal prothallus, and the more greenish color of the cyanobacterial photobiont. The tiny fibrils of
D. metallicum are reminiscent of those of Cyphellostereum phyllogenum (Müll. Arg.) Lücking, Dal-Forno
& Lawrey, comb. nov. [Mycobank #805396; bas.: Dichonema phyllogenum Müll. Arg., Flora 66: 352 (1883);
syn.: Dictyonema phyllogenum (Müll. Arg.) Zahlbr., Cat. Lich. Univ. 7: 746 (1931); lectotype (Parmasto
1978: 124): Malaysia (Borneo: Sarawak), Beccari 1624 (G!)] and C. nitidum (Lücking) Lücking (Lücking
2008; Yánez et al. 2012), but those two species have a Cyphellostereum-type thallus lacking a distinct hyphal
sheath and also lacking haustoria.
Additional specimens examined:—ECUADOR. Pichincha: Río Guajalito Protected Forest; 0°09’S,
78°39’W, 1800 m; montane rainforest, on tree trunk; September 2008, Lücking 26255 (F).
Dictyonema obscuratum Lücking, Spielmann & Marcelli, sp. nov. (Fig. 12)
Mycobank #805386
Genbank ITS barcoding sequence: KF443223
Differing from Dictyonema phyllophilum and D. schenckianum s.lat. in the densely and irregularly interwoven, dark
olive-green fibrils and the absence of a distinct hypothallus.
Holotype:—BRAZIL. São Paulo: Mogi-Graçu, Martinho Prado Jr., Mogi-Guaçu Ecological Reserve,
Fazenda Campininha; 22° 15' S, 47° 10' W, 635 m; interior of dense Cerrado; 7 November 2007, Lücking
23025 (F; isotype: SP).
Thallus epiphytic on tree trunks, appressed filamentous, individual patches up to 5 cm across but
eventually covering large areas of the substrate, forming a strongly compressed mat of irregularly arranged,
densely interwoven, very dark olive-green fibrils resting on a very thin, often indistinct, sordid pale brown,
byssoid hypothallus. Thallus in section 200–400 µm thick, composed of an upper photobiont layer 150–250
µm thick and a lower medulla (forming the hypothallus) 50–100 µm thick; photobiont layer composed of
numerous cyanobacterial filaments wrapped in a closed hyphal sheath formed by jigsaw puzzle-shaped cells,
connected to loose hyphae towards the medulla; medulla composed of a loose network of interwoven hyphae
sparsely intermingled with cyanobacterial filaments; cyanobacterial filaments composed of 20–25 µm wide
and 6–8 µm high, dark green cells (becoming orange-yellow towards the tips) penetrated by tubular fungal
hyphae, often longitudinally divided, heterocytes sparse, pale yellow, 15–20 µm wide and 6–9 µm high; cells
of hyphal sheath wavy in lateral outline, 3–4 µm thick; medullary hyphae and those associated with hyphal
sheath straight, 4–6 µm thick, lacking clamp connections but often sparsely and finely papillose.
Hymenophore developed as bulging, stereoid patches from the underside of the thallus margins, white;
hymenophore in section 200–400 µm thick, composed of a paraplectenchymatous layer connected to loose
medullary hyphae; hymenium composed of numerous, palisade-like basidioles and scattered basidia;
basidioles 20–30 × 5–7 µm; basidia 30–40 × 5–8 µm, 4-sterigmate; basidiospores ellipsoid to narrowly drop-
shaped, non-septate, hyaline, 7–9 × 3–4 µm.
Chemistry: no substances detected by TLC.
Distribution and Ecology:—This species is thus far known from Cerrado ('Cerrado denso') vegetation in
the state of São Paulo, Brazil, where it grows on the corky bark of characteristic Cerrado trees.

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DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
Etymology:—The epithet refers to the very dark color of the thallus, at first glance not at all resembling a
lichen.
FIGURE 12. Dictyonema obscuratum. A. Specimen in the field (photograph A. Spielmann). B. Thallus surface enlarged showing
filaments. C. Filaments with heterocytes in microscopic view. Scale in B = 1 mm, in C = 20 µm.
Remarks:—This is another new species in the complex formerly recognized as just a single species,
Dictyonema sericeum (Parmasto 1978). It differs from superficially similar species such as D. phyllophilum

LÜCKING ET AL.26 • Phytotaxa 139 (1) © 2013 Magnolia Press
and D. schenckianum (Müll. Arg.) Zahlbr. (Zahlbruckner 1931: 748) in the very dark color of the thallus and
the very broad, dark green rather than bluish green, irregularly arranged fibrils in which the photobiont cells
of the cyanobacterial filaments tend to divide longitudinally, giving them partially a 'muriform' appearance.
This feature is reminiscent of D. moorei (Nyl.) Henssen (Henssen 1963: 109; Parmasto 1978), in which the
hyphal sheath usually contains two filaments, but in D. obscuratum no distinct separate filaments are formed
within a single sheath. Also, the surface of the filaments in D. moorei is different and more similar to the
genus Acantholichen.
Additional specimens examined:—BRAZIL. São Paulo: Mogi-Graçu, Mogi-Guaçu Biological
Reserve, Fazenda Campininha, Cerrado Seco; 22° 15' S, 47° 10' W, 650 m; interior of dense Cerrado; 7
November 2007, Lücking 23025, 23204 (F, SP).
Key to currently accepted genera of Dictyonema s.lat.
1. Thallus composed of distinct fibrils including cyanobacterial filaments, either appressed to substrate or forming hor-
izontally projecting, semicircular lobes ........................................................................................................................2
- Thallus microsquamulose to foliose, no distinct fibrils visible, photobiont instead forming clusters of short, irregu-
larly coiled threads inside the thallus ............................................................................................................................3
2. Photobiont cells narrow (5–7 µm broad), lacking haustoria; hyphal sheath around photobiont filaments composed of
irregular hyphae leaving interspaces; basidiomata (hymenophores) if present stipitate and erect, only at the base con-
nected to lichenized thallus ............................................................................................. Cyphellostereum D. A. Reid
- Photobiont cells broad (7–20 µm broad), with tubular intracellular haustoria; hyphal sheath around photobiont fila-
ments composed of paraplectenchymatous, jigsaw-puzzle-shaped cells forming a completely closed layer; basidi-
omata (hymenophores) if present stereoid-corticioid, without stipe, their dorsal portion partially overgrown with the
lichenized thallus or completely formed on the thallus underside ........................ Dictyonema C. Agardh ex Kunth
3. Thallus microsquamulose; thallus underside in microscope view forming apically thickened, distinctly spinulose
hyphae (acanthohyphae) .................................................................................................... Acantholichen P. M. Jørg.
- Thallus macrosquamulose; acanthohyphae absent .......................................................................................................4
4. Upper cortex thin, distinctly paraplectenchymatous; upper surface color dark blue-green or olive-brown when dry;
isidioid propagules sometimes present; hymenophores unknown........................................................... Corella Vain .
- Upper cortex thick, composed of an upper, periclinal layer of loosely packed hyphae supported by a layer of anticli-
nal hyphal bundles leaving large interspaces; soredioid propagules sometimes present; hymenophores mostly pres-
ent..................................................................................................................................................................... Cora Fr.
Key to currently recognized species of Cyphellostereum (excluding species not belonging in this clade)
1. Thallus with distinct white prothallus........................................................................................................................... 2
- Thallus lacking distinct prothallus................................................................................................................................ 3
2. Fibrils irregularly appressed; hyphal sheath around cyanobacterial filaments dense.....................................................
........................................................................ C. imperfectum Lücking, Barillas & Dal-Forno (Yánez et al. 2012)
[Illustration in Yánez et al. 2012: 227, fig. 1d–f]
- Fibrils strongly appressed; hyphal sheath around cyanobacterial filaments loose ..... C. nitidum (Lücking) Lücking
[Illustration in Lücking 2008: 784, fig. 257D]
3. Thallus terrestrial; basidiomata common............ C. pusiolum (Berk. & M. A. Curtis) D. A. Reid (Reid 1965: 342)
[Syn.: Stereum cyphelloides Berk. & M. A. Curtis (Berkeley & Curtis 1868: 331); Stereophyllum pallens P. Karst.
(Karsten 1889: 223); Thelephora uleana Henn. (Hennings 1897: 194); Podoscypha minutula Pat. (Patouillard 1924:
33); illustration in Dal-Forno et al. 2013, fig. 3A; synonymy is based on current species concept but possibly some
of the synonyms represent distinct species]
- Thallus epiphytic; basidiomata unknown ..................C. phyllogenum (Müll. Arg.) Lücking, Dal-Forno & Lawrey
[Illustration in Lücking 2008: 784, fig. 257C]

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DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
Key to currently recognized species of Dictyonema s.str.
1. Thallus appearing applanate microfruticulose, each branch including 2–3 cyanobacterial filaments ...........................
............................................................................................................................................. D. moorei (Nyl.) Henssen
[Syn.: Dictyonema japonicum Asahina (Asahina 1944); Dictyonema confusum Henssen in herb. (nom. inval.); illus-
tration in Henssen 1963: taf. 28d, 30c–e].
- Thallus distinctly filamentous, cyanobacterial filaments always solitary ....................................................................2
2. Thallus forming semicircular lobes projecting horizontally from the substrate ...........................................................3
- Thallus appressed-filamentous, forming a crust over the substrate..............................................................................6
3. Thallus surface with coarse, finger-like outgrowths; clamp connections present ........................................................ 4
- Thallus surface plane; clamp connections present or absent ........................................................................................ 5
4. Fibrils narrow, very densely arranged, giving the lobes an almost smooth appearance; lobe surface intensely blue-
green ............................................................................................................................ D. scabridum (Vain.) Lücking
- Fibrils broad, more loosely and irregularly arranged and leaving interspaces, giving the lobes a rough appearance;
lobe surface mottled white and blue-green ...................................................................D. irpicinum Mont. (Fig. 13A)
5. Fibrils narrow, very densely arranged, giving the lobes an almost smooth appearance; lobe surface intensely blue-
green; clamp connections present .................... D. ligulatum (Kremp.) Zahlbr. (Zahlbruckner 1908: 239; Fig. 13B)
[Dictyonema laxum Müll. Arg., Bot. Jahrb. 4: 57 (1883)].
- Fibrils broad, more loosely and irregularly arranged and leaving interspaces, giving the lobes a rough appearance;
lobe surface mottled white and blue-green; clamp connections absent ........................ D. sericeum (Sw.) Berk. s.lat.
[This is a collective taxon comprising several distinct lineages, but more data are required to establish exact species
boundaries; possibly distinct species are Dictyonema sericeum s.str., described from the Caribbean; D. aeruginosum
(Blume & T. Nees) Berk. (Berkeley 1872), described from Indonesia (Java); D. excentricum C. Agardh (Kunth
1822: 1), with thick, horizontally arranged bundles of fibrils (Fig. 13C), described from French Guiana; and D. spon-
giosum Berk. & M. A. Curtis (Berkeley & Curtis 1868: 335), with a thick, spongiose upper surface composed of
bundles of vertically projecting fibrils (Fig. 13D), described from Cuba. The type material of D. sericeum is rather
small and consists of three lobes with more or less appressed, aeruginous fibrils and a whitish to cream-colored,
marginal zone lacking photobiont filaments; it appears most similar to the lineage labeled D. sericeum 1 in Dal-
Forno et al. (2013); the type of D. aeruginosum is extremely small and cannot be identified with certainty and must
be considered a nomen dubium; D. excentricum has not been recollected by us; and the fourth taxon, D. spongiosum,
was gathered in Guatemala (Fig. 13E–F) and was sequenced and was found to represent the lineage labeled D. seri-
ceum 3 in Dal-Forno et al. (2013)]
6. Fibrils distinctly 'combed' (oriented in a single direction) or embedded in a gelatinous matrix, forming a regular or
smooth surface with the fibrils horizontally arranged and closely appressed............................................................... 7
- Fibrils neither combed nor embedded in a gelatinous matrix, forming a more or less irregular, rough surface, with
the fibrils irregularly arranged to ascending or erect .................................................................................................. 11
7. Fibrils distinctly 'combed' .............................................................................................................................................8
- Fibrils embedded in a gelatinous matrix or closely appressed to substrate .................................................................. 9
8. Fibrils olive-green; prothallus indistinct; hyphal sheath papillose towards the tips of the filaments.............................
..................................................................... D. pectinatum Dal Forno, Yánez & Lücking (Yánez et al. 2012: 234)
[Illustration in Yánez et al. 2012: 235, fig. 3d–f]
- Fibrils distinctly blue-green; prothallus distinct; hyphal sheath smooth ........................................................................
..................................................................................................... D. schenckianum (Müll. Arg.) Zahlbr. (Fig. 14A)
[This name was used in a broader sense by Chaves et al. (2004) and Yánez et al. (2012), including also specimens
with rather thick, appressed thalli with irregularly arranged fibrils, often being fertile; revision of type material
revealed that these specimens come closer to D. irrigatum (differing by the lack of clamp connections), whereas the
fibrils of D. schenckianum s.str. have a combed appearance]
9. Fibrils closely appressed but not embedded in gelatinous matrix; over bryophyes .......................................................
....................................................................................................... D. diducens Nyl. ex Lücking, sp. nov. (Fig. 14B)
[Mycobank #805387. This taxon was not validly described by Nylander (1885) and a brief description follows: Dif-
fering from the morphologically similar Dictyonema thelephora in the closely appressed fibrils forming an almost
continuous crust. Holotype: Peru, unknown locality and date, Krause s.n. (BM-001084450!). Thallus epiphytic on bry-

LÜCKING ET AL.28 • Phytotaxa 139 (1) © 2013 Magnolia Press
ophytes, appressed filamentous and forming a more or less smooth crust of irregularly arranged to nearly parallel, aeruginous
fibrils, lacking a distinct hypothallus and prothallus. Thallus in section 20–50 µm thick, of numerous cyanobacterial filaments
wrapped in a closed hyphal sheath formed by jigsaw puzzle-shaped cells; cyanobacterial filaments composed of 8–14 µm wide
and 4–5 µm high, blue-green cells penetrated by tubular fungal hyphae; heterocytes sparse, pale yellow, 7–12 µm wide and 3–4
µm high; cells of hyphal sheath wavy in lateral outline, 3–4 µm thick; free hyphae associated with hyphal sheath straight, 4–6 µm
thick, lacking clamp connections.]
- Fibrils embedded in gelatinous matrix; on bark ......................................................................................................... 10
FIGURE 13. A. Dictyonema irpicinum (holotype: PC). B. D. ligulatum (Papua New Guinea, Sands 1918: BM). C. D. excentricum
(isotype: PC). D–F. D. spongiosum (D, syntype: PC; E–F, Guatemala, Lücking 25561: F). Scale in A = 5 mm, in B–D, F = 10 mm, in E
= 50 mm.

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DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
FIGURE 14. A. Dictyonema schenkianum (isotype of Laudatea schenkiana: S). B. D. diducens (holotype: BM). C. D. caespitosum
(holotype of Laudatea caespitosa: S). D. D. irrigatum (holotype of Corticium irrigatum: PC). E. D. phyllophilum (holotype of D.
sericeum f. phyllophilum: W). F. D. aff. irrigatum (Costa Rica, Lücking s.n.: F). Scale in A, C = 1 mm, in B, F = 10 mm, in D = 5 mm.
10. Thallus thick, with thick, gelatinous, zonate prothallus, opaque when dry; fibrils light aeruginous .............................
......................................................................................................... D. hernandezii Lücking, Lawrey & Dal-Forno
[Illustration in Lumbsch et al. 2011: 47, fig. 10C]
- Thallus thin, lacking distinct prothallus, with metallic shimmer when dry; fibrils dark greenish blue .........................
............................................................................................ D. metallicum Lücking, Dal-Forno & Lawrey (Fig. 11)

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11. Thallus with coarse, finger-like outgrowths .................. D. aeruginosulum Lücking, Nelsen & Will-Wolf (Fig. 10)
- Thallus plane............................................................................................................................................................... 12
12. Clamp connections present .........................................................................................................................................13
- Clamp connections absent ..........................................................................................................................................14
13. Thallus usually over bryophytes, thin with appressed, mostly horizontal fibrils, pale greenish blue, rarely fertile ......
.............................................................................................................. D. caespitosum (Johow) Lücking (Fig. 14C)
[Dictyonema caespitosum (Johow) Lücking, comb. nov.; Mycobank #805392; bas.: Laudatea caespitosa Johow,
Jahrb. Wiss. Bot. 15: 386 (1884); holotype: Brazil, Dusén s.n. (S!)]
- Thallus usually on tree trunks, thick with irregularly appressed to ascending or short-erect fibrils, dark blue-green,
often fertile with stereoid hymenophores......................... D. irrigatum (Berk. & M. A. Curtis) Lücking (Fig. 14D)
[Dictyonema irrigatum (Berk. & M. A. Curtis) Lücking, comb. nov.; Mycobank #805393; bas.: Corticium irriga-
tum Berk. & M. A. Curtis, Proc. Amer. Acad. Arts & Sci. 4: 123 (1860); holotype: China, Wright 108 (PC!)]
14. Thallus usually on living leaves; prothallus distinct, white ............................................................................................
................................................................. D. phyllophilum (Parmasto) Lücking, Dal-Forno & Lawrey (Fig. 14E)
- Thallus on tree trunks or bryophytes; prothallus indistinct or absent......................................................................... 15
15. Thallus on tree trunks; often fertile............................................................................................................................. 16
- Thallus on mosses or liverworts; usually sterile......................................................................................................... 17
16. Thallus dark blue-green to brownish; cells of the cyanobacterial filaments often longitudinally divided.....................
........................................................................................ D. obscuratum Lücking, Spielmann & Marcelli (Fig. 12)
- Thallus light to dark blue-green; cells of the cyanobacterial filaments not divided.......................................................
................................................................................... D. aff. irrigatum (Berk. & M. A. Curtis) Lücking (Fig. 14F)
17. Thallus dark blue-green, shiny; western Europe ..... D. coppinsii Lücking, Barrie & Genney (Lücking et al. 2014)
[Dictyonema interruptum auct., non (Carmich. ex Hook.) Parmasto (= Rhizonema interruptum Lücking & Barrie);
illustration in Lücking et al. 2014, fig. 1]
- Thallus light greyish blue-green; tropics ....................................................................................................................18
18. Fibrils irregularly appressed ........................................... D. thelephora (Spreng.) Zahlbr. (Zahlbruckner 1931: 748)
- Fibrils irregularly erect .......................... D. galapagoense Yánez, Dal Forno & Bungartz (Yánez et al. 2012: 234)
[Illustration in Yánez et al. 2012: 235, fig. 3a–c]
Key to currently recognized species of Cora
1. Upper or lower lobe surface with distinct tomentum or tufts of hairs .......................................................................... 2
- Upper lobe surface glabrous, lower surface glabrous or minutely arachnoid ..............................................................7
2. Lobes with irregular, dark, sorediate margins contrasting with the white lobe surface; upper lobe surface glabrous;
lower lobe surface with long tufts of hairs................................................... C. inversa Lücking & Moncada (Fig. 7)
- Lobes with rounded, non-sorediate margins; upper lobe surface at least partially arachnoid-tomentose; lower lobe
surface minutely arachnoid ...........................................................................................................................................3
3. Upper lobe surface with rather long, erect to horizontally combed setae of agglutinated hairs................................... 4
- Upper lobe surface with short, arachnoid tomentum or concentric zones of hairs formed by simple hyphae............. 5
4. Upper surface tomentose up to the margin and setae longest along the margin; surface of hymenophore minutely
arachnoid ..................................................................................... C. strigosa Lücking, E. Paz & L. Salcedo (Fig. 9)
- Upper surface tomentose with a narrow, glabrous submarginal zone; surface of hymenophore glabrous.....................
........................................................................................... C. hirsuta (Moncada & Lücking) Moncada & Lücking
[Illustration in Lumbsch et al. 2011: 47, fig. 10D]
5. Upper surface glabrous except for concentrical, sometimes inconspicuous zones of short hairs ..................................
......................................................................................................... C. aspera Wilk, Lücking & E. Morales (Fig. 4)
- Upper surface arachnoid-tomentose throughout ...........................................................................................................6

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DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
6. Lobes up to 5 cm broad, brown when fresh; thallus mostly terrestrial...........................................................................
.......................................................................................................... C. arachnoidea J. E. Hern. & Lücking (Fig. 3)
- Lobes up to 2 cm broad, white when fresh; thallus usually epiphytic..... C. byssoidea Lücking & Moncada (Fig. 5)
7. Lobes up to 2 cm broad; upper cortex compacted, lacking distinct supporting medullary layer .................................8
- Lobes up to 7 cm broad; upper cortex with distinct supporting medullary layer formed by bundles of anticlinal
hyphae separated by large interspaces ........................................................................................................................10
8. Lobes white with dark, granular margins; thallus epiphytic ... C. minor (Lücking, E. Navarro & Sipman) Lücking
[Illustration in Chaves et al. 2004: 245, fig. 1B–D]
- Lobes dark with paler, minutely arachnoid margins; thallus terrestrial between bryophytes....................................... 9
9. Lobes up to 1 cm broad, with plane surface ................ C. squamiformis Wilk, Lücking & Yánez-Ayabaca (Fig. 8)
- Lobes up to 2 cm broad, with concentrically undulate surface ......................................... C. bovei Speg. (Fig. 2C–D)
10. Thallus usually epiphytic, grey to blue-grey or aeruginous when fresh..................................................................... 11
- Thallus usually terrestrial, more or less grey-brown when fresh................................................................................ 13
11. Thallus light aeruginous when fresh; lobe surface pitted; hymenophore stereoid-cyphelloid .......................................
......................................................................................... C. cyphellifera Dal-Forno, Bungartz & Lücking (Fig. 6)
- Thallus grey to blue-grey when fresh; lobe surface not pitted; hymenophore corticioid...........................................12
12. Lobe surface concentrically undulate; tropical Africa (Mauritius) ................................................... C. gyrolophia Fr.
[Syn.: Gyrolophium elegans Kunze (G. 'mauritianum' Kunze)].
- Lobe surface more or less plane; tropical America ......................... C. aspera Wilk, Lücking & E. Morales (Fig. 4)
13. Lobes up to 7 cm broad; lobe surface strongly concentrically undulate ......................... C. pavonia (Sw.) Fr. (Fig. 1)
Syn.: Cora pavonia (Weber & D. Mohr) Fr. [nom. illeg.]; Wainiocora ciferrii Tom a s.
- Lobes up to 3 cm broad; lobe surface plane to shallowly concentrically undulate .................................................... 14
14. Lobe surface shallowly concentrically undulate; hymenophore finely reticulate, very regularly arranged, with even
or slightly downturned margins even when dry ........................................................ C. reticulifera Vain. (Fig. 2E–F)
- Lobe surface plane; hymenophore irregular, with slightly upturned margins, especially when dry ..............................
......................................................................................................................... C. glabrata (Spreng.) Fr. (Fig. 2A–B)
Key to currently recognized species of Corella
1. Thallus forming irregular, isidioid to finger-like outgrowths .........................................................................................
.............................................................. C. melvinii (Chaves, Lücking & Umaña) Lücking, Dal-Forno & Lawrey
[Corella melvinii (Chaves, Lücking & Umaña) Lücking, Dal-Forno & Lawrey, comb. nov.; Mycobank #805394;
bas.: Dictyonema melvinii Chaves, Lücking & Umaña in Chaves et al., Bryologist 107: 244 (2004); holotype: Costa
Rica, Chaves 122 (INB-3762769); illustration in Chaves et al. 2004: 245, fig. 1E–F]
- Thallus lacking isidioid outgrowths but sometimes forming irregular lobules ............................ C. brasiliensis Va in .
[Syn.: Corella tomentosa Vain. (Vainio 1899); Corella zahlbruckneri Schiffn. (Zahlbruckner 1909); illustration in
Dal-Forno et al. 2013, fig. 3O–P]
The following names have not yet been checked since type material was not located, and hence their
taxonomic status remains uncertain:
Dichonema aeruginosum Blume & T. Nees, Nova Acta Acad. Caes. Leop.-Carol. 13: 12 (1826); ≡ Cora
neesiana Lév., Ann. Sci. Nat., Bot., Sér. 3, 5: 154 (1846) [nom. illeg.]; Indonesia (Java).
Dictyonema expansum Pouls., Vidensk. Medd. Naturhist. Foren København 1899: 280 (1899); Indonesia
(Java).
Dictyonema membranaceum C. Agardh, Syst. Alg.: 85 (1824); Mariana Islands.

LÜCKING ET AL.32 • Phytotaxa 139 (1) © 2013 Magnolia Press
Dictyonema membranaceum var. guadalupense Rabenh., Hedwigia 13: 7 (1874); Guadeloupe.
Dictyonema sericeum f. membranaceum P. Metzner, Ber. Deutsch. Bot. Ges. 52: 238 (1934); Indonesia (Java).
Rhipidonema crustaceum P. Metzner, Ber. Deutsch. Bot. Ges. 52: 232 (1934); Indonesia (Java).
Rhipidonema puiggarii Speg., Boln Soc. Cienc. Córdoba 23(3-4): 70 [reprint] (1919); Brazil.
The name Dictyonema sericeum f. laminosum Har., Bull. Soc. Mycol. Fr. 7: 41 (1891), listed in Index
Fungorum, is a lapsus. Hariot (1891) did not describe a taxon with that name, but instead divided Dictyonema
into two groups corresponding to series, Sericea (species with shelf-like thallus) and Laminosa (species with
appressed thallus).
The name Thelephora textilis Spreng. suggests another representative of Dictyonema. This name is cited
in Fries (1825) as type of the new genus Cilicia Fr.; however, it appears that Sprengel never validly described
a species under that name. Fries (1825) gave as a typical species of his new genus Auricularia reflexa Bull.,
which is considered a synonym of Stereum hirsutum (Willd.) Pers. (Smith et al. 1824; Streinz 1862; Saccardo
1888a). The latter is superficially similar to Cora, but is a completely unrelated, non-lichenized fungus. In the
absence of a valid description of Thelephora textilis, the genus name Cilicia Fr. should be considered a
synonym of Stereum Hill ex Pers., but certainly not a synonym of Chrysothrix Mont., as suggested by
Zahlbruckner (1923). The name Cilicia aeruginosa Fr. is mentioned in the literature (e.g. Parmasto 1978) as
described in the protologue of Cilicia Fr. (Fries 1825: 301), but no such name was described by Fries (1825)
in that work.
Conclusions
The results of our study reinforce the idea that Dictyonema s.lat., previously considered to represent only a
few species in a single genus, actually comprises an unexpectedly high diversity of species in several, distinct
genera, differing in morphology, anatomy, substrate ecology, and distribution. This applies even considering
that Parmasto (1978) did not take into account Cyphellostereum pusiolum, which was first recognized as
lichenized by Aptroot & Sipman (1991), nor Acantholichen pannarioides, which was not yet described at the
time. Of the 40 species distinguished here, 38 would be included in Parmasto's concept of Dictyonema s.lat.,
although he questioned the placement of D. phyllogenum (now in Cyphellostereum) in the genus. Based on
our limited sampling focusing on the wet northern Andes, we suspect that many more species will eventually
be discovered, in addition to the four species of Cyphellostereum, at least 20 species of Dictyonema s.str., one
species of Acantholichen, two species of Corella, and 14 species of Cora. Among the material collected by us
that remains to be sequenced and characterized morphologically and anatomically, we already anticipate at
least three more species each of Cyphellostereum and Dictyonema, one each of Acantholichen and Corella,
and at least two of the genus Cora. Together with the remaining type material requiring revision, in particular
of names described from the Paleotropics, this is a dramatic, more than ten-fold increase compared to the five
"lichen" and two "fungal" species recognized by Parmasto (1978) in this group. One possible reason why
species of this group have not been properly recognized before, particularly in the genus Cora, is the
observation that, similar to macrolichens in the order Peltigerales, such as Leptogium, Peltigera and Sticta and
relatives, the correct identification of species requires field experience and preferably images of specimens
taken in situ before being collected, since some of the diagnostic characters, such as color and shape of fresh
hymenophores, cannot be readily observed in herbarium material.
Acknowledgements
This study was supported by three grants from the National Science Foundation: TIC O LICHE N (DEB
0206125 to The Field Museum; PI Robert Lücking), Neotropical Epiphytic Microlichens – An Innovative

Phytotaxa 139 (1) © 2013 Magnolia Press • 33
DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
Inventory of a Highly Diverse yet Little Known Group of Symbiotic Organisms (DEB 0715660 to The Field
Museum; PI R. Lücking), and Phylogenetic Diversity of Mycobionts and Photobionts in the Cyanolichen
Genus Dictyonema, with Emphasis on the Neotropics and the Galapagos Islands (DEB 0841405 to George
Mason University; PI J. Lawrey; Co-PIs: R. Lücking, P. Gillevet). Research by K. Wilk was funded by the W.
Szafer Institute of Botany, Polish Academy of Sciences, through a statutory fund. The Universidad Distrital
Francisco José de Caldas is thanked for the support to the lichen herbarium and the curatorial work of the
UDBC collections, and we especially acknowledge the invaluable help of laboratory assistant Alejandra
Suárez. The curators of the herbaria cited, in particular Harrie Sipman (B), Holger Thüs (BM), Philippe Clerc
(G), Soili Stenroos (H), Bruno Dennetière (PC), Marianne Hamnede and Anders Tehler (S), Roland Moberg
(UPS), Gregory McKee and Rusty Russell (US), and Anton Igersheim (W), were extremely helpful in
providing access to type material and other relevant collections. Marcela Cáceres helped to locate type
material at BM. Linda in Arcadia is thanked for advice concerning the nomenclature of Cora pavonia.
Adriano Spielmann and Marcelo Marcelli acknowledge the support from FAPESP, CNPq and FUNDECT.
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DICTYONEMA AND CORA (AGARICALES: HYGROPHORACEAE)
Index to Scientific Names
aeruginosa (Cilicia) ........................................................................................................................................................... 32
aeruginosulum (Dictyonema)....................................................................................................................................... 20, 30
aeruginosum (Dichonema)................................................................................................................................................. 31
aeruginosum (Dictyonema) ................................................................................................................................................ 27
arachnoidea (Cora)........................................................................................................................................................ 6, 31
aspera (Cora).......................................................................................................................................................8, 9, 30, 31
bovei (Cora) ..................................................................................................................................................... 3, 5, 6, 18, 31
brasiliensis (Corella)...................................................................................................................................................... 2, 31
byssoidea (Cora)........................................................................................................................................ 11, 12, 16, 20, 31
caespitosa (Laudatea) ..................................................................................................................................................29, 30
caespitosum (Dictyonema) ...........................................................................................................................................29, 30
ciferrii (Wainiocora)....................................................................................................................................................... 3, 31
confusum (Dictyonema) ..................................................................................................................................................... 27
coppinsii (Dictyonema) ...................................................................................................................................................... 30
crustaceum (Rhipidonema) ................................................................................................................................................ 32
cyphellifera (Cora)................................................................................................................................................. 12, 13, 31
cyphelloides (Stereum) ....................................................................................................................................................... 26
diducens (Dictyonema)................................................................................................................................................. 27, 29
elegans (Gyrolophium)...............................................................................................................................................3, 6, 31
excentricum (Dictyonema) ...........................................................................................................................................27, 28
expansum (Dictyonema)..................................................................................................................................................... 31
galapagoense (Dictyonema)............................................................................................................................................... 30
glabrata (Cora) ..................................................................................................................................................2, 3, 5, 6, 31
glabrata (Thelephora).................................................................................................................................................. 3, 5, 6
glabratum (Dictyonema) .................................................................................................................................................. 2, 3
gyrolophia (Cora).......................................................................................................................................................3, 6, 31
hernandezii (Dictyonema)............................................................................................................................................24, 29
hirsuta (Cora)...............................................................................................................................................8, 12, 16, 20 ,30
hirsutum (Dictyonema).....................................................................................................................................................3, 8
hirsutum (Stereum) .............................................................................................................................................................32
imperfectum (Cyphellostereum) .........................................................................................................................................26
inversa (Cora)........................................................................................................................................................14, 15, 30
irpicinum (Dictyonema) ......................................................................................................................................... 22, 27, 28
irpicinum f. scabridum (Rhipidonema) ..............................................................................................................................22
irrigatum (Corticium)...................................................................................................................................................29, 30
irrigatum (Dictyonema) .........................................................................................................................................27, 29, 30
japonicum (Dictyonema).................................................................................................................................................... 27
laxum (Dictyonema) ........................................................................................................................................................... 27
ligulatum (Dictyonema) ...............................................................................................................................................27, 28
ligulatum f. scabridum (Dictyonema) ................................................................................................................................ 22
mauritianum (Gyrolophium) ...................................................................................................................................... 3, 6, 31
melvinii (Corella) ............................................................................................................................................................... 31
melvinii (Dictyonema) ........................................................................................................................................................ 31
membranaceum (Dictyonema) ........................................................................................................................................... 31
membranaceum var. guadalupense (Dictyonema) ............................................................................................................. 32
metallicum (Dictyonema) ....................................................................................................................................... 22, 23, 29
minor (Cora) ................................................................................................................................................................16, 31
minus (Dictyonema) ....................................................................................................................................................... 3, 16
minutula (Podoscypha) ...................................................................................................................................................... 26
montana (Ulva) ................................................................................................................................................................3, 4
moorei (Dictyonema).................................................................................................................................................... 26, 27
neesiana (Cora).................................................................................................................................................................. 31
nitidum (Cyphellostereum)........................................................................................................................................... 24, 26
obscuratum (Dictyonema)......................................................................................................................................24, 25, 30
pallens (Stereophyllum)...................................................................................................................................................... 26
pavonia (Cora)................................................................................................................................................. 2, 4, 6, 14, 31
pavonia (Thelephora).......................................................................................................................................................4, 6

LÜCKING ET AL.38 • Phytotaxa 139 (1) © 2013 Magnolia Press
pectinatum (Dictyonema) ...................................................................................................................................................27
phyllogenum (Cyphellostereum)...................................................................................................................................24, 26
phyllogenum (Dichonema) .................................................................................................................................................24
phyllophilum (Dictyonema)....................................................................................................................................22, 29, 30
puiggarii (Rhipidonema).................................................................................................................................................... 32
pusiolum (Cyphellostereum)............................................................................................................................................... 26
reflexa (Auricularia)........................................................................................................................................................... 32
reticulifera (Cora)......................................................................................................................................................5, 6, 31
scabridum (Dictyonema)..............................................................................................................................................22, 27
schenckianum (Dictyonema) ........................................................................................................................................ 26, 27
sericeum (Dictyonema) ...................................................................................................................................... 2, 22, 25, 27
sericeum f. laminosum (Dictyonema) ................................................................................................................................ 32
sericeum f. membranaceum (Dictyonema) ........................................................................................................................ 32
sericeum f. phyllophilum (Dictyonema)....................................................................................................................... 22, 29
spongiosum (Dictyonema)............................................................................................................................................27, 28
squamiformis (Cora)....................................................................................................................................................16, 31
strigosa (Cora)............................................................................................................................................................. 18, 30
textilis (Thelephora) ........................................................................................................................................................... 32
thelephora (Dictyonema) ................................................................................................................................................... 30
tomentosa (Corella)............................................................................................................................................................ 31
uleana (Thelephora)........................................................................................................................................................... 26
zahlbruckneri (Corella)...................................................................................................................................................... 31