Ten new species of lichenized Basidiomycota in the genera Dictyonema and Cora (Agaricales:...

PHYTOTAXA

ISSN 1179-3155 (print edition)

ISSN 1179-3163 (online edition)Copyright copy 2013 Magnolia Press

Phytotaxa 139 (1) 1ndash38 (2013) wwwmapresscomphytotaxa

Article

httpdxdoiorg1011646phytotaxa13911

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 LUumlCKING1 MANUELA DAL-FORNO2 JAMES D LAWREY2 FRANK BUNGARTZ3

MARIacuteA E HOLGADO ROJAS4 JESUacuteS E HERNAacuteNDEZ M5 MARCELO P MARCELLI6 BIBIANA

MONCADA7 EDUARDO A MORALES8 MATTHEW P NELSEN19 ELIAS PAZ10 LUIS SALCEDO11

ADRIANO A SPIELMANN12 KARINA WILK13 SUSAN WILL-WOLF14 amp ALBA YAacuteNEZ-AYABACA3

1Integrative Science and Collections The Field Museum 1400 South Lake Shore Drive Chicago Illinois 60605 USA

email rluckingfieldmuseumorg2Department of Environmental Science and Policy George Mason University 4400 University Drive Fairfax VA 22030-4444 USA3Biodiversity Assessment Charles Darwin Foundation (AISBL) Puerto Ayora Santa Cruz Galaacutepagos Ecuador4Facultad de Ciencias Biologicas Universidad Nacional de San Antonio Abad del Cusco Av de la Cultura Nro 733 Cusco Peruacute5Fundacioacuten Instituto Botaacutenico de Venezuela Divisioacuten de Plantas no Vasculares Seccioacuten Hongos y Liacutequenes Ave Salvador Allende

Jardiacuten Botaacutenico de Caracas Universidad Central de Venezuela Apartado 2156 Caracas 1010-A Venezuela6Instituto de Botacircnica Nuacutecleo de Pesquisa em Micologia Caixa Postal 68041 Satildeo PauloSP CEP 04045-972 Brazil7Licenciatura en Biologiacutea Universidad Distrital Francisco Joseacute de Caldas Cra 4 No 26B-54 Torre de Laboratorios Herbario

Bogotaacute Colombia8Herbario Criptogaacutemico Universidad Catoacutelica Boliviana San Pablo Calle M Maacuterquez esq Plaza Jorge Trigo sn PO Box 5381

Cochabamba Bolivia9Committee on Evolutionary Biology University of Chicago 1025 E 57th Street Chicago Illinois 60637 USA10Facultad de Ciencias Biologicas Universidad Nacional de San Antonio Abad del Cusco Av de la Cultura Nro 733 Cusco Peruacute11Laboratorio de Ecofisiologiacutea Vegetal Facultad de Ciencias Naturales y Matemaacutetica Universidad Nacional Federico Villarreal

Lima Peruacute12Laboratoacuterio de Botacircnica Departamento de Biologia Centro de Ciecircncias Bioloacutegicas e da Sauacutede Universidade Federal de Mato

Grosso do Sul Cidade Universitaacuteria Caixa Postal 549 CEP 79070-900 Campo Grande Mato Grosso do Sul Brazil13Laboratory of Lichenology W Szafer Institute of Botany Polish Academy of Sciences Lubicz 46 PL-31-512 Krakoacutew Poland14Department of Botany University of Wisconsin-Madison 430 Lincoln Dr Madison Wisconsin 53706 USA

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 amp Luumlcking sp nov C aspera

Wilk Luumlcking amp E Morales sp nov C byssoidea Luumlcking amp Moncada sp nov C cyphellifera Dal-Forno Bungartz

amp Luumlcking sp nov C inversa Luumlcking amp Moncada sp nov C squamiformis Wilk Luumlcking amp Yaacutenez-Ayabaca sp

nov C strigosa Luumlcking E Paz amp L Salcedo sp nov Dictyonema aeruginosulum Luumlcking Nelsen amp Will-Wolf sp

nov D diducens Nyl ex Luumlcking sp nov D metallicum Luumlcking Dal-Forno amp Lawrey sp nov and D obscuratum

Luumlcking Spielmann amp 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 amp Luumlcking) Moncada amp Luumlcking

comb nov C minor (Luumlcking E Navarro amp Sipman) Luumlcking comb nov Corella melvinii (Chaves Luumlcking amp

Umantildea) Luumlcking Dal-Forno amp Lawrey comb nov Cyphellostereum phyllogenum (Muumlll Arg) Luumlcking Dal-Forno amp

Lawrey comb nov Dictyonema caespitosum (Johow) Luumlcking comb nov D irrigatum (Berk amp M A Curtis)

Luumlcking comb nov D phyllophilum (Parmasto) Luumlcking Dal-Forno amp Lawrey comb et stat nov and D scabridum

(Vain) Luumlcking 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

Accepted by Mohammad Sohrabi 15 Sept 2013 published 24 Oct 2013 1

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 amp Luumlcking 2013) The family Hygrophoraceae (Agaricales) contains two diverse lichenized clades Lichenomphalia Redhead Lutzoni Moncalvo amp Vilgalys (Redhead et al 2002 38) slat and Dictyonema C Agardh ex Kunth (Kunth 1822 1) slat (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 slat 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 (Moumlller 1893 Oberwinkler 1970 Parmasto 1978 Larcher amp 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 slat 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 1987andashc Fritz-Sheridan amp Portecop 1987 Iacomini et al 1987 Fritz-Sheridan 1988 Larcher amp 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 Luumlcking 2008 Lawrey et al 2009 Yaacutenez 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 slat can be divided into five genera Cyphellostereum D A Reid (Reid 1965 336) Dictyonema sstr Acantholichen P M Joslashrg (Joslashrgensen 1998 444) Cora and Corella (Lawrey et al 2009 Dal-Forno et al 2013) and this concept was applied to a treatment of Galaacutepagos basidiolichens (Yaacutenez 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

sstr Cyphellostereum has cyphelloid basidiomata emerging from the undifferentiated lichenized thallus whereas Dictyonema sstr 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 amp Vicente 1979) this taxon in reality forms a separate genus sister to Acantholichen (Dal-Forno et al 2013) supported by its different cortex structure

LUumlCKING ET AL2 bull Phytotaxa 139 (1) copy 2013 Magnolia Press

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 slat 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 Joseacute 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 Kotildeljalg 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 Luumlcking E Navarro amp Sipman (Chaves et al 2004 247) and D hirsutum Moncada amp Luumlcking (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 slat (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 amp D Mohr (Weber amp Mohr 1805 Fries 1838 Saccardo 1888b) but this is an error since Weber amp Mohr (1805) did not publish a separate name or species but referred to Swartzs (1806) upcoming publication on the replacement name Thaelaephora pavonia oder ehemaligen Ulva montana Swartz (Weber amp 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 (equiv 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 [equiv C pavonia (Weber amp 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 352) In 1838 he cited the wrong

Phytotaxa 139 (1) copy 2013 Magnolia Press bull 3DICTYONEMA AND CORA (AGARICALES HYGROPHORACEAE)

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 413) especially since the name mentioned in Weber amp Mohr (1805) as mentioned above is an indirect reference to Swartz (1806)

FIGURE 1 Cora pavonia A Specimen in the field (Colombia Luumlcking sn) B Lobe underside enlarged showing hymenophore with involute margins (Venezuela Hernaacutendez 1778) C Part of the lectotype of Ulva montana (equiv Thelephora pavonia) in BM showing undulate lobe surface and underside with hymenophore Scale in A = 10 mm in BndashC = 1 mm


FIGURE 2 Cora spp AndashB Cora glabrata (lectotype of Thelephora glabrata in UPS) C Cora bovei (isotype in NY) DndashF C

reticulifera D Lobe underside with hymenophore (isotype in BM) E Lobe showing upper side (Ecuador Cole 123) F Lobe underside with hymenophore (Ecuador Luumlcking 26201) Scale in AndashC = 5 mm in D F = 1 mm in E = 10 mm

The type of Thelephora glabrata is not well-developed (Fig 2AndashB) 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 5ndash10 semicircular lobes per thallus with the lobes 1ndash2 cm wide and 1ndash3 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 03


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 200ndash300 microm thick in section with the upper cortex formed by a 25ndash50 microm thick layer of rather loosely packed irregular hyphae supported by a 20ndash30 microm high medullary layer of spaced groups of densely packed anticlinal hyphae The photobiont layer is 100ndash150 microm 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 2DndashF) 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 1987andashc Fritz-Sheridan amp Portecop 1987 Iacomini et al 1987 Fritz-Sheridan 1988 Hawksworth 1988 Larcher amp 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 Lis ting of Interesting Plants of the World [http wwwnewcropsuqeduaulistingspecies_pages_DDictyonema_glabratumhtm] 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 amp Luumlcking sp nov (Fig 3)Mycobank 805376Genbank 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

HolotypemdashVENEZUELA Meacuterida Parque Nacional Sierra Nevada surroundings of Laguna de Mucubajiacute 8ordm 47 N 70ordm 49 W 3626 m 6 December 2009 Hernaacutendez 1780 (VEN)

Thallus terricolous between bryophytes rarely epiphytic on bryophyte-laden branches foliose up to 10 cm across composed of 5ndash10 semicircular lobes per thallus lobes 1ndash3(ndash5) cm wide and 1ndash5(ndash7) 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 02ndash03 mm long and


25ndash50 microm 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 250ndash350 microm thick with upper cortex photobiont layer and medulla upper cortex formed by a 25ndash50 microm thick layer of rather loosely

FIGURE 3 Cora arachnoidea A Specimen in the field (Colombia Luumlcking 32700) B Lobe enlarged showing tomentose upper surface (holotype) C Lobe underside showing hymenophore with involute byssoid margins (Venezuela Hernaacutendez 1782) Scale in A C = 5 mm in B = 1 mm


packed periclinal 4ndash5 microm thick hyphae supported by a 20ndash30 microm high medullary layer of spaced groups of densely packed anticlinal 3ndash5 microm thick hyphae photobiont layer 50ndash150 microm thick irregular composed of clusters of short coiled cyanobacterial filaments wrapped in a dense paraplectenchymatous hyphal sheath formed by jigsaw puzzle-shaped cells clusters 20ndash30 microm diam individual photobiont cells 10ndash12 microm broad and 6ndash8 microm long dark blue-green to yellow-green in upper portions penetrated by tubular fungal hyphae heterocytes sparse hyaline to pale yellow 8ndash10 microm wide and 5ndash6 microm long cells of hyphal sheath wavy in lateral outline 3ndash4 microm thick medulla 50ndash100 microm thick composed of loosely woven irregularly arranged to more or less periclinal hyphae 4ndash5 microm thick clamp connections not observed

Hymenophore developed as irregular to angular or elongate resupinate patches dispersed on the underside patches 3ndash10 mm diam with pale yellow smooth surface and byssoid margins hymenophore in section 50ndash100 microm thick composed of a paraplectenchymatous layer resting on loose 4ndash6 microm thick generative medullary hyphae and supporting the hymenium hymenium composed of numerous palisade-like basidioles and scattered basidia basidioles 20ndash30 times 5ndash6 microm basidia 25ndash35 times 5ndash7 microm 4-sterigmate basidiospores (few seen) ellipsoid non-septate hyaline 7ndash8 times 25ndash35 microm

Chemistry no substances detected by TLCDistribution and EcologymdashThis 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

EtymologymdashThe epithet refers to the arachnoid tomentum on the upper surfaceRemarksmdashCora arachnoidea is the second species known with a tomentose surface after Cora hirsuta

(Moncada amp Luumlcking) Moncada amp Luumlcking comb nov [Mycobank 805388 bas Dictyonema hirsutum

Moncada amp Luumlcking in Lumbsch et al Phytotaxa 18 48 (2011) holotype Colombia Luumlcking 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 examinedmdashMEXICO Jalisco Parque Nacional Volcaacuten Nevado de Colima beyond entrance station in La Joya area near the campground 19deg 35 N 103deg 36 W 3415 m 26 December 2006 Egan 17538 (OMA) COSTA RICA Saacuten Joseacute Los Santos Forest Reserve Cerro de la Muerte (Paciacutefico Central Conservation Area) Talamanca Ridge km 90 on road (ruta 2) from Cartago to San Isidro access road to towers on summit 83deg 45 W 9deg 34 N 3400ndash3500 m upper montane cloud forest and subalpine paramo zone disturbed low paramo shrub with Chusquea on bryophyte exposed September 2007 Luumlcking R18 (F) COLOMBIA Cundinamarca Paacuteramo de Sumapaz Laguna de Chizacaacute 4deg 17 N 74deg 12 W 3700ndash3750 m wet paramo with Espeletia August 2010 Luumlcking 32700 (F UDBC) VENEZUELA Venezuela Meacuterida Parque Nacional Sierra Nevada surroundings of Laguna de Mucubajiacute 8ordm 47 N 70ordm 49 W 3626 m 6 December 2009 Hernaacutendez 1779 1782 (VEN) BOLIVIA Santa Cruz Caballero Siberia region near La Palma 17deg 49 S 64deg 40 W 2582 m Yungas cloud forest epiphytic on bark 12 December 2004 Wilk 2780a (KRAM)

Cora aspera Wilk Luumlcking amp E Morales sp nov (Fig 4)Mycobank 805377Genbank 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


HolotypemdashBOLIVIA Santa Cruz Caballero Siberia region near La Palma 17deg 49 S 64deg 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 Luumlcking 33332) B Lobe enlarged showing rough upper surface (holotype) C Lobe underside showing hymenophore with slightly involute smooth margins (holotype) Scale in AndashB = 10 mm in C = 1 mm

Thallus epiphytic on twigs and branches of trees foliose up to 7 cm across composed of 1ndash5 semicircular lobes per thallus lobes 1ndash5 cm wide and 1ndash5 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


in thin concentric lines but lacking a continuous distinct tomentum trichomes where present in concentric lines free and irregularly arranged 01ndash015 mm long and 5ndash10 microm 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 200ndash300 microm thick with upper cortex photobiont layer and medulla upper cortex formed by a 25ndash50 microm thick layer of rather loosely packed irregularly arranged to nearly periclinal 4ndash5 microm thick hyphae supported by an indistinct 20ndash30 microm high medullary layer of spaced groups of densely packed anticlinal 3ndash5 microm thick hyphae photobiont layer 70ndash120 microm thick composed of clusters of short coiled cyanobacterial filaments wrapped in a dense paraplectenchymatous hyphal sheath formed by jigsaw puzzle-shaped cells clusters 20ndash30 microm diam individual photobiont cells 10ndash13 microm broad and 5ndash8 microm long dark blue-green to lighter green in upper portions penetrated by tubular fungal hyphae heterocytes sparse hyaline to pale yellow 9ndash12 microm wide and 5ndash6 microm long cells of hyphal sheath wavy in lateral outline 3ndash4 microm thick medulla 50ndash100 microm thick composed of loosely woven irregularly arranged to more or less periclinal hyphae 4ndash5 microm thick clamp connections not observed

Hymenophore developed as elongate resupinate patches forming more or less concentric ridges on the underside patches 1ndash10 mm long and 05ndash1 mm broad with pale yellow smooth surface and smooth involute margins hymenophore in section 50ndash100 microm thick composed of a paraplectenchymatous layer resting on loose 4ndash6 microm thick generative medullary hyphae and supporting the hymenium hymenium composed of numerous palisade-like basidioles and scattered basidia basidioles 20ndash35 times 5ndash6 microm basidia 25ndash40 times 6ndash7 microm 4-sterigmate basidiospores not observed


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

EtymologymdashThe epithet refers to the rough appearance of the surface especially when dryRemarksmdashParmasto (1978) and other authors (Mitidieri et al 1964 Feige 1969 Oberwinkler 1970

1984 2001 Parmasto 1978 Coxson 1987andashc Fritz-Sheridan amp Portecop 1987 Iacomini et al 1987 Fritz-Sheridan 1988 Hawksworth 1988 Larcher amp 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 examinedmdashCOSTA RICA Puntarenas Coto Brus San Vito Las Cruces Biological Station and Botanical Garden September 2007 Luumlcking 21016 (F) BOLIVIA La Paz Murillo Valle de Zongo Laguna de Viscachani a las orillas de la laguna 16ordm 12 S 68ordm 08 W 3805 m piso altoandino con pajonales y vegetacioacuten baja 13 November 2007 Luumlcking 23564 (F LPB) Cochabamba Chapare Incachaca 17deg 13 S 65deg 50 W 2018 m 7 July 2009 Luumlcking 29128 (F HCUCB) Cochabamba Chapare Corani 17deg 16 S 65deg 54 W 3262 m 7 July 2009 Luumlcking 29356 29364 (F HCUCB) PERU Cuzco Aguas Calientes near Machu Picchu August 2009 Vera sn (F)


Cora byssoidea Luumlcking amp Moncada sp nov (Fig 5)Mycobank 805378Genbank 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 Bogotaacute B Lobe enlarged showing arachnoid-byssoid upper surface (holotype) C Lobe underside showing hymenophore with strongly involute margins (holotype) Scale in BndashC = 1 mm


HolotypemdashCOLOMBIA Cundinamarca Choachiacute Paacuteramo El Verjoacuten 4ordm 33 N 74ordm 00 E 3200 m 18 August 2008 Luumlcking 25901 (F)

Thallus epiphytic on thin branches and twigs of paramo shrubs foliose up to 3 cm across composed of 1ndash3 semicircular lobes per thallus lobes 1ndash2 cm wide and 1ndash2 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 01ndash02 mm long and 5ndash6 microm 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 250ndash400 microm thick with upper cortex photobiont layer and medulla upper cortex formed by a 50ndash100 microm thick layer of rather loosely woven irregularly arranged 4ndash6 microm thick hyphae supported by a 30ndash50 microm high medullary layer of irregularly arranged to anticlinal 4ndash6 microm 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 100ndash200 microm thick irregular composed of clusters of short coiled cyanobacterial filaments wrapped in a dense paraplectenchymatous hyphal sheath formed by jigsaw puzzle-shaped cells clusters 30ndash50 microm diam individual photobiont cells 10ndash12 microm broad and 6ndash8 microm long dark blue-green to yellow-orange in upper portions penetrated by tubular fungal hyphae heterocytes sparse hyaline to pale yellow 8ndash10 microm wide and 5ndash6 microm long cells of hyphal sheath wavy in lateral outline 3ndash4 microm thick medulla 30ndash50 microm thick composed of loosely woven irregularly arranged to more or less periclinal hyphae 4ndash5 microm thick clamp connections not observed

Hymenophore developed as irregular to elongate resupinate patches dispersed on the underside patches 1ndash3 mm long and 05ndash1 mm broad with pale yellow smooth surface and strongly involute smooth margins hymenophore in section 50ndash100 microm thick composed of a paraplectenchymatous layer resting on loose 4ndash6 microm thick generative medullary hyphae and supporting the hymenium hymenium composed of numerous palisade-like basidioles and scattered basidia basidioles 25ndash30 times 5ndash7 microm basidia 25ndash35 times 5ndash8 microm 4-sterigmate basidiospores ellipsoid non-septate hyaline 7ndash9 times 3ndash4 microm

Chemistry no substances detected by TLCDistribution and EcologymdashThis species is known from a single collection growing on a shrub in the

Colombian paramo regions Due to its small size it is certainly overlookedEtymologymdashThe epithet refers to the arachnoid-byssoid submarginal tomentum on the upper surfaceRemarksmdashThis 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 amp Luumlcking sp nov (Fig 6)Mycobank 805379Genbank 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

HolotypemdashECUADOR Imbabura Andes Cantoacuten Cotacachi 22ordm 298 N 78ordm 27 246 W 2053 m small


entrance driveway towards the Reserva Alto Chocoacute 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 AndashB = 10 mm

Thallus epiphytic on twigs and branches of trees foliose up to 15 cm across composed of 20ndash30 semicircular lobes per thallus lobes 3ndash5 cm wide and 2ndash3 cm long lacking branching sutures light


aeruginous with slight concentric color zonation when fresh with shallow concentric ridges (8ndash11 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 285ndash400 microm thick with upper cortex photobiont layer and medulla upper cortex formed by a 25ndash35 microm thick layer of rather densely packed periclinal 4ndash5 microm thick hyphae supported by an indistinct 80ndash120 microm high medullary layer of spaced groups of densely packed anticlinal 3ndash5 microm thick hyphae photobiont layer 60ndash80 microm thick composed of clusters of short coiled cyanobacterial filaments wrapped in a dense paraplectenchymatous hyphal sheath formed by jigsaw puzzle-shaped cells clusters 40ndash70 microm diam individual photobiont cells 8ndash11 microm broad and 6ndash8 microm long bluish green to orange-yellow in upper portions penetrated by tubular fungal hyphae heterocytes sparse hyaline to pale yellow 9ndash12 microm wide and 5ndash6 microm long cells of hyphal sheath wavy in lateral outline 3ndash4 microm thick medulla 100ndash200 microm thick composed of loosely woven irregularly arranged to more or less periclinal hyphae 4ndash5 microm thick clamp connections not observed

Hymenophore developed as stereoid to cyphelloid structures irregularly dispersed along the margins on the underside 5ndash10 mm long and 10ndash15 mm broad with white smooth surface and smooth margins hymenophore in section 70ndash100 microm thick composed of a paraplectenchymatous layer resting on loose 4ndash6 microm thick generative medullary hyphae and supporting the hymenium hymenium composed of numerous palisade-like basidioles and scattered basidia basidioles 20ndash35 times 5ndash8 microm basidia 18ndash25 times 7ndash9 microm 4-sterigmate basidiospores ellipsoid to lacrymoid non-septate hyaline 7ndash8 times 25ndash35 microm

Chemistry no substances detected by TLCDistribution and EcologymdashThis 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 patchesEtymologymdashThe epithet refers to the unusual type of hymenophoreRemarksmdashThis 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 Moumlller (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 Luumlcking amp Moncada sp nov (Fig 7)Mycobank 805380Genbank 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



FIGURE 7 Cora inversa A Specimen in the field showing irregular-fuzzy lobe margins and clusters of trichomes projecting from lower surface (Colombia Luumlcking 33340) B Lobes enlarged showing marginal soredia (Colombia Luumlcking 33308) C Lobe underside showing arachnoid surface (Colombia Luumlcking 25902) Scale in AndashB = 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 1ndash5(ndash10) semicircular lobes per thallus lobes 1ndash3 cm wide and 1ndash3 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


forming soredia composed of cy anobacte r ia l g ranules 30ndash50 microm 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 200ndash300 microm thick with upper cortex photobiont layer and medulla upper cortex formed by a 50ndash100 microm thick layer of loosely woven irregularly arranged 4ndash6 microm thick hyphae covered by a thin layer of distinctly periclinal compacted hyphae and supported by a 30ndash50 microm high medullary layer of irregularly arranged to anticlinal 4ndash6 microm thick hyphae photobiont layer 50ndash100 microm thick irregular composed of clusters of short coiled cyanobacterial filaments wrapped in a dense paraplectenchymatous hyphal sheath formed by jigsaw puzzle-shaped cells clusters 30ndash50 microm diam individual photobiont cells 9ndash12 microm broad and 5ndash6 microm long yellow-orange to olive-yellow in upper portions penetrated by tubular fungal hyphae heterocytes sparse hyaline to pale yellow 8ndash10 microm wide and 4ndash5 microm long cells of hyphal sheath wavy in lateral outline 3ndash4 microm thick medulla 30ndash50 microm thick composed of loosely woven irregularly arranged to more or less periclinal hyphae 4ndash5 microm thick clamp connections not observed

Hymenophore not observedChemistry no substances detected by TLCDistribution and EcologymdashThis species known from several collections growing on shrubs

particularly at their base in the Colombian paramoEtymologymdashThe epithet refers to the partially strigose underside as opposed to a similar tomentum

formed on the upper side by Cora hirsutaRemarksmdashCora inversa is one of several species producing soredia in the genus most of which being

undescribed Sorediate margins are also known from Cora minor (Luumlcking E Navarro amp Sipman)

Luumlcking comb nov [Mycobank 805389 bas Dictyonema minus Luumlcking E Navarro amp 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 examinedmdashCOLOMBIA Cundinamarca Choachiacute Paacuteramo El Verjoacuten 4ordm 33 N 74ordm 00 E 3200 m 18 August 2008 Luumlcking 25903 (F)

Cora squamiformis Wilk Luumlcking amp Yaacutenez-Ayabaca sp nov (Fig 8)Mycobank 805382Genbank 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

HolotypemdashBOLIVIA La Paz Franz Tamayo Madidi National Park Sanchez Pass between Pelechuco and Keara 14deg 43 S 69deg 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 3ndash5(ndash10) semicircular lobes per thallus lobes 05ndash1 cm wide and 05ndash1 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


(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 200ndash300 microm thick with upper cortex photobiont layer and medulla upper cortex much reduced formed by a 20ndash50 microm thick layer of loosely woven irregularly arranged 4ndash6 microm thick hyphae and a 15ndash25 microm thick layer of strongly compacted periclinal hyphae 3ndash5 microm

FIGURE 8 Cora squamiformis A Specimen in the field (Colombia Luumlcking sn) B Specimen with strongly bent and sinuose margins (Colombia Luumlcking sn) C Specimen in the herbarium (holotype) Scale in A = 10 mm in BndashC = 5 mm


thick and with brownish color medullary layer absent photobiont layer 50ndash100 microm thick irregular composed of clusters of short coiled cyanobacterial filaments wrapped in a dense paraplectenchymatous hyphal sheath formed by jigsaw puzzle-shaped cells clusters 30ndash50 microm diam individual photobiont cells 9ndash13 microm broad and 5ndash6 microm long green to yellow-orange in upper portions penetrated by tubular fungal hyphae heterocytes sparse hyaline to pale yellow 8ndash10 microm wide and 4ndash5 microm long cells of hyphal sheath wavy in lateral outline 3ndash4 microm thick medulla 30ndash80 microm thick composed of loosely woven irregularly arranged to more or less periclinal hyphae 4ndash5 microm thick clamp connections not observed

Hymenophore not observedChemistry no substances detected by TLCDistribution and EcologymdashThis species is known from several collections growing on soil between

bryophytes in the Ecuadorian and Bolivian high AndesEtymologymdashThe epithet refers to the thallus appearing squamulose rather than folioseRemarksmdashCora 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 examinedmdashECUADOR Napo Papallacta 3300 m disturbed wet paramo vegetation 4 July 2010 Luumlcking 32300 (F) BOLIVIA La Paz Franz Tamayo Madidi National Park Sanchez Pass between Pelechuco and Keara 14deg 43 S 69deg 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 15deg 15 S 69deg 03 W 4536 m high mountain vegetation (puna) on ground between mosses 12 October 2007 Wilk 7446 (F KRAM)

Cora strigosa Luumlcking E Paz amp L Salcedo sp nov (Fig 9)Mycobank 805383Genbank ITS barcoding sequence KF443241

Differing from the morphologically similar Cora hirsuta and the closely related C byssoidea in the strigose tomentum

developed mostly submarginally

HolotypemdashPERU Cuzco Piscacucho 13deg 10 S 72deg 21 W 2700ndash3800 m disturbed montane rainforest and pasture 4 August 2009 Paz amp Salcedo 3 (F)

Thallus on rocks associated with other lichens (Hypotrachyna and Rimelia) foliose up to 10 cm across composed of 1ndash3 semicircular lobes per thallus lobes 1ndash3 cm wide and 1ndash2 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) 1ndash15 mm long and 25ndash50 microm 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 250ndash350 microm thick with upper cortex photobiont layer and medulla upper cortex formed by a 25ndash50 microm thick layer of rather loosely packed to indistinctly periclinal 4ndash5 microm thick hyphae supported by a 25ndash50 microm high medullary layer of spaced groups of densely packed anticlinal 3ndash5 microm thick hyphae photobiont layer 50ndash150 microm thick irregular composed of clusters of short coiled cyanobacterial filaments wrapped in a dense paraplectenchymatous hyphal sheath formed by jigsaw puzzle-shaped cells clusters 20ndash


30 microm diam individual photobiont cells 10ndash13 microm broad and 5ndash7 microm long dark blue-green to orange-yellow in upper portions penetrated by tubular fungal hyphae heterocytes sparse hyaline to pale yellow 8ndash10 microm wide and 5ndash6 microm long cells of hyphal sheath wavy in lateral outline 3ndash4 microm thick medulla 50ndash80 microm thick composed of loosely woven irregularly arranged to more or less periclinal hyphae 4ndash5 microm 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


Hymenophore developed as irregular to elongate resupinate patches arranged in reticulate pattern or more or less concentric zones on the underside patches 1ndash3 mm long and 05ndash1 mm broad with white to pale yellowish finely arachnoid surface and slightly involute finely byssoid margins hymenophore in section 50ndash100 microm thick composed of a paraplectenchymatous layer resting on loose 4ndash6 microm 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 20ndash50 microm long and 4ndash5 microm thick basidioles 25ndash35 times 5ndash6 microm basidia 30ndash40 times 5ndash7 microm 4-sterigmate basidiospores not observed

Chemistry no substances detected by TLCDistribution and EcologymdashThis species is known from a single locality in a heavily disturbed montane

rain forest in Peru near Machu PicchuEtymologymdashThe epithet refers to the radially projecting conspicuous trichomesRemarksmdashThis 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 examinedmdashPERU Cuzco Piscacucho 13deg 10 S 72deg 21 W 2700ndash3800 m disturbed montane rainforest and pasture 4 August 2009 Diacuteaz amp Jihuallanco sn (F)

Dictyonema aeruginosulum Luumlcking Nelsen amp Will-Wolf sp nov (Fig 10)Mycobank 805384Genbank 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

HolotypemdashCOSTA RICA Alajuela Volcaacuten Tenorio National Park Piloacuten Biological Station Arenal-Tempisque Conservation Area Tilaraacuten Ridge 140 km NW of San Joseacute 25 km NNW of Tilaraacuten near Bijagua access road to station and river 84deg 59 W 10deg 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 300ndash800 microm thick (excluding the projections) composed of an upper photobiont layer 200ndash400 microm thick and a lower medulla (forming the hypothallus) 100ndash400 microm 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 8ndash12 microm wide and 4ndash5 microm high blue-green cells penetrated by tubular fungal hyphae heterocytes sparse pale yellow 7ndash11 microm wide and 3ndash4 microm high cells of hyphal sheath wavy in lateral outline 3ndash4 microm thick medullary hyphae and those associated with hyphal sheath straight 4ndash6 microm thick lacking clamp connections Projections in section formed by a network of medullary


hyphae 4ndash6 microm 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 50ndash100 microm thick composed of a paraplectenchymatous layer resting on loose medullary hyphae and


supporting the hymenium hymenium composed of numerous palisade-like basidioles and scattered basidia basidioles 10ndash20 times 5ndash7 microm basidia 15ndash25 times 5ndash8 microm 4-sterigmate basidiospores (few seen) ellipsoid to narrowly drop-shaped non-septate hyaline 7ndash9 times 3ndash4 microm

Chemistry no substances detected by TLCDistribution and EcologymdashThis species is thus far known from montane rain forest in the northern

Cordillera de Tilaraacuten 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

EtymologymdashThe epithet refers to the characteristic blue-green color of this species while most other species are either more bluish or greenish

RemarksmdashDictyonema 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) Luumlcking Dal-

Forno amp 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) Luumlcking 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 examinedmdashCOSTA RICA Alajuela Volcaacuten Tenorio National Park Piloacuten Biological Station Arenal-Tempisque Conservation Area Tilaraacuten Ridge 140 km NW of San Joseacute 25 km NNW of Tilaraacuten near Bijagua access road to station and river 84deg 59 W 10deg 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 Luumlcking Dal-Forno amp Lawrey sp nov (Fig 11)Mycobank 805385Genbank 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

HolotypemdashECUADOR Pichincha Riacuteo Guajalito Protected Forest 0deg09rsquoS 78deg39rsquoW 1800 m montane rainforest on tree trunk September 2008 Luumlcking 26255 (QCNE isotype F)

Thallus epiphytic on tree trunks and overgrowing nearby bryophytes appressed filamentous in irregular dispersed to confluent patches each 1ndash5 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 25ndash50 microm thick composed of an irregular photobiont layer but lacking a discernible medulla photobiont layer


composed of numerous cyanobacterial filaments wrapped in a closed hyphal sheath formed by jigsaw puzzle-shaped cells cyanobacterial filaments composed of 10ndash13 microm wide and 4ndash6 microm high dark aeruginous blue cells penetrated by tubular fungal hyphae heterocytes sparse hyaline 8ndash12 microm wide and 4ndash6 microm high cells of hyphal sheath wavy in lateral outline 3ndash4 microm thick hyphae associated with hyphal sheath straight 4ndash6 microm thick lacking clamp connections compacted prothallus mostly found by densely arranged empty hyphal sheaths admixed with straight hyphae

FIGURE 11 Dictyonema metallicum (Ecuador Luumlcking 26203) AndashB Specimen in the field C Thallus surface enlarged showing filaments Scale = 1 mm


Hymenophore not observed Chemistry no substances detected by TLCDistribution and EcologymdashThis 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

EtymologymdashThe epithet refers to the metallic shimmer of the thallus when dryRemarksmdashDictyonema metallicum is similar to the recently described D hernandezii Luumlcking Lawrey

amp 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 (Muumlll Arg) Luumlcking Dal-Forno

amp Lawrey comb nov [Mycobank 805396 bas Dichonema phyllogenum Muumlll Arg Flora 66 352 (1883) syn Dictyonema phyllogenum (Muumlll Arg) Zahlbr Cat Lich Univ 7 746 (1931) lectotype (Parmasto 1978 124) Malaysia (Borneo Sarawak) Beccari 1624 (G)] and C nitidum (Luumlcking) Luumlcking (Luumlcking 2008 Yaacutenez et al 2012) but those two species have a Cyphellostereum-type thallus lacking a distinct hyphal sheath and also lacking haustoria

Additional specimens examinedmdashECUADOR Pichincha Riacuteo Guajalito Protected Forest 0deg09rsquoS 78deg39rsquoW 1800 m montane rainforest on tree trunk September 2008 Luumlcking 26255 (F)

Dictyonema obscuratum Luumlcking Spielmann amp Marcelli sp nov (Fig 12)Mycobank 805386Genbank ITS barcoding sequence KF443223

Differing from Dictyonema phyllophilum and D schenckianum slat in the densely and irregularly interwoven dark

olive-green fibrils and the absence of a distinct hypothallus

HolotypemdashBRAZIL Satildeo Paulo Mogi-Graccedilu Martinho Prado Jr Mogi-Guaccedilu Ecological Reserve Fazenda Campininha 22deg 15 S 47deg 10 W 635 m interior of dense Cerrado 7 November 2007 Luumlcking

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 200ndash400 microm thick composed of an upper photobiont layer 150ndash250 microm thick and a lower medulla (forming the hypothallus) 50ndash100 microm 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 20ndash25 microm wide and 6ndash8 microm high dark green cells (becoming orange-yellow towards the tips) penetrated by tubular fungal hyphae often longitudinally divided heterocytes sparse pale yellow 15ndash20 microm wide and 6ndash9 microm high cells of hyphal sheath wavy in lateral outline 3ndash4 microm thick medullary hyphae and those associated with hyphal sheath straight 4ndash6 microm 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 200ndash400 microm thick composed of a paraplectenchymatous layer connected to loose medullary hyphae hymenium composed of numerous palisade-like basidioles and scattered basidia basidioles 20ndash30 times 5ndash7 microm basidia 30ndash40 times 5ndash8 microm 4-sterigmate basidiospores ellipsoid to narrowly drop-shaped non-septate hyaline 7ndash9 times 3ndash4 microm

Chemistry no substances detected by TLCDistribution and EcologymdashThis species is thus far known from Cerrado (Cerrado denso) vegetation in

the state of Satildeo Paulo Brazil where it grows on the corky bark of characteristic Cerrado trees


EtymologymdashThe 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 microm

RemarksmdashThis 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


and D schenckianum (Muumlll 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 examinedmdashBRAZIL Satildeo Paulo Mogi-Graccedilu Mogi-Guaccedilu Biological Reserve Fazenda Campininha Cerrado Seco 22deg 15 S 47deg 10 W 650 m interior of dense Cerrado 7 November 2007 Luumlcking 23025 23204 (F SP)

Key to currently accepted genera of Dictyonema slat

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 (5ndash7 microm 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 (7ndash20 microm 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 Joslashrg

- 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 Luumlcking Barillas amp Dal-Forno (Yaacutenez et al 2012)

[Illustration in Yaacutenez et al 2012 227 fig 1dndashf]

- Fibrils strongly appressed hyphal sheath around cyanobacterial filaments loose C nitidum (Luumlcking) Luumlcking

[Illustration in Luumlcking 2008 784 fig 257D]

3 Thallus terrestrial basidiomata common C pusiolum (Berk amp M A Curtis) D A Reid (Reid 1965 342)

[Syn Stereum cyphelloides Berk amp M A Curtis (Berkeley amp 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 (Muumlll Arg) Luumlcking Dal-Forno amp Lawrey

[Illustration in Luumlcking 2008 784 fig 257C]


Key to currently recognized species of Dictyonema sstr

1 Thallus appearing applanate microfruticulose each branch including 2ndash3 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 30cndashe]

- 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) Luumlcking

- 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)


green clamp connections present D ligulatum (Kremp) Zahlbr (Zahlbruckner 1908 239 Fig 13B)

[Dictyonema laxum Muumlll Arg Bot Jahrb 4 57 (1883)]


lobe surface mottled white and blue-green clamp connections absent D sericeum (Sw) Berk slat

[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 sstr described from the Caribbean D aeruginosum

(Blume amp 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 amp M A Curtis (Berkeley amp 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 13EndashF) 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 Yaacutenez amp Luumlcking (Yaacutenez et al 2012 234)


- Fibrils distinctly blue-green prothallus distinct hyphal sheath smooth

D schenckianum (Muumlll Arg) Zahlbr (Fig 14A)

[This name was used in a broader sense by Chaves et al (2004) and Yaacutenez 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 sstr have a combed appearance]

9 Fibrils closely appressed but not embedded in gelatinous matrix over bryophyes

D diducens Nyl ex Luumlcking 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 sn (BM-001084450) Thallus epiphytic on bry-


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 20ndash50 microm thick of numerous cyanobacterial filaments wrapped in a closed hyphal sheath formed by jigsaw puzzle-shaped cells cyanobacterial filaments composed of 8ndash14 microm wide and 4ndash5 microm high blue-green cells penetrated by tubular fungal hyphae heterocytes sparse pale yellow 7ndash12 microm wide and 3ndash4 microm high cells of hyphal sheath wavy in lateral outline 3ndash4 microm thick free hyphae associated with hyphal sheath straight 4ndash6 microm 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) DndashF D spongiosum (D syntype PC EndashF Guatemala Luumlcking 25561 F) Scale in A = 5 mm in BndashD F = 10 mm in E = 50 mm


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 Luumlcking sn 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 Luumlcking Lawrey amp 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 Luumlcking Dal-Forno amp Lawrey (Fig 11)


11 Thallus with coarse finger-like outgrowths D aeruginosulum Luumlcking Nelsen amp 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) Luumlcking (Fig 14C)

[Dictyonema caespitosum (Johow) Luumlcking comb nov Mycobank 805392 bas Laudatea caespitosa Johow

Jahrb Wiss Bot 15 386 (1884) holotype Brazil Duseacuten sn (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 amp M A Curtis) Luumlcking (Fig 14D)

[Dictyonema irrigatum (Berk amp M A Curtis) Luumlcking comb nov Mycobank 805393 bas Corticium irriga-

tum Berk amp M A Curtis Proc Amer Acad Arts amp Sci 4 123 (1860) holotype China Wright 108 (PC)]

14 Thallus usually on living leaves prothallus distinct white

D phyllophilum (Parmasto) Luumlcking Dal-Forno amp 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 Luumlcking Spielmann amp Marcelli (Fig 12)

- Thallus light to dark blue-green cells of the cyanobacterial filaments not divided

D aff irrigatum (Berk amp M A Curtis) Luumlcking (Fig 14F)

17 Thallus dark blue-green shiny western Europe D coppinsii Luumlcking Barrie amp Genney (Luumlcking et al 2014)

[Dictyonema interruptum auct non (Carmich ex Hook) Parmasto (= Rhizonema interruptum Luumlcking amp Barrie)

illustration in Luumlcking 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 Yaacutenez Dal Forno amp Bungartz (Yaacutenez et al 2012 234)

[Illustration in Yaacutenez et al 2012 235 fig 3andashc]

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 Luumlcking amp 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 Luumlcking E Paz amp L Salcedo (Fig 9)

- Upper surface tomentose with a narrow glabrous submarginal zone surface of hymenophore glabrous

C hirsuta (Moncada amp Luumlcking) Moncada amp Luumlcking

[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 Luumlcking amp E Morales (Fig 4)

- Upper surface arachnoid-tomentose throughout 6


6 Lobes up to 5 cm broad brown when fresh thallus mostly terrestrial

C arachnoidea J E Hern amp Luumlcking (Fig 3)

- Lobes up to 2 cm broad white when fresh thallus usually epiphytic C byssoidea Luumlcking amp 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 (Luumlcking E Navarro amp Sipman) Luumlcking

[Illustration in Chaves et al 2004 245 fig 1BndashD]

- 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 Luumlcking amp Yaacutenez-Ayabaca (Fig 8)

- Lobes up to 2 cm broad with concentrically undulate surface C bovei Speg (Fig 2CndashD)

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 amp Luumlcking (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 Luumlcking amp 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 amp D Mohr) Fr [nom illeg] Wainiocora ciferrii Tomas

- 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 2EndashF)

- Lobe surface plane hymenophore irregular with slightly upturned margins especially when dry

C glabrata (Spreng) Fr (Fig 2AndashB)

Key to currently recognized species of Corella

1 Thallus forming irregular isidioid to finger-like outgrowths

C melvinii (Chaves Luumlcking amp Umantildea) Luumlcking Dal-Forno amp Lawrey

[Corella melvinii (Chaves Luumlcking amp Umantildea) Luumlcking Dal-Forno amp Lawrey comb nov Mycobank 805394

bas Dictyonema melvinii Chaves Luumlcking amp Umantildea in Chaves et al Bryologist 107 244 (2004) holotype Costa

Rica Chaves 122 (INB-3762769) illustration in Chaves et al 2004 245 fig 1EndashF]

- Thallus lacking isidioid outgrowths but sometimes forming irregular lobules C brasiliensis Vain

[Syn Corella tomentosa Vain (Vainio 1899) Corella zahlbruckneri Schiffn (Zahlbruckner 1909) illustration in

Dal-Forno et al 2013 fig 3OndashP]

The following names have not yet been checked since type material was not located and hence their taxonomic status remains uncertain

Dichonema aeruginosum Blume amp T Nees Nova Acta Acad Caes Leop-Carol 13 12 (1826) equiv Cora

neesiana Leacutev Ann Sci Nat Bot Seacuter 3 5 154 (1846) [nom illeg] Indonesia (Java)Dictyonema expansum Pouls Vidensk Medd Naturhist Foren Koslashbenhavn 1899 280 (1899) Indonesia

(Java)Dictyonema membranaceum C Agardh Syst Alg 85 (1824) Mariana Islands


Dictyonema membranaceum var guadalupense Rabenh Hedwigia 13 7 (1874) GuadeloupeDictyonema 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 Coacuterdoba 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 (eg 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 slat 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 amp Sipman (1991) nor Acantholichen pannarioides which was not yet described at the time Of the 40 species distinguished here 38 would be included in Parmastos concept of Dictyonema slat 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 sstr 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 TICOLICHEN (DEB 0206125 to The Field Museum PI Robert Luumlcking) Neotropical Epiphytic Microlichens ndash An Innovative


Inventory of a Highly Diverse yet Little Known Group of Symbiotic Organisms (DEB 0715660 to The Field Museum PI R Luumlcking) 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 Luumlcking 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 Joseacute 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 Suaacuterez The curators of the herbaria cited in particular Harrie Sipman (B) Holger Thuumls (BM) Philippe Clerc (G) Soili Stenroos (H) Bruno Dennetiegravere (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 Caacuteceres 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

References

Aptroot A amp Sipman HJM (1991) New lichens and lichen records from New Guinea Willdenowia 20 221ndash256

Asahina Y (1944) Lichenologische Notizen (XXV) Journal of Japanese Botany 20 129ndash134

Azenha G Iturriaga T Michelangeli FI amp Rodriguez E (1998) Ethnolichenology biological activity and

biochemistry of Amazonian lichen species Emanations from the Rainforest 1 8ndash14

Berkeley MJ (1843) Notices of some Brazilian fungi London Journal of Botany 2 629ndash643

Berkeley MJ (1872) Australian Fungi received principally from Baron F von Mueller and Dr R Schomburgk Journal

of the Linnean Society Botany 13 155ndash177

httpdxdoiorg101111j1095-83391872tb02397ax

Berkeley MJ amp Curtis MA (1868) Fungi Cubenses (Hymenomycetes) [cont] Journal of the Linnean Society Botany

10 321ndash341

httpdxdoiorg101111j1095-83391868tb00648x

Carbonero ER Sassaki GL Gorin PAJ amp Iacomini M (2002) A (1gt6)-linked -mannopyrananan pseudonigeran

and a (1gt4)-linked -xylan isolated from the lichenised basidiomycete Dictyonema glabratum FEMS Microbiology

Letters 206 175ndash178


Chaves JL Luumlcking R Sipman HJM Umantildea L amp Navarro E (2004) A first assessment of the ticolichen

biodiversity inventory in Costa Rica the genus Dictyonema (Polyporales Atheliaceae) The Bryologist 107 242ndash

249

httpdxdoiorg1016390007-2745(2004)107[0242afaott]20co2

Coxson DS (1987a) Effects of desiccation on net photosynthetic activity in the basidiomycete lichen Cora pavonia E

Fries from the cloudmist zone of the tropical volcano La Soufriere (Guadeloupe) The Bryologist 90 241ndash245

Coxson DS (1987b) Net photosynthetic response patterns of the basidiomycete lichen Cora pavonia (Web) E Fries

from the tropical volcano La Soufriere (Guadeloupe) Oecologia 73 454ndash458

httpdxdoiorg101007bf00385264

Coxson DS (1987c) The temperature dependence of photoinhibition in the tropical basidiomycete lichen Cora pavonia

E Fries Oecologia 73 447ndash453


Dal-Forno M Lawrey JD Sikaroodi M Bhattarai S Gillevet PM Sulzbacher M amp Luumlcking R (2013) Starting

from scratch evolution of the lichen thallus in the basidiolichen Dictyonema (Agaricales Hygrophoraceae) Fungal

Biology 117 584ndash598

httpdxdoiorg101016jfunbio201305006

Elifio SL Da Silva MLCC Iacomini M amp Gorin PAJ (2000) A lectin from the lichenized Basidiomycete

Dictyonema glabratum New Phytologist 148 327ndash334

httpdxdoiorg101046j1469-8137200000758x

Feige B (1969) Stoffwechselphysiologische Untersuchungen an den tropischen Basidiolichene Cora pavonia (Sw) Fr

Flora 160 169ndash180

Fries E (1821) Systema Mycologicum Sistens Fungorum Ordines Genera et Species Huc Usque Cognitas Quas ad

Normam Methodi Naturalis Determinavit Vol 1 Lund


httpdxdoiorg105962bhltitle5378

Fries E (1825) Systema Orbis Vegetabilis Primas lineas novae constrictionis periclitatur Elias Fries Pars I Plantae

homonemeae Lund

Fries E (1838) Epicrisis Systematis Mycologici Uppsala

Fritz-Sheridan RP (1988) Nitrogen fixation on a tropical volcano La Soufriere nitrogen fixation by the pioneer lichen

Dictyonema glabratum Lichenologist 20 96ndash100

httpdxdoiorg101017s002428298800012x

Fritz-Sheridan RP amp Portecop J (1987) Nitrogen fixation on the tropical volcano La Soufriere (Guadeloupe) 1 A

survey of nitrogen fixation by blue-green algal microepiphytes and lichen endophytes Biotropica 19 194ndash199

Hariot P (1891) Observations sur les espegraveces du genre Dictyonema Bulletin de la Socieacuteteacute Mycologique de France 7

32ndash41

Hariot P (1892) Observations sur les espegraveces du genre Dictyonema Beihefte zum Botanischen Centralblatt 1892 19

Hawksworth DL (1988) A new name for Dictyonema pavonium (Swartz) Parmasto Lichenologist 20 101

Hennings PC (1897) Beitraumlge zur Pilzflora Suumldamerikas 2 Hedwigia 36 190ndash246

Henssen A (1963) Eine Revision der Flechtenfamilien Lichinaceae und Ephebaceae Symbolae Botanicae Upsalienses

18(1) 1ndash123

Hibbett DS Binder M Bischoff JF Blackwell M Cannon PF Eriksson OE Huhndorf S James T Kirk

PM Luumlcking R Lumbsch HT Lutzoni F Matheny PB McLaughlin DJ Powell MJ Redhead S Schoch

CL Spatafora JW Stalpers JA Vilgalys R Aime MC Aptroot A Bauer R Begerow D Benny GL

Castlebury LA Crous PW Dai YC Gams W Geiser DM Griffith GW Gueidan C Hawksworth DL

Hestmark G Hosaka K Humber RA Hyde KD Ironside JE Kotildeljalg U Kurtzman CP Larsson KH

Lichtwardt R Longcore J Miadlikowska J Miller A Moncalvo JM Mozley-Standridge S Oberwinkler F

Parmasto E Reeb V Rogers JD Roux C Ryvarden L Sampaio JP Schuumlssler A Sugiyama J Thorn

RG Tibell L Untereiner WA Walker C Wang Z Weir A Weiss M White MM Winka K Yao YJ amp

Zhang N (2007) A higher-level phylogenetic classification of the Fungi Mycological Research 111509ndash547

httpdxdoiorg101016jmycres200703004

Hodkinson BP amp Luumlcking R (2013) Lepidostromatales a new order of lichenized fungi (Basidiomycota

Agaricomycetes) with two new genera Ertzia and Sulzbacheromyces and one new species Lepidostroma

winklerianum Fungal Diversity (in press)

Iacomini M Zanin SMW amp Fontana JD (1987) Isolation and characterization of B-D-glucan heteropolysaccharide

and trehalose components of the basidiomycetous lichen Cora pavonia Carbohydrate Research 168 55ndash65

httpdxdoiorg1010160008-6215(87)80006-x

Johow F (1884) Die Gruppe der Hymenolichenen Ein Beitrag zur Kenntnis basidiosporer Flechten Pringsheims

Jahrbuumlcher fuumlr Wissenschaftliche Botanik 15 361ndash409

Joslashrgensen PM (1998) Acantholichen pannarioides a new basidiolichen from South America The Bryologist 101 444ndash

447

Karsten PA (1889) Fungi aliquot novi in Brasilia a Dre Edw Wainia anno 1885 lecti Hedwigia 28 190ndash195

Kotildeljalg U Nilsson RH Abarenkov K Tedersoo L Taylor AFS Bahram M Bates ST Bruns TD Bengtsson-

Palme J Callaghan TM Douglas B Drenkhan T Eberhardt U Duentildeas M Grebenc T Griffith G W

Hartmann M Kirk PM Kohout P Larsson E Lindahl BD Luumlcking R Martiacuten MP Matheny B Nguyen

NH Niskanen T Oja J Peay KG Peintner U Peterson M Oldmaa KP Saag L Saar R Schuumlssler A

Scott JA Seneacutes C Smith ME Suija A Taylor DL Telleria MT Weiss M amp Larsson K-H (2013)

Towards a unified paradigm for sequence-based identification of fungi Molecular Ecology (online first)

httpdxdoiorg101111mec12481

Kunth CS (1822) Synopsis plantarum quas in itinere circa plagas Orbis Novi colleg Humboldt et Bonpland Paris

Lange OL Buumldel B Zellner H Zotz G amp Meyer A (1994) Field Measurements of water relations and CO2

exchange of the tropical cyanobacterial basidiolichen Dictyonema glabratum in a Panamanian rainforest Botanica

Acta 107 279ndash290

Larcher W amp Vareschi V (1988) Variation in morphology and functional traits of Dictyonema glabratum from

contrasting habitats in the Venezuelan Andes Lichenologist 20 269ndash277

httpdxdoiorg101017s0024282988000301

Lawrey JD Luumlcking R Sipman HJM Chaves JL Redhead SA Bungartz F Sikaroodi M amp Gillevet PM

(2009) High concentration of basidiolichens in a single family of agaricoid mushrooms (Basidiomycota Agaricales

Hygrophoraceae) Mycological Research 113 1154ndash1171


Lightfoot J (1777) Flora Scotica or a Systematic Arrangement in the Linnaean Method of the Native Plants of

Scotland and the Hebrides Vol II White at Horaces Head London

Luumlcking R (2008) Foliicolous lichenized fungi Flora Neotropica Monograph 103 1ndash866

httpdxdoiorg1016390007-2745-1131224


Luumlcking R Barrie F amp Genney D (2014) Dictyonema coppinsii a new name for the European species known as

Dictyonema interruptum (Basidiomycota Agaricales Hygrophoraceae) with a validation of its photobiont

Rhizonema (Cyanoprokaryota Nostocales Rhizonemataceae) The Lichenologist 46 (in press)

Lumbsch HT (2002) Analysis of phenolic products in lichens for identification and taxonomy In Kranner I Beckett

RP amp Varma AK (eds) Protocols in Lichenology Culturing Biochemistry Ecophysiology and Use in

Biomonitoring 281ndash295 Springer Berlin Heidelberg

httpdxdoiorg101016s0031-9422(02)00238-8

Lumbsch HT Ahti T Altermann S Amo De Paz G Aptroot A Arup U Baacutercenas Pentildea A Bawingan PA

Benatti MN Betancourt L Bjoumlrk CR Boonpragob K Brand M Bungartz F Caacuteceres MES Candan M

Chaves JL Clerc P Common R Coppins BJ Crespo A Dal Forno M Divakar PK Duya MV Elix

JA Elvebakk A Fankhauser JD Farkas E Ferraro LI Fischer E Galloway DJ Gaya E Giralt M

Goward T Grube M Hafellner J Hernaacutendez M JE Herrera Campos MA Kalb K Kaumlrnefelt I Kantvilas

G Killmann D Kirika P Knudsen K Komposch H Kondratyuk S Lawrey JD Mangold A Marcelli

MP Mccune B Ines Messuti M Michlig A Miranda Gonzaacutelez R Moncada B Naikatini A Nelsen MP

Oslashvstedal DO Palice Z Papong K Parnmen S Peacuterez-Ortega S Printzen C Rico VJ Rivas Plata E

Robayo J Rosabal D Ruprecht U Salazar Allen N Sancho L Santos De Jesus L Santos Vieira T Schultz

M Seaward MRD Seacuterusiaux E Schmitt I Sipman HJM Sohrabi M Soslashchting U Zeuthen Soslashgaard M

Sparrius LB Spielmann A Spribille T Sutjaritturakan J Thammathaworn A Thell A Thor G Thuumls H

Timdal E Truong C Tuumlrk R Umantildea Tenorio L Upreti DK Van Den Boom P Vivas Rebuelta M Wedin

M Will-Wolf S Wirth V Wirtz N Yahr R Yeshitela K Ziemmeck F Wheeler T amp Luumlcking R (2011) One

hundred new species of lichenized fungi a signature of undiscovered global diversity Phytotaxa 18 1ndash127

Mattirolo O (1881) Contribuzioni allo studio del genere Cora Fries Nuovo Giornale Botanico Italiano 13 245ndash267

Metzner P (1934) Zur Kenntnis der Hymenolichenen Berichte der Deutschen Botanischen Gesellschaft 51 231ndash240 2

pl

Mitidieri J Joly S amp Ferraz EC (1964) Teste de antibiose exercida pelo extrato do liquens Parmelia tinctorum Desp

e Cora pavonia (Web) E Fries Revista de Agronomiacutea [Piracicaba] 39 119ndash121

Moumlller A (1893) Ueber die eine Thelephoree welche die Hymenolichenen Cora Dictyonema und Laudatea bildet


Montagne C (1848) Sixiegraveme centurie de plantes cellulaires exotiques nouvelles Deacutecades 1ndash2 Annales des Sciences

Naturelles 10 106ndash136

Nylander W (1885) Arthoniae novae America borealis Continuatio Flora 68 447ndash449

Oberwinkler F (1970) Die Gattungen der Basidiolichenen Vortraumlge aus dem Gesamtgebiet der Botanik NF 4 139ndash

169

Oberwinkler F (1980) Symbiotic relationships between fungus and alga in basidiolichens In Schwemmler W amp

Schenk HEA (eds) Endocytobiology Endosymbiosis and Cell Biology pp 305ndash315 Walter de Gruyter Berlin

Oberwinkler F (1984) Fungus-alga interactions in basidiolichens Beiheft zur Nova Hedwigia 79 739ndash774

Oberwinkler F (2001) Basidiolichens In Hock B (ed) The Mycota Vol IX Fungal Associations 211ndash225 Springer

Berlin Heidelberg New York

Oberwinkler F (2012) Basidiolichens In Hock B (ed) The Mycota Second Edition Vol IX Fungal Associations

341ndash362 Springer Berlin Heidelberg New York

httpdxdoiorg101007978-3-642-30826-0_16

Orange A James PW amp White FJ (2001) Microchemical Methods for the Identification of Lichens British Lichen

Society London

Parmasto E (1978) The genus Dictyonema (Thelephorolichenes) Nova Hedwigia 29 99ndash144

Patouillard N (1924) Quelques champignons du Tonkin Suite Bulletin Trimestrel de la Socieacuteteacute Mycologique de

France 40 29ndash37

Piovano M Chamy MC Garbarino JA amp Quilhot W (1995) Studies on Chilean lichens XXIV Secondary products

from Dictyonema glabratum (Basidiomycotina) Boletin Sociedad Chilena de Quiacutemica 40 163ndash165


Redhead SA Lutzoni F Moncalvo J-M amp Vilgalys R (2002) Phylogeny of agarics partial systematics solutions

for core omphalinoid genera in the Agaricales (Euagarics) Mycotaxon 83 19ndash57

Reid DA (1965) A monograph of the stipitate stereoid fungi Beihefte zur Nova Hedwigia 18 1ndash388

Saccardo PA (1888a) Sylloge Fungorum Omnium Hucusque Cognitorum Vol 4

Saccardo PA (1888b) Sylloge Fungorum Omnium Hucusque Cognitorum Vol 7

Spegazzini CL (1888) Fungi fuegiani Boletin de la Academiacutea Nacional de Ciencias de Coacuterdoba 11 135ndash311

Sprengel C (1820) Plantarum cryptogamicarum tropicarum pugillus Kongliga Svenska Vetenskaps-Akademiens

Handlingar Stockholm 1820 46ndash53

Swartz O (1788) Nova Genera et Species Plantarum seu Prodromus Descriptionum Vegetabilium Maximam Partem

Incognitarum quae sub Itinere in Indiam Occidentalem Annis 178387 Digessit Holmiae


Swartz O (1806) Flora Indiae Occidentalis London

Thomas MA Nash III TH amp Gries C (1997) Ecophysiological comparison of two tropical subtropical lichen

species Dictyonema glabratum from an alpine habitat and Coenogonium interplexum from a lowland forest

Bibliotheca Lichenologica 67 183ndash195

httpdxdoiorg101046j1469-8137200200360x

Tomaselli R (1950) Appunti sulla sistematica e distribuzione geografica dei Basidiolicheni Archivio Bot 28(2)[Terza

Ser 10(2)] 3ndash19

Trembley ML Ringli C amp Honegger R (2002a) Differential expression of hydrophobins DGH1 DGH2 and DGH3

and immunolocalization of DGH1 in strata of the lichenized basidocarp of Dictyonema glabratum New Phytologist

154 185ndash195

Trembley ML Ringli C amp Honegger R (2002b) Hydrophobins DGH1 DGH2 and DGH3 in the lichen-forming

basidiomycete Dictyonema glabratum Fungal Genetics and Biology 35 247ndash259

httpdxdoiorg101006fgbi20011325

Vainio EA (1890) Etude sur la classification et la morphologie des lichens du Breacutesil I Acta Societatis pro Fauna et

Flora Fennica 7 VndashXXIX 1ndash247


Vainio EA (1899) Lichenes novi rarioresque Ser III Hedwigia 38(Beiblatt) 253ndash259

Von Krombholz JV (1831) Naturgetreue Abbildungen und Beschreibungen der essbaren schaumldlichen und verdaumlchtigen

Schwaumlmme Vol 1 Prag

httpdxdoiorg101080037454809495193

Weber F amp Mohr DMH (1805) Einige Worte uumlber unsre bisherigen hauptsaumlchlich carpologischen Zergliederungen

von kryptogamischen Seegewaumlchsen Beitraumlge zur Naturkunde 1 204ndash329

Wolf JHD (1993) Epiphyte communities of tropical montane rain forests in the northern Andes I Lower montane

communities Phytocoenologia 22 1ndash52

Xavier Filho L amp Vicente C (1979) Observaciones morfoloacutegicas sobre Corella Boletim da Sociedade Broteriana

Serie 2 53 7ndash13

Yaacutenez A Dal-Forno M Bungartz F Luumlcking R amp Lawrey JD (2012) A first assessment of Galapagos

basidiolichens Fungal Diversity 52 225ndash244

httpdxdoiorg101007s13225-011-0133-x

Zahlbruckner A (1908) Flechten (Lichenes) In Engler HG amp Prantl KA Die Natuumlrlichen Pflanzenfamilien I(1)

239

Zahlbruckner A (1909) Lichenes (Flechten) Denkschriften der Kaiserlichen Akademie der Wissenschaften

Mathematisch-Naturwissenschaftliche Klasse 83 85ndash211

httpdxdoiorg101007978-3-662-24755-6

Zahlbruckner A (1931) Catalogus Lichenum Universalis Borntraeger Leipzig


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


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


Abstract

Introduction


Taxonomic Treatment

Conclusions

Acknowledgements

References


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 amp Luumlcking 2013) The family Hygrophoraceae (Agaricales) contains two diverse lichenized clades Lichenomphalia Redhead Lutzoni Moncalvo amp Vilgalys (Redhead et al 2002 38) slat and Dictyonema C Agardh ex Kunth (Kunth 1822 1) slat (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 slat 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 (Moumlller 1893 Oberwinkler 1970 Parmasto 1978 Larcher amp 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 slat 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 1987andashc Fritz-Sheridan amp Portecop 1987 Iacomini et al 1987 Fritz-Sheridan 1988 Larcher amp 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 Luumlcking 2008 Lawrey et al 2009 Yaacutenez 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 slat can be divided into five genera Cyphellostereum D A Reid (Reid 1965 336) Dictyonema sstr Acantholichen P M Joslashrg (Joslashrgensen 1998 444) Cora and Corella (Lawrey et al 2009 Dal-Forno et al 2013) and this concept was applied to a treatment of Galaacutepagos basidiolichens (Yaacutenez 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

sstr Cyphellostereum has cyphelloid basidiomata emerging from the undifferentiated lichenized thallus whereas Dictyonema sstr 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 amp Vicente 1979) this taxon in reality forms a separate genus sister to Acantholichen (Dal-Forno et al 2013) supported by its different cortex structure






(2013)

Taxonomic Treatment



























































































formed soredia





















































































































ent Cora Fr

















































































- Thallus plane 12


























































































Conclusions


Acknowledgements





References










10 321ndash341








249
















httpdxdoiorg101046j1469-8137200000758x








homonemeae Lund








32ndash41





18(1) 1ndash123
















httpdxdoiorg1010160008-6215(87)80006-x




447















httpdxdoiorg101017s0024282988000301








httpdxdoiorg1016390007-2745-1131224








httpdxdoiorg101016s0031-9422(02)00238-8

















pl









169








httpdxdoiorg101007978-3-642-30826-0_16


Society London



France 40 29ndash37



















httpdxdoiorg101046j1469-8137200200360x


Ser 10(2)] 3ndash19



154 185ndash195










httpdxdoiorg101080037454809495193






Serie 2 53 7ndash13





239



httpdxdoiorg101007978-3-662-24755-6










bovei (Cora) 3 5 6 18 31





















hirsuta (Cora) 8 12 16 20 30



















minor (Cora) 16 31



montana (Ulva) 3 4


neesiana (Cora) 31






























Abstract

Introduction


Taxonomic Treatment

Conclusions

Acknowledgements

References






(2013)

Taxonomic Treatment



























































































formed soredia





















































































































ent Cora Fr

















































































- Thallus plane 12


























































































Conclusions


Acknowledgements





References










10 321ndash341








249
















httpdxdoiorg101046j1469-8137200000758x








homonemeae Lund








32ndash41





18(1) 1ndash123
















httpdxdoiorg1010160008-6215(87)80006-x




447















httpdxdoiorg101017s0024282988000301








httpdxdoiorg1016390007-2745-1131224








httpdxdoiorg101016s0031-9422(02)00238-8

















pl









169








httpdxdoiorg101007978-3-642-30826-0_16


Society London



France 40 29ndash37



















httpdxdoiorg101046j1469-8137200200360x


Ser 10(2)] 3ndash19



154 185ndash195










httpdxdoiorg101080037454809495193






Serie 2 53 7ndash13





239



httpdxdoiorg101007978-3-662-24755-6










bovei (Cora) 3 5 6 18 31





















hirsuta (Cora) 8 12 16 20 30



















minor (Cora) 16 31



montana (Ulva) 3 4


neesiana (Cora) 31






























Abstract

Introduction


Taxonomic Treatment

Conclusions

Acknowledgements

References



















































































formed soredia





















































































































ent Cora Fr

















































































- Thallus plane 12


























































































Conclusions


Acknowledgements





References










10 321ndash341








249
















httpdxdoiorg101046j1469-8137200000758x








homonemeae Lund








32ndash41





18(1) 1ndash123
















httpdxdoiorg1010160008-6215(87)80006-x




447















httpdxdoiorg101017s0024282988000301








httpdxdoiorg1016390007-2745-1131224








httpdxdoiorg101016s0031-9422(02)00238-8

















pl









169








httpdxdoiorg101007978-3-642-30826-0_16


Society London



France 40 29ndash37



















httpdxdoiorg101046j1469-8137200200360x


Ser 10(2)] 3ndash19



154 185ndash195










httpdxdoiorg101080037454809495193






Serie 2 53 7ndash13





239



httpdxdoiorg101007978-3-662-24755-6










bovei (Cora) 3 5 6 18 31





















hirsuta (Cora) 8 12 16 20 30



















minor (Cora) 16 31



montana (Ulva) 3 4


neesiana (Cora) 31






























Abstract

Introduction


Taxonomic Treatment

Conclusions

Acknowledgements

References




















































































































ent Cora Fr

















































































- Thallus plane 12


























































































Conclusions


Acknowledgements





References










10 321ndash341








249
















httpdxdoiorg101046j1469-8137200000758x








homonemeae Lund








32ndash41





18(1) 1ndash123
















httpdxdoiorg1010160008-6215(87)80006-x




447















httpdxdoiorg101017s0024282988000301








httpdxdoiorg1016390007-2745-1131224








httpdxdoiorg101016s0031-9422(02)00238-8

















pl









169








httpdxdoiorg101007978-3-642-30826-0_16


Society London



France 40 29ndash37



















httpdxdoiorg101046j1469-8137200200360x


Ser 10(2)] 3ndash19



154 185ndash195










httpdxdoiorg101080037454809495193






Serie 2 53 7ndash13





239



httpdxdoiorg101007978-3-662-24755-6










bovei (Cora) 3 5 6 18 31





















hirsuta (Cora) 8 12 16 20 30



















minor (Cora) 16 31



montana (Ulva) 3 4


neesiana (Cora) 31






























Abstract

Introduction


Taxonomic Treatment

Conclusions

Acknowledgements

References


































































- Thallus plane 12


























































































Conclusions


Acknowledgements





References










10 321ndash341








249
















httpdxdoiorg101046j1469-8137200000758x








homonemeae Lund








32ndash41





18(1) 1ndash123
















httpdxdoiorg1010160008-6215(87)80006-x




447















httpdxdoiorg101017s0024282988000301








httpdxdoiorg1016390007-2745-1131224








httpdxdoiorg101016s0031-9422(02)00238-8

















pl









169








httpdxdoiorg101007978-3-642-30826-0_16


Society London



France 40 29ndash37



















httpdxdoiorg101046j1469-8137200200360x


Ser 10(2)] 3ndash19



154 185ndash195










httpdxdoiorg101080037454809495193






Serie 2 53 7ndash13





239



httpdxdoiorg101007978-3-662-24755-6










bovei (Cora) 3 5 6 18 31





















hirsuta (Cora) 8 12 16 20 30



















minor (Cora) 16 31



montana (Ulva) 3 4


neesiana (Cora) 31






























Abstract

Introduction


Taxonomic Treatment

Conclusions

Acknowledgements

References















































Conclusions


Acknowledgements





References










10 321ndash341








249
















httpdxdoiorg101046j1469-8137200000758x








homonemeae Lund








32ndash41





18(1) 1ndash123
















httpdxdoiorg1010160008-6215(87)80006-x




447















httpdxdoiorg101017s0024282988000301








httpdxdoiorg1016390007-2745-1131224








httpdxdoiorg101016s0031-9422(02)00238-8

















pl









169








httpdxdoiorg101007978-3-642-30826-0_16


Society London



France 40 29ndash37



















httpdxdoiorg101046j1469-8137200200360x


Ser 10(2)] 3ndash19



154 185ndash195










httpdxdoiorg101080037454809495193






Serie 2 53 7ndash13





239



httpdxdoiorg101007978-3-662-24755-6










bovei (Cora) 3 5 6 18 31





















hirsuta (Cora) 8 12 16 20 30



















minor (Cora) 16 31



montana (Ulva) 3 4


neesiana (Cora) 31






























Abstract

Introduction


Taxonomic Treatment

Conclusions

Acknowledgements

References




References










10 321ndash341








249
















httpdxdoiorg101046j1469-8137200000758x








homonemeae Lund








32ndash41





18(1) 1ndash123
















httpdxdoiorg1010160008-6215(87)80006-x




447















httpdxdoiorg101017s0024282988000301








httpdxdoiorg1016390007-2745-1131224








httpdxdoiorg101016s0031-9422(02)00238-8

















pl









169








httpdxdoiorg101007978-3-642-30826-0_16


Society London



France 40 29ndash37



















httpdxdoiorg101046j1469-8137200200360x


Ser 10(2)] 3ndash19



154 185ndash195










httpdxdoiorg101080037454809495193






Serie 2 53 7ndash13





239



httpdxdoiorg101007978-3-662-24755-6










bovei (Cora) 3 5 6 18 31





















hirsuta (Cora) 8 12 16 20 30



















minor (Cora) 16 31



montana (Ulva) 3 4


neesiana (Cora) 31






























Abstract

Introduction


Taxonomic Treatment

Conclusions

Acknowledgements

References




homonemeae Lund








32ndash41





18(1) 1ndash123
















httpdxdoiorg1010160008-6215(87)80006-x




447















httpdxdoiorg101017s0024282988000301








httpdxdoiorg1016390007-2745-1131224








httpdxdoiorg101016s0031-9422(02)00238-8

















pl









169








httpdxdoiorg101007978-3-642-30826-0_16


Society London



France 40 29ndash37



















httpdxdoiorg101046j1469-8137200200360x


Ser 10(2)] 3ndash19



154 185ndash195










httpdxdoiorg101080037454809495193






Serie 2 53 7ndash13





239



httpdxdoiorg101007978-3-662-24755-6










bovei (Cora) 3 5 6 18 31





















hirsuta (Cora) 8 12 16 20 30



















minor (Cora) 16 31



montana (Ulva) 3 4


neesiana (Cora) 31






























Abstract

Introduction


Taxonomic Treatment

Conclusions

Acknowledgements

References








httpdxdoiorg101016s0031-9422(02)00238-8

















pl









169








httpdxdoiorg101007978-3-642-30826-0_16


Society London



France 40 29ndash37



















httpdxdoiorg101046j1469-8137200200360x


Ser 10(2)] 3ndash19



154 185ndash195










httpdxdoiorg101080037454809495193






Serie 2 53 7ndash13





239



httpdxdoiorg101007978-3-662-24755-6










bovei (Cora) 3 5 6 18 31





















hirsuta (Cora) 8 12 16 20 30



















minor (Cora) 16 31



montana (Ulva) 3 4


neesiana (Cora) 31






























Abstract

Introduction


Taxonomic Treatment

Conclusions

Acknowledgements

References






httpdxdoiorg101046j1469-8137200200360x


Ser 10(2)] 3ndash19



154 185ndash195










httpdxdoiorg101080037454809495193






Serie 2 53 7ndash13





239



httpdxdoiorg101007978-3-662-24755-6










bovei (Cora) 3 5 6 18 31





















hirsuta (Cora) 8 12 16 20 30



















minor (Cora) 16 31



montana (Ulva) 3 4


neesiana (Cora) 31






























Abstract

Introduction


Taxonomic Treatment

Conclusions

Acknowledgements

References









bovei (Cora) 3 5 6 18 31





















hirsuta (Cora) 8 12 16 20 30



















minor (Cora) 16 31



montana (Ulva) 3 4


neesiana (Cora) 31






























Abstract

Introduction


Taxonomic Treatment

Conclusions

Acknowledgements

References

























Abstract

Introduction


Taxonomic Treatment

Conclusions

Acknowledgements

References


Ten new species of lichenized Basidiomycota in the genera Dictyonema and Cora (Agaricales:...

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Transcript of Ten new species of lichenized Basidiomycota in the genera Dictyonema and Cora (Agaricales:...