Phytotaxa 18: 1–127 (2011)
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Copyright © 2011 Magnolia Press
Monograph
ISSN 1179-3155 (print edition)
PHYTOTAXA
ISSN 1179-3163 (online edition)
PHYTOTAXA
18
One hundred new species of lichenized fungi:
a signature of undiscovered global diversity
H. THORSTEN LUMBSCH1*, TEUVO AHTI2, SUSANNE ALTERMANN3, GUILLERMO AMO DE PAZ4, ANDRÉ
APTROOT5, ULF ARUP6, ALEJANDRINA BÁRCENAS PEÑA7, PAULINA A. BAWINGAN8, MICHEL N.
BENATTI9, LUISA BETANCOURT10, CURTIS R. BJÖRK11, KANSRI BOONPRAGOB12, MAARTEN BRAND13,
FRANK BUNGARTZ14, MARCELA E. S. CÁCERES15, MEHTMET CANDAN16, JOSÉ LUIS CHAVES17, PHILIPPE
CLERC18, RALPH COMMON19, BRIAN J. COPPINS20, ANA CRESPO4, MANUELA DAL-FORNO21, PRADEEP K.
DIVAKAR4, MELIZAR V. DUYA22, JOHN A. ELIX23, ARVE ELVEBAKK24, JOHNATHON D. FANKHAUSER25,
EDIT FARKAS26, LIDIA ITATÍ FERRARO27, EBERHARD FISCHER28, DAVID J. GALLOWAY29, ESTER GAYA30,
MIREIA GIRALT31, TREVOR GOWARD32, MARTIN GRUBE33, JOSEF HAFELLNER33, JESÚS E. HERNÁNDEZ
M.34, MARÍA DE LOS ANGELES HERRERA CAMPOS7, KLAUS KALB35, INGVAR KÄRNEFELT6, GINTARAS
KANTVILAS36, DOROTHEE KILLMANN28, PAUL KIRIKA37, KERRY KNUDSEN38, HARALD KOMPOSCH39,
SERGEY KONDRATYUK40, JAMES D. LAWREY21, ARMIN MANGOLD41, MARCELO P. MARCELLI9, BRUCE
MCCUNE42, MARIA INES MESSUTI43, ANDREA MICHLIG27, RICARDO MIRANDA GONZÁLEZ7, BIBIANA
MONCADA10, ALIFERETI NAIKATINI44, MATTHEW P. NELSEN1, 45, DAG O. ØVSTEDAL46, ZDENEK PALICE47,
KHWANRUAN PAPONG48, SITTIPORN PARNMEN12, SERGIO PÉREZ-ORTEGA4, CHRISTIAN PRINTZEN49,
VÍCTOR J. RICO4, EIMY RIVAS PLATA1, 50, JAVIER ROBAYO51, DANIA ROSABAL52, ULRIKE RUPRECHT53,
NORIS SALAZAR ALLEN54, LEOPOLDO SANCHO4, LUCIANA SANTOS DE JESUS15, TAMIRES SANTOS
VIEIRA15, MATTHIAS SCHULTZ55, MARK R. D. SEAWARD56, EMMANUËL SÉRUSIAUX57, IMKE SCHMITT58,
HARRIE J. M. SIPMAN59, MOHAMMAD SOHRABI 2, 60, ULRIK SØCHTING61, MAJBRIT ZEUTHEN SØGAARD61,
LAURENS B. SPARRIUS62, ADRIANO SPIELMANN63, TOBY SPRIBILLE33, JUTARAT SUTJARITTURAKAN64,
ACHRA THAMMATHAWORN65, ARNE THELL6, GÖRAN THOR66, HOLGER THÜS67, EINAR TIMDAL68,
CAMILLE TRUONG18, ROMAN TÜRK69, LOENGRIN UMAÑA TENORIO17, DALIP K. UPRETI70, PIETER VAN
DEN BOOM71, MERCEDES VIVAS REBUELTA4, MATS WEDIN72, SUSAN WILL-WOLF73, VOLKMAR WIRTH74,
NORA WIRTZ75, REBECCA YAHR20, KUMELACHEW YESHITELA28, FRAUKE ZIEMMECK14, TIM WHEELER76,
ROBERT LÜCKING1
(full contract addresses of 103 authors on pages 3–5)
Magnolia Press
Auckland, New Zealand
Accepted by W.-Y. Zhuang: 8 Jan. 2011; published: 18 Feb. 2011
H. THORSTEN LUMBSCH ET AL.
One hundred new species of lichenized fungi: a signature of undiscovered global diversity
(Phytotaxa 18)
127 pp.; 30 cm.
18 February 2011
ISBN 978-1-86977-649-7 (paperback)
ISBN 978-1-86977-650-3 (Online edition)
FIRST PUBLISHED IN 2011 BY
Magnolia Press
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other than private research use.
ISSN 1179-3155
(Print edition)
ISSN 1179-3163
(Online edition)
2 •
Phytotaxa 18 © 2011 Magnolia Press
LUMBSCH ET AL.
Full contact addresses and emails of 103 authors
1
Department of Botany, The Field Museum, 1400 South Lake Shore Drive, Chicago, Illinois 60605-2496, U.S.A.
Corresponding author, e-mail: tlumbsch@fieldmuseum.org. *Other e-mail addresses see footnote.
2
Botanical Museum, P. O. Box 7, FI-00014, Helsinki University, Finland.
3
Department of Ecology and Evolutionary Biology, University of California, Santa Cruz, California 95064, U.S.A.
4
Universidad Complutense de Madrid, Departamento de Biología Vegetal II, Plaza de Ramón y Cajal s/n, 28040 Madrid,
Spain.
5
ABL Herbarium, G. v. d. Veenstraat 107, NL-3762 XK Soest, The Netherlands.
6
Botanical Museum, Lund University, Östra Vallgatan 18, SE-223 61 Lund, Sweden.
7
Universidad Nacional Autónoma de México (UNAM), Instituto de Biología, Departamento de Botánica, Apdo. Postal 70233, Coyoacán 04360, México D.F., Mexico.
8
College of Natural Sciences, Saint Louis University, Baguio City, The Philippines.
9
Instituto de Botânica, Núcleo de Pesquisa em Micologia, Caixa Postal 3005, 01031-970, São Paulo, Brazil.
10
Licenciatura en Biología, Universidad Distrital Francisco José de Caldas, Cra. 4 No. 26B-54, Torre de Laboratorios,
Herbario, Bogotá, Colombia.
11
Stillinger Herbarium, University of Idaho, Moscow, Idaho 83843, U.S.A.
12
Biology Department, Faculty of Science, Ramkhamhaeng University, Hua Mak, Bangkok 10240, Thailand.
13
Klipperwerf 5, NL-2317 DX Leiden, The Netherlands.
14
Fundación Charles Darwin, Puerto Ayora, Santa Cruz, Galápagos, Ecuador; mailing address: Charles Darwin Foundation
for the Galápagos Islands, Av. 6 de Diciembre N 36-109 y Pasaje California, Post Box 17-01-3891, Quito, Ecuador.
15
Departamento de Biociências, Universidade Federal de Sergipe, CEP: 49.500-00, Itabaiana, Sergipe, Brazil.
16
Department of Biology, Faculty of Science, Anadolu University, 26470 Eskişehir, Turkey.
17
Laboratorio de Hongos, Instituto Nacional de Biodiversidad (INBio), Apdo. 22-3100, Santo Domingo de Heredia, Costa
Rica.
18
Conservatoire et Jardin botaniques, Case postale 60, CH-1292 Chambésy/Genève, Switzerland.
19
534 Fenton St., Lansing, Michigan 48910, U.S.A.
20
Royal Botanic Garden, Edinburgh, EH3 5LR, Scotland, U.K.
21
Department of Environmental Science and Policy, George Mason University, Fairfax, Virginia 22030-4444, U.S.A.
22
Conservation International Philippines, Number 6 Maalalahangin Street, Teachers Village, Diliman 1101, Quezon City, The
Philippines.
23
Research School of Chemistry, Building 33, Australian National University, Canberra, ACT 0200, Australia.
24
Tromsø University Museum, University of Tromsø, N-9037 Tromsø, Norway.
25
Department of Plant Biology, University of Minnesota, St. Paul, MN 55108, U.S.A.
26
Institute of Ecology and Botany, Hungarian Academy of Sciences, H-2163 Vácrátót, Hungary.
27
Instituto de Botánica del Nordeste (IBONE), Casilla del Correo 209, 3400 Corrientes, Argentina.
28
Institute for Integrated Natural Sciences, Department of Biology, University of Koblenz-Landau, Universitätstraße 1, D56070 Koblenz, Germany.
29
Landcare Research, Private Bag 1930, Dunedin 9054, New Zealand.
30
Department of Plant Biology (Botany Unit), Facultat de Biologia, Universitat de Barcelona, Av. 7 Diagonal 645, 08028
Barcelona, Spain.
31
Departament de Bioquímica i Biotecnologia (Àrea de Botànica), Facultat d'Enologia de Tarragona, Universitat Rovira i
Virgili, Marcellí Domingo s/n, 43007, Tarragona, Spain.
32
Herbarium, Department of Botany, University of British Columbia, Vancouver, British Columbia V6G 2B1, Canada;
mailing address: Edgewood Blue, Box 131, Clearwater, British Columbia V0E 1N0, Canada.
33
Institut für Pflanzensystematik, Universität Graz, Holteigasse 6, A-8010 Graz, Austria.
34
Fundación Instituto Botánico de Venezuela, División de Plantas no Vasculares, Sección Hongos y Líquenes, Ave. Salvador
Allende, Jardín Botánico de Caracas, Universidad Central de Venezuela, Apartado # 2156, Caracas 1010-A, Venezuela.
35
Lichenologisches Institut Neumarkt, Im Tal 12, D-92318 Neumarkt, Germany.
36
Tasmanian Herbarium, GPO Box 252-04, Hobart, Tasmania 7001, Australia.
37
Botany Department, East Africa Herbarium, National Museums of Kenya, Nairobi, Kenya.
38
The Herbarium, Deptartment of Botany & Plant Sciences, University of California, Riverside, California 92521, U.S.A.
39
OIKOS – Institute for Applied Ecology and Basic Research, Hartbergerstr. 40, 8200 Gleisdorf, Austria.
40
M. G. Kholodny Institute of Botany, Tereschenkivs'ka 2, 252601 Kiev-4, Ukraine.
41
Ossastrasse 6, D-12045 Berlin, Germany.
42
Department of Botany and Plant Pathology, Oregon State University, 2082 Cordley Hall, Corvallis, Oregon 97331-2902,
U.S.A.
ONE HUNDRED NEW SPECIES OF LICHENIZED FUNGI
Phytotaxa 18 © 2011 Magnolia Press
• 3
43
Instituto de Investigaciones en Biodiversidad y Medioambiente (INIBIOMA) – CONICET, Universidad del Comahue,
Quintral 1250, 8400 Sán Carlos de Bariloche, Río Negro, Argentina.
44
Institute of Applied Science, Faculty of Science and Technology, University of the South Pacific, Private Bag, Laucala
Campus, Suva, Fiji.
45
Committee on Evolutionary Biology, University of Chicago, 1025 E. 57th Street, Chicago, Illinois 60637, U.S.A.
46
Bergen Museum, DNS, P.O.Box 7803, 5020 Bergen, Norway.
47
Institute of Botany, Czech Academy of Science, CZ-252 43 Pruhonice & Department of Botany, Faculty of Natural Sciences,
Charles University, Benátská 2, CZ-128 01 Prague, Czech Republic.
48
Department of Biology and Natural Medicinal Mushroom Museum, Faculty of Science, Mahasarakham University
Khamrieng, Kantarawichai Maha Sarakham Province, 44150 Thailand.
49
Department of Botany and Molecular Evolution, Forschungsinstitut Senckenberg, Senckenberganlage 25, D-60325
Frankfurt am Main, Germany.
50
Department of Biological Sciences, University of Illinois-Chicago, 845 West Taylor Street (MC 066), Chicago, Illinois
60607, U.S.A.
51
Coordinador del Bosque Protector Río Guajalito, Mariana de Jesús E6-09 y Hungría, Quito, Ecuador.
52
Departamento de Biología. Universidad de Oriente. Patricio Lumumba s/n, CP. 90500 Cuba.
53
Fachbereich Organismische Biologie, Universität Salzburg, Hellbrunnerstrasse 34, 5020 Salzburg, Austria.
54
Smithsonian Tropical Research Institute, Box 0843-03092, Balboa, Ancon, Panama.
55
Biocenter Klein Flottbek and Botanical Garden, University of Hamburg, Ohnhorststrasse 18, D-22609 Hamburg, Germany.
56
Department of Environmental Science, The University, Bradford, Yorkshire, BD7 1DP, England, U.K.
57
Plant Taxonomy & Conservation Biology Unit, University of Liège, Sart Tilman B22, B-4000 Liège, Belgium
58
Biodiversity and Climate Research Centre (BiK-F), Goethe University and Senckenberg, Senckenberganlage 25, D-60325
Frankfurt, Germany.
59
Botanischer Garten & Botanisches Museum Berlin Dahlem, Königin-Luise-Strasse 6–8, D-14195 Berlin, Germany;
60
Department of Plant Science, University of Tabriz, 51666, Tabriz, Iran.
61
Department of Biology, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen Ø, Denmark.
62
Vrijheidslaan 27, NL-2806 KE Gouda, The Netherlands.
63
Laboratório de Botânica, Departamento de Biologia, Centro de Ciências Biológicas e da Saúde, Universidade Federal de
Mato Grosso do Sul, Cidade Universitária, Caixa Postal 549, CEP 79070-900, Campo Grande, Minas Gerais, Brazil.
64
King Mongkut' Institute Of Technology Ladkrabang, Chumphon Campus, 17/1 Moo 6, Chumcho, Patiew, Chumphon,
86160, Thailand.
65
Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen Province, 40002, Thailand.
66
Department of Ecology, Swedish University of Agricultural Sciences, P.O. Box 7044, SE-750 07 Uppsala, Sweden.
67
Botany Department, Natural History Museum, Cromwell Road, London, SW7 5BD, England, U.K.
68
Natural History Museum, University of Oslo, PO Box 1172 Blindern, N-0318 Oslo, Norway.
69
Fachbereich Organismische Biologie, Universität Salzburg, Hellbrunnerstrasse 34, 5020 Salzburg, Austria.
70
Lichenology Laboratory, National Botanical Research Institute, Rana Pratap Marg, P. B. No. 436, Lucknow-226001, India.
71
Arafura 16, NL-5691 JA Son, The Netherlands.
72
Department of Cryptogamic Botany, Swedish Museum of Natural History, P.O. Box 50007, SE-104 05 Stockholm, Sweden.
73
University of Wisconsin-Madison, Department of Botany, 430 Lincoln Dr., Madison, Wisconsin 53706, U.S.A.
74
Staatliches Museum für Naturkunde, Erbprinzenstr. 13, D-76133 Karlsruhe, Germany.
75
48 Wood Lane, Leeds LS6 2BA, England, U.K.
76
6606 Jocko Canyon Road, Arlee, MT 59821 USA
*E-mail addresses of authors (resp.): tlumbsch@fieldmuseum.org, teuvo.ahti@helsinki.fi, salter@biology.ucsc.edu,
g_amo@farm.ucm.es, andreaptroot@gmail.com, ulf.arup@botmus.lu.se, abarcenas@ibunam2.ibiologia.unam.mx,
pbawingan@slu.edu.ph, michel_benatti@yahoo.com.br, macualu@gmail.com, crbjork@gmail.com, kansri@ru.ac.th,
fbungartz@fcdarwin.org.ec, mscaceres@hotmail.com, mecandan@anadolu.edu.tr, jchaves@inbio.ac.crm, philippe.clerc@ville-ge.ch,
common@msu.edu, lichensEL@btopenworld.com, acrespo@farm.ucm.es, manudalforno@hotmail.com, pdivakar@farm.ucm.es,
mvduya@conservation.org, john.elix@anu.edu.au, arve.elvebakk@uit.no, fank0006@umn.edu, efarkas@botanika.hu,
itati_liq@yahoo.com.ar, efischer@uni-koblenz.de, gallowayd@xtra.co.nz, eb62@duke.edu, mireia.giralt@urv.cat,
tgoward@interchange.ubc.ca, martin.grube@uni-graz.at, josef.hafellner@uni-graz.at, jesus.hernandez@ucv.ve,
mahc@ibiologia.unam.mx, klaus.kalb@arcor.de, ingvar.karnefelt@botmus.lu.se, gintaras.kantvilas@tmag.tas.gov.au, killmann@unikoblenz.de, paulkirika@yahoo.com, knudsen@ucr.edu, harald.komposch@utanet.at, ksya_net@ukr.net, jlawrey@gmu.edu,
arminmangold@gmail.com, mpmarcelli@msn.com, bruce.mccune@science.oregonstate.edu, mmessuti@crub.uncoma.edu.ar,
andreamichlig@hotmail.com, mirandar_g@yahoo.com.mx, lbmoncada@udistrital.edu.co, naikatini_a@usp.ac.fj,
mpnelsen@gmail.com, dag.ovstedal@bm.uib.no, zpalice@yahoo.com, khwanruan.p@msu.ac.th, sparnmen@gmail.com,
sperezortega@farm.ucm.es, christian.printzen@senckenberg.de, rico@farm.ucm.es, erivasplata@fieldmuseum.org,
4 •
Phytotaxa 18 © 2011 Magnolia Press
LUMBSCH ET AL.
javierobayo@gmail.com, drl@cnt.uo.edu.cu, ulrike.carl@sbg.ac.at, salazarn@si.edu, sancholg@farm.ucm.es,
lucianaufsbio@hotmail.com, tamy_phn@hotmail.com, schultzm@botanik.uni-hamburg.de, m.r.d.seaward@bradford.ac.uk,
e.serusiaux@ulg.ac.be, imke.schmitt@senckenberg.de, h.sipman@bgbm.org, mohammad.sohrabi@helsinki.fi, ulriks@bio.ku.dk,
mzsogaard@snm.ku.dk, sparrius@biodiv.nl, adrianospielmann@yahoo.com.br, toby.spribille@uni-graz.at, sea_flora@hotmail.com,
achara@kku.ac.th, arne.thell@botmus.lu.se, goran.thor@ekol.slu.se, h.thues@nhm.ac.uk, einar.timdal@nhm.uio.no,
camille.truong@ville-ge.ch, roman.tuerk@sbg.ac.at, lumana@inbio.ac.cr, upretidk@rediffmail.com, pvdboom@kpnmail.nl,
mpvivas@farm.ucm.es, mats.wedin@nrm.se, swwolf@wisc.edu, volkmar.wirth@online.de, norawirtz@googlemail.com,
r.yahr@rbge.org.uk, kumeyesh@yahoo.com, frauke.ziemmeck@fcdarwin.org.ec, timothybwheeler@gmail.com,
rlucking@fieldmuseum.org.
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
The 100 New Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Acarospora flavisparsa V.J.Rico & Candan, sp. nov. (Fig. 2A–B) Mycobank MB 518279 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Acarospora janae K.Knudsen, sp. nov. (Fig. 2C) Mycobank MB 516796 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Aderkomyces thailandicus Papong, Boonpragob & Lücking, sp. nov. (Fig. 2D) Mycobank MB 517730. . . . . . . . . . . . . . . . . . 11
Amandinea maritima Giralt, van den Boom & Elix, sp. nov. (Fig. 2E–F) Mycobank MB 517731 . . . . . . . . . . . . . . . . . . . . . . . 13
Ampliotrema cocosense Lücking & Chaves, sp. nov. (Fig. 3A) Mycobank MB 517732 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Anomomorpha lecanorina Sipman, sp. nov. (Fig. 3B–C) Mycobank MB 517733 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Anomomorpha tuberculata Lücking, Umaña & Will-Wolf, sp. nov. (Fig. 3D) Mycobank MB 517734 . . . . . . . . . . . . . . . . . . . . 17
Aspicilia mansourii Sohrabi, sp. nov. (Fig. 3E–F) Mycobank MB 518502 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Bacidina sorediata Seaward & Lücking, sp. nov. (Fig. 4A) Mycobank MB 517735 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Badimia multiseptata Papong & Lücking, sp. nov. (Fig. 4B) Mycobank MB 517736 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Badimia vezdana Lücking, Farkas & Wirth, sp. nov. (Fig. 4C–D) Mycobank MB 517737 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Biatora epirotica Printzen & T.Sprib., sp. nov. (Fig. 4E–F) Mycobank MB 519464 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
Buellia sulphurica Bungartz & Aptroot, sp. nov. (Fig. 5A–B) Mycobank MB 517738 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Bunodophoron pinnatum Wedin, sp. nov. (Fig. 5C) Mycobank MB 517739 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Byssoloma spinulosum Sérus., sp. nov. (Fig. 5D) Mycobank MB 517740 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Calopadia cinereopruinosa Bungartz & Lücking, sp. nov. (Fig. 5E–F) Mycobank MB 517741 . . . . . . . . . . . . . . . . . . . . . . . . . 27
Calopadia editae Vĕzda ex Chaves & Lücking, sp. nov. (Fig. 6A) Mycobank MB 517742 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Caloplaca brownlieae S.Y.Kondr., Elix & Kärnefelt, sp. nov. (Fig. 6B) Mycobank MB 517743 . . . . . . . . . . . . . . . . . . . . . . . . 28
Caloplaca decipioides Arup, sp. nov. (Fig. 6C) Mycobank MB 517744 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Caloplaca digitaurea Søgaard, Søchting & Sancho, sp. nov. (Fig. 6D–E) Mycobank MB 517745 . . . . . . . . . . . . . . . . . . . . . . . 31
Caloplaca magnussoniana S.Y.Kondr., Kärnefelt & A.Thell, sp. nov. (Fig. 6F) Mycobank MB 517746 . . . . . . . . . . . . . . . . . . 32
Caloplaca mereschkowskiana S.Y.Kondr. & Kärnefelt, sp. nov. (Fig. 7A) Mycobank MB 517747 . . . . . . . . . . . . . . . . . . . . . . . 33
Caloplaca yorkensis S.Y.Kondr. & Kärnefelt, sp. nov. (Fig. 7B) Mycobank MB 517748 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
Calvitimela uniseptata G.Thor, sp. nov. (Fig. 7C) Mycobank 517749 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Chapsa microspora Kalb, sp. nov. (Fig. 7D) Mycobank MB 517750 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Chapsa psoromica M.Cáceres, Santos de Jesus & Santos Vieira, sp. nov. (Fig. 7E) Mycobank MB 517751 . . . . . . . . . . . . . . . 37
Chapsa rubropulveracea Hale ex Mangold, Lücking & Lumbsch, sp. nov. (Fig. 7F) Mycobank MB 517753 . . . . . . . . . . . . . . 38
Chapsa thallotrema Lücking & N.Salazar, sp. nov. (Fig. 8A–B) Mycobank MB 517753 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Chiodecton pustuliferum Aptroot, sp. nov. (Fig. 8C–D) Mycobank MB 517754 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Cladonia mongkolsukii Parnmen & Ahti, sp. nov. (Fig. 8E–F) Mycobank MB 517755 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Clypeopyrenis porinoides Komposch, J.E.Hern. & Rosabal, sp. nov. (Fig. 9A) Mycobank MB 517756 . . . . . . . . . . . . . . . . . . . 43
Coccocarpia delicatula Bungartz, Ziemmeck & Lücking, sp. nov. (Fig. 9B) Mycobank MB 517757 . . . . . . . . . . . . . . . . . . . . 45
Coenogonium flammeum L.I.Ferraro, Michlig & Lücking, sp. nov. (Fig. 9C–D) Mycobank MB 517758 . . . . . . . . . . . . . . . . . . 45
Cresponea ancistrosporelloides Sparrius & Sipman, sp. nov. (Fig. 9E–F) Mycobank MB 517759 . . . . . . . . . . . . . . . . . . . . . . . 46
Crocynia microphyllina Aptroot, sp. nov. (Fig. 10A–B) Mycobank MB 517760 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Dictyonema hernandezii Lücking, Lawrey & Dal-Forno, sp. nov. (Fig. 10C) Mycobank MB 517761 . . . . . . . . . . . . . . . . . . . . 47
Dictyonema hirsutum Moncada & Lücking, sp. nov. (Fig. 10D) Mycobank MB 517762 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Diorygma microsporum M.Cáceres & Lücking, sp. nov. (Fig. 10E) Mycobank MB 517763 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . 0
Diorygma sticticum Sutjaritturakan, Kalb & Lücking, sp. nov. (Fig. 10F) Mycobank MB 517764 . . . . . . . . . . . . . . . . . . . . . . . 50
Echinoplaca pernambucensis Øvstedal & Elix, sp. nov. (Fig. 11A) Mycobank MB 517765 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Echinoplaca schizidiifera J.E.Hern. & Lücking, sp. nov. (Fig. 11B–C) Mycobank MB 517766 . . . . . . . . . . . . . . . . . . . . . . . . . 52
Eremithallus marusae R.Miranda, Gaya & Lücking, sp. nov. (Fig. 11D) Mycobank MB 517767 . . . . . . . . . . . . . . . . . . . . . . . . 52
Everniastrum constictovexans Sipman, sp. nov. (Fig. 11E) Mycobank MB 517768 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
ONE HUNDRED NEW SPECIES OF LICHENIZED FUNGI
Phytotaxa 18 © 2011 Magnolia Press
• 5
Fellhanera borbonica Sérus., van den Boom & Brand, sp. nov. (Fig. 11F) Mycobank MB 517769 . . . . . . . . . . . . . . . . . . . . . . 55
Fibrillithecis sprucei Mangold, Lücking & Lumbsch, sp. nov. (Fig. 12A) Mycobank MB 517770 . . . . . . . . . . . . . . . . . . . . . . . 56
Fissurina astroisidiata Herrera-Campos & Lücking, sp. nov. (Fig. 12B–C) Mycobank MB 517771 . . . . . . . . . . . . . . . . . . . . . 57
Fissurina nigrolabiata Rivas Plata, Bawingan & Lücking, sp. nov. (Fig. 12D) Mycobank MB 517772 . . . . . . . . . . . . . . . . . . . 58
Fissurina subcomparimuralis Common & Lücking, sp. nov. (Fig. 12E) Mycobank MB 517773 . . . . . . . . . . . . . . . . . . . . . . . . 59
Graphis caribica Lücking, sp. nov. (Fig. 12F) Mycobank MB 517774 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
Graphis cerradensis Marcelli, Benatti & Lücking, sp. nov. (Fig. 13A–B) Mycobank MB 517775 . . . . . . . . . . . . . . . . . . . . . . . 60
Graphis itatiaiensis Nelsen, Lücking & Spielmann, sp. nov. (Fig. 13C) Mycobank MB 517776 . . . . . . . . . . . . . . . . . . . . . . . . 62
Graphis marusae B.Peña & Lücking, sp. nov. (Fig. 13D) Mycobank MB 517777 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Gyalideopsis chicaque Moncada & Lücking, sp. nov. (Fig. 13E) Mycobank MB 517778 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Gyrotrema papillatum Lücking, sp. nov. (Fig. 13F) Mycobank MB 517779 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Harpidium gavilaniae Amo, Pérez-Ortega & A. Crespo, sp. nov. (Fig. 14A–B) Mycobank MB 517780 . . . . . . . . . . . . . . . . . . 65
Hypogymnia amplexa Goward, Björk & Wheeler, sp. nov. (Fig. 14C) Mycobank MB 517781 . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Hypotrachyna guatemalensis Elix & van den Boom, sp. nov. (Fig. 14D) Mycobank MB 517782 . . . . . . . . . . . . . . . . . . . . . . . 68
Hypotrachyna indica Divakar, Lumbsch, Upreti & A.Crespo, sp. nov. (Fig. 14E–F) Mycobank MB 518238 . . . . . . . . . . . . . . 68
Hypotrachyna lueckingii Sipman, sp. nov. (Fig. 15A) Mycobank MB 517783 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Hypotrachyna paracitrella Sipman & Palice, sp. nov. (Fig. 15B) Mycobank MB 517784 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Hypotrachyna paraphyscioides Sipman, sp. nov. (Fig. 15C) Mycobank MB 517785 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
Hypotrachyna parasinuosa Sipman & Palice, sp. nov. (Fig. 15D) Mycobank MB 517786 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Icmadophila eucalypti Kantvilas, sp. nov. (Fig. 15E–F) Mycobank MB 517787 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Krogia microphylla Timdal, sp. nov. (Fig. 16A–B) Mycobank MB 517788 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
Lecanora mugambii Kirika, I.Schmitt, Fankhauser & Lumbsch, sp. nov. (Fig. 16C–D) Mycobank MB 517789 . . . . . . . . . . . . 76
Lecanora printzenii Pérez-Ortega, Vivas & Hafellner, sp. nov. (Fig. 16E) Mycobank MB 517790 . . . . . . . . . . . . . . . . . . . . . . . 77
Lecanora xanthoplumosella Lumbsch & Elix, sp. nov. (Fig. 16F) Mycobank MB 517791 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Lecidea lygommella Elix, sp. nov. (Fig. 17A) Mycobank MB 517792 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Lecidella greenii U.Ruprecht & Türk, sp. nov. (Fig. 17B–C) Mycobank MB 517793 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
Lempholemma corticola M.Schultz & T.Sprib., sp. nov. (Fig. 17D–E) Mycobank MB 517794 . . . . . . . . . . . . . . . . . . . . . . . . . 81
Lepraria sekikaica Elix, sp. nov. (Fig. 17F) Mycobank MB 517795 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
Lobariella sipmanii Moncada, Betancourt & Lücking, sp. nov. (Fig. 18A–B) Mycobank MB 519082 . . . . . . . . . . . . . . . . . . . . 83
Megalospora austropacifica Lumbsch, Naikatini & Lücking, sp. nov. (Fig. 18C) Mycobank MB 517796 . . . . . . . . . . . . . . . . . 84
Megalospora galapagoensis Bungartz, Ziemmeck & Lücking, sp. nov. (Fig. 18D–E) Mycobank MB 517797 . . . . . . . . . . . . . 84
Menegazzia endocrocea Kantvilas, sp. nov. (Fig. 18F) Mycobank MB 517798 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
Myriotrema endoflavescens Hale ex Lücking, sp. nov. (Fig. 19A–B) Mycobank MB 517799 . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
Ocellularia albobullata Lücking, Sipman & Grube, sp. nov. (Fig. 19C) Mycobank MB 517800 . . . . . . . . . . . . . . . . . . . . . . . . 88
Ocellularia vizcayensis Rivas Plata, Duya & Lücking, sp. nov. (Fig. 19D) Mycobank MB 517801 . . . . . . . . . . . . . . . . . . . . . . 89
Ochrolechia insularis Kantvilas & Elix, sp. nov. (Fig. 19E–F) Mycobank MB 517802 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Opegrapha viridipruinosa B.J.Coppins & R.Yahr, sp. nov. (Fig. 20A–B) Mycobank MB 517715 . . . . . . . . . . . . . . . . . . . . . . . 91
Pannaria phyllidiata Elvebakk, sp. nov. (Fig. 20C) Mycobank MB 517803 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Parmelia asiatica A.Crespo & Divakar, sp. nov. (Fig. 20D–E) Mycobank MB 518288 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Pertusaria conspersa Messuti, sp. nov. (Fig. 20F) Mycobank MB 517804 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Phlyctis psoromica Elix & Kantvilas, sp. nov. (Fig. 21A) Mycobank MB 517805 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Placopsis imshaugii D.J.Galloway, sp. nov. (Fig. 21B) Mycobank MB 516034 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Platismatia wheeleri Goward, Altermann & Björk, sp. nov. (Fig. 21C) Mycobank MB 517806 . . . . . . . . . . . . . . . . . . . . . . . . 100
Porina huainamdungensis Papong, Thammathaworn & Lücking, sp. nov. (Fig. 21D) Mycobank MB 517807 . . . . . . . . . . . . 101
Ramalina hyrcana Sipman, sp. nov. (Fig. 21E–F) Mycobank MB 517808 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Ramalina stoffersii Sipman, sp. nov. (Fig. 22A) Mycobank MB 517809 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
Relicina colombiana Elix & Sipman, sp. nov. (Fig. 22B) Mycobank MB 517810 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
Rhizocarpon diploschistidina McCune, sp. nov. (Fig. 22C–D) Mycobank MB 516796 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Sagenidiopsis isidiata G.Thor, Elix, Lücking & Sipman, sp. nov. (Fig. 22E–F) Mycobank MB 517811 . . . . . . . . . . . . . . . . . . 106
Sticta venosa Lücking, Moncada & Robayo, sp. nov. (Fig. 23A–C) Mycobank MB 517812 . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
Tapellaria albomarginata Lücking, sp. nov. (Fig. 23D–E) Mycobank MB 517813 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
Thelotrema fijiense Lumbsch, Lücking & Naikatini, sp. nov. (Fig. 23F) Mycobank MB 517814 . . . . . . . . . . . . . . . . . . . . . . . 109
Tricharia nigriuncinata Yeshitela, Eb.Fischer, Killmann & Sérus., sp. nov. (Fig. 24A–B) Mycobank MB 517815 . . . . . . . . . 110
Usnea galapagona Truong & P.Clerc, sp. nov. (Fig. 24C–F) Mycobank MB 518904 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
Usnea pallidocarpa Wirtz & Lumbsch, sp. nov. (Fig. 25A–B) Mycobank MB 517816 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
Verrucaria rhizicola Aptroot & Thüs, sp. nov. (Fig. 25C–D) Mycobank MB 517817 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
Xanthomendoza rosmarieae S.Y.Kondr. & Kärnefelt, sp. nov. (Fig. 25E–F) Mycobank MB 517818 . . . . . . . . . . . . . . . . . . . . 115
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
6 •
Phytotaxa 18 © 2011 Magnolia Press
LUMBSCH ET AL.
Abstract
The number of undescribed species of lichenized fungi has been estimated at roughly 10,000. Describing and cataloging
these would take the existing number of taxonomists several decades; however, the support for taxonomy is in decline
worldwide. In this paper we emphasize the dire need for taxonomic expertise in lichenology. We bring together 103
colleagues from institutions worldwide to describe a total of 100 new species of lichenized fungi, representing a wide
taxonomic and geographic range. The newly described species are: Acarospora flavisparsa, A. janae, Aderkomyces
thailandicus, Amandinea maritima, Ampliotrema cocosense, Anomomorpha lecanorina, A. tuberculata, Aspicilia
mansourii, Bacidina sorediata, Badimia multiseptata, B. vezdana, Biatora epirotica, Buellia sulphurica, Bunodophoron
pinnatum, Byssoloma spinulosum, Calopadia cinereopruinosa, C. editae, Caloplaca brownlieae, C. decipioides, C.
digitaurea, C. magnussoniana, C. mereschkowskiana, C. yorkensis, Calvitimela uniseptata, Chapsa microspora, C.
psoromica, C. rubropulveracea, C. thallotrema, Chiodecton pustuliferum, Cladonia mongkolsukii, Clypeopyrenis
porinoides, Coccocarpia delicatula, Coenogonium flammeum, Cresponea ancistrosporelloides, Crocynia microphyllina,
Dictyonema hernandezii, D. hirsutum, Diorygma microsporum, D. sticticum, Echinoplaca pernambucensis, E.
schizidiifera, Eremithallus marusae, Everniastrum constictovexans, Fellhanera borbonica, Fibrillithecis sprucei,
Fissurina astroisidiata, F. nigrolabiata, F. subcomparimuralis, Graphis caribica, G. cerradensis, G. itatiaiensis, G.
marusa, Gyalideopsis chicaque, Gyrotrema papillatum, Harpidium gavilaniae, Hypogymnia amplexa, Hypotrachyna
guatemalensis, H. indica, H. lueckingii, H. paracitrella, H. paraphyscioides, H. parasinuosa, Icmadophila eucalypti,
Krogia microphylla, Lecanora mugambii, L. printzenii, L. xanthoplumosella, Lecidea lygommella, Lecidella greenii,
Lempholemma corticola, Lepraria sekikaica, Lobariella sipmanii, Megalospora austropacifica, M. galapagoensis,
Menegazzia endocrocea, Myriotrema endoflavescens, Ocellularia albobullata, O. vizcayensis, Ochrolechia insularis,
Opegrapha viridipruinosa, Pannaria phyllidiata, Parmelia asiatica, Pertusaria conspersa, Phlyctis psoromica,
Placopsis imshaugii, Platismatia wheeleri, Porina huainamdungensis, Ramalina hyrcana, R. stoffersii, Relicina
colombiana, Rhizocarpon diploschistidina, Sticta venosa, Sagenidiopsis isidiata, Tapellaria albomarginata, Thelotrema
fijiense, Tricharia nigriuncinata, Usnea galapagona, U. pallidocarpa, Verrucaria rhizicola, and Xanthomendoza
rosmarieae. In addition, three new combinations are proposed: Fibrillithecis dehiscens, Lobariella botryoides, and
Lobariella pallida.
Key words: Darwin declaration, Encyclopedia of Life, Global Taxonomy Initiative, lichens, taxonomy, taxonomic
impediment
Introduction
The number of species inhabiting our planet is estimated to be between 2–10 and 30–100 million (May 1988,
Curtis et al. 2002, Rosenzweig et al. 2003). About 2.2 million are recorded, including 400,000 plants, 100,000
fungi (including 17,500 lichens), 1.5 million animals, 200,000 protists, and 10,000 bacteria (Prance et al.
2000, Govaerts 2001, Pitman & Jørgensen 2002, Scotland & Wortley 2003, Schloss et al. 2004, Williamson &
Day 2007, Feuerer & Hawksworth 2007, Kirk et al. 2008). The total number of fungi is estimated at 700,000
to 1.5 million species (Hawksworth 1991, 2001, Mueller & Schmit 2007). The majority of the undescribed
fungal species are expected in poorly studied areas, such as tropical forests or in under-explored habitats, for
example living on or in insects, plants or lichens (Hawksworth & Rossman 1997, Frohlich & Hyde 1999,
Taylor et al. 2000a, Sipman & Aptroot 2001, Lawrey & Diederich 2003, Arnold & Lutzoni 2007). There is
also a growing body of evidence suggesting that the approach to current species recognition, which is based
largely on morphology and chemistry, significantly underestimates the number of species. Phylogenetic
studies indicate that numerous distinct lineages may be hidden under a single species name (Roy et al. 1998,
Grube & Kroken 2000, Kroken & Taylor 2001, Koufopanou et al. 2001, Peterson et al. 2001, Cruse et al.
2002, Molina et al. 2004, Douhan & Rizzo 2005, Pringle et al. 2005, Geml et al. 2006, Kauserud et al. 2006,
Matute et al. 2006, Arguello et al. 2007b, Giraud et al. 2008, Wirtz et al. 2008, Baloch & Grube 2009).
Even with a conservative estimate of ten million existing species, including perhaps around one million
fungi, the number of eight million remaining undiscovered species, including 900,000 fungi, is mindboggling. Assuming that modern taxonomic revisions were produced mainly in the past 50 years, it would
ONE HUNDRED NEW SPECIES OF LICHENIZED FUNGI
Phytotaxa 18 © 2011 Magnolia Press
• 7
take the same number of taxonomists another 200 years to describe the remaining undiscovered species
(Hebert et al. 2003, Lücking 2009b). This appears an almost impossible task, especially given the rapid and
dramatic decline in numbers of employed taxonomists in the past 20 years (Butler et al. 1998, Suarez &
Tsutsui 2004, Graham 2005). In light of these huge gaps in our knowledge, the shortage of trained
taxonomists and museum curators, the taxonomic impediment (http://flyaqis.mov.vic.gov.au/chaec/
taximp.html) now has a significant negative impact on our ability to manage and utilize our biological
diversity. In addition, the advent of molecular phylogeny has created a misleading impression that taxonomy
is an outdated discipline. The Barcode of Life initiative (http://www.barcoding.si.edu) is sometimes seen as a
way to replace taxonomists by handheld DNA scanners (Hebert et al. 2003, Stoeckle 2003). Identification of
species using DNA signature sequences is indeed a fascinating concept, but for this to become reality, a huge
reference database is required that contains the signature sequences of all species (Lücking 2009b). This is an
enormous task that encompasses not only sequencing itself, but also requires targeted collecting, taxonomic
revision and assembly of reference voucher collections and with it a large number of well-trained taxonomists
who are not only specialists in their respective groups but proficient in a variety of methods, including
nomenclature, morphology, and molecular phylogeny, and are capable of attracting and training students.
The task of cataloging our planet's diversity is even more imperative considering estimates that 15–50%
of the species will go extinct in the next fifty years due to increasing pressure from a growing world
population (Hughes et al. 1997, Woodruff 2001, Thomas et al. 2004, Pimm et al. 2005, Ehrlich & Pringle
2008). We will not only lose biodiversity of unknown ecological and economic importance, we will also alter
the dynamic equilibrium of our global ecosystems, something that has happened repeatedly during the long
history of our planet, but never within a timeframe of a few hundred years (Ehrlich & Pringle 2008). Global
initiatives such as the United Nations Convention on Biological Diversity CBD (http://www.cbd.int), the
Darwin Declaration (http://www.environment.gov.au/biodiversity/abrs/publications/other/darwin/index.html),
and the Global Taxonomy Initiative GTI (http://www.cbd.int/gti) are attempting to reverse this trend by
publicizing the need for increased efforts in cataloging biodiversity before it vanishes, and the need for shifts
in human behavior to protect our ecosystems. However, in reality not much is happening to support a
sustainable level of well-trained taxonomists; in fact we are in a paradoxical situation where politics has
acknowledged a problem that science refuses to deal with appropriately: without proper taxonomic expertise,
the sound management of biodiversity is not possible (House Of Lords Report 1991, Aylward et al. 1993,
Janzen 1993, Systematics Agenda 2000 1994, Graham 2005). Taxonomists therefore must actively make a
strategic change to work effectively on the huge task at hand (Janzen 1993, Lücking 2009b). The degree to
which collaborative research can be effective in accumulating a huge amount of knowledge in a short time is
demonstrated by the National Center for Biotechnology Information, commonly known as GenBank (http://
www.ncbi.nlm.nih.gov/guide). Within two decades, it has become the most important resource in biosciences
(McEntyre & Lipman 2001). Further, the NSF-funded Tree of Life projects, including AFTOL (Assembling
the Fungal Tree of Life; http://aftol.org), which began in 2002 and is currently in its second phase until 2011,
have been extremely effective in enhancing global collaboration to elucidate the phylogeny of the fungal
kingdom (Lutzoni et al. 2004, James et al. 2006, Matheny et al. 2006, Miadlikowska et al. 2006, Spatafora et
al. 2006, Schoch et al. 2006, 2009, Hibbett et al. 2007).
Traditionally, systematic papers and monographic revisions have just one or a few authors, but the
aforementioned collaborative initiatives have initiated a mind-shift with a multi-authored, community-wide
classification for the fungal kingdom (Hibbett et al. 2007). In addition, the mycological community has
several, regularly updated fundamental resources for species and higher level classification and nomenclature
available, such as Myconet (http://www.fieldmuseum.org/myconet), Index Fungorum (http://
www.indexfungorum.org) and other CABI Bioscience databases, and the Dictionary of the Fungi (Kirk et al.
2008). The organizing authors of the present paper, H. T. Lumbsch and R. Lücking, extended this successful
collaborative concept to promote and organize a community-wide monographic revision of the lichenized
fungal family Graphidaceae (http://www.nsf.gov/awardsearch/showAward.do?AwardNumber=1025861).
This concept has been labelled ATM – Assembling a Taxonomic Monograph. In the present paper, we
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Phytotaxa 18 © 2011 Magnolia Press
LUMBSCH ET AL.
elucidate this idea by bringing together 102 colleagues from 35 countries to describe 100 new species of
lichenized fungi, representing a wide taxonomic and geographic range covering 33 families of Ascomycota
and Basidiomycota and 37 countries around the globe (Fig. 1).
FIGURE 1. Map showing origin of new species (red dots) and geographic affiliation of contributing authors (blue dots).
The 100 New Species
The new species are described below in alphabetical order. They represent a total of 70 genera in 33 families,
15 orders, five classes in the Ascomycota and one genus, family, and order in the Basidiomycota. The
individual taxon entries are standardized as far as possible but reflect in style and content the diversity of
participating authors.
Acarospora flavisparsa V.J.Rico & Candan, sp. nov. (Fig. 2A–B) Mycobank MB 518279
Sicut Acarospora microcarpa sed cum Protoparmelia montagnei consociata, squamulis et ascomatibus majoribus,
squamulis latere carbonato nigro, paraphysibus crassibus, ascosporis elongate-cylindraceis et epanorinam
continente differt.—http://www.eol.org/pages/Acarospora flavisparsa
Type:—SPAIN. Madrid: Sán Lorenzo de El Escorial, near Silla de Felipe II, 40º 35’N, 04º 09’W, 1080 m,
associated with Protoparmelia montagnei on granite, July 1986, Florido & Rico 1133/1 (holotype MAF-Lich.
16463).
Thallus areolate to squamulose (subsquamulose), forming dispersed patches of one to few areoles
associated with thalli of, among areoles or overgrowing it, Protoparmelia montagnei (Fr.) Poelt & Nimis and
occasionally with nearby Dimelaena oreina (Ach.) Norman. Areoles irregular, rounded to angular by
pressure, sometimes lobulate, 0.5–3.5 mm in diameter, up to 2.25 mm thick, commonly forming a short stipe
and becoming squamulose; upper surface yellow to pale yellow or greenish yellow, usually dull, convex to
concave, smooth to rough, sometimes fissured; lateral/lower surface ± smooth, yellow-brown to dark brown
or carbonized black. Upper cortex paraplectenchymatous, (35–)40–125 µm thick depending on development
of areoles, cells usually indistinct, ± globular, in several layers, upper layers brown-yellow pigmented, lower
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layers hyaline; layers containing granules of various sizes; syncortex lacking or up to 25 µm; lateral/lower
cortex continuous with upper cortex, but with brown or dark brown to commonly carbonized black pigmented
upper layers. Algal layer continuous, up to 200 µ m thick, photobiont chlorococcoid, cells 6–15 µ m in
diameter. Medulla hyaline to dirty white, continuous with broad to forming stipe attaching tissue. Apothecia
immersed, aspicilioid, up to 0.7 mm in diameter, 1–8 per areole, sometimes confluent, punctiform to round or
irregular; disk brown to red, concave to ± plane, rough. Proper excipulum cupular, surrounded by algae, ±
prosoplechtenchymatous, 12–30 µm wide, hyaline to yellowish. Hymenium hyaline to yellowish, 125–250(–
300) µm tall; paraphyses 1.5–3 µm wide at the base, up to 4.5 µm at slightly pigmented apices; epihymenium
brownish-yellow, coherent, with yellowish granules, up to 20 µm tall. Hypothecium, including subhymenium,
hyaline to slightly yellowish, up to 150 µm thick. Asci clavate, 70–130 × 18–30 µm, more than 100-spored,
outer wall K/I + blue, tholus K/I–. Ascospores hyaline, simple, elongate to cylindrical, 4–6.01–7.5 × 1.5–
2.78–3.5 µm, length/width ratio 1.66–3, l/w mean value 2.21 (n = 150). Pycnidia immersed, punctiform, ±
spherical, up to 210 µm in diameter, ostiole inconspicuous or yellow to red-brown, wall hyaline to yellowish,
surrounded by algae. Conidia hyaline, bacilliform, 2.5–5 × 1–2 µm. Spot tests in cortex and medulla: K–, C–,
KC–, P–, I–; thallus: UV ± orange. Secondary chemistry: epanorin (major) with or without rhizocarpic acid
(traces) (8 TLC, 3 HPLC).
Distribution and habitat:—Currently, Acarospora flavisparsa is known from continental mediterranean
areas in Portugal (Tras os Montes) and Spain (Madrid, València and Zamora provinces). The species occurs
on acid rocks in inland areas, at 550–1480 m, on exposed ± vertical rock walls from continental dry to subhumid places, associated with Protoparmelia montagnei and sometimes with Dimelaena oreina.
Etymology:—The epithet refers to the scattered yellow squamules growing among other lichens.
Additional specimens examined (paratypes):—PORTUGAL. Tras os Montes e Alto Douro: Bragança,
IP4 road to the south, Sabor river valley, 3 km to Lagoa, Rico 3649/2 (MAF). SPAIN. Zamora: Pereruela, Las
Enillas, September 1998, Crespo & Blanco s.n. (MAF). Madrid: between El Berrueco and Berzosa del
Lozoya, El Villar dam, Barreno & Rico 275/1, 276/3 (MAF). La Cabrera, Sierra de La Cabrera, El Mojón
summit, Rico & Florido 380/2, 385/1, 434/2, 451/2, 452/1 (MAF). Manzanares el Real, La Pedriza, Collado
de Valdealcones, Rico & Barreno 589/4, 590/1, 621/3, 632/1 (MAF); ibid., El Chivato stream, Rico &
Barreno 477/3, 489/2 (MAF). Lozoya, El Chaparral, Rico & Manrique 770/1 (MAF). Collado-Mediano,
Cerro del Castillo, Rico 863/2, 901/6, 903/3 (MAF). Robledillo de la Jara, Collado de Fragüela, Rico &
Manrique 1093/1 (MAF). Robledo de Chavela, El Almojón summit, Rico 805/1, 807/3 (MAF). Zarzalejo,
December 1988, Valladares s.n. (MAF). València: Serra, Font del Berro, December 1990, Calatayud s.n.
(VAB). Castelló: Alfondeguilla, Nevera summit, June 1989, Calatayud s.n. (VAB).
The new taxon falls into the group of mediterranean Acarospora species (Clauzade & Roux, 1981) with
yellow, non-effigurate margin, silicolous thalli, associated with other lichens, brown to red apothecia and K–,
C–, white medulla. Acarospora flavisparsa is probably closest to A. microcarpa (Nyl.) Wedd. (Magnusson
1929, 1936, Poelt & Vĕzda 1977, Clauzade & Roux 1981, Egea & Llimona 1981), apparently a juvenile or
specific parasite on silicolous Diploschistes actinostomus (Ach.) Zahlbr. also growing on nearby lichens
(Poelt 1958, Clauzade & Roux 1981, 1985, Sipman & Rausch 1999, Roux et al. 2006). Furthermore, A.
microcarpa differs in having: areoles never lobulate, squamulose or shortly stalked and with yellowish to pale
lateral surface, smaller areoles and apothecia (up to 2 and 0.4 mm respectively), thinner cortex, algal layer,
hymenium and hypothecium (up to 70, 100, 140 and 35 µm respectively), narrow paraphyses at the base (up
to 2 µm), ellipsoidal ascospores [3–6 × 2.5–4 µm, length/width ratio 1.28–1.85, l/w mean value 1.6 (n = 80)],
rhizocarpic acid as major secondary substance (5 TLC, 8 HPLC; Leuckert & Buschardt 1978) and occurring
below 700 m elevation in France, Greece, Italy, Morocco and Spain, with optimum in the ThermoMediterranean belt (Egea & Rowe 1987, Nimis & Poelt 1987). Although A. flavisparsa grows primarily
dispersed within the areoles of P. montagnei and D. oreina, it does not appear to be an obligatory or hostspecific juvenile parasite, but a close association with those lichens is assumed. On the contrary, in some
specimens of A. microcarpa the thallus was observed growing endokapylically at first, becoming successively
epikapylic and independent and forming little thalli up to 2 cm diameter on D. actinostomus [Follmann, Lich.
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Exs. Sel. Cass. 182 (MAF)]. Acarospora heufleriana Körb., frequently invades other lichens but differs in its
ellipsoid to subglobose ascospores, rhizocarpic with or without norstictic acid as major substances, and ±
continuous thalli often with pruina (Clauzade & Roux 1981, Knudsen 2007). Acarospora charidema
(Clemente ex Colmeiro) Llimona is a thermophilic pioneer, not a parasitic lichen, with robust convex
squamules up to 4 mm wide, apothecia up to 1.5 mm wide and with a hymenium up to 140 µm tall (Clauzade
& Roux 1981, Egea & Rowe 1987). Acarospora epithallina H.Magn. is also similar to A. flavisparsa, but
grows as a juvenile parasite on Acarospora hilaris (Dufour) Arnold, forming up to 5 mm wide convex
squamules, ± globose ascospores and producing rhizocarpic and gyrophoric acids (Leuckert & Buschardt
1978, Clauzade & Roux 1981).
Acarospora janae K.Knudsen, sp. nov. (Fig. 2C) Mycobank MB 516796
Sicut Acarospora obpallens sed differt thallo striato.—http://www.eol.org/pages/Acarospora janae
Type:—U.S.A. New Mexico: Sán Miguel County, Las Vegas, 'aux fous', on sandstone, January 1927, Bro.
Arsène Brouard 19568 (holotype FH).
Thallus of dispersed epruinose areoles, dark brown to light brown, with a single immersed apothecium, at
first punctiform, becoming dilated and the areole rarely lobulate. Lacking faveoles but upper surface often
with fine striations. Lacking a stipe, broadly attached, lower surface white and ecorticate. Disc epruinose,
same colour or darker than the thallus, rough, sometimes with umbos and ridges, these disappearing as disc
fully dilates. Algal layer continuous. Exciple and hypothecium indistinct. Subhymenium 30–40 µ m high.
Hymenium ca. 100 µm high, paraphyses 2 µm in diameter. Asci clavate, ca. 60 × 20 µm. Ascospores about
100 per ascus, 3–4 × 2 µm. Secondary chemistry: gyrophoric acid.
Distribution and habitat:—On siliceous rock. Currently known from historical collections from Las
Vegas, New Mexico and a modern collection by Gary B. Perlmutter from North Carolina.
Etymology:—The species is named in honor of my colleague, frequent co-author and fiancé, Jana
Kocourková.
Additional specimens examined (paratypes):—U.S.A. New Mexico: Sán Miguel County, Las Vegas,
Bro. Arsène Brouard 19936 (FH). North Carolina: Wake County, Marks Creek county conservation area,
Perlmutter 2216 (NCU).
Acarospora is a large, cosmopolitan genus with at least 45 species known in southwestern North America
(Knudsen 2007, 2008, Knudsen & Morse 2009). The genus is characterized by polyspory (50–200 ascospores
per ascus), simple hyaline ascospores, aspicilioid or pseudo-lecanorine apothecia, and bitunicate but nonfissitunicate asci with a non-amyloid tholus. The new species is distinguished by its epruinose and finely
striated areoles, each with a single immersed and eventually dilated apothecium, and containing gyrophoric
acid. Though the author was aware of the taxon for some time, it was not found during research on the genus
in the greater Sonoran desert region (Knudsen 2007). It is expected to be at least infrequent at middle
elevations on sandstone, possibly from Utah and the Colorado Plateau south into non-Sonoran Mexico and
east to North Carolina. Acarospora fuscescens H.Magn. (Knudsen 2008) is similar, but has larger, verruciform
areoles with punctiform apothecia. When its apothecia are fully dilated, A. janae resembles A. obpallens (Nyl.
ex Hasse) Zahlbr. (Knudsen 2007), but that species is persistently faveolate with small pits, especially on the
thalline rim around the disc.
Aderkomyces thailandicus Papong, Boonpragob & Lücking, sp. nov. (Fig. 2D) Mycobank MB 517730
Sicut Aderkomyces gomezii sed thallo verrucoso et ascosporis minoribus differt.—http://www.eol.org/pages/
Aderkomyces thailandicus
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FIGURE 2. A–B. Acarospora flavisparsa (holotype). A. Habit growing among areoles of Protoparmelia montagnei
(photograph V. J. Rico). B. Detail of squamules with apothecia, showing dark lateral surface (photograph V. J. Rico). C.
Acarospora janae (holotype), habit (photograph M. Schmull). D. Aderkomyces thailandicus (holotype), habit
(photograph K. Papong). E. Amandinea maritima (van den Boom 19164), general habit showing the crustose, creamy
thallus; the young apothecia, erumpent or lecideine with pseudothalline margins; and the mature apothecia, typically
lecideine. F. Amandinea maritima (holotype), ascospore ontogeny and ascospore variability. Scale in A, C, D = 1 mm, in
B, E = 0.5 mm, in F = 10 µm.
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Type:—THAILAND. Chiang Mai Province: Huai Nam Dung National Park, 19°18’N, 98°36’E, 1368 m,
lower montane rainforest, forest margin, February 2006, Papong 2654 (holotype KKU).
Thallus epiphyllous, crustose, verrucose, greenish grey, 10–15 µm thick, with white, long sterile setae 2–
2.5 mm long. Apothecia at first immersed-erumpent with thin marginal lobules but finally broadly sessile to
adnate, with slightly prominent margin, light grey-brown, sometimes with orange tinge, 0.2–0.5 mm diameter.
Excipulum composed of radiating hyphae embedded in a gelatinous matrix, colourless, 25–30 µm broad.
Hymenium colourless, 35–50 µm high. Paraphyses anastomosing. Asci annelasceous, clavate, colourless, 35–
40 × 18–20 µm, tholus I–. Ascospores 2–4 per ascus, colourless, ellipsoid, muriform, 17–22 × 12–14 µm.
Secondary chemistry: no substances detected by TLC.
Distribution and habitat:—Known from several collections in Thai lower montane rainforest; found in
the understory in more illuminated situations.
Etymology:—Referring to the country in which the species was discovered.
Additional specimens examined (paratypes):—THAILAND. Chiang Mai Province: Huai Nam Dung
National Park, Papong 2686 (F), 2692 (KKU), 2694 (KKU); ibid., 1410 m, Papong 2812 (KKU), 2824
(KKU), 2830 (F), 2845 (KKU), 2916 (KKU), 2995 (KKU).
Aderkomyces thailandicus is recognized by its initially immersed-erumpent apothecia (similar to those of
several Gyalideopsis species) and its submuriform ascospores, 2–4 per ascus, in combination with the
verrucose thallus. No other species of Aderkomyces has a similar combination of characters or comes close to
this new taxon. The neotropical A. gomezii Lücking is most similar but has a smooth thallus and much larger
ascospores. Aderkomyces thailandicus agrees with several species of Gyalideopsis in the verrucose thallus
(Lücking 2008), but Gyalideopsis always lacks sterile setae.
Amandinea maritima Giralt, van den Boom & Elix, sp. nov. (Fig. 2E–F) Mycobank MB 517731
Sicut Amandinea fouquieriensis, sed differt thallo crustaceo, albido vel cremeo, hypothecio et parte interni excipuli
semper incoloribus, ascosporis typo Physconia.—http://www.eol.org/pages/Amandinea maritima
Type:—ITALY. Sardinia: Sassari, Capo Punta Falcone, Felsabbrüche, Dornpolster-Heiden, Juniperus
phoenicea-Bestände, July 1985, Nimis & Poelt (holotype GZU).
Thallus corticolous, episubstratal, crustose, continuous, thin and smooth to moderately thickened, rugose
to verrucose, whitish to creamy, covering large areas. Prothallus absent. Medulla not amyloid (I–), lacking
calcium oxalate (H2SO4–). Photobiont chlorococcoid, 10–15 µ m diameter. Apothecia initially immersederumpent and apparently lecanorine, surrounded by a pseudothalline margin concolorous with the thallus,
entire or cracked and discontinuous, becoming lecideine and sessile but remaining broadly attached, often
contiguous, up to 0.6 mm diameter. Disc black, epruinose, concave at first, then plane, rarely subconvex.
Proper margin thick in young apothecia, becoming thinner, always prominent, persistent. Proper excipulum ±
aethalea-type (Bungartz et al., 2007), 35–50 µm wide, prosoplectenchymatous, inner cells elongate, hyaline,
outermost cells short, swollen and brown. Hymenium colourless, 50–70(–80) µm high, not inspersed with oil
droplets. Hypothecium colourless, 75–100(–120) µ m high. Apical cells of the paraphyses 3–5(–6) µ m
diameter, with dark brown cap. Asci 8-spored, Bacidia-type (Rambold et al., 1994). Ascospores Physconiatype, (9–)10.7–13.3(–15) × (4–)5.2–6.5(–7) µm (M = 12; 5.8; SD = 1.3; 0.6; n = 100), ellipsoid, straight or
slightly curved, not constricted at septum, faintly microrugulate at 1000×; ontogeny of type A (Giralt 2001).
Pycnidia subimmersed, abundant, conidiophores type III (Vobis 1980). Conidia filiform, curved, 12–20 × 0.5–
1 µm. Secondary chemistry: nil or lichesterinic acid by HPLC (Elix et al. 2003).
Distribution and habitat:—Amandinea maritima is known only from coastal areas, including the
western Mediterranean coastal areas, from where it extends to the Atlantic coast of southern Portugal and the
Canary Islands. It grows in coastal-dune areas, mainly on Juniperus spp., accompanied by other typically
maritime Mediterranean-Atlantic-Macaronesian species such as Caloplaca aegatica Giralt, Nimis & Poelt,
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Diploicia canescens (Dicks.) A.Massal., Endohyalina kalbii (Giralt & Matzer) Giralt, van den Boom & Elix,
Rinodina anomala (Zahlbr.) H.Mayrhofer & Giralt and R. nimisii Giralt & H.Mayrhofer. Further associated
species on Juniperus in the Canary Islands locality include Bactrospora patellarioides (Nyl.) Almq.,
Lecanora sabinae Hern.-Padr. & Vänskä, Labrocarpon canariensis (D.Hawksw.) Etayo & Pérez-Ortega (on
Pertusaria) and Thelopsis isiaca Stizenb.
Etymology:—The name reflects the typical habitat of this species.
Additional specimens examined (paratypes):—SPAIN. Canary Islands: El Hierro, W of Sabinosa, Eside along road H1-500, SW of Montaña del Escobar, van den Boom 43015 (hb. van den Boom). Andalucía:
Málaga, Costa del Sol, Playas de Manilva, Egea s.n. (BCN). PORTUGAL. Alentejo: Porto Covo, 1 km S of
village, van den Boom 19164 (hb. van den Boom). Algarve: NW of Lagos, N of Carrapateira, van den Boom
32560 (hb. van den Boom).
The genus Amandinea was segregated from the genus Buellia on the basis of its filiform, curved conidia
(Choisy 1950, Scheidegger 1993). Additional diagnostic characters have been proposed for the genus
(Rambold et al. 1994, Sheard & May 1997, Marbach 2000, Mayrhofer & Sheard 2002). It is a mid-sized,
cosmopolitan genus of ca. 50 species which is particularly well-represented in tropical and subtropical areas,
where it commonly occurs on bark and lignum rather than on rocks. Amandinea shows considerable variation
in apothecium-type, hypothecium pigmentation and chemistry. Amandinea maritima is mainly distinguished
by the initially erumpent apothecia with a pseudothalline margin, the colourless hypothecium and inner part of
the proper excipulum and the small Physconia-type ascospores, with a fine microrugulate ornamentation.
Related species include A. dakotensis (H.Magn.) P.May & Sheard and Buellia fouquieriensis Bungartz,
corticolous species from North America. Like A. maritima, both have apothecia with an initial pseudothalline
margin. However, the former is distinguished by the significantly wider (Bungartz et al. 2007), Buellia-type
ascospores and the brown hypothecium (Sheard & May 1997, Bungartz et al. 2007). Buellia fouquieriensis is
mainly separated by the brown, subsquamulose to distinctly squamulose thallus and the brown hypothecium
(Bungartz et al. 2007; see B. fouquieriensis).
Ampliotrema cocosense Lücking & Chaves, sp. nov. (Fig. 3A) Mycobank MB 517732
Sicut Ampliotrema lepadinoides sed ascosporis muriformibusque differt.—http://www.eol.org/pages/Ampliotrema
cocosense
Type:—COSTA RICA. Puntarenas: Cocos Island National Park, Wafer Bay, 500 km SW of Costa Rica, trail
above ranger station, 87°04’W, 5°31’N, 100 m, lowland rainforest zone, closed primary forest, on bark
(branch), partially shaded, April 1992, Lücking s.n. (holotype CR, isotypes F, INB, USJ).
Thallus grey-olive-yellow, verrucose, with dense, paraplectenchymatous cortex; photobiont layer and/or
medulla with clusters of calcium oxalate crystals. Apothecia sessile, rounded, 1–2 mm diameter; disc partially
covered by 0.3–0.6 mm wide pore, brown-black, yellow- to orange-pruinose; margin entire, fused, yellowish
white, covered by thalline layer. Columella absent. Excipulum paraplectenchymatous, carbonized;
periphysoids absent. Hymenium 150–200 µm high, inspersed; paraphyses apically branched. Ascospores 4–8/
ascus, richly muriform, 80–100 17–22 µm, oblong, with thick septa and rounded lumina, colourless, I+
violet-blue (amyloid). Secondary chemistry: protocetraric and virensic acid and satellites; thallus (medulla)
C–, K–, P+ orange-red.
Distribution and habitat:—Known from a rich collection from lower montane rainforest on Cocos
Island.
Etymology:—The epithet refers to the type locality.
Ampliotrema cocosense is known from one well-developed collection made on Cocos Island. The lichens
of this island are not well-known except for foliicolous species (Lücking & Lücking 1995), and the fact that
the only two thelotremoid Graphidaceae collected there are new to science (Sipman et al. unpubl. data)
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indicates that for certain groups, Cocos Island harbors a unique lichen biota. Ampliotrema cocosense is the
only species in the genus with large, muriform ascospores, and hence easy to recognize. It is most closely
related to A. lepadinoides (Leight.) Kalb, which has ascospores of similar length but which are much narrower
and with transverse septa only. From species of Ocellularia s.lat. with large muriform ascospores, the new
species is set apart by the combination of a wide pore, absence of a columella, paraplectenchymatous
excipulum, inspersed hymenium, and protocetraric acid as main secondary substance.
Anomomorpha lecanorina Sipman, sp. nov. (Fig. 3B–C) Mycobank MB 517733
Anomomorpha ascomatibus rotundis margine lecanorino provisis, ad basin constrictis; hymenio insperso, ascosporis
submuriformibus 10–15 × 7 µm, 4 × 1–2 loculis.—http://www.eol.org./pages/Anomomorpha lecanorina
Type:—ECUADOR. Zamora-Chinchipe: Estacion Cientifico Sán Francisco Nature Reserve, S of road LojaZamora, ca. 40 km from Loja, 79°04’W, 3°58’S, 2025 m, primary montane forest on steep slope, tree near 5/I27, Elaeagia obovata, ca. 40 cm dbh, on canopy branches, June 2004, Sipman 52804 (holotype B, isotype
LOJA).
Thallus crustose, corticolous, forming large patches over 5–10 cm wide, pale grey, irregularly fissured,
delimited by a dark-brown prothallus line; upper surface flat to depressed-verrucose, smooth; cortex strong,
15–25 µm thick, of densely conglutinated, periclinal hyphae; medulla ca. 200 µm thick, of loose hyphae,
filled with many small crystals showing white in polarised light, sometimes including old, overgrown cortical
layers. Photobiont trentepohlioid, cells ca. 10 µm in diameter. Ascomata frequent, rounded with prominent
thalloid margin, 1–3 mm wide, sessile with constricted, wide base, with pale to dark grey-brown, pruinose,
flat disc surrounded by a thin, ca. 0.05 mm wide, white proper margin, which is often separated by a split from
the thallus-coloured, ca. 0.2 mm wide thalline margin. Hymenium 75–100 µm tall, inspersed with ca. 1–2 µm
wide, irregular droplets disappearing in K, I–. Paraphyses ca. 1.2 µm thick, septate, unbranched except near
the tip; terminal locules subglobose and to 3 µm thick, not spinulose. Hypothecium yellow-brown, I–. Asci
cylindrical with attenuated base, ca. 90 × 10 µm. Ascospores hyaline, 8/ascus, uniseriate, submuriform, with
rounded lumina, I+ dark blue-violet, 10–15 7 µ m, with 4 × 1–2 locules, halonate. Pycnidia not seen.
Secondary chemistry: thallus K+ orange, C–, KC–, UV–. Secondary metabolites: constictic acid (TLC).
Distribution and habitat:—Known so far only from three specimens from the Reserva Biológica Sán
Francisco, prov. Zamora-Chinchipe, southern Ecuador, where it was found in montane primary forest at
2000–2500 m. Two specimens were collected in the canopy of Elaeagia obovata trees, which have a rather
acid bark.
Etymology:—The epithet reflects the similarity of the ascocarps with the genus Lecanora, quite unusual
in Graphidaceae.
Additional specimens examined (paratypes):—ECUADOR. Zamora-Chinchipe: Cordillera Numbala,
Sán Francisco Biological Reserve, S of road Loja-Zamora, transecto 1, near Refugio, Sipman & Mandl 51464
(B, LOJA). Estación Científica Sán Francisco Nature Reserve, S of road Loja-Zamora, ca. 40 km from Loja,
Sipman 53142 (B, LOJA).
The family Graphidaceae is long known as very diverse in the tropics, but for a long time the large
number of species described on limited and often unconvincing characters was in most cases prohibitive for a
careful identification. Recent studies by Staiger (2002) and Lücking et al. (2008) have greatly improved the
situation and currently many additional species are being recognized, e.g. Dal-Forno & Eliasaro (2010).
Anomomorpha lecanorina is a very unusual species, because its ascomata look much like species of
Lecanora. However, the I-negative hymenium and I+ dark-violet, muriform spores are a certain indication for
Graphidaceae, and the species matches well the genera Anomomorpha and Platythecium (Staiger 2002).
Ascocarps in these genera are usually lirelloid, but may be rather variable in shape. Apart from the rounded
apothecia, A. lecanorina differs from other species of Anomomorpha, which mostly have norstictic acid, in the
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unusual chemistry. The region where the species was found appears to be particularly rich in locally endemic
lichens. For instance, Sipman et al. (2009) report three endemic Hypotrachyna species from the area.
FIGURE 3. A. Ampliotrema cocosense (holotype), apothecia. B–C. Anomomorpha lecanorina (holotype). B. Thallus
with apothecia. C. Hymenium with asci and ascospores. D. Anomomorpha tuberculata (holotype), thallus with lirellae
and tubercles. E–F. Aspicilia mansourii. E. General thallus morphology in situ. F. Rounded thallus lobes and tiny
rhizomorph-like extensions. Scale in A, D = 1 mm, in B, F = 2 mm, in C = 10 µm, in E = 20 mm.
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Anomomorpha tuberculata Lücking, Umaña & Will-Wolf, sp. nov. (Fig. 3D) Mycobank MB 517734
Anomomorpha speciei apotheciis sessilibus et thallo tuberculato differt.—http://www.eol.org/pages/Anomomorpha
tuberculata
Type:—COSTA RICA: Puntarenas: La Amistad International Park, Cerro Biolley Section, Sabanas
Esperanza (La Amistad Pacífico Conservation Area), Terraba Valley, 83°03’W, 9°04’N, 1300–1400 m,
lowland to lower montane moist forest savanna zone, disturbed savanna vegetation with abundant shrubs and
trees, on bark (lower trunk), July 2002, Lücking 15309c (holotype INB, isotype F).
Thallus corticolous, up to 10 cm diameter, 50–100 µm thick, continuous; surface uneven, grey to pale
brownish or yellowish grey, with numerous large tubercles; prothallus absent. Tubercles 0.8–1.5 mm diameter
and up to 1.5 mm high, orbicular with the base constricted, white, consisting of a wall of numerous corticate,
angular plates (resembling a soccer ball) which eventually break off as angular-rounded schizidia (especially
at the apex) to expose a soredia-like surface composed of an irregular mass of hyphae intermingled with
photobiont cells. Thallus in section with cartilaginous upper cortex, irregular algal layer and large clusters of
crystals. Apothecia angular-rounded to elongate and then flexuose, unbranched, sessile with the base strongly
constricted, with thick, prominent thalline margin, 1–3 mm long, 0.5–0.1 mm wide, 0.4–0.5 mm high; disc
exposed, grey-brown with white pruina and therefore appearing grey-pruinose; proper margin indistinct, labia
entire; thalline margin conspicuous, thick, prominent. Excipulum uncarbonized, orange-brown, 100–150 µm
wide; hypothecium prosoplectenchymatous, 30–50 µm high, colourless to pale yellow; hymenium 120–150
µm high, colourless, strongly inspersed, inspersion partly disappearing in K; epithecium granulose, 5–10 µm
high, grey-brown. Paraphyses unbranched; asci fusiform to clavate, 100–130 × 10–15 µm. Ascospores 8 per
ascus, ellipsoid, 3-septate, 7–10 × 5–6 µm, 1.5 times as long as wide, colourless. Secondary chemistry: no
substances detected by TLC.
Distribution and habitat:—The new species was found in a rare cerrado vegetation in the southern part
of Costa Rica.
Etymology:—The epithet refers to the conspicuous tubercles of the thallus.
Additional specimens examined (paratypes):—COSTA RICA. Alajuela: Volcán Tenorio National Park,
Pilón Biological Station (Arenal-Tempisque Conservation Area), Will-Wolf 12772 (INB, WIS), Sipman 51789
(B-600173357, INB). Puntarenas: Parque Internacional La Amistad (AC Amistad Pacifico), Cerro Biolley, 30
km NNW of Sán Vito near Biolley, July 2002, Sipman 48109c (B-600173358, INB). VENEZUELA: Bolivar:
Cerro Guaiquinima, in central part of upper plateau (near camp 4); February 1990, Sipman 26569 (B600083625, VEN); ibid., near NE edge of upper plateau (near camp 2), February 1990, Sipman 26782 (B600083627, VEN). GUYANA: Upper Mazaruni District: N-slope of mount Roraima (campsite 5), February
1985, Sipman & Aptroot 18760 (B-600113247), 18808 (B-60 0113248).
Anomomorpha is a small genus in Graphidaceae characterized by usually uncarbonized lirellae with
strongly inspersed hymenium and minute ascospores (Staiger 2002). The new species is placed in this genus
as it agrees in the non-carbonized lirellae with strongly inspersed hymenium and minute ascospores.
However, it deviates from other species of the genus in the strongly sessile lirellae and the lack of secondary
substances. In addition, the large tubercles formed on the thallus are unique within the family. Anomomorpha
tuberculata is probably closely related to the preceding species, since the apothecia are partly rounded and
resemble those of a Lecanora species. However, distinctly elongate lirellae are also present, the thallus
features large tubercles, the ascospores are 3-septate, and no secondary substances appear to be present.
Aspicilia mansourii Sohrabi, sp. nov. (Fig. 3E–F) Mycobank MB 518502
Thallus terricola, squamulosus vel subfruticulosus, subtus rhizomorphis obtectus. Similis Aspiciliae filiformis sed thallo
plus squamuloso, lobis mas irregularibus, ultimis simplicioribus, apice non nigrescentibus. Aspicilinam
continens.—http://www.eol.org/pages/Aspicilia mansourii
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• 17
Type:—IRAN: Golestan National Park, Mirzabaylou towards Almeh valley, 37°21’N, 56°12’E, 1300 m, May
2008, Sohrabi 15077 & Ghobad-Nejhad (holotype IRAN-MS015088, isotypes H-MS016188, GZUMS016189, hb. Sohrabi-MS016192).
Thallus terricolous, on soil or plant debris, subsquamulose to appressed sub-fruticose, attached by tiny
rhizomorph-like extensions, forming small patches up to 3–5(–8) cm wide, lobes discrete, more or less
stringy, continuous, warty, rimose, to areolate-verruculose to irregularly developed on the substrate, central
part subgranular to subsquamulose, mostly irregular, sometimes angular to rounded lobes, 0.3–0.9(–1.3) mm
in diameter, prothallus absent. Surface with small cracks or depressions, smooth to roughened, whitepruinose, sometimes diffusely granulose, when dry, dull, light grey, when wet, whitish grey to dark grey, ashy
grey. Cortex paraplectenchymatous, (10–)20–30(–35) µm thick, filled with granules and irregular globose to
round cells 5–8(–10) µm wide, uppermost part brown; cortex usually covered with a thick epinecral layer,
(10–)12–17(–22) µm thick, with crystals. Photobiont green, chlorococcoid, unicellular, cells 5–16(–22) µm
wide, distributed in regular to irregular layer, occasionally fragmented in small patches 30–70 µm wide.
Medulla ± loose to paraplectenchymatous, variable in thickness, lower parts somewhat interrupted by
prosoplectenchymatous tissue, originated from the small rhizomorph-like extensions. Lower surface white to
pale yellow, sometimes dark grey to ochraceous, without cortex, basally with 1–3 ± isodiametric cells, some
partly belonging to rhizomorph-like extensions. Rhizomorph-like extensions from the lower surface short, up
to 0.5–1 mm long, hypha 5–8(–10) µm thick, delicate, lacking algal cells, pale, white to yellowish, sometimes
light brown, usually attached to plant debris, occasionally visible under the margins. Apothecia and pycnidia
not known. Secondary chemistry: aspicilin and unknown fatty acid detected in some thalli by TLC; thallus
(medulla) K–, C–, P–, KC–, CK–, I–.
Distribution and habitat:—Aspicilia mansourii has been found on soil and dead plant debris, often on
dead tufts of Poa bulbosa L. and other perennial grass. It grows on calcareous soil in open situation in the
mountain areas with steppe-like habitat. It is so far known from Iran.
Etymology: The species is named in honor of Reza Mansouri, Iranian physicist who made significant
contributions to the development of science in Iran.
Additional specimens examined (paratypes):—IRAN: Golestan Prov., Golestan National Park,
Mirzabaylou towards Almeh valley, Sohrabi 15078 & Ghobad-Nejhad (hb. Sohrabi). East Azerbaijan Prov.,
Jolfa district 1 km S of Daran village, E of Hadishahr, Sohrabi et al. 10119, 10123 (hb. Sohrabi).
Aspicilia is a relatively large genus, comprising ca. 200 species. The majority of the species are
saxicolous, but several taxa are known to be terricolous, and a few lignicolous or corticolous species have also
been described. Recently, non-vagrant-terricolous species were revised (Sohrabi et al. 2010). Aspicilia
mansourii is distinguished from the terricolous North American species, A. californica Rosentreter, by its
subsquamulose to appressed subfruticose thallus, small and short lobes and tiny rhizomorph-like extensions,
lacking black lobe tips, as well as in negative reactions to K, C, and P in both medulla and cortex. It differs
from A. filiformis Rosentreter, another North American terricolous species, in lacking black tips on the lobes
and a different thallus morphology (Rosentreter 1998). There is another poorly known species, A. reptans
(Looman) Wetmore, which has a crustose, subsquamulose to appressed sub-fruticose thallus and its type is
fertile, with brown-black discs, growing directly on soil. However, rhizomorph-like extensions and chemistry
are lacking in A. reptans. Aspicilia crespina V.J.Rico is a Mediterranean species, known from Spain. It is also
fertile, squamulose and attached with large rhizomorph-like extensions to the substrate and usually grows
among mosses, on siliceous rocks and on soil, and lacks chemistry. Aspicilia tibetica Sohrabi & Owe-Larss.
was recently described from Tibet in China (Sohrabi et al. 2010). It has a well-developed fertile thallus and
commonly grows on soil and plant debris in the very high mountains of the Tibetan Plateau. Aspicilia tibetica
differs in certain morphological characters (e.g. apothecia very common), lacking secondary chemistry, and
whitish colour thallus, and its ecological requirements. Additional information can be found at the Myco-Lich
website http://www.myco-lich.com (Sohrabi & Ghobad-Nejhad 2010).
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LUMBSCH ET AL.
Bacidina sorediata Seaward & Lücking, sp. nov. (Fig. 4A) Mycobank MB 517735
Sicut Bacidina apiahica sed thallo sorediato differt.—http://www.eol.org/pages/Bacidina sorediata
Type:—SEYCHELLES. Mahé: Path to Le Niol Road, 70 m, January 1974, Norkett 18668B (holotype BM,
isotypes F, hb. Seaward 115567).
Thallus foliicolous, 5–10 mm across and 15–20 µm thick, dispersed into minute, corticate, granulose to
microsquamulose patches 0.05–0.1 × 0.05 mm, giving the entire thallus a granulose appearance, pale olivegreen, sorediate; soredia at first discrete, maculiform, becoming in part confluent, farinose, 0.2–0.3(–0.5) mm
diameter, yellowish white. Photobiont chlorococcoid, cells 5–10 µm diameter. Apothecia rounded, 0.25–0.5
mm diameter and 130–180 µm high; disc plane to slightly convex, pale yellow to orange-yellow; margin thin,
evanescent, same colour as disc or somewhat paler. Excipulum paraplectenchymatous, 30–50 µm broad.
Hypothecium 20–30 µm high, colourless to very pale yellow. Hymenium 40–50 µm high, colourless. Asci
clavate, 35–45 × 6–7 µm. Ascospores arranged in bundle, bacillar to very narrowly clavate, tapering towards
the proximal end, 25–35 × 1.2–1.7 µm, 17–22 times as long as broad, 3-septate. Pycnidia not observed.
Secondary chemistry: no substances detected by TLC.
Distribution and habitat:—Known from a rich collection from the Seychelles.
Etymology:—The epithet refers to the sorediate thallus, an unusual feature in the genus.
Bacidina is a small genus of crustose lichens with usually granulose to microsquamulose thallus and pale
yellow to orange apothecia (Ekman 1996, Lücking 2008). In several species, including Bacidina varia
S.Ekman, the farinose to granulose thallus can become diffusely sorediate. However, the combination of a
distinctly microsquamulose thallus consisting of corticate squamules with discrete, maculiform soralia of a
different colour is not yet known from the genus. Bacidina sorediata otherwise combines features of three
other common, foliicolous species: B. apiahica (Müll.Arg.) Vĕzda has the same type of ascospores but a
farinose to granulose thallus lacking distinct soralia, whereas B. defecta Vĕzda and B. scutellifera (Vĕzda)
Vĕzda have the same thallus squamules as in the new species and also share the apothecium and ascospore
type, but instead of soralia produce scutelliform isidia (Lücking 2008).
Badimia multiseptata Papong & Lücking, sp. nov. (Fig. 4B) Mycobank MB 517736
Sicut Badimia pallidula sed ascosporis maioribus multiseptatis differt.—http://www.eol.org/pages/Badimia multiseptata
Type:—THAILAND. Nakhorn Ratchasima Province: Khao Yai National Park, 14°11’N, 101°37’E, 650 m,
lower montane rainforest, shady understory, on leaves, December 2005, Papong 5888 (holotype KKU).
Thallus foliicolous, continuous or irregularly dispersed, 20–25 µm thick, densely verrucose, pale greenish
grey; verrucae wart-shaped, white. Photobiont chlorococcoid. Apothecia rounded, 0.4–0.8 mm diameter; disc
pale yellow to orange; margin slightly prominent, paler than the disc. Excipulum paraplectenchymatous,
lacking crystals, 30–40 μm broad. Hypothecium prosoplectenchymatous, 20–30 μm high, colourless to pale
yellow, K–. Epithecium indistinct. Hymenium 45–55 µm high, colourless, I+ blue. Paraphyses unbranched, 1–
1.5 µm thick. Asci clavate, 40–50 × 8–10 µm, I+ blue. Ascospores 8 per ascus, oblong to fusiform, (3–)5–7(–
9)-septate, without or with slight constrictions at septa, 25–32 × 4–5 µ m, 6–7 times as long as broad,
colourless. Campylidia sessile, halfmoon-shaped, chamois-coloured to white. Conidia filiform, 5–7-septate,
50–60 × 1–1.5 µm, with short lateral appendages at most segments, colourless. Chemistry: usnic acid and
zeorin.
Distribution and habitat:—Known from several collections from lowland rain forest in Thailand.
Etymology:—The epithet refers to the multiseptate ascospores as compared to other species of the genus.
Additional specimens examined (paratypes):—THAILAND. Chanthaburi Province: Namtok Phlio
National Park, Papong 882, 884-1, 887-1 (KKU). Loei Province: Phu Kradueng National Park, Papong 3269
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• 19
(KKU). Nakhon Si Thammarat Province: Khao Lung National Park, Papong 3526 (F), 3543 (KKU). Nakhorn
Ratchasima Province: Khao Yai National Park, Papong 5905-2 (F).
Badimia is a small genus of foliicolous lichens characterized by comparatively large, vividly coloured
apothecia and unique, campylidioid conidiomata (Vĕzda 1986, Lücking 2008). The genus is pantropical and
restricted to lowland rain forest and appears to be an excellent indicator of undisturbed rain forest (Lücking &
Kalb 2001). Two groups are distinguished which differ in apothecial colour and excipulum structure: the core
group of the genus with crystalline excipulum and opaque apothecia, and the pallidula-group with noncrystalline excipulum and slightly translucent, gyalectoid apothecia (Lücking 2008). Badimia multiseptata
belongs in the latter group and is characterized by its multiseptate, comparatively large ascospores. It is
morphologically similar to B. pallidula (Kremp.) V ĕ zda and relatives, but differs in its ascospores: B.
pallidula and B. polillensis (Vain.) Vĕzda have 3-septate ascospores, whereas those of B. verrucosa (Vĕzda)
Lücking & Vĕzda are submuriform.
Badimia vezdana Lücking, Farkas & Wirth, sp. nov. (Fig. 4C–D) Mycobank MB 517737
Sicut Badimia galbinea sed thallo continuo regulariter verrucoso differt.—http://www.eol.org/pages/Badimia vezdana
Type:—COSTA RICA. Limón: Tortuguero National Park, 10°32’N, 83°30’W, sea level, lowland rainforest
zone, primary rainforest close to village, June 1988, Lücking 88-56 (holotype F).
Thallus foliicolous, continuous, 10–40 mm across and 15–25 µm thick, strongly verrucose, pale bluish
grey; verrucae hemispherical, 0.1–0.15(–0.2) mm diameter, pale bluish grey to white, filled with yellow
crystals. Apothecia rounded, 0.3–0.8 mm diameter and 150–220 µm high; disc plane to slightly concave,
bright yellow to orange-yellow; margin distinct, slightly prominent, yellow to orange. Excipulum with
indistinct structure due to strong incrustation with yellow crystals, 20–40 µm broad, grey, in K under pressure
dissolving into free, moniliform hyphae with globose cells and constricted septa. Hypothecium 20–30 µm
high, colourless to pale yellow, K–. Epithecium 5–10 µm high, yellow granular. Hymenium 40–60 µm high,
colourless. Asci 35–50 × 8–12 µm. Ascospores ellipsoid to fusiform, 3-septate, with slight constrictions at
septa, 10–16 × 3.5–5 µ m, 3–3.5 times as long as broad, colourless. Campylidia 0.7–1.6 mm broad,
ferrugineous brown, wall incrusted with yellow crystals. Conidia filiform, 3–9-septate, 50–70 × 1.2–1.6 µm,
with lateral appendages up to 10 µm long and 0.7 µm thick, colourless. Chemistry: usnic acid, zeorin.
Distribution and habitat:—Neotropical and restricted to the shady understory of undisturbed lowland
rainforests.
Etymology:—The epithet honors the late Antonín Vĕzda, one of the prominent lichen taxonomists of the
20th century (Wirth 2009, Farkas 2010, Farkas et al. 2010a, b).
Additional specimens examined (paratypes):—COSTA RICA. Limón: Barra del Colorado National
Wildlife Refuge, Lücking 92-2386 (B, CR, GZU, LG, M, NY, STU, UPS), 92-2414 (B, CR, GZU, LG, M,
STU, UPS). PANAMA. Colón: Barro Colorado Island, Lücking 92-581 (hb. Lücking). COLOMBIA. Nariño:
La Espriella Forest Station, Sipman et al. 29591, 29665 (B). GUYANA. Potaro/Siparuni: Paramakatoi,
Lücking 96-3779 (BRG), 96-3781 (US). Upper Takutu: 50 km S of Aishalton, trail to Kassikaityu river,
Sipman 37086 (B). BRAZIL. Pará: Caxiuanã, 300 km W of Belém, Lücking 95-74 (F, Lücking, Lich. Fol.
Exs. 57).
This species was believed to be conspecific with the paleotropical B. galbinea (Kremp.) Vĕzda (Lücking
2008). However, restudy of the available material confirmed the differences previously observed between the
neotropical and paleotropical populations: the neotropical populations have a continuous, strongly and
regularly verrucose thallus with white verrucae and yellow to yellow-orange apothecia, whereas in the
paleotropical material, the irregularly laciniate thallus produces scattered and irregular, pale yellow verrucae
and the apothecia are ferrugineous-orange. We therefore describe the neotropical populations under a separate
name.
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LUMBSCH ET AL.
FIGURE 4. A. Bacidina sorediata (isotype), microsquamulose thallus with apothecia. B. Badimia multiseptata
(holotype), thallus with apothecia and one campylidium. C–D. Badimia vezdai (holotype). C. Thallus with apothecia. D.
Thallus with campylidia. E–F. Biatora epirotica (holotype). E. Habit. F. Cross section through apothecium stained with
lactic blue, note gelatinized hypothecium with "lacunae" (arrow). Scale in A–D = 1 mm, in E = 0.5 mm, in F = 10 µm.
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• 21
Biatora epirotica Printzen & T.Sprib., sp. nov. (Fig. 4E–F) Mycobank MB 519464
Biatorae pallentis similis sed ascosporis majoribus et hymenio altiore differt.—http://www.eol.org/pages/Biatora
epirotica
Type:—GREECE. Epirus, Nomos Ioanninon, Tzoumerka, near Palaiochorion, 39°30’N, 21°05’E, 776 m,
corticolous on twigs of Abies borisii-regis, April 2006, Spribille 19735 (holotype FR, isotypes ATHU, GZU,
further isotypes to be distributed as exsiccatae).
Thallus ecorticate, smooth to rimose, surface beige to gray, often with a greenish tinge, matt, esorediate,
up to 10 mm in diameter. Photobiont trebouxioid. Apothecia sessile with a broad base or slightly constricted
basally, 0.3–0.6 mm diameter; margin usually not visible, rarely as a lighter rim around the disc; disc weakly
to moderately convex, white to pale ochre, matt, epruinose. Excipulum hyaline, 30–40 µm laterally, 20–45(–
90) µm wide basally, of strongly gelatinized, diverging hyphae with 1–2.5 µm wide lumina, lumina of apical
cells 1.5–3.5 µm wide, partly free and with thick gelatinous outer walls. Hymenium hyaline, 50–60 μm high.
Paraphyses colourless, weakly to moderately branched and anastomosing, mostly in basal parts of the
hymenium and the subhymenium, lumina 0.8–1.2(–1.5) µm wide, apical cells with lumina of 1–3(–4) µm.
Subhymenium hyaline, 20–40(–55) µ m high. Hypothecium 20–40(–80) µ m high, of densely entangled,
narrow hyphae with lumina 0.8–1.2 µm wide, strongly gelatinised, but with characteristic "lacunae" 3–10 µm
wide. Asci Biatora-type, 8-spored. Ascospores (0–)3-septate, (11–)12.4–16.5(–19.5) × (3.3–)4.2–4.8(–5.5)
µ m, hyaline, narrowly ellipsoid, sometimes slightly bent. Pycnidia not found. Secondary chemistry: no
substances detected by TLC.
Distribution and habitat:—Biatora epirotica has been found on twigs of Abies borisii-regis, A.
nordmanniana, Laurus nobilis, Ostrya carpinifolia, Quercus coccifera, and trunks and dead twigs of young
Picea orientalis on the southwest Balkans and along the Turkish Black Sea coast. It mostly occurs in
mountain areas with high humidity and extensive cyanolichen communities at elevations of 450 to 1500 m.
Etymology:—The epithet is derived from Epirus, the geographical region on the Balkans where most of
the material was collected.
Additional specimens examined (paratypes):—GREECE. Epirus, Nomos Ioanninon, Dimos
Tzoumerka, near Kataraktis, shrine to Profitis Ilias, Spribille s.n. (hb. Spribille); ibid., near Palaiochorion,
Spribille 16283 (GZU); ibid., above Pramantha, Spribille 19818 (GZU); ibid., W of Arachthos River above
Politsa, Spribille 19861 (GZU). Epirus, Nomos Ioanninon, Dimos Koinotita Vovousas, Zagori region, Pindos
Mts., above Aoos River, S of Vovousa, Spribille 16001 (GZU). TURKEY. Bolu vilayet, S of Karacasu,
Spribille 23136 (GZU). Kars vilayet, lake Karagöl NE of Savşat, Printzen 6468, 6495 (BG).
Biatora epirotica most closely resembles B. pallens (Kullh.) Printzen in having 3-septate ascospores and
excipular hyphae with broad lumina. Biatora pallens is easily distinguished by its narrower ascospores (10.5–
17.0 × 2.5–4.0 µm), a more shallow hymenium (30–40 µm tall) and the presence of usnic acid. Three more
species of Biatora with predominantly 4-celled spores are known: B. aegrefaciens Printzen, B. nobilis
Printzen & Tønsberg and B. rufidula (Graewe) S.Ekman & Printzen. All three species have excipular hyphae
with narrowly elongate lumina and orange-brown apothecia. In B. aegrefaciens and B. rufidula, the exciple
reacts I+ violaceous. Biatora nobilis and B. aegrefaciens also have broader ascospores (5.0–8.5 µm wide).
The known distributional range of Biatora epirotica is centred around the Black Sea region, a well known
Tertiary and Pleistocene refugial area. Other species of Biatora with a similar distributional range include B.
bacidioides Printzen & Tønsberg, B. pontica Printzen & Tønsberg and B. longispora (Degel.) Lendemer &
Printzen. The latter two are also known from eastern North America and East Asia, and B. epirotica should be
looked for in those regions.
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LUMBSCH ET AL.
FIGURE 5. A–B. Buellia sulphurica (holotype). A. Thallus with apothecia. B. Section through apothecium. C.
Bunodophoron pinnatum (holotype), fertile branches to the left; note black mazaediate terminal apothecia. Sterile
branches with distinct pinnate branching to the right (photograph R. Ubral-Hedenberg). D. Byssoloma spinosum
(holotype), apothecia on the leaf surface and pycnidia on the leaf margin. E–F. Calopadia cinereopruinosa (E holotype, F
Bungartz 5480). E. Apothecia. F. Thallus with campylidia. Scale in A, D = 2 mm, in B = 50 µm, in C = 10 mm, in E, F =
1 mm.
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Buellia sulphurica Bungartz & Aptroot, sp. nov. (Fig. 5A–B) Mycobank MB 517738
Sicut Buellia saxicola sed thallo sulphureo, apotheciis minutis et hypothecio hyalino—http://www.eol.org/pages/Buellia
sulphurica
Type:—ECUADOR. Galápagos: Isabela Island, Volcán Alcedo, upper NNW-exposed slope inside the crater,
0°27’S, 91°7’W, 1055 m, open vegetation with Adianthus concinnum and scattered shrubs of Tournefortia
rufosericea among basalt blocks in the vicinity of the sulfur vents, on basalt, March 2006, Aptroot 64881
(holotype CDS-31458, isotype ABL).
Thallus crustose, thin, of dispersed groups of areoles, marginal ones ± subsquamulose, epilithic;
prothallus absent; thallus surface matt, bright neon yellow, finely whitish pruinose, phenocorticate; thallus
densely filled with fine crystals (ca. 5 µ m in diameter, dissolving in 10% KOH), and few large mineral
crystals (irregular in diameter, ca 30–50 µm, not dissolving). Apothecia lecideine, initially inconspicuously
lecanorine; (0.10–)0.15–0.25(–0.30) mm in diameter; soon adnate to sessile; proper margin black, soon
excluded, when emerging typically covered by coarse, thick thalline fragments; disc black, epruinose, initially
plane, but soon strongly convex; thalline exciple, if present, poorly developed and reduced to very few
adglutinated hyaline hyphae densely packed with photobiont cells, ± merging with the surrounding thallus;
proper exciple similar to the aethalea-type sensu Scheidegger (1993), but strongly reduced to few, parallel
hyphae, almost indistinct from the paraphyses, apically moderately swollen (textura oblita) and reddish brown
(cf. elachista-brown, HNO3–), pigmentation continuous with the epihymenium; excipular hyphae downward
soon losing pigmentation and not clearly differentiated from either hymenium or hypothecium, hymenium
hyaline, not inspersed with oil droplets; paraphyses simple to moderately branched, apically swollen, with a
brown pigment cap (cf. elachista-brown). Asci 8-spored, clavate, Bacidia-type. Ascospores oblong, not
constricted with age, with obtuse ends, not curved, (9.8–)10.7–[12.1]–12.7(–13.7) × (5.9–)6.0–[6.6]–6.7(–6.8)
µ m (n = 25), 1-septate, proper septum briefly thickened during spore ontogeny (Physconia-type);
ornamentation absent (not visible in DIC). Pycnidia not found. Secondary chemistry: rhizocarpic acid; thallus
P−, K−, C−, KC−, CK−; thallus surface and medulla not amyloid (always test with concentrated Lugol's
iodine or in the compound microscope; thallus reactions can be very weak!), the hymenium reacts amyloid in
Lugol's.
Distribution:—Known only from Galápagos, on Isabela Island (Volcán Alcedo: north-northwest exposed
inner caldera rim; Volcán Chico: along northern outer slope of Sierra Negra); on basalt (HCl–), close to
volcanic sulphur vents.
Etymology:—The epithet refers both to the thallus colour and the close approximation to sulphur vents.
Additional specimens examined (paratypes):—ECUADOR. Galápagos: Isabela Island, Volcán Alcedo,
upper NNW-exposed slope inside the crater, Aptroot 64815, 64798, 64800, 64797 (CDS). Volcán Chico, on
the N outer slope of the caldera of Sierra Negra, Bungartz 8732 (CDS-44070).
Despite its minute size, it is a very conspicuous species, unlikely to be overlooked in the field because of
its bright neon colour. Found only near sulphur vents. Both the conspicuous colour and its unusual habitat
suggest that the species could be truly endemic to the archipelago. The species is very unusual not only
because of its habitat and presumed high tolerance to sulphur. It contains rhizocarpic acid, a substance very
rarely encountered in Buellia and known only from few species of uncertain taxonomic placement, e.g.
Buellia centralis H.Magn. (Lamb 1968, Obermayer et al. 2004). The apothecia have a hyaline hypothecium
that is typically regarded as diagnostic for Rinodina rather than Buellia; as very few species of Buellia are
known to possess a weakly or completely unpigmented hypothecium. The ascospores show Physconia-type
ontogeny, i.e., they initially have a thickened septum, becoming reduced with age. This spore type is seen both
in Buellia and Rinodina. The apothecia are unusually small, have a strongly reduced proper exciple, and,
when emerging from the thallus, large chunks of thalline material initially remain attached. Nevertheless they
never develop a distinctly lecanorine margin. The main reason why the species is described here in Buellia
s.lat. is the typical Bacidia-type ascus, which in Lugol has the characteristic deep blue flanks merging at the
tip and a thin, unstained inner tholus axis.
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Bunodophoron pinnatum Wedin, sp. nov. (Fig. 5C) Mycobank MB 517739
Species insignis thallo elongato, subtereti, pinnato et sporis parvis, brunneis diagnoscenda. Sphaerophorinam et acidum
protocetraricum continens.—http://www.eol.org/pages/Bunodophoron pinnatum
Type:—PAPUA NEW GUINEA. Northern Province: English Peaks, 147°29’E, 8°45’S, 3600 m, 1988,
Lambley 1643 (holotype BM, isotype UPNG 17591, not seen).
Thallus elongated, narrow, up to 7 cm long. Fertile branches narrowly flattened to almost terete, 30–50
mm long and 1–2 mm wide; major branch sparingly branched but typically with abundant, perpendicular,
terete branchlets along the sides. Upper surface pale grey to white, smooth or rugose to slightly wrinkled
above the apothecia. Lower surface white. Ascomata sparse, terminal, 2–4 mm wide. Mazaedia subapically
exposed; thalline receptacle ruptured early but remaining and slightly widening at maturity. Ascospores ±
globose, 6.5–7.5 µ m in diameter, greyish brown, with an irregular ornamentation of mazaedial material.
Pycnidia common, along the sides and in the apices of terminal branchlets. Conidia oblong to obovoid, (3.0–
)3.5–4.5 × 1.5 µm. Secondary chemistry: sphaerophorin and protocetraric acid (no ascomata were tested due
to the scarce material).
Distribution and habitat:—Bunodophoron pinnatum is known from three localities in the Northern
Province of Papua New Guinea only. According to the label data, on Mt. Kenevi, B. pinnatum grows on trunks
of Dacrycarpus in stunted Dacrycarpus-dominated forests. The type does not carry any further information
on substrate or forest type.
Etymology:—The epithet refers to the pinnate branching of the thallus.
Additional specimens examined (paratypes):—PAPUA NEW GUINEA. Northern Province: S of Mt
Kenevi, Lambley 1349 (BM, UPNG 17167, not seen); ibid., Mt Kenevi, Myola, Lambley 1625 (BM, UPNG
17573, not seen).
Wedin (1993) reinstated the genus Bunodophoron A.Massal. (about 30 species) following a hypothesis on
the phylogeny of the mazaedia-producing family Sphaerophoraceae (Lecanorales). Bunodophoron includes
the majority of the species formerly classified in Sphaerophorus Pers. and differs in the spore shape and spore
ontogeny: the globose spores in Bunodophoron have an irregular ornamentation consisting of an amorphous
material adhering to the spore wall after the spores have been released from the asci (Tibell 1981).
Bunodophoron is further characterised by having a β-orcinol depsidone-based chemistry (compared to βorcinol depsides in Sphaerophorus) and rod-shaped conidia (compared to ellipsoidal in Sphaerophorus). A
detailed discussion of taxonomic concepts, relevant characters, and terminology is found in Wedin (1993,
1995). Bunodophoron pinnatum is a very characteristic species with no apparent close relatives. There are few
protocetraric acid-containing species in the tropics, and among these B. pinnatum is easily recognised by its
almost terete main branches, with a unique pinnate branching pattern, and the comparatively small, brown
ascospores.
Byssoloma spinulosum Sérus., sp. nov. (Fig. 5D) Mycobank MB 517740
A Byssolomate humboldtiano differt pycnidiis evolutis ad marginem foliis, apotheciis magnis et hypothecio reagenti K+
purpureo.—http://www.eol.org/pages/Byssoloma spinulosum
Type:—PAPUA NEW GUINEA. Northern Province: Owen Stanley Range, Myola, right bank of Iora river,
9°09’S, 147°46’E, 2100–2400 m, primary montane forest on slope, October 1995, Sérusiaux s.n. (holotype
LG, isotype F).
Thallus mostly developed along margins and scars of living leaves, over 1–1.5 cm in diameter, usually
continuous near the margins (not developing on the under surface of the leaves, not even in a non-lichenized
stade) but typically patchy, patches irregular in shape and size, whitish to very pale bluish, seemingly cottony
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at high magnification, withish prothallus sometimes seen between the patches (hyphae distinct at high
magnification). Apothecia present or not, rounded, (0.3–)0.4–0.5(–0.6) mm in diameter (incl. the margin),
with a pale yellowish to whitish margin, typically byssoid, especially in young apothecia, getting thinner
when old but never excluded, disc first slightly concave and eventually flat, bluish black. Excipulum made of
densely interwoven, almost cylindrical hyphae, without constrictions, inspersed with minute, pale orange
brown crystals (not disappearing in K). Hypothecium and apothecia basal center brown to dark brown, K+
dark brown or usually purple brown to bluish black. Asci clavate, 40–45 5–10 µm, of the Byssoloma-type.
Paraphyses simple or slightly branched, not broadened at apices. Ascospores 8/ascus, ellipsoid, 3-septate,
with rounded ends and without constrictions at the septa, 12–15 × 3.5–4 µm, with a thin halo around them.
Pycnidia mostly but not exclusively developed at the leaves margins (natural margins or along scars),
typically in a single row at the margins, conical or almost so, with a short but nevertheless distinct beak, 0.15–
0.2(–0.25) mm high, ca. 0.2 mm in diameter, basis covered by the thallus (sometimes up to the ostiole margin)
which can be slightly byssoid, upper part bluish grey to dark bluish, sometimes very pale, K+ pale bluish to
aeruginose, a pale yellow orange glut of conidia sometimes present. Conidia numerous, rarely ellipsoid,
usually enlarged at one end or typically pyriform, simple, colourless, 5×6 × 2–2.5 µm.
Distribution and habitat:—Byssoloma spinulosum is typically a foliicolous species in the understory of
montane rainforest. So far, it has been found in two localities in the mountains of Papua New Guinea.
Etymology:—The epithet refers to the characteristic pycnidia frequently located at the leaves margins
and that look like tiny spinules.
Additional specimen examined (paratype):—PAPUA NEW GUINEA. Madang: S side of Ramu river,
Bundi village, on slope towards Mt Pizetara, November 1995, Sérusiaux s.n. (LG).
Several species of foliicolous lichens in Lecanorales have beaked pycnidia. They belong to two families
(Ramalinaceae for the first genus mentioned and Pilocarpaceae for all others) and different genera (Lücking,
2008): Bacidina pseudohyphophorifera (Lücking & Sérus.) Lücking (probably pantropical), Byssoloma
humboldtianum Lücking & Kalb (Neotropics), B. kalbii Sérus. (Macaronesia), Fellhanera africana (Vĕzda)
Lücking (Paleotropics), F. pilomarginata Lücking (Central America), F. tricharioides Lücking & R.Sant.
(Neotropics), F. vandenberghenii (Sérus.) Vĕzda (Neotropics and Africa), Eugeniella corallifera (Lücking)
Lücking, Sérus. & Kalb (Neotropics), and Szczawinskia tsugae Fink (Central and North America). Two
different types of pycnidia are involved: (type 1): the conidiogenous chamber is located at the tip of a pedicel
(Bacidina and Szczawinskia) or (type 2): the conidiogenous chamber is sessile (thus forming a typical
pycnidium) but its ostiole is extended into a distinct beak or pedicel and thus opens at its tip (other genera
mentioned above). A further case (type 3): is provided by the rare Bacidia clauzadei Sérus. & Lambinon, so
far known only from Central Africa, where the upper surface of cilia is covered with proliferating
conidiogenous cells (Sérusiaux & Lambinon 1994). We here report the discovery of a further foliicolous
species with beaked pycnidia, found in montane forests of Papua New Guinea. It is easily characterized by the
position of the pycnidia, which are most developed at the margins of the leaves, either the natural margins or
along scars. The ellipsoid, small and 3-septate ascospores and a byssoid margin (densely interwoven hyphae
with cylindrical cells without constrictions, incrusted with minute brownish granules) point to the speciose
and diverse genus Byssoloma in the Pilocarpaceae. Phylogenetic studies with statistical treatment of several
loci sequences are very much needed to circumscribe the genera within that family; meanwhile, the generic
delimitation of Lücking (2008) is the most appropriate. Within the genus, Byssoloma spinosum is close to the
Amazonian endemic B. humboldtianum (Lücking & Kalb 2000) and, besides the main development of its
pycnidia at leaves margins and scars, differs in the following characters: apothecia (0.3–)0.4–0.5(–0.6) mm in
diameter (vs 0.15–0.25 mm in B. humboldtianum) with a thick byssoid margin inspersed with minute orange
granules (vs thin and without granules in B. humboldtianum), and hypothecium and apothecia basal center
brown to dark brown, K+ purple brown to bluish black (vs hypothecium sordid aeruginous and apothecial
base dark brown, both being K– in B. humboldtianum). Apart from the conspicuous, beaked pycnidia, the new
species is morphologically close to B. confusum Farkas & Vĕzda and B. discordans var. flavescens G.Thor,
Lücking & Tat.Matsumoto, which agree in the pale thallus with a bluish tinge and compact, crystalline
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LUMBSCH ET AL.
apothecial margins, but B. confusum has colourless and B. discordans var. flavescens has yellow crystals in
the margin.
Calopadia cinereopruinosa Bungartz & Lücking, sp. nov. (Fig. 5E–F) Mycobank MB 517741
Sicut Calopadia subcoerulescens sed apotheciis cinereopruinosis differt.—http://www.eol.org/pages/Calopadia
cinereopruinosa
Type:—ECUADOR. Galápagos: Isla Sán Cristóbal, area W of Cerro Pelado on the way to El Ripioso,
0°52’S, 89°28’W, 400 m, transition zone, open Psidium guajava shrubland with Macraea laricifolia and
dominant annual herb Malachra capitata, on bark and wood, dead twigs of Psidium guajava, sunny, windand rain-exposed, August 2008, Bungartz 8489 (holotype CDS-41135).
Thallus corticolous or between bryophytes, continuous or marginally dispersed into rounded, confluent
patches, up to 30 mm across and 20–40 µm thick, ecorticate or with an indistinct cortical layer, smooth, pale
grey to greenish grey. Apothecia rounded, 0.4–1 mm diameter and 250–350 µm high; disc plane, grey-black
and with distinct, white pruina; margin distinct, rather thick and prominent when young, grey-black and
white-pruinose. Excipulum 50–70 µm broad. Hypothecium 50–100 µm high, aeruginous. Apothecial base
aeruginous. Epithecium 5–10 µm high, granular, grey-brown. Hymenium 120–190 µm high, colourless. Asci
100–170 × 20–30 µm. Ascospores single, oblong-ellipsoid, muriform, 45–80 × 15–25 µm, 3–3.5 times as long
as broad, colourless. Campylidia sessile, 0.4–0.7 mm broad; lobe well-developed, hood-shaped, grey but
white-pruinose; socle not apparent. Conidia filiform with clavate apex, 3–7-septate, 40–55 × 1–1.5 µm.
Distribution and habitat:—The new species was found fertile only on Isla Sán Cristóbal in the transition
zone towards the humid upper parts of the island. Collections with campylidia were also found on other
islands mostly in the humid zone. The species usually grows on bark of stems and twigs.
Etymology:—The epithet refers to the grey-black, white-pruinose apothecia.
Additional specimens examined (paratypes):—ECUADOR. Galápagos: Isla Santa Cruz, N of El
Puntudo, Bungartz 7295 (CDS). Isla Santa Cruz, near Los Gemelos craters, Bungartz 5480 (CDS).
Calopadia cinereopruinosa closely resembles C. subcoerulescens (Zahlbr.) Vĕzda in the grey-black
apothecia and aeruginous hypothecium. However, the apothecia are thickly and persistently pruinose, a
feature not observed in C. subcoerulescens. The new species is similar to C. editae, in forming pruinose
apothecia and ascospores shorter than 100 µm, but C. editae has brownish apothecia often with a yellowish
tinge and the hypothecium is light brown. The two paratype specimens bear campylidia only and their
identification is not absolutely certain, however, the pruinose lobes indicate that they belong to the new
species, as other species in the genus with non-pruinose apothecia also have non-pruinose campylidia.
Calopadia editae Vĕzda ex Chaves & Lücking, sp. nov. (Fig. 6A) Mycobank MB 517742
Sicut Calopadia perpallida sed ascosporis brevioribus differt.—http://www.eol.org/pages/Calopadia editae
Type:—TANZANIA. Morogoro Region: Submontane rainforest on the SE slope of Mt. Kanga, along Mkange
stream, 850–1200 m, foliicolous, March 1989, Farkas 89110 (holotype PRA).
Thallus foliicolous or corticolous, continuous or marginally dispersed into rounded, confluent patches, up
to 50 mm across and 20–40 µm thick, ecorticate, smooth, pale grey to brownish grey. Apothecia rounded, 0.5–
1 mm diameter and 250–350 µm high; disc initially plane but becoming convex in old apothecia, light brown
and with thick, pale yellowish to creme-coloured pruina; margin distinct, rather thick and prominent when
young, creme-coloured. Excipulum 40–70 µm broad. Hypothecium 50–100 µm high, light brown. Apothecial
base sordid brown to aeruginous. Epithecium thin, 5–10 µm high, granular, pale yellowish brown. Hymenium
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100–130 µm high, colourless. Asci 90–110 × 30–35 µm. Ascospores single, oblong-ellipsoid, muriform, 50–
80 × (15–)20–30 µm, 2.5–4 times as long as broad, colourless. Campylidia sessile, 0.4–0.6 mm broad; lobe
well-developed, hood-shaped, grey but white to pale yellow pruinose; socle not apparent. Conidia filiform
with clavate apex, 3–7-septate, 40–60 × 1–1.5 µm.
Distribution and habitat:—The new species appears to have a wide tropical distribution, as it has also
been found in collections from Costa Rica and the Galápagos Islands (Bungartz et al. unpubl. data). The
available collections are either foliicolous or corticolous and found in more or less exposed microsites at low
to mid elevations.
Etymology:— Antonín Vĕzda wished to dedicate this beautiful species to the Hungarian lichenologist
Edit Farkas, but due to illness did not have the opportunity to formally publish it before he died in 2008. Since
we have, meanwhile, found the species in several localities across the tropics, we take up Vĕzda's original
name for this taxon and validate it here.
Additional specimen examined (paratype):—COSTA RICA. Limón: Gandoca-Manzanillo Wildlife
Refuge, Manzanillo Section (La Amistad Caribe Conservation Area), Manzanillo, Lücking 17098a (F, INB).
Calopadia editae is externally similar to C. perpallida (Nyl.) Vĕzda, another pantropical species.
However, the ascospores in the latter are longer and mostly exceed 100 µm in length. The ascospores of C.
editae are similar in size to those of C. fusca (Müll.Arg.) Vĕzda and relatives, but these species lack pruina on
the apothecia (Lücking 2008).
Caloplaca brownlieae S.Y.Kondr., Elix & Kärnefelt, sp. nov. (Fig. 6B) Mycobank MB 517743
Caloplacae cinnabarinae similis, sed thallo obscure roseo vel obscure rubello, interdum inconspicue albo-pruinato,
ascosporis latioribus (6–7 µm latis) et latius septatis (3–5 µm), praesentia acido gyrophorici multo, etiam acido
ovoico et acido lecanorico differt.—http://www.eol.org/pages/Caloplaca brownlieae
TYPE:—AUSTRALIA. New South Wales: Above Lake Eucumbene, on granite boulder; April 1969,
Brownlie (holotype MEL 1023635).
Thallus 3–5 cm wide or aggregated in larger colonies, crustose, distinctly areolate, dull pink, dirty whitish
pink to pinkish orange or dull brownish orange; areoles 0.3–0.8 mm wide, somewhat smaller in the centre, but
1.0–1.5 mm wide in the peripheral zone, angular or more or less rounded with somewhat raised and slightly
darker coloured edges, upper surface dull pink. Apothecia (0.2–)0.3–0.6 mm diameter, immersed at first then
slightly raised above the level of the areole, usually 1–3(–5) per areole, rounded to distorted by mutual
pressure, rarely aggregating in irregular, compound apothecia, often with a very thin, obscure, paler pinkish
thalline rim ca. 40–50 µm wide; disc plane to subconvex, dark reddish brown to dark brick orange, sometimes
with a sparse whitish pruina; hymenium ca. 50 µm high; uppermost cells of paraphyses swollen, to 4 µm
diameter; asci (7–)8–spored, very small, 27–30(–40) × 12–15 µm, various sized ascospores sometimes present
in the same ascus; ascospores broadly ellipsoid, slightly widened at the septum, (8–)9–11(–14) × (5.5–)6–7
µm in water and (9–)10–13(–14) × 7–9 µm in K, septum 3–5 µm wide in water and 3–5(–6) µm wide in K.
Secondary chemistry: parietin (major), gyrophoric acid (major), ovoic acid (minor/trace), lecanoric acid
(minor), xanthorin (minor), erythroglaucin (minor).
Distribution and habitat:—Scattered on siliceous rocks in Western Australia, Northern Territory,
Queensland, Australian Capital Territory, New South Wales, and South Australia.
Etymology:—The species is named in honour of the Australian botanist Sue Brownlie, who kindly
provided us with the type collection.
Additional specimens examined (paratypes):—AUSTRALIA. Western Australia: Wokatharra Hill, W
Moresby Range, August 1983, Cranfield s.n. (PERTH). Lake Argyle Road, 31 km south-east of Kununurra,
Elix 27797, Lumbsch & Streimann 39419 (CANB). Northern Territory: Newcastle Range, 12 km SSW of
Timber Creek, Streimann 42016 (CANB); escarpment, Newcastle Range, 8 km SSW of Timber Creek,
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Streimann 42016 (CANB). Queensland: Mt. Walker, 15 km S of Hughenden, Elix & Streimann 20697
(CANB). East Leichhardt River Reservoir, presumably near Mt Isa, September 1969, Brownlie s.n. (MEL).
First Turkey, Mt Archer Environmental Park, 7 km NE of Rockhampton, Elix 34518 (CANB). Australian
Capital Territory: 11 km S of Tharwa, Booroomba rocks, Filson 19336 (MEL). Brindabella Ranges, 9 km SW
of Tharwa, Booroomba Rocks, Filson 16736 (MEL). New South Wales: 'Bush Bottoms', 32 km NE of
Goulburn, Streimann 38875 (B, CANB). Armidale Hills, Elix 2694 (CANB). South Australia: Kangaroo
Island, just S of Wisanger Hills H. S., 7 km WSW of Emu Bay, Elix & Elix 19644 (CANB).
Caloplaca brownlieae is similar to C. cinnabarina (Ach.) Zahlbr. in having dark reddish brown to dark
brick orange discs but differs in its dull pink to dull brownish orange thallus, wider paraphysis tips, wider
ascospores and septa. Caloplaca australiensis S.Y.Kondr. & Kärnefelt is also similar. The latter differs in its
distinctly areolate, crustose thallus, while C. montisfracti S.Y.Kondr. & Kärnefelt has a thicker, somewhat
subconvex thallus with slightly raised edges (Kondratyuk et al. 2007). A fourth similar species is C. eos
S.Y.Kondr. & Kärnefelt, from which C. brownlieae is distinguished by having more immersed, somewhat
biatorine apothecia. Finally, Caloplaca brownliae differs from all four species in the high concentration of
gyrophoric acid and presence of ovoic and lecanoric acids (Kondratyuk et al. 2007).
Caloplaca decipioides Arup, sp. nov. (Fig. 6C) Mycobank MB 517744
Caloplacae decipienti similis, sed lobi magis plani, soralia ex isidiis demum fractis formata, nec in calce crescit.—http:/
/www.eol.org/pages/Caloplaca decipioides
Type:—SOUTH KOREA. Gangwon Province: Inje-gun, Buk-myun, Yongdae-ri, Sorak-san National Park,
inner part of the massif Sorak Mts, along the road in Backdam Valley from Backdam (Paekdam) temple
towards the village Yongdae-ri, where the road crosses the river ca. 1.5 km NW Backdam (Paekdam) temple,
38°10’N, 128°22’E, 410–420 m, on almost vertical rock, shaded from running water, probably at least partly
with a higher pH than true siliceous rocks, exposed to S at the river, October 2006, Thor 20768 (holotype
UPS, isotypes NIBR, UPS).
Thallus to 1.5 cm, single or coalescing with other thalli, lobate, growing radiately in rosettes with
marginal lobes and the centre covered by smaller, irregularly arranged, often imbricate lobes; lobes 0.5–2.3 ×
0.2–0.6 mm, slightly convex to rather flat, slightly wider toward tips, irregularly branched one to three times,
divided by narrow but distinct furrows, 0.1–0.2 mm wide, surface ± smooth to finely granular near lobe tips,
yellow (in shade) to dark orange or red-orange, slightly paler at margin and partly with very thin white pruina;
soralia ± punctiform or irregular in outline, laminal or terminal on central lobes, initiating as globose to
elongate isidia that soon dissolve into soredia; isidia 75–125 µm when globose and up to 200 µm long when
elongate, in dense clusters, of the same colour as the thallus; soredia 35–60 µm, rather compact with rather
smooth surface, yellow to orange-yellow, contrasting against the darker thallus; cortex 20–40 µm thick,
paraplectenchymatous with thin-walled cells 4–8 µ m in diameter, without epinecral layer; algal layer ±
continuous with trebouxioid algal cells 7–12 µ m in diameter; prothallus not seen; pycnidia not seen.
Apothecia not seen. Secondary chemistry (HPLC): Parietin (major), fallacinal (major), vicanicin (major,
isofulgidin (major), teloschistin (major) and traces of emodin, parietinic acid and caloploicin. This
chemosyndrome corresponds rather well with A+3 according to Søchting & Frödén (2002).
Distribution and habitat:—The new species was collected under an overhang of a tall rock with a river
running below. The rock face was exposed to the south and there were no trees giving any shade. The lichen
flora is very rich at the locality and C. decipioides was very abundant in suitable places away from both rain
and running water. The rock was very hard and probably of some kind of greenstone, but did not contain any
lime. The new species is so far known only from the type locality. The locailty is situated in the Sorak-san
National Park and many habitats included mainly oldgrowth forests on steep mountain slopes dominated by
deciduous trees. The climate is somewhat continental with warm summers (up to 36°C) and cold winters
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(down to –17°C). The annual precipitation is 1300 mm and the mountains are snow-covered from November
to April. The species was collected at just above 400 m elevation. The distribution range is not yet known for
this species but it is probaly rare since the collector has not seen it elsewhere in Korea or Japan on his
exursions in the region.
FIGURE 6. A. Calopadia editae (holotype), apothecia. B. Caloplaca brownlieae (holotype), general habit. C. Caloplaca
decipioides (holotype), habitus. D–E. Caloplaca digitaurea (D, holotype; E, Elvebakk 97-468). D. Vertical view of
thallus. E. Thallus with apothecia. F. Caloplaca magnussoniana (holotype), general habit. Scale in A–E = 1 mm, in F = 2
mm.
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Etymology:—The epithet refers to the similarity with C. decipiens.
This new species of Caloplaca was collected in South Korea by G. Thor who kindly put the material at the
author's disposal. The lobate species turned out to be difficult to determine using appropriate keys and works.
Neither Gaya's recent work on the C. saxicola group (Gaya 2009) nor those from North America (Arup 1995,
Wetmore & Kärnefelt 1998, Wetmore 2007), nor Kondratyuk's papers on Australian species (Kondratyuk et
al. 2007a, 2007b, 2009a, 2009b), nor Poelt & Hinteregger's (1993) book on Himalayan Caloplaca helped in
determining the species. None of the older papers from Asia were useful either (Nylander 1890, Magnusson
1940, 1944). Finally, the new species failed to match any of the many genetically investigated species of
Caloplaca in the possesion of the author and U. Søchting or in Genbank. The new species is characterized by
the orange lobate thallus with clusters of isidia that dissolve into soredia. It is similar to C. decipiens (Arnold)
Blomb. & Forssell, a rather common lichen on anthropogenic calcareous substrates in Europe, but differs in
the flatter lobes and in the soredia that develop from breakdown of globose isidia. In addition, the isidia and
soralia are laminal or terminal on horizontal lobes rather than borne on ascending lobe tips as in C. decipiens.
Finally, C. decipioides differs in its ecology, as it grows on hard, non-calcareous rocks and not on limestone,
concrete or mortar. C. decipens sometimes produces apothecia, but the new species has not yet been found
fertile. Molecular data also show that these two species are not closely related, as C. decipiens belongs to the
C. saxicola group (Gaya et al. 2008) and C. decipioides belongs to quite another clade within the family
(unpublished data). The new species could also be confused with C. cirrochora (Ach.) Th.Fr., but differs from
that species in the same way as it differs from C. decipiens.
Caloplaca digitaurea Søgaard, Søchting & Sancho, sp. nov. (Fig. 6D–E) Mycobank MB 517745
Sicut Caloplaca wetmorei sed apothecia majores et thallus non parasitcus.—http://www.eol.org/pages/Caloplaca
digitaurea
Type:—CHILE. XII Región de Magellanes y de la Antártica Chilena: 15 km N of Puerto Natales, Cueva del
Milodón, 51°34’S, 72°37’W, 145 m, January 2005, Søchting 10.401 (holotype C).
Thallus fruticulose, composed of mostly vertical, terete to rarely flattened, moderately branched, isidialike lobules dispersed among mosses or cyanobacteria and sometimes forming up to 2 mm high, compact
cushions; vertical lobules 60–100 µm thick and up to 1500 µm high, slightly moniliform, at the thallus edge
slightly flattened and more horizontal, colour vivid orange yellow to vivid orange with uneven, matt surface.
Photobiont trebouxioid. Apothecia sparse, embedded between the lobes, 0.8–1.5 mm with regular to
flexuouse margin, initially zeorine with a proper margin concolorous with disc and an isidiate thalline margin
concolorous with thallus, later lecanorine with an elevated margin, which is often obscured by isidia; disc
deep orange, darker than thallus, flat; episamma medium coarse. Exciple fan shaped, laterally ca. 80 µ m,
centrally ca. 20 µm. Hymenium 60–100 µm; paraphyses branched with slightly thickened end cells, up to 4
µm. Asci clavate with 8 spores. Spores ellipsoid, polarbilocular, (12.5–)13–16(–18) × (5–)6–7.5(–8) µm;
septum narrow, (1–)1.5–2.5(–3.5) µm. Pycnidia not observed. Secondary chemistry (HPLC): parietin (major),
teloschistin (minor), fallacinal (minor), parietinic acid (minor), emodin (minor); conforms to chemosyndrome
A according to Søchting (1997).
Distribution and habitat:—Chile, Province Última Esperanza and Province Antárctica Chilena, Isla
Navarino. On rock, detritus and over other lichens on exposed outcrops in shrub vegetation, and grassland by
coast, but not maritime, 5–170 m elevation.
Etymology:—The epithet digitaurea refers to the golden, finger-like isidia with a slight reference to the
James Bond blockbuster movie Goldfinger from 1964. Its English vernacular name would be "Goldfinger Fire
Dot Lichen".
Additional specimens examined (paratypes):—CHILE. XII Región de Magellanes y de la Antártica
Chilena. Última Esperanza Province, Parque Nacional Torres del Paine, Søgaard 38 (C). Parque Nacional
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Torres del Paine. Walk from Lago Grey, Søgaard 43, 48 (C), Parque Nacional Torres del Paine, Lago Pehoe,
Río Paine, 2 km NE of Laguna Linda, Elvebakk 97:468 (C). Antártica Chilena Province, Isla Navarino, 37 km
W of Pto. Williams, Caleta Honde, Søchting 10.175 (C).
The genus Caloplaca is one of the largest lichen genera, with probably close to 1000 species, particularly
in cool regions, favoring especially calcareous or nutrient rich substrata or maritime rocks. The cool parts of
the Southern Hemisphere are still poorly studied. Caloplaca digitaurea is one of a number of undescribed
species discovered during recent field work in Southern Patagonia as part of the Spanish ANTRAX project. It
is characterized by the vivid orange, thick and terete isidia-like lobules, often in microfruticose clusters. The
lobules are mostly distinct, always growing ± vertically upwards. The species can resemble C. isidioclada
Zahlbr., but C. digitaurea has thicker lobules (up to 100 µm) than C. isidioclada (30–60 µm). Based on
available ITS sequences and morphology, Caloplaca wetmorei Nimis, Poelt & Tretiach from the Sonoran
Desert seems to be the closest relative. It has, however, smaller apothecia (0.3–0.4 mm) and a parasitic life
habit (Nimis et al. 1994). Also, even though C. wetmorei may be a relative, there is only a 93 % similarity
between the ITS region of the nuclear rDNA of the two species (C. digitaurea Genbank number HQ 317922,
C. wetmorei, isotype Genbank number HQ 317923). Therefore it is unlikely that they belong to the same
taxon.
Caloplaca magnussoniana S.Y.Kondr., Kärnefelt & A.Thell, sp. nov. (Fig. 6F) Mycobank MB 517746
Caloplacae lithophilae similis, sed apotheciis in thallo centrali aggregatioribus, paraphysibus cum cellulis oleosis, et
septo ascosporae angustiore (3–4 µm lato) differt.—http://www.eol.org/pages/Caloplaca magnussoniana
Type: AUSTRALIA. Tasmania: Mt Barrow Chalet; 41°23’S, 147°25’E, 600 m, on dolerite, sheltered and
rare, January 1966, Bratt & Bratt 3226 (holotype HO 38775).
Thallus 1–2 cm wide, usually aggregated into larger colonies, appearing dull yellowish or dull brownish
orange due to numerous apothecia, thallus areoles absent or rare, minute, to 0.1 mm wide, whitish grey
greenish-yellow or whitish-yellowish, in the peripheral zone or among the apothecia in the centre of thallus,
rarely seen as undulating fragments of a continuous yellowish film, ca. 0.3 mm wide. Apothecia 0.2–0.4(–
0.55) mm diameter, 0.2 mm thick, usually numerous and densely aggregated, biatorine, dull yellowish, proper
margin margin 0.03–0.05 mm wide, becoming excluded, disc plane to subconvex, dull orange to dull
brownish orange; in section biatorine; true exciple to 70 µm thick in the uppermost lateral portion, 10–15 µm
thick in the basal portion; hymenium 70–75 µ m high; paraphyses swollen towards the tips, to 4–5 µ m
diameter, with numerous oil cells, 4–5 µm diameter (better seen in K!); ascospores broadly ellipsoid, (8–)9–
12 × (5.5–)6–7 µm in water and 10–13(–15) × 7–10 µm in K, septum 3–4 µm wide in water and (3–)4–5(–6)
µm wide in K. Secondary chemistry: epihymenium K+ dark purple, becoming pale, in K brownish.
Distribution and habitat:—Known from scattered localities in Victoria and Tasmania, but probably
overlooked. Australian records of C. lithophila or epilithic collections of C. pyracea or C. holocarpa may
represent this new species. It grows on siliceous or calcareous rocks.
Etymology:—It is named for the well known Swedish lichenologist A. H. Magnusson, who made
significant contributions to our knowledge of Caloplaca and described C. lithophila, which is closely related
to this new species.
Additional specimens examined (paratypes):—AUSTRALIA. Victoria: Gippsland Plain, Highett
(suburb of Melbourne), CSIRO, March 1985, Martin s.n. (MEL). 5 km E of Tragalgon, Flynn’s Creek Road,
heavily grazed paddock, Filson 15219 (MEL). 26 km E of Murrayville on the Mallee Highway, 20 km S of
Walpeup, Kärnefelt 992601, 992602 (LD). Tasmania: Clarkes Island, ca. 5.1 km at 2 degrees NW of Black
Point, Whinray 1442 (MEL).
The key characters indicate that C. magnussoniana is closely related to C. lithophila H.Magn. and C.
approximata (Lynge) H.Magn. It differs from C. approximata in having wider ascospores and septa and from
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LUMBSCH ET AL.
C. lithophila by its narrower ascospores and by the presence of oil cells, not only oil droplets, in the
paraphyses. It differs from both these species in lacking distinct thalline areoles and in having apothecia more
aggregated towards the centre (Kondratyuk et al. 2004).
Caloplaca mereschkowskiana S.Y.Kondr. & Kärnefelt, sp. nov. (Fig. 7A) Mycobank MB 517747
Caloplacae brachysporae similis, sed apotheciis aggregatis, paraphysibus supra cum cellulis et guttulis oleosis, ascis
saepe 1–2–4–6-sporis, ascosporis latius septatis (3–4(–6) µm), et hypothallo obscure cinereo deficiente differt.—
http://www.eol.org/pages/Caloplaca mereschkowskiana
Type: AUSTRALIA. Western Australia: North of Northampton, road to Horrocks, Bower River Road, on
calcareous rocks, 28°24’S, 114°27’E, January 2004, Kärnefelt et al. 20041503 (holotype PERTH, isotypes
CANB, LD, KW).
Thallus 0.5–2 cm wide, usually aggregated into larger colonies, appearing dull orange or brownish orange
due to the numerous apothecia, thallus areoles yellowish white to greenish yellow-white or brownish, very
thin, immersed and indistinct in the peripheral zone to rather thick and areolate in the centre, areoles
immersed and almost indiscernible at the periphery to distinct in the centre, (0.2–)0.3–0.8(–1.3) mm wide,
0.3–0.4 mm thick. Apothecia 0.2–0.5(–0.7) mm diameter, 0.15–0.22 mm thick, numerous, immersed at first,
becoming sessile, 1–2 per areole, aggregated but regularly rounded, rarely crowded; disc plane to subconvex,
dull orange, brownish yellow to brownish orange or somewhat orange-pink, ± with sparse whitish pruina;
biatorine in section with a slightly paler proper margin, 0.03–0.05 mm wide, becoming excluded or very
rarely zeorine; thalline exciple developed only at the base, 70–80 µm thick, cortical layer 10–15 µm thick;
hymenium 50–60 µ m high, hyaline, epihymenium brownish orange; subhymenium 80–100 µ m thick;
paraphyses swollen towards the tips to 4-6(–7) µm diameter, usually with oil droplets 1–2(–4) µm diameter,
very rarely with oil cells 5–6(–7) µm diameter present; asci (1–2–4–6–)8-spored, bipolar and simple spores
seen in the same ascus; ascospores distinctly widened at the septum, with rounded ends, varying from
elongate-ellipsoid to almost spherical, (9–)10–12(–15) × 6–7 µm in water and 10–13(–14) × (5–)6–7(–8) µm
in K (often with wrinkled surface in K), and rather narrow septa, 3–4(–6) µm wide in water and (4–)5–7 in K.
Secondary chemistry: thallus K+ purple, epihymenium K+ purple or blackish purple in places, then dark
cherry blossom pink to dark cherry blossom red; containing parietin (major), parietinic acid (minor), fallacinal
(trace), teloschistin (trace).
Distribution and habitat:—Common on limestone outcrops in Western Australia, South Australia,
Victoria and Tasmania.
Etymology:—This species is named after the Russian biologist, Konstantin Sergejewicz Mereschkowsky
(1855–1921), known for his hypothesis on symbioses, who described C. brachyspora belonging to this
species group.
Additional specimens examined (paratypes):—AUSTRALIA. Western Australia: E. Wallabi, N end of
airstrip WA, April 1980, Green s.n. (PERTH). Nullarbor Plain, Eyre Highway, near Eucla Pass, 1 km W of
Eucla, Curnow 4753 (CANB). N of Northampton, road to Horrocks, Bower River Road, Kärnefelt 20041502
(LD). 0.25 km E of mouth of Bower River, S of Horrocks Beach, Kondratyuk et al. 20421 (KW). South
Australia: Salt Creek, Bratt 67/138 (HO 64728). 15 km W of Peake along Highway 12, Elix 8727 (MEL).
Eyre Peninsula, 28.3 km E of Cowell on the Lincoln Highway, Filson 11795 (MEL). Nullarbor Homestead,
January 1952, Kempsey s.n. (MEL). Victoria: Buchan-Gelantipy road, 6 km NNE of Buchan, Streimann
39787 (CANB). Tasmania: Bass Strait, Hogan Group, Hogan Island, 4 m, December 1973, Whinray s.n.
(MEL).
Caloplaca mereschkowskiana belongs to the Caloplaca lactea group, characterized by a whitish thallus
and yellow apothecia with small ascospores. It is similar to C. lactea (A.Massal.) Zahlbr. from which it differs
by wider ascospore septa and absence of a well-developed thalline margin of the apothecia (Kondratyuk et al.
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2004). The whitish thallus of C. mereschkowskiana resembles C. brachyspora Mereschk., a species that
differs in having less aggregated apothecia, in lacking oil droplets and oil cells in the upper portion of
paraphyses and in having narrower ascospore septa and the presence of a dark grey hypothallus (Kondratyuk
et al. 2004). Further, C. mereshkowskiana differs from C marmorata (Bagl.) Jatta and C. lacteoides Nav.-Ros.
& Hladun, two species growing on limestone outcrops with reddish apothecia, by its smaller ascospores
(Kondratyuk et al. 2004). Caloplaca yorkensis often grows together with C. mereschkowskiana but differs in
having narrower ascospores and a more immersed thallus and apothecia without the pink orange-pink tinge
observed in C. mereschkowskiana.
Caloplaca yorkensis S.Y.Kondr. & Kärnefelt, sp. nov. (Fig. 7B) Mycobank MB 517748
Caloplacae lacteae similis, sed thallo tenuissimo–crassiusculo, albido-cinereo, apotheciis biatorinis, obscure luteis,
subhymenio minus evoluto et ascosporis angustissimis ((3.5–5(–5.5[–6]) µm latis) differt.—http://www.eol.org/
pages/Caloplaca yorkensis
Type: AUSTRALIA. South Australia: Yorke Peninsula, Coodowie, 5 km E of Editburgh along the coast,
35°03’S, 137°44’E, on calcareous rocks, growing together with Caloplaca johnwhinrayi, January 1999,
Kärnefelt 995207 (holotype CANB).
Thallus (0.3–)1–2 cm wide or aggregated into larger colonies forming grey to greyish brown, somewhat
indistinct areas of different thickness, from indistinct, almost endolithic or very thin and immersed in the
substrate, cracked together with the substrate forming areoles, 0.4–0.7 mm wide, to rather thick and welldeveloped, (to 0.3 mm thick), plane or subconvex, immersed to ± exfoliating; areoles grey or greyish white to
brownish grey with sparse, often minute, biatorine, dull yellow orange apothecia. Apothecia to 0.3–0.6(–0.7)
mm diameter, 0.2–0.25 mm thick in section, biatorine, usually scattered and regularly rounded, rarely
irregular due to mutual pressure; proper margin pale yellowish or concolorous with disc, becoming excluded;
disc plane to subconvex, dull yellow orange to brownish yellow-orange, in section biatorine, true exciple 30–
50 µm thick in uppermost lateral part and to 40 µm thick in lower lateral part, 30–70 µm thick at the base,
pseudoprosoplectenchymatous to somewhat mesodermatous paraplectenchymatous with a well-developed
matrix in the centre; hymenium 50–60 µm high; subhymenium 20–25(–70) µm thick; epihymenium bright
yellow; paraphyses 1.5–2 µm diameter, distinctly swollen to 4–5 µm diameter towards the tips; asci 2–4–6–8spored, often mature bipolar and simple undeveloped or abortive ascospores present; ascospores small and
very narrow, slightly swollen at the septa, 9–11(–12) × 3.5–5(–6) µm in water and (8–)9–12 × (4–)4.5–6(–7)
µm in K, with rather thin septum, (1.5–)2–3.5 µm wide in water and 2.5–3.5(–4) µm wide in K. Secondary
chemistry: epihymenium K+ purple to crimson purple, becoming brownish crimson on standing.
Distribution and habitat:—Caloplaca yorkensis is known from scattered localities in Western Australia,
South Australia, Victoria and Tasmania, where it occurs on limestone outcrops.
Etymology:—This new species is named after the type locality, Yorke Peninsula in South Australia.
Additional specimens examined (paratypes):—AUSTRALIA. Western Australia: Point Peron, ca. 80
km S of Perth, Marchant 5578 (MEL 234720). 5 km S of Dongara, Kärnefelt et al. 20041103 (LD). 0.25 km E
of mouth of Bow River, S of Horrocks Beach, Kondratyuk et al. 20421 (CANB, LD, KW, PERTH). Victoria:
Portland region, Cape Bridgewater, Kärnefelt 996101 (LD). Tasmania: Bass Strait, Hogan Group, Hogan
Island, 4 m, December 1973, Whinray s.n. (MEL). Bass Strait, Furneaux Group, Kangaroo Island, ca. 90 m
SSW of the summit, February 1972, Whinray s.n. (MEL).
Caloplaca yorkensis, with its grey to greyish brown thallus and yellow to orange apothecia with small
ascospores resembles species in the Caloplaca lactea group. This new species is similar to C. lactea (A.
Massal.) Zahlbr., but differs in having a thinner and more immersed thallus. The subhymenium is thinner in C.
yorkensis, the ascospores considerably narrower (3.5–5.5 µm vs, 6–7 µm wide), and the apothecia lack a welldeveloped thalline margin. Caloplaca brachyspora Mereschk. has a similar morphology, and also grows on
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limestone However, the thallus is better developed and greenish-brown in C. brachyspora and the ascospores
are broader (6–8 µm; Kondratyuk et al. 2004). Caloplaca yorkensis differs from Caloplaca lithophila by its
grey, mainly immersed thallus and smaller, narrower ascospores. Caloplaca oxfordensis Fink and C.
mereschkowskiana, described above, differ in having orange apothecia and larger or broader ascospores.
FIGURE 7. A. Caloplaca mereschkowskiana (holotype), general habit. B. Caloplaca yorkensis (MEL 234720), general
habit. C. Calvitimela uniseptata (holotype), thallus with apothecia. (photograph by Vítězslav Maň ák). D. Chapsa
microspora (holotype), thallus with apothecia. E. Chapsa psoromica (isotype), thallus with apothecia. F. Chapsa
rubropulveracea (holotype), apothecium. Scale in A–E = 1 mm, in F = 0.5 mm.
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Calvitimela uniseptata G.Thor, sp. nov. (Fig. 7C) Mycobank MB 517749
Thallus squamulosus, squamulis adscendentibus, verrucoso-coralloideis, cinereis vel cinereo-viridibus. Ascosporae
hyalinae, subcurvatae usque ad ellipsoideae, 1-septatae vel raro non septatae, 9–16 µm longae et 4–5 µm latae.—
http://www.eol.org/pages/Calvitimela uniseptata
Type:—ANTARCTICA. Dronning Maud Land: Vestfjella, the nunatak Basen, 1000 m NW of the Swedish
station Wasa, 73°02’S, 13°25’W, 505 m, steep SW facing slope 50 m E of the precipice, at old Pagodroma
nivea nest, February 1992, Thor 10792 (holotype S, isotype UPS).
Thallus of erect, wart-like to coralloid squamules, grey to greenish grey, not pruinose; prothallus diffuse
to clearly visible, whitish; medulla whitish to greyish, with calcium oxalate crystals, C–, K–, PD–, UV–, I–,
K/I–; hyphae 2–4 µm wide; soralia not seen. Photobiont chlorococcoid, cells 5–12 µm in diameter. Apothecia
frequent, clearly constricted at base, up to 1.1 mm in diameter; margin thin to thick, often partly lacking, of
the same colour as the thallus; disc flat to convex, black to brown to colourless with pink tinge (in deep
shade), not pruinose; hypothecium colourless; hymenium colourless, without oil-drops, 40–70 µm tall, I+
blue, K/I+ blue, K–; epithecium colourless to dark olive-green, N+ reddish violet; paraphyses sparsely
branched, thin-walled, without distinct gallertic sheaths, 2 µm wide; paraphyses apices colourless to dark
olive-green, 3–5 µm wide. Asci of the Lecanora-type, 50–55 × 15–20 µm. Ascospores 8/ascus, colourless,
slightly curved to ellipsoid, without perispore, 1-septate or rarely simple, (9–)10–13(–16) × (4–)4–5(–5) µm
(Length: X = 11.7 µm, SD = 1.4 µm, n = 120; Width: X = 4.8 µ m, SD = 0.5 µm, n = 120). Pycnidia few,
immersed; wall colourless; conidia thread-like, curved (14–)17–21(–22) × (1–)1(–1) µm (Length: X = 18.6
µ m, SD = 2.0 µm, n = 120; Width: X = 1.0 µm, SD = 0.0 µ m, n = 120). Secondary chemistry (TLC): no
substances.
Distribution and habitat:—Calvitimela uniseptata is only known from the type locality, where it was
abundant. It was searched for on numerous other localities in Heimefrontfjella and Vestfjella without success.
The species grows on basalt rocks and on adjacent old snow petrel Pagodroma nivea feathers at an old snow
petrel nest under a boulder. Thus, the locality was probably rich in nitrogen and phosphorous.
Etymology:—The new species is named after the 1-septate ascospores.
The genus Calvitimela is separated from Tephromela on the bases of, e.g., the greenish epithecium, which
reacts N+ red, and asci of the Lecanora-type (Hafellner & Türk 2001). Tephromela s. str. is distinguished by a
reddish brown hymenium, reacting K+ violet and by having asci of the Bacidia-type (Hafellner & Türk 2001).
Calvitimela uniseptata is recognized by the wart-like to coralloid thallus squamules and the 1-septate spores.
The immersed and colourless pycnidia are easily overlooked. The species is not included in Hertel (1984,
1987) or Øvstedal & Lewis Smith (2001).
Chapsa microspora Kalb, sp. nov. (Fig. 7D) Mycobank MB 517750
Similis Chapsae aggregatae sed differt thallo non corticato, ascosporis minoribus, hyalinis et materia chemica.—http://
www.eol.org/pages/Chapsa microspora
Type:—BRAZIL. Amazonas: Rainforest along Rio Negro, 150 km upstream Manaus, 02°30’S, 61°10’W, 40
m, October 1980, Kalb s.n. (holotype hb. Kalb 26934).
Thallus corticolous, light grey to whitish, smooth, continuous, ca. 20–50 µm thick, dull, ecorticate to
endoperidermal. Photobiont layer indistinct, 10–15 µ m thick, with numerous photobiont cells and the
inclusion of decomposed periderm cells and of a few small calcium oxalate crystals. Medulla endophloeodal.
Apothecia erumpent to prominent, roundish, 0.6–1.2 mm in diameter, usually 1–3 aggregate. Margin strongly
raised, deeply fissured to lobed, upright to usually strongly recurved, with a thin, pale-grey to white-felty and
sometimes slightly crystalline inner surface; proper exciple free. Disc pale brown, covered by a thin to rather
thick white pruina. Periphysoids distinct, ca. 20 µm long. Hymenium 50–60 µm high, clear. Epihymenium
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unpigmented or partially greyish, ca. 5–8 µ m high. Paraphyses simple, straight, 2–2.5 µ m wide; tips
indistinctly moniliform to occasionally branched, adspersed with fine greyish granules. Ascospores 6–8/
ascus, 2–3 seriate, hyaline, with (2–)3 transverse septae, 7–9 × 4 µm, thick-walled, with rounded ends, I–.
Pycnidia not seen. Secondary chemistry (TLC): stictic acid (major), constictic acid (major), hypostictic acid
(trace) and hypoconstictic acid (trace).
Distribution and habitat:—Chapsa microspora is known from a single but ample collection from a
more or less virgin tropical rainforest in the Amazon basin. It was collected from the smooth and slightly
decaying bark of an old deciduous tree in a rather shady situation.
Etymology:—The epithet refers to the relatively small ascospores.
Chapsa microspora is a conspicuous species being characterised by rather large roundish and aggregate
apothecia with a clear hymenium, a free proper exciple, paraphyses with indistinctly moniliform or slightly
branched tips and hyaline, transversely septate, small, non-amyloid ascospores with a well-developed
endospore. The thallus is pale grey to whitish and contains substances of the stictic acid aggregate. C.
aggregata (Hale) Sipman & Lücking from Dominica looks similar, but that species is distinctly corticate,
without secondary lichen products, and has brown ascospores, ca. 16 × 8 µm. Chapsa albomaculata (Sipman)
Sipman & Lücking also has aggregate apothecia, but the thallus is corticate and the ascospores are more
septate with (4–)5–7(–8) septa, and the chemistry is slightly different [constictic acid (major), stictic acid
(trace)]. Chapsa platycarpa (Tuck.) A.Frisch, another species with stictic acid aggr., has an epiperidermal,
olive-green thallus with a prosoplectenchymatous cortex and brown ascospores. The same chemistry and a
similar habitus as C. microspora, together with a free exciple, is found in Thelotrema porinoides Mont. &
Bosch, but that species differs by large, multiseptate, I+ violet ascospores. Another species of Chapsa with
very small (4.5 × 2.5 µm) and I– ascospores, namely C. bicellularis Sipman & Lücking, has a clear hymenium
too, but differs in the absence of secondary lichen products and the hardly discernable thallus.
Chapsa psoromica M.Cáceres, Santos de Jesus & Santos Vieira, sp. nov. (Fig. 7E) Mycobank MB 517751
Sicut Chapsa elabens sed thallo olivaceo corticato, acidum psoromicum continente et ascosporis I+ violaceis differt.—
http://www.eol.org/pages/Chapsa psoromica
Type:—BRAZIL. Rondônia: Porto Velho, Estação Ecológica de Cuniã, 8º 04’S, 63º 31’W, 100 m, July 2009,
Cáceres 7595 (holotype ASE, isotype F).
Thallus corticolous, up to 5 cm diameter, 70–100 µm thick, continuous; surface smooth to uneven, olivegreen. Thallus in section with well-developed, prosoplectenchymatous upper cortex with internal splitting and
irregular algal layer; clusters of calcium oxalate crystals absent. Apothecia erumpent, angular-rounded, 0.4–
0.6 mm diameter; disc more or less covered by a narrow pore, flesh-coloured and slightly translucent, nonpruinose; proper margin indistinct; thalline margin fissured to lobulate with 3–6 inclined to erect lobules.
Columella absent. Excipulum paraplectenchymatous, 30–60 µ m thick, yellow; periphysoids present but
indistinct. Hymenium 150–200 µm high, strongly inspersed and appearing nubilous grey; inspersion of two
types: primary inspersion quickly dissolving in KOH and then hymenium appearing translucent, but
secondary inspersion persistent; paraphyses unbranched. Ascospores 8 per ascus, 9–11-septate, 35–45 × 9–12
µm, oblong, with thickened septa and lenticular lumina, colourless, I+ dark violet-blue. Secondary chemistry:
psoromic acid.
Distribution and habitat:—This species is thus far only known from the type collection from the state of
Rondônia in the western part of the Amazon forest. It appears to be a forest understory species on the bark of
large trees.
Etymology:—The epithet refers to the, for this genus, unusual chemistry.
Chapsa psoromica at first glance resembles a species of Fissurina because of its olive-green, corticate
thallus with a split-cortex and fissured apothecia. However, the apothecia are rounded with a well-developed
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disc and therefore the species is here placed in Chapsa. Inspersion is very rare in Chapsa and only three
species with hyaline, transversely septate ascospores show this feature: C. elabens (Müll.Arg.) Rivas Plata &
Mangold, C. lueckingii Kalb, and C. pseudoschizostoma (Hale) Sipman (Kalb 2009, Rivas Plata et al. 2010).
The latter has an ecorticate, whitish thallus and much smaller ascospores; also, the apothecia are
pseudocolumellate. Chapsa lueckingii produces stictic acid and has much larger apothecia with thick white
pruina and smaller ascospores. Chapsa elabens differs from C. psoromica in the ecorticate, endoperidermal
thallus and the non-amyloid, narrower ascospores with more numerous septa. Both species also lack psoromic
acid. Chapsa psoromica resembles other fissurinoid species in the genus, namely C. dissuta (Hale) Mangold,
C. lassae Mangold, and C. zahlbruckneri (Redinger) A.Frisch. The first two are externally almost
indistinguishable but have a non-inspersed hymenium and much smaller, 3-septate ascospores, and lack
psoromic acid. The latter agrees with C. psoromica especially in the indistinct periphysoids, but its hymenium
is also non-inspersed and the ascospores are muriform; in addition, the apothecia are much larger and more
distinctly chroodiscoid; also here, psoromic acid is lacking. The unusual psoromic acid chemistry may
suggests that C. psoromica is not a genuine Chapsa; however, in the related genus Thelotrema, there is also a
single species with psoromic acid, T. saxicola (Vain.) Salis. (Rivas Plata et al. 2010).
Chapsa rubropulveracea Hale ex Mangold, Lücking & Lumbsch, sp. nov. (Fig. 7F) Mycobank MB 517753
Sicut Chapsa farinosa sed apotheciis rubris ascosporis minoribus differt.—http://www.eol.org/pages/Chapsa
rubropulveracea
Type:—DOMINICA. Barbers Block, May 1972, Hale 46121 (holotype US).
Thallus corticolous, up to 5 cm diameter, 50–100 µm thick, continuous; surface distinctly farinose, whitegrey. Thallus in section lacking upper cortex, with irregular algal layer and abundant clusters of calcium
oxalate crystals. Apothecia erumpent, angular-rounded, 0.5–1 mm diameter; disc exposed, red-pruinose;
margin lobulate, fused, white-pruinose. Columella absent. Excipulum paraplectenchymatous, 10–20 µm thick,
colourless; periphysoids present. Hymenium 50–70 µm high, with a layer of crystalline red pigment above
which dissolves in KOH; paraphyses unbranched. Ascospores 8 per ascus, 5–7-septate, 15–20 × 5–6 µ m,
oblong, with thin to slightly thickened septa and rectangular lumina, colourless, I–. Secondary chemistry: no
substances detected with TLC except red pigment on apothecial disc (K+ purple).
Distribution and habitat:—This species is thus far only known from the type collection from Barber's
Block, a 400 m high hill about 8 km N of Grand Savanna on Dominica.
Etymology:—The epithet refers to the red-pruinose apothecial disc.
Chapsa is a large genus of chiefly tropical, crustose lichens in Graphidaceae, usually with large apothecia
bordered by a lobulate margin (Rivas Plata et al. 2010). The genus is quite variable morphologically,
chemically, and ecologically, although most species are found in lowland to lower montane rain forests.
Chapsa rubropulveracea was intended to be described as a new Thelotrema by the late Mason Hale, but
apparently never published, and is validated here in the genus Chapsa. There are only few other species of
Chapsa with transversely septate, hyaline ascospores and pigmented apothecial discs. Chapsa waasii (Hale)
Sipman & Lücking from southeast Asia agrees in most aspects, but has apothecia with a purple-violet disc and
distinct, recurved, felty-pruinose lobules, as well as an endoperidermal thallus (Rivas Plata et al. 2010).
Chapsa rubropruinosa Messuti & Codesal from Argentina (Messuti et al. 2010) differs in the red-brown
apothecial disc and the loosely corticate thallus, as well as in the presence of stictic acid. Chapsa rivas-platae
Kalb & Lücking from Brazil (Kalb 2009) is another similar species; it also lacks a cortex but the apothecia are
ochraceous to orange-pruinose, the ascospores are much larger (45–80 µm) and I+ violet-blue, and stictic acid
is present. Finally, there is C. magnifica (Berk. & Broome) Rivas Plata & Mangold, with a dense thallus
cortex, inclined to erect apothecial lobules, and large, multiseptate ascospores (70–100 µm). Another species
of Chapsa with pigmented apothecial disc, C. neei (Hale) Mangold & Lücking, differs in having brown
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ascospores and lepadinoid apothecia with a double margin. Except for the apothecial pigment, C.
rubropulveracea is most similar to C. farinosa Lücking & Sipman from Costa Rica (Sipman et al. 2010),
which agrees in the farinose thallus and general apothecial morphology, but lacks a pigment and has larger
ascospores (20–30 µm).
Chapsa thallotrema Lücking & N.Salazar, sp. nov. (Fig. 8A–B) Mycobank MB 517753
Sicut Chapsa sublilacina sed thallo sorediis instructo differt.—http://www.eol.org/pages/Chapsa thallotrema
Type:—PANAMA. Panamá: Altos de Campana National Park, 50 km W of Panama City near the town of
Capira, 8º 42’N, 79º 57’W, 500–600 m, submontane rainforest, on bark of large trees in the shaded understory,
Lücking 27305 (holotype F, isotype PMA).
Thallus grey-olive, smooth to uneven, with dense, prosoplectenchymatous cortex with internal splitting,
sorediate; soralia usually numerous but discrete, capitate, finely granular, 1–2 mm diameter, white to bluish or
yellowish white, strongly contrasting with the surrounding thallus; photobiont layer and/or medulla with
clusters of calcium oxalate crystals. Apothecia erumpent, angular-rounded, 1–3(–5) mm diameter; disc
exposed, flesh-coloured to pink-purple; margin lobulate to recurved, often layered, flesh-coloured to beige.
Columella absent. Excipulum paraplectenchymatous, colourless; periphysoids present. Hymenium 150–200
µm high; paraphyses unbranched. Ascospores 2–4/ascus, 15–19-septate, 60–120 × 10–15 µm, cylindrical,
with thick septa and lens-shaped lumina, colourless, I+ violet-blue (amyloid). Secondary chemistry: stictic
acid and satellites (K+ yellow).
Distribution and habitat:—Apparently a widely distributed neotropical species restricted to the shady
understory of lowland to lower montane rainforests.
Etymology:—This common and widespread taxon was long ago recognized by Henry Imshaug and his
students from collections in the Caribbean and provisionally named ‘Thallotrema antillarum’ but was never
validly published. The chosen epithet honours this earlier name.
Additional specimens examined (paratypes):—COSTA RICA. Alajuela: Volcán Tenorio National Park,
Pilón Biological Station (Arenal-Tempisque Conservation Area), Tilarán Ridge, 140 km NW of Sán José, 25
km NNW of Tilarán, near Bijagua, Nelsen 3745 (INB, WIS), Lücking 17219 (F); ibid., main trail through
forest to crater, Will-Wolf 12745b (CR, INB, WIS). Puntarenas: Corcovado National Park, Sirena Section,
Sirena Biological Station (Osa Conservation Area), Osa Pensinsula, 160 km SSE of Sán José and 50 km
WSW of Golfito, Lücking 16204 (F, USJ); 16228 (F, INB). PANAMA. Panamá: Altos de Campana National
Park, 50 km W of Panama City near the town of Capira, Lücking 27303, 27304 (F, PMA).
This new species is part of a complex of species centered around Chapsa sublilacina (Ellis & Everh.)
Sipman & Lücking, all sharing the same thallus morphology and ascospore type. Four forms are
distinguished: with or without stictic acid and satellite substances, and with either large, compact, finely
granular soralia or with small, irregular, coarse soralia. The form with large soralia, described here as C.
thallotrema, contains stictic acid, whereas the form with small soralia lacks secondary substances. Thus, C.
thallotrema is considered the sorediate counterpart of C. sublilacina.
Chiodecton pustuliferum Aptroot, sp. nov. (Fig. 8C–D) Mycobank MB 517754
Chiodecton thallo sterili pustulato, pustulis sorediatis albidis, hypothallo extenso brunneo, acidum roccellicum
continens.—http://www.eol.org/pages/Chiodecton pustuliferum
Type:—MADAGASCAR. Tamatave: Périnet, Andasibe, 18º 56’S, 48º 16’E, May 1984, Aptroot & Hensen
13406 (holotype ABL).
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FIGURE 8. A–B. Chapsa thallotrema (holotype). A. Thallus with apothecia. B. Thallus with soralia. C–D. Chiodecton
pustuliferum (holotype). C. Thallus with soralia. D. Hyphae of hypothallus. E–F. Cladonia mongkolsukii (holotype). E.
Thallus with podetia. F. Thallus with podetia. Scale in A–C = 1 mm, in D = 50 µm, in E, F = 10 mm.
Thallus corticolous, crustose, pale brownish along the margins, whiter at the centre of the areoles, dull,
consisting of ca. 0.4–4 mm diameter crenate thallus areoles of irregular surface and outline on a continuous
layer of brown arachnoid hypothallus that is underlying the whole thallus, covering areas of up to 7 cm
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diameter, margins delimited by a continuous hypothallus, forming a line of up to 1 mm. Hypothallus filaments
2–3 µm wide, surface partly rough from crystals. Pustules numerous, erumpent, starting round, up to 3.0 mm
diameter and generally ca. 1.0 mm high, scarcely confluent, white (contrasting with the thallus), globular to
irregular. Soredia formed inside the pustules, granular, ca. 100–200 µ m diameter, partly clustering into
aggregates, internally formed of branched and gnarled hyphae with copious crystals. Algae trentepohlioid,
cells ellipsoid, ca. 10 × 15 µm, identical in thallus and soredia. Apothecia and pycnidia unknown. Secondary
chemistry: roccellic acid.
Distribution and habitat:—The new species is only known from the type from primary tropical
mountain forest at 950 m elevation in Madagascar.
Etymology:—The epithet refers to the pustules.
Chiodecton is a small genus in the family Roccellaceae characterized by stromatic ascomata with
punctiform discs, superficially resembling the perithecial stromata of Trypethelium or other pyrenocarpous
taxa (Thor 1990). This species is the first Chiodecton to be described with pustules (Thor 1990), and even in
the order Arthoniales pustules were previously unknown. Although the material is sterile, the dull, yet not
byssoid thallus on the arachnoid hypothallus and the presence of roccellic acid, make a classification in
Chiodecton possible.
Cladonia mongkolsukii Parnmen & Ahti, sp. nov. (Fig. 8E–F) Mycobank MB 517755
Cladoniae awasthianae similis sed podetiis esorediatis et microsquamulosis differt.—http://www.eol.org/pages/Cladonia
mongkolsukii
Type:—THAILAND. Pitsanulok Province: State Authority Agency areas, Phu Hin Rong Kla National Park,
16°59’N, 100°60’E, 1110 m, on rocks in lower montane scrub forest, May 2005, Parnmen SP271RAMK
(holotype RAMK, isotype H).
Primary thallus persistent, irregularly lobed to deeply laciniate, 0.7–1.8 mm long, 0.2–0.8 mm wide,
lower surface white, ecorticate, upper surface greenish, corticate; podetia 10–65 mm tall, 0.3–3 mm thick,
clavate, whitish grey to brownish, unbranched or slightly branched at apex, blunt, at first ascyphose but soon
becoming tipped with very narrow scyphi, 1–3 mm in diameter, 6–7 tiers of scyphi produced per podetium;
podetial wall 87.5–150 µ m: medulla 12.5–50 µ m, stereome 70–100 µ m; surface rough, ecorticate to
discontinuously corticate with microsquamules, especially densely squamulate near tips; podetial squamules
0.6–2 mm long, 0.2–0.5 mm wide, upper part green, corticate, lower part whitish, ecorticate; apothecia pale
brown to orange or brownish, terminal on podetia; ascospores simple, hyaline, oblong to ellipsoid, (6.4–)9–
9.5–10(–14) × (1.6–)2.7–2.9–3.1(–4.2) µm, 8/ ascus; conidiomata not seen. Secondary chemistry: podetia: P+
orange turning red, K± yellow, C–, KC–, containing fumarprotocetraric acid (major) and homosekikaic acid
(major).
Distribution and habitat:—Cladonia mongkolsukii is known from lower montane scrub forests in
northeast Thailand.
Etymology:—The specific epithet honors the Thai lichenologist Pachara Mongolsuk (Bangkok).
Additional specimens examined (paratypes):—THAILAND. Pitsanulok Province: Phu Hin Rong Kla
National Park, Lan Hin Pum, July 2002, Dangphui s.n. (RAMK); ibid., the office of the National Park and
guesthouse, 400 m, February 2003, Dangphui s.n. (RAMK); ibid., the sideway to Ban CP house and the State
Authority Agency, February 2003, Buarueng s.n. (RAMK); ibid., natural trail 100 m from car park, February
2003, Chaiyabutr s.n. (RAMK); ibid., Lan Hin Pum, February 2003, Homchantara & Diangsa s.n. (RAMK);
ibid., the State Authority Agency areas, June 2003, Homchantara s.n. (RAMK); ibid., natural trail 100 m from
car park, June 2003, Chaiyabutr s.n. (RAMK); ibid., Pha Chu Thong, June 2003, Homchantara & Diangsa
s.n. (RAMK); ibid., the sideway to Lan Hin Pum, June 2003, Dangphui s.n. (RAMK); ibid., natural trail about
10 m from the State Authority Agency, June 2003, Chaiyabutr s.n. (RAMK); ibid., areas between the air raid
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shelter of the State Authority Agency, June 2003, Homchantara & Diangsa s.n. (RAMK); ibid., along the
sideway to Lan Hin Teak, June 2003, Mongkolsuk s.n. (RAMK); ibid., along the sideway to Hmun Daeng
waterfall, March 2004, Parnmen s.n. (RAMK); ibid., kilometer post 28, April 2004, Parnmen s.n. (RAMK);
ibid., the State Authority Agency areas, May 2005, Parnmen s.n. (RAMK); ibid., Phu Luang Wildlife
Sanctuary, August 2005, Parnmen s.n. (RAMK); ibid., the sideway to TV station, August 2005, Parnmen s.n.
(RAMK).
Morphologically this new species resembles the Himalayan Cladonia awasthiana Ahti & Upreti (2004),
but that species differs in having sorediate podetia and microsquamules that are distributed over the entire
length of the podetia and not densely squmulose near the tips. In our phylogenetic analysis (Parnmen et al.
2008) based on ITS sequence data, C. mongkolsukii [as cf. awasthiana; specimens SP271RAMK (EU113290)
and SP283RAMK (EU113289)] was placed near Cladonia singhii Ahti & Dixit in the same clade with
Cladonia fimbriata (L.) Fr., Cladonia gracilis (L.) Willd. subsp. gracilis and Cladonia ochrochlora Flörke.
Stenroos et al. (2002) placed the last three species in their “supergroup” Cladonia.
Clypeopyrenis porinoides Komposch, J.E.Hern. & Rosabal, sp. nov. (Fig. 9A) Mycobank MB 517756
Sicut Clypeopyrenis microsperma sed peritheciis thallo obtectis differt.—http://www.eol.org/pages/Clypeopyrenis
porinoides
Type:—COSTA RICA. Puntarenas: Sán Vito de Coto Brus, Las Cruces Biological Station, 82°58’W, 8°47’N,
1200 m, on ridge beyond Río Java, on trunks and undergrowth of primary forest, September 2007, Komposch
et al. A6J2T3 (holotype INB, isotype F)
Thallus corticolous, up to 10 cm diameter, continuous; surface smooth to uneven, light greyish green.
Thallus in section with thick, paraplectenchymatous upper cortex and thick algal layer. Perithecia solitary or
2–3 aggregate, erumpent to prominent, wart-shaped, 0.4–0.6 mm diameter, 0.3–0.4 mm high, completely
covered by thalline layer up to the ostiole, light greyish green to yellowish green, ostiolar area with red-brown
to dark brown spot. Involucrellum and excipulum not well separated, basally up to 25 µm thick and laterally
and apically 50–100 µm thick, fully carbonized; covered by corticate algiferous thallus up to the ostiole;
hypothecium prosoplectenchymatous, 5–10 µm high, colourless; hamathecium colourless, clear, I–, KI–.
Paraphyses unbranched, glabrous; periphyses not observed. Asci clavate, 40–50 × 6–8 µm. Ascospores 8 per
ascus, irregularly arranged, ellipsoid with rounded ends, 1-septate, with slight constriction at septum, 5–6 ×
2.5–3 µ m, 2 times as long as wide, with slightly thickened septum and angular lumina, dark brown, I–.
Secondary chemistry: no substances detected by TLC.
Distribution and habitat:—The new species was found abundantly in the understory of a primary rain
forest at mid elevation but is not known outside that area.
Etymology:—The species closely resembles a species of Porina.
Additional specimens examined (paratypes):—COSTA RICA. Puntarenas: Sán Vito de Coto Brus, Las
Cruces Biological Station, Komposch et al. A4AT1, A6HT1, A7J2T1 (F).
Clypeopyrenis was proposed as a monospecific genus in Pyrenulaceae by Aptroot (1991), to
accommodate a species with very small, dark brown ascospores without distinct endospore, C. microsperma
(Müll.Arg.) Aptroot. The type material originates from Costa Rica, very close to the locality of the new
species, but C. microsperma is also known from the Caribbean and South America. While C. microsperma is
characterized by completely exposed, black perithecia with rather massive involucrellum (clypeus), resting on
a dark green to olive thallus, the new species, C. porinoides, has more delicate perithecia completely covered
by a thalline layer up to the ostiole. It therefore closely resembles a Porina, and only microscopical
examination will reveal the fundamental anatomical differences. Besides the morphological differences,
Clypeopyrenis porinoides also has smaller asci and ascospores than C. microsperma (the smallest known to
date in the family), although at this size, it is difficult to ascertain whether these differences are significant.
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The excipulum and involucrellum are also more delicate (up to 300 µm thick in C. microsperma). At the type
locality, both species grew together, although C. porinoides was much more abundant, and the morphological
differences were easily observed. The new species was collected during the first OTS (Organization for
Tropical Studies) course on tropical lichenology held at Las Cruces Biological Station in 2007.
FIGURE 9. A. Clypeopyrenis porinoides (holotype), thallus with perithecia. B. Coccocarpia delicatula (holotype), part
of thallus with isidia. C–D. Coenogonium flammeum (holotype), thallus with apothecia. E–F. Cresponea
ancistrosporelloides (holotype). E. Detail of type specimen showing apothecioid ascomata and areolate thallus. F. Ascus
with non-tailed spores next to one with tailed spores and paraphysoid tips. Scale in A–E = 1 mm, in F = 25 µm.
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Coccocarpia delicatula Bungartz, Ziemmeck & Lücking, sp. nov. (Fig. 9B) Mycobank MB 517757
Sicut Coccocarpia domingensis sed isidiis cylindricis laminalibus differt.—http://www.eol.org/pages/Coccocarpia
delicatula
Type:—ECUADOR. Galápagos: Isla Sán Cristóbal, area W of Cerro Pelado on the way to El Ripioso,
0°52’S, 89°28’W, 400 m, transition zone, open Psidium guajava shrubland with Macraea laricifolia and
dominant annual herb Malachra capitata, on bark and wood, dead twigs of Psidium guajava, sunny, windand rain-exposed, August 2008, Bungartz 8496 (holotype CDS-41142).
Thallus foliicolous, foliose, rounded, up to 10 mm diameter. Lobes thin, flat, 0.1–0.2 mm wide, linear but
slightly widened towards the apex, isotomically branched and dissected, rather close. Upper surface bluegreen when moist, plumbeous to bluish grey when dry, with thin, longitudinal, white striae. Lower surface
pale, with pale, unbranched, tapering, 0.2–0.4 mm long rhizines partly protruding from under the lobe margins
and visible from above. Lobes isidiate; isidia laminal, dense, uniformly cylindrical, mostly unbranched to
bifurcate, 0.1–0.3 mm long and 0.05–0.1 mm thick, in mature thalli covering most of the thallus interior.
Apothecia not observed. Pycnidia not observed. Secondary chemistry: no substances detected by TLC.
Distribution and habitat:—Thus far only known from the Galápagos Islands, in the transition zone on
Sán Cristóbal Island.
Etymology:—The epithet refers to the delicate thallus compared to the common Coccocarpia palmicola
(Spreng.) Arv. & D.J.Galloway, which has the same type of isidia.
This new species is close to Coccocarpia domingensis Vain. and the recently described C. neglecta
Aptroot & Lücking (Lücking et al. 2007). All three species have very narrow, linear thallus lobes with fine,
white longitudinal striae on the surface, and differ principally in the shape and arrangement of their isidia. In
C. neglecta, the isidia are mostly marginal and rounded-squamiform, whereas C. domingensis has marginal,
cylindrical to often slightly flattened isidia. C. delicatula produces laminal, always cylindrical isidia, identical
to the type of isidia found in C. palmicola. The latter, however, has much broader, rounded thallus lobes.
Coenogonium flammeum L.I.Ferraro, Michlig & Lücking, sp. nov. (Fig. 9C–D) Mycobank MB 517758
Sicut Coenogonium luteum sed apotheciis fusco-rubris differt.—http://www.eol.org/pages/Coenogonium flammeum
Type:—ARGENTINA. Salta: El Rey National Park, March 2007, Ferraro 7977 (holotype CTES, isotype F).
Thallus crustose, corticolous, continuous, thin, smooth, pale olive-grey, 2–5 cm diameter, with
cartilaginous corticiform layer, shiny; white prothallus present. Photobiont Trentepohlia, cells angularrounded, in irregular plates or short threads, 6–10 µm diameter. Apothecia sessile, rounded, 0.6–1.2 mm
diameter and 150–200 µm high; disc at first plane but becoming slightly convex, deep orange-brown to redbrown; margin thin, not prominent, minutely denticulate (especially when young), creme-coloured.
Excipulum paraplectenchymatous with radiating cell rows in outer parts, 70–150 µm broad, colourless to pale
orange peripherally, I+ sordid yellow-brown; outermost cells elongated and thin-walled. Hypothecium 15–25
µm high, colourless to pale yellowish. Hymenium 90–100 µm high, colourless, I+ (pale) blue then quickly
sordid green, then reddish brown. Asci 70–80 × 5–6 µm. Ascospores uniseriate, ellipsoid, 1-septate, 8–11 ×
2.5–3.5 µm, 3–3.5 times as long as broad. Pycnidia not observed. Chemistry: no substances detected by TLC.
Distribution and habitat:—The new species was found in northwestern Argentina, in montane rainforest
of the "yungas", at localities in the provinces of Salta and Jujuy. The forest is dominated by Myrtaceae,
Lauraceae, and Podocarpus, with a mean annual temperature of 20 ºC and annual precipitation of up to 2000
mm.
Etymology:—The epithet refers to the deep orange-brown to red-brown apothecia.
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Additional specimens examined (paratypes):—ARGENTINA. Salta: El Rey National Park, Ferraro
8018 (CTES). Santa Victoria, Los Toldos, Quebrada El Nogalar, Palací 433 (CTES). Jujuy: Ledesma, Parque
Nacional Calilegua, Ferraro 7519 (CTES).
Coenogonium flammeum is characterized by the unusually dark, deep red-brown apothecia. Anatomically,
including the ascospore size and length to width relation, it comes closest to C. luteum (Dicks.) Kalb &
Lücking, but that species has a pale-orange disc and a thicker apothecial margin, and the apothecia are larger
on average. Another similar taxon is C. siquirrense f. denticulatum Rivas Plata & Lücking (Rivas Plata et al.
2006), which agrees in the size of the apothecia and the minutely denticulate margin, but the disc is vividly
orange with a darker margin and the ascospores are smaller on average.
Cresponea ancistrosporelloides Sparrius & Sipman, sp. nov. (Fig. 9E–F) Mycobank MB 517759
Cresponea saxicola apotheciis parvis 0.3-0.6 mm diametro, ascosporis (6-)8-locularibus basaliter frequenter uncinatis
et torsivis, 35–50 x 5 µm.—http://www.eol.org/pages/Cresponea ancistrosporelloides
Type:—AUSTRALIA. Western Australia: Trail to Toolbrunup Peak, Stirling Ranges, Stirling Range National
Park, 40 km SW of Borden, 118°3’E, 34°23’S, 740 m, dry sclerophyll forest with pockets of denser shrub
vegetation, on volcanic rocks, September 1994, Elix 41495 (holotype B, isotype CANB).
Thallus crustose, saxicolous, over 10 cm wide, flat, pale grey, finely areolate, up to 150 µm thick; areoles
irregular in size and outline, 0.1–0.6 mm wide, separated by thin and irregularly winding fissures; cortical
layer up to 30 µm, of dense, winding hyphae; medulla ca. 100 µm, white, with loose hyphae and ca. 8 µm
diameter, thick-walled trentepohlioid photobiont cells. Apothecia frequent, black, slightly glossy, 0.3–0.6 mm
in diameter, with flat disc and thin margin not extending above the disc. Excipulum dark-brown, strongly
conglutinated and without individual hyphae visible. Hypothecium pale. Hymenium 60–65 µm tall, clear, I+
reddish throughout; paraphysoids 2 µm thick, septate, hardly branched between the asci, apically swollen to
ca. 4 µm and furcate; terminal cell or two uppermost cells clavate, 3–4 × 6–10 µm, covered by a brown layer
with tiny granules. Epithecium dark-brown, ca. 20 µm, composed of densely agglutinated paraphyses tips.
Asci pyriform, ca. 50–60 × 20–22 µm. Ascospores 8/ascus, (6–)8-loculate, fusiform, with thin walls and septa
without thickened edges, at base usually attenuated into a ca. 20 µm long tail which is spiralled in the ascus,
(30–)45–50 × 5 µ m. Pycnidia not seen. Secondary chemistry: thallus K–, C–, KC–, UV–; no substances
found.
Distribution and habitat:—Known so far only from the type collection, from volcanic rock at 740 m in
sclerophyll scrub in West Australia.
Etymology:—The epithet reflects the tailed spores which resemble those of the genus Ancistrosporella.
The genus Cresponea was monographed by Egea & Torrente (1993). Since then only few additional
species were recognized, C. apiculata Egea et al. (1996) and C. litoralis Elix (2007). Cresponea
ancistrosporelloides differs markedly from all known species by the long basal tails of the spores. Only in C.
apiculata are basally attenuated spores reported, but the attenuated part is short and the spores are 11–14loculate. The attenuated spores may be an overlooked feature because they were recently described from
several taxa: Ancistrosporella australiensis (G.Thor) G.Thor (1990, as Ancistrospora), Opegrapha curvata
Aptroot [Aptroot et al. (1997) = Ancistrosporella curvata (Aptroot) Komposch] and Ancistrosporella
psoromica Komposch, Aptroot & Hafellner (Komposch et al. 2002). These species differ in their lirelloid
ascomata and partly byssoid thallus. Also in these species, the asci are much longer than the spores, so that the
spore tails are probably not spiralled below the main body of the spores in the asci, as in C.
ancistrosporelloides. Therefore the new species is provisionally placed in Cresponea rather than in
Ancistrosporella. However, the genus Cresponea seems to be not completely fitting since the ascospores lack
the thick walls and thickened septum edges illustrated by Egea & Torrente (1993).
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Crocynia microphyllina Aptroot, sp. nov. (Fig. 10A–B) Mycobank MB 517760
Crocynia thallo sterili microphyllino albido, hypothallo extenso nigro, acidum protocetraricum continens.—http://
www.eol.org/pages/Crocynia microphyllina
Type:—PAPUA NEW GUINEA. Central Province: Owen Stanley Range, Kagi village, along Kokoda Trail
towards Gap, 9º 08’S, 147º 40’E, October 1995, Aptroot 39483a (holotype ABL).
Thallus corticolous, crustose, pale greenish white, byssoid, dull, consisting of ca. 0.4–2.0 mm diameter
crenate thallus areoles of irregular surface and outline on a continuous layer of black arachnoid hypothallus
that is underlying the whole thallus, covering areas of up to 10 cm diameter, margins delimited by a
continuous hypothallus, forming a line of up to 1 mm. Hypothallus filaments 2–3 µm wide, branching mostly
perpendicular, surface partly rough from crystals. Phyllidia numerous, ascending, starting at the margins of
the areoles, up to 0.3 mm diameter and generally ca. 0.2 mm high, scarcely confluent, pale greenish white (not
contrasting with the thallus), flattened to globular to irregular, internally formed of branched and gnarled
hyphae with copious crystals. Algae chlorococcoid, cells ellipsoid, ca. 5 × 7 µm. Apothecia and pycnidia
unknown. Secondary chemistry: protocetraric acid.
Distribution and habitat:—The new species is only known from the type from primary tropical
mountain forest dominated by Lithocarpus on a ridge at 1700 m elevation in Papua New Guinea.
Etymology:—The epithet refers to the phyllidiate thallus.
Crocynia is a small genus with only two widely accepted species (Cáceres 2007), which are both often
fertile and differ in chemistry, lobe configuration, hypothallus colour, and apothecium characters. This species
is the first Crocynia to be described with phyllidia. Although the material is sterile, the byssoid thallus on the
arachnoid hypothallus and the presence of chlorococoid algae make a classification in Crocynia possible. The
chance that it represents an aberrant morph of one of the known species of Crocynia is low, because the only
other species with a dark hypothallus, C. pyxinoides, is restricted to the Neotropics.
Dictyonema hernandezii Lücking, Lawrey & Dal-Forno, sp. nov. (Fig. 10C) Mycobank MB 517761
Sicut Dictyonema phyllophilum sed thallo et prothallo cartilagineo differt.—http://www.eol.org/pages/Dictyonema
hernandezii
Type:—COSTA RICA. Puntarenas: Las Cruces Biological Station and Wilson Botanical Garden (La Amistad
Pacífico Conservation Area), 8°47’N, 82°57’W, 1200 m, montane rainforest zone, disturbed rainforest
patches, on branches, July 2007, Hernández s.n. (holotype F).
Thallus starting out as small, circular crust but soon covering large areas of the substrate, appressedfilamentous, composed of a strongly compressed mat of densely interwoven, blue-green cyanobacterial
filaments embedded in a yellowish white thallus matrix which forms an often zonate, yellowish white to
brown prothallus free of cyanobacteria; the entire structure appearing gelatinous when wet. Thallus in section
300–500 µm thick, composed of a dominant medullary layer with numerous, interwoven hyphae, above which
is a strongly compacted photobiont layer with more or less periclinally arranged cyanobacterial filaments;
upper part strongly compacted and forming a thin, pale yellow, cortex-like layer. Cyanobacterial filaments
blue-green to yellow-green with cells 10–12 µm wide and 3–5 µm high; surrounded by a thin fungal sheath
forming jigsaw puzzle-shaped cells, the same cells extending above the photobiont layer to form the thin
cortex. Clamps not observed on the medullary hyphae. Basidiocarps not observed.
Distribution and habitat:—The new species is known from two collections in the montane rainforest of
southern Costa Rica. It was found growing in the same habitat as Dictyonema glabratum (Spreng.)
D.Hawksw. and D. sericeum (Sw.) Berk.
Etymology:—We dedicate this new species to our colleague and friend, Venezuelan lichenologist Jesús
Hernández, who also collected the type material.
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Additional specimen examined (paratype):—COSTA RICA. Puntarenas: Las Cruces Biological Station
and Wilson Botanical Garden (La Amistad Pacífico Conservation Area), Lücking s.n. (F).
FIGURE 10. A–B. Crocynia microphyllina (holotype). A. Thallus. B. Hyphae of hypothallus. C. Dictyonema
hernandezii (paratype), thallus with zonate prothallus. D. Dictyonema hirsutum (holotype), thallus lobes with surface
hairs. E. Diorygma microsporum (holotype), thallus with lirellae. F. Diorygma sticticum (holotype), clustered lirellae.
Scale in A, E, F = 1 mm, in B = 50 µm, in C, D = 5 mm.
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This remarkable new species belongs in the group formed by Dictyonema sericeum and relatives,
characterized by a more or less filamentous thallus having a characteristic fungal sheath around the
cyanobacterial filaments composed of jigsaw puzzle-shaped cells (Chaves et al. 2004, Lawrey et al. 2009). It
is distinguished from all other species in this group by its peculiar thallus morphology and anatomy: the
thallus is strongly compacted and gelatinous when wet, with a smooth and nitidous surface when dry, and in
section the photobiont layer is strongly compacted in the upper part of the thallus and resting on a thick
medulla. This heteromerous structure is otherwise known only from D. glabratum and relatives, which differ
from species of the D. sericeum group in the foliose thallus with the photobiont formed by individual cells
rather than filaments. Yet, phylogenetically, D. hernandezii is closely related to species in the D. sericeum
group.
Dictyonema hirsutum Moncada & Lücking, sp. nov. (Fig. 10D) Mycobank MB 517762
Sicut Dictyonema glabratum sed thallo hirsuto differt.—http://www.eol.org/pages/Dictyonema hirsutum
Type:—COLOMBIA. Cundinamarca: Municipio de Choachí, Vereda el Verjón, Reserva Natural
Matarredonda, vía Laguna El Verjón, 3220 m, paramo vegetation, August 2008, Lücking s.n. (holotype
UDBC, isotype F).
Thallus terricolous or between bryophytes, foliose, composed of semicircular to flabellate lobes 0.5–1 cm
diameter wide and 200–300 µm thick, with thickened, involute margins; upper surface densely hirstute and
covered with obliquely oriented, white trichomes 0.3–0.5 mm long and 20–30 µm thick at the base, composed
of densely packed parallel hyphae; zone close to the lobe margin usually glabrous and olive-green, surface
otherwise white; lower surface ecorticate, finely felty-byssoid, mottled pale brownish to bluish grey. Thallus
in section with upper corticiform layer composed of loosely packed, intricate, 3–4 µ m thick, colourless
hyphae, irregular photobiont layer with groups of cyanobacterial cells wrapped in a dense, partially
paraplectenchymatous, hyphal sheath, and thin medulla of loosely woven hyphae also forming the lower
thallus surface; clamps not observed. Photobiont a chroococcoid morph of Rhizonema (Lücking et al. 2009),
cells in groups of 4–8, groups 20–40 µm diameter, individual cells ovoid-ellipsoid, 10–13 × 6–10 µm, olive
green. Basidiocarps on lower side, forming pieces of flat crust, each 1–3 mm in diameter; hymenophore
attached by basal layer to lower side of thallus lobes, beige; margin slightly involute when dry, pale beige to
white. Basal layer byssoid, composed of loosely woven, 4–5 µm thick hyphae, 200–300 µm high, colourless;
subhymenial layer composed of densely woven, short-celled hyphae, 50–90 µm high, colourless; hymenium
composed of densely arranged, palisade-like, 30–40 × 4–5 µm, slightly clavate basidioles, colourless, nonamyloid. Basidiospores not observed.
Distribution and habitat:—The new species is known from the paramo region near Bogotá at over 3000
m elevation.
Etymology:—The epithet refers to the conspicuously hirsute upper surface of the thallus lobes.
The foliose forms of the genus Dictyonema are the most commonly collected basidiolichens; they usually
grow at higher elevations on soil or between bryophytes but also occur as epiphytes on trees (Chaves et al.
2004). A recent phylogenetic study suggested that the single foliose species distinguished by Parmasto (1978)
as D. glabratum (Spreng.) D.Hawksw., actually comprises several different species that are distinct
morphologically and genetically (Chaves et al. 2004, Lawrey et al. 2009). Dictyonema hirstutum differs from
D. glabratum s.str. in the dense layer of trichomes produced on the upper surface, together with the overall
smaller lobes. Cora pavonia f. villosa Tomas. might represent the same taxon, but that epithet has not been
used at the species level. Because of its small thalli and its growth over soil and between bryophytes, D.
hirsutum is easily overlooked between vegetation but very conspicuous up close for its peculiar morphology.
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Diorygma microsporum M.Cáceres & Lücking, sp. nov. (Fig. 10E) Mycobank MB 517763
Sicut Diorygma poitaei sed ascosporis minoribus et thallo acidum norsticticum continente differt.—http://www.eol.org/
pages/Diorygma microsporum
Type:—BRAZIL. Pernambuco: Igarassu, Charles Darwin Ecological Refuge, 34°54’W, 7°50’S, 20 m,
undisturbed, closed lowland rain forest (Mata Atlântica), on bark, 2000, Cáceres & Lücking 00-531 (holotype
ASE, isotype F).
Thallus corticolous, up to 5 cm diameter, 50–120 µm thick, continuous; surface smooth to uneven, light
greyish green; white prothallus sometimes present. Thallus in section with 5–10 µm thick loose upper cortex,
irregular algal layer and clusters of calcium oxalate crystals. Lirellae in dense clusters, stellately branched,
erumpent, with thick complete thalline margin, 1–5 mm long, 0.15–0.25 mm wide, 0.1 mm high; disc partly
exposed, flesh-coloured to light red-brown but thickly white-pruinose; labia conspicuous, white, the proper
excipulum white-pruinose and crumbling and usually separated from the thalline margin by a split; thalline
margin white. Excipulum entire, orange-brown, 15–25 µm thick; laterally covered by corticate algiferous
thallus including clusters of crystals; hypothecium prosoplectenchymatous, 10–15 µ m high, yellowish;
hymenium 60–70 µm high, colourless, clear; epithecium granulose, 5–10 µm high, yellow-brown. Paraphyses
unbranched; asci clavate, 60–70 × 12–15 µ m. Ascospores 8 per ascus, ellipsoid, submuriform with 3
transverse and 0–1 longitudinal septa per segment, 12–15 6–7 µm, 2 times as long as wide, with thickened
septa and rounded lumina, colourless, strongly I+ violet-blue. Secondary chemistry: norstictic acid.
Distribution and habitat:—Diorygma microsporum appears to have a rather wide neotropical
distribution, as indicated by the disjunct collections from northeastern Brazil, Colombia, and Florida.
Etymology:—The epithet refers to the unusually small ascospores, thus far the smallest known in the
genus.
Additional specimens examined (paratypes):—U.S.A. Florida: Collier Co., Fakahatchee Strand
Preserve State Park, Lücking & Rivas Plata 26504 (F). COLOMBIA: Tolima: Purificacion, Vereda El Tambo,
Finca El Santuario, Cruz & Sandoval s.n. (UDBC).
The genus Diorygma is a mid-sized genus in Graphidaceae (Kalb et al. 2004). The genus is essentially
tropical and restricted to lowland and lower montane rain forest. Species typically grow on tree trunks in the
shady understory of undisturbed forest and hence are good indicators of forest health. The new species
presented here was first collected in Brazil and identified as Anomomorpha aggregans (Nyl.) Staiger (Cáceres
2007), because of the very small ascospores and norstictic acid chemistry. A second collection was recently
made in Florida, and restudy of the material revealed that hymenial inspersion, an important character in
Anomomorpha (Staiger 2002) is lacking; also, the morphology differs from typical species in Anomomorpha
being more reminescent of Diorygma. DNA sequencing of the Florida collection confirmed placement in
Diorygma (Rivas Plata et al. unpubl. data), where the species is quite unusual due to its very small ascospores
and compact thallus (Kalb et al. 2004). Superficially, the new species is similar to D. poitaei (Nyl.) Kalb,
Staiger & Elix.
Diorygma sticticum Sutjaritturakan, Kalb & Lücking, sp. nov. (Fig. 10F) Mycobank MB 517764
Sicut Diorygma poitaei sed ascosporis minoribus differt.—http://www.eol.org/pages/Diorygma sticticum
Type:—THAILAND. Nakhon Ratchasima Province: Khao Yai National Park, in front of Ban To Or, on
unidentified tree, 1999, Sutjaritturakan 12683 (holotype RAMK).
Thallus corticolous, up to 5 cm diameter, 70–150 µm thick, continuous; surface smooth to uneven, light
green; white prothallus sometimes present. Thallus in section with 10–15 µ m thick loose upper cortex,
irregular algal layer and clusters of calcium oxalate crystals. Lirellae in dense clusters, stellately branched,
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erumpent, with thick, complete thalline margin, 1–3 mm long, 0.15–0.3 mm wide, 0.1 mm high; disc
concealed to partly exposed, pale brown but thickly white-pruinose; labia conspicuous, white, the proper
excipulum white-pruinose; thalline margin white. Excipulum entire, orange-brown, 15–30 µm thick; laterally
covered by corticate algiferous thallus including clusters of crystals; hypothecium prosoplectenchymatous,
10–15 µm high, yellowish; hymenium 60–80 µm high, colourless, clear; epithecium granulose, 5–10 µm high,
yellow-brown. Paraphyses unbranched; asci clavate, 60–70 × 10–12 µm. Ascospores 8 per ascus, ellipsoid,
submuriform with 3 transverse and 0–1 longitudinal septa per segment, 10–13 × 6–8 µm, 1.5–2 times as long
as wide, with thickened septa and rounded lumina, colourless, strongly I+ violet-blue. Secondary chemistry:
stictic, hypostictic, and cryptostictic acids.
Distribution and habitat:—Diorygma sticticum is so far known from several collections from lowland
rain forest in Thailand.
Etymology:—The epithet refers to the secondary chemistry.
Additional specimens examined (paratypes):—THAILAND. Nakhon Ratchasima Province: Khao Yai
National Park, Sutjaritturakan 13556 (RAMK). Saraburi Province: Sutjaritturakan 20215 (RAMK). Loei:
Phuluang Wildlife Sanctuary, Phuluang Animal Research Station, Poensungnoen RU-VP00413 (RAMK).
This is a second new species in the genus Diorygma, similar to the preceding one in having extremely
small ascospores, but differing in the secondary chemistry and the geographic distribution.
Echinoplaca pernambucensis Øvstedal & Elix, sp. nov. (Fig. 11A) Mycobank MB 517765
Speciei Echinoplacae verruciferae similis, sed sine setis, et acidum gyrophoricum, acidum lecanoricum et acidum
subgyrophoricum continens. Apothecia non vidi.—http://www.eol.org/pages/Echinoplaca pernambucensis
Type:—BRAZIL. Pernambuco: Recife, Dois Irmãos, January 2010, Silva s.n. (holotype BG).
Thallus crustose, the basal crust whitish with a green hue, 2–3 cm wide, 100–115 µm thick, corticate but
some places cracked and granular. Photobiont a species of Chlorococcaceae, cells 4–6 µm wide (similar to
that in E. similis). Calcium oxalate crystals abundant. Numerous conical areolae present on the upper surface,
up to 0.2 mm high and 0.2 mm wide, pinkish white, without algae, with a more compact texture than the basal
crust, with a central hyphophore. Hyphophores whitish, up to 0.4 mm long, mostly curved, 30–40 µm wide,
slightly broader at the apices. No diahyphae, setae or apothecia seen. Secondary chemistry: gyrophoric acid
(major), lecanoric acid (minor) and subgyrophoric acid (minor).
Distribution and habitat:—On bark of undetermined tree in remnant Mata Atlantica, ca 1.5 m above
ground.
Etymology:—The specific epithet derives from the locality of the type specimen.
The genus Echinoplaca (Gomphillaceae) is usually epiphyllous, but a few species are corticolous
(Cáceres 2007, Kalb & Vĕzda 1988, Lücking 2008). It is characterized by apothecia which are flush with the
thallus and frequently the presence of hyphophores and setae. A more detailed description of the genus and
family is found in Lücking (2008). The combination of hyphophores, the presence of calcium oxalate crystals
and the type of photobiont is clearly consistent with Gomphillaceae. In the absence of apothecia, the generic
placement of this species is tentative. At least three genera seem likely possibilities: Echinoplaca,
Gyalideopsis and Calenia. Cáceres (2007) found no species of Gyalideopsis on bark in Pernambuco.
Although there are some epiphyllous species of Gyalideopsis with white hyphophores present in the
Neotropics (Lücking 2008), all have some characteristics which distinguish them from the present
species.There are no similar corticolous species reported in a recent review of the genus (Lücking et al. 2006).
In contrast, Cáceres (2007) found three species of Echinoplaca on bark in Pernambuco, E. bispora Kalb &
Vĕzda, E. caruaruensis Cáceres & Lücking, and E. leucotrichoides (Vain.) R.Sant. However, E. caruaruensis
lacks hyphophores, E. leucotrichoides has tapered hyphophores with a darkened apex, and E. bispora has a
thin and shiny thallus, small, white, tapered setae and hyphophores with a pale base and dark brown apices. A
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further corticolous Brazilian species, E. similis Kalb & Vĕzda, has a thick white thallus, white tapered setae
and white hyphophores which are dark on the dorsal side. It contains no lichen compounds (TLC, type
studied). At present only one corticolous species of Calenia, viz. C. corticola (Henssen) Ferraro, Lücking &
Séru. is known, but that species lacks hyphophores. As to the epiphyllous species of Calenia, several have
hyphophores (e.g. C. lueckingii Hartmann, C. monospora Vĕzda) like those present in E. pernambucensis but
their thalli differ and they lack lichen substances (Lücking 2008). There are parasitic members of
Gomphillaceae (Lücking 2008) and one could envisage that the hyphophore-bearing areolae belong to a
parasite on another species, but as the areolae bearing hyphophores react C+ red, there appears to be only one
species present. No mention of lichen products in Gomphillaceae is found in Vĕzda & Poelt (1987), Kalb &
Vĕzda (1988), Lücking (1997, 2008), and Lücking et al. (2005, 2007).
Echinoplaca schizidiifera J.E.Hern. & Lücking, sp. nov. (Fig. 11B–C) Mycobank MB 517766
Echinoplaca species thallo schizidiis disintegrantibus differt.—http://www.eol.org/pages/Echinoplaca schizidiifera
Type:—VENEZUELA. Distrito Federal: Parque Nacional El Ávila, zonas aledañas a Lagunazo, 10º 33’N,
66º 52’E, 2000 m, bosques nublados costeros de la Cordillera de la Costa, bosque siempreverde de media
altura (20–25 m) con árboles emergentes que pueden alcanzar hasta 40 m, foliicolous, August 2008, Lücking
26084 (holotype VEN, isotype F).
Thallus foliicolous, up to 50 mm across and 20–30 µm thick, with cartilaginous, corticiform layer, uneven
to coarsely verrucose due to incrustation with clusters of calcium oxalate crystals, with very scattered sterile
setae, pale greenish grey to white; verrucae rather flat, irregular, 0.1–0.3 mm diameter, white; setae 0.2–0.3
mm long, white or apically darkened. Thallus especially in inner parts dissolving into angular schizidia by
disintegration; schizidia with the same anatomy as the thallus, 0.3–0.5(–1) mm diameter, at first visible by
irregular splits but eventually becoming involute and falling off, leaving behind small, rounded structured
marking the attachment points. Apothecia not observed. Hyphophores not observed (but sterile setae might
represent postmature hyphophores lacking diaphyphae). Secondary chemistry: no substances detected by
TLC.
Distribution and habitat:—Echinoplaca schizidiifera was found inhabiting the leathery leaves of Clusia
trees in upper montane cloud forest in Venezuela near the capital Caracas.
Etymology:—The epithet refers to the thallus dissolving into schizidia.
This new species is distinguished from all other species of Echinoplaca by the unusual vegetative
reproduction via schizidia. Schizidia are otherwise unknown in the family Gomphillaceae but very similar
schizidia have been described from species of Coenogonium in the related family Coenogoniaceae (Lücking
2008). The schizidia of E. schizidiifera should not be confused with the disc-shaped propagules found in a
species of the related genus Actinoplaca, A. gemmifera (Lücking) Lücking. In the latter, the discs produced on
the thallus surface are homologous to hyphophores, whereas in the new species, the schizidia represent parts
of the original vegetative thallus.
Eremithallus marusae R.Miranda, Gaya & Lücking, sp. nov. (Fig. 11D) Mycobank MB 517767
Sicut Eremithallus costaricensis sed ascosporis minoribus differt.—http://www.eol.org/pages/Eremithallus marusae
Type:—MEXICO. Jalisco: Estación de Biología Chamela, tropical deciduous dry forest, on bark of
Heliocarpus pallidus, 19° 29’N, 105°02W, 60 m, June 2009, Miranda 766 (holotype MEXU).
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FIGURE 11. A. Echinoplaca pernambucensis (holotype), hyphophores. B–C. Echinoplaca schizidiifera (isotype),
thallus dissolving into schizidia that leave behind circular traces where the stipe originally attached to the leaf surface. D.
Eremithallus marusae (holotype), apothecia in different stages of development. E. Everniastrum constictovexans
(holotype), detail of thallus. F. Fellhanera borbonica (holotype), thallus with apothecia. Scale in A = 0.3 mm, in B–D, F
= 1 mm, in E = 3 mm.
Thallus superficially delimited by yellowish-grey colour and partially a thin, brown-black prothallus, but
proper thallus structures (cortex, algal layer, medulla) absent; photobiont a species of Trentepohlia, cells 7–12
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× 7–9 µm, located within the periderm cells of the tree bark and connected to the base of the apothecia by
hyaline fungal hyphae. Apothecia erumpent from periderm and finally almost appearing sessile, rounded, up
to 1 mm diameter, with brown-black disc and irregular marginal, grey-white lobules (remnants of the initially
covering periderm). Proper excipulum 20–40 µ m broad, red-brown, lateral towards the hymenium with
anastomosing periphysoids and paraphysoids. Hymenium 55–70 µ m high, non-amyloid, composed of
unbranched to rarely anastomosing paraphyses; asci ellipsoid to clavate, 55–65 × 15–18 µm, non-amyloid,
with thick tholus and distinct ocular chamber when young (probably fissitunicate). Ascospores 8 per ascus,
ellipsoid, 1-septate, 15–20 × 6–8 µm, hyaline, non-amyloid, constricted at septum when mature. Secondary
chemistry: no substances detected by TLC.
Distribution and habitat:—Eremithallus marusae is known from Chamela-Cuixmala Biosphere Reserve
in Jalisco, Mexico, which is one of the last protected areas of tropical deciduous dry forest remaining in the
Neotropics. The reserve bears a unique and highly diverse lichen biota with dominance of crustose taxa
(Miranda et al. 2009). Both known species of Eremithallus seem to prefer (semi-)deciduous dry forest
vegetation.
Etymology:—We honour our mentor and colleague, María de los Angeles Herrera-Campos, for her
contributions to lichenology in Mexico.
Additional specimen examined (paratype):—MEXICO. Jalisco: Estación de Biología Chamela,
Miranda 767, 1328, 1331 (MEXU).
Eremithallus is a recently described genus that appears to represent a distinctive lineage within the higher
Ascomycota (Lücking et al. 2008b). Morphologically and anatomically it resembles Ostropales
(Lecanoromycetes), but genetically clusters outside that class. Unique features of the genus are the absence of
a lichen thallus, the photobiont cells instead clustered in distinct, discrete groups within dead periderm cells of
the tree bark, the laterally strongly anastomosing periphysoids, and the ascospores containing folds or
crystals, at least in the type species. The genus superficially resembles non-lichenized fungi such as Dothidea
and Melaspilea, but these differ anatomically in the structure of the hymenia and/or stromata. The new species
differs from the type species mainly in the smaller ascospores (20–30 µm long in E. costaricensis) which lack
folds. Also, the hymenium is lower and the asci are smaller on average. Otherwise, the apothecial anatomy,
especially the thin brown excipulum bordering a layer of strongly anastomosing periphysoids (or lateral
paraphyses), as well as the peculiar position of the photobiont, agree in both taxa. While in the type species,
only mature asci were observed, the new species also showed young asci with a typical fissitunicate anatomy.
It is not unlikely that an older name is available for this genus, but thus far we have not been able to detect
anything similar described in the literature or among revised herbarium collections.
Everniastrum constictovexans Sipman, sp. nov. (Fig. 11E) Mycobank MB 517768
Everniastrum corticola lobis linearibus, isidiis ciliatis provisum, acida consticticum et protolichesterinicum in medulla
continens.—http://www.eol.org/pages/Everniastrum constictovexans
Type:—PERU. Cuzco: Along the Cuzco-Pilcopata-Paucartambo Rd. at a point ca 135 km from the
intersection of the Cuzco-Puno Rd., on soil in road cut, July 1976, Dumont et al. 1653 (holotype NY).
Thallus foliose to subfruticose, corticolous, over 5 cm wide, not coriaceous, dichotomously lobate. Lobes
long-linear, separate, 0.6–1.5 mm wide, over most of their length strongly convex and subtubular. Upper
surface pale grey, slightly shiny, epruinose, weakly maculate, isidiate, lacking soredia, pustules, dactyls or
lobules. Isidia cylindrical, scattered over the surface, concolorous with the thallus, when young often browncapped, when old often with an erect, apical, black cilium. Medulla white. Lower surface black, towards the
lobe tips brown. Cilia black, slender, 1–4 mm long, simple or occasionally once dichotomously branched.
Apothecia and Pycnidia not seen. Chemistry: upper cortex K+ yellow, C–, KC–, P–, UV–, medulla K+
orange, C–, KC–, P+ orange-red, UV–. Secondary metabolites: Upper cortex with atranorin, medulla with
constictic and protolichesterinic acids (TLC).
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Distribution and habitat:—Known so far only from one specimen from the highlands of Peru.
Etymology:—The epithet reflects the morphological similarity to Everniastrum vexans (Zahlbr. ex
W.Culb. & C.Culb.) Hale ex Sipman and the presence of constictic acid.
A key to all known species of Everniastrum was given in Sipman (1986a). Since then only few additional
species were published, from East and Southeast Asia (Chen et al. 1989, Jiang & Wei 1989, 1993, Pooprang et
al. 1999). The genus was included in Cetrariastrum by, e.g. Culberson & Culberson (1981), considered
different by Sipman (1986b) and of undecided status following DNA analyses (Divakar et al. 2006, 2010).
Here the traditional classification based on morphology is followed provisionally, awaiting further taxonomic
rearrangements resulting from DNA analyses. Everniastrum constictovexans is morphologically very similar
to E. vexans, at least the common morph of the Neotropics, with which it shares the often very elongate and
ciliate superficial isidia. It differs only in the presence of constictic acid instead of salazinic acid as main
medullary substance. Thus it might be considered as a chemical strain. However, in recent taxonomic
treatments (Sipman et al. 2009) such chemical differences are given species rank. The chemical difference is
not easily demonstrated with spot tests. E. constictovexans has a non-isidiate relative in E. peruvianum Hale
ex W.Culb. & C.Culb.
Fellhanera borbonica Sérus., van den Boom & Brand, sp. nov. (Fig. 11F) Mycobank MB 517769
Ab omnibus speciebus generis Fellhanerae differt ascosporis submuriformibus et muriformibus et conidiis biclavatis.—
http://www.eol.org/pages/Fellhanera borbonica
Type:—LA RÉUNION (French Mascarenes). Forêt de Bélouve, near entrance of Sentier de Mazerin;
21°04’S, 55°33’E, 1500 m; subvertical shaded basaltic outcrop; van den Boom et al. 39866 (holotype LG,
isotype hb. van den Boom).
Thallus saxicolous, crustose, greyish green to pale grey, somewhat rimose and cracked (and then medulla
seen at high magnification); prothallus indistinct. Photobiont chlorococcoid with cells green and globose, 5–8
µ m diameter. Apothecia rare, round, 0.5–0.7 mm in diameter and ca. 0.2 mm high, sessile and strongly
constricted at the base, disc flat and eventually slightly convex, dark brown, with a violet hue (especially
when fresh), margin thin but distinct, first slightly prominent and eventually flexuose and almost excluded,
pale orange brown to brown; outer parts usually downy or with short whitish hairs, in young apothecia some
connecting the apothecia to the thallus. Excipulum thin (c. 20–30 µ m), inner parts typically
paraplectenchymatous, becoming less typical towards the margin, and then made of densely interwoven
hyphae with short hyphae protruding. Hypothecium 50–70 µm thick, dark brown to blackish, K–. Paraphyses
strongly gelatinized, branched and not inflated at apices. Asci clavate, 50–60 × 10 µm, of the Byssoloma-type.
Ascospores 8/ascus, ellipsoid, submurifom to muriform, 3(–4) transverse septa and (1–)2(–3) longitudinal or
oblique ones, 18–20 × 5–7 µm, with a distinct halo (best seen in K). Pycnidia half-immersed in the thallus, ca.
0.1 mm in diameter, wall pale orange, K–, C–, with a distinct gaping ostiole but no glut of conidia. Conidia
numerous, typically biclavate (almost all conidia with both ends typically inflated and almost spherical),
colourless, simple, 4–5 × 1.5–2 µm (width measured at extremities).
Distribution and habitat:—Fellhanera borbonica is a saxicolous species, only known from the type
collection in the montane forest of La Réunion ("forêt de Bois de Couleur des Hauts"). It was found on a
shaded basaltic outcrop within a disturbed forest.
Etymology:—The epithet has been choosen after the historical name of La Réunion: Île Bourbon
(Borbonia).
Lücking (1997) provided a comprehensive survey of the variation in Fellhanera (Pilocarpaceae), mostly
represented by foliicolous species, including two species with muriform ascospores. The genus is
characterized by: biatorine apothecia with paraplectenchymatous exciple, Byssoloma-type asci, Bilimbia- type
ascospores and bacillar to pyriform conidia. This concept still prevails (Lücking 2008), awaiting eventual
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support from molecular phylogenetic studies. In their survey of the Fellhanera silicis group in eastern North
America, Harris & Lendemer (2009) recorded variation in conidia shape and size and used these characters to
support delimitation of several new species. Although known only from a single collection, Fellhanera
b o r bo n i ca f u rt h er d e m o n st ra t es t h e va r ia ti o n e nc o u n te r ed i n th a t g ro u p a s it co m bi n e s a
paraplectenchaymatous excipulum (becoming less organized and almost byssoid at the margins) with
submuriform to muriform ascospores and biclavate conidia, this shape not earlier reported in the genus.
Otherwise, the new species is most similar to F. elliottii (Vain.) Vĕzda and F. paradoxa (Vĕzda) Vĕzda, both
of which have a similar morphology and also (sub-)muriform ascospores, but the ascospores are distinctly
longer and the conidia are bacillar to narrowly pyriform (Lücking 2008).
Fibrillithecis sprucei Mangold, Lücking & Lumbsch, sp. nov. (Fig. 12A) Mycobank MB 517770
Sicut Fibrillithecis dehiscens sed ascosporis muriformibus majoribusque differt.—http://www.eol.org/pages/
Fibrillithecis sprucei
Type:—BRAZIL: Amazonas: Rio Negro Sán Carlos, Spruce 395 (holotype BM, two larger pieces on left
side).
Thallus corticolous, up to 10 cm diameter, 200–500 µm thick, continuous; surface uneven to bumpy, light
greyish green to olive-green. Thallus in section with 20–30 µm thick prosoplectenchymatous upper cortex,
thick, irregular algal layer, and white medulla; calcium oxalate crystals not apparent. Apothecia erumpent,
rounded to angular-rounded, 0.7–1.2 mm diameter; disc hidden by a narrow, 0.05–0.1 mm wide pore; proper
margin invisible; thalline margin thick, coarsely crenulate to almost lobulate, exposing the white medulla.
Excipulum pale yellow, 30–50 µm thick; laterally covered by a thick layer of algiferous, apically with radially
elongate, partially free fibrils; hypothecium prosoplectenchymatous, 10–20 µ m high, colourless to pale
yellowish; hymenium 180–250 µm high, colourless, strongly inspersed; epithecium indistinct. Paraphyses
unbranched, glabrous; periphysoids absent; asci fusiform to clavate, 150–200 × 30–40 µm. Ascospores single,
oblong, richly muriform, 120–180 × 25–35 µm, 4–5 times as long as wide, with comparatively thin septa and
angular lumina, colourless, I+ violet-blue. Secondary chemistry: psoromic acid and satellite substances (P+
yellow).
Distribution and habitat:—This species is so far only known from a single historic collection from the
Amazon rainforest.
Etymology:—The epithet honours Richard Spruce (1817–1893), collector of the type material and many
other interesting Amazonia lichens.
This new species was completely overlooked previously. It is part of the type collection of another species
of the same genus, Fibrillithecis dehiscens (Leight.) Mangold, Lücking & Lumbsch, comb. nov.
[Basionym: Thelotrema dehiscens Leight., Trans. Linn. Soc. London 25: 448 (1866); lectotype (here selected):
BRAZIL: Amazonas: Rio Negro Sán Carlos, Spruce 395 (lectotype BM, two smaller pieces on upper right
side)]. Both species look very similar externally and share the same apothecial morphology and anatomy, the
inspersed hymenium, and the psoromic acid chemistry. However, of the four pieces of bark in the collection,
two differ in having much larger apothecia. These are mostly sterile, but a few contain ascospores that are
large, muriform, and single in each ascus. The other two specimens have smaller apothecia with ascospores
being 11–15-septate and 40–60 × 8–12 µ m. This material corresponds to the description of Thelotrema
dehiscens, whereas the material with large apothecia and muriform ascospores is here described as new.
Fibrillithecis sprucei is the first species in the genus with large, muriform ascospores; all other species have
transversely septate or submuriform ascospores that are very small to medium-sized, not exceeding 60 × 12
µm (Rivas Plata et al. 2010).
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Fissurina astroisidiata Herrera-Campos & Lücking, sp. nov. (Fig. 12B–C) Mycobank MB 517771
Sicut Fissurina incrustans sed thallo isidiato, lirellis stellatis et ascosporis minoribus differt.—http://www.eol.org/pages/
Fissurina astroisidiata
FIGURE 12. A. Fibrillithecis sprucei (holotype), thallus with apothecia. B–C. Fissurina astroisidiata (B, isotype; C,
paratype). B. Thallus with lirellae. C. Isidia. D. Fissurina nigrolabiata (holotype), thallus with lirellae. E. Fissurina
subcomparimuralis (holotype), thallus with lirellae. F. Graphis caribica (holotype), thallus with lirellae. Scale = 1 mm.
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Type:—MEXICO: Veracruz: Los Tuxtlas Biosphere Reserve, Volcán Sán Martín Tuxtla, 95°13’N, 18°34',
1000–1100 m, selva alta perennifolia perturbada, February 2003, Herrera Campos et al. RLD051 (holotype
MEXU, isotype F).
Thallus corticolous, up to 10 cm diameter, 100–150(–300) µm thick, continuous but typically cracked,
isidiate; surface uneven to bumpy (large bumps up to 300 µm thick), light greyish green to olive-green; isidia
coarse, cylindrical, basally constricted and tapering towards the apex, 1–2 mm long and 0.1–0.15 mm thick,
yellowish white but often darkened at the tip. Thallus in section with 10–15 µm thick prosoplectenchymatous
upper cortex, irregular algal layer and clusters of calcium oxalate crystals. Lirellae stellately branched, the
stellate clusters concentrated on the elevated thallus bumps, erumpent, fissurine, with thick complete thalline
margin, 1–3 mm long, 0.2–0.3 mm wide, 0.1–0.15 mm high; disc partly exposed; labia thin but conspicuous,
yellowish white; thalline margin light yellowish green to olive-green. Excipulum entire, colourless to orangebrown, 15–25 µ m thick; laterally covered by corticate algiferous thallus including clusters of crystals;
hypothecium prosoplectenchymatous, 5–10 µm high, colourless to pale yellowish; hymenium 90–100 µm
high, colourless, clear; epithecium granulose, 5–10 µm high, grey-brown. Paraphyses unbranched, glabrous;
periphysoids not observed; asci fusiform to clavate, 80–90 × 12–15 µm. Ascospores 8 per ascus, ellipsoid,
submuriform with 3 transverse and 0–1 longitudinal septa per segment, 12–15 × 6–8 µm (including 1–1.5 µm
thick wall), 2 times as long as wide, with angular-rounded septa and lumina, colourless, I–. Secondary
chemistry: no substances detected by TLC.
Distribution and habitat:—Fissurina astroisidiata is only known from Los Tuxtlas Biosphere Reserve
in Veracruz, Mexico, which represents the northernmost extension of neotropical rain forest. The new species
grows on trunks of larger trees in the rain forest understory and hence belongs to the group of species of
Graphidaceae which can be used as bioindicators of ecological continuity or forest health (Rivas Plata et al.
2008).
Etymology:—The epithet refers to the stellately branched lirellae and the isidiate thallus.
Additional specimen examined (paratype):—MEXICO: Veracruz: Los Tuxtlas Biosphere Reserve,
Volcán Sán Martín Tuxtla, Herrera Campos et al. RLD057 (F).
The genus Fissurina belongs in Graphidaceae, the largest family of tropical crustose lichens, and includes
species with more or less inconspicuous lirellae that usually appear as fissures in the thallus (Staiger 2002).
Species of Fissurina are typically found in undisturbed tropical to subtropical rain forests and grow on tree
trunks in the shady understory, less frequently on branches. Currently, c. 65 species are known (Lücking
unpubl. data), being distinguished on ascospore septation, chemistry, and lirellae and thallus morphology.
Fissurina astroisidiata is the first species in the genus known to produce isidia; similar isidia are known from
unrelated species in the family, such as Myriotrema hartii (Müll.Arg.) Hale and Reimnitzia santensis (Tuck.)
Kalb. (Rivas Plata et al. 2010, Sipman et al. 2010). Apart from the isidia, F. astroisidiata is morphologically
most similar to F. dumastii Fée, F. incrustans Fée, and F. karnatakensis Makhija & Adaw. (Staiger 2002,
Makhija & Adawadkar 2007); all three species share the gaping lirellae with the disc partially exposed and
thin but distinct labia, similar to species of the genus Platythecium (Staiger 2002). However, all lack isidia,
and F. dumastii differs in its 3-septate ascospores and irregularly branched lirellae, whereas F. incrustans has
larger, muriform ascospores and irregularly branched lirellae. The lirellae of F. karnatakensis look very
similar to those of F. astroisidiata, but that species has 3-septate ascospores and produces protocetraric and
fumarprotocetraric acid. The two available collections of the new species were sequenced (mtSSU and
nuLSU) and cluster at the base of Fissurina s.str., including also F. crassilabra Mont. & Bosch, F. dumastii, F.
elaiocarpa (A.W.Archer) A.W.Archer, and F. triticea (Nyl.) Staiger (Rivas Plata et al. unpubl. data).
Fissurina nigrolabiata Rivas Plata, Bawingan & Lücking, sp. nov. (Fig. 12D) MycobankMB 517772
Sicut Fissurina comparimuralis sed labiis ventricosis et ascosporis I– differt.—http://www.eol.org/pages/Fissurina
nigrolabiata
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Type:—PHILIPPINES. Luzon Island, Nueva Vizcaya: Mount Palali, 1400 m, March 2007, Rivas Plata 1198B
(holotype F, isotype CAHUP).
Thallus corticolous, 3–5 cm diameter, 30–50 µm thick, continuous; surface smooth to uneven, green,
shiny. Thallus in section with prosoplectenchymatous upper cortex, irregular algal layer and large clusters of
crystals. Lirellae straight to curved, unbranched to sparsely branched, erumpent to prominent, with welldeveloped labia, with lateral thalline margin, 1–3 mm long, 0.3–0.4 mm wide, 0.3–0.4 mm high; disc
concealed; labia conspicuous, thickened, greyish black to brown-black, apically covered by translucent
cortical layer; thalline margin cream-coloured, sometimes with slight redding tinge. Excipulum entire,
completely and massively carbonized including below the hypothecion, black, 80–120 µm wide laterally and
up to 200 µm thick below; laterally covered by corticate algiferous thallus including large clusters of crystals
at the base; hypothecium prosoplectenchymatous, 5–10 µm high, colourless to pale yellowish; hymenium
120–180 µm high, colourless, clear; epithecium indistinct. Paraphyses unbranched, glabrous; periphysoids not
observed; asci fusiform, 80–100 × 15–20 µm. Ascospores 8 per ascus, ellipsoid, muriform with 3–5 × 1–2
septa, 12–17 × 5–8 µ m, with comparatively thin outer wall, 2–2.5 times as long as wide, I–. Secondary
chemistry: no substances detected by TLC.
Distribution and habitat:—This new species is thus far only known from pristine lower montane
rainforest in northern Luzon.
Etymology:—The epithet refers to the thick, carbonized labia.
Fissurina nigrolabiata was first believed to belong in Carbacanthographis, because of its massively
carbonized labia and I– ascospores. However, molecular data (Rivas Plata et al. unpubl. data) place this taxon
in the Fissurina clade, quite distant from Carbacanthographis. The new species is therefore provisionally
placed in Fissurina, with which it agrees in the distinctly corticate, shiny thallus and the lirellate ascomata.
However, Fissurina is possibly a collective genus and more data are required to divide this genus into natural
groups. Within Fissurina s.lat., several species are known to form lirellae with thick labia and well-developed
excipulum (subcontexta-type according to Staiger 2002). However, a completely and massively carbonized
excipulum was not formerly reported from this genus. The shape and anatomy of the lirellae are reminiscent
of Graphis, but species in that genus have I+ violet-blue ascospores and the thallus is mostly white-grey.
Fissurina subcomparimuralis Common & Lücking, sp. nov. (Fig. 12E) Mycobank MB 517773
Sicut Fissurina comparimuralis sed ascosporis I– et periphysoidis spinulosis differt. Thallus corticatus. Lirellae
fissurinae, 1–3 mm longae, 0.25 mm latae. Ascosporae muriformes, 15–25 × 6–8 µm, I–. Acidi lichenici desunt.—
http://www.eol.org/pages/Fissurina subcomparimuralis
Type:—U.S.A. Florida. Collier County, Fakahatchee Strand Preserve State Park, Common 7323A (holotype
MSC, isotype hb. Common).
Thallus corticolous, 1–2 cm diameter, 30–50 µm thick, continuous; surface smooth, yellowish-brown to
olive-brown. Thallus in section with prosoplectenchymatous upper cortex, irregular algal layer and scattered
clusters of crystals. Lirellae straight to curved, unbranched to sparsely branched, erumpent, fissurine, with
complete thalline margin, 1–3 mm long, 0.2–0.25 mm wide, 0.12–0.15 mm high; disc concealed; labia
conspicuous but not thickened, greyish black to brown-black below corticate thalline margin; thalline margin
olive-brown. Excipulum entire, brown-black, 20–40 µ m wide; covered by corticate algiferous thallus
including clusters of crystals; hypothecium prosoplectenchymatous, 5–10 µ m high, colourless to pale
yellowish; hymenium 70–90 µm high, colourless, clear; epithecium granulose, 5–10 µm high, grey-brown.
Paraphyses unbranched, glabrous; periphysoids present, apically spinulose; asci fusiform, 70–80 × 15–20 µm.
Ascospores 8 per ascus, ellipsoid, muriform with 5–7 × 1–3 septa, 15–25 × 6–8 µm (including 1–1.5 µm thick
wall), 2.5–3 times as long as wide, with ± trypethelioid septa and lumina, colourless, I–. Secondary chemistry:
no substances detected by TLC.
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Distribution and habitat:—Fissurina subcomparimuralis is known from several rich collections from
Fakahatchee Strand Preserve State Park in Florida.
Etymology:—The epithet refers to the close similarity with F. comparimuralis.
Additional specimens examined (paratypes):—U.S.A. Florida. Collier County, Fakahatchee Strand
Preserve State Park, Common 7276A, 7276J (hb. Common).
This new species was at first identified with F. comparimuralis Staiger, until we discovered two
significant differences: F. comparimuralis has I+ distinctly violet-blue ascospores and lacks spinulose
periphysoids. Since we consider these characters taxonomically important, we describe the Florida material as
a separate species.
Graphis caribica Lücking, sp. nov. (Fig. 12F) Mycobank MB 517774
Sicut Graphis fournieri sed excipulo carbonisato in parte apicali et ascosporis transversaliter septatis differt.—http://
www.eol.org/pages/Graphis caribica
Type:—DOMINICA. Bois Serpé (trail from South Chiltern to Soufrière Bay) Parishes of St Luke and St.
Marks, 300 m, November 1963, Imshaug & Imshaug 32777-A2 (holotype MSC).
Thallus corticolous, up to 5 cm diameter, 30–70 µm thick, continuous; surface uneven, yellowish green;
prothallus absent. Thallus in section with cartilaginous upper cortex, irregular algal layer and large clusters of
crystals below the algal layer. Apothecia lirelliform, flexuose, sparsely to irregularly branched, prominent,
lacking thalline margin, 1–5 mm long, 0.3–0.4 mm wide, 0.2–0.3 mm high; disc concealed; proper margin
thick, labia striate; thalline margin absent. Excipulum apically crenulate, apically to peripherally carbonized,
50–100 µ m wide, apically black but laterally and basally pale yellow to orange-brown; hypothecium
prosoplectenchymatous, 10–15 µ m high, colourless to pale yellowish; hymenium 120–150 µ m high,
colourless, clear; epithecium granulose, 10–15 µm high, olive brown. Paraphyses unbranched; asci fusiform
to clavate, 120–150 × 20–30 µm. Ascospores 8 per ascus, oblong, transversely 9–15-septate, 50–65 × 9–12
µm, 5–6 times as long as wide, colourless. Secondary chemistry: no substances detected by TLC.
Distribution and habitat:—The new species is known from several collections throughout the
Neotropics, centered around the Caribbean, being typically found in drier and subtropical forests.
Etymology:—The epithet refers to the apparently Caribbean distribution of this species.
Additional specimens examined (paratypes):—USA. Florida: Manatee, s.dat., Reasoner s.n. (F1204901). COSTA RICA. Alajuela: La Palma de Sán Ramón, Brenes 84 (FH). Volcán Tenorio National Park,
Pilón Biological Station (Arenal-Tempisque Conservation Area), Tilarán Ridge, Lücking 17265ab (F).
Guanacaste: Barra Honda National Park, Barra Honda Biological Station (Tempisque Conservation Area),
Barra Honda Mountains, Lücking 17550e, 17557a (F). Monte Alto Forest Reserve, Monte Alto Biological
Station (Tempisque Conservation Area), Nicoya Peninsula, Lücking 17511e (F, INB).
This new species, collected first by the late Henry Imshaug, was already discussed in detail in a
monographic revision of Graphis from Costa Rica but not yet described formally (Lücking et al. 2008).
Graphis caribica differs from most other species of the genus in its yellowish green, uneven thallus. It is
morphologically very similar to G. fournieri Lizano & Lücking, but the latter has a more or less completely
carbonized excipulum and muriform ascospores.
Graphis cerradensis Marcelli, Benatti & Lücking, sp. nov. (Fig. 13A–B) Mycobank MB 517775
Sicut Graphis inspersostictica sed ascosporis muriformibus differt.—http://www.eol.org/pages/Graphis cerradensis
Type:—BRAZIL. São Paulo: Municipality of Mogi-Guaçu, Martinho Prado Jr. District, Mogi-Guaçu
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Biological Reserve, 22°15’S, 47°11’W, dense, humid Cerrado vegetation, on bark, November 2007, Lücking
23027 (holotype SP, isotype F).
FIGURE 13. A–B. Graphis cerradensis (isotype). Thallus with lirellae. C. Graphis itatiaiensis (isotype), thallus with
lirellae. D. Thallus with lirellae. E. Gyalideopsis chicaque (isotype), thallus with apothecia. F. Gyrotrema papillatum
(holotype), thallus with apothecia. Scale = 1 mm.
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Thallus corticolous, up to 5 cm diameter, 30–70 µm thick, continuous; surface smooth to uneven, matte,
pale greenish grey; prothallus absent. Thallus in section with cartilaginous upper cortex, irregular algal layer
and large clusters of crystals. Apothecia lirelliform, flexuose, unbranched to sparsely branched, prominent,
with thick complete thalline margin, 1–3 mm long, 0.5–0.6 mm wide, 0.25–0.35 mm high; disc concealed;
proper margin thick, labia striate; thalline margin thick, completely covering the labia, white. Excipulum
apically crenulate, completely carbonized, 70–150 µm wide, black; laterally covered by corticate algiferous
thallus including large clusters of crystals; hypothecium prosoplectenchymatous, 10–15 µm high, colourless
to pale yellowish; hymenium 150–200 µm high, colourless, strongly inspersed with small granules along the
paraphyses (paraphyses and asci hardly discernible), inspersion rapidly disappearing in KOH (type B);
epithecium granulose, 10–15 µm high, olive brown. Paraphyses unbranched; asci fusiform to clavate, 130–
180 × 30–40 µm. Ascospores 4–8 per ascus, oblong, muriform with 15–25 transverse and 1–3 longitudinal
septa per segment, 80–100 × 15–20 µm, 6–7 times as long as wide, colourless. Secondary chemistry: stictic
acid.
Distribution and habitat:—The new species is known from a single collection in a dense, humid
Cerrado vegetation in southern Brazil.
Etymology:—The epithet refers to the type of vegetation in which the species grows.
Graphis cerradensis belongs to the G. cinerea group, characterized by robust, prominent lirellae with
often striate labia, massively carbonized excipulum, and strongly inspersed hymenium (Lücking 2009,
Lücking et al. 2008, 2009). Thus far, no stictic-acid containing species with muriform ascospores was known
in this group. Most similar is G. argentata Lücking & Umaña, which lacks secondary substances and has
slightly broader ascospores. Graphis inspersostictica Sipman & Lücking also appears to be closely related; it
agrees with G. cerradensis in morphology and chemistry, but the lirellae are narrower with entire labia and the
ascospores are transversely septate or very rarely provided with a longitudinal septum in one of the segments.
Graphis itatiaiensis Nelsen, Lücking & Spielmann, sp. nov. (Fig. 13C) Mycobank MB 517776
Sicut Graphis rhizocola sed thallo isidioso et lirellis astroideis aggregatis differt.—http://www.eol.org/pages/Graphis
itatiaiensis
Type:—BRAZIL. Minas Gerais: Municipality of Itamonte, Parque Nacional do Itatiaia, Estrada das
Prateleiras, 22°22’S, 44°44’W, July 2009, Nelsen 4185 (holotype SP, isotype F).
Thallus corticolous, up to 7 cm diameter, 30–70 µm thick, continuous; surface smooth to unevenverrucose, greenish grey, with scattered isidioid outgrowths; isidoid outgrowths globose to irregular, 0.05–0.1
mm diameter. Thallus in section with cartilaginous upper cortex, irregular algal layer and large clusters of
crystals; photobiont trentepohlioid. Apothecia lirelliform, flexuose, stellately branched and aggregate into
pseudostromatic clusters, prominent to sessile, with thick, almost complete thalline margin leaving only the
uppermost part of the labia exposed, 1–3 mm long, 0.3–0.5 mm wide, 0.25–0.35 mm high; disc concealed;
proper margin thick, labia entire, grey-black; thalline margin thick, covering the labia except for the
uppermost part, white-grey and contrasting with the darker surrounding thallus. Excipulum completely
carbonized, 50–100 µm wide, black; laterally covered by algiferous thallus lacking crystals; hypothecium
prosoplectenchymatous, 10–15 µ m high, colourless to pale yellowish; hymenium 150–200 µ m high,
colourless, clear; epithecium granulose, 10–15 µm high, olive brown. Paraphyses unbranched; asci fusiform
to clavate, 130–180 × 30–40 µm. Ascospores 2–4 per ascus, oblong to cylindrical, 15–25-septate, 100–150 ×
14–17 µm, 7–9 times as long as wide, colourless, with thin gelatinous caps at both ends. Secondary chemistry:
no substances detected by TLC.
Distribution and habitat:—The new species is known from a few collections in lower montane Atlantic
rainforest in southeastern Brazil.
Etymology:—The epithet refers to the type locality.
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Additional specimens examined (paratypes):—BRAZIL. Minas Gerais: Municipality of Itamonte,
Parque Nacional do Itatiaia, Estrada das Prateleiras, Nelsen 4186 (F).
Graphis is a large genus with over 300 species found primarily on tree bark in semi-open to open
conditions in midelevation sites, such as lower montane rainforests (Lücking et al. 2008, Lücking 2009,
Lücking et al. 2009). The genus is typically characterized by a white-grey, corticate thallus, with erumpentprominent lirellae, a partly to fully carbonized excipulum, a hymenium mostly lacking inspersion and
transversely septate to muriform ascospores reacting I+ violet-blue (Staiger 2002, Lücking et al. 2008,
Lücking 2009). G. itatiaiaensis is distinguished from G. rhizocola (Fée) Lücking & Chaves by its coarsely
isidiate thallus and stellately branched lirellae. G. itatiaiaensis differs from G. isidiata (Hale) Lücking in that
the new species has elongate, stellately branched lirellae and septate rather than muriform ascospores. G.
itatiaiaensis differs from G. patwardhanii C.R.Kulk, another isidiate species with stellately branched lirellae,
in having prominent-sessile lirellae covered with thallus, slightly larger ascospores with a greater number of
septa per ascospore, a completely carbonized excipulum, and labia lacking striations.
Graphis marusae B.Peña & Lücking, sp. nov. (Fig. 13D) Mycobank MB 517777
Sicut Graphis longula sed lirellis nudis differt.—http://www.eol.org/pages/Graphis marusae
Type:—MEXICO. Veracruz: Municipio Catemaco, Los Tuxtlas Biosphere Reserve, Ejido Adolfo López
Mateos Ecological Reserve, 18º 26’N, 94º 28’W, 345 m, lowland rainforest, on bark of Astrocaryum
mexicanum, April 2009, Bárcenas Peña 1077 (holotype MEXU).
Thallus corticolous, up to 5 cm diameter, 50–100 µm thick, continuous; surface smooth, pale greenish
grey, shiny; prothallus absent. Thallus in section with cartilaginous upper cortex, irregular algal layer and
clusters of crystals. Apothecia lirelliform, flexuose, usually unbranched, prominent, lacking or with indistinct
basal to lateral thalline margin, 1–5 mm long, 0.2–0.3 mm wide, 0.2–0.25 mm high; disc concealed; proper
margin thick, labia for a long time entire but eventually becoming striate; thalline margin indistinct, thin, basal
to almost lateral, greenish grey, the grey-black labia almost fully exposed. Excipulum entire to apically
crenulate, completely carbonized, 80–120 µm wide, black; basally covered by thin thallus layer; hypothecium
prosoplectenchymatous, 10–20 µ m high, colourless; hymenium 100–150 µ m high, colourless, clear;
epithecium indistinct. Paraphyses unbranched; asci fusiform, 90–120 × 25–35 µm. Ascospores 8 per ascus,
oblong, 9–13-septate, 50–70 × 10–15 µm, 4–5 times as long as wide, colourless. Secondary chemistry: no
substances detected by TLC.
Distribution and habitat:—The new species is known from the Biosphere Reserve Los Tuxtlas in
Veracruz, Mexico, a relict tropical lowland rainforest. It was found on bark of Astrocaryum mexicanum and
Pseudolmedia oxyphyllaria in exposed understory at 200–345 m elevation.
Etymology:—This new species is dedicated to our mentor and colleague, María de los Angeles HerreraCampos for her important contributions to lichenology in Mexico and especially the taxonomy of the genus
Usnea.
Because of its completely carbonized excipulum, clear hymenium, and mid-sized, transversely septate
ascospores, this new species would key out under Graphis cognata Müll.Arg. with entire labia and under G.
longula Kremp., G. asterizans Nyl., and G. glauconigra Vain. with striate labia (Lücking et al. 2009). The
latter species agrees in the mostly entire labia which eventually become striate, but differs in the shorter
lirellae with distinct, conspicuous lateral thalline margin. Graphis asterizans likewise also has a distinct
thalline margin and the lirellae are stellately branched. Graphis longula differs in the soon striate labia and the
distinct lateral thalline margin which also forms white lines between the labial striae. The most closely related
species seems to be G. cognata, but that species has a white-grey thallus and shorter lirellae with jet-black
labia and a distinct basal thalline margin.
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Gyalideopsis chicaque Moncada & Lücking, sp. nov. (Fig. 13E) Mycobank MB 517778
Sicut Gyalideopsis tuerkii sed apotheciis minoribus et ascosporis brevioribus differt.—http://www.eol.org/pages/
Gyalideopsis chicaque
Type:—COLOMBIA. Cundinamarca: Sán Antonio del Tequendama, Chicaque Natural Park, 4°36’N,
74°18’W, 2300 m, montane rainforest, on compacted soil, August 2008, Lücking s.n. (UDBC, isotype F).
Thallus terricolous over strongly compacted soil, crustose, continuous, 10–30 mm across and 10–20 µm
thick, with a cartilaginous, corticiform layer, uneven to irregularly verrucose with clusters of calcium oxalate
crystals, pale greenish grey, nitidous. Photobiont trebouxioid, cells 5–8 µm diameter. Apothecia applanate to
broadly sessile, 0.25–0.5 mm diameter and 80–100 µm high; disc plane, dark brown, non-pruinose; proper
margin distinct, dark brown, non-pruinose. Excipulum composed of radiating and anastomosing hyphae
embedded in a gelatinous matrix, 30–50 µ m broad, colourless but externally covered by a dark brown,
granular layer. Hypothecium 5–10 µm high, colourless; epithecium 5–10 µm high, dark brown granulose.
Hymenium 70–90 µm high, colourless. Asci clavate, 60–80 × 12–15 µm. Ascospores 4–8/ascus, ellipsoid,
submuriform with 3–5 transverse septa and 0–1 longitudinal septa per segment, with slight constrictions at the
septa, 12–16 × 6–8 µm, 1.8–2.2 times as long as broad, colourless. Hyphophores not observed. Secondary
chemistry: no substances detected with TLC.
Distribution and habitat:—Gyalideopsis chicaque is known from a single but well-developed collection
found in a montane rainforest over compacted soil.
Etymology:—The epithet is a noun referring to the type locality.
Gyalideopsis chicaque is characterized by its small, broadly sessile, dark-brown apothecia lacking pruina
and very small, submuriform ascospores. There is no similar species growing on rock or soil. Gyalideopsis
nepalensis Vĕzda & Poelt has pruinose apothecia and larger ascospores, G. modesta Vĕzda & Poelt has palebrown apothecia, and G. lecideina Kalb & Vĕzda has black apothecia and larger ascospores (Lücking et al.
2006). Gyalideopsis mexicana Tretiach, Giralt & Vĕ zda, growing over bryophytes, has much larger, redbrown apothecia. The alpine, muscicolous G. tuerkii V ĕ zda is most similar to the new species but has
consistently larger apothecia and longer ascospores (15–25 µm, about 3 times as long as broad; Vĕzda 2003).
Gyrotrema papillatum Lücking, sp. nov. (Fig. 13F) Mycobank MB 517779
Sicut Gyrotrema wirthii sed apotheciis majoribus et thallo papilloso differt.—http://www.eol.org/pages/Gyrotrema
papillatum
Type:—COSTA RICA. Puntarenas: Corcovado National Park, Los Patos Section, Los Patos Station (Osa
Conservation Area), Osa Pensinsula, 160 km SSE of Sán José and 40 km WSW of Golfito, access trail to
station and first, hilly part (5 km) of trail to Sirena, 83°29’W, 8°34’N, 100–300 m, lowland rainforest zone,
closed primary forest, on bark (lower trunk), partially shaded, April 2003, Lücking 16301 (holotype F, isotype
INB).
Thallus grey-green to olive-green, with numerous white papillae, with dense, prosoplectenchymatous
cortex; photobiont layer and/or medulla with clusters of calcium oxalate crystals. Apothecia erumpent,
rounded to lobate, 1–1.5 mm diameter; disc exposed, cinnabar-red; margin lobulate to recurved, fused,
cinnabar-red inside. Columella absent but concentric rings of excipular tissue separating rings of old hymenia,
with youngest hymenium ring next to margin. Excipulum prosoplectenchymatous, carbonized; periphysoids
absent. Hymenium 80–100 µm high; paraphyses unbranched. Ascospores 8/ascus, 5–9-septate, 25–30 × 6–8
µ m, ellipsoid, with thick septa and lens-shaped lumina, colourless, I+ violet-blue (amyloid). Secondary
chemistry: apothecial disc with unidentified anthraquinone.
Distribution and habitat:—Gyrotrema papillatum is known from a single, well-developed collection
found in the lowland rainforest of southern Costa Rica.
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Etymology:—The epithet refers to the papillose thallus.
Gyrotrema papillatum closely resembles G. wirthii and has the same apothecial pigment. However, its
apothecia are larger and more irregular and its thallus is densely white-papillose. Whereas G. wirthii has been
found in the southern part of Osa Pensinsula, G. papillatum is known from a rich collection from the central
part close to the Golfo Dulce. It thus seems that in this case, also a microgeographic differentiation supports
the separation of these two species.
Harpidium gavilaniae Amo, Pérez-Ortega & A. Crespo, sp. nov. (Fig. 14A–B) Mycobank MB 517780
Sicut Harpidium rutilans sed thallus effiguratus et areolae majores..—http://www.eol.org/pages/Harpidium gavilaniae
Type:—SOUTH AFRICA. North Cape Province: Namaqualand, 57 km E of Springbok, Kangna's Farm,
29º34’S, 18º20’E, 1036 m, quartzite creek area, June 2005, Crespo & Amo de Paz SA-80b (holotype MAFLich 16488, isotype MAF-Lich 16489).
Thallus crustose, effigurate. Thallus rounded to ellipsoid, up to 2.5 cm in diameter. Areoles independent,
flat to bullate or sub-stipitate, affixed to substrate over most of lower surface; marginal zone usually free,
maroon-purplish to copper-red. Peripheral areoles enlongated, forming lobes, 0.4–0.9(–1.3) mm in diameter.
Central areoles rounded, bearing immersed apothecia, 0.3–0.6(–0.8) mm in diameter (n = 35). Areoles
stratified in section, 0.2–0.4(–0.7) mm high. Upper cortex paraplectenchymatous, to 25 µm high; Photobiont
layer continuous, 90–200 µ m high; Photobiont chlorococcoid, Trebouxia-like, 6–15 µ m in diameter.
Apothecia irregularly rounded, usually one per areole, immersed as in typical Aspicilia species, disc blackish,
0.1–0.3(–0.4) mm in diameter. Thalline exciple up to 20 µm thick. Epihymenium reddish, K–, ca. 10 µm
thick; hymenium 50–70 µm high; KI+ blue, paraphyses, moniliform, 4–6 µm wide, slightly wider at the apex.
Asci unitunicate-rostrate, clavate, 8-spored, 24–27 × 11–14 µm (n = 30), ascus wall thick. Ascospores hyaline,
simple, crescent form with the two apices rotated 90º, 9–13 × 6–7 µm (n = 35), with thick wall (ca. 0.8 µm).
Pycnidia frequent, laminal, immersed; 70–120 µm in diameter. Conidia cylindrical to ellipsoidal, 3 x 1 µm.
Secondary chemistry: no secondary metabolites detected by TLC.
Distribution and habitat:—Harpidium gavilaniae is so far only known from the type locality, growing
on quartzites in vertical furrows (dihedral) with seasonal water flows. The area belongs to Namaqualand
Hardevel biome (Mucina & Rutherford 2006) and it is very dry (below 100 mm per year).
Etymology:—This species is named for Rosario Gavilán, Botanist at the Universidad Complutense de
Madrid (Spain) who took part in our lichenologial expedition to South Africa in 2005.
Harpidium is a small genus, with only two species described so far. Harpidium rutilans is known from
few localities in the Eurasian mountains (Sancho & Crespo 1983) and H. nashii from four localities of SW
North America (Schultz et al. 2000). Harpidium and Euopsis together comprise the family Harpidiaceae
(Scheidegger & Schultz 2004). The new taxon represents the first record of the genus in the Southern
Hemisphere. The new species differs from the other two known species of Harpidium in its thallus shape. It
forms effigurate thalli, up to 2.5 cm in diameter. Harpidium rutilans is the most similar species but we have
seen elongated areoles only in young thalli; subsequently the thallus becomes irregular with areoles dispersed
over the surface. On the contrary, H. gavilaniae remains ellipsoid in shape and has elongated peripheral lobes.
Harpidium nashii differs in having umbilicate areoles and a more dispersed thallus. These differences,
together with the allopatric distribution and the genetic distance between populations (unpubl. data), in our
opinion justify description of the South African population of Harpidium as a new species.
Hypogymnia amplexa Goward, Björk & Wheeler, sp. nov. (Fig. 14C) Mycobank MB 517781
Sicut Hypogymnia imshaugii sed lobis peripheralibus adnatis, thallo centrali cerebriformi et acido physodalico
decipienti differt. Corticola.—http://www.eol.org./pages/Hypogymnia amplexa
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Type:—U.SA. Washington: Spokane County, Riverside State Park, 6.5 km NW of downtown Spokane,
47˚42’N, 117˚30’W, 525 m, on branches of Pinus ponderosa, July 2009, Björk 19340 (holotype UBC,
isotypes B, CANL, GZU).
FIGURE 14. A–B. Harpidium gavilaniae (holotype). A. Habit. B. Hymenium detail. C. Hypogymnia amplexa
(holotype), thallus with apothecia. D. Hypotrachyna guatemalensis (holotype), thallus with apothecia. E–F.
Hypotrachyna indica (holotype). E. Habit and showing fragile and flaking upper cortex. F. Detail showing whitemaculate upper surface. Scale in A, C, D = 1 mm, in B = 10 µm, in E, F = 2 mm.
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Thallus foliose, up to 6–8(–10) cm across, corticate above and below; lobes hollow, usually readily
divisible into two morphologically distinct lobe types: peripheral and central. Peripheral lobes short, usually
about as broad as long, nearly or quite contiguous, usually flaring outward to the (rounded) tips, tending to
wrap around the supporting branch, broadest ones 2–3(–4) mm wide, sometimes with small subapical
perforations, these usually over the upper surface, but sometimes also at the lobe tip. Central lobes sparse to
numerous, separate, more or less elongate, at least partly ascending, branching at first isotomic dichotomous,
but later predominantly anisotomic, terminating at least in part in rather pointed tips, broadest ones 1.5–2.5
mm wide, occasionally with small lateral or axillary perforations (check near margin of lower surface). Upper
surface whitish to pale pastel green, rather dull, becoming brownish where exposed, without black mottling,
more or less rugose throughout, often becoming cerebriform toward thallus centre. Medullary ceiling white
(except weakly brown near ruptures in lower surface). Lower surface black; thin, easily torn, sharply wrinkled
or folded, without rhizines. Soredia and isidia absent. Photobiont trebouxioid. Apothecia laminal, sparse to
more often numerous, stalked, the stalks pale, often scabrid, the excipulum not much expanded (except
sometimes expanded in age), often incurled at the rim; disk brown, the largest ones (2–)3–6(–8) mm across,
deeply cup-shaped, the outer portions usually vertically oriented; ascospores ellipsoid, averaging (6.8–)8.0(–
9.0) × (4.8–)5.7(–7.0) µm, L/W ratio (1.11–)1.41(–1.67); pycnidia copious over the upper surface forming
tiny black dots; conidia (6.5–)6.9(–8.0) × 1 µm. Secondary chemistry (TLC): atranorin, physodic acid (major,
constant), unknown 7 (present, occasional); cortex K+ yellow, medulla KC+ abruptly pink.
Distribution and habitat:—Currently known only from western intermontane North America, especially
eastern Washington, north and central Idaho, western Montana, rarely north to southern intermontane British
Columbia. Most specimens seen were collected from the branches of Pinus ponderosa, with occasional
material also from Abies lasiocarpa, Artemisia tridentata, Pinus albicaulis and Pseudotsuga menziesii.
Usually restricted in well-illuminated sites. This is the most frequently encountered Hypogymnia in some
portions of eastern Washington and adjacent Idaho, sometimes a dominant macrolichen of conifer branches in
open, dry forests.
Etymology:—Amplexa is from Latin, meaning "embracing" or "clasping": a reference to the peripheral
lobes, which tend to wrap around the supporting twig or branch.
Additional specimens examined (paratypes):—CANADA. British Columbia: Okanagan River
Drainage, Three Lakes area, 8 km N of Oliver, Björk 15123 (UBC). U.S.A. Idaho: Shoshone County, Coeur
d'Alene Mountains, Bloom Peak, Björk 16742a (UBC). Montana: Lake Co., Jocko Canyon Road, Wheeler
1834 (UBC). St. Marys Peak, Wheeler 836 (UBC). Missoula County, N fork of Finley Creek, Wheeler 2379
(UBC). Whelp Lake, Mission Mtns., Wheeler 1418 (UBC). Washington: Grant Co., Northup Canyon, Grand
Coulee, Björk 17713 (UBC). Lincoln County, Telford Road, 16 km SE of Creston, Björk 19095 (UBC).
Spokane Co., Kentuck Trails Road, 9.5 km W of Fairfield, Björk 19182 (UBC). Spokane Co., Turnbull
National Wildlife Refuge, 11 km S of Cheney, Björk 8946 (UBC). Spokane County, Riverside State Park, 7
km NW of downtown Spokane, Björk 19334 (UBC). Spokane County, Peone Prairie, near Moffat Road, 17
km NE of downtown Spokane, Björk 19196 (UBC). Spokane County, Dishman Hills County Park, 1 km SW
Dishman, Björk 10129 (UBC).
Hypogymnia is a well known genus of about 90 foliose lichens found primarily at temperate to boreal
latitudes, mostly on trees. Most species are whitish above, hollow within, black below, and lack rhizines.
Within the genus, H. amplexa is characterized by its combination of ascending, irregularly branched central
lobes that taper to a blunt point, and appressed, flattened, apically rounded peripheral lobes that tend to wrap
around the supporting branch, its strongly rugose (cerebriform) central portions, its deeply cup-shaped
apothecia, and by its white medullary ceiling and P– medulla. Hypogymnia amplexa appears to be most
closely allied with H. imshaugii Krog, with which it sometimes co-occurs. Hypogymnia imshaugii is a
morphologically variable species described from British Columbia (Krog 1968), and endemic to western
North America (Brodo et al. 2001). Both it and H. amplexa lack soredia, bear copious apothecia, and have
long, narrow, divergently branched, ascending lobes, at least in central portions of the thallus. In H imshaugii
Krog, however, the lobes are evenly branched, the apothecia tend to be shallowly saucer-shaped, and the
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medulla is P+ orange. Additionally the central lobes of H. imshaugii are smooth to at most weakly rugose
above and scarcely differ in habit from the (typically ascending) peripheral lobes. Subalpine populations of H.
amplexa might be confused with the "swollen form" of H. wilfiana Goward, T.Sprib. & Ahti (Goward et al.
2010) which likewise has appressed lobes and a white medullary ceiling. Such material, however, lacks
subapical perforations and ascending central lobes, has shallowly saucer-shaped apothecia, and contains
apinnatic acid. An unusual feature of H. amplexa is the occasional presence of perforations in the upper
cortex, especially in the vicinity of the lobe tips. Laminal perforations in Hypogymnia have hitherto been
reported only in H. magnifica X.L.Wei & McCune, a southwest Asian endemic lichen clearly unrelated to our
new species (Wei et al. 2010).
Hypotrachyna guatemalensis Elix & van den Boom, sp. nov. (Fig. 14D) Mycobank MB 517782
Sicut Hypotrachyna bahiana sed diminuta, medulla albida, rhizinis densis et ramosissimis et acidum
conhypoprotocetraricum et acidum hypoprotocetraricum continente differt.—http://www.eol.org/pages/
Hypotrachyna guatemalensis
Type:—GUATEMALA. Quezaltenango: S of Llano del Pinal, N slope of volcano Santa María, 14°47’N,
91°33’E, 2700 m, on Alnus arguta along path among small agriculture fields with small forests, shrubs, trees
and outcrops, July 2004, van den Boom & van den Boom 32993 (holotype CANB, isotype hb. van den Boom).
Thallus foliose, 4–5 cm wide, adnate to loosely adnate, coriaceous, irregularly lobate. Lobes contiguous to
sparingly imbricate, sublinear to subirregular, 1–4 mm wide, with entire to crenulate margins and subrotund
apices, eciliate but with rhizines projecting beyond the margins in part. Upper surface grey-white to yellowish
grey, emaculate, shiny and black-margined at the apices but dull and rugulose in the centre, lacking soredia,
isidia, pustules, dactyls, and lobules. Medulla white. Lower surface black but brown at the apices, rhizines
dense, richly dichotomously branched. Apothecia common, sessile to substipitate, 1–4 mm diameter, disc pale
brown, exciple smooth, margins thick and persistent. Ascospores broadly ellipsoid, 7–9 × 5–6 µm. Pycnidia
common, black, immersed. Conidia weakly bifusiform, 6–8 × 1 µm. Secondary chemistry: atranorin (minor),
chloroatranorin (minor), conhypoprotocetraric acid (major), hypoprotocetraric acid (minor), 4-Odemethylnotatic acid (trace).
Distribution and habitat:—At present, H. guatemalensis is known only from the type collection.
Etymology:—The specific epithet derives from the Latin -ensis (place of origin) and the location of the
type.
Morphologically this new species resembles Hypotrachyna bahiana (Nyl.) Hale (1975), but is
distinguished in having a white medulla (partially pigmented orange or yellow-brown in H. bahiana), smaller
thallus (4–5 cm wide versus 6–20 cm) with dense, richly dichotomously branched rhizines (branched 3–5
times), whereas the rhizines in H. bahiana are sparse to moderately dense and moderately dichotomously
branched (branched 1–3 times). The two species also differ chemically, since H. bahiana contains atranorin
(minor), chloroatranorin (minor), protocetraric acid (major), virensic acid (minor or trace), ±gyrophoric acid
(trace), ±conprotocetraric acid (trace) and the pigments skyrin (minor) ±pigmentosin A (major or minor)
(Sipman et al. 2009). In contrast, H. guatemalensis lacks pigments and contains atranorin (minor),
chloroatranorin (minor), conhypoprotocetraric acid (major), hypoprotocetraric acid (minor), 4-Odemethylnotatic acid (trace). The rare depsidone, conhypoprotocetraric acid, was first isolated from a
Panamanian collection of Relicina cf. incongrua Hale (Elix et al. 1995). Conhypoprotocetraric acid is not
known to occur as a major component in any other taxon in Parmeliaceae.
Hypotrachyna indica Divakar, Lumbsch, Upreti & A.Crespo, sp. nov. (Fig. 14E–F) Mycobank MB 518238
Sicut Hypotrachyna exasplendens sed differt acidum barbaticum, 4-O-demethylbarbaticum, obtusaticum, et
norobtusaticum continente, et pustulata, sorediis destituta..— http://www.eol.org/pages/Hypotrachyna indica
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Type:—INDIA. Tamil Nadu: Western Ghat, Nilgiri Hills, Dodabetta, trail from Samer to Tiger Hills, 11º
24’N, 76º 44’E, 2607 m, on coniferous tree trunk, Divakar et al. 19715K (holotype MAF-Lich, isotypes F,
LWG).
Thallus corticolous, adnate, small, to 5 cm across. Lobes short, sublinear, dichotomously branched with
truncate apices, 0.5–2 mm wide. Upper surface greenish grey, dull, strongly white-maculate, sometimes
reticulate near lobe apices, pustulate, without soredia and isidia; upper cortex fragile and flaking, with
blackened, eroded areas. Pustules subterminal, erupting apically, not forming soredia but may shed corticated
granules from open pustule margins. Medulla white. Lower surface black with narrow brown margins,
rhizinate, rhizines richly dichotomously branched, to 1.5 mm long. Apothecia and pycnidia not seen.
Secondary chemistry: Cortex K+ yellow; medulla K–, C–, KC+ orange, P–; containing atranorin, barbatic
acid, 4-O-demethylbarbatic acid, obtusatic acid and norobtusatic acid (trace).
Distribution and habitat:—The species grows on tree trunk in open and humid coniferous forest at 2607
m elevation. At present the taxon is only known from type locality.
Etymology:—The specific epithet refers to the name of the locality.
Additional specimens examined (paratypes):—INDIA. Tamil Nadu: Western Ghat, Nilgiri Hills,
Dodabetta, trail from Samer to Tiger Hills, Divakar et al. 19715O, 19718P (MAF-Lich).
Hypotrachyna indica is characterized by a strongly white-maculate upper surface, flaking upper cortex,
pustules that do not form soredia and the presence of the barbatic acid complex. This new species resembles
Hypotrachyna exasplendens (Hale) Hale, reported from Central America and South Africa (Hale 1975), but
differs in containing barbatic acid, 4-O-demethylbarbatic acid, obtusatic acid and norobtusatic acid instead of
alectoronic acid; and in having pustulate upper surface, which break open apically but do not produce soredia
rather than capitate soralia. It is also similar to Hypotrachyna exsecta (Taylor) Hale, known from Australasia
and South East Asia (Elix 1994a, Divakar & Upreti 2003, 2005), but the latter has an emaculate, shiny, and
sorediate upper surface, a continuous upper cortex (not flaking), and broad (2–5 mm wide) lobes.
Hypotrachyna lueckingii Sipman, sp. nov. (Fig. 15A) Mycobank MB 517783
Hypotrachyna corticola thallo flavo, sine propagulis vegetativis, acida protocetraricum et gyrophoricum in medulla
continens.— http://www.eol.org/pages/Hypotrachyna lueckingii
Type:—COSTA RICA. Cartago: Cordillera de Talamanca, Cerro de la Muerte, Cerro de la Asunción, along
Carretera Interamericana near sideway to the peak, 83°46’W, 9°34’N, 3350 m, subpáramo scrub, ca. 2 m tall,
epiphyte on shrub branches, April 1998, Sipman et al. 43698 (holotype B, isotype INB).
Thallus foliose, corticolous, 5–10 cm wide, loosely adnate, not coriaceous, dichotomously to
subdichotomously lobate. Lobes sublinear to subirregular, mostly separate, (0.5–)1.5–3 mm wide, plane to
slightly convex, with subtruncate apices. Upper surface greenish yellow, slightly shiny, epruinose, emaculate,
lacking soredia, isidia, pustules, dactyls or lobules, and with intact upper cortex. Medulla white, with a
continuous algal layer above. Lower surface black, peripherally dark brown. Rhizines black, rather dense,
sometimes forming a marginal fringe, ca. 0.5 mm long, 1–4 times dichotomously branched. Apothecia
frequent, 2–7 mm in diameter, with concave, finally flat, brown, glossy discs and crenulated, incurved
margins. Ascospores ellipsoid, 10–13 × 6–7 µm. Pycnidia frequent, laminal, immersed. Conidia weakly
bifusiform, ca. 6 × 0.5 µm. Chemistry: upper cortex K–, C–, KC+ yellow, P–, UV–, medulla K–, C–, KC–, P+
orange-red, UV–. Secondary metabolites: upper cortex with usnic acid (major), medulla with protocetraric
acid (major) and gyrophoric acid (minor).
Distribution and habitat:—Known so far only from two specimens from the highest parts of the
Cordillera de Talamanca in Costa Rica, on scattered trees in páramo scrub at 3300–3500 m.
Etymology:—The epithet honours Robert Lücking (Chicago, U.S.A.), who contributed greatly to the
lichenological knowledge of Costa Rica by, e.g., his detailed studies of foliicolous lichens and the
organisation of the TICOLICHEN project.
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FIGURE 15. A. Hypotrachyna lueckingii (holotype), substrate with thalli. B. Hypotrachyna paracitrella (holotype),
thallus lobes enlarged with apothecium. C. Hypotrachyna paraphyscioides (holotype), thallus lobes enlarged. D.
Hypotrachyna parasinuosa (holotype), thallus lobes enlarged. E–F. Icmadophila eucalypti (E, holotype; F, Kantvilas 21/
04). E. Habit (photograph J. Jarman). F. Detail of lobe apices; the swellings resemble apothecial initials of other members
of the Icmadophilaceae (photograph J. Jarman). Scale in A = 10 mm, in B–D = 3 mm, in E = 1 mm, in F = 0.5 mm.
Additional specimens examined (paratypes):—COSTA RICA. Sán José: Cerro Buenavista, Hafellner
7747 (GZU).
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During the recent revision of Hypotrachyna (Sipman et al. 2009), the number of species was doubled in
comparison with an earlier revision (Hale 1975). Nevertheless the diversity of this conspicuous genus in the
Neotropics is still not completely understood and additional taxa continue to be described (Yánez-Ayabaca &
Eliasaro 2009). Hypotrachyna lueckingii is very similar to H. flavida (Zahlbr.) Hale, with which it shares
cortical usnic acid and medullary protocetraric acid as major substances, and the absence of vegetative
reproduction (soredia, isidia, pustules, flaking cortex, deciduous rhizines). It is clearly distinguished by the
presence of gyrophoric acid in the medulla, and its less linear thallus, which resembles more the isidiate H.
microblasta (Vain.) Hale or the sorediate H. sinuosa (Smith) Hale. Different also is its corticolous habitat,
while H. flavida is mostly found on rock or on moss covering rock. Of the latter only one corticolous
specimen is known from Colombia (Aguirre & Sipman 5989 (B), which differs clearly from the new species
in the absence of gyrophoric acid and the sublinear lobes. Further, H. lueckingii is allopatric with H. flavida,
which is not known further north than Colombia. So far morphological analysis has not given clear indications
for an infrageneric classification of Hypotrachyna, and Sipman et al. (2009) did not recognize any
infrageneric categories. However, recent DNA analyses (Divakar et al. 2006) suggest that several distantly
related groups are involved. In the absence of molecular data, the position of H. lueckingii within
Hypotrachyna s.lat. remains uncertain. However, its morphological similarity with H. sinuosa would indicate
an affinity with clade D of Divakar et al. (2006: fig. 1), while H. flavida might belong to clade B.
Hypotrachyna paracitrella Sipman & Palice, sp. nov. (Fig. 15B) Mycobank MB 517784
Hypotrachyna sine rhizinis, cum ciliis furcatis, corticola thallo flavo, sine propagulis vegetativis, acida barbaticum,
obtusaticcum, 4-O-demethylbarbaticum et norobtusaticum in medulla continens.—http://www.eol.org/pages/
Hypotrachyna paracitrella
Type:—ECUADOR. Tungurahua. Llanganates National Park, Cordillera Llanganates, ca. 5 km WSW of
Cerro Hermoso, 01°15’S, 78°20’W, 3750–3800 m, well-lit dwarfy forest at steep, E-facing slope above
unnamed lagoon, March 2003, Palice 8487 (holotype PRA, isotype B, QCA).
Thallus foliose, corticolous, ca. 5 cm wide, loosely adnate, not coriaceous, dichotomously to
subdichotomously lobate, ciliate. Lobes sublinear to subirregular, mostly separate, 2–3 mm wide, plane to
slightly convex, with subtruncate apices. Upper surface greenish yellow with usually a black marginal rim,
slightly shiny, epruinose, emaculate, lacking soredia, isidia, pustules, dactyls or lobules. Medulla white, with a
continuous algal layer above. Lower surface black, paler and brown at the lobe tips. Rhizines absent. Marginal
cilia present, black, somewhat dendroid, 1–2 mm long, near the base about 0.2 mm thick, 1–3 times furcately
branched. Apothecia frequent, 2–5 mm in diameter, with strongly constricted base, with concave, finally flat,
brown, glossy discs, and with crenulated, incurved margins. Ascospores ellipsoid, 12–13 × 6–7 µm. Pycnidia
frequent, laminal, immersed. Conidia not seen. Secondary chemistry: upper cortex K–, C–, KC+ yellow, P–,
UV–; medulla K+ yellow turning red, C–, KC–, P+ orange-red, UV–; upper cortex with usnic acid (minor),
medulla with salazinic acid (major), consalazinic acid (minor), norstictic acid (trace) and protocetraric acid
(trace) (HPLC kindly provided by J.A. Elix).
Distribution and habitat:—Known so far only from a single specimen in Tungurahua province,
Ecuador, in humid dwarfed forest at 3750–3800 m.
Etymology:—The epithet reflects the similarity to Hypotrachyna citrella (Kurokawa) Hale (Sipman et al.
2009).
After revisions of neotropical representatives of the genera Everniastrum (Sipman 1986a) and
Hypotrachyna (Sipman et al. 2009), these can now be identified routinely, which increases the chances for the
discovery of rare species. H. paracitrella is apparently one such species. It is known only from a single
collection, although the macrolichens of Ecuador have been sampled extensively by several trained
lichenologists, in particular L. Arvidsson from Gothenburg, Sweden. As expressed by its name, H.
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paracitrella is most similar to H. citrella. It agrees in lobe configuration, absence of vegetative reproduction,
and chemistry, and differs in the absence of rhizines and the presence of furcately branched cilia.
Hypotrachyna as currently circumscribed is polyphyletic as was shown/confirmed by recent molecular studies
(Divakar et al. 2006, 2010). If the genus is to be divided, then the two newly described taxa would perhaps
deserve affiliation to a separate genus because of the combination of unique features. Hypotrachyna
paracitrella and two other species treated here, H. paraphyscioides and H. parasinuosa, are unusual because
they lack rhizines and have furcately branched cilia instead. In this way they fall outside the traditional limit
of the genus Hypotrachyna and fit rather the genus Everniastrum. Therefore they were left outside the recent
revision of neotropical Hypotrachyna (Sipman et al. 2009). However, by their lobe configuration and their
chemistry they deviate greatly from other representatives of Everniastrum and resemble more closely
particular species of Hypotrachyna, as expressed by their names. As a consequence more characters suggest
their affinity with species of Hypotrachyna rather than with species of Everniastrum species and they are
therefore consigned here to Hypotrachyna rather than to Everniastrum. A tendency to develop a naked lower
surface occurs also in H. everniastroides Sipman (Sipman et al. 1986b), where rhizines are restricted to a
marginal zone. Hypotrachyna as currently circumscribed is polyphyletic as was shown/confirmed by recent
molecular studies (Divakar et al. 2006, 2010). It is noteworthy that all species of Hypotrachyna with cilia
instead of rhizines known so far are very rare, and that two are known only from a single localityt. This is
somewhat reminiscent of hybrids in phanerogams, which show an unusual mix of characters from two
parents, and are usually much rarer than either. It may also indicate relict endemism. It is also noteworthy that
the specimens of H. paracitrella and H. parasinuosa grow mixed on the same tree.
Hypotrachyna paraphyscioides Sipman, sp. nov. (Fig. 15C) Mycobank MB 517785
Hypotrachyna corticola, sine rhizinis, cum ciliis furcatis, thallo cinereo, sine propagulis vegetativis, acida barbatica,
obtusatica, 4-O-demethylbarbatica et norobtusatica in medulla continens.—http://www.eol.org/pages/
Hypotrachyna paraphyscioides
Type:—COLOMBIA. Magdalena: Sierra Nevada de Santa Marta, transecto del Alto Buritaca, 2880 m, on
branch, August 1977, Rangel et al. 1083b (holotype L).
Thallus foliose, corticolous, 5–10 cm wide, loosely adnate, not coriaceous, dichotomously to
subdichotomously lobate, ciliate. Lobes sublinear to subirregular, mostly separate, (0.5–)1.5–3 mm wide,
plane or slightly convex, with subtruncate apices. Upper surface pale grey with black marginal rims, slightly
shiny, epruinose, maculate, lacking soredia, isidia, pustules, dactyls or lobules, and with intact upper cortex.
Medulla white. Lower surface black, pale brown towards the lobe tips. Rhizines absent. Marginal cilia
present, black, 1–3 mm long, somewhat flexuous and 1–2 times furcately branched at rather wide angles,
occasionaly also present away from the margin on the lower side. Apothecia frequent, 2–7 mm in diameter,
with concave, finally flat, brown, glossy discs and crenulated, incurved margins. Ascospores ellipsoid, 10–13
× 6–7 µm. Pycnidia frequent, laminal, immersed. Conidia weakly bifusiform, ca. 6 × 0.5 µm. Secondary
chemistry: upper cortex K+ yellow, C–, KC–, P–, UV–; medulla K–, C+ yellow-orange, KC+ orange, P–,
UV–; upper cortex with atranorin, medulla with barbatic acid (major), obtusatic acid (trace), 4-Odemethylbarbatic acid (major) and norobtusatic acid (trace) (TLC).
Distribution and habitat:—Known so far only from one specimen from the N-slope of the Sierra
Nevada de Santa Marta in Colombia, on a branch in montane forest at 2880 m.
Etymology:—The epithet reflects the similarity to Hypotrachyna physcioides (Nyl.) Hale (Sipman et al.
2009).
Hypotrachyna paraphyscioides resembles H. physcioides closely in lobe configuration, presence of
soralia, and chemistry (Sipman et al. 2009), and differs in the presence of furcately branched cilia and the
absence of rhizines.
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Hypotrachyna parasinuosa Sipman & Palice, sp. nov. (Fig. 15D) Mycobank MB 517786
Hypotrachyna corticola sine rhizinis, cum ciliis furcatis, thallo flavo, soraliis capitatis, acidum salazinicum in medulla
continens.— http://www.eol.org/pages/Hypotrachyna parasinuosa
Type:—ECUADOR. Tungurahua. Llanganates National Park, Cordillera Llanganates, ca. 5 km WSW of
Cerro Hermoso, 01°15’S, 78°20’W, 3750–3800 m, well-lit dwarfy forest at steep, E-facing slope above
unnamed lagoon, March 2003, Palice 8487b (holotype PRA).
Thallus foliose, corticolous, ca. 5 cm wide, loosely adnate, not coriaceous, dichotomously to
subdichotomously lobate, ciliate. Lobes sublinear to subirregular, mostly separate, 2–3 mm wide, plane to
slightly convex, with subtruncate apices. Upper surface greenish yellow with usually a black marginal rim,
slightly shiny, epruinose, emaculate, sorediate, lacking isidia, pustules, dactyls or lobules. Soralia apical or
subapical, more or less capitate or spreading over the surface, pale yellow but becoming blackened with age,
with granular soredia ca. 60 µm in diameter. Medulla white. Lower surface black, paler and brown at the lobe
tips. Rhizines absent. Marginal cilia present, black, somewhat dendroid, 1–2 mm long, near the base about 0.2
mm thick, 1–3 times furcately branched. Apothecia and pycnidia not seen. Secondary chemistry: upper cortex
K–, C–, KC+ yellow, P–, UV–; medulla K+ yellow turning red, C–, KC–, P+ orange-red, UV–; upper cortex
with usnic acid (minor), medulla with salazinic acid (major), consalazinic acid (minor), norstictic acid (trace)
and protocetraric acid (trace) (HPLC by J. A. Elix).
Distribution and habitat:—Known so far only from a single specimen in prov. Tungurahua, Ecuador, on
humid, dwarfed forest at 3750–3800 m.
Etymology:—The epithet reflects the similarity to Hypotrachyna sinuosa (Taylor) Hale (Sipman et al.
2009).
Hypotrachyna parasinuosa resembles H. sinuosa closely in lobe configuration, presence of soralia, and
chemistry (Sipman et al. 2009), and differs in the presence of furcately branched cilia and the absence of
rhizines. For further comments see under H. paracitrella.
Icmadophila eucalypti Kantvilas, sp. nov. (Fig. 15E–F) Mycobank MB 517787
Species corticola, squamulosa vel subfruticulosa, albida, acidum thamnolicum continens, sorediata, squamis 1–3.5(–5)
µm latis, flabellatis, a thallo primario effuso, ephemero, albido enatis, ascomata conidiomataque ignota sed huic
generi ob materiam chemicam, anatomiam morphologiamque adsignata.—http://www.eol.org/pages/Icmadophila
eucalypti
Type:—AUSTRALIA. Tasmania: Hartz Road near entrance to the National Park, 43°12’S, 146°47’E, 570 m,
on moist trunks of old Eucalyptus obliqua in mixed forest, July 2007, Kantvilas 285/07 (holotype HO, isotype
BM).
Thallus squamulose, whitish to pale grey, sometimes with a pale bluish, creamish or beige tinge, forming
extensive, irregular colonies to 50 cm wide. Individual squamules 1–3.5(–5) µm wide, unevenly 130–350 µm
thick and densely inspersed with crystals that fluoresce in polarised light and dissolve in KOH, arising from
an effuse, very thin and transient, white primary thallus, scattered and discrete, or imbricate, rarely fusing
together, stellate, rosette-like or, more typically, flabellate and with one side tightly adnate to the substratum
and the other ascending, initially with crenulate, rather thickened margins, soon becoming nodulose, lobulate
or palmately lobed and ± subfruticulose, sorediate; upper surface matt, smooth or minutely granular-scabrid
(lens), in section with a pseudocortex 20–30 µm thick comprising randomly orientated, short-celled hyphae
3–5 µm wide, interspersed with occasional dead algal cells; lower surface white, ecorticate; soredia coarsely
granular, 40–80 µm diameter, concolorous with the thallus, originating at the lower surface of the squamule
margins, soon eroding the margin somewhat and spreading acrosss the upper surface. Photobiont a unicellular
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green alga with globose cells 5–11 µm diameter. Ascomata not seen. Pycnidia not seen. Secondary chemistry:
thamnolic acid; thallus K+ vivid yellow, P+ orange, KC–, C–, UV–.
Distribution and habitat: Icmadophila eucalypti is widespread in Tasmania where it occurs in wet
eucalypt rainforest and cool temperate rainforest from lowland to subalpine elevations. It occurs on the lower,
shaded parts of trunks of very large trees with a thick, fibrous bark. Eucalyptus obliqua L'Hérit., a common
dominant in many Tasmanian wet eucalypt forest communities, is by far the preferred host species, although
the fire-sensitive, rainforest conifer, Athrotaxis selaginoides Don, is also sometimes colonised. The new
species forms extensive patches on the moist sides of such trees, where it associates with several other lichens
with a similar squamulose growth form: Cladonia rigida (Hook.f. & Taylor) Hampe, Neophyllis melacarpa
(F.Wilson) F.Wilson and Cladia schizopora (Nyl.) Nyl. Its predeliction for very old trees suggests that it may
serve as an old forest indicator in ecological studies.
Etymology: The specific epithet refers to the preferred host of the new species.
Additional specimens examined (paratypes):—AUSTRALIA. Tasmania: Algonkian Mountain,
Kantvilas 145/90 (HO); West of Tahune Bridge, Kantvilas 21/98 (BM, HO); Weindorfers Forest, Mar1998,
Kantvilas s.n. (HO); Mueller Road, Kantvilas 11/98 (HO); track to Wylds Craig, Kantvilas 616/03 (HO); Lake
Helen, Kantvilas 21/04 (HO, K); Florentine Road, Kantvilas 306/05 (HO, K); W of Tahune Bridge, Coupe 8I,
Kantvilas 275/07 (BM, HO, MSC).
Icmadophila is a small, well-circumscribed genus of four species, characterised, inter alia, by: a crustose
to squamulose thallus containing thamnolic acid, a Coccomyxa-type photobiont, sessile to stipitate ascomata
and Icmadophila-type asci (Rambold et al. 1993). In the absence of fertile material, the inclusion of I.
eucalypti in this genus was based on an evaluation of morphological, anatomical, ecological and chemical
characters. Several attempts to extract DNA for molecular confirmation of this classification (by Dr H.
Döring, Kew) were unsuccessful. Thus for the time being, this classification is provisional. However, the
general morphology, anatomy and chemistry of the species are all consistent with what is found in
Icmadophila as exemplified by the type species, I. ericetorum (L.) Zahlbr., I. japonica (Zahlbr.) Rambold &
Hertel and the Australasian endemic, I. splachnirima (Hook.f. & Taylor) D.J.Galloway. The interpretation of
the primary thallus of the new species is equivocal. Thin sections suggest that it is lichenised, albeit very
weakly, although it does not seem to contain thamnolic acid. There is a possibility that this feature is better
interpreted as a prothallus rather than a true thallus horizontalis, although the latter is found frequently in the
family Icmadophilaceae. Occasional thalli of I. eucalypti also develop pinkish, gall-like thickenings on the
surface and at the margins which are highly reminiscent of the ascomatal initials of other members of this
family, including Dibaeis and Siphulella. However, sections did not reveal any early development of
ascomatal tissues.
Krogia microphylla Timdal, sp. nov. (Fig. 16A–B) Mycobank MB 517788
Krogiae coralloidi et K. antillari affinis, sed squamis minoribus, hypothecio obscure olivaceo et thallo acidis lichenosis
destituto.—http://www.eol.org/pages/Krogia microphylla
Type:—DOMINICAN REPUBLIC. El Ceibo: 25 km N of El Ceibo on road to Miches, 18 km S of Miches,
18°55’N, 69°09’W, January 1991, Harris 26749 (holotype NY).
Thallus squamulose, effuse. Squamules to 0.3 mm wide when young but soon coalescing into a
continuous, microphyllinous crust, deeply and irregularly divided into ca. 0.1 wide lobes, ascending,
irregularly imbricate, pale brown (old herbarium material), with scattered patches of orange (K+ purple)
spots, epruinose, dull; margin concolorous with upper side, not fibrillose; lower side pale brown; soredia and
isidia absent. Upper cortex poorly defined, composed of a thin, rather loose layer of irregularly oriented
hyphae. Algal layer filling the inner part of the squamule. Medulla and/or lower cortex poorly defined. No
crystals in thallus section (polarized light), PD–, K–, K/I–. Prothallus lacking. Apothecia biatorine, to 0.7 mm
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FIGURE 16. A–B. Krogia microphylla (holotype). A. Habit. B. Ascus in K/I. C–D. Lecanora mugambii (holotype). C.
Thallus with apothecia. D. Apothecia. E. Lecanora printzenii (holotype), apothecia on thalli of Umbilicaria. F. Lecanora
xanthoplumosella (holotype), thallus with apothecia. Scale in A, C–F = 1 mm, in B = 10 µm.
diameter when simple, but often forming up to 1 mm diameter aggregates, with a rusty brown, more or less
plane disc and a grayish brown, slightly shiny, not fibrillose, often flexuous margin. Proper exciple dark
olivaceous brown, K+ green in the inner part, pale brown to colourless in the rim, composed of radiating,
closely conglutinated, thick-walled hyphae with cylindrical lumina, not containing crystals. Hypothecium
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continuous with proper exciple, dark olivaceous brown, K+ green, composed of closely conglutinated, thickwalled hyphae with angular to cylindrical lumina, not containing crystals. Hymenium and epithecium
colourless, not containing crystals. Paraphyses simple or sparingly branched, moderately conglutinated, rather
thin-walled; apical cell slightly swollen, colourless. Ascus clavate, surrounded by a thin amyloid sheet; tholus
well-developed, not amyloid or with a wide, faintly amyloid tube, with a well-developed ocular chamber
when young. Ascospores eight in the ascus, colourless, thin-walled, filiform, curved, spirally arranged, simple
or with 1–3 pseudosepta, 25–35 × ca. 1.0 µm (n = 20). Secondary chemistry: no lichen substances (by TLC).
Distribution and habitat:—Krogia microphylla is known only from the holotype. The collecting site
was, according to the label, "cloud forest in ravine and scattered trees in pasture" at ca. 450 m elevation. Two
species of Phyllopsora were collected at the same site: P. chodatinica Elix (Harris 26683, NY) and P.
furfuracea (Pers.) Zahlbr. (Harris 26725, NY).
Etymology:—The species is named after the tiny squamules.
The genus Krogia and its originally sole species, K. coralloides Timdal, was described from one locality
in Mauritius (Timdal 2002). A second species, K. antillarum Timdal, was described from six localities in the
West Indies (Timdal 2008), and a third species is described here. Krogia is a corticolous genus occurring in
subtropical rainforests and closely resembling Phyllopsora. It differs from Phyllopsora in having a weak or
absent amyloid reaction in the tholus and filiform, curved ascospores which are spirally arranged in the ascus.
The new species differs from Krogia coralloides and K. antillarum in forming smaller squamules which do
not contain secondary substances, in having rusty brown apothecia with a grayish brown margin, and in
having a dark olivaceous brown, K+ green hypothecium and inner part of the proper exciple. Krogia
coralloides forms up to 1 mm wide squamules and contains boninic acid (Timdal 2002), and K. antillarum
forms up to 1.5 mm wide squamules and contains 4-O-methylcryptochlorophaeic acid (Timdal 2008); both
species have medium brown apothecia with a margin concolorous with or slightly paler than the disc, and a
pale-brown, K– hypothecium and inner part of the proper exciple. The ascus and ascospores of K. microphylla
are indistinguishable from those of the two other species.
Lecanora mugambii Kirika, I.Schmitt, Fankhauser & Lumbsch, sp. nov. (Fig. 16C–D) Mycobank MB
517789
A Lecanora glaucodea thallis crassis, ascosporis latis et acidum alectoronicum continente differt.—http://www.eol.org/
pages/Lecanora mugambii
Type:—KENYA. Western Province: Bukusu District, Mt. Elgon Forest, Kimilili Rd., 0°55’N, 34°38’E, large
boulders in remnant cloud forest, on siliceous rocks, January 2007, Lumbsch et al. 19559a (holotype EA,
isotypes F, MIN).
Thallus thin to thick, verrucose to verruculose-bullate, whitish grey. Thallus surface smooth, epruinose,
isidia and soredia absent. Prothallus not visible. Apothecia sessile, abundant, dispersed, plane or slightly
convex, roundish, 0.6–1.5 mm diameter. Apothecial margin lecanorine, concolourous with the thallus,
persistent, thick, entire when young, becoming slightly verruculose or remaining entire, disc grey to greybrown with a thick, granulose, whitish grey to bluish grey pruina. Amphithecial cortex distinct, gelatinous,
hyaline, inspersed with small crystals, uniform, 15–20 µ m thick. Amphithecium with large crystals.
Parathecium hyaline, 10–15 µm thick, with small crystals. Epihymenium red-brown to brown, 15–20 µm
thick, granulose with coarse crystals soluble in KOH. Hymenium, subhymenium and hypothecium hyaline.
Paraphyses sparingly branched and not thickened apically. Asci of Lecanora-type, 8-spored. Ascospores nonseptate, hyaline, broadly ellipsoid to subglobose, 11–15 × 7–10.5 µ m. Pycnidia immersed in verrucae,
blackish with filiform conidia. Spot tests: K+ yellow, C–; KC–, P+ yellow, UV+ blue. Secondary chemistry:
alectoronic acid [major], atranorin [submajor], chloroatranorin [minor], physodic acid [minor], usnic acid
[submajor], and unidentified alectoronic acid derivatives [minors].
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Distribution and habitat:—This species occurs on siliceous rocks in exposed habitats at the edge of a
remnant cloud forest. It is only known from the type locality at 2500 m elevation, where it is very common.
Associated species included Diploschistes scruposus, Heterodermia spp., Ingvariella bispora, Pertusaria
subventosa, Punctelia spp., Tephromela sp., and Xanthoparmelia spp.
Etymology:—The species is named after our colleague and friend George Mugambi (Nairobi), who has
made significant contributions to our knowledge of East African ascomycetes.
Lecanora mugambii is characterized by bluish grey pruinose apothecial discs, broadly ellipsoid to
subglobose ascospores, and the presence of alectoronic and usnic acids in addition to the atranorin
chemosyndrome. Alectoronic acid was not previously been reported from the Lecanora subfusca group
(Guderley 1999, Lumbsch 1994). The species may be confused with the neotropical Lecanora glaucodea Nyl.
The latter species is distinguished by a thin, never bullate thallus, smaller apothecia, an epihymenium with
small crystals, narrower ascospores, a basally thickened amphithecial cortex, and a different chemistry (Feige
et al. 2000, Guderley 1999). Another similar species is Lecanora formosula Lumbsch, which, however is
readily distinguished by having a thin thallus, only slightly pruinose apothecial discs, smaller ascospores, and
a different chemistry (Lumbsch et al. 1995).
Lecanora printzenii Pérez-Ortega, Vivas & Hafellner, sp. nov. (Fig. 16E) Mycobank MB 517790
Species habitu Lecanorae variae similis, sed in thallis Umbilicariarum lichenicola et in sequentia regionis ITS differt.—
http://www.eol.org/pages/Lecanora printzenii
Type:—SPAIN. Castilla y León: Burgos, Neila, Parque Natural de las Lagunas de Neila, 42º 03’N, 03º 03’W,
1868 m, on Umbilicaria cylindrica, August 2007, Pérez-Ortega 1204 (holotype MAF-Lich).
Thallus composed of areoles, ± dispersed or aggregated, esorediate; areoles irregularly rounded, up to 0.3
mm in diameter and height, convex, surface yellowish green, dull. Photobiont Trebouxia-like, cells up to 20
µm in diameter. Apothecia rounded, flexuose when mature, rarely single, often crowded; sessile, constricted
at the base, especially when mature, (0.3–)0.5–1(–1.3) mm in diameter; disc greenish when young but soon
becoming blackish, often pruinose when mature; margin prominent, persistent, matt and greenish when young
but soon at least partly becoming melanized, black and shiny, with the exception of a narrow band, the closest
part to the disc that remains greenish. Amphithecium corticated, up to 200 µm in the middle part, composed of
a medulla of irregularly entangled hyphae and crowded algae; cortex composed of hyphae with thick walls,
strongly gelatinized, with narrow lumina 0.5–1 µm; cortex 5–40 µm in the upper part, 20–70 µm in the middle
part, 25–50 µm in the lower part, colourless within, outer part melanized. Hypothecium 20–40 µ m high,
hyaline; subhymenium 5–15 µ m high; hymenium 50–65 µ m; colourless; epihymenium ca. 10 µ m, dark
blackish green, K–; paraphyses weakly branched, anastomosed, 1–2 µm wide, expanded at the apex up to 5
µm, with a blackish green pigment surrounding the apices. Asci clavate, of Lecanora–type and with rostrate
dehiscence, 35–40 × 9–14 µm (n = 10). Ascospores hyaline, simple, 7–11 × 4–6 µm (n = 40). Pycnidia not
seen. Secondary chemistry: K–, C–, P+ yellow (amphithecium) or P–.
Distribution and habitat:—The new species is so far known from two localities in Spain and one in
Austria. Lecanora printzenii lives lichenicolous on different species of Umbilicaria, causing serious damages
in the areas of the lichen where it grows.
Etymology:—The new taxon is named after Christian Printzen (Frankfurt, Germany) for his contribution
to the taxonomy of yellow Lecanoras in particular and to lichenology in general.
Additional specimens examined (paratypes):—SPAIN. Castilla y León: Salamanca, Sierra de Francia,
June 2009, Lumbreras & Roca s.n. (MAF). AUSTRIA. Steiermark: Niedere Tauern, Seckauer Tauern, Groβer
Ringkogel NW von Knittelfeld, NE-Abhänge ober dem Sundlsee, Hafellner 75167 (GZU, MAF; to be
distributed in Hafellner, Lichenicolous Biota).
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The taxonomy of the Lecanora varia group was recently revised in several papers (Printzen 2001, Śliwa
& Wetmore 2000, Laundon 2003, Martínez & Aragón 2004) and a phylogeny of the group was established
(Pérez-Ortega et al. 2010). The new species is closely related to other species of the group, such as L. varia, L.
burgaziae, and L. densa, but differs in its lichenicolous habitus, the at least partly melanized excipulum, the
finally blackish disc, and the ITS sequence. A Blast search (Altschul et al. 1997) with the ITS sequence of L.
printzenii as template returned L. densa, L. burgaziae and L. varia as the closest results, with a 93%, 92% and
92% of similarity respectively. A NJ tree made with species of the L. varia group (data not shown) returned L.
printzenii as basal regarding the other three taxa. Other lichenicolous species of Lecanora from species of
Umbilicariaceae are L. lasalliae on Lasallia pustulata (Pérez-Ortega & Etayo 2008) and L. gyrophorina on U.
hyperborea (Poelt et al. 1996) but these do not belong to the Lecanora varia group and differ in their habitus,
anatomical characters and chemistry.
Lecanora xanthoplumosella Lumbsch & Elix, sp. nov. (Fig. 16F) Mycobank MB 517791
Sicut Lecanora margarodes sed thallis cremeis, ascosporis minoribus et pannarinam destituto differt.—http://
www.eol.org/pages/Lecanora xanthoplumosella
Type:— AUSTRALIA. Queensland: Razorback Range, 3 km NW of Mount Morgan, 23°28’S, 150°22’E, dry
sclerophyll forest on steep slope with Cycas and Macrozamia, on metamorphic rocks, August 1993, Elix
34643 (holotype CANB).
Thallus thin to thick, verrucose-areolate to verruculose, cream coloured or creamish yellow. Thallus
surface smooth, epruinose, isidia and soredia absent. Prothallus well-developed, blackish brown. Apothecia
sessile, abundant, dispersed, plane or slightly convex to convex, roundish, 0.5–1.2 mm diameter. Apothecial
margin lecanorine, concolourous with the thallus, persistent, thick, entire when young, becoming strongly
verruculose, disc bluish grey to grey with a thick, granulose, yellowish to yellowish grey pruina.
Amphithecial cortex distinct, gelatinous, hyaline, inspersed with small crystals, 10–15 µ m thick.
Amphithecium with large and small crystals. Parathecium hyaline, 10–15 µ m thick, lacking crystals.
Epihymenium orange-brown to reddish brown, 15–20 µm thick, granulose with coarse crystals soluble in
KOH. Hymenium, subhymenium and hypothecium hyaline. Paraphyses sparingly branched and slightly
thickened apically. Asci of Lecanora-type, 8-spored. Ascospores non-septate, hyaline, broadly ellipsoid to
ellipsoid, 9.5–13.5 × 6.0–7.5 µm. Pycnidia immersed in verrucae, bluish grey with filiform conidia. Spot tests:
K+ yellow, C+ orange; KC+ orange, P+ yellow, UV+ dark orange. Secondary chemistry: thiophanic acid
[major], arthothelin [minor], 2’-O-methylperlatolic acid [minor], atranorin [trace], chloroatranorin [trace], 5chloro-2'-O-methylanziaic acid [minor].
Distribution and habitat:—This new species occurs on siliceous rocks in semi-exposed to exposed
habitats in dry sclerophyll forests in Queensland. It is only known from two localities between 300 and 860 m
elevation.
Etymology:—The name refers to the superficially similar neotropical species Lecanora xanthoplumosa
Guderley.
Additional specimens examined (paratypes):—AUSTRALIA. Queensland: Razorback Range, 3 km
NW of Mount Morgan, Streimann 52399 (CANB, F), Elix 34639 (CANB). Watsonville on the HerbetonPetford Road, 10 km W of Herberton, Mayrhofer 11899 (GZU).
Lecanora xanthoplumosella is characterized by a cream-coloured to creamish yellow thallus, the pruinose
apothecial discs with the thalline margins that become strongly verruculose, broadly ellipsoid to ellipsoid
ascospores and the presence of the thiophanic acid chemosyndrome, atranorin, 2'-O-methylperlatolic acid and
5-chloro-2'-O-methylanziaic acid. The latter substance is present as a minor constituent in L. lividocinerea
Bagl. (Elix et al. 1997), a corticolous species occurring in Western Australia that lacks xanthones (Lumbsch
1994, Lumbsch & Elix 2004). The species may be confused with Lecanora margarodes (Körber) Nyl. and L.
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pseudogangaleoides ssp. verdonii Lumbsch. The latter species is distinguished by having black apothecial
discs, an egranulose epihymenium, and the presence of usnic acid in addition to the presence of the arthothelin
chemosyndrome (Lumbsch 1995, Lumbsch & Elix 2004). Lecanora margarodes is readily distinguished in
having epruinose apothecial discs, longer ascospores, and the presence of pannarin in addition to chlorinated
xanthones (Lumbsch 1994, Lumbsch & Elix 2004). The neotropical L. xanthoplumosa Guderley is only
superficially similar and differs in having smaller apothecia with epruinose or slightly pruinose apothcial
discs, longer ascospores, and the presence of the chodation chemosyndrome (Guderley 1999, Guderley et al.
2000).
Lecidea lygommella Elix, sp. nov. (Fig. 17A) Mycobank MB 517792
Sicut Lecidea lygomma sed acida lichenum deficiente differt.—http://www.eol.org/pages/Lecidea lygommella
Type:—AUSTRALIA. Victoria: Alpine National Park, Bogong High Plains, Basalt Hill, 20 km SE of Mt.
Beauty, 36°53’S, 147°18’E, 1650 m, on basalt rocks in exposed alpine grassland with basalt outcrops,
February 1994, Elix & Streimann 40407 (holotype CANB, isotypes Elix, Lichenes Australasici Exsiccati 284,
as Lecidea lygomma).
Thallus saxicolous, verrucose to evanescent, up to 7 cm wide, 0.2–0.7 mm thick. Upper surface whitish to
pale grey or rusty red-brown, areolate, areoles irregular to angular, 0.5–1.0 mm wide, flat, surface roughened
or ±smooth; cortex 20–50 µm thick, upper cell layer unpigmented to grey-brown. Photobiont layer 70–100
µm thick, algal cells 9–16 µm wide. Medulla I−. Hypothallus dark grey, sometimes obvious at the thallus
margin but often not developed. Apothecia scattered or crowded, rounded to angular, subimmersed to adnate,
rarely basally constricted, 0.5–2.3 mm wide; disc black, flat to slightly convex, matt, epruinose or with rusty,
red-brown pruina, margin concolorus with disc, distinct at first, persistent or excluded with age, rarely shiny.
Hypothecium dark brown to black-brown in basal part, K+ violet-brown, 120–250 µ m thick. Hymenium
colourless, 60–85 µ m high, I+ blue. Epihymenium brownish green, 10–15 µ m high, N+ red-violet
(cinereorufa-green); paraphyses occasionally branched and anastomosing, ca. 2 µm wide, apices thickened to
3–4 µm wide. Asci 8-spored, 50–65 × 11–13 µm. Ascospores simple, ellipsoidal, colourless, 10–17 × 5–8 µm.
Pycnidia immersed, ostiole black, punctiform; conidia bacilliform, 9–12 × 1 µm. Secondary chemistry: nil.
Distribution and habitat:—At present this distinctive new species is known from rocks in alpine areas
of New South Wales and Victoria.
Etymology:—The specific epithet refers to the similarity of this species to Lecidea lygomma Nyl.
Additional specimens examined (paratypes):—AUSTRALIA. New South Wales: Bulee Gap, 8 km NE
of Nerriga, just S of Morton National Park, Elix 39721 (CANB). Kosciuszko National Park, Blue Lake, 7 km
NE of Mt Kosciuszko, Streimann 9591 (CANB). Victoria: Alpine National Park, Bogong High Plains, Basalt
Hill, 20 km SE of Mt. Beauty, Elix & Streimann 40443 (CANB), Elix & Streimann 40447 (B, CANB). Alpine
National Park, Bogong High Plains, 20 km SSE of Mt. Beauty, Elix & Streimann 40477 (CANB). 'Ruined
Castle', Bogong High Plains, 16.5 km SSE of Mt. Beauty, Elix & Streimann 40616 (CANB).
This species is characterized by the areolate, whitish to grey or rusty red-brown thallus, the I− medulla,
the black, subimmersed to adnate apothecia, the dark brown to black-brown hypothecium and brownish green
epihymenium; the relatively large ascospores and by the absence of lichen substances. Lecidea lygommella is
morphologically very similar to L. lygomma, but the latter differs chemically in containing norstictic acid
(major) and connorstictic acid (minor) (Rambold 1989, Hertel 1997). In the latter paper, Hertel commented
that “Lecidea lygomma with no TLC-detectable substance … presently must be treated as L. lygomma sensu
lato”.
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FIGURE 17. A. Lecidea lygomella (holotype), thallus with apothecia. B–C. Lecidella greenii (holotype). B. Thallus
with apothecia. C. Ascus with ascospores. D–E. Lempholemma corticola (holotype). D. Central part of thallus with
irregularly shaped areoles composed of densely aggregated thallus granules and numerous apothecia with thin, receding
thalline margin (arrow). E. Section of thallus granule showing Nostoc photobiont (arrow bottom) and enlarged hyphal
cell at thallus surface (arrow top). F. Lepraria sekikaica (holotype), thallus. Scale in A = 2 mm, in B, D, F = 1 mm, in C
= 10 µm, in E = 25 µm.
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Lecidella greenii U.Ruprecht & Türk, sp. nov. (Fig. 17B–C) Mycobank MB 517793
Thallus crustosus, areolatus, albidus. Apothecia lecideina, Asci ad typum Lecidella pertinens, Ascosporae hyalinae, nonseptatae, ellipsoidae, 10–11(–12) × 6–7(–8) µm.—http://www.eol.org/pages/Lecidella greenii
Type:—ANTARCTICA. Ross Dependency: Transantarctic Range, Granite Harbour, 77°00’S, 162°31’E,
January 2008, Türk 43015 (holotype SZU, isotype A).
Thallus crustose, whitish, grey, beige, well-developed, rimose to areolate and sometimes granulose;
chasmolithic and epilithic, up to 1 mm high and 5 cm in diameter. Pycnidia infrequent, up to 0.2 mm;
photobiont Trebouxia sp. Apothecia in dense groups; visible margin when young, inconspicuous when old and
large; black, matt disk, flat to convex, up to 1.2 mm in diameter, sessile, constricted at the base. Epihymenium
up to 10 µm, olive to dark green; hymenium 80–90 µm, hyaline to light brownish when mature, inspersed;
Paraphyses simple, 1–2 µm in diameter, rarely anastomosing, flexuous, slightly adglutinated, end cell dark
green to olive and slightly thickened to 4 × 4 µ m; subhymenium up to 40 µ m, hyaline to light green;
hypothecium hyaline, inspersed with brown crystals; excipulum to 200 µm broad, I–, hyaline, dark green
margin, of radiate hyphae; asci of the Lecidella-type, 40–50 µ m long; ascospores 8/ascus, non-septate,
hyaline, 10–11(–12) × 6–7(–8) µ m, broadly ellipsoid, length-width index 1.5–1.6. Secondary chemistry:
chemotype I: nil, chemotype II: stictic acid chemosyndrome; spot tests: K– or K+ yellow, I–, C–.
Distribution and habitat:—So far endemic to continental Antarctica, on granite rock and sandy soil.
Etymology:—Dedicated to T. G. Allan Green, University of Waikato, New Zealand.
Additional specimens examined (paratypes):—ANTARCTICA: Ross Dependency: Transantarctic
Range, Granite Harbour, Türk 43007, 43013, 43014, 43015, 43017, 43029 (SZU); Taylor Valley, Türk 33611,
33612, 33620, 33628, 33586, 33630, 33649, 33659, 33700, 33706 (SZU); Mt Suess, Türk 42999 (SZU) .
The widespread genus Lecidella comprises c. 50 species worldwide. These crustose lichens occur on tree
bark as well as on mosses, sand and rocks. The genus is characterized by small black lecideoid ascomata and
Lecidella-type asci (Knoph & Leuckert 2004). The new species is one of two species of Lecidella occurring in
continental Antarctica. It is identified by the shape and size of its ascospores, its exclusively saxicolous habit
and its secondary chemistry that clearly sets this lichen apart from all other species of Lecidella so far
described.
Lempholemma corticola M.Schultz & T.Sprib., sp. nov. (Fig. 17D–E) Mycobank MB 517794
Thallus corticola, nigrescente-olivaceus, ecorticatus, madefacte gelatinosus, granulosus vel irregulariter areolatus,
granulis effusis, deinde aggregatis areolae irregulares formantes. Hyphae reticulum formantes. Apothecia thallo
concolore, parva, 0.25–0.37 mm lata, adnata vel stipitata, margine thallino tenui, excipulo proprio pallido, disco
pallide rufescenti, plano. Hymenium iodo caerulescens. Ascosporae octonae, simplices, ellipsoideae, (10.5–)13–
19(–20.5) × (6–)6.5–8(–9) µm. Cyanobacterium symbioticum Nostoc. Pycnidia et pycnoconidia non visa.—http://
www.eol.org/pages/Lempholemma corticola
Type:—GREECE. Crete: Fassas Valley, between Langos and Nea Roumata, 35°24'N, 23°53’E, 350 m,
corticolous on Platanus orientalis, May 2004, Spribille 13286 (holotype GZU, isotype HBG).
Thallus corticolous, blackish olivaceous, ecorticate or thallus granules and apothecia pseudocorticate,
gelatinous when wet, granulose to irregularly areolate, granules ca. 65–165 µ m wide, effuse to later
aggregated forming irregularly shaped areoles, 0.6–2.5 mm wide giving the thallus a very uneven appearance,
fixed to the substrate by pale rhizohyphae. Isidia and soralia lacking. Apothecia small, 0.25–0.37 mm,
subglobose, adnate with slightly constricted base to stipitate, disc at first punctiform, eventually opened, pale
brownish, plane, thalline margin thin, 25–30 µm thick, soon receding. Excipulum proprium thin but distinct,
composed of few interwoven hyphae, pale, basally 5 µm thick, laterally ca. 10 µm thick, apically widened and
seen from above as pale ring. Hymenium clear, 110–120 µ m high, KOH/IKI+ blue, paraphyses distinct,
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sparsely branched, apically widened. Asci broad cylindrical to obclavate, 75–80 × 12–15 µm, wall thin, not
amyloid, tip not thickened. Ascospores 8/ascus, simple, ellipsoid to broad ellipsoid, hyaline, (10.5–)13–19(–
20.5) × (6–)6.5–8(–9) µ m. Pycnidia and pycnoconidia not seen. Ascomata developing from tangle of
generative hyphae. Photobiont Nostoc in bead-like chains. Secondary chemistry: nil.
Distribution and habitat:—So far known only from the bark of old Platanus orientalis trees in the
Fassas Valley in western Crete, in the vicinity of a stream. The riverine Platanus forests are in severe decline
and threatened by encroachment of developments and orange plantations.
Etymology:—Referring to the substrate, which is most unusual in the genus.
Additional specimen examined (paratype):—GREECE. Crete: Langos, “Straßenrand, i.d.N. des
Baches”, May 2004, Wagner s.n. (UPS).
The new species is characterized by its corticolous habit and granulose to irregularly areolate growth
form. The thallus is essentially ecorticate as in all species of Lempholemma but especially at the base of the
thallus granules as well as towards the base of the apothecia a "primitive pseudocortex" (Degelius 1954: 46,
47) may be observed. The hyphae forming this pseudocortex are quite robust whereas those attacking the
photobiont are more delicate. Hitherto, no species of Lempholemma was knownto grow directly on bark. Dry
or moist rock, mosses and soil being preferred substrata. Growth form varies considerably among species of
Lempholemma. Whereas membanaceous to subfoliose-plicate and umbilicate, squamulose-lobate thalli are
common, granulose thalli are rare . L. minutulum (Born.) Zahlbr. and “Psorotichia” lutophila Arnold, both
pioneer species on soil, are examples of the latter. Lempholemma is a mid-sized genus of c. 40 species
occurring worldwide. The genus is in the family Lichinaceae and is distinguished from all other members of
the family by the presence of Nostoc photobionts. The genus Collema also contains Nostoc photobionts but
differs in the presence of septate ascospores, a thickened, amyloid ascus apex, a different type of pycnidia and
conidiophores and the lack of finger-like fungal haustoria invaginating the cells of the photobiont.
Lepraria sekikaica Elix, sp. nov. (Fig. 17F) Mycobank MB 517795
Sicut Lepraria lobificans sed acidum sekikaicum continente differt.—http://www.eol.org/pages/Lepraria sekikaica
Type:—AUSTRALIA. Western Australia: Near summit of Mt. Brown, 3 km SE of York, 31°53’S, 116°47’E,
295 m, on soil below granite overhang in remnant Acacia woodland with scattered Eucalyptus and granite
outcrops, April 2006, Elix 38221 (holotype PERTH, isotypes CANB, HO).
Thallus leprose-sorediate, powdery, creamy white, grey-white, pale yellow-grey or brownish grey,
forming extensive, irregularly spreading patches to 10 cm wide, or in small, irregularly roundish, colonies
0.5–2 cm wide that eventually coalesce; not delimited, well-defined lobes absent but sublobes often present,
0.4–1.0 mm wide, compacted at the margins; thin or thick (up to 250 µm thick), medulla absent; hyphae 1.0–
2.5 µm thick; granules (soredia) farinose, dispersed or forming a thick, continuous layer, ± roundish, 15–40
µm wide, commonly aggregated in roundish to irregular clumps (consoredia) 100–200 µm wide; rarely with
short projecting hyphae (10–20 µm long) but usually not apparent; photobiont chlorococcoid, more or less
spherical, with individual cell 6–13 µm diameter. Hypothallus thin, dispersed, pale brown to white or orangebrown in part, sometimes not apparent. Secondary chemistry: atranorin (minor), sekikaic acid (major),
homosekikaic acid (minor), 4'-O-demethylsekikaic acid (minor), fragilin (minor), ± di-O-methylstrepsilin
(trace); thallus K+ yellow, C–, KC–, P+ pale yellow.
Distribution and habitat:—At present this new species is known from several localities in south-western
Western Australia where it occurs on soil in sheltered rock ledges.
Etymology:—The epithet derives from the unique chemistry of this species.
Additional specimen examined (paratype):—AUSTRALIA. Western Australia: Boyagin Rock,
Boyagin Nature Reserve, 20 km NW of Pingelly, Elix 38939 (CANB, PERTH).
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This is the first reported occurrence of sekikaic acid as a major secondary substance in the genus
Lepraria. Morphologically this new species resembles the very common L. lobificans Nyl., with similar
soredia, consoredia and occasional sublobes at the margins. However, L. lobificans differs in chemistry
(atranorin, zeorin and the stictic acid complex), in having a distinct medulla and consoredia with long
projecting hyphae up to 100 µm long.
Lobariella sipmanii Moncada, Betancourt & Lücking, sp. nov. (Fig. 18A–B) Mycobank MB 519082
Sicut Lobariella botryoides sed apotheciis instructis differt.—http://www.eol.org/pages/Lobariella sipmanii
Type:—COLOMBIA. Cundinamarca: Bogotá D.C., Páramo de Sumapaz, Laguna de Chisacá, 4°18’N,
74°13’W, 3725 m, paramo vegetation with with Diplostephium revolutum S.F.Blake, Espeletia spp. and
Pentacalia spp., August 2010, Lücking et al. s.n. (holotype UDBC, isotype F).
Thallus growing on trunks and stems of small trees, up to 8 cm diameter, very loosely attached; individual
lobes up to 3 cm long, irregular to rounded, 5–15 mm wide, irregularly branched. Photobiont cyanobacterial
(Nostoc). Upper surface pale blue-grey, glabrous, densely white-reticulate through the formation of elongatelinear maculae; lower surface white to yellowish white, white-tomentose; tomentum up to 1.5 mm thick,
formed by branched hairs with globose cells. Lobe margins entire to coarsely crenulate; isidia and soralia
absent. Upper cortex paraplectenchymatous, consisting of 4–6 cell layers, 30–45 µ m thick, hyaline;
cyanobacterial layer 80–130 µm thick; medulla composed of longitudinally oriented hyphae, up to 180 µm
thick; lower cortex paraplectenchymatous, consisting of 2–6 cell layers from which the tomentum emerges,
12–25 µ m thick, hyaline. Apothecia abundant, cup-shaped, up to 8 mm diameter, with thick, strongly
prominent, more or less entire to irregular but not distinctly lobulate, white margins; disc concave, vividly
orange-brown. Ascospores fusiform to acicular, 55–115 × 4–5 µ m, 5–9-septate, hyaline. Pycnidia not
observed. Secondary chemistry: gyrophoric acid (major) and satellite substances; medulla C+ rose to red, K–,
P–.
Distribution and habitat:—The species is known from several collections at the type locality, growing
in partial shade on the lower to mid-trunks and stems of small trees of Diplostephium revolutum S. F. Blake,
associated with Lobariella pallida and mosses of the family Dicranaceae.
Etymology:—We dedicate this species to our colleague, mentor, and friend Harrie Sipman for his
invaluable contributions to lichenology in the Neotropics.
Additional specimens examined (paratypes):—COLOMBIA. Cundinamarca: Bogotá D.C., Páramo de
Sumapaz, Laguna de Chisacá, Mondada et al. 4084, 4098 (UDBC).
Lobariella is a small genus in Lobariaceae, currently comprising five species that are characterized by
forming pseudocyphellae on the upper surface and a more or less continuous, white tomentum on the lower
surface (Yoshimura 1984, 1998, 2002, Yoshimura & Arvidsson 1994). The species of this group, first
designated as L. crenulata group, were subsequently separated in the genus Durietzia Yoshim., which turned
out to be an illegitimate later homonym of Durietzia Gyeln. Yoshimura (2002) therefore introduced the
replacement name Lobariella for this group. Only three species were formally recombined in Lobariella
(Yoshimura 2002) and the following two species are recombined here: Lobariella botryoides (Yoshim. &
Arv.) Moncada & Lücking, comb. nov. [Lobaria botryoides Yoshim. & Arv., Acta Bot. Fenn. 150: 226
(1994); Durietzia botryoides (Yoshim. & Arv.) Yoshim., Recollecting Edvard August Vainio (São Paulo): 91
(1998)] and Lobariella pallida (Hook.f.) Moncada & Lücking, comb. nov. [Sticta pallida Hook.f. in Kunth,
Syn. Plant. Aequinoct. Orb. Novi 1: 28 (1822); Lobaria pallida (Hook.f.) Trevis., Lichenotheca Veneta: no. 75
(1869); Durietzia pallida (Hook.f.) Yoshim., Recollecting Edvard August Vainio (São Paulo): 91 (1998)]. The
new species is only the second one in the genus having a cyanobacterial photobiont after L. botryoides; all
other species have green photobionts. Lobariella sipmanii differs from L. botryoides in the more or less
smooth surface which lacks reticulate ridges; instead, the surface is very strongly reticulate-maculate. In
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addition, isidia and soralia are absent and apothecia are abundantly formed. Because of the different lobe
morphology, L. sipmanii cannot be considered the fertile counterpart of L. botryoides. Except for the
photobiont, the new species resembles L. crenulata (Hook.f.) Yoshim. and L. pallida in the richly apotheciate
thallus with the white, reticulate pattern of the lobe surface, but differs from both latter species in several
details: L. crenulata has a more strongly attached thallus with conspicuous pseudocyphellae that only partly
form a reticulate pattern; in addition, the apothecial margins are strongly lobulate. Lobaria pallida shares with
L. sipmanii the loosely attached thallus and the formation of reticulate maculae, but the maculae are usually
less distinct and also in that species, the apothecial margin is strongly lobulate. Therefore, L. sipmanii cannot
be considered a photomorph of either L. crenulata or L. pallida.
Megalospora austropacifica Lumbsch, Naikatini & Lücking, sp. nov. (Fig. 18C) Mycobank MB 517796
Similis Megalosporae javanicae sed ascosporis majoribus et angustatis differt.—http://www.eol.org/pages/Megalospora
austropacifica
Type:—FIJI. Taveuni: Access road to summit of Devo Peak, montane relict forest at roadside, 16°51’S,
179°58’W, April 2008, Lumbsch 19805h (holotype SUVA, isotype F).
Thallus yellowish grey to whitish grey, glossy, thick, slightly rugulose to continuous, often irregularly
cracked, sometimes with small papillae containing conidiomata. Isidia and soredia absent. Apothecia
common, orbicular, up to 4.5 mm in diameter, up to 1 mm high, disc concave when young, becoming flat to
slightly convex, orange-brown to red-brown, blackish towards the margins, glossy, epruinose. Margin
prominent, thick, inner part (proper exciple) yellowish brown to pale brown, matt, epruinose, outer part
(thalline margin) concolorous with thallus, glossy, epruinose. Exciple with an orange-brown inner part and a
hyaline ectal part, K–. Epihymenium orange-brown, 20–35 µ m thick. Hypothecium hyaline to orange.
Hymenium 200–250 µ m high, hyaline, inspersed, amyloid. Asci 8-spored. Ascospores hyaline, 2-celled,
slightly curved (sulphurata-type, fide Sipman 1983: 44), 60–85 × 22–26 µm, spore walls 2–3 µm thick with
thin, smooth epispore. Secondary chemistry: usnic acid and zeorin.
Distribution and habitat:—So far only known from the islands of Taveuni and Viti Levu in Fiji, where
the species occurs in relict or secondary montane forests on bark.
Etymology:—Named after the occurrence of the new species in the South Pacific.
Additional specimens examined (paratypes):—FIJI. Taveuni, Access road to summit of Devo Peak,
montane relict forest at roadside, Lumbsch 19804t (F, SUVA). Viti Levu: Nadarivatu Nature Reserve,
Secondary montane forest at road to Koro'o, Lumbsch 19850 g (F, SUVA).
This is an additional species in the Megalospora sulphurata complex (Sipman 1983), which is
characterized by asci with more than one, 2-celled, and usually curved ascospores. Megalospora sulphurata
Meyen was recognized in a wide sense by Sipman (1983), but recently Untari (2006) separated additional
species in the complex. The new species is distinguished from M. flavoexcipulata Untari and M. sulphurata in
having a hyaline to orange hypothecium and from M. javanica Untari in having paler margins, regularly
zeorine apothecia, a paler hypothecium, a hyaline ectal exciple, and longer and narrower ascospores.
Megalospora galapagoensis Bungartz, Ziemmeck & Lücking, sp. nov. (Fig. 18D–E) Mycobank MB 517797
Sicut Megalospora porphyritis sed acidum usnicum continente differt.—http://www.eol.org/pages/Megalospora
galapagoensis
Type:—ECUADOR. Galápagos: Isla Sán Cristóbal, trail from Cerro Pelado to El Ripioso, 0°52’S, 89°28’W,
392 m, transition zone, Psidium guajava forest with some old Hippomane mancinella trees and dense
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understory of Rubus niveus, Tournefortia rufosericea and Zanthoxylum fagara, on bark, S-exposed side of
inclined Hippomane mancinella trunk (ca. 20 cm in diameter), semi-shaded, wind- and rain-sheltered, August
2008, Bungartz 8516 (holotype CDS-41162, isotype F).
FIGURE 18. A–B. Lobariella sipmanii (A, holotype; Moncada 4084), thallus with apothecia. C. Megalospora
austropacifica (holotype), thallus with apothecia. D–E. Megalospora galapagoensis (D, holotype; E, Bungartz 3987),
thallus with apothecia and soredia. F. Menegazzia endocrocea (holotype), thallus with apothecia (photograph J. Jarman).
Scale in A, B = 10 mm, in C–E = 1 mm, in F = 5 mm.
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Thallus yellowish grey to whitish grey, glossy, thick, uneven to slightly rugulose, up to 20 cm diameter.
Soredia present, diffuse, starting out as coarse, corticated granules but soon becoming confluent and maculate.
Apothecia round, 0.5–1.5 mm in diameter, up to 0.6 mm high, disc concave when young, becoming flat to
slightly convex when mature, grey-black to black, glossy, epruinose. Margin prominent, thick, black.
Excipulum brownish, K–. Epihymenium reddish brown, 10–20 µm thick. Hypothecium brown, 100–130 µm
high. Hymenium 200–250 µm high, hyaline, strongly inspersed, amyloid. Ascospores single, hyaline, (3–)5septate, 45–75 × 15–25 µm. Secondary chemistry: usnic acid and zeorin.
Distribution and habitat:—So far only known from the Galápagos Islands, found on both Santa Cruz
and Sán Cristóbal Island in the transition and the humid zone.
Etymology:—The epithet refers to the known geographical range of the new species.
Additional specimen examined (paratype):—ECUADOR. Galápagos: Isla Santa Cruz, along the dirt
road from Bellavista to Media Luna, Bungartz 3987 (CDS).
This species belongs in the Megalospora tuberculosa aggregate and could be included in that species in a
wide sense (Sipman 1983). However, Harris (1984, 1986) reported that populations with pannarin instead of
usnic acid, a sorediate thallus, and smaller ascospores represent an autonomous taxon which he recognized as
M. porphyritis (Tuck.) R.C.Harris, a position adopted by other authors (Lücking 2007). The new species is
intermediate between M. tuberculosa s. str. and M. porphyritis and represents a third species in this group.
Thallus chemistry and the non-pruinose apothecia are as in M. tuberculosa s. str., but the (granular) soralia and
the comparatively small ascospores with few septa recall M. porphyritis. The new species is potentially
endemic to the Galápagos Islands, as no such material has yet been found among neotropical mainland
populations of Megalospora.
Menegazzia endocrocea Kantvilas, sp. nov. (Fig. 18F) Mycobank MB 517798
Ad saxa granitica incolens, Menegazziae elongatae M. corrugataeque aliquantum similis et item acidum sticticum
continens sed hoc habitu singulari et pigmento pallide croceo medulloso differens.—http://www.eol.org/pages/
Menegazzia endocrocea
Type:—AUSTRALIA. Tasmania: Mt Cameron, 40°59’S, 147°56’E, 550 m, on granite outcrops in heathland,
July 1995, Kantvilas & Crittenden 42/95 (holotype HO, isotype BM).
Thallus rather loosely adnate, forming irregular rosettes to 100 mm wide, or occurring as scattered,
smaller clumps of lobes, lacking soredia and isidia. Lobes hollow, inflated, cylindrical, (1–)1.5–3(–4) mm
wide, ± dichotomously branched, slightly constricted at the axils; central lobes densely imbricate, generally
lacking secondary lobules; marginal lobes radiating, contiguous, with apices somewhat decumbent. Upper
surface perforate, pale grey to cream-grey, pale brownish grey at the lobe apices, smooth, matt, often patchily
discoloured blackish in older or abraded portions, lightly whitish-pruinose, especially in younger parts of the
thallus. Perforations numerous, scattered, roundish to broadly ellipsoid, 0.5–1.7 mm wide, with margins
usually flush to the the surface or, occasionally, raised and the perforations appearing volcano-like. Lower
surface wrinkled, dull to glossy black, becoming brownish near the lobe tips. Medullary cavity black and
often with a loose cobweb of whitish hyphae in older lobes, faintly pale orange in younger lobes and towards
the lobe apices. Apothecia scattered, 1–4 mm wide, roundish, subpedicillate; thalline margin initially thick
and inrolled, later to 0.1–0.3 mm wide, mostly smooth and entire or with occasional radial cracks and
crenulations; disc reddish brown, undulate, often eroded. Hymenium 90–140 µm thick, colourless in the lower
part, overlain by a reddish brown, non-granular epithecial layer 5–10 µm thick, becoming fuscous brown in
KOH. Asci 2-spored, broadly ellipsoid, 80–110 × 30–55 µm, rather rarely encountered, mostly nestled deeply
within a reticulum of anastomosing paraphyses 2–3 µm thick, with capitate, brown apices 3–8 µ m thick.
Ascospores simple, hyaline although sometimes brownish when over-mature, subglobose to broadly ellipsoid,
(38–)40–51.5–66(–72) × (24–)28–34.8–44(–56) µ m; wall 4–8 µ m thick. Pycnidia abundant, scattered,
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immersed in the upper surface and evident as black specks; ostiole 0.08 mm wide; conidia fusiform, 6–8 × 1–
1.2 µ m. Secondary chemistry: atranorin (trace), chloroatranorin (trace), stictic acid, constictic acid,
cryptostictic acid (trace), peristictic acid (trace), plus the pigment isopigmentosin A and traces of
isopigmentosin B and isopigmentosin C (pigments determined by J. Elix using HPLC); cortex K+ yellow;
medulla K+ yellow, KC–, C–, P+ orange, UV+ orange.
Distribution and habitat:—Menegazzia endocrocea is restricted to the east coast of Tasmania and the
islands of Bass Strait. The species occurs exclusively on Devonian granite, typically in sheltered, fireprotected crevices on rocky, heathy summits overlooking the sea. It is rarely abundant and most colonies are
very fragmented, suggesting a relict distribution and a species in decline. Fire in this heathy, sclerophyllous
vegetation represents a very significant threatening factor to many lichens. The granite of Tasmania's eastern
coastal areas supports a diverse flora rich in highly restricted and/or endemic taxa, not only of lichens but also
of vascular plants (Kantvilas & Elix, 2008). This new species adds to the already high conservation value of
this region.
Etymology:—The specific epithet refers to the orange pigmentation of the medullary cavity of young
lobes.
Additional specimens examined (paratypes):—AUSTRALIA. Tasmania: Mt. Amos, Kantvilas 289/91
(HO). Mt. Freycinet, Kantvilas 151/95 (HO). Mt. Dove, Kantvilas 135/95 (HO). Mt. Mayson, Kantvilas 3328/
97 (HO). Summit of Mt. Freycinet, Kantvilas 359/99 (HO). Flinders Island, Mt. Leventhorpe summit,
Kantvilas 48/07 (HO).
With at least 25 species including four endemics (McCarthy 2009), Menegazzia is well represented in
Tasmania. Most species are epiphytic and occur in high rainfall or highland areas, especially in Nothofagus- or
conifer-dominated cool temperate rainforest, scrub or heathland in western parts of the island. In that regard
alone, this species is rather unusual because, in lower rainfall areas, the only taxa likely to be encountered are
the sorediate M. subpertusa P.James & D.J.Galloway, M. nothofagi (Zahlbr.) P.James & D.J.Galloway or M.
caesiopruinosa P.James, or the brown-lobed M. aeneofusca (Müll.Arg.) R.Sant. Morphologically and
chemically, the new species is very well-characterised. There are numerous esorediate, two-spored species,
just as there are species containing stictic acid, but few combine these features with relatively broad lobes and
the presence of an orange pigment. In the Tasmanian flora, somewhat similar species, both restricted to
corticolous habitats in high rainfall areas, include M. corrugata P.James, which has a wrinkled thallus and a
very thick, inflated, corrugated apothecial margin, and M. elongata P.James, which is chemically identical but
and has markedly constricted, sausage-like lobes. Orange or yellow pigments occur in numerous species of
Menegazzia. Secalonic acid-type compounds give a suffused yellow colour and are K–; they occur in M.
pertransita (Stirt.) R.Sant. Intensely orange-yellow skyrin, co-occurring with pigmentosin, reacts K+ purple,
UV+ orange and is found in M. jamesii Kantvilas & Louwhoff. Then there are K+ purple, UV+ orange
emodin-type pigments that occur in M. caliginosa P.James & D.J.Galloway and M. foraminulosa (Kremp.)
Bitter (Kantvilas & Louwhoff 2004). The pigment combination found in M. endocrocea is also known in M.
elongata (endemic to Tasmania), M. lordhowensis Elix (endemic to Lord Howe Island) and M. grandis
P.James (endemic to New South Wales).
Myriotrema endoflavescens Hale ex Lücking, sp. nov. (Fig. 19A–B) Mycobank MB 517799
Sicut Myriotrema album sed medulla flava et apotheciis ecolumellatis differt.—http://www.eol.org/pages/Myriotrema
endoflavescens
Type:—PANAMA. Veraguas: 4–8 km from Alto Piedra along road to Río Calovebora, 800 m, primary rain
forest, February 1975, Hale 44859 (holotype US).
Thallus light yellow-olive, smooth to uneven, in parts bullate to almost gall-forming, with dense,
prosoplectenchymatous cortex with internal splitting; medulla pale yellow; photobiont layer and/or medulla
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with clusters of calcium oxalate crystals. Apothecia immersed, rounded, 0.2–0.4 mm diameter; disc covered
by narrow, 0.1–0.2 mm wide pore, flesh-coloured, white-pruinose; margin entire, fused, light yellow.
Columella absent. Excipulum prosoplectenchymatous, yellow-brown; periphysoids absent. Hymenium 60–80
µm high; paraphyses unbranched. Ascospores 8/ascus, 3–5-septate, 15–20 × 5–7 µm, ellipsoid, with thick
septa and lens-shaped lumina, colourless, I+ violet-blue (amyloid). Secondary chemistry: no substances (all
spot tests negative); medulla with pale yellow pigment (K+ yellow).
Distribution and habitat:—Known from a single collection in a lower montane rainforest in Panama.
Etymology:—The specific epithet refers to the pale yellow medullary pigmentation.
Mason Hale intended to describe this very characteristic species as new but never published it formally. It
displays a very typical Myriotrema morphology and appears to be most closely related to M. album Fée, M.
myrioporum (Tuck.) Hale, and M. subconforme (Nyl.) Hale, all lacking secondary substances. No species of
Myriotrema having a pigmented medulla is known to date. Most species with pigmented medulla belong in
Ocellularia s.str. (Rivas Plata et al. unpubl. data). Except for the pigmented medulla, M. endoflavescens
resembles M. album, which also differs by its often present irregular pseudocolumella, and M. myrioporum
and M. subconforme, respectively, which differ in their either 1-septate or submuriform ascospores.
Ocellularia albobullata Lücking, Sipman & Grube, sp. nov. (Fig. 19C) Mycobank MB 517800
Ocellularia thallo albo-cinereo, acido psoromico; apothecia non carbonisata, verrucis bullatis immersa,
pseudocolumella instructa; ascosporae hyalinae, transverse 3–5-septatae, ad 20 × 9 µm, I+ violaceae.—http://
www.eol.org/pages/Ocellularia albobullata
Type:—COSTA RICA. Puntarenas: Corcovado National Park, Sirena Section, Sirena Biological Station (Osa
Conservation Area), Osa Pensinsula, 160 km SSE of Sán José and 50 km WSW of Golfito, round trail NW of
station (Sendero Sirena and Sendero Guanacaste), 83°35’W, 8°29’N, sea level, lowland and coastal rain forest
zone, oldgrowth coastal secondary forest and primary forest remnants on sandy soil, on bark (lower trunk),
exposed, April 2003, Grube 11580 (holotype INB-4066756, isotype GZU).
Thallus white-grey, bullate, with dense, prosoplectenchymatous cortex; photobiont layer and/or medulla
with clusters of calcium oxalate crystals. Apothecia prominent, angular-rounded to irregular, 0.7–1.2 mm
diameter; disc covered by narrow, 0.2–0.3 mm wide pore, filled by brown, white-pruinose pseudocolumella;
margin annulate, white, laterally covered by thick thalline layer. Pseudocolumella present, irregular, brown,
white-pruinose. Excipulum prosoplectenchymatous, orange-brown; periphysoids absent. Hymenium 60–80
µm high; paraphyses unbranched. Ascospores 8/ascus, 3–5-septate, 10–20 × 6–9 µm, ellipsoid, with thick
septa and lens-shaped lumina, colourless, I+ violet-blue. Secondary chemistry: psoromic, subpsoromic and 2'O-demethylpsoromic acids (C–, K–, P+ yellow).
Distribution and habitat:—Known from several collections from the coastal rainforest in southern
Costa Rica.
Etymology:—The specific epithet refers to the white-bullate thallus.
Additional specimens examined (paratypes):—COSTA RICA. Puntarenas: Corcovado National Park,
Sirena Section, Sirena Biological Station (Osa Conservation Area), Osa Pensinsula, 160 km SSE of Sán José
and 50 km WSW of Golfito, beach south of station, Sipman 51123 (B, INB).
This new species is easily recognized by its irregularly bullate thallus with white-grey colour. It belongs to
a small number of species of Ocellularia with psoromic acid and where the thallus does not have the
characteristic olive-green colour but is more or less white. Other such species include O. antillensis Hale and
the Australian O. wirthii Mangold, Elix & Lumbsch (Mangold et al. 2008, 2009), which, among other
characters, are separated from O. albobullata by their non-bullate thallus and carbonized excipulum.
Ocellularia calvescens (Fée) Müll.Arg. is similar to O. albobullata in chemistry and ascospores but has a nonbullate thallus, lacks a columella, and has a carbonized excipulum.
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Ocellularia vizcayensis Rivas Plata, Duya & Lücking, sp. nov. (Fig. 19D) Mycobank MB 517801
A Ocellularia mordenii medulla alba differt.—http://www.eol.org/pages/Ocellularia vizcayensis
FIGURE 19. A–B. Myriotrema endoflavescens (holotype). A. Thallus with apothecia. B. Thallus showing yellow
medulla. C. Ocellularia albobullata (holotype), thallus with apothecia. D. Ocellularia vizcayensis (holotype), thallus
with apothecia. E–F. Ochrolechia insularis (F, holotype). E. Habit on coastal granite boulders (photograph G. Kantvilas).
F. Thallus (photograph J. Jarman). Scale in A–D = 1 mm, in F = 2 mm.
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Type:—PHILIPPINES. Luzon Island, Nueva Vizcaya: Mount Palali, 1400 m, March 2007, Rivas Plata 1170
(holotype F, isotype CAHUP).
Thallus white, distinctly areolate, with loose, irregular cortex; photobiont layer and/or medulla with large
clusters of calcium oxalate crystals. Apothecia immersed to erumpent in thalline warts, round, 0.8–1.5 mm
diameter; disc covered by very narrow, 0.05–0.1 mm wide pore; margin with a broad, red-brown to purplish
brown zone around the ostiole. Columella absent. Excipulum strongly carbonized laterally, jet-black, 100–200
µm wide; periphysoids absent. Hymenium 250–400 µm high, clear; paraphyses unbranched. Ascospores 1–2
per ascus, 15–29-septate, 100–300 × 15–30 µm, oblong-fusiform, with thick septa and lens-shaped lumina,
colourless, I+ violet-blue. Secondary chemistry: no substances detected by TLC.
Distribution and habitat:—Known from a single, well-developed collection in pristine lower montane
rainforest in northern Luzon.
Etymology:—The epithet refers to the province of the type locality.
Ocellularia contains a number of species with very large, transversely septate ascospores. Among these,
only two lack secondary substances including pigments. Ocellularia dolichotata (Nyl.) Zahlbr. has a pale
yellowish-brown, minutely verrucose thallus with columnar clusters of calcium oxalate crystals in the
photobiont layer and medulla and a paraplectenchymatous cortex; in addition, the apothecia are columellate.
Ocellularia jamesii (Patw. & C.R.Kulk.) D.D.Awasthi differs in the smaller, pale brown ascospores, among
other features. Most closely related to the new species appears to be O. mordenii Hale, which agrees in the
whitish, more or less areolate thallus; however, that species produces a vivid, cinnabar-red pigment in the
medulla and the apothecial margin is very irregularly and coarsely verrucose and lacks the brown zone around
the ostiole. Ocellularia vizcayensis superficially resembles Thelotrema monosporum Nyl. and allies, which
also has a whitish thallus and brown-rimmed apothecia, but the internal anatomy of the apothecia is very
different, being uncarbonized and featuring periphysoids and brown ascospores in the Thelotrema species.
Ochrolechia insularis Kantvilas & Elix, sp. nov. (Fig. 19E–F) Mycobank MB 517802
Species littoralis, saxicola, thallo albido, crustaceo, usque ad 4.5 mm crasso, papillato vel nodulato vel plicato, acidum
gyrophoricum continenti recognita; apothecia ignota, igitur eam huic generi cautione aliqua adsignamus.—http://
www.eol.org/pages/Ochrolechia insularis
Type:—AUSTRALIA. South Australia: Kangaroo Island, Cape Willoughby, 35°50’S, 138°08’E, 50 m, on
sunny, exposed faces of granite outcrops overlooking the sea, September 2009, Kantvilas 367/09 (holotype
HO, isotypes AD, CANB).
Thallus crustose, intensely papillate, nodular or plicate, 0.3–4.5 mm thick, with upper surface white,
smooth, matt to rather glossy, in places minutely speckled with white maculae, not pruinose or crystalline,
forming irregular, undelimited patches to 10 cm across, not lobate or placodioid at the margins; prothallus
absent; individual nodules 0.5–1 mm wide, very densely crowded together in clumps to 5 mm wide, rather
loosely attached, in section with a poorly developed cortex 15–30 µ m thick composed of irregularly
orientated, branched and anastomosing, short-celled hyphae 2–4 µm thick, overlain by a hyaline, epinecral
layer; photobiont cells generally concentrated in a subcortical layer 40–100 µm thick, with individual cells
globose, 6–15 µm diameter; medullary hyphae loosely interwoven, 3–5 µm thick. Pycnidia immersed, visible
as minute, pale brownish dimples or short cracks; conidia bacilliform, 4–6.5 × 1 µm. Secondary chemistry:
gyrophoric acid [major], lecanoric acid [minor or trace], 2’-O-methyllecanoric acid [± trace]; cortex K–, KC+
red, C+ red, P–, UV–; medulla K–, KC+ red, C+ red, P–, UV–.
Distribution and habitat:—Ochrolechia insularis is a seemingly rare species, known only from the type
locality where it occurs on large, sunny, windswept, granite boulders overlooking the sea but well above the
normal extent of sea spray. It is associated with the typical community that dominates coastal granite rocks in
southern Australia and Tasmania: Rinodina blastidiata Matzer & H.Mayrhofer, Tylothallia pahiensis (Zahlbr.)
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Hertel & Kilias, Xanthoria ligulata (Körber) P.James and species of Buellia, Caloplaca and Xanthoparmelia,
especially X. conranensis (Elix) Elix and X. subprolixa (Nyl. ex Kremp.) Blanco et al. The species is
extremely localised, and has been found on only a few large boulders; searches in similar habitats in the
general vicinity, as well as further afield such as on Flinders Island in Bass Strait, have failed to reveal any
further localities. Thus it represents an extremely uncommon taxon of high conservation value, especially as
much of the site where it grows is heavily degraded by the impact of sheep grazing and other disturbance.
Etymology:—The specific epithet refers to the habitat of the new species on Kangaroo Island off the
southern coast of mainland Australia.
Additional specimens examined (paratypes):—AUSTRALIA. South Australia: Kangaroo Island, Cape
Willoughby, 35°50’S, 138°08’E, 50 m, on sunny, exposed faces of granite outcrops overlooking the sea,
September 2008, Kantvilas 332/08 (AD, HO).
In the absence of apothecia, the generic assignment of this species is tentative; however, its highly
localised distribution warrants its description, and Ochrolechia is considered the most appropriate genus. Its
position there is supported by its general appearance, in combination with its habitat ecology, chlorococcoid
photobiont, secondary chemistry and bacilliform conidia. However, it is conceded that these characters may
all be found in several other genera. From the thallus chemistry (cortex C+ red, medulla C+ red) this species
appears to belong to O. africana Vain. group (Brodo, 1991), but morphologically it closely resembles O.
tartarea (L.) A. Massal. Both O. tartarea and O. insularis are saxicolous and may develop an irregularly
warty to nodulate-papillate upper surface. However, O. insularis differs in having a smooth and glossy upper
surface (granular to powdery-tartareous in O. tartarea) and in lacking a prothallus (O. tartarea is surrounded
by a paler prothallus) (Fletcher et al., 2009). In addition, the colour of the upper surface of O. tartarea varies
from pale to dark grey (white in O. insularis) and the medulla of O. insularis reacts C+ red (C– in O.
tartarea). At first glance, the new species is most reminiscent of members of the Pertusaria dactylina
complex, which, in the southern Australian region, is well-represented on granite in such habitats (Kantvilas
& Elix, 2008). However, these lichens are all isidiate rather then papillate, and contain salazinic, norstictic or
hypothamnolic acids. One fragment of the new species (Kantvilas 332/08A, HO) supports a small cluster of
stalked mazaedia of Sphinctrina leucopoda Nyl., a lichenicolous fungus previously unrecorded for South
Australia. This widespread species is commonly known to parasitise the genera Pertusaria, as well as
Diploschistes and Rinodina (Tibell, 1987). It has not been reported previously from Ochrolechia.
Opegrapha viridipruinosa B.J.Coppins & R.Yahr, sp. nov. (Fig. 20A–B) Mycobank MB 517715
Sicut Opegrapha varia sed thallo tenui, griseoviridi, pulverulento, saepe sulphureo suffuso, ascosporis brevioribus, 4-5
septatis, pycnidiis et conidiis parvioribis differt.— http://www.eol.org/pages/Opegrapha viridipruinosa
Type:—ENGLAND. East Suffolk, Needham Market, 52°9’N, 1°4’E, on Sambucus bark in sheltered, disused
chalk pit, June 2004, Earland-Bennett & Hitch s.n. (holotype E).
Thallus thin, grey-green powdery, effuse, turning lemon-yellow in herbarium or almost absent except as
thin veil at margins of excipulum. Pigments in thallus dissolving K+ yellow in section (best observed adjacent
to or below lirellae). Photobiont trentepohlioid. Lirellae erumpent-sessile, mostly simple, few- or stellatebranched, scattered evenly over the thallus, with thin powdery lateral thalline cover, 0.3–0.6(–1) mm long,
0.1–0.3(–0.4) mm broad. Exciple margins completely carbonized and raised above disc, generally slit-like at
first and mostly gaping to broadly exposed later, brownish-yellow at inside margins, K– in section. N+ red.
Disc mostly exposed, dark brown and often with green (yellow in herbarium) pruina, dissolving in K to give
bright yellow. Hymenium 40–60 µm, I+ red. Ascospores 8/ascus, (3–)4–5 septate, (14)–15–19(–22) × (3–)4–
5(–6) µm, hyaline, excluding perispore (1 µm), with medial cells somewhat enlarged, and infrequent slight
constriction below these. Perispore of old ascospores turning brown and roughly wrinkled. Pycnidia frequent,
pale-green pruinose, sessile 0.07–0.12 mm diameter, with short bacilliform conidia 2.5–3 × 1–1.3 µm.
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Distribution and habitat:—Britain and Ireland, on bark and lignum of hardwood trees (Acer
platanoides, A. pseudoplatanus, Betula, Fraxinus, Quercus, Sambucus and Ulmus).
FIGURE 20. A–B. Opegrapha viridipruinosa (holotype). A. Habit showing ascomata and soredia-covered pycnidia (top
right and lower right). B. Mature ascospores (centre of image) and overmature ascospores (below and to the right). C.
Pannaria phyllidiata (holotype), photographed in the field. D–E. Parmelia asiatica (holotype). D. Habit. E. Detail
showing terminal soralia and marginal linear pseudocyphellae. F. Pertusaria conspersa (holotype), thallus with soralia.
Scale in A, F = 1 mm, in B = 20 µm, in C–E = 2 mm.
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Etymology:—From the yellowish-green powdery pruina, conspicuous on the discs and pycnidia.
Additional specimens examined (paratypes):—ENGLAND. East Suffolk: Sudbourne, January 2002,
Earland-Bennett & Hitch s.n. (E); Pettistree, Byng Hill Lane, March 1997, Earland-Bennett (E).
NORTHERN IRELAND. Londonderry, SSW of Dungiven, Banagher Woods, Coppins et al. 14438 (E).
SCOTLAND. East Lothian: Oak Wood, Coppins 22638 (E); Binning Wood (NW), Coppins 22505 (E);
Peebles: West Linton, Coaly Burn, Coppins 22819 (E).
Opegrapha is a large genus of ca. 300 species, with both lichenized and lichenicolous species (Kirk et al.
2008). The genus is generally characterized by having crustose, non-corticate thalli and carbonized, elongate
to rounded lirellae, transversely-septate and I– ascospores, but the large size of the genus results in few
reliable morphological features common across the genus (Ertz 2009). The new species is most likely to be
confused with O. varia Pers., but is distinguished by smaller and fewer septate ascospores (mostly 5–6-septate
in O. varia), generally smaller lirellae and pycnidia, and smaller conidia. In O. varia the pycnidia are mostly
(0.1–)1.2–2(–2.5) mm diameter, with conidia that are either ellipsoid to pyriform, 3.5–4.5 × 1.5–2 µm, or
bacilliform, 3.8–5.5(–6.5) × 1–1.5 µm. Opegrapha varia generally has a whiter thallus, lacking the yellowish
colouration present in many collections of the new species, even when the apothecia or pycnidia are greenish
pruinose. The same is true for O. herbarum Mont, and is very evident in collections on lignum where it grows
side-by-side with O. viridipruinosa. Opegrapha herbarum differs further in having 3-septate ascospores, and
apparently never producing pycnidia. Opegrapha xerica Torrente & Egea differs in having a K+ green
epithecium and exciple, and longer conidia. The new species is most frequently found in the eastern, drier
parts of Britain and Northern Ireland, and is found in both pycnidiate and sexual forms. It is so far known
from SE England, E Scotland, W Scotland (rare), and Northern Ireland. The earliest collection dates only
from 1991, and it is tempting to suggest that it has only recently entered the British Isles or, at least, expanded
greatly in E Scotland and E England following the amelioration of high SO2 levels since the 1980s. It grows
on a range of hardwood trees, mainly in woodland or woodland edge situations, and associated species
include Anisomeridium biforme, A. polypori, Bacidia delicata, Chaenotheca brachypoda, Chrysothrix
candelaris, Cliostomum griffithii and, on lignum, Opegrapha herbarum and O. ochrocheila. It occurs on dry
bark and does not seem to enter the Xanthorion communities, as can the similar O. herbarum and O. varia. It
is frequent in secondary woodland and is not strongly linked to old-growth woodland or veteran trees, as is O.
xerica.
Pannaria phyllidiata Elvebakk, sp. nov. (Fig. 20C) Mycobank MB 517803
Pannariae sphinctrinae similis, sed praesentia circiter 0.5 mm longorum, ramosorum, ascendentium phyllidiorum et
perisporarum nodulosis apicalibus extensionibus instructarum differt.—http://www.eol.org/pages/Pannaria
phyllidiata
Type:—AUSTRALIA. New South Wales: Brown Mountain, 40 km NNE of Bombala, Rutherford's Creek (S
of Piper's Lookout), 36°35’S, 149°27’E, 90 m, on Southern Sassafras (Atherosperma moschatum), April
2008, Elvebakk 08:160 (holotype NSW, isotypes CANB, UPS, TROM).
Thallus foliose, corticolous, forming rosettes 3–15 cm in diameter, closely attached to the substrate,
unless growing over bryophytes or other uneven substrates. Lobes irregularly to subdichotomously branched,
discrete in peripheral parts, imbricate to centrally coalescent, 0.7–1.2 mm wide and up to 10 mm long,
flattened to weakly concave, margins entire, narrowly swollen, 120–200 µm thick. Upper surface pale greyish
green when fresh and dry, salad green when fresh and moist, turning gradually chestnut brown after storage in
herbaria, glabrous and glossy. Epicortex 30–50 µm thick, upper third developing brown pigmentation after
storage and almost sclerenchymatic near the surface, below paraplectenchymatic with luminae globose to
irregularly ellipsoid, 8–15 µm long, walls 1.5–3 µm thick. Photobiont layer 20–25 µm thick, of globose to
subglobose cf. Myrmecia cells, 5–15 µm in diameter. Medulla 50–100 µm thick, white, dark brown in lower
part. Lower cortex lacking; rhizines common, brown and simple to sparingly branched; hypothallus felted,
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brown, and sometimes forming a blackish prothallus, particularly when growing on smooth bark. Cephalodia
common, laminal on the upper surface, globose to subglobose when young, later becoming irregularly
pulvinate, and finally placodioid-nodulose and up to 2 mm in size, occasionally also developed on the
hypothallus and the lower side; epicortex as in the chlorobiont, but luminae 5–10 µm in size; cyanobiont
Nostoc, cells greyish green, subglobose to irregularly ellipsoid, 3–4.5 × 4–7 µm in size, organized within
indistinct spherical glomeruli, ca. 20 µm in size, and without visible chain structures. Apothecia scattered,
laminal, substipitate, 0.7–2.5 mm broad, discs rufous brown, flat,; thalline excipulum crenate-striate, with
phyllidia; epithecium light brown, 15–25 µm thick, hymenium colourless, but strongly IKI+ blue, ca. 100 µm
thick; hypothecium light brown, ca. 80 µm thick, IKI negative; paraphyses simple to weakly branched,
septate, with slightly swollen apices; asci clavate, 15 × 70–90 µm in size, no internal IKI+ amyloid structures
observed, with eight ascospores. Proper ascospores hyaline, non-septate, ellipsoid to weakly citriform, 7.5–10
× 12–16 µm in size; perispores ellipsoid to weakly citriform, 8–11 × 14–19 µm in size, distinctly verrucose
when mature. Secondary chemistry: vicanicin (major) and unidentified terpenoids (trace).
Distribution and habitat:—Australia, New South Wales, with single occurrences in Victoria and
Queensland. The species is common on tree or tree fern trunks in rainforests from elevations between 200 and
1500 m, occasionally on rocks near streams. Other specified phorophytes include Nothofagus, Eucalyptus,
Olearia, Pomaderis, Trochocarpa laurina, Lomatia ilicifolia, and Bedfordia salicina.
Etymology:—Named after its numerous large phyllidia.
Additional specimens examined (paratypes):—AUSTRALIA. New South Wales: Clyde Mountain, Elix
1305, 1306, 1767, 1770, 1785, 1796, 1809, 1810 (CANB); Brown Mountain, below lookout, Elix 2025, 2028,
9647 (CANB); Brown Mountain, Rutherford's Creek, Streimann 16714 (CANB); ibid., Elix 40822, Elix &
Kalb 40859 (CANB); ibid., Elvebakk 08:166, 08:167, 08:168 (CANB, TROM). 15 km W of Dorrigo along
the Armidale Road, Elix 2331 (CANB). Mount Warning State Park, Elix 4044 (CANB). New England
National Park, Point Lookout, 34 km SW of Dorrigo, McVean 67189 (CANB). New England National Park, E
of Armidale Knights Lookout road, Weber & McVean s.n. (CANB-L49343). New England National Park,
Weeping Rocks Track, Elix 33918 (CANB). New England National Park, track to Wright’s Lookout, 72 km E
of Armidale, Streimann 47880 (CANB). South Coast, Maxwell's Road, Nadgee State Forest, 41 km SSW of
Eden, Elix 21521, 21526, Elix & Streimann 21528 (CANB). Big Badja Hill, 44 km SW of Araluen, Streimann
39881A (CANB). Gibraltar Range National Park, 56 km SE of Tenterfield, Streimann 43568 (CANB). Moppy
Lookout, Barrington Tops State Forest, 40 km WNW of Gloucester, Streimann 44421, 44475, 44480 (CANB).
Dilgry River, Dilgry Circle Road, Barrington Tops State Forest, 40 km WNW of Gloucester, Streimann 44619
(CANB). Pinkwood Creek, Hanging Mountain Forest Reserve, 25 km SW of Moruya, Streimann 44925
(CANB). 3 km SW of Monga, along the Milo Forest road, Elix 6491 (CANB). Queensland: Bunya Mountains,
between Westcott Picnic Area and Long Plain Bald, Kantvilas 167/95 (HO). Victoria: Along the Bonang
River, 7 km S of Bonang township, Elix 5186 (CANB).
This species belongs to the Pannaria sphinctrina (Mont.) Tuck. complex, as circumscribed by Elvebakk
(2007). Among the foliose tripartite species of Pannaria previously accommodated in Psoroma, this group is
characterized by having thin lobes and by vicanicin as its major secondary compound. Pannaria sphinctrina is
a panaustral primarily fertile species. Similar specimens with vegetative propagules are very common both in
southern South America, southeastern Australia and New Zealand, and have generally been interpreted as P.
microphyllizans (Nyl.) P.M.Jørg., in Australian herbaria also filed under P. sphinctrina sensu lato. Pannaria
microphyllizans is probably not a part of this complex, and that material instead represents several
undescribed species. Pannaria phyllidiata is characterized by its numerous ascendent, branched phyllidia.
These are c. 0.5 mm in size and much larger than the propagules of the other still undescribed species, but
shorter than the simple ascendent lobules of Pannaria lobulifera Elvebakk, another new species in the
complex recently described from New Caledonia (Elvebakk 2007). Pannaria phyllidiata has a remarkably
restricted distribution. It is very common in the eastern moist forests of New South Wales, and almost
restricted to this area. So far just a single occurrence has been confirmed about 20–25 km into neighbouring
Victoria, and one locality is known from Queensland.
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Parmelia asiatica A.Crespo & Divakar, sp. nov. (Fig. 20D–E) Mycobank MB 518288
Similis Parmeliae protosulcatae sed differt acidum salazinicum continente et distributione in Asia austro-orientali..—
http://www.eol.org/pages/Parmelia asiatica
Type:—CHINA. Yunnan: Jianchuan County, Shi Bao Shan Park, 75 km S of Lijiang, 26º 22’N, 99º 50’E,
2490 m, on Rhododendron tree trunk, Crespo et al. s.n. (holotype MAF-Lich 16478).
Thallus corticolous, adnate, small, to 3 cm across. Lobes sublinear, separate, 1–2 mm wide. Upper surface
pale greenish to whitish grey, brownish towards lobe apices, smooth, pseudocyphellate, sorediate, without
isidia. Pseudocyphellae marginal, linear, appearing as continuous white rim (to 2 mm long) (Fig. 20 E).
Soralia terminal, developing on lobe apices (Fig. 20 D), usually orbicular; soredia granular. Medulla white.
Lower surface black, densely rhizinate, rhizines densely squarrosely branched, to 1 mm long. Apothecia and
pycnidia not seen. Secondary chemistry (TLC): atranorin and salazinic acid; cortex K+ yellow; medulla K+
yellow turning red, C–, KC–, P+ orange-red.
Distribution and habitat:—The species grows on Rhododendron tree trunk in humid Rhododendron
forest at 2490 m elevation. Parmelia asiatica occurs in temperate regions of China and at present is known
from type locality.
Etymology:—The specific epithet refers to its Asian distribution.
Parmelia asiatica is distinguished by: terminal soralia, marginal, linear pseudocyphellae and atranorin
and salazinic acid. This species recalls Parmelia protosulcata, reported from Australia, New Zealand, South
America and the Falkland Islands (Hale 1987, Elix 1994b), but differs in having salazinic acid, and a
temperate Southeast Asian distribution. In containing salazinic acid, and having a sorediate upper surface and
squarrose rhizines it resembles other related species with wide distributions such as Parmelia sulcata (Hale
1987, Divakar et al. 2005) and Parmelia hygrophiloides known from the Indian subcontinent (Divakar et al.
2003). However Parmelia asiatica is easily distinguished in having terminal soralia and marginal, linear
pseudocyphellae. Molecular studies based on nuITS rDNA show that Parmelia asiatica and Parmelia sulcata
are not closely related (Crespo et al. unpubl. data).
Pertusaria conspersa Messuti, sp. nov. (Fig. 20F) Mycobank MB 517804
Thallus crustaceus, avellaneus, saxicola, areolatus usque ad rugoso-areolatus, tenuis vel crassus, sorediatus. Apothecia
ignota. Acidum haemathamnolicum in magna summa continens.—http://www.eol.org/pages/Pertusaria conspersa
Type:—CHILE. V. Región (Valparaíso): Provincia Sán Felipe de Aconcagua, Cuesta La Dormida, north of
Santiago, October 1960, Mattick 232d (holotype B).
Thallus crustose, light to moderate yellowish brown (hazel), areolate to rugose-areolate, thin to thick,
surface smooth and dull, epruinose or with a white pruina, saxicolous, areolae flattened to slightly bullate,
orbicular to angular or irregular, margins of the areoles thick and slighty effuse, contiguous or separated by
thick, deep cracks, lacking isidia, sorediate. Soralia cream coloured, orbicular, flattened to convex, numerous,
conspicuous, solitary or crowded, sometimes confluent (2–4), sessile, somewhat resembling apothecia,
erumpent, with a smooth or irregular margin, concolorous with the thallus, light to moderate yellowish brown,
0.2–1.5 mm wide; soredia granular, 0.10–0.15 mm diameter. Apothecia not seen. Pycnidia not seen.
Secondary chemistry (determined by HPLC): haemathamnolic acid (major), thamnolic acid (major, minor or
trace).
Distribution and habitat:—The species was found only in the central region of Chile (Chile Central), in
the V Región de Valparaíso (Valparaíso and Sán Felipe de Aconcagua Provinces). It is an uncommon
saxicolous species known only from three localities in Chile. Reported substrates include rocks with quartz
and also sandstone. The species grows at an elevation of 1000 to 1200 m in open and exposed areas.
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Etymology:—The specific epithet conspersa is derived from the Latin conspersus, sprinkled, a reference
to the appearance of the cream coloured soralia on the brownish thallus.
Additional specimens examined (paratypes):—CHILE. V. Región (Valparaíso): Provincia Sán Felipe
de Aconcagua, Cuesta de Chacabuco, N to Santiago, Mattick 266 (B). Cuesta La Dormida, N of Santiago,
Mattick 232b (B).
Pertusaria is a large, cosmopolitan genus of ca. 700 species (Archer 1997) found in subarctic to tropical
regions colonising bark, dead wood, non-calcareous rocks, and rarely soil. The genus is characterized by a
crustose thallus, hemiangiocarpous apothecia, a cupulate true exciple, paraphysoids, thick-walled asci, and
one celled, often thick-walled, hyaline to brownish ascospores, and includes numerous substance classes
(Lumbsch et al. 1999). This species is readily identified by its moderately yellowish brown thallus, the
numerous, erumpent soralia, the absence of apothecia and the presence of haemathamnolic acid. This species
may be confused with other sterile, saxicolous Pertusaria species, but is readily distinguished by its
chemistry. Morphologically the new species resembles the saxicolous P. subventosa Malme var. subventosa
but the latter is distinguished chemically by the presence of lichexanthone, thamnolic acid and picrolichenic
acid as major substances (Archer 1997). The presence of haemathamnolic acid, with varying concentrations of
thamnolic acid, in this new species also distinguishes it from the similar saxicolous P. kinigiensis A.W.Archer
et al. from Central Africa (Archer et al. 2009). Pertusaria kinigiensis has a pale-grey to pale yellowish grey
thallus and white soralia. Other saxicolous species with soralia, are P. coccopoda Vain. (TUR-Vain 06667,
lectotype, selected here, TUR-Vain 06668, duplicate) (= P. xantholeucoides Müll.Arg,) from Brazil, which
also contains haemathamnolic acid but differs in having slightly stipitate soralia and in containing
lichexanthone, and P. sordida A.W.Archer from Australia, but this species contains atranorin and
fumarprotocetraric (Archer 1991).
Phlyctis psoromica Elix & Kantvilas, sp. nov. (Fig. 21A) Mycobank MB 517805
Phlyctidi subuncinatae, P. sordidae P. uncinataeque similis sed acidum psoromicum continens et ascosporis minoribus,
30–52 µm longis, 4–6 µm latis.—http://www.eol.org/pages/Phlyctis psoromica
Type:—AUSTRALIA. New South Wales: Cottan-Bimbang National Park, Stockyard Creek Rest Area, c. 83
km E of Walcha, 31°24’S, 152°07’E, 685 m, on Acacia in wet Eucalyptus forest, August 2008, Elix 43095
(holotype CANB, isotypes HO, NSW).
Thallus crustose, 40–80 µm thick, whitish to very pale blue-grey to greenish grey, rimose, scurfy or
furfuraceous, lacking isidia and soredia but often becoming ulcerous, eroded and rather granular in the
vicinity of the apothecia, forming extensive, irregularly spreading patches 3–10 cm wide; prothallus white,
arachnoid, sometimes forming a narrow, ± byssoid margin; photobiont chlorococcoid, with cells globose to
subglobose, 6–12(–15) µm wide. Apothecia 0.1–0.4 mm wide, usually aggregated in clusters of 2–6(–10),
rarely solitary, immersed to adnate; thalline margin slightly raised, at first entire, soon becoming abraded and
at length reduced to white, granular blotches on the surface of the thallus; disc irregularly roundish, plane to
concave, pale grey to grey-brown, densely covered with a fine, white pruina; proper exciple grey to greyblack, in section cupular, 10–40 µ m, brownish, composed of interwoven, short-celled, branched and
anastomosing hyphae 2–4 µm thick. Hypothecium brownish, ca. 25–40 µm thick, poorly differentiated from
the excipulum. Hymenium colourless, separating readily in K, 75–120 µm thick, overlain by a brownish, K±
olive-brown, granular epihymenial layer that partially dissolves in K; paraphyses simple in the lower part,
sparsely branched towards the apices, septate, 1.5–2.5 µm wide, with apices slightly thickened to 2–4 µm
wide; asci (4–)8–spored, cylindrical-clavate, 70–100 × 10–12 µm, with wall 1.5–2 µm thick, weakly amyloid
but with a thin, more intensely amyloid outer coat, especially near the ascus apex. Ascospores hyaline,
fusiform, transversely (3–)7-septate, coiled in the ascus, straight or sigmoid-curved, 30–39.3–50(–52) × 4–
5.2–6 µ m. Pycnidia not seen. Spot tests: cortex K–, KC–, C– P+ intense yellow. Secondary chemistry:
psoromic acid.
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Distribution and habitat:—This new species is known at present only from several localities in New
South Wales. It occurs on smooth bark in shaded habitats on the trunks of understory trees in wet upland
forests from 680 to 1500 m elevation. Frequent hosts include Doryphora sassafras, Atherosperma moschatum
and species of Elaeocarpus and Pomaderris.
FIGURE 21. A. Phlyctis psoromica (holotype), thallus with apothecia. B. Placopsis imshaugii (holotype), thallus with
apothecia and cephalodia. C. Platismatia wheeleri (holotype), thallus with soralia. D. Porina huinamdungensis
(holotype), thallus with perithecia. E–F. Ramalina hyrcana (Maassoumi 2105). E. Apical branches. F. Detail with
marginal soralia, mainly on the lower side. Scale in A, C–F = 1 mm, in B = 3 mm.
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Additional specimens examined (paratypes):—AUSTRALIA. New South Wales: Mt William,
Kantvilas 328/88 (HO, NSW); New England National Park, c. 3 km W of Point Lookout, Kantvilas 661/88
(HO, NSW); Mount Hyland Nature Reserve, 20 km N of Hernani, Elix 36611, 36652, 36658 (CANB, HO).
Morphologically, Phlyctis psoromica closely resembles several other species of Phlyctis that share a
whitish, often scurfy thallus, ± immersed, often clustered apothecia, 8-spored asci, and fusiform, transversely
septate ascospores. It differs from these taxa mainly by its chemical composition. Phlyctis subuncinata Stirt.,
which is widespread in Australasia, differs in containing stictic, constictic and cryptostictic acids, as well as
having somewhat larger ascospores, 40–72 × 5–7 µm (Galloway 2007). Other related species, also with larger
ascospores, include: P. uncinata Stirt., containing norstictic acid but currently unrecorded for Australia; and P.
sordida C.Knight, which contains hypostictic acid and is here recorded from Australia for the first time (New
South Wales: Gloucester Tops, 32°04’S, 151°34’E, Kantvilas 406/88A, HO). Psoromic acid is known from at
least two other species of the genus. Phlyctis megalospora (P.James) D.J.Galloway & G.Guzmán occurs in
New Zealand, also contains conpsoromic acid and has ascospores 285–390 × 75–95 µm occurring singly in
the ascus. There is also a closely related, similarly large-spored and as yet unidentified species in alpine areas
of Tasmania.
Placopsis imshaugii D.J.Galloway, sp. nov. (Fig. 21B) Mycobank MB 516034
Sicut Placopsis baculigera sed differt in characteribus sequentibus: lobi marginales pallide cinerei sine maculis albidis,
effiguratis; hymenium alterum (250–300 µm) et sporae parum latiores et conidia filiformia (27–45 × 1 µm); acida
gyrophoricum (major) lecanoricumque (minor) et atranorinam (minor) continens.—http://www.eol.org/pages/
Placopsis imshaugii
Type:—CHILE. XII Región: Isla Desolación, treeless ridge in cirque on W side of Fondeadero Nassau,
Puerto Churruca, 53º 04’S, 73º 56’W, October 1969, Imshaug & Ohlsson 44877 (holotype MSC).
Thallus rosette-forming, to subirregular, sometimes forming coalescing patches, ±lobate or effigurate at
margins, rarely lobate from margins to centre, more commonly areolate centrally, closely attached to
substratum from margins to centre, (15–)20–55(–75) mm diameter, prothallus absent. Lobes not swollen,
rather flattened to subconvex, 0.5–2 (rarely to 4) mm wide, parallel, radiating, contiguous from margins to
centre, or only in a broad marginal zone and areolate-cracked centrally, separated by narrow to deep and
sometimes gaping cracks; margins entire, flabellate, shallowly rounded or scalloped to occasionally minutely
lobulate. Upper surface pale grey brown to pinkish brown at margins, orange-brown to rust-coloured
centrally, smooth, subnitid and somewhat coriaceous at or near margins, uneven to minutely papillate to
±cerebriform (×10 lens) in central areolae, without isidia, maculae, pruina, pseudocyphellae or soredia.
Cephalodia widely scattered centrally, to occasionally developing in groups or clustered in bands towards
thallus margins, rounded, 1–5(–9) mm diameter, purplish blue when wet, rusty brown to brown-black when
dry, shallowly hemispherical to somewhat flattened, margins plicate-crenulate, surface very shallowly ridged
at first becoming deeply ridged and/or with gaping, radiating fissures at maturity; cyanobiont in chains, cells
rounded to laterally compressed, 5–8.5 µ m diameter. Apothecia subimmersed to ±aspicilioid at first,
becoming sessile and appearing subthelotremoid at maturity, 0.8–1.5(–2.5) mm diameter, scattered to
crowded centrally, solitary to 2–4-together, rounded to slightly constricted through mutual pressure. Thalline
margin prominent, obscuring disc at first, swollen, 0.3–0.6 mm thick, concolorous with thallus or paler, entire
to striate-cracked, roughened or wrinkled, without pruina or soredia. Proper margin very thin to scarcely
apparent, concolorous with disc or slightly darker; disc deeply urceolate, sometimes reduced to a slit or a pore
by the swollen thalline margin, especially when young, yellow-brown to dark-brown, waxy, glossy, epruinose.
Epithecium pale pinkish brown, granular, 25–30 µm thick. Hymenium colourless, (200–)250–300 µm tall.
Hypothecium pale yellow-brown or pinkish, opaque, 150–200 µm thick. Asci cylindrical, 200–250 × 8–10
µ m. Ascospores uniseriate in ascus, broadly ellipsoidal, colourless, 25–30 × 15–18(–20) µ m. Pycnidia
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common, scattered on thallus from near margins to central areolae, immersed, ostiole minute, punctiform,
pale whitish to red-brown to black. Conidia filiform, 27–32.5(–35) × 1 µ m. Chemistry (J. A. Elix, pers.
comm. 2010): medulla K–, C+ red, KC+ red, P–; containing gyrophoric acid (major), lecanoric acid (trace)
and atranorin (trace).
Distribution and habitat:—Placopsis imshaugii appears to be a rare Chilean endemic and is presently
known only from several remote coastal sites in Region XII, in the canals region of western Patagonia. It
colonises hard siliceous rocks on scattered outcrops on ridges and open hillsides and associates occasionally
with Aspicilia sp., Placopsis gelida (L.) Linds., and with rust-coloured species of Porpidia. The rather few
collections seen give only a sketchy idea of its ecology, and since it is a distinctive rust-coloured species, it is
easily seen and should be more carefully looked for, so that its ecological requirements can be more clearly
ascertained.
Etymology:—The specific epithet honours the major contribution to temperate South American
lichenology made by its collector, the late Prof. Henry A. Imshaug, 1925–2010 (Fryday & Prather 2001).
Additional specimens examined (paratypes):—Chile. XII Región: Isla Wellington, Puerto Charrúa,
Imshaug & Ohlsson 43586, 44635 (MSC, CANB). Isla Grant, Puerto del Morro, Canal Trinidad, Imshaug &
Ohlsson 44707, 44709 (MSC, CANB). Isla Pilot, Puerto del Morro, Canal Trinidad, Imshaug & Ohlsson
44713 (MSC, CANB). Isla Desolación, moorland on ridge and summit of hill at head of Brazo Lobo, Imshaug
& Ohlsson 44842 (MSC).
Placopsis imshaugii is characterized by: the closely attached, rosette-forming thallus without a marginal
prothallus, with a grey-brown to pinkish brown marginal zone and an orange-brown or rust-brown upper
surface centrally, without isidia, maculae, pruina, pseudocyphellae or soredia; apothecia that are aspicilioid at
first then becoming subthelotremoid, with a prominent thalline margin often obscuring the disc; a colourless
hymenium 200–300 µm tall; cylindrical asci, 200–250 × 8–10 µm; broadly ellipsoidal ascospores, 25–30 ×
15–18(–20) µ m; filiform conidia, 27–32.5(–35) × 1 µ m; and gyrophoric acid as principal secondary
metabolite (together with traces of lecanoric acid and atranorin). A rust-coloured thallus is characteristic of
several Southern Hemisphere species of Placopsis with the ability to accumulate iron from the substratum, a
situation found also, though more widespread, in the genus Porpidia (Schwab 1986, Gowan 1989a, 1989b,
Gowan & Ahti 1993, Fryday 2005), with taxa of the two genera often forming sympatric communities.
Placopsis imshaugii is distinct from the following rust-coloured species: Placopsis baculigera I.M.Lamb is a
Chilean endemic having distinctly swollen, convex lobes that are pale to dark cinnamon-buff to orangebrown, minutely papillate and with white, effigurate maculae; sessile apothecia with a thick, entire to striatecracked or wrinkled thalline margin, a waxy, glossy, yellow-brown to dark-brown disc; and characteristic rodshaped conidia 5–7.5(–10) × 0.5–1 µm. Placopsis bicolor (Tuck.) B.de Lesd. is an austral species known from
southern Chile, Îles Kerguelen, Îles Crozet, Heard Island, Prince Edward Islands, North and South Islands of
New Zealand and from South Georgia (Lamb 1947, Dodge 1948, Galloway 2001, 2002, 2007, Galloway et al.
2005, Øvstedal & Gremmen 2006, 2007). It is separated from P. imshaugii by differences in thallus
morphology: P. bicolor has rather more convex lobes that are conspicuously rusty, orange-yellow to redbrown and lack a pale, grey-brown or pinkish brown marginal zone; the hymenium is shorter (160–240 µm);
ascospores are smaller [(18–)20–24 × 10–15 µ m]; and the chemistry is different, with minor amounts of
gyrophoric acid and the major metabolites being either porphyrilic acid or cryptostictic acid, with differing
trace amounts of pannaric acid, pannaric acid 2-methyl ester, stictic acid and alternariol methyl ether (J.A.
Elix pers. comm. 2010). Placopsis elixii D.J.Galloway (Galloway 2001) is a New Zealand endemic that is
often markedly rusty, orange-brown, especially when growing in exposed habitats. It is distinguished from P.
imshaugii by the white, laminal pseudocyphellae, the distinctive surface maculation, the patches of glistening,
white pruina on the lobe apices and on the surface of the cephalodia, and by differences in apothecial anatomy
and ascospore size, those of P. elixii being (30–)31.5–36.5(–40) × (8.5–)10–12(–13.5) µ m. Placopsis
lateritioides I.M.Lamb is an austral species (Galloway 2007) that differs from P. imshaugii in having
characteristic, grey-blue, scattered, laminal soralia, and it is very rarely fertile. It also has a differing chemistry
of gyrophoric (major), stictic (minor) and norstictic (trace) acids.
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Platismatia wheeleri Goward, Altermann & Björk, sp. nov. (Fig. 21C) Mycobank MB 517806
Platismatiae glaucae similis sed thallii linearibus et soraliis marginalibus latis densisque differt. Isidia absunt.
Apothecia et conidiomata ignotae. Corticola.—http://www.eol.org./pages/Platismatia wheeleri
Type:—U.S.A. Montana: Missoula County, Mount Sentinel, 1 km E of the University of Montana, 45°51’N,
113°57’W, 1050 m, on branches of Pseudotsuga menziesii, March 2008, Wheeler 2985 (holotype UBC).
Thallus foliose, up to 5–10 cm across, corticate above and below; lobes ascending, loosely attached from
the base, proportionately rather broad, rounded to irregular in outline, mostly 6–20 mm wide, thin. Upper
surface whitish, smooth to weakly pitted, without pseudocyphellae, matt or at any rate not distinctly shiny, I–.
Soredia present, fine, powdery, uniform in size, borne in continuous long marginal soralia, also sometimes
present over upper or lower surface in more or less circular soralia; marginal soralia strongly sinuous even
when young. Lower surface varying from brown to more often black, usually with a reticulum of narrow
raised ridges, dull to occasionally shiny toward the tips, lacking rhizines or with sparse unbranched rhizines.
Upper cortex 16–26 µm thick, prosoplectenchymatous; medulla white, 100–170(–200) µm thick, lower cortex
100–150 µ m thick, prosoplectenchymatous, I+ purplish; photobiont cells Trebouxioid. Apothecia and
pycnidia unknown. Secondary chemistry: atranorin (cortex K+ yellow) and caperatic acid (by TLC).
Distribution and habitat:—Currently known only from western intermontane North America, north into
southern British Columbia, and south to Washington, Idaho and Oregon. All specimens seen by us were
collected from the branches of Pinus ponderosa and Pseudotsuga menziesii.
Etymology:—Platismatia wheeleri is named in honour of lichen photographer and taxonomist Timothy
B. Wheeler, who first brought this species to our attention, and who contributed both the holotype and the
accompanying photograph.
Additional specimens examined (paratypes):—CANADA. British Columbia: Fraser River Drainage,
Siska Valley, 10 km S of Lytton, Goward 09-88 (UBC). Idaho: Kootenay County, Q'emlin Park, 1 km W of
downtown Post Falls, Björk 14186 (UBC). Bonner County, University of Idaho Field Campus, trail to
Antelope Lake, 1.5 km SE of town of Clark Fork, Björk 14255 (UBC). Washington: Spokane Co., Riverside
State Park, 6.5 km NW of downtown Spokane, Björk 14132 (UBC). Shortly east of Highway 195, 43 km S of
Spokane, Björk 19109 (UBC). Near intersection of Cahill and Kentuck Trails Roads, 31 km SSW of Spokane,
Björk 19187 (UBC). Riverside State Park, slopes above Deep Creek, 13 km NW of downtown Spokane, Björk
19198 (UBC). Peone Prairie, near Moffat Road, 17 km NE of downtown Spokane, Björk 20081 (UBC).
Platismatia is a well-known genus of 11 foliose lichens found primarily at temperate latitudes, mostly on
trees (Culberson & Culberson 1968). It is characterized by the presence of large, whitish, more or less basally
attached lobes and a white to black lower surface usually without rhizines. Platismatia wheeleri was formerly
included in P. glauca (L.) W.L.Culb. & C.F.Culb., a highly variable lichen recognized by its broad lobes
usually with at least some marginal isidia. In some specimens the isidia grade into soralia, leading to
confusion with P. wheeleri. In P. glauca, however, the soredia vary in size and at least when young bear tiny
dark apical initials (“eye spots”), giving the supporting soralia a minutely “speckled” appearance at ×40. The
soredia of P. wheeleri are more uniform and lack “eye spots”; when exposed to light, they darken uniformly
across the upper surface. Additionally, soredia of P. wheeleri are borne in more or less continuous marginal
soralia that are strongly sinuous from the first, while in P. glauca soralia tend to be more discontinuous and
become strongly sinuous, if at all, only later during their development. Both species also bear laminal soralia,
though only P. glauca produces, in addition, "isidioid fruticose branches" as described by Culberson &
Culberson (1968) and illustrated in Brodo et al. (2001: fig. 693). The holotype specimen of P. wheeleri was
sequenced in connection with a separate study on P. glauca s. lat. (Goward & Altermann unpubl. data). The
resulting sequence (ITS only) differed by eleven or more base pairs (10 sites) from 15 sequenced specimens of
P. glauca available on GenBank. Platismatia wheeleri is also superficially similar to Parmotrema
praesorediosum (Nyl.) Hale which differs, however, in the presence of smaller, more appressed lobes up to 10
mm across, a smooth lower surface, and production of medullary praesorediosic acid.
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Porina huainamdungensis Papong, Thammathaworn & Lücking, sp. nov. (Fig. 21D) Mycobank MB
517807
Sicut Porina cupreola sed peritheciis pallide aurantiacis differt.—http://www.eol.org/pages/Porina huainamdungensis
Type:—THAILAND. Chiang Mai Province: Huai Nam Dung National Park, 19°14’N, 98°41’E, 1474 m,
lower montane rainforest, February 2006, Papong 2697 (holotype KKU).
Thallus epiphyllous, crustose, glabrous, dispersed into rounded, confluent patches, shiny, greenish grey,
without visible prothallus, 15–20 µm thick. Photobiont Phycopeltis, with rectangular cells in radiating rows.
Perithecia applanately lens-shaped, basal part spreading, pale yellow-orange, 0.2–0.3 mm diameter.
Involucrellum light yellow, K+ orange; excipulum colourless, K+ yellowish; no algal layer between
excipulum and involucrellum. Paraphyses unbranched. Asci fusiform, 65–75 × 8–10 µm, colourless, tholus I–
. Ascospores 8 per ascus, colourless, fusiform, 7–septate, 24–28 × 3–4 µm.
Distribution and habitat:—Known from several collections in the lower montane rainforest of Thailand;
typical of the forest understory.
Etymology:—Referring to the type locality in Huai Nam Dung National Park.
Additional specimen examined (paratypes):—THAILAND. Chiang Mai Province: Huai Nam Dung
National Park, Papong 2707 (KKU), 2714 (KKU), 2719 (KKU). Loei Province: Phu Kradueng National Park,
Papong 2061 (KKU).
Porina huinamdungensis is similar to P. cupreola (Müll.Arg.) F.Schill. in its small, lens-shaped perithecia
and 7-septate ascospores, but the latter has dark red-brown perithecia and slightly longer ascospores. The
Valdivian P. fulvelloides Lücking agrees with the new species in general appearance but has 3-septate
ascospores. Porina subapplanata Malcolm, Vĕzda, P.M.McCarthy & Kantvilas is very similar but has larger
perithecia and ascospores and appears to be a cool-temperate rainforest species of southeastern Australia,
Tasmania, and New Zealand.
Ramalina hyrcana Sipman, sp. nov. (Fig. 21E–F) Mycobank MB 517808
A Ramalina farinacea simili differt soraliis lateralibus deorsum versis, acidum sekikaicum continens.—http://
www.eol.org/pages/Ramalina hyrcana
Type:—IRAN. Golestan: Gonbad-e-Kavus district, along road from Khan Bebin to Shirabad waterfall, 120
m, deciduous forest with Parrotia persica and Carpinus, on Pterocarya trunk, October 2007, Sipman et al.
55201 (holotype IRAN, isotypes B, hb. Sohrabi 9432).
Thallus forming bushy cushions 1–4 cm thick, densely and often dichotomously branched from a simple
base into strap-shaped, 0.2–1 mm wide lobes; lobes flat with rounded margin, surface sometimes slightly
striate; pseudocyphellae absent; often with tiny, 0.5–1 mm long and ca. 0.1 mm wide, fragile branchlets with
hooked tip, mainly towards the lobe tips, sometimes also on the soralia; soralia present, mainly marginal,
round to elongate with somewhat raised thalline rim, to ca. 0.5 × 0.2 mm in size, the larger ones turned
downward so that the soredia are mainly produced on the underside of ± horizontal lobes. Apothecia and
pycnidia unknown. Secondary chemistry: usnic acid (trace, cortical), sekikaic acid with traces of related
substances (medullary).
Distribution and habitat:—So far Ramalina hyrcana is known only from the Hyrcanian forest area
along the Caspian coast in Iran, between 10 m (sea level) and ca. 400 m, where it grows on tree trunks and can
stand considerable shade.
Etymology:—The epithet refers to the forest type where this species is found.
Additional specimens examined (paratypes):—IRAN. Gilan: Prope Enseli ad mare Hyrcanum, April
1902, Bornmüller (Iter Persicum alterum) s.n. (B). Mazandaran: Zirab, between Khalil Kola and Zebh Valley,
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Valadbeigi 19 (B). Tonekabon, She-hezar towards Lireh-sar, Maassoumi 2105 (B, TARI). Zirab, Amlak
Valley, Valadbeigi 11, 17 (B). Northern Khorasan: Darkesh, Nargesli, Hadjmoniry 1905 (B, FUMH).
Ramalina hyrcana is sympatric with R. farinacea and R. pollinaria and strongly reminds one of these
species by its deeply divided thallus with strap-like lobes and marginal soralia. It is distinguished from both
by the presence of sekikaic acid and down-turned soralia. The soredia-producing surface of the soralia is
directed downward, at least in the larger soralia, giving the branches a dorsiventral aspect uncommon in
Ramalina. In R. farinacea, soralia are lateral and remain so, not becoming turned downward; in Iran strains
with protocetraric, salazinic and norstictic acid are known. R. pollinaria, also known from Iran, contains
evernic acid and has less regularly divided thalli with often wider lobes, to 3 mm; its soralia, which may
sometimes be turned downward, become often granular. The new species agrees with R. nervulosa
(Müll.Arg.) Des Abb. and R. peruviana Ach. in its chemistry, thallus shape and presence of soralia (Krog &
Swinscow 1976, Swinscow & Krog 1988; own TLC observations). It differs from both in its down-turned
soralia. R. nervulosa is mainly confined to tropical regions, although it is reported from Yemen (Sipman
2002). R. peruviana has mostly cylindrical lobes (Stevens 1987, Swinscow & Krog 1988).
Ramalina stoffersii Sipman, sp. nov. (Fig. 22A) Mycobank MB 517809
A Ramalina dendroide persimili differt soraliis farinosis, acido salacinico absente et acido divaricatico presente.—http:/
/www.eol.org/pages/Ramalina stoffersii
Type:—NETHERLANDS ANTILLES. Saba: Lower Hell's Gate, trail head of trail to Sulphur Mine,
17°39’N, 63°14’W, 300 m, steep slope with scrub and rock outcrops, on andesite rock, March 2007, Buck
54767 (holotype NY, isotype B).
Thallus pale yellowish green, 3–10 cm high, forming ca. 3–20 cm wide, rather intricate, subpendulous
cushions; prothallus spreading and giving rise to swarms of thallus lobes, but often decayed and the thallus
attached by secondary attachment spots on the thallus lobes; thallus lobes linear, rather regularly
dichotomously branched up to over 10 times over their whole length, applanate, ca. 1–2 mm wide and ca. 0.3
mm thick, with flat to slightly convex sides, rarely in part canaliculate or subcylindrical; pseudocyphellae
present, linear, mainly on the lobe margins and there to over 1 cm long and causing the lobes to become
rectangular in section, also frequent on the sides but there shorter and rarely over 2 mm long; lobe upper parts
and tips sometimes with minute lobules, often densely branched, about 0.1 mm wide, to over 1 mm long, with
hooked tips; soralia present, marginal, farinose, rounded to elongate, ca. 0.2–0.5 mm in diameter, sometimes
warty, with ca. 50 µ m wide soredia. Apothecia and pycnidia unknown. Secondary chemistry: usnic and
divaricatic acids.
Distribution and habitat:—Known from the Lesser Antillean islands of Saba and St. Maarten/St. Martin
and from Puerto Rico (Landron 1972). The species occurs in maritime environment below ca. 500 m, mainly
on andesite boulders and stone walls among cultivated fields, once one a trunk of a Mammea americana tree.
Perhaps it is more commonly epiphytic in more natural vegetation.
Etymology:—The new species is named after the late A. L. Stoffers (Utrecht, The Netherlands), who
raised my interest in Antillean lichens.
Additional specimens examined (paratypes):—NETHERLANDS ANTILLES. Saba: Windwardside,
17°38’N, 63°14’W, 400 m, Sipman 15176 (B). Along path Windwardside-Rendez Vous, Sipman 15185 (B),
15238 (B). Hell’s Gate, on steep N-facing slope above the sulphur mine, Sipman 15384 (B). North Coast trail
from parking area in Lower Hell's Gate to Crab Rock, mostly within Saba National Park, Buck 50882 (NY),
50884 (NY). Village of Windwardside, Buck 50942 (NY). St. Maarten: Dutch Cul de Sac, Emilio Wilson
estate, Sentry Hill, Buck 50963 (NY).
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FIGURE 22. A. Ramalina stoffersii (holotype), lobe fragment with soralia. B. Relicina colombiana (holotype), thallus.
C–D. Rhizocarpon diploschistidina (McCune 29900). C. Habit. D. Hymenium in water. E–F. Sagenidiopsis isidiata
(holotype), thallus with pseudoisidia. Scale in A–C, E, F = 1 mm, in D = 10 µm.
Ramalina is among the largest genera of macrolichens but nevertheless new species are still being
discovered regularly. The Antilles, where R. stoffersii was discovered, are promising for diversity in the genus
Ramalina, because the genus as a whole shows considerable insular endemism, e.g., on the Canary Islands
(Krog & Osthagen 1980), the Galápagos Islands (Aptroot & Bungartz 2007), and St. Helena (Aptroot 2008).
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Moreover, the Neotropics in general, in spite of some regional treatments (Kashiwadani 1987, Kashiwadani &
Kalb 1993) appear to be very incompletely explored, as suggested by, e.g., the published results from
unfinished studies in Venezuela by Morales Méndez and collaborators (Marcano & Morales Méndez 1994).
The representation of the genus on the Antillean Island of Puerto Rico has received considerable attention
(Landron 1972, Harris 1989), and most of the specimens collected by the author on the islands of Saba and St.
Eustatius fit the indicated species well. Only one group of specimens lacked description because it was
misunderstood, and it is described here. Ramalina stoffersii is morphologically very similar to R. dendroides
Nyl., which grows in the same localities. In fact, Landron (1972) was already aware of its existence but
treated it as a chemically deviant population. The specimens collected on the Antillean islands Saba and St.
Eustatius made clear that it involves two distinct species with a different mode of vegetative reproduction. R.
stoffersii is a truly sorediate species producing soredia in rounded, marginal, sometimes slightly tuberculate
soralia. R. dendroides produces larger, corticate granules on top of short, ca. 1–2 mm long, cylindrical, lateral
branchlets. These morphological differences may be obscured in poorly developed specimens and much
abraded herbarium material. The combination of linear thallus lobes with marginal soralia and divaricatic acid
seems to occur otherwise only in R. nervulosa (Müll.Arg.) Des Abb. (Stevens 1987), but that species lacks the
prominent marginal peudocyphellae and has larger, more elongate and excavate soralia. The primary species
related to both R. dendroides and R. stoffersii is probably R. peranceps Nyl., or if one prefers to consider this
a chemical strain, R. anceps Nyl. The presence of divaricatic acid was tested in all listed specimens and
confirmed by cochromatography with Evernia divaricata (L.) Ach. using standard TLC procedure (Orange et
al. 2001; solvents A, B' and C).
Relicina colombiana Elix & Sipman, sp. nov. (Fig. 22B) Mycobank MB 517810
Sicut Relicina subabstrusa sed diminuta, lobis angustioribus et lobulatis et acidum sticticum et acidum consticticum
continente differt.—http://www.eol.org/pages/Relicina colombiana
Type:—COLOMBIA. Bogotá D.C.: Páramo de Chisacá, along the road Usmé-Nazareth, highest point of road
at Laguna Grande, 3700 m, on rock outcrops on hill slope with Espeletia vegetation, September 1984, Aguirre
& Sipman 5032 (holotype B, isotypes CANB, COL).
Thallus small foliose, adnate to tightly adnate, 1–3.5 cm wide. Lobes separate or rarely sparingly
imbricate, dichotomously branched, linear-elongate, 0.2–1.0 mm wide, lobulate; lobules marginal, sublinear,
simple or rarely dichotomously branched, 0.05–1.0 mm wide; apices pointed to subtruncate; bulbate cilia
conspicuous, forming a dense marginal fringe, simple or often furcate, black. Upper surface pale yellowgreen, ±flat, shiny at the apices, smooth to rugulose, emaculate, without isidia. Medulla white. Lower surface
black; rhizines sparse, black, simple then becoming densely entangled. Apothecia not seen. Pycnidia
common, black, immersed; ostiole punctiform, to 0.1 mm wide. Conidia bacilliform, ca. 3 × 1 µm. Secondary
chemistry: usnic acid (major), norstictic acid (major), stictic acid (submajor), constictic acid (minor), salazinic
acid (trace), cryptostictic acid (trace), peristictic acid (trace), connorstictic acid (trace).
Distribution and habitat:—Restricted to saxicolous substrata in the paramos of the Eastern Cordillera of
Colombia (3700–4315 m elevation).
Etymology:—The epithet refers to the geographic distribution of this species.
Additional specimen examined (paratype):—COLOMBIA. Boyaca: Sierra Nevada del Cocuy,
Boqueron de Cusirí, Cleef 8816b (B, COL).
In overall morphology and chemistry this new species resembles Relicina subabstrusa (Gyeln.) Hale, but
it is distinguished in having a smaller thallus (1–3.5 cm wide versus 4–8 cm) with narrower lobes (0.2–1.0
mm wide versus 0.5–2.0 mm) which bear dense marginal lobules (R. subabstrusa is elobulate), and in
containing medullary stictic and constictic acids in substantial amounts. By contrast, R. subabstrusa contains
only norstictic acid and connorstictic acids in the medulla (Elix 1995). Relicina subabstrusa is common and
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widely distributed in north-eastern Australia, Southeast Asia, South America (Brazil, Guyana and Paraguay)
and the Indian Ocean (Comoro Is.) where it occurs on the trunks and branches of trees in a broad range of
habitats, from mangroves at sea level to forests at mid elevation (850 m). By contrast, R. colombiana is
restricted to quartzitic rock in the páramo zone of the Eastern Cordillera of Colombia (3700–4315 m
elevation). Yellow and white species of Hypotrachyna, such as H. flavida (Zahlbr.) Hale and H. physcioides
(Nyl.) Hale, are very common on the mossy quartzite rocks in the páramo belt of the Colombian Andes, and
R. colombiana was at first confused with this genus. It differs in having stout, bulbate cilia which form a
marginal fringe. Ciliate margins may occur on lobules of some Hypotrachyna species, but the the cilia are
never bulbate. Further, the new species differs chemically from known species of Hypotrachyna (Sipman et
al. 2009).
Rhizocarpon diploschistidina McCune, sp. nov. (Fig. 22C–D) Mycobank MB 516796
Species lichenicola, hospes Diploschistes muscorum; sicut Rhizocarpon malenconianum sed thallo crasso flavo et
acidum rhizocarpicum continente differt.—http://www.eol.org/pages/Rhizocarpon diploschistidina
Type:—U.S.A. Oregon: Gilliam County, Fourmile Allotment of Bureau of Land Management, SW of
Fourmile Road, 45°37’N, 120°03’W, 258 m, with Pseudoroegneria spicata, Poa secunda, and Festuca
octoflora, on Diploschistes muscorum on silty soil, summit of low ridge, May 2009, McCune et al. 29990
(holotype OSC, isotypes UPS, US).
Thallus areolate, very pale yellowish tan to an intense fluorescent yellow, forming small yellow mounds
of areoles on Diploschistes; areoles first appearing as a yellowing surface of the Diploschistes, later
protruding as a convex mound with a diffuse margin, ultimately forming discrete but contiguous areoles;
apothecia black, lacking a thalline margin; to about 1 mm diameter, developing centrally or marginally on the
areoles, circular to irregular in outline, becoming compound or confluent; spores dark brown, 16–18 × 7–8 µm
, mostly 4-celled (occasionally more or less), the septation and outline often slightly asymmetrical or
irregular; epihymenium dark brown; hymenium with POL+ granules; hypothecium dark brown. Secondary
chemistry: rhizocarpic acid (lecanoric and diploschistesic acids also detected by TLC in two of three
specimens, but these are presumably from the host); cortex P–, K–, C–, KC–, UV+ orange; medulla P-, K-, C, KC- or KC+ pink (from underlying Diploschistes?), IKI– or partly IKI+ bluish (from underlying
Diploschistes?), UV–.
Distribution and habitat:—So far known only from shrub steppe and grasslands in central Washington
and north-central Oregon, overgrowing Diploschistes muscorum on soil and grass stubble in little-grazed
areas with well-developed biotic crusts.
Etymology:—The epithet refers to the apparently obligate host, Diploschistes.
Additional specimens examined (paratypes):—U.S.A. Oregon: Deschutes Co., from Highway 97 in
Terrebonne, Root 1820 (OSC). Gilliam Co., above Eightmile Canyon, low ridge, McCune 29999 (OSC).
Wheeler Co., 6 km E of the John Day River, off Highway 218, near John Day Fossil Beds State Park,
Rosentreter 6518 (herb. Rosentreter). Washington: Yakima Co., Bureau of Land Management, Ponzetti 1582
(OSC). Benton Co., Horse Heaven Hills, Ponzetti 955a, 1149a, 1189, 1245, 1285a, 1291b, 1308b, McCune
24291 (all OSC).
Spores and apothecia of R. diploschistidina are very similar to those of R. malenconianum (Llimona &
Werner) Hafellner & Mayrh. In fact, R. diploschistidina was first reported from North America as R.
malenconianum (McCune & Ponzetti 2005, McCune & Rosentreter 2007), but that species apparently should
be removed from the North American checklist. The author thanks Josef Hafellner for pointing out the
consistent absence of a distinct thallus in R. malenconianum. Furthermore, R. malenconianum occurs on
Diploschistes diacapsis on gypsiferous soils in Spain and North Africa (Llimona & Werner 1975, CasaresPorcel et al. 1994), while R. diploschistidina occurs on D. muscorum on noncalcareous silt loams and loams
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in North America. It is similar in appearance to Epilichen scabrosus (Ach.) Clem. ex Hafellner, but that
species occurs on Baeomyces rather than Diploschistes. These differ in that spores of Epilichen are 2-celled,
while R. diploschistidina is usually 4-celled and thickly halonate.
Sagenidiopsis isidiata G.Thor, Elix, Lücking & Sipman, sp. nov. (Fig. 22E–F) Mycobank MB 517811
Differt a Sagenidiopse undulata pseudoisidis numerosis, simplicibus, usque ad 0.8 × 0.5 mm.—http://www.eol.org/pages/
Sagenidiopsis isidiata
Type:—GUATEMALA. Baja Verapaz: Biotopo del Quetzal, road CA-14, 3 km SE of Purulha, Sendero Corto
(Los Helechos) trail, 15°13'N, 90°13'W, 1600–1800 m, montane rain forest with Podocarpus oleifolius,
Hieronia guatemalensis, Oreomunnia guatemalensis, Ocotea spp., May 2008, Lücking & Rivas Plata 25511
(holotype F, isotypes BIGU, UPS).
Thallus up to 10 cm diameter, usually several thalli of different size growing intermixed, rather loosely
appressed to the substrate, byssoid, not flaking off, cream-coloured to greyish; in section up to 0.4 mm thick,
hyphae 3–5 µm wide, without calcium oxalate crystals. Hypothallus below the entire thallus, byssoid, dark
brown to blackish brown, composed of 3–4 µ m wide hyphae. Prothallus prominent, up to 5 mm wide,
byssoid, composed of interwoven and radiating hyphae, dark chocolate brown. Pseudoisidia numerous,
cylindrical, unbranched or sparsely branched, fluffy-felty with projecting hyphae, of same colour as thallus or
slightly paler, up to 0.8 × 0.5 mm. Photobiont cells single or in short, irregular threads, 23–31 × 17–25 µm.
Ascomata and pycnidia not seen. Secondary chemistry: C+ red, K–, PD–, UV+ white, I–, K/I–; diploschistesic
acid [major] and lecanoric acid [minor] (P. & B. v.d. Boom & V. Freire 33413, HPLC by J. Elix).
Distribution and habitat:—Sagenidiopsis isidiata is known from a few but widely distributed localities
in Central and South America. The species is corticolous in tropical montane rainforests.
Etymology:—The new species is named after the pseudoisidia frequently occurring on the thallus
surface. Note that Sagenidiopsis undulata incorrectly is referred to as undulatum, e.g. in Egea et al. (1995),
Cáceres (2007) and Index Fungorum (http://www.indexfungorum.org/Names/Names.asp).
Additional specimens examined (paratypes):—GUATEMALA. Baja Verapaz: SSE of Coban, SE of
Purulhá, Biotopo Mario Dary Rivera (Biotopo del Quetzal), Fern trail, including the area of the biological
station, 1700 m, August 2004, P. & B. v.d. Boom & V. Freire 33413 (herb. P. v.d. Boom, UPS). HAITI. Dept.
De l’Ouest: ridge north of Foret des Pins (Shada station), near border of Dominican Republic, 1770 m, July
1958, Imshaug & Wetmore 2900 (H). VENEZUELA. Mérida: Finca San Eusebio, La Carbonera, cercanías de
Mérida, 2200 m, March 1978, Lopez-Figueiras & M. Keogh 15530 (S). ECUADOR. Zamora-Chinchipe:
Nature Reserve of Estacion Cientifico San Francisco, S of road Loja-Zamora, ca. 40 km from Loja, 2025 m,
June 2004, Sipman 52623 (B-600148395, LOJA). Cordillera Numbala, Reserva Biologica San Francisco, S of
road Loja-Zamora, June 2004, Sipman 52600 (B-600148375, LOJA); ibid., July 2004, Sipman 53148 (B600148848, LOJA).
This puzzling species has been known to the first author for at least 20 years. As the generic position has
been unclear, it has until now remained undescribed. It is superficially similar to other byssoid genera in the
Arthoniales like Crypthonia, Dichosporidium, and Herpothallon sensu lato. Crypthonia and Herpothallon
both have thalli reacting K/I+ patchily blue in 0.2% and 1% aqueous iodine solution (Frisch & Thor 2010,
Frisch et al. 2010) and Dichosporidium has thalli that at least sometimes are sparsely squamulose, a chemistry
with depsidones of the β-orcinol series and when pseudoisidia are present, these are more compact (Thor
1990). Otherwise, Dichosporidium nigrocinctum (Ehrenb.Fr.) G.Thor resembles the new species in thallus
morphology and coloration. The genus Sagenidiopsis in South America until now only includes S. undulata
(Fée) Egea, Tehler, Torrente & Sipman (Egea et al. 1995, Cáceres 2007), a species previously included in
Byssophoropsis (Tehler 1993). Sagenidiopsis undulata has apothecia, lacks pseudoisidia and has a similar
secondary chemistry including lecanoric acid and an unidentified pinkish spot (Tehler 1993). The presence of
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diploschistesic acid in S. isidiata is surprising as this substance previously has only been found in
Diploschistes species.
Sticta venosa Lücking, Moncada & Robayo, sp. nov. (Fig. 23A–C) Mycobank MB 517812
Sicut Sticta filicinella sed thallo foveolato isidiate et tomento venoso differt.—http://www.eol.org/pages/Sticta venosa
Type:—ECUADOR. Pichincha: Río Guajalito Protected Forest, 0°09’S, 78°39’W, 1800 m, montane
rainforest, on ground between mosses, September 2008, Lücking 26252 (holotype QCNE, isotype F).
Thallus growing on the ground between mosses and hepatics, up to 15 cm diameter, individual lobes up to
10 cm long, linear to slightly flabellate, 3–5 mm wide, much branched especially towards the tip; upper and
lower surface strongly and reticulately ridged, appearing faveolate. Photobiont cyanobacterial (Nostoc).
Upper surface blue-grey with brownish tinge when fresh, pale brownish grey when dry, glabrous; lower
surface white to yellowish white, glabrous except for thin lines of dark brown tomentum growing on top of the
ridges and forming distinct, thin, radiating veins; tomentum up to 2 mm thick, formed by strongly branched
and partly conglutinated hairs with cylindrical cells; cyphellae scattered, minute, up to 0.15 mm diameter,
with pore-like opening (thelotremoid), white, K–. Lobe margins much incised and crenulate, with tufts of up
to 0.5 mm long, dark brown hairs growing from the incisions especially at the lobe tips; lobe margins
otherwise and towards the thallus center forming numerous marginal isidia, isidia terete to flattenend,
unbranched to branched and partly resembling phyllidia, 0.05 mm wide and up to 0.5 mm long, of the same
colour as the upper lobe surface. Upper cortex paraplectenchymatous, consisting of 1–2 cell layers, 15–25 µm
thick, hyaline; cyanobacterial layer 25–50 µm thick; medulla composed of longitudinally oriented, more or
less compacted hyphae, up to 100 µm thick; lower cortex paraplectenchymatous, consisting of a single cell
layer from which the tomentum emerges, 15–25 µm thick, brown, cells with rounded outer surface. Apothecia
not observed. Secondary chemistry: no substances detected by TLC; thallus and medulla C–, K–, P–.
Distribution and habitat:—The species is known from a rich collection from the type locality, growing
in the shaded understory of a montane rainforest.
Etymology:—The epithet refers to the distinct veins formed by the ridges and the tomentum following
the ridges on the lower side.
This new species is similar and closely related to Sticta filicinella Nyl., which agrees in the substrate and
general lobe morphology including the cyanobacterial photobiont. We examined an isotype of the collection
cited in the protologue of Sticta filicinella and the specimen also exhibits the thin white veins on the lower
surface partially covered with lines of brown tomentum; that material lacks isidia but bears apothecia instead.
Sticta venosa can therefore be considered the isidiate counterpart of S. filicinella. The distinctly ridged and
foveolate lobe surface forming veins on both sides and the tomentum on the lower side following these veins
give the appearance of a Peltigera or a Lobaria of the L. peltigera group. The new species somewhat
resembles the enigmatic submersed lichen formerly known as Hydrothyria venosa J.L.Russell (Peltigera
hydrothyria Miadl. & Lutzoni), but can immediately be separated by its terrestrial ecology and the presence of
cyphellae on the lobe underside.
Tapellaria albomarginata Lücking, sp. nov. (Fig. 23D–E) Mycobank MB 517813
Sicut Tapellaria leonorae sed
albomarginata
ascosporis minoribus 3-septatis differt.—http://www.eol.org/pages/Tapellaria
Type:—COSTA RICA. Puntarenas: Isla del Caño Biological Reserve, 8°42’N, 83°52’W, sea level, lowland
rainforest, foliicolous, July 1992, Lücking 92-3797 (F).
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FIGURE 23. A–C. Sticta venosa (holotype). A. Thallus in situ. B. Thallus underside. C. Marginal isidia. D–E.
Tapellaria albomarginata (holotype). D. Thallus with apothecia. E. Thallus with campylidia. F. Thelotrema fijiense
(holotype), thallus with apothecia. Scale in A, B = 5 mm, in C = 0.5 mm, in D–F = 1 mm.
Thallus dispersed into rounded, partly confluent patches, 10–25 mm across and 10–15 µm thick,
ecorticate, smooth, pale grey to white. Apothecia rounded to strongly irregular in outline, 0.3–0.5 mm
diameter and 100–150 µm high; disc plane to slightly concave, black; margin thin, pale grey pruinose (in
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necrotic specimens pure black). Excipulum 20–30 µm broad, pale brown. Hypothecium 20–50 µm high, dark
brown, K+ purplish. Apothecial base dark purplish brown. Epithecium distinct, 5–10 µ m high, dark
aeruginous brown. Hymenium 50–60 µm high, colourless. Asci 40–50 × 10–15 µm. Ascospores (6–)8 per
ascus, ellipsoid, 3(–4)-septate, 14–20 × 3–6 µm, 3–5 times as long as broad, colourless. Campylidia sessile,
0.3–0.5 mm broad; lobe well-developed, hood-shaped, pale grey to white pruinose; socle not apparent.
Conidia filiform with slightly clavate apex, usually enrolled, 3–7-septate, 20–40 × 2–2.5 µm, associated with
scattered algal cells. Chemistry: no substances detected by TLC.
Distribution and habitat:—Neotropics, apparently confined to coastal lowland areas.
Etymology:—The epithet refers to the white-pruinose apothecial margin.
This species was considered conspecific with the paleotropical Tapellaria bilimbioides R.Sant. (Lücking
2008). However, after further revision of the available material, it appears that the neotropical populations
consistently have a slightly prominent and irregular, white-pruinose apothecial margin, whereas the few
available paleotropical collections have apothecia with a non-pruinose, black, flush margin. Tapellaria
albomarginata thus differs from T. bilimbioides in the same way as T. nana (Fée) R.Sant. differs from T.
epiphylla (Müll.Arg.) R.Sant. or T. leonorae M.Cáceres & Lücking from T. nigrata (Müll.Arg.) R.Sant.
(Lücking 2008). The new species is externally most similar to T. leonorae, which has much larger, 7-septate
ascospores.
Thelotrema fijiense Lumbsch, Lücking & Naikatini, sp. nov. (Fig. 23F) Mycobank MB 517814
Sicut Thelotrema subtile sed ascomatis brunneis et ascosporis minutis differt. http://www.eol.org/pages/Thelotrema
fijiense
Type:— FIJI. Taveuni Island, access road to summit of Devo Peak, 16°51’S, 179°58’E, April 2008, Lumbsch
19813c (holotype SUVA, isotype F).
Thallus epi- to hypophloedal, thin, whitish grey, pale olive to brown around ascomata. Thallus dull to
shiny, smooth, continuous. Protocortex discontinuous, up to 20 µm thick. Algal layer continuous, crystals
sparse, small. Vegetative propagules not seen. Ascomata 0.2–0.4 mm in diameter, roundish, apothecioid,
solitary to rarely marginally fused, semi-emergent to emergent, flattened-hemispherical. Disc often becoming
partly visible, greyish to pale flesh-coloured, slightly pruinose. Pores small, up to 0.25 µ m in diameter,
roundish to slightly irregular, entire to slightly split, proper exciple apically visible from surface, whitish to
off-white, incurved. Thalline rim margin thin, becoming wide to rarely gaping with age, roundish to irregularroundish, entire, olive-brown to brown. Proper exciple becoming free, at least partly, hyaline to pale yellowish
internally, yellowish brown marginally, amyloid at the base. Hymenium up to 140 µ m high, clear,
conglutinated, paraphyses moderately interwoven, unbranched to slightly branched, tips slightly thickened,
lateral paraphyses present, inconspicuous, up to 30 µ m long, columellar structures absent. Epihymenium
hyaline to brown, granulose. Asci 8-spored, tholus thick narrowing at maturity. Ascospores transversely
septate, cell walls thick, smooth, non-halonate, hyaline when young, becoming brown at maturity, faintly
amyloid, fusiform, ends roundish to acute, loci angular to roundish, 19–27 × 5–7 µm with (7–)9(–11) septa.
Pycnidia not seen. Secondary chemistry (HPTLC): no substances (all spot tests negative).
Distribution and habitat:—This new species is only known from the type locality in Taveuni Island
(Fiji), where it grows on twigs in a cloud forest at 800 m elevation.
Etymology:—The name refers to the archipelago of Fiji, where the type was collected.
Thelotrema is a large genus of crustose, mostly tropical-montane lichens in Graphidaceae, characterized
by apothecia with a double margin and periphysoids projecting from the excipulum (Frisch 2006, Rivas Plata
et al. 2010). Thelotrema fijiense is characterized by its whitish thallus with brown ascomata, and small,
transversely septate ascospores that turn brown when mature. The species lacks secondary metabolites and is
similar to T. subtile and T. suecicum (Lumbsch et al. 2008, Mangold et al. 2009). The latter differs in having
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larger ascospores that remain hyaline and less emergent ascomata that are concolorous with the thallus.
Thelotrema subtile agrees with the new species in having ascospores that turn brown, but is readily
distinguished by having ascomata that are concolorous with the thallus and larger, halonate ascospores with
more or less crenate cell walls. A further species of the T. subtile-group that may be confused with T. fijiense
is T. defossum (Lumbsch et al. 2008). This species, however, differs in having halonate, hyaline ascospores
that never turn brown, and in having inconspicuous, immersed ascomata.
Tricharia nigriuncinata Yeshitela, Eb.Fischer, Killmann & Sérus., sp. nov. (Fig. 24A–B) Mycobank MB
517815
Ab omnibus speciebus generis Trichariae differt nigris setis in medio uncinatis.—http://www.eol.org/pages/Tricharia
nigriuncinata
Type:—UGANDA. Budongo forest, 01°43’N, 31°32’E, ca. 1000 m, on living leaves on Argomuellera
macrophylla, October 2005, Yeshitela 16 (holotype LG).
Thallus epiphyllous, continuous, smooth, lacking crystals, thin, greyish green, up to 5 mm wide, furnished
with abundant sterile setae. Photobiont chlorococcoid with cells green and globose, 5–8 µm diameter. Setae
setiform, black, simple or usually furnished with 1–2(–6) horizontal or usually downwards bending hooks
near their middle, slightly bending, up to 1.5 mm long and 0.04 mm thick at their base, black. Hyphophores
not found. Apothecia usually absent, sometimes abundant, sessile, round, 0.1–0.2 mm diameter, ca. 75 μm
high; disc light orange to almost translucid, opaque, plane without pruina; margin thin, non or slightly
prominent, smooth, pale brown, usually darker than the disc. Excipulum gelatinized, prosoplectenchymatous,
20–30 µm thick, light brown. Hypothecium ca. 10 µm thick, hyaline. Epithecium 5–8 µm high, pale brown.
Hymenium hyaline, 40–50 µm thick. Paraphyses branched and anastomosing, 1 µ m thick. Asci broadly
clavate, 1(–2)-spored, 50–65 × 35–50 µm. Ascospores muriform with numerous cells, ellipsoid, 40–55 × 15–
25 µm, hyaline.
Distribution and habitat:—Tricharia nigriuncinata is typically a foliicolous species in the understory of
dense rainforests; it is confined to undisturbed forests along the Albertine Rift, a major biodiversity hotspot in
East Africa (Plumptre et al. 2007). It seems to be most common on the Western side of the Rift in Congo
RDC.
Etymology:—The epithet refers to the characteristic hooks on the setae.
Additional specimens examined (paratypes): CONGO RDC. South Kivu: Irangi, rainforest by river
Luhoho, Lambinon 78/263, 269, 272, 273 (all LG). Kahuzi-Biega National Park, km 58 of Bukavu-Walikale
road, Lambinon 78/320 (LG). RWANDA. Nyungwe Forest National Park, near Kamiranzovu swamp,
Lambinon s.n. (LG). Gisakura waterfall, December 2009, Fischer s.n. (KOBL). UGANDA. Budongo forest,
Yeshitela 65, 66, 71 & 75 (KOBL), Sérusiaux s.n. (LG).
Within the very diverse lichen family Gomphillaceae, the genus Tricharia is a well-delimited and
coherent taxon (Lücking et al. 2005), with black setae as a distinct character. Indeed, black setae are otherwise
known only in Gyalectidium nashii, Paratricharia paradoxa and in rare forms of Arthotheliopsis floridensis
(Lücking et al. 2007, Lücking 2008). Tricharia nigriuncinata is easily recognized by its long and rather stiff
setae usually furnished with 1–2(–6) hooks near their middle. Tricharia demoulinii Sérus. (known from Papua
New Guinea and Thailand), T. elegans Sérus. (known from Papua New Guinea), and T. substipitata Vězda
(known from Congo RDC) are the only other three species in the genus with hooked setae (Sérusiaux 1984,
V ě zda 1979, Papong et al. 2007). However, the hooks of these species are at the tip of the setae; in T.
substipitata, the hooked setae may represent immature hyphophores as the thallus is otherwise furnished with
very long, black hairs. The setae of T. nigriuncinata are similar to the white setae of Aderkomyces armatus
(Vězda) Lücking, Sérus. & Vězda (Vězda 1975). Tricharia nigriuncinata is very similar with the widespread
and pantropical T. vainioi R.Sant. and differs only in the hooks on the setae. We first considered it might be an
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aberrant form of the latter, but we finally came to the conclusion that it represents a different species as both
are sympatric in a restricted area in the Tropics (e. g. pristine forests in the Albertine Rift in Africa) and as T.
nigriuncinata has never been observed in the large collections examined either from the Neotropics (Lücking
2008) or from Papua New Guinea (E. Sérusiaux unpubl. data).
Usnea galapagona Truong & P.Clerc, sp. nov. (Fig. 24C–F) Mycobank MB 518904
Thallus erectus; basi leviter rubescens; ramuli irregulares; soralia punctiformia, interdum agregata vel conjuncta;
cortex nitidus et valde crassus (proportione 16–18%); medulla valde tenuis (3–4.5%) et K– (substantiam
incognitam et UV+ viridis continente).—http://www.eol.org/pages/Usnea galapagona
Type:—ECUADOR. Galápagos Islands: Isla Sán Cristóbal, Cerro Mundo, at the top of the rock cliffs on the S
side close to the summit, 00°53’S, 89°34’W, 282 m, transition zone with Bursera graveolens, Croton scouleri
and Jasminocereus thouarsii, on Jasminocereus thouarsii on the ridge, August 2008, Clerc & Truong 08-405
(holotype CDS, isotypes G, CMA: 16/3/61.5; chemistry: usnic acid, unknown medullary metabolite reacting
UV+ green after charring).
Thallus fruticose, up to 8 cm long, erect and strongly shrubby, often with several attachment points. Base
concolorous with thallus, usually with a reddish tinge. Branching distinctly anisotomic-dichotomous.
Branches irregular to tapering, not constricted at their attachment point. Section of branches terete to slightly
flattened or ridged. Terminal branches with few divisions and thickened at the tips. Fibrils few or absent.
Papillae hemispherical, often numerous. Soralia minute, often aggregating in small packs or fusing and thus
looking like a large single soralium, developing on top of papillae or directly on the cortex ad initio,
remaining plane with a distinct cortical margin, rarely becoming slightly stipitate or efflorescent, but never
excavating. Isidiomorphs present, very small. Apothecia not seen. Cortex thick (15.5–19.5% of branch width,
n = 10), very hard and vitreous in longitudinal section of branch. Medulla compact, usually very thin and
almost indistinct (3–4.5%), rarely thicker (up to 10%) close to the base. Axis very thick (54.5–62%). Medulla
K–, C–, P–. Secondary metabolites detected by thin layer chromatography (n = 9): usnic acid and an unknown
compound reacting UV+ green after charring (RF classes: solvent A = 6; B = 3; C = 5).
Distribution and habitat:—Endemic to the Galápagos Islands, so far known from Isabela, Sán Cristóbal
and Santa Cruz Islands. This species usually grows on exposed habitats, for instance along ridges or on the
slopes of the volcan’s crater. It was found in the transition vegetation zone, seldom in the arid and humid
zones. Substrate: on rocks, on cactus and more rarely on bark (for exemple trunks of Bursera graveolens).
Etymology:—The epithet refers to the location where this species was discovered.
Additional specimens examined (paratypes):— ECUADOR. Galápagos Islands: Isabela, Volcán
Alcedo, Aptroot 64769A (CDS). Isabela, Volcán Darwin, SW slope, Bungartz 7862B (CDS). Sán Cristóbal,
sector of the “gotera de agua”, Clerc & Truong 08-330 (CDS, G). Santa Cruz, N side of the island, Aptroot
64568 (CDS
Usnea is a large genus of c. 350 mainly corticolous species Swinscow & Krog 1979, Clerc 1987, 2006,
2008, Halonen et al. 1998, Herrera-Campos et al. 1998, Ohmura 2001, Stevens 2004). The genus is
characterized by the fruticose thallus, branches holding a central axis and the presence of usnic acid in the
cortex (Clerc 1998). This new species, so far endemic to the Galápagos, is characterized by anisotomicdichotomous branching, terminal branches with few divisions, the absence of fibrils and the minute soralia
developing from low papillae or on the cortex of branches, often aggregating in packs or fusing. Anatomy in
longitudinal section of branch is very characteristic, with a thick, very hard and vitreous cortex, a thick axis
and a very thin, almost indistinct medulla. TLC analysis revealed the presence of a new unknown substance in
the medulla, reacting UV+ green after charring. The cortex of U. subscabrosa Motyka (Herrera-Campos et al.
1998) is similar to U. galapagona, but the former species is pendulous, with a very different morphology,
chemistry and ecology.
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FIGURE 24. A–B. Tricharia nigriuncinata (A, holotype; B, Yeshitela 71), thalli with apothecia and setae. C–F. Usnea
galapagona (holotype). C. Thallus showing irregular branches with few ramifications. D. Concolorous basis with a
reddish tinge. E. Detail showing minute soralia, aggregating in small packs and developing on top of papillae. F. Detail
showing thick and vitreous cortex, compact and almost indistinct medulla, and very large axis. Scale in A–D = 1 mm, in
E, F = 500 µm.
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Usnea pallidocarpa Wirtz & Lumbsch, sp. nov. (Fig. 25A–B) Mycobank MB 517816
A Usnea perpusilla ascomatis croceo-brunneis vel brunneis differt. http://www.eol.org/pages/Usnea pallidocarpa
Type:—ARGENTINA. Río Negro: Sán Carlos de Bariloche, Cerro Catedral, 41°16’S, 71°20’W, 1900 m,
December 2003, Wirtz & Messuti PA-25b (holotype F).
Thallus approx. 5–7 cm, arising from a proliferating, unpigmented or brownish holdfast with main
branches tapered towards the holdfast; erect, shrubby, subdichotomous to richly branched with terete
branches; thallus surface yellow-green, conspicuously smooth and glossy, sparsely faveolated; main branches
unpigmented, side branches ± variegated with bands of black pigment. “Compressed” morphotypes with
thick, clawed, dark pigmented side branches occurring (as in U. perpusilla). Cortex annulations occurring. No
papillae. Fibrils very rare. Medulla dense. Axis thick, occupying (34)–53–(71)% of branch diameter Soredia
and isidiomorphs unknown. Apothecia frequent, subterminal, rarely terminal, occasionally in series; cupular,
becoming mostly flat or undulate with maturity. Disc light yellow to brownish sometimes with black blotches
or shaded; excipulum smooth, margin thin, often excluded, with few rays. Usneoid ascospores 8/ascus,
simple, ellipsoid, hyaline, with no taxonomic value. Photobiont trebouxioid. Secondary chemistry (TLC):
inconsistent hypostrepsilic acid chemosyndrome (Elix et al. 2007).
Distribution and habitat:—Known from just a single locality in the Andean Cordillera in southern
South America (Argentina). It is an alpine species found at high elevations, about 1800 m, on rocks, in
communities with U. perpusilla, U. sphacelata and U. lambii.
Etymology:—The epithet refers to the pale apothecial discs.
Additional specimens examined (paratypes):—ARGENTINA. Río Negro: Sán Carlos de Bariloche,
Cerro Catedral, 41°16’S, 71°20’W, 1850 m, December 2003, Wirtz & Messuti PA-24 (F).
Usnea pallidocarpa belongs to a small group of polar-alpine species formerly summarized in the
subgenus Neuropogon (Walker 1985, Wirtz et al. 2008). This group is characterized by its fruticose growth
form, a yellow-green thallus with varying black pigmentation, a black or rarely brown apothecial disc and
saxicolous habitat (Walker 1985). The new species (referred to as U. aff. perpusilla in Wirtz et al. 2006 and as
Usnea sp. 1 in Wirtz et al. 2008) is closely related to U. perpusilla, but is distinguished by its smooth almost
glossy thallus surface, a thick central cord, a dense medulla and a yellow apothecial disc with a smooth
excipulum. Given just the single collection of the species and having not collected at sites with similar
habitats in the near surroundings, especially north of Sán Carlos de Bariloche, it is very likely that the species
occurs more frequently in the Andean Cordillera. However, we were unable to find U. pallidocarpa and U.
perpusilla about 1000 km further south in the El Chalten region in Santa Cruz.
Verrucaria rhizicola Aptroot & Thüs, sp. nov. (Fig. 25C–D) Mycobank MB 517817
Verrucaria corticola ad radicem Alnis saepe in aqua submergenti, thallo viridi, ascomatis superficialibus nigris,
involucrellum dimidiatum, ascosporis reniformibus (10-) 12-15 µm longis. http://www.eol.org/pages/Verrucaria
rhizicola
Type:—France. Pas de Calais, Forêt de Boulogne, plot 99, 1 May 2009, Aptroot 68737 (holotype ABL).
Thallus corticolous, crustose, continuous with finely uneven surface appearing smooth from a distance,
green, dull, remaining opaque when wet (not subgelatinous), 25–50 µm thick, covering areas of up to 2 cm
diameter at the upper surface of roots, margins not delimited by a hypothallus, without blackened basal layer
and without back dots. Cortex absent. Algae chlorococcoid, cells round or more often ellipsoid to angular,
irregularly arranged in the thallus, ca. 5–10 × 7–15 µ m. Ascomata superficial, black, shiny, 0.1–0.2 mm
diameter, with black clypeus above that extends sideways as a saucer-like flat disc with a smooth surface,
covering the upper 1/3 to 2/3 of the exciple in some perithecia or reaching the exciple base; exciple hyaline to
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pale brown below. Ostiole 10–25 µm wide. Hamathecium consisting of unbranched periphyses of 7–10 × 2–3
µ m in gel near the ostiole. Asci clavate, 25–40 × 9–15 µm. Ascospores 8/ascus, hyaline, long ellipsoid,
asymmetrical with one side flattened, in some views kidney- or D-shaped, (10–)12–15 × 4.5–6 µ m, on
average 12 × 5 µm (25 measured). Pycnidia unknown.
FIGURE 25. A–B. Usnea pallidocarpa (holotype). A. Thallus with apothecia. B. Apothecium enlarged. C–D.
Verrucaria rhizicola (holotype). C. Thallus with perithecia. D. Perithecia enlarged. E–F. Xanthomendoza rosmarieae
(isotype). E. General habit. F. Enlarged fragments with soredia. Scale in A = 10 mm, in B = 5 mm, in C, F = 1 mm, in D
= 0.5 mm, in E = 3 mm.
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Distribution and habitat:—Only known from the type from roots of Alnus glutinosa along small rivulet
in temperate forest. Irregularly submersed in fresh water.
Etymology:—After the substrate.
Numerous aquatic species have been described in Verrucaria, but only 20 corticolous species are known
(Lendemer & Breuss 2009). The special habitat is not a character per se, as for instance of the 22 lichen
species found on submerged alder roots in Lithuania (Motiejunaite 2003) nine are known to be more
widespread on rocks (Smith et al. 2009), amongst them distinctive species such as Bacidia arnoldiana Körb.
and B. inundata (Fr.) Körb. Furthermore, some terrestrial saxicolous Verrucaria species, e.g. V. muralis Ach.,
are occasionally found on bark It remains to be seen whether not some of the Verrucaria which are described
as strictly corticolous, like V. viridigrana Breuss, are not in fact occurrences of species usually growing on
other substrates (in this case probably V. bryoctona (Th.Fr.) A.Orange, a species known from soil, moss,
lignum, leather and other waste materials). However, the species described here is unique among all known
species of Verrucaria, whether aquatic, terrestrial, corticolous or saxicolous, because of the prominent, tiny,
shiny ascomata with a distinctive dimidiate involucrellum and the asymmetrically kidney-shaped ascospores
with a size which is larger than e.g. in V. aquatilis and some of the corticolous species but smaller than other
Verrucaria species which generally occur on bark or rocks. In freshwater habitats Hydropunctaria
rheithrophila (Zschacke) Keller, Gueidan & Thüs can have similar sized ascospores, but this species has
immersed perithecia with a usually rough surface and a subgelatinous thallus, which often contains black dots
(Thüs & Schultz 2008). Verrucaria cinereolutescens Zschacke is a poorly known taxon with only slightly
shorter ascospores but a yellowish, thick thallus and immersed perithecia with a dark brown exciple
(Zschacke 1927). Verrucaria madida Orange has a subgelatinous thallus, perithecia covered by a thallus
mantle and an olive-grey pigment in the cortex (Orange 2004). Among the corticolus species treated by
Lendemer & Breuss (2009) Verrucaria lignicola Zschacke comes closest but has larger ascospores (15–19 ×
7–9 µm). Verrucaria lignicola is regarded as a synonym of the mostly saxicolus Verrucaria sublobulata in
Thüs & Schultz (2008). Re-examination of the type material (Bouly de Lesdain: Spa, racines d’Alnus dans un
ruisseau. 10.08.1904. W: Aqcu-No. 1921-718) has comfirmed the range of ascospore size quoted by Zschacke
which extends the documented minimual range for V. sublobulata marginally from 16 to 15 µm. The thallus of
the type material of V. lignicola is sharply delimited, in parts with a distinctly cracked surface and half
immersed perithecia. A section reveiled a thin involucrellum that is thin and fading downwards, making the
type material indistinguishable from V. sublobulata Eitner ex Servit. The new species grows together on the
roots with Porina aenea (Wallr.) Zahlbr. Further accompanying species in this lichen-rich forest are
mentioned in Van den Broeck et al. (2009).
Xanthomendoza rosmarieae S.Y.Kondr. & Kärnefelt, sp. nov. (Fig. 25E–F) Mycobank MB 517818
Xanthomendozae gallowayi similis, sed sorediis isidiosis, rhizinis latioribus brevioribusque (0.06–0.12 × 0.3–0.4 mm),
ascosporis angustioribus (6–8(–11) µm latis), et excipulo vero , e typo 'textura intricata', minus evoluto differt.—
http://www.eol.org/pages/Xanthomendoza rosmarieae
Type: U.S.A. Delaware: Sussex Co., Georgetown, northern entrance drive of Stockley Centre (on bark of
aspen growing together with Parmelia sulcata), April 2000, Crichton 57 (holotype ZH, isotype K).
Thallus of numerous semierect, very narrow, richly branched and anastomosing, yellowish to whitish
orange, sorediate lobes; soralia becoming isidiose, either marginal, of fallax-type, (0.2–)0.3–1(–1.2) mm long,
or on the underside, of ulophyllodes-type, as a greenish spherical mass, (28–)35–50(–60) µ m diameter.
Sometimes 2–3 soredia aggregated in irregular groups, to 90(–100) µm across, where true conblastidia are
absent. Lobes mainly horizontally orientated and somewhat raised above the substrate; main lobes 2–2.5(–4)
mm long and 0.1–0.2 mm wide, becoming narrower towards the tips, 0.4–0.5(–0.6) mm wide at the base,
usually only 1 mm long fragments seen; secondary lobules ca. 0.1 mm wide throughout and 1–1.5 mm long;
the total width of a lobe including secondary lobules are 2–2.5 mm. Rhizines variable in size but generally
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LUMBSCH ET AL.
rather short, 0.05–)0.06–0.12 mm diameter and (0.2–)0.3–0.4(–0.5) mm long. Thallus in section (90–)110–
120(–150) µ m thick, upper cortical layer to 25(–30) µ m thick, more or less mesodermatous
paraplectenchymatous, lower cortical layer to 12(–30) µ m thick, leptodermatous to mesodermatous
paraplectenchymatous with vertically elongated cell lumina; algal layer 30–40 µm thick; medulla to 40–50(–
70) µm thick, usually with a few scattered hyphae, sometimes with paraplectenchymatous parts, sometimes
completely filled with algal clusters. Apothecia to 1.5(–2.2) mm diameter, 0.35–0.4 mm high, lecanorine,
thalline margin to 0.1–0.15(–0.2) mm wide; cortical layer on the underside 25–30(–40) µ m thick,
mesodermatous paraplectenchymatous; true exciple often indistinct or poorly developed, ‘textura intricata’;
hymenium 70–90 µm high; subhymenium 20–30 µm thick; paraphyses ca. 2 µm diameter in lower portion,
and to 4–5 µm diameter towards the tips, very often with oil droplets (better seen in K); asci 8-spored but
often with 2–4–6 mature bipolar ascospores together with aborted two-celled spores; ascospores from
elongated ellipsoid to broadly ellipsoid or ovoid with a wide septum; (8–)10–16 × 6–8(–11) µm in water and
(10–)11–14(–16) × 7–10 µm in K, septum rather wide, (3–)5–8 µm wide in water and (4–)7–8 µm wide in K;
spermogonia to 360 µm diameter, punctiform, reddish; spermatia very narrow, bacilliform, 3.1–4.0 × 0.6–0.8
µm.
Distribution and habitat:—This new species is only known from the type locality where it grows on
bark of trees.
Etymology:—Xanthomendoza rosmarieae is named for the distinguished Swiss lichenologist, Rosmarie
Honegger, in recognition of her unique contributions to xanthorioid lichens.
Xanthomendoza S.Y.Kondr. & Kärnefelt is a genus of 20 species found mainly on tree trunks in the
Northern Hemisphere and on rocks in both North and South America. The genus is characterized by mainly
foliose thalli with well-developed rhizines (Kondratyuk & Kärnefelt 1997, 2003a, Søchting et al. 2002,
Fedorenko et al. 2009, Kondratyuk et al. 2011). This new species is distinguished by the presence of isidiate
soredia, rather wide and short rhizines, narrow ascospores, and a poorly developed true exciple of textura
intricata type. The rhizines of X. rosmarieae are broader (90–100 µ m wide) in comparison most similar
species of the X. gallowayi group (50–60 µ m wide) and the genus Oxneria (Kondratyuk & Poelt 1997)
Furthermore, X. rosmarieae differs in lacking a strongly gelatinized pseudoprosoplectenchymatous true
exciple. In addition to the wider rhizines, X. rosmarieae differs from X. gallowayi (syn. X. weberi) and X.
sogdiana in having a higher hymenium and narrower ascospores (Kondratyuk & Kärnefelt 2003b,
Kondratyuk et al. 2011). The apothecial anatomy of X. rosmarieae is similar to that of the Southern
Hemisphere genus Jackelixia S.Y.Kondr. et al., belonging to the same phylogenetic branch as Xanthodactylon
(Fedorenko et al. 2009), in having the Jackelixia-type of true exciple, i.e. poorly developed in the basal
portion and its textura intricata nature.
Acknowledgements
Giralt et al. are grateful to Harrie Sipman for his assistance with the Latin diagnosis of Amandinea maritima.
The Galápagos National Park Service provided specimens permits for analysis of the new species in the
genera Buellia, Calopadia, Coccocarpia, and Megalospora by Bungartz et al. Specimens were collected in
the Galápagos Islands with support from a National Geographic Research Grant #8162-07 to F. Bungartz.
This publication is contribution number 2030 of the Charles Darwin Foundation for the Galapagos Islands.
María Inés Messuti thanks Universidad Nacional del Comahue and CONICET for financial support. ZP
acknowledges support of a project of Academy of CR (AV0Z60050516) and the Ministry of Education, Youth
and Sports of CR (no. 0021620828). P. Lambley kindly supplied interesting Bunodophoron-collections from
Papua New Guinea. M. Wedin acknowledges financial support from The Swedish Research Council (VR 6212006-3760, VR 621-2009-5372). U. Arup thanks G. Thor for placing the material of Caloplaca decipioides at
his disposal and letting him describe this new species. He also thanks P. Lassen for help with the Latin and L.
Christiansen for help with the chemical analysis. Søgaard et al. (authors of Caloplaca digitaurea) appreciate
ONE HUNDRED NEW SPECIES OF LICHENIZED FUNGI
Phytotaxa 18 © 2011 Magnolia Press
• 115
the assistance of L. Knudsen for production of the ITS sequences, L. Christiansen for operating the HPLC
analyzer, and A. Elvebakk, Tromsø, for the possibility to study his collections from Torres del Paine; the work
was supported by grant 2008_01_0645 from the Carlsberg Foundation. A. Frisch kindly wrote the Latin
diagnosis for Calvitimela uniseptata. L. Ferraro and A. Michlig, authors of Coenogonium flammeum, are
grateful to the Myndel Botanica Foundation for financial support of our fieldwork and to the authorities of the
National Park of Argentina for collection permits. D. Øvstedal and J. A. Elix are indepted to Mrs. B. Helle and
Mr. J. Berge, both Bergen, for help with the illustration, and to Mr. L. Silva, Recife, for donation of the
material of Echinoplaca pernambucensis. Goward et al. thank T. Spribille and C. Miller for performing TLC
of Hypogymnia samples and Platismatia wheeleri, and Tim Wheeler for providing a photograph. E. Timdal
would like to thank the curator of NY for the loan of the West-Indian Phyllopsora material in which the new
species of Krogia was discovered. A. Elvebakk would like to thank the staff at CANB for kind help and for
possibility to study their material of Pannaria phyllidiata, P. P. Aspaas, the University Library of Tromsø, for
translating the diagnosis, and J. A. Elix, National University of Australia for hospitality and company in the
field, together with G. Kantvilas and P. McCarthy. D. Galloway is grateful to P. M. Jørgensen for providing the
Latin diagnosis of Placopsis imshaugii; to J. A. Elix for chemical help, and to A. Fryday (MSC) for the loan
of specimens. A. Nordin (Uppsala) kindly wrote the latin diagnosis of Sagenidiopsis isidiata. Bibiana
Moncada and Robert Lücking thank David Galloway for valuable comments on the taxonomy of Sticta
venosa. Fieldwork on the genus Usnea on the Galápagos Islands by C. Truong and P. Clerc was supported by
a grant "Augustin Lombard" from the Société de Physique et d’Histoire Naturelle de Genève. The work was
made in collaboration with F. Bungartz and the Charles Darwin Foundation, and María de los Angeles
Herrera-Campos participated in the field trip. The ministry of environment of the Parque Nacional Galápagos
provided collection permits. N. Wirtz thanks M. I. Messuti for organizing and accompanying her on a field
trip to Tierra del Fuego. André Aptroot and Holger Thüs thank U. Passauer and A. Igersheimer (W) for the
loan of the type of Verrucaria lignicola Research in Hungary on foliicolous lichens is supported by the
Hungarian Scientific Research Fund (OTKA K81232). Edit Farkas is also grateful for the loan of type
specimens to the director Dr. Jan Kirschner, the curator Dr Zdeněk Palice and the staff at herbarium PRA-V
(Průhonice, Czech Republic). K. Papong thanks Mahasarakham University for support of her study on Thai
foliicolous lichens. M. A. Herrera-Campos, A. Barcenas, R. Miranda, and R. Lücking are grateful for grants
by CONACYT (#35008-V) and PAPIIT-UNAM (#IN025808). Víctor J. Rico acknowledges support from the
Spanish Ministerio de Educación y Ciencia (Flora Liquenológica Ibérica project, CGL2007-66734-C03-01)
and from BSCH-UCM (GR58/08-910773). This publication was also supported by a grant of EOL initiative
to support a meeting entitled “Parmeliaceae-Improving our understanding of taxonomy, classification and
biogeography of the largest family of lichen-forming fungi”. Material of some of the new species was
collected and studied in the frame of the NSF-funded projects TICOLICHEN (DEB 0206125 to The Field
Museum; PI Robert Lücking), "Phylogeny and Taxonomy of Ostropalean Fungi, with Emphasis on the
Lichen-forming Thelotremataceae" (DEB 0516116 to The Field Museum; PI H. T. Lumbsch; Co-PI R.
Lücking), and "Neotropical Epiphytic Microlichens – An Innovative Inventory of a Highly Diverse yet Little
Known Group of Symbiotic Organisms" (DEB 715660 to The Field Museum; PI R. Lücking). The
Caterpillar® company provided funds to study lichens and other cryptogams from Panama. Maarten
Christenhusz is thanked for editing the final version of this manuscript. This publication initiates our worldwide and community-wide revision of the lichen family Graphidaceae in the frame of the NSF-funded project
"ATM – Assembling a taxonomic monograph: The lichen family Graphidaceae" (DEB-1025861 to The Field
Museum; PI T. Lumbsch, CoPI R. Lücking).
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