Folia Cryptog. Estonica, Fasc. 41: 1–4 (2005)
EDITORIAL
The 5th Symposium of the International Association for
Lichenology LICHENS IN FOCUS
held in Tartu (Estonia) 16–21 August 2004
This was a historical moment, at least for us
in the Local Organising Committee – to receive
hundreds of lichenologists in our small university town Tartu. It was unbelievable but true – you
could meet a lichenologist on every street corner
in the center of Tartu during the dinner break,
or in the Gunpowder Cellar and Wilde Pub every
evening (or perhaps even at night time). The concentration of lichenologists in the population of
Tartu (which had somewhat diminished due to
the summer holidays) was surely highest of all
times, and could be perceived also by eye – blue
bags represented the most popular trend among
any bags worn at that time in Tartu.
But let’s speak through numbers. There were
exactly 250 registered participants from 36
countries; six scientific sessions (both oral and
poster sessions) took place as well as three discussion sessions; 65 lectures and 153 posters
were presented in the following proportions:
I session “Systematics and Evolution” – 11
lectures and 44 posters;
II session “Quality and Quantity: maintaining
biological diversity in space and time” – 13
lectures and 34 posters;
III session “Genes, Physiology and Structure”
– 14 lectures and 16 posters;
IV session “Contributions of Lichen Ecology
to a better understanding of lichens in ecosystems” – 10 lectures and 29 posters;
V session “Lichen Photobionts – physiological, ecological and phylogenetic aspects of
their diversity” – 7 lectures and 9 posters;
VI session “Lichen Uses” – 10 lectures and
21 posters.
Furthermore, we listened to the exciting opening lecture by Angela Douglas “Symbiosis:
cooperation, slavery or domestification?” and
could present everyone’s opinion during the
discussion sessions “Translation of phyloge-
netic analyses into classification” (convened
by Thorsten Lumbsch), “In search of model
organisms” (convened by Rosmarie Honegger)
and “Phylogenetic methods” (convened by
Francois Lutzoni).
It is a well-known difficulty in preparing
any meeting, symposium or congress – how
to design the schedule of presentations. Quite
normally every participant is eager to introduce
his or her latest studies and new results. Some
persons are willing to give a lecture and others
prefer to prepare posters. The usual opinion is
that lecturers belong among Very Important
Persons and authors of posters should be students. This time it was different, not vice versa,
but just different: there were VIPs and students
among both categories – the lecturers and the
authors of posters. Later several participants
of respectable age commented: the significant
proportion of young persons among the participants of the symposium was the most striking
news of IAL5.
For this thanks are due to the Scientific
Committee which consisted of “triumvirates”
(Chairman, Convener and Posterman) of every
scientific session + conveners of discussion session. Those 21 persons carried out the difficult
job of revising abstracts that had been submitted
to the symposium, and designed the content of
sessions.
The meeting offered us also a good opportunity to introduce our countryside and nature.
Altogether 64 persons participated in three
excursions which were organised to different
parts of Estonia. The participants of the longest, pre-symposium excursion visited Lahemaa
and Soomaa National Parks, drove through the
Russian old orthodox villages, travelled to the
three greatest western islands – Saaremaa,
Muhumaa and Hiiumaa. The post-symposium
excursion to the eastern and northern parts of
Estonia took the participants to the swamp in
Endla Nature Reserve, demonstrated the wood-
2
Folia Cryptog. Estonica
Participants of the 5th IAL Symposium (August 17, 2004 in
Vanemuise Concert Hall)
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Hiroyuki Kashiwadani (Japan)
Kansri Boonpragob (Thailand)
Kawinnat Buaruang (Thailand)
Kwang Hee Moon (Korea)
Fenja Brodo (Canada)
Tatiana Makry (Russia)
Ave Suija (Estonia)
Tiiu Tõrra (Estonia)
Palmira Carvalho (Portugal)
Mauro Tretiach (Italy)
Rui Figuera (Portugal)
Elena Molodtsova (Russia)
Carolina Cornejo (Switzerland)
Laurens B. Sparrius
(Netherlands)
Lauri Saag (Estonia)
Inga Jüriado (Estonia)
Åsa Dahlkild (Sweden)
Derek Persoh (Germany)
Otto L. Lange (Germany)
Mark Seaward (UK)
Yoshiaki Kon (Japan)
Gennadii Urbanavichus
(Russia)
Irina Urbanavichene (Russia)
Svetlana Ektova (Russia)
Evgenia Mouchnik (Russia)
Susan Will-Wolf (USA)
Silvana Munzi (Italy)
Mari-Liis Rebane (Estonia)
Katrin Kolnes (Estonia)
Vagn Alstrup (Denmark)
Guido Incerti (Italy)
Juri Nascimbene (Italy)
Mikhail Piskaryov (Russia)
Daniela Csencics (Switzerland)
Erast Parmasto (Estonia)
Anna Zalewska (Poland)
Katarzyna Jando (Poland)
Beata Guzow-Krzeminska
(Poland)
Agniezka Kowalewska (Poland)
Eric Steen Hansen (Denmark)
Ulf Schiefelbein (Germany)
Frank Bungartz (Germany)
Irwin Brodo (Canada)
Sanja Savic (Sweden)
Harrie Sipman (Germany)
Jouko Rikkinen (Finland)
Thilo Hasse (Germany)
Natalia Davydova (Russia)
Evgeny Davydov (Russia)
Line Balschmidt (Denmark)
Piret Lõhmus (Estonia)
Evelyn Silvet (Estonia)
Bettina Staiger (Germany)
Christoph Scheidegger
(Switzerland)
Alessia Fappiano (Italy)
Renato Benesperi (Italy)
Andres Saag (Estonia)
Roland Moberg (Sweden)
Paula de Priest (USA)
Lucia Muggia (Italy)
Magdalena Opanowicz (Poland)
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Knut Asbjorn Solhaug (Norway)
Per Gerhard Ihlen (Sweden)
Asuncion de los Rios (Spain)
Nadezhda Alexeeva (Russia)
Guillermo Amo de Paz (Spain)
Francois Lutzoni (USA)
Jolanta Miadlikowska (USA)
Jana Kocourkova (Czech Republic)
Valerie Hofstetter (USA)
Martin Kukwa (Poland)
Beata Krzewicka (Poland)
Pawel Czarnota (Poland)
Paola Adamo (Italy)
Paolo Giordani (Italy)
Stefano Martellos (Italy)
Cecile Gueidan (USA)
Rebecca Yahr (USA)
Christopher Ellis (UK)
Anna Crewe (Sweden)
Valerie Reeb (USA)
Torbjorg Bjelland (Norway)
William Purvis (UK)
Cameron Williams (USA)
Helga Bültmann Germany)
Ljudmilla Martin (Estonia)
Bruce McCune (USA)
Begona Aguirre-Hudson (UK)
Victor Jimenez Rico (Spain)
Ruth del Prado (USA)
Ineke Beltman (Netherlands)
Alexey Zavarzin (Russia)
Olga Merkulova (Russia)
Christine Keller (Switzerland)
Maarja Nõmm (Estonia)
Ede Leppik (Estonia)
Edit Farkas (Hungaary)
Peter Scholtz (Germany)
Starri Heidmarsson (Iceland)
Imke Schmitt (USA)
Zdenek Palice (Czech Republic)
Stepanka Bayerova (Czech Republic)
Silke Werth (Switzerland)
Ondrej Peksa (Czech Republic)
Anna Guttova (Slovakia)
David Svoboda (Czech Republic)
Mahroo Haji Moniri Anbaran (Iran)
Tiina Randlane (Estonia)
Suzanne Joneson (USA)
Alexandro Caruso (Sweden)
Heath O’Brian (USA)
Shyam Nyatti (Switzerland)
Sandra Scherrer (Switzerland)
Christof Eichenberger (Switzerland)
David Hawksworth (Spain)
Jurga Motiejunaite (Lithuania)
Holger Thüs (Germany)
Milos Bartak (Czech Republic)
Ulrik Sochting (Denmark)
Alexander Paukov (Russia)
Helmut Mayrhofer (Austria)
Josef Hajek (Czech Republic)
Elisabeth Baloch (Austria)
Sabine Wornik (Austria)
Alexander Taran (Russia)
Svetlana Tchabanenko (Russia)
4
Folia Cryptog. Estonica
138.
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Juliane Blaha (Austria)
Irina Mikhailova (Russia)
Franc Batic (Slovenia)
Margarita Magomedova
(Russia)
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Isao Yoshimura (Japan)
Oleg Blum (Ukraine)
Jüri Martin (Estonia)
Zvonka Jeran (Slovenia)
Louise Lindblom (Sweden)
Rene Spigelberg Larsen
(Denmark)
Thomas Nash III (USA)
Frank Kauff (USA)
Armin Mangold (USA)
Mats Wedin (Sweden)
Anders Nordin (Sweden)
Elfie Stocker-Wörgötter
(Austria)
André Aptroot (Netherlands)
Francoise Rolley (France)
Silvia Stofer (Switzerland)
Wanaruk Saipunkaew
(Thailand)
Teuvo Ahti (Finland)
Astri Botnen (Norway)
Pat Wolseley (UK)
Katherine Glew (USA)
165. Christian Printzen
(Germany)
166. Scott LaGreca (UK)
167. Tor Tonsberg (Norway)
168. Leena Myllys (Finland)
169. Robert Lücking (USA)
170. Laura Kivistö (Finland)
171. Rosa Perez Perez (Mexico)
172. Heini Hyvärinen (Finland)
173. Heidi Döring (Sweden)
174. Philippe Clerc (Switzerland)
175. Filip Högnabba (Finland)
176. Henrik Hedenås (Sweden)
177. Lucyna Sliwa (Poland)
178. Geir Hestmark (Norway)
179. William Sanders (Spain)
180. Ingrid Berney (Switzerland)
181. Diane Fahselt (USA)
182.
183. Arne Thell (Sweden)
184. Enrica Matteucci (Italy)
185. Elena Pittao (Italy)
186. Paola Crisafulli (Italy)
187. Jacob Garty (Israel)
188. Deborah Isocrono (Italy)
189.
190. Sergio Enrico Favero-Longo
(Italy)
191. Lauro Xavier-Filho (Brazil)
192. Mohammad Sohrabi (Iran)
193. Domenico Puntillo (Italy)
land key-habitat, wooded meadow and limestone
fields. Another post-symposium excursion was
organised to visit the beautiful sites in southern Estonia such as Devonian sandstone cliffs
in Taevaskoja, raised bog in Meenikunno and
sandy dunes close to Värska. The choice of visited sites turned out to be the most successful – as
it can be read from the contribution by Aptroot
et al. in the present volume – during these three
excursions 30 species of lichens or lichenicolous
fungi new to the country were collected!
Contributions presented during IAL5 will be
recorded in the various proceedings of the
symposium. This volume of Folia Cryptogamica
Estonica is only one of them; special volumes
or special parts of ordinary volumes of such
periodicals as Lichenologist, Environmental
Pollution and Journal of Vegetation Science
will follow. In this issue you will find 13 papers
based on poster presentations, plus a summary
of lichenised or lichenicolous taxa new or rare
in Estonia, which were found during the excursions or the symposium.
The organisation of the 5th Symposium of the International Association for Lichenology “Lichens
in Focus” was a joint effort of the Local Organis-
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Pier Luigi Nimis (Italy)
Nora Wirtz (Germany)
Thorsten Lumbsch (USA)
Rikke Reese Naesborg (Sweden)
Jan-Eric Mattsson (Sweden)
Matthias Schultz (Germany)
Tassilo Feuerer (Germany)
Anders Tehler (Sweden)
Håkan Lättman (Sweden)
Orvo Vitikainen (Finland)
Emmanuël Sérusiaux
(Belgium)
Gintaras Kantvilas
(Australia)
Peter Crittenden (UK)
Rikard Sundin (Sweden)
Rosmarie Honegger
(Switzerland)
Michaela Schmull
(Germany)
Manfred Jennsen (Germany)
Elisabeth Lay (USA)
Maria Angeles HerreraCampos (Mexico)
Angela Douglas (UK)
Fabian Seymour (UK)
Jutta Buschbom (Germany)
Ana Pintado (Spain)
Stefano Loppi (Spain)
Alar Suija (Estonia)
ing Committee, IAL5 Scientific Committee and
the IAL Council which served during 2000–2004.
The main principles for the organisation of this
event as well as numerous small details of even
marginal problems were thoroughly discussed
and decided first of all by the IAL Council. The
Local Organising Committe felt the support
of the Council all the time during these four
preparatory years. Therefore we would like to
express our sincere gratitude to the people who
formed this fantastic IAL Council:
Pier Luigi Nimis
Irwin Brodo
Leopoldo Sancho
Francois Lutzoni
Christoph Scheidegger
Martin Grube
Jack Elix
Rosmarie Honegger
Gintaras Kantvilas
The orchids which we received at the IAL dinner
are still in blossoms – reminding us of these days
in August 2004 that now, in retrospect, seem to
have been nothing but pure pleasure.
Tiina Randlane & Andres Saag
Folia Cryptog. Estonica, Fasc. 41: 5–12 (2005)
Lichens from islands in the Russian part of the Gulf of Finland
Nadezhda Alexeeva
Department of Botany, St. Petersburg State University, Universitetskaya emb. 7/9, 199034 St. Petersburg, Russia
E-mail: baltic@teia.org
Abstract: The present paper is based on material from several recent collections made between 1993 and 2003 on islands
in the central part of the Gulf of Finland (Gogland, Bolshoy Tuters, Maly Tuters, Seskar, Moschny, Maly, Sommers) and
several islands situated near the Russian-Finnish border (Bolshoy Pogranichny and surrounding islands). The list of lichen
species includes 170 species and 3 subspecies. Amandinea cacuminum, Buellia badia, Caloplaca cerinella, C. diphyodes, C. microthallina,
Cladonia borealis, C. diversa, C. metacorallifera, Lecanora marginata, L. persimilis, L. rimicola, L. salina, Lepraria caesioalba, Melanelia
disjuncta, Ramalina polymorpha, Rhizocarpon richardii, Rinodina gennarii, Stereocaulon incrustatum and S. rivulorum have been reported
for the Leningrad region only from these islands. Bryoria subcana, Cladonia macrophylla, Flavocetraria nivalis, Melanelia hepatizon,
M. stygia, Neofuscelia pulla, Umbilicaria hirsuta, U. hyperborea, Peltigera scabrosa and Ramalina fraxinea are red-listed in the Leningrad
region.
Kokkuvõte: Samblikud Venemaale kuuluvatel Soome lahe saartel.
Töö põhineb aastatel 1993 kuni 2003 Soome lahe keskosa (Suursaar, Suur Tütarsaar, Väike Tütarsaar, Seiskari, Lavansaari,
Penisaari, Sommers) ja Vene-Soome piiri lähedastelt saartelt (Paatio jt.) kogutud kollektsioonidel. Samblikuliikide nimekiri
koosneb 170 liigist ja 3 alamliigist. Leningradi oblastis on liike Amandinea cacuminum, Buellia badia, Caloplaca cerinella, C. diphyodes,
C. microthallina, Cladonia borealis, C. diversa, C. metacorallifera, Lecanora marginata, L. persimilis, L. rimicola, L. salina, Lepraria caesioalba,
Melanelia disjuncta, Ramalina polymorpha, Rhizocarpon richardii, Rinodina gennarii, Stereocaulon incrustatum ja S. rivulorum leitud ainult
nendelt saartelt. Bryoria subcana, Cladonia macrophylla, Flavocetraria nivalis, Melanelia hepatizon, M. stygia, Neofuscelia pulla, Umbilicaria
hirsuta, U. hyperborea, Peltigera scabrosa ja Ramalina fraxinea kuuluvad Leningradi oblasti punasesse nimekirja.
INTRODUCTION
Many islands situated in the Russian part of
the Gulf of Finland have been closed for visiting during the last 60 years due to the Soviet
military requirements. Before World War ll this
area belonged to Finland and on the largest islands there were several settlements. Nevertheless, neither large-scale agriculture nor forestry
have been practised on the islands. Thus the
natural complexes and landscape diversity of
these territories nearby St. Petersburg have
mainly been left undisturbed. During the war
civil inhabitants were forced to leave this area.
In 1944 the islands became a part of USSR and
were included into the so called “border zone”,
where access was practically forbidden.
The first notes on richness and uniqueness
of the natural complexes of the islands were
published by zoologists and botanists who visited these territories in early 1990-s after almost
50-years period of strict protection. Recent inventories of the flora revealed that these islands
are extraordinary rich in species and support
populations of many red-listed plants (e.g. Glazkova, 2002). The current lichen diversity of the
islands remained poorly studied until recently.
The lichen flora of Gogland (Hogland) island was
investigated by Brenner (1886), but modern detailed review of his collections is still needed. No
further inventories of lichens were made on the
islands for over a century. In 2002 a list of 120
lichen species collected in 1994 from Bolshoy
Tuters island was published by Andreyev (2002).
There are also scanty collections of lichens made
between 1993 and 2003 on other islands in the
central part of the Gulf of Finland (Gogland,
Bolshoy Tuters, Maly Tuters, Seskar, Moschny,
Maly, Sommers, administratively belonging to
Kingiseppsky district of the Leningrad region)
and several islands situated near the Russian-Finnish border (Bolshoy Pogranichny and
surrounding islands, Vyborgsky district of the
Leningrad region).
Here I present the first summary of the lichens found on the remote islands of the Gulf
of Finland in recent years. Hopefully this will
help to attract lichenologists’ attention to these
highly interesting areas.
The islands under consideration are situated between 59o48’–60o32’N and 26o55’–28o25’E
(Fig.1) and differ considerably in their area,
6
Folia Cryptog. Estonica
height and appearance. Gogland and Bolshoy
Tuters are high islands (176 and 56 m a.s.l.
respectively), with rocks, cliffs and stone terraces, formed by gneisses, granites, gneissesgranites, quarz-porphyries and quarzite. Sommers island consists of low granite outcrops,
so-called “bullheads”. In contrast, Maly Tuters,
Moschny, Maly and Seskar are low islands (5–15
m a.s.l.); their surface cover consists of moraine
and marine deposits of sand, rubble, shingle
and gravel. Along with Bolshoy Tuters, they
are famous for their dune systems and beach
ridges (Glazkova, 2001). Bolshoy Pogranichny
and surrounding islands are formed by rocky
(predominantly rapakivi granites) ridges up to
20–30 m high. Most of the islands are covered
with forests. Pine (Pinus sylvestris) forests are
prevailing. Spruce forests formed by Picea
abies and secondary small-leaved forests with
Betula pendula, B. pubescens, Populus tremula
and Sorbus aucuparia are also present. Alnus
glutinosa stands occur along the seacoast.
Broad-leaved trees are mainly restricted to the
previous settlements. Various types of meadows
and sandy heathlands, as well as fens, raised
bogs and transitory bogs can be found on
particular islands (Glazkova, 2001; Noskov &
Botch, 1999).
MATERIALS AND METHODS
The present paper is based on material from
several collections made on the islands between
1993 and 2003 (names of the islands, year of collection and number of samples see in Table 1). In
the course of the present studies collections of N.
Balashova, E. Glazkova and A. Gaginskaya were
revised. The specimens were determined with
routine techniques. Secondary compounds of
Cladonia borealis were analysed according to the
standardised TLC method. Herbarium samples
are kept at the department of Botany of St. Petersburg State University (LECB). Other records
used include list of species for Bolshoy Tuters
island published by M. Andreyev; his collections
are deposited in the herbaria of Komarov Botanical Institute (LE), Uppsala University (UPS) and
Bergen Universiry (BG) (Andreyev, 2002).
RESULTS
A total of 170 species and 3 subspecies are
reported for the islands. The majority of these
species are common in the Leningrad region. Still,
some species identified are of special interest.
Amandinea cacuminum, Buellia badia, Caloplaca
cerinella, C. diphyodes, C. microthallina,
FINLAND
B. Pogranichny
Sommers
Gogland
Maly
RUSSIA
St. Petersburg
Seskar
Moschny
B. Tuters
M. Tuters
ESTONIA
Fig. 1. Location of the islands Gogland, Bolshoy Tuters, Maly Tuters, Seskar, Moschny, Maly,
Sommers, Bolshoy Pogranichny in the Gulf of Finland.
7
Table 1. The area of the islands, collector name and year of collection
Russian names
Finnish names
Suursaari
Abbreviations
Gog
Area,
(km2)
20.64
Gogland
Bolshoy Tuters
Suur Tutersaari
BTu
9.2
Maly Tuters
Seskar
Moschny
Maly
Sommers
Bolshoy Pogranichny
(and complex of surrounding islands)
Pieni Tutersaari
Seiskari
Lavansaari
Peninsaari
Sommers
Paatio
MTu
Ses
Mo
Mal
Som
BPo
3.0
4.5
16.5
3.5
0.14
9.0
Cladonia borealis, C. diversa, C. metacorallifera,
Lecanora marginata, L. persimilis, L. rimicola, L.
salina, Lepraria caesioalba, Melanelia disjuncta,
Ramalina polymorpha, Rhizocarpon richardii,
Rinodina gennarii, Stereocaulon incrustatum
and S. rivulorum have been reported for the
Leningrad region only from these islands.
Bryoria subcana, Cladonia macrophylla,
Flavocetraria nivalis, Melanelia hepatizon, M.
stygia, Neofuscelia pulla, Umbilicaria hirsuta,
U. hyperborea, Peltigera scabrosa and Ramalina
fraxinea are red-listed in the Leningrad region
(Tzvelev, 2000). The characteristic feature of the
territory is the occurrence of oceanic and suboceanic species, such as Caloplaca scopularis,
Cladonia ramulosa, C. scabriuscula, Peltigera
hymenina and Ramalina polymorpha.
List of species
The following list is presented in alphabetical
order. The nomenclature follows Santesson et
al. (2004). Abbreviations of islands see in Table
1. Species mentioned only by Andreyev (2002)
and not found in other collections are marked
with asterisk (*). Notes on ecology mentioned by
M. Andreyev are presented in brackets.
*ACAROSPORA FUSCATA (Schrad.) Th. Fr. – BTu;
(Andreyev: on granite pebbles on the road,
granite boulders on dunes).
AMANDINEA CACUMINUM (Th. Fr.) H. Mayrhofer &
Sheard – MTu; on siliceous pebble.
Collector name, year of collection
and number of samples
N. Balashova, 1993, 12 samples
A. Gaginskaya, 2003, 32 samples
N. Balashova, 1993, 26 samples
M. Andreyev, 1994, 240 samples
(Andreyev, 2002)
N. Balashova, 1993, ca 60 samples
N. Balashova, 1993, 16 samples
N. Balashova, 1993, 12 samples
N. Balashova, 1993, 37 samples
E. Glazkova, 1995, ca 50 samples
E. Glazkova, 2002, ca 140 samples
A. CONIOPS (Wahlenb.) M. Choisy ex Scheid. & H.
Mayrhofer – MTu; on siliceous pebble.
A. PUNCTATA (Hoffm.) Coppins & Scheid. – Mal,
BTu; on bark of deciduous trees (Andreyev:
Padus avium, Quercus robur near the ruins
of old village).
*ASPICILIA CAESIOCINEREA (Nyl. ex Malbr.) Arnold
– BTu; (Andreyev: on granite rocks and
boulders on the coast, granite pebbles on
the road).
A. CINEREA (L.) Körb. – Som; on siliceous pebble.
BRYORIA CAPILLARIS (Ach.) Brodo & D. Hawksw.
– BTu, Gog; on bark of coniferous trees
(Andreyev: Picea abies in the young spruce
forest).
*B. FUSCESCENS (Gyeln.) Brodo & D. Hawksw.
– BTu; (Andreyev: on bark of Picea abies in
the spruce forest).
B. SUBCANA (Nyl. ex Stizenb.) Brodo & D. Hawksw.
– Mal, MTu; on bark of coniferous trees.
*BUELLIA BADIA (Fr.) A. Massal. – BTu; (Andreyev:
on granite boulders on dunes).
*CALOPLACA CERINELLA (Nyl.) Flagey – BTu; (Andreyev: on bark of Acer platanoides near
the ruins of old village).
*C. DIPHYODES (Nyl.) Jatta – BTu; (Andreyev: on
granite rocks and boulders on the coast).
C. HOLOCARPA (Hoffm. ex Ach.) A.E. Wade – BTu,
BPo; on bark of deciduous trees (Andreyev:
Fraxinus excelsior near the ruins of old village).
8
*C.
Folia Cryptog. Estonica
(Wedd.) Zahlbr. – BTu; (Andreyev: on
granite rocks and boulders on the coast).
*C. MICTR OTHALLINA (Wedd.) Zahlbr. – BTu;
(Andreyev: on granite rocks and boulders
on the coast).
*C. SAXICOLA (Hoffm.) Nordin – BTu; (Andreyev: on
limestone in the ruined old village).
C. SCOPULARIS (Nyl.) Lettau – Som; on concrete.
CANDELARIELLA AURELLA (Hoffm.) Zahlbr. – Som,
BTu; on concrete (Andreyev: on limestone
in the ruined old village).
C. CORALLIZA (Nyl.) H. Magn. – MTu; on siliceous
pebble.
C. VITELLINA (Hoffm.) Müll. Arg. – BTu, MTu; on
siliceous pebbles (Andreyev: on granite rocks
and boulders on the coast, granite boulders
on dunes).
CETRARIA ACULEATA (Schreb.) Fr. – Mal; on soil.
C. ISLANDICA (L.) Ach. – Som, Mal, BTu, Gog, BPo;
on soil (Andreyev: on sand dunes).
C. MURICATA (Ach.) Eckfeldt – Som, Mal; on soil.
C. SEPINCOLA (Ehrh.) Ach. – BPo; on bark of
deciduous trees.
*CHAENOTHECA FERRUGINEA (Turner ex Sm.) Mig.
– BTu; (Andreyev: on wood in the pine
forest).
*CLADONIA AMAUROCRAEA (Flörke) Schaer. – BTu;
(Andreyev: on granite rocks in the pine
forest).
C LADONIA ARBUSCULA (Wallr.) Flot. ssp. MITIS
(Sandst.) Ruoss – Som, BTu; on soil (Andreyev: on sand dunes).
C. ARBUSCULA (Wallr.) Flot. ssp. SQUARROSA (Wallr.)
Ruoss – Som, Mal, BTu, Gog; on soil (Andreyev: on sand dunes, granite rocks in the
pine forest).
*C. BELLIDIFLORA (Ach.) Schaer. – BTu; (Andreyev:
on wood on dunes).
C. BOREALIS S. Stenroos – Mal, Gog; on soil.
C. BOTRYTES (K.G. Hagen) Willd. – BTu, BPo; on
soil, (Andreyev: on sand dunes).
*C. CENOTEA (Ach.) Schaer. – BTu; (Andreyev: on
wood in the pine forest).
*C. CHLOROPHAEA (Flörke ex Sommerf.) Spreng.
– BTu; (Andreyev: on granite rocks in the
pine forest).
*C. COCCIFERA (L.) Willd. – BTu; (Andreyev: on
granite rocks in the pine forest).
C. CONIOCRAEA (Flörke) Spreng. – Som, BTu,
BPo; on soil, wood, (Andreyev: on wood on
dunes).
C. CORNUTA (L.) Hoffm. ssp. CORNUTA – BTu, BPo;
on soil, (Andreyev: on wood in the pine forest
and on sand dunes).
MARINA
C.
(Ach.) Flot. – BTu, BPo; on soil (Andreyev: on wood in the pine forest and on
sand dunes).
*C. DEFORMIS (L.) Hoffm. – BTu; (Andreyev: on
wood on dunes).
C. DIGITATA (L.) Hoffm. – BPo; on decaying
wood.
C. DIVERSA Asperges – Som; on soil.
C. FIMBRIATA (L.) Fr. – Som, BTu, BPo; on soil
(Andreyev: on sand dunes).
C. FLOERKEANA (Fr.) Flörke – Som, BTu, BPo; on
soil (Andreyev: on wood on dunes).
C. FURCATA (Huds.) Schrad. – Som, Mal, BTu,
Gog, BPo; on soil (Andreyev: on sand dunes,
granite rocks in the pine forest).
C. GRACILIS (L.) Willd. ssp. GRACILIS – Som, BTu,
Gog, BPo; on soil (Andreyev: on granite rocks
in the pine forest).
*C. GRACILIS (L.) Willd. ssp. TURBINATA (Ach.) Ahti
– BTu; (Andreyev: on wood in the pine forest
and on sand dunes).
C. MACILENTA Hoffm. – Som, BTu, Mal; on soil
(Andreyev: on wood on dunes and in the
pine forest).
C. MACROPHYLLA (Schaer.) Stenh. – BPo; on soil.
C. METACORALLIFERA Asahina – Gog; on soil.
C. PHYLLOPHORA Hoffm. – BTu, Gog, BPo; on soil
(Andreyev: on granite rocks in the pine forest).
C. PLEUROTA (Flörke) Schaer. – BTu, Gog, BPo; on
soil (Andreyev: on granite rocks and wood
in the pine forest).
C. PYXIDATA (L.) Hoffm. – Som, Mal, BTu, BPo;
on soil.
*C. PORTENTOSA (Dufour) Coem. – BTu; (Andreyev:
on sand dunes).
C. RAMULOSA (With.) J.R. Laundon – Som; on
soil.
C. RANGIFERINA (L.) F.H. Wigg. – Som, Mal, BTu,
Gog, BPo; on soil (Andreyev: on sand dunes,
granite rocks in the pine forest).
*C. REI Schaer. – BTu; (Andreyev: on wood in the
pine forest).
C. SCABRIUSCULA (Delise) Nyl. – BTu; on soil.
C. SQUAMOSA Hoffm. – Som, BTu, Gog, BPo; on
soil (Andreyev: on granite rocks in the pine
forest).
C. STELLARIS (Opiz) Pouzar & Vězda – Mal, BTu;
on soil (Andreyev: on sand dunes, granite
rocks in the pine forest).
C. SUBULATA (L.) Weber ex F.H. Wigg. – BTu, BPo;
on soil (Andreyev: on wood and sand on
dunes).
CRISPATA
9
C. TURGIDA Hoffm. – Mal, BTu; on soil (Andreyev:
on soil in the young spruce forest, granite
rocks in the pine forest).
C. VERTICILLATA (Hoffm.) Schaer. – BTu, BPo; on
soil (Andreyev: on sand dunes).
C. UNCIALIS (L.) Weber ex F.H. Wigg. ssp. BIUNCIALIS
(Hoffm.) M.Choisy – Som, BTu; on soil.
C. UNCIALIS (L.) Weber ex F.H. Wigg ssp. UNCIALIS
– BTu, Gog; on soil (Andreyev: on granite
rocks in the pine forest).
FLAVOCETRARIA NIVALIS (L.) Kärnefelt & A. Thell
– Mal; on soil.
*HYPOCENOMYCE SCALARIS (Ach.) M. Choisy – BTu;
(Andreyev: on wood in the pine forest).
HYPOGYMNIA PHYSODES (L.) Nyl. – Som, Mal, BTu,
Gog, Ses, Mo, BPo; on bark of deciduous
and coniferous trees, rocky outcrops (Andreyev: on bark of Alnus sp., Picea abies,
wood, granite rocks and boulders, sand – in
spruce and pine forests, on the coast, on
sand dunes).
H. TUBULOSA (Schaer.) Hav. – Gog; on bark of
coniferous trees.
IMSHAUGIA ALEURITES (Ach.) S.L.F. Meyer – Gog; on
bark of Pinus sylvestris.
LASALLIA PUSTULATA (L.) Mérat – Som, BPo; on
siliceous rocks.
L ECANORA ALLOPHANA Nyl. – Ses; on bark of
deciduous trees.
*L. ARGENTATA (Ach.) Malme – BTu; (Andreyev: on
bark of Alnus sp. on the coast).
*L. CAMPESTRIS (Schaer.) Hue – BTu; (Andreyev:
on granite rocks in the pine forest).
L. CARPINEA (L.) Vain. – Mal, BTu, Gog, Ses, Mo;
on bark of deciduous trees (Andreyev: Acer
platanoides, Betula sp., Fraxinus excelsior,
Padus avium, Quercus robur, Sorbus aucuparia on the coast and near the ruins of
old village).
*L. CHLAROTERA Nyl. – BTu; (Andreyev: on bark
of Quercus robur near the ruins of old village).
L. DISPERSA (Pers.) Sommerf. – Som, BTu; on concrete (Andreyev: on limestone in the ruined
old village).
L. HAGENII (Ach.) Ach. – MTu, BPo; on bark of
deciduous trees.
*L. HELICOPIS (Wahlenb.) Ach. – BTu; (Andreyev: on
granite rocks and boulders on the coast).
*L. MARGINATA (Schaer.) Hertel & Rambold – BTu;
(Andreyev: on granite boulders on dunes).
*L. PERSIMILIS (Th. Fr.) Nyl. – BTu; (Andreyev: on
bark of Alnus sp. on the coast).
L.
(Ehrh. ex Hoffm.) Rabenh. – BTu,
MTu; on siliceous pebbles (Andreyev: on
granite pebbles on the road, granite boulders on dunes).
*L. PULICARIS (Pers.) Ach. – BTu; (Andreyev: on
bark of Picea abies, Sorbus aucuparia on
the coast).
*L. RIMICOLA H. Magn. – BTu; (Andreyev: on
granite boulders on the coast).
L. RUPICOLA (L.) Zahlbr. – Som, BTu, MTu; on
siliceous pebbles (Andreyev: on granite
boulders on dunes).
*L. SALINA H. Magn. – BTu; (Andreyev: on granite
rocks and boulders on the coast).
L. SULPHUREA (Hoffm.) Ach. – MTu; on siliceous
pebbles.
L. SYMMICTA (Ach.) Ach. – Ses, BTu, Mo; on bark of
deciduous trees (Andreyev: Alnus sp., Betula
sp., Sorbus aucuparia on the coast).
*L. VARIA (Hoffm.) Ach. – BTu; (Andreyev: on wood
on dunes).
*LECIDEA FUSCOATRA (L.) Ach. – BTu; (Andreyev: on
granite boulders on dunes).
L. LAPICIDA (Ach.) Ach. var. PANTHERINA Ach. – Som,
BTu, MTu; on siliceous pebbles (Andreyev:
on granite rocks and boulders on the coast,
granite boulders on dunes, granite pebbles
on the road).
*LECIDELLA ACHRISOTERA (Nyl.) Hertel & Leuckert –
BTu; (Andreyev: on bark of Acer platanoides,
Betula sp., Fraxinus excelsior, Quercus robur
near the ruins of old village).
*L. CARPATHICA Körb. – BTu; (Andreyev: on granite
rocks in the pine forest).
L. ELAEOCHROMA (Ach.) M. Choisy – Mal, BTu, Gog,
Ses; on bark of deciduous trees.
*L. EUPHOREA (Flörke) Hertel. – BTu; (Andreyev:
on bark of Padus avium near the ruins of
old village).
*LEPRARIA CAESIOALBA (de Lesd.) J.R. Laundon
– BTu; (Andreyev: on granite boulders in
the young spruce forest).
*MELANELIA DISJUNCTA (Erichsen) Essl. – BTu;
(Andreyev: on granite rocks and boulders
on the coast).
MELANELIA EXASPERATA (De Not.) Essl. – BPo; on
bark of deciduous trees.
M. EXASPERATULA (Nyl.) Essl. – Mal, BTu, Gog, Ses,
BPo; on bark of deciduous trees and dry
twigs of Picea abies (Andreyev: on Alnus
sp. on the coast).
*M. HEPATIZON (Ach.) A. Thell – BTu; (Andreyev:
on granite boulders on dunes).
POLYTROPA
10
M.
Folia Cryptog. Estonica
(L.) Essl. – Mal, BTu, Ses; on bark
of deciduous trees (Andreyev: on Alnus sp.,
Sorbus aucuparia on the coast).
*M. STYGIA (L.) Essl. – BTu; (Andreyev: on granite
boulders on dunes).
*NAETROCYMBE PUNCTIFORMIS (Pers.) R.C. Harris
– BTu; (Andreyev: on bark of Alnus sp.,
Betula sp. on the coast).
NEOFUSCELIA PULLA (Ach.) Essl. – Som, BTu, MTu;
on siliceous pebbles (Andreyev: on granite
rocks on the coast and in the pine forest,
granite boulders on dunes).
*N. VERRUCULIFERA (Nyl. ) Essl. – BTu; (Andreyev:
on granite rocks and boulders on the
coast).
*PACHYPHIALE FAGICOLA (Hepp) Zwackh – BTu; (Andreyev: on bark of Acer platanoides near the
ruins of old village).
*PARMELIA OMPHALODES (L.) Ach. – BTu; (Andreyev:
on granite pebbles on the road).
P. SAXATILIS (L.) Ach. – Som, Mal, BTu, MTu, Gog,
Ses, BPo; on siliceous pebbles and rocks
(Andreyev: on bark of Alnus sp., Sorbus aucuparia on the coast, on granite rocks on the
coast and in the pine forest, granite boulders
on dunes, granite pebbles on the road).
P. SULCATA Taylor – Mal, BTu, Gog, Ses, Mo, BPo;
on bark of deciduous trees (Andreyev: on
Sorbus aucuparia on the coast, Quercus
robur near the ruins of old village).
PARMELIOPSIS AMBIGUA (Wulfen) Nyl. – Mal, BTu, Mo,
BPo; on bark of deciduous and coniferous
trees, wood (Andreyev: on wood in the pine
forest).
P. HYPEROPTA (Ach.) Arnold – Gog, BPo; on bark
of deciduous and coniferous trees.
PELTIGERA APHTHOSA (L.) Willd. – BPo; on soil.
P. CANINA (L.) Willd. – Som, Mal, BTu, BPo; on
soil (Andreyev: on soil in the ruined old
village).
P. DIDACTYLA (With.) J.R. Laundon – BTu, BPo;
on soil.
P. HYMENINA (Ach.) Delise – BPo; on soil.
P. LEUCOPHLEBIA (Nyl.) Gyeln. – Som; on soil.
P. MALACEA (Ach.) Funk – BTu, BPo; on soil.
P. MEMBRANACEA (Ach.) Nyl. – BPo; on soil.
P. NECKERI Hepp. ex Müll. Arg. – Som, Mal, BPo;
on soil.
P. POLYDACTYLON (Neck.) Hoffm. – Som, BPo; on
soil.
*P. PREATEXTATA (Flörke ex Sommerf.) Zopf – BTu;
(Andreyev: on soil in the young spruce
forest).
OLIVACEA
P.
(Weiss) Humb. – Som, Mal, BTu,
BPo; on soil.
P. SCABROSA Th. Fr. – BPo; on soil.
*P HAEOPHYSCIA SCIASTRA (Ach.) Moberg – BTu;
(Andreyev: on granite pebbles on the road,
granite rocks and boulders on the coast).
*PHLYCTIS ARGENA (Spreng.) Flot. – BTu; (Andreyev:
on bark of Alnus sp. on the coast).
PHYSCIA ADSCENDENS H. Olivier – BTu, Mo, BPo; on
bark of deciduous trees.
P. CAESIA (Hoffm.) Fürnr. – MTu, BPo; on siliceous
pebbles and rocks.
P. DUBIA (Hoffm.) Lettau – BTu, BPo; on bark of
deciduous trees (Andreyev: on Acer platanoides, Quercus robur near the ruins of old
village, on granite rocks and boulders on the
coast, granite boulders on dunes).
*P. LEPTALEA (Ach.) DC. – BTu; (Andreyev: on bark
of Alnus sp. on the coast).
P. STELLARIS (L.) Nyl. – BTu, Ses, Mo, BPo; on bark
of deciduous trees (Andreyev: on Fraxinus
excelsior, Quercus robur near the ruins of
old village).
P. TENELLA (Scop.) DC. – Mal, BTu, Ses, Mo; on
bark of deciduous trees (Andreyev: on Quercus robur near the ruins of old village, on
granite rocks and boulders on the coast).
PLATISMATIA GLAUCA (L.) W.L. Culb. & C.F. Culb.
– Mal, BTu, Gog, BPo; on bark of deciduous
and coniferous trees, rocky outcrops
(Andreyev: on bark of Picea abies in spruce
forests, wood on dunes).
*PORPIDIA CRUSTULATA (Ach.) Hertel & Knoph – BTu;
(Andreyev: on granite boulders on dunes).
*P. FLAVOCAERULESCENS (Hornem.) Hertel & A.J.
Schwab – BTu; (Andreyev: on granite
boulders in the young spruce forest).
*PROTOPARMELIA BADIA (Hoffm.) Hafellner – BTu;
(Andreyev: on granite boulders on dunes).
PROTOPARMELIOPSIS MURALIS (Schreb.) M. Choisy –
MTu, BPo; on siliceous pebbles and wood.
PSEUDEVERNIA FURFURACEA (L.) Zopf – Mal, BTu, Gog,
BPo; on bark of deciduous and coniferous
trees (Andreyev: on Picea abies in the spruce
forest, on wood and granite rocks and boulders in the pine forest and on dunes).
RAMALINA FARINACEA (L.) Ach. – Mal, BTu, BPo; on
bark of deciduous trees (Andreyev: on Alnus
sp. on the coast).
RAMALINA FRAXINEA (L.) Ach – Mal; on bark of
deciduous trees.
*R. POLLINARIA (Westr.) Ach. – BTu; (Andreyev: on
bark of Alnus sp., Sorbus aucuparia on the
RUFESCENS
11
coast, bark of Quercus robur near the ruins
of old village).
R. POLYMORPHA (Liljeblad) Ach. – Som; on siliceous
rocks.
RHIZOCARPON DISTINCTUM Th. Fr. – Som, BTu, MTu;
on siliceous pebbles (Andreyev: on granite
pebbles on the road).
R. GEOGRAPHICUM (L.) DC. – Som, BTu; on siliceous
pebbles (Andreyev: on granite rocks on the
coast and in the pine forest).
R. GRANDE (Flörke) Arnold – MTu; on siliceous
pebble.
*R. LECANORINUM Anders – BTu; (Andreyev: on
granite rocks on the coast and in the pine
forest, granite boulders on dunes).
*R. RICHARDII (Lamy ex Nyl.) Zahlbr. – BTu; (Andreyev: on granite rocks and boulders on
the coast).
*RINODINA GENNARII Bagl. – BTu; (Andreyev: on
limestone in the ruined old village).
*R. PYRINA (Ach.) Arnold – BTu; (Andreyev: on
bark of Quercus robur near the ruins of old
village).
*R. SOPHODES (Ach.) A. Massal. – BTu; (Andreyev:
on bark of Sorbus aucuparia on the coast).
*SCHAERERIA FUSCOCINEREA (Nyl.) Clauzade & Cl.
Roux – BTu; (Andreyev: on granite rocks on
the coast and in the pine forest).
SCOLICIOSPORUM CHLOROCOCCUM (Graewe ex Stenh.)
Vězda – BTu, Gog; on bark of deciduous and
coniferous trees (Andreyev: Alnus sp., Picea
abies, Sorbus aucuparia on the coast).
*S. UMBRINUM (Ach.) Arnold – BTu; (Andreyev: on
granite rocks and boulders on the coast).
S PHAEROPHORUS FRAGILIS (L.) Pers. – BPo; on
rocks.
STEREOCAULON ALPINUM Laurer – Mal; on soil.
*S. INCRUSTATUM Flörke – BTu; (Andreyev: on sand
dunes).
*S. RIVULORUM H. Magn. – BTu; (Andreyev: on
sand dunes).
*S. SAXATILE H. Magn. – BTu; (Andreyev: on sand
dunes).
*S. TOMENTOSUM Fr. – BTu; (Andreyev: on sand
dunes).
*T EPHROMELA ATRA (Huds.) Hafellner – BTu;
(Andreyev: on granite rocks and boulders
on the coast, granite boulders on dunes).
*TREMOLECIA ATRATA (Ach.) Hertel – BTu; (Andreyev:
on granite rocks and boulders on the coast,
granite boulders on dunes).
T UCKER MANNOPSIS CHLOROPHYLLA (Willd.) Hale
– Mal, BTu, Gog, Ses, Mo, BPo; on bark of
deciduous and coniferous trees (Andreyev:
Picea abies in the spruce forest).
UMBILICARIA DEUSTA (L.) Baumg. – Som, Mal, BTu,
Gog, BPo; on siliceous rocks (Andreyev: on
granite boulders on sand dunes).
U. HIRSUTA (Sw. ex Westr.) Hoffm. – Gog, BPo; on
siliceous rocks.
*U. HYPERBOREA (Ach.) Hoffm. – BTu; (Andreyev:
on granite boulders in the young spruce
forest).
U. POLYPHYLLA (L.) Baumg. – Som, BTu; on
siliceous rocks (Andreyev: on granite boulders on dunes).
*U. TORREFACTA (Lightf.) Schrad. – BTu; (Andreyev:
on granite boulders on dunes).
USNEA HIRTA (L.) Weber ex F.H. Wigg. – Mal, Gog;
on bark of coniferous trees.
*VERRUCARIA MAURA Wahlenb. – BTu; (Andreyev: on
granite rocks and boulders on the coast).
VULPICIDA PINASTRI (Scop.) J.-E. Mattsson & M.J.
Lai – Ses, Mo, BPo; on bark of deciduous
and coniferous trees, wood.
XANTHOPARMELIA CONSPERSA (Ach.) Hale – Som,
BTu, MTu, Ses, BPo; on siliceous pebbles
and rocks (Andreyev: on granite boulders
on dunes).
X. SOMLOËNSIS (Gyeln.) Hale – BTu, BPo; on
siliceous pebbles and rocks (Andreyev: on
granite boulders on dunes).
XANTHORIA CANDELARIA (L.) Th. Fr. – BTu, BPo; on
siliceous rocks, wood (Andreyev: on granite
boulders on dunes).
X. ELEGANS (Link) Th. Fr. – BPo; on siliceous
rocks.
X. PARIETINA (L.) Th. Fr. – Som, Mal, BTu, MTu,
Ses, Mo, BPo; on siliceous pebbles and
rocks, concrete, bark of deciduous trees
(Andreyev: Fraxinus excelsior, Padus avium,
Sorbus aucuparia near the ruins of old village, on granite rocks and boulders on the
coast).
X. POLYCARPA (Hoffm.) Th. Fr. ex Rieber – Mal,
BTu, MTu, Gog, Ses, Mo, BPo; on bark of
deciduous trees, dry twigs of Picea abies,
wood (Andreyev: on bark of Alnus sp. on the
coast, Quercus robur near the ruins of old
village, wood on dunes).
ACKNOWLEDGEMENTS
The author is sincerely grateful to Teuvo Ahti
and Olga Kataeva for identification and confirmation of some specimens; Dmitry Himelbrant
12
Folia Cryptog. Estonica
and Alexei Zavarzin for identification of specimens and discussion, and Irina Shapiro for help
with TLC. Thanks are due to Natalia Balashova
and Anna Gaginskaya for collections and Elena
Glazkova for collections and valuable information about the islands.
REFERENCES
Andreyev, M. 2002. Lichens of Bolshoi Tyuters
island in Gulf of Finland, Leningrad region (in
Russian). Novitates Systematicae Plantarum non
Vascularium 36: 73–79.
Brenner, M. 1886. Bidrag till kanedom af Finska vikensovegetation. IV Hoglands lafvar. Medd. Soc.
Fauna et Flora Fennica 3: 1–144.
Glazkova, E. 2001. Vascular flora of the islands of the
eastern Gulf of Finland: structure and analysis (in
Russian). St. Petersburg. 346 pp.
Noskov, G. & Botch, M. (eds) 1999. Red Data Book of
Nature of the Leningrad region. Vol. 1. – Protected
Areas. St. Petersburg. 352 pp.
Santesson, R., Moberg, R., Nordin, A., Tønsberg,
T. & Vitikainen, O. 2004. Lichen-forming and
lichenicolous fungi of Fennoscandia. Uppsala.
359 pp.
Tzvelev, N. (ed.) 2000. Red Data Book of Nature of
the Leningrad region. Vol 2. Plants and fungi. St.
Petersburg. 627 pp.
Folia Cryptog. Estonica, Fasc. 41: 13–22 (2005)
New or interesting lichens and lichenicolous fungi found during
the 5th IAL Symposium in Estonia
Andre Aptroot1, Paweł Czarnota2, Inga Jüriado3, Jana Kocourková4, Martin
Kukwa5, Piret Lõhmus3, Zděnek Palice6, Tiina Randlane3, Lauri Saag3, Emmanuel
Sérusiaux7, Harrie Sipman8, Laurens B. Sparrius9, Ave Suija3 & Holger Thüs10
ABL Herbarium, Gerrit van der Veenstraat 107, NL-3762 XK, Soest, the Netherlands. E-mail: andreaptroot@wanadoo.nl
2
Scientific Laboratory of the Gorce National Park, Poręba Wielka 590 PL-34-735 Niedźwiedź, Poland.
E-mail: pawel.czarnota@gpn.pl
3
Institute of Botany and Ecology, University of Tartu, Lai 38, Tartu, Estonia. E-mail: ave.suija@ut.ee
4
Mycological Department of National Museum Prague, Václavské náměstí 68, CZ-115 79 Praha 1, Czech Republic.
E-mail: jana.kocourkova@nm.cz
5
Department of Plant Taxonomy and Nature Protection, University of Gdańsk, Al. Legionów 9, PL-80-441 Gdańsk,
Poland. E-mail: dokmak@univ.gda.pl
6
Botanical Institute of the Czech Academy of Sciences, CZ-252 43 Průhonice, Czech Republic. E-mail: palice@ibot.cas.cz
7
Plant Taxonomy and Conservation Biology, University of Liège. Sart Tilman B22, B4000, Liège, Belgium.
E-mail: E.Serusiaux@ulg.ac.be
8
Botanic Garden and Botanical Museum Berlin-Dahlem, Freie Universität Berlin, Königin-Luise-Strasse 6-8, D-14191
Berlin, Germany. E-mail: h.sipman@bgbm.org
9
BIO.DIV, Kongsbergstraat 1, NL-2804 XV, Gouda, the Netherlands E-mail: sparrius@biodiv.nl
10
General Biology, University of Kaiserslautern, Erwin-Schrödinger-Strasse 13, A67663 Kaiserslautern, Germany.
E-mail: hthues@senckenberg.de
1
Abstract: Altogether 74 species of lichens and lichenicolous fungi found during IAL5 (Estonia, August 2004) are listed; 30
of them are new to the country. New lichens: Absconditella celata, A. delutula, A. sphagnorum, Bacidia adastra, B. caligans, Bacidina
chloroticula, Cladonia monomorpha, Dirina massiliensis f. sorediata, Lecanora subcarpinea, Lecidella flavosorediata, Micarea anterior, M. lutulata,
M. micrococca, Ramonia aff. nigra, Rinodina degeliana, Thelocarpon lichenicola, Trapelia involuta, Verrucaria bryoctona, V. dolosa. New
lichenicolous fungi: Arthonia digitatae, A. galactinaria, Cercidospora macrospora, Cornutispora lichenicola, Marchandiomyces aurantiacus,
Merismatium heterophractum, Phoma epiphyscia, Pronectria xanthoriae, Telogalla olivieri, Trichonectria anisospora, T. hirta.
Kokkuvõte: IAL5 sümpoosioni ajal Eestist leitud uued ning huvipakkuvad samblikud ja lihhenikoolsed
seened.
Rahvusvahelise Lihhenoloogide Assotsiatsiooni 5nda sümpoosioni ajal 2004. a-l leiti Eestist 74 haruldast või seni leidmata
taksonit. Eestile uued samblikuliigid on (19) on: Absconditella celata, A. delutula, A. sphagnorum, Bacidia adastra, B. caligans, Bacidina chloroticula, Cladonia monomorpha, Dirina massiliensis f. sorediata, Lecanora subcarpinea, Lecidella flavosorediata, Micarea anterior,
M. lutulata, M. micrococca, Ramonia aff. nigra, Rinodina degeliana, Thelocarpon lichenicola, Trapelia involuta, Verrucaria bryoctona, V.
dolosa. Eestile uued lihhenikoolsed seened (11) on: Arthonia digitatae, A. galactinaria, Cercidospora macrospora, Cornutispora lichenicola, Marchandiomyces aurantiacus, Merismatium heterophractum, Phoma epiphyscia, Pronectria xanthoriae, Telogalla olivieri, Trichonectria
anisospora, T. hirta.
INTRODUCTION
Lichenological field trips and symposia which
have been arranged in Estonia and attended
by colleagues from other countries have always
been very generous in new floristic data. In 1989,
several Swedish lichenologists visited the western part of the country and presented then 317
taxa, 75 of them new to Estonia (Ekman et al.,
1991). Two years later, our Swedish colleagues
Göran Thor and Anders Nordin investigated
wooded meadows in West-Estonia and some
localities in the vicinity of Tartu and identified
16 new species (Thor & Nordin, 1998). The XIV
Symposium of Baltic Mycologists and Lichenologists which took place in the south-eastern part
of Estonia in 1999 was not an exception either:
43 rare lichen and lichenicolous fungus species,
17 of which were previously unknown from the
country, were reported (Halonen et al., 2000).
14
Folia Cryptog. Estonica
The present paper contains notes on 74 lichenized and lichenicolous fungi found during
the 5th IAL symposium “Lichens in Focus”. As
a well-entrenched tradition, the number of new
species was high again – 30. Most of the specimens were collected during three excursions to
different parts of Estonia; however, some interesting taxa were also found in Tartu where the
symposium took place.
The longest, five days lasting pre-symposium
excursion covered various parts of the mainland
and western islands of Estonia. 23 participants
visited natural forests in Lahemaa National Park
and clint forest in northern Estonia, broadleaved deciduous forest and wooded meadow
in western Estonia, and a raised bog in Soomaa
National Park in south-western part. Thereafter the trip continued in the islands Muhumaa,
Saaremaa (Fig. 1) and Hiiumaa. Different forest
types, alvars, oak wood and sandy dunes were
visited. Beside these, some places with cultural
implications e.g. Kaali meteorite crater and the
medieval castle in Kuressaare (both in Saaremaa), Kuremäe orthodox nunnery and Palmse
estate (both on the mainland) were included into
the excursion program as well.
The most popular trip (30 participants),
two days lasting post-symposium excursion
took place in the eastern and northern part of
Estonia. The tour consisted of a trip to the raised
bog in Endla Nature Reserve, visits to a forest
stand with old aspens and to a wooded meadow
(both near Rakvere), and a stop in a karst field in
Kostivere (Fig. 2). A short visit to Sagadi manor
house and Museum of Forestry in Lahemaa
National Park was also arranged.
19 participants of the one-day post-symposium excursion were taken to South-Estonia.
The longer lichenological stops were arranged on
the high sandstone banks of the primeval valley
of Ahja River, on a raised bog and on the sandy
dunes in Värska close to the border of Russia.
In addition, a visit to Setu Farm Museum was
a tourist event of the day.
Fig. 1. Participants of IAL5 pre-symposium excursion on Atla alvar, island Saaremaa (photo
I. Jüriado).
15
Fig. 2. Karst field in Kostivere, northern part of Estonia (photo I. Jüriado).
Localities
(see also Fig. 3)
1. Lääne-Virumaa, Oandu forest trail
(59°33’45’’N 26°00’39’’E). Spruce forest on
peat soil with scattered deciduous trees. 11
Aug 2004.
2. Ida-Virumaa, coast near Ontika (59°26’39’’N
27°19’22’’E). Deciduous, broad-leaved clint
forest. 12 Aug 2004.
3. Ida-Virumaa, Kuremäe cloister (59°11’56’’N
27°32’03’’E). Trees along footways. 12 Aug
2004.
4. Pärnumaa, Soomaa National Park, Riisa
bog (58°29’23’’N 24°59’33’’E). Raised bog
and spruce forest. 13 Aug 2004.
5. Pärnumaa, Koonga community, Nedrema
wooded meadow (58°32’45’’N 24°04’23’’E).
Woodland with scattered deciduous trees.
13 Aug 2004.
6. Saaremaa, Muhu Island, Tupenurme cliffs
(58°38’45’’N 23°13’10’’E). Inland limestone
cliffs, ca 2 m high. 13 Aug 2004.
7. Saaremaa, Vilsandi National Park, Atla alvar
(58°17’06”N 21°54’46”E). Limestone fields.
14 Aug 2004.
8. Saaremaa, Sõrve Peninsula, Lõo alvar
(58°06’09’’N 22°10’50’’E). Coastal limestone
fields. 14 Aug 2004.
9. Saaremaa, Kaali Meteorite Crater
(58°27’21’’N 22°39’48’’E). Park around
crater lake. 14 Aug 2004.
10. Hiiumaa, Kõpu peninsula, Rebastemäe
Nature Track (58°55’22’’N 22°15’02’’E).
Spruce and pine forest on steep sand dunes,
alt. ca 70 m. 15 Aug 2004.
11. Hiiumaa, Pihla-Kaibaldi Nature Reserve
(58°58’24’’N 22°39’51’’E). Kaibaldi heath
pine forest and inland dunes. 15 Aug
2004.
12. Tartumaa, nature trail N of the city along
Emajõgi river (58°25’20’’N 26°40’32’’E).
Riverine deciduous and coniferous forest.
19 Aug 2004.
13. Tartu, parks in city center (58°23’N 26°43’E).
22 Aug 2004.
14. Tartu, Aleksandri Street (59°22’45’’N
26°43’49’’E). Deciduous trees along the
street. 20 Aug 2004.
15. Jõgevamaa, Tooma village near the border
of Endla Nature Reserve (58°52’19’’N
16
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
Folia Cryptog. Estonica
26°16’16’’E). Trees along footpath. 22 Aug
2004.
Jõgevamaa, Endla Nature Reserve,
Männikjärve bog trail (58°52’21’’N
26°14’56’’E). Raised bog and spruce forest.
22 Aug 2004.
Lääne-Virumaa, around Emumäe tower, alt.
166 m (58°56’17’’N 26°22’23’’E). Coniferous
forest. 22 Aug 2004.
Ida-Virumaa, Oonurme village, Kautvere
(key habitat site) (59°10’28’’N 26°57’36’’E).
Coniferous forest with scattered aspen trees.
22 Aug 2004.
Lääne-Virumaa, Rakvere town (59°21’03’’N
26°21’11’’E). Deciduous trees. 23 Aug
2004.
Lääne-Virumaa, Mädapea wooded meadow
(59°19’31’’N 26°15’60’’E). Woodland with
scattered deciduous trees. 23 Aug 2004.
Lääne-Virumaa, Lahemaa National Park,
Sagadi manor (59°32’11’’N 26°05’11’’E).
Trees in the vicinity of Museum of Forestry.
23 Aug 2004.
Harjumaa, Kostivere karst field (59°26’07’’N
25°06’45’’E). 23 Aug 2004.
Põlvamaa, N of Põlva, Taevaskoja along Ahja
river (58°06’34’’N 27°03’06’’E). Forest dominated by Pinus sylvestris and Picea abies.
22 Aug 2004.
Põlvamaa, SE of Värska on Russian border,
Mustoja kame field (57°53’40’’N 27°42’40’’E).
Sand dunes with sparse pine forest and
small open sand patches. 22 Aug 2004.
Põlvamaa, Meenikunno raised bog
(57°56’23’’N 27°19’40’’E). 22 Aug 2004.
26. Põlvamaa, Värska, yard of Setu Farm Museum (57°56’34’’N 27°38’57’’E). 22 Aug
2004.
LIST OF SPECIES
New species to Estonia are typed in bold; all
changes in frequency classes (Randlane & Saag,
1999) are indicated with → and are also typed
in bold.
Abbreviations and symbols: # = lichenicolous
fungus; + non-lichenized species. Frequency
classes: rr = very rare (1–2 localities); r = rare
(3–5); st r = rather rare (6–10). Collectors: AA =
Andre Aproot; AS = Ave Suija; ES = Emmanuel
Sérusiaux, HS = Harrie Sipman; HT = Holger
Thüs; IJ = Inga Jüriado; JK = Jana Kocourková;
LS = Laurens Sparrius; LSaag = Lauri Saag;
MK = Martin Kukwa; PC = Paweł Czarnota, ZP
= Zděnek Palice, TR = Tiina Randlane. Herbaria: ABL = Adviesbureau voor Bryologie en
Lichenologie, the Netherlands; B = Botanischer
Garten und Botanisches Museum Berlin-Dahlem, Zentraleinrichtung der Freien Universität
Berlin, Germany; GPN = Gorce National Park,
Poland; Herb. Palice = private herbarium of
Zděnek Palice, Czech Republic; Herb. Sparrius
= private herbarium of Laurens Sparrius, the
Netherlands; Herb. Thüs = private herbarium of
Holger Thüs, Germany; LG = Université de Liège,
Belgium; PRM = Mycological Department of National Museum, Prague, Czech Republic; TU =
Fig. 3. Location of the
sampling points:
– pre-symposium excursion;
– post-symposium excursion lasting two
days
– post-symposium excursion of one day;
– Tartu, during symposium.
17
University of Tartu, Estonia; UGDA-L = lichen
herbarium of Gdansk University, Poland.
ABSCONDITELLA CELATA Döbbeler & Poelt – 23:
on lying decaying trunks of Picea abies and
on decaying wood of bridge over brook, JK
(PRM); 25: on wood, cutting flat of stump,
ZP (Herb. Palice). Freq.: rr.
ABSCONDITELLA DELUTULA (Nyl.) Coppins & H. Kilias – 16: on wood, MK (UGDA-L). Freq.: rr.
ABSCONDITELLA LIGNICOLA Vězda & Pišút – 23: on
decaying wood of bridge over brook; in
spruce forest, on lying decaying trunk of
Picea abies, JK (PRM). Freq.: rr. – This is
the second locality, earlier record from the
same region (Halonen et al., 2000).
ABSCONDITELLA SPHAGNORUM Vězda & Poelt – 4:
on peat, LS (Herb. Sparrius), TR (TU); 25:
on Sphagnum cf. magellanicum, JK (PRM).
Freq.: rr. – This species is more or less restricted to patches of dead Sphagnum in
raised bogs in the Northern Hemisphere.
It probably lives partly as saprob. The species is visible as minute (0.1–0.3 mm diam.)
flesh-coloured, marginate apothecia on dry,
decaying Sphagnum which then has more or
less the same colour.
AGONIMIA GLOBULIFERA Brand & Diederich – 7: on
calcareous soil with Fulgensia bracteata,
LS (Herb. Sparrius, TU). Freq.: rr. – This
is the second record from Estonia. Steffen
Boch (Germany) found Agonimia globulifera
two months earlier from another alvar in
Saaremaa rather close to this locality (Nõmm
et al., 2005).
ANISOMERIDIUM BIFORME (Borrer & Sowerb.) R.C.
Harris – 20: on Quercus robur, LS (Herb.
Sparrius). Freq.: rr. – This is the second
locality, so far was recorded in southwestern region of Estonia (Randlane &
Saag, 2004).
ANISOMERIDIUM POLYPORI (Ellis & Everh.) M.E. Barr
– 12: on Quercus robur, AA (ABL); 13: on
Salix sp., AA (ABL); 16: on Ulmus sp., ES
(LG). Freq.: rr → r. – Known until now from
two localities in western islands and northwestern region of Estonia (Randlane & Saag,
2004), probably overlooked.
ARTHONIA MUSCIGENA Th. Fr. – 18: on Populus
tremula, LS (Herb. Sparrius). Freq.: rr.
– Earlier was known from only one locality in
western islands (Randlane & Saag, 2004).
# ARTHONIA DIGITATAE Hafellner – 23: on Cladonia
digitata on bark of Pinus sylvestris, JK (PRM).
Freq.: rr. – The species seems to be known
until now only from Austria (Hafellner 1999),
Luxemburg (Sérusiaux et al. 2003) and
the Czech Republic (Kocourková & Boom,
2005).
# ARTHONIA GALACTINARIA Leight. – 22: on Lecanora
dispersa agg. on concrete, MK (UGDA-L).
Freq.: rr.
# ARTHRORHAPHIS AERUGINOSA R. Sant. & Tønsberg
– 23: on squamules of Cladonia polydactyla
and C. digitata growing at base of Picea
abies, JK (PRM); 24: on C. phyllophora,
JK (PRM). Freq.: rr. – The fungus is easily
recognized by the characteristic deep bluegreen pigmentation of basal squamules of
the host species (Santesson & Tønsberg,
1994).
BACIDIA ADASTRA Sparrius & Aptroot – 13: on
Malus sp., AA (ABL, TU). Freq.: rr. – This
species was found on a few fruit trees in
the urban environment. It is only recently
described taxon (Sparrius & Aptroot, 2003)
which may have been overlooked as it is often
sterile. The samples collected in Estonia are
unfortunately sterile. The species should
belong to the genus Bacidina mainly by the
characteristics of the fruit-bodies.
BACIDIA BIATORINA (Körb.) Vain. – 18: on Populus
tremula, LS (Herb. Sparrius), HS (B). Freq.: r.
– Previously reported from three localities in
Pärnumaa, south-western region of Estonia
(Randlane & Saag, 2004).
BACIDIA CALIGANS (Nyl.) A.L. Sm. – 13: botanical
garden, on Pyrus sp., AA (ABL, TU). Freq.:
rr. – This species should belong to the genus
Bacidina (Ekman, 1996 : 125; Santesson et
al., 2004), however, no valid combination
has been made.
BACIDIA INCOMPTA (Borrer ex Hook.) Anzi – 23:
on rotten wood of bridge over brook, JK
(PRM). Freq.: r. – Previously known from
three scattered localities (Randlane & Saag,
2004).
BACIDINA CHLOROTICULA (Nyl.) Vězda & Poelt – 12:
on Betula sp. and Quercus robur, AA (ABL),
LS (Herb. Sparrius, TU); 13: on Quercus
robur, LS (Herb. Sparrius); 25: on wood,
cutting flat of stump, ZP (Herb. Palice).
Freq.: r. – B. chloroticula is mainly found
in disturbed, artificial habitats on both
acid rock, wood and bark. All collected
18
Folia Cryptog. Estonica
specimens have the characteristic grey to
dark green, areolate to coarsely granular
thallus; however, specimens collected on
bark (loc. 12 & 13) deviate from typical
B. chloroticula by the rather dark, reddish
brown pigmentation of the excipulum. The
specimen collected on wood (loc. 25) has
distinctive colourless excipulum.
BIATORA CHRYSANTHA (Zahlbr.) Printzen – 18: on
Populus tremula, MK (UGDA-L), PC (GPN).
Freq.: rr → r. – Until now only two localities
were known from north- and south-western regions of Estonia (Randlane & Saag,
2004).
BUELLIA SCHAERERI De Not. – 20: on Quercus robur,
LS (Herb. Sparrius). Freq.: r. – Previously
known from four scattered localities (Randlane & Saag 2004).
CALICIUM PINASTRI Tibell – 24: on peeling bark of
young Pinus sylvestris, ZP (Herb. Palice).
Freq.: rr → r. – Earlier only two localities
were known in south-eastern and -western
regions (Randlane & Saag, 2004).
CALOPLACA OBSCURELLA (Körb.) Th. Fr. – 13: on
Ulmus sp., LS (Herb. Sparrius). Freq.: rr.
– Only one record from western islands was
known before (Randlane & Saag, 2004).
# CERCIDOSPORA MACROSPORA (Uloth) Hafellner
& Nav.-Ros. – 22: on Lecanora muralis on
granite stone, AS (TU), MK (UGDA-L). Freq.:
rr. – C. macrospora was lately found also
from Kadakalaid islet near Hiiumaa (Jüriado
& Suija, unpubl.)
CLADONIA MONOMORPHA Aptroot, Sipman & Herk
– 24: on acid soil, AA (ABL), IJ (TU), ZP (Herb.
Palice). Freq.: rr. – A recently described species (Aptroot et al., 2001) which has been
confused with Cladonia pyxidata. The latter
species is generally more southern growing
predominantly on boulders, not on open
sandy soils. C. monomorpha is morphologically distinct by large bullate plates on the
cups, both on the inside and outside.
CLADONIA POLYDACTYLA (Flörke) Spreng. – 16: on
Betula sp., AS (TU); 23: on Pinus sylvestris,
AA & IJ (TU) and on rotten stump, JK (PRM,
sub Arthrorhaphis aeruginosa); 25: on the
shore of a lake, on wood of Pinus sylvestris,
JK (PRM). Freq.: r → st r. – C. polydactyla
has earlier been found from three scattered
localities in Estonia (Randlane & Saag,
1999). However, the species might be overlooked and confused with C. digitata and C.
coniocraea which can grow in rather similar
habitats.
# CLYPEOCOCCUM HYPOCENOMYCES D. Hawksw. – 20:
on Hypocenomyce scalaris on wood, MK
(UGDA-L). Freq.: r. – Previously reported
from north-eastern and south-eastern regions of Estonia (Randlane & Saag, 2004).
# CORNUTISPORA LICHENICOLA D. Hawksw. & B. Sutton – 23: on Cladonia digitata at the base of
Pinus sylvestris, JK (PRM) Freq.: rr.
CYPHELIUM TIGILLARE (Ach.) Ach. – 16: on wooden
footpath in Männikjärve bog, AS (TU). Freq.:
r → st r. – Five scattered localities were
known before (Randlane & Saag, 2004).
DIRINA MASSILIENSIS Durieu & Mont. f. SOREDIATA
(Müll. Arg.) Tehler – 6: on soft limestone,
LS (Herb. Sparrius), TR (TU). Freq.: rr. – The
genus Dirina has a predominantly Mediterranean distribution in Europe while the
sorediate form of D. massiliensis occurs
also in northern Europe. It favours shaded
overhangs and vertical walls of basic (natural and artificial) rock and also occurs on
acid rock near the coast with direct influence of salt spray making the substratum
pH slightly higher (Tehler, 1983). The record
in Estonia is the north-easternmost record
ever. It occurs at slightly higher latitudes
only in Norway (Botnen & Tønsberg 1988)
and in Scotland, and is distributed more
eastern only in Bulgaria, Yemen (Socotra)
and along the Mediterranean coast (Tehler,
1983).
GYALECTA TRUNCIGENA (Ach.) Hepp. – 18: on Populus
tremula, ES (LG), PC (GPN). Freq.: r. – Earlier
known from four scattered localities (Randlane & Saag, 2004).
# ILLOSPORIOPSIS CHRISTIANSENII (B. L. Brady & D.
Hawksw.) D. Hawksw. – 13: on Physcia tenella on Quercus robur, LS (Herb. Sparrius);
15: on P. cf. stellaris on Malus domestica, MK
(UGDA-L); 22: on P. tenella on hard wood,
LS (Herb. Sparrius), on P. dubia over granitic
boulder in meadow, PC (GPN) and on P. cf.
stellaris on Malus domestica, MK (UGDA-L).
Freq.: r → st r. – Until now, the species was
known only from three localities in Estonia
(Randlane & Saag, 2004).
LECANORA SUBCARPINEA Szatala – 5: on Quercus
robur, LS (Herb. Sparrius); 13: on Populus
tremula AA (ABL), LS (Herb. Sparrius); 20:
on Quercus robur, LS (Herb. Sparrius). Freq.:
r. – This species is widespread in central
19
and southern Europe (e.g. Lumbsch et al.,
1997; Wirth, 1995; van Herk & Aptroot,
2004), however, not recorded from northern
Europe. It is similar to Lecanora carpinea,
but has fewer and smaller apothecia and
a P+ yellow disc margin additional to the
C+ orange reaction of the disc pruina. The
thallus is bright white and therefore rather
conspicuous. It occurs mostly on acid and
neutral bark of well-lit trees.
LECIDEA HYPOPTA Ach. – 23: on Pinus sylvestris, AA
(ABL). Freq.: r. – This is the fourth record
from Estonia, before was known from western islands only (Randlane & Saag, 2004).
L ECIDELLA FLAVOSOREDIATA (Vězda) Hertel &
Leuckert – 3: on Quercus robur, LS (Herb.
Sparrius, TU); 12: on Populus tremula AA
(ABL); 13: on Quercus robur, LS, on P.
tremula, AA (ABL); 20: on Quercus robur,
LS (Herb. Sparrius) and on Betula sp., MK
(UGDA-L). Freq.: r. – This sorediate and
usually sterile species is rather common
in northern Europe on well-lit, acid bark,
but much overlooked. It was found on
wayside oak trees in parkland, in a community with predominantly crustose species: Haematomma ochroleucum, Lecanora
expallens, Melanelia glabratula, Pertusaria
coccodes, Phlyctis argena and Pyrrhospora
quernea. Lecidella flavosorediata is similar
in appearance to Pyrrhospora quernea, but
has a darker green, fine soredia (+/- isidiate
in Pyrrhospora) on a grey areolate thallus
(yellowish or absent in Pyrrhospora).
LECIDELLA SUBVIRIDIS Tønsberg – 16: on Picea abies,
MK (UGDA-L). Freq.: rr → r. – Earlier two
records were from western islands and
south-eastern region of Estonia (Randlane
& Saag, 2004).
LEPRARIA ATLANTICA Orange – 23: on sandstone
rock, LSaag (TU). Freq.: rr. – This is the
second record from Estonia (Jüriado et al.,
2002). TLC: atranorin & porphyrilic acid.
LEPTOGIUM GELATINOSUM (With.) J.R. Laundon – 8:
on calciferous soil, LS (Herb. Sparrius).
Freq.: r. – The species occurs in western
islands and north-eastern region of the
mainland (Randlane & Saag, 1999).
LEPTOGIUM PLICATILE (Ach.) Leight. – 22: on limestone, HS (B). Freq.: rr. – L. plicatile was considered extinct in Estonia, the only known
record is from the end of 19th century in
the north-eastern region of the mainland
(Randlane & Trass, 1994).
LOPADIUM DISCIFORME (Flot.) Kullh. – 18: on trunk
of old Populus tremula, ES (LG). Freq.: r.
– This is the fourth locality of this species
in Estonia (Randlane & Saag, 2004).
# MARCHANDIOMYCES AURANTIACUS (Lasch) Diederich – 15: on Physcia tenella on Malus domestica, MK (UGDA-L); 19: on Physcia sp.
on Acer platanoides, MK (UGDA-L); 20: on
P. tenella on Quercus robur, MK (UGDA-L).
Freq.: r. – This species was recently found
also from Kadakalaid islet near Hiiumaa
(Jüriado & Suija, unpubl.).
MELANELIA ELEGANTULA (Zahlbr.) Essl. – 20: on
Quercus robur, LS (Herb. Sparrius). Freq.:
rr. – This species was recorded as doubtful
in Estonia, known only according to the literature data (Mereschkowsky, 1913).
MELANELIA SEPTENTRIONALIS (Lynge) Essl. – 12: on
Betula sp., AA (ABL) Freq.: r → st r. – This
is the sixth record of the species, previously
has been reported also from Tartu (Trass &
Randlane, 1994).
# MERISMATIUM HETEROPHRACTUM (Nyl.) Vouaux
– 23: on Lecanora sp. on twigs of Pinus sylvestris, JK (PRM). Freq.: rr.
MICAREA ANTERIOR (Nyl.) Hedl. – 20: on rotting wood of a fallen deciduous tree, PC
(GPN). Freq.: rr. – A rare lignicolous species
reported only from a few European countries:
Sweden and Finland (Coppins, 1983), Komi
Republic in Russia (Hermansson et al.,
1998), Germany (Wirth, 1995) and Czech
Republic (Palice, 1999) and recently from
Poland (Czarnota, in press). M. anterior is
inconspicuous, epixylic lichen forming in
the most cases only stipitate pycnidia and
probably for this reason it is overlooked
during field work or mistakenly identified
as a non-lichenized fungus. In many
respects darker specimens of M. anterior
are very similar to M. misella (Nyl.) Hedl.,
and moreover they sometimes grow together.
In the Estonian case the two species are
growing also together, but M. misella forms
only stipitate pycnidia, whose walls react
K+ violet. M. anterior produces a lot of small
pale brownish apothecia. The brown tinge
in epithecium and narrow excipulum is still
present after KOH and moreover the species
has also small 2–(3)septate ascospores.
This species may be difficult to recognize
20
Folia Cryptog. Estonica
due to a few other lignicolous species and
unidentified facultative lichenicolous fungus
growing together with it.
MICAREA LUTULATA (Nyl.) Coppins – 11: on shingle
on sandy plane, AS & IJ (TU). Freq.: rr.
– M. lutulata was recently found also from
Vohilaid islet near Hiiumaa (Jüriado & Suija,
unpubl.).
MICAREA MICROCOCCA (Körb.) Gams ex Coppins
– 1: on rotten wood, LS (Herb. Sparrius); 10:
on Pinus sylvestris, LS (Herb. Sparrius); 16:
on P. sylvestris and Picea abies, PC (GPN);
18: on Populus tremula, PC (GPN). Freq.:
r. – This species has been included in M.
prasina s. lato a few times, but recently it
was distinguished from M. prasina s. str. by
the presence of methoxymicareic acid while
M. prasina produces micareic acid. There
are also some ecological differences between
the species: M. micrococca mainly grows on
bark of many tree species, in forests and also
small woods, even near industrial regions
and seems to be more acidophilic than M.
prasina which occurs in most cases on soft
wood of decaying stumps within larger,
shaded woodlands. The morphological
variability of M. micrococca seems to be quite
large. Typical specimens have whitish or
cream-coloured apothecia but some others
have dark gray or even blackish fruit-bodies
and could be similar to M. melanobola. For
this reason the two species and also the
whole M. prasina group seem to need more
taxonomical study based on molecular
analyses. The species is surely widespread
in Europe, but mostly treated as synonym
of M. prasina.
MICAREA NITSCHKEANA (J. Lahm ex Rabenh.) Harm.
– 16: on wood of fallen pine log within pine
bog forest, PC (GPN); 24: peeling bark of
young Pinus sylvestris, ZP (Herb. Palice)
Freq.: st r. – The species was not recorded
in the southern part of Estonia until now
(Randlane & Saag, 2004).
PELTIGERA SCABROSA Th. Fr. – 24: on acid soil, AA
(ABL). Freq.: rr. – This is the second locality
of the species in Estonia, previously known
only from the island Aegna on the northern
coast of the mainland (Trass & Randlane,
1994).
# P HAEOPYXIS PUNCTUM (A. Massal.) Rambold,
Triebel & Coppins – 18: on Cladonia ochrochlora on Populus tremula, MK (UGDA-L).
Freq.: r. – This lichen parasite, restricted
to Cladonia species, was previously known
from three scattered localities (Randlane &
Saag, 2004; Suija, unpubl.).
# PHAEOSPOROBOLUS USNEAE D. Hawksw. & Hafellner – 23: on Pseudevernia furfuracea on
Pinus sylvestris, JK (PRM); 25: on shore
of a lake, on Usnea filipendula on bark of
P. sylvestris, JK (PRM); on P. furfuracea on
Betula pubescens, JK (PRM). Freq.: rr → r.
– Although P. usneae is a widespread species
(Hawksworth & Hafellner, 1986), the taxon
was recorded from Estonia only once before
(Randlane & Saag, 2004).
# PHOMA EPIPHYSCIA Vouaux – 15: on Xanthoria
parietina on Malus domestica, MK (UGDAL). Freq.: rr.
# PRONECTRIA XANTHORIAE Lowen & Diederich
– 15: on Xanthoria parietina on Malus domestica, MK (UGDA-L). Freq.: rr.
RAMONIA AFF. NIGRA Coppins – 18: on trunk of old
Populus tremula, ES (LG). Freq.: rr. – The
material is scanty but definitely a Ramonia
with dark brown apothecia and muriform
ascospores; still, these are much too small
to be the spores of R. nigra.
RINODINA DEGELIANA Coppins – 20: mixed with
Scoliciosporum chlorococcum on Betula sp.,
MK (UGDA-L). Freq.: rr. – TLC: atranorin and
zeorin were found as major substances.
+ SAREA DIFFORMIS (Fr. ) Fr.– 23: on resin of
Picea abies, JK (PRM). Freq.: r. – This is
the fourth locality in Estonia (Randlane &
Saag, 2004).
SCOLICIOSPORUM SAROTHAMNI (Vain.) Vězda – 13:
on Quercus robur, AA (ABL); 16: on twigs
of Picea abies, PC (GPN); 20: on branches
of Quercus robur, MK (UGDA-L); 23: on
P. abies, AA (ABL). Freq.: r → st r. – The
species was known from three localities
before (Randlane & Saag, 2004); probably
overlooked.
STAUROTHELE FRUSTULENTA Vain. – 22: on limestone,
HT (Herb. Thüs). Freq.: rr → r. – This is
the third locality of the species in Estonia
(Randlane & Saag, 2004).
STEINIA GEOPHANA (Nyl.) Stein – 2: on clay, LS
(Herb. Sparrius); 25: on wood, cutting flat
of stump, ZP (Herb. Palice). Freq.: rr → r.
– This ephemeral lichen species has been
reported only once before in Estonia (Randlane & Saag, 2004).
21
# STIGMIDIUM XANTHOPARMELIARUM Hafellner – 26:
on Xanthoparmelia conspersa on granite
boulder, JK (PRM) Freq.: rr. – This is the
second record of the species (Randlane &
Saag, 2004).
# SYZYGOSPORA PHYSCIACEARUM Diederich – 17: on
Physcia tenella on Sorbus aucuparia, PC
(GPN); 20: on Physcia sp. on Quercus robur,
MK (UGDA-L); 22: on P. stellaris on Malus
domestica, MK (UGDA-L). Freq.: rr → r.
– This fungus was previously reported from
two localities (Randlane & Saag, 2004).
# TAENIOLELLA BESCHIANA Diederich – 23: on Cladonia polydactyla on rotten stump, JK (PRM);
25: on squamules of C. polydactyla on wood
of Pinus sylvestris at bank of lake, JK (PRM).
Freq.: rr → r. – The species was until now
known from three scattered localities (Randlane & Saag, 2004; Suija, unpubl.).
# TELOGALLA OLIVIERI (Vouaux) Nik. Hoffmann
& Hafellner – 15: on thallus of Xanthoria
parietina on twigs of Malus domestica, PC
(GPN), MK (UGDA-L); 21: on X. parietina on
Acer platanoides, MK (UGDA-L). Freq.: rr.
THELIDIUM DECIPIENS (Nyl.) Kremp. – 9: on soft
limestone, LS (Herb. Sparrius). Freq.: r.
– This perithecioid lichen has been previously known from three localities in Estonia
(Randlane & Saag, 2004), probably it has
been overlooked.
+ THELOCARPON LICHENICOLA (Fuckel) Poelt &
Hafellner – 23: on lying decaying trunks of
Picea abies, JK (PRM). Freq.: rr.
TRAPELIA INVOLUTA (Taylor) Hertel – 11: on limestone shingle, AS (TU). Freq.: rr.
T RAPELIA PLACODIOIDES Coppins & P. James – 8: on
humus rich soil with Candelariella vitellina,
LS (Herb. Sparrius); 24: on granite boulder
in pine forest, AA (ABL), IJ & LSaag (TU);
on granite boulder in pine forest, JK (PRM).
Freq.: rr → r. – The species was known from
two scattered localities before (Randlane &
Saag, 2004; Suija, unpubl.), probably has
been overlooked.
T RAPELIOPSIS GLAUCOLEPIDEA (Nyl.) Gotth. Schneid. – 24: on peat/humus among roots of
eradicated tree, ZP (Herb. Palice). Freq.: rr.
– Czarnota and Kukwa (2004) mentioned
the occurrence of this species in Estonia,
Tartumaa.
# T REMELLA LICHENICOLA Diederich – 23: on Mycoblastus fucatus on Betula sp., AA (ABL).
Freq.: rr → r. – The species was known until
now from two localities (Randlane & Saag
2004).
# TREMELLA PHAEOPHYSCIAE Diederich & M.S. Christ.
– 14: on Phaeophyscia orbicularis on Acer
platanoides, PC (GPN); 21: on P. orbicularis on A. platanoides, MK (UGDA-L); 22:
on P. orbicularis on Malus domestica, MK
(UGDA-L). Freq.: rr → r. – Until now, only
one locality (in Tartu) was known (Randlane
& Saag, 2004).
# TREMELLA RAMALINAE Diederich – 20: on Ramalina
fraxinea on Betula sp., MK (UGDA-L). Freq.:
rr. – This is the second record from Estonia
(Halonen et al., 2000).
# TRICHONECTRIA ANISOSPORA (Lowen) van den
Boom & Diederich – 23: on Hypogymnia
physodes and on H. tubulosa on branches of
Picea abies and Pinus sylvestris, JK (PRM).
Freq.: rr. – Until now the species has been
known only on H. physodes; H. tubulosa represents a new host species for this fungus.
# TRICHONECTRIA HIRTA (Bloxam) Petch. – 24: on
Cladonia phyllophora, Placynthiella icmalea
and on other dead lichens on plant debris
and twigs lying on ground, JK (PRM). Freq.:
rr.
VERRUCARIA BRYOCTONA (Th. Fr.) Orange – 22: on
calciferous soil, LS (Herb. Sparrius, TU).
Freq.: rr.
VERRUCARIA DOLOSA Hepp – 16: on asbestos pipe
under a bridge over a little creek in forest, HT
(Herb. Thüs); 17: at the +/- shaded, bottom
parts of granitic boulders, HT (Herb. Thüs).
Freq.: rr. – This species is easily recognized
by its superficial perithecia, a thin thallus
and rather small elongated ascospores
(16–18 × 4–7 µm). Its outer appearance is
quite similar to some morphs of V. muralis
but the almost “sessile” perithecia and its
microscopical features make V. dolosa one
of the few Verrucaria species that are rather
easy to identify.
VERRUCARIA VIRIDULA (Schrad.) Ach. – 9: on soft
limestone, LS (Herb. Sparrius). Freq.: rr →
r. – This is the third record of this species in
Estonia (Randlane & Saag, 2004).
# XANTHORIICOLA PHYSCIAE (Kalchbr.) D. Hawksw.
– 20: on Xanthoria parietina on Quercus
robur, MK (UGDA-L). Freq.: rr. – The species
has been recorded from Estonia only once,
from island Saaremaa (Randlane & Saag,
2004). The infection of this hyphomycete is
22
Folia Cryptog. Estonica
easily recognized through the host apothecia
which are turned black.
ACKNOWLEDGEMENTS
Martin Kukwa thanks dr. Urszula Bielczyk
(Kraków, Poland) for confirming the identification
of Absconditella delutula; the participation in
post-symposium excursion and collaboration of
Paweł Czarnota was supported by Committee
for Scientific Research in Poland (grant KBN No.
3 P04C 040 23). Estonian co-authors received
financial support from the grant No. 5823 of the
Estonian Science Foundation.
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Thor, G. 1991. New or interesting lichens from
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Steiermark. Linzer Biol. Beitr. 31(1): 507–532.
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Estonia. Folia Cryptog. Estonica 36: 17–21.
Hawksworth, D. L. & Hafellner, J. 1986.
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lichenicolous deuteromycete. Nova Hedwigia 43:
525–530.
Hermansson, J., Pystina N.T. & Kudrejasheva, D.I.
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Komi Republic (in Russian, English summary).
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39: 62.
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some new records for the country. Herzogia 18
(in press).
Lumbsch, H. T., Plümper, M., Guderley, R. & Feige,
G. B. 1997. The corticolous species of Lecanora
sensu stricto with pruinose apothecial discs.
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Uppsala, 131–161.
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T. & Vitikainen, O. 2004. Lichen-forming and
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Folia Cryptog. Estonica, Fasc. 41: 23–34 (2005)
Ecological analysis of lichens in the Teberda State Biosphere
Reserve (North-Western Caucasus, Russia)
Olga Blinkova1 & Gennadiy Urbanavichus2
Lomonosov Moscow State University, Vorobjovy Gory, 119899 Moscow, Russia
E-mail: blinok@yandex.ru
2
Polar-Alpine Botanical Garden-Institute of the Kola Sci. Center of RAS,
184256 Kirovsk-6, Murmansk region, Russia
E-mail: urban@aprec.ru
1
Abstract: The distribution and general ecology of 389 species of lichens are recorded for ca 694 km2 of mountainous
terrain in the Teberda State Biosphere Reserve located on the northern side of the Great Caucasus mountain range. 56 taxa
are documented for the first time for North Caucasus, 36 for Caucasus and 3 for Russia (Aspicilia szechenyi, Bacidina delicata,
Rhizocarpon carpaticum). There are more crustose species (48,7%) than foliose (31,4%) or fruticose species (19,8%). Corticolous
lichens were found on 24 tree species, the greatest biodiversity being recorded on Abies nordmanniana, Acer platanoides, Pinus
sylvestris and Fagus orientalis.
Kokkuvõte: Teberda Riikliku Biosfääri Kaitseala (Loode-Kaukasus, Venemaa) samblike ökoloogiline
analüüs.
Käsitletakse Teberda Riikliku Biosfääri Kaitseala 389 samblikuliigi levikut ja põhilisi ökoloogilisi nõudlusi. Proovid koguti
umbes 694 km2 suuruselt mägiselt maastikult Suur-Kaukasuse aheliku põhjaküljel. 56 taksonit mainitakse esmakordselt PõhjaKaukasuse, 36 Kaukasuse ja 3 Venemaa jaoks. Sagedasemad on kooriksamblikud (48,7%), vähemsagedased leht- (31,4%) ja
põõsassamblikud (19,8%). Epifüütseid samblikke leiti 24 puuliigil, suurim oli bioloogiline mitmekesisus forofüütidel Abies
nordmanniana, Acer platanoides, Pinus sylvestris ja Fagus orientalis.
INTRODUCTION
The Great Caucasus is a huge mountain area
on the border between Europe and Asia. Its rich
flora and varied vegetation are very attractive
for naturalists and scientists. The complicated
and dramatic history of the area and the harsh
alpine environment are favorable to nature
protection, with several large national reserves
on the northern slopes of the Great Caucasus
(North-Osetia, Kabardino-Balkaria, Teberda,
and the Caucasian Reserve in the Krasnodar
region) were established in the 20th century.
The lichen flora of Teberda Reserve, although one of the richest in the Great Caucasus, has attracted relatively little attention with
only 79 species known before our investigations
(Barhalov, 1983; Novruzov & Onipchenko, 1985;
Onipchenko, 1987; Vorobyova & Onipchenko,
1994). The investigation of lichens in this region is part of an integrated study of biodiversity
which has been carried out in this mountain
region since 1936.
The present study has two primary objectives: first, to provide a floristic account of the
lichens and increase our knowledge of lichens
in the Teberda Reserve; second, to indicate the
salient features of ecological distribution of the
species.
THE STUDY AREA
The Teberda State Biosphere Reserve is located
on the northern side of the Great Caucasus
mountain range (Fig. 1). The main section of
the reserve occupies the upper reaches of the
Teberda river valley between 41°35’–41°55’ E,
and 43°13’–43°28’ N, and has a total area of ca
694 km2. Elevations inside the reserve range
from 1259 m to 4046 m a.s.l.; about 83% of the
reserve’s territory lies above 2000 m a.s.l. and
can be considered to be truly alpine. Siliceous
rocks predominate in the reserve, but local outcrops of limestone exist two km to the north of
the reserve boundary (Tushinskii, 1957).
The climate of different parts of the reserve
varies considerably due to the two main factors:
elevation and distance from the Great Caucasus
Range, which forms the southern boundary of
the reserve. Overall, the mean annual tempera-
24
Folia Cryptog. Estonica
ture decreases by 0.5–0.6 °C per 100 m increase
in altitude. Precipitation increases with altitude
up to about 2200–2500 m. On the whole, the
climate of the Teberda Reserve is temperatecontinental. In the region of the town of Teberda,
the average annual temperature is +1,2 °C, the
average relative humidity is 79%, and the average annual rainfall is 1400 mm.
The total number of vascular plant species
found in the Teberda Reserve is ca 1150 (Vorobyova & Onipchenko, 2001); 35% of the reserve
is forested, 28% is covered with meadows, and
more than 10% is occupied by glaciers (Pavlov,
2004). There are more than 100 tree species in
the reserve, the most frequent being Pinus sylvestris, Betula litwinowii, Abies nordmanniana,
Picea orientalis and Fagus orientalis (Kononov,
1957). Vegetation units (life zones) are based
on the altitudinal zonation of the northwestern
Caucasus proposed by Kuznetsov (1909). Six
altitudinal belts have been recognized in the
Teberda Reserve: middle forest (1250–1600 m
a.l.s), upper forest (1600–2000 m), subalpine
(2000–2500 m), alpine (2500–3200 m), subnival
(3200–3400 m) and nival (from 3200–3800 m).
RUSSIA
Caspian Sea
GEORGIA
ARMENIA
TURKEY
RESULTS AND DISCUSSION
The full list of lichens in the Teberda Reserve
(Appendix 1) consists of 389 species, distributed
in 125 genera, 45 families, 11 orders, 3
subclasses and one class – Ascomycetes (Kirk
et al., 2001). 56 taxa are documented for the
first time for North Caucasus, 36 for Caucasus,
and 3 for Russia (Aspicilia szechenyi, Bacidina
delicata, Rhizocarpon carpaticum).
Growth forms of lichens
TEBERDA
RESERVE
Black Sea
2002). For an ecological summary we report
lichen species by growth forms, substrate
groups, and distribution on tree species.
Epiphyte samples were collected from branches,
trunks and bases of trees to a height of 2 m; for
these analyses only trees with 20 specimens or
more were sampled.
Subnival and nival belts were combined
into one because of the small number of lichen
samples and species represented.
The collected specimens have been deposited
in the herbarium of the Biological Faculty
of the Moscow State University (MW) and in
the Herbarium of the Polar-Alpine Botanical
Garden-Institute (KPABG).
The names of taxa are given according to
the latest lichen checklist of Austria (Hafellner
& Türk, 2001) and authors' abbreviations are
according to Kirk and Ansell (1992).
AZERBAIJAN
IRAN
Fig. 1. Location of Teberda Reserve in Caucasus area.
MATERIAL AND METHODS
24 canyons with a total length of about 150 km
at elevations ranging from 1259 to 3800 m a.s.l.
were investigated. About 2000 specimens of
lichens were collected in the different altitudinal
belts and plants communities between 2000–
2003. Some data from the literature were
also interpreted (Barhalov, 1983; Novruzov &
Onipchenko, 1985; Vorobyova & Onipchenko,
1994; Onipchenko et al., 1999; Onipchenko,
Three growth forms of lichens are distinguished:
crustose, foliose and fruticose; the first group
includes also effigurate, squamulose and
peltate taxa (Oksner, 1974). Crustose species
(e.g. Aspicilia caesiocinerea, Bellemerea alpina,
Candelariella vitellina, Lecanora intricata,
Rhizocarpon geographicum) predominate in the
investigated territory (189 species or 48,7%); this
is usual for all lichen floras of arctic, mountain
and arid areas (Urbanavichus, 1998). They play
an important role at all elevations, but show the
maximum species diversity (89 species or 45,2%)
in the upper forest belt (Fig. 2). A wide variety
of microhabitats creates favorable conditions
here for different species of all growth forms.
In the alpine and nival belts crustose lichens
predominate over other growth forms (Table 1).
The rigorous conditions of these altitudes are
evidently less favorable for foliose and fruticose
lichens and vascular plants.
25
subnival and nival belts
46
27
25
alpine belt
31
subalpine belt
41
42
43
31
upper forest belt
crustose
foliose
fruticose
66
71
37
middle forest belt
the number of species 0
10
20
30
41
40
50
60
70
89
76
80
80
90
100
Fig. 2. The distribution of fruticose, foliose and crustose lichen species in different altitude
belts.
Table 1. Percentage of crustose, foliose and
fruticose lichens in the different altitude belts.
Altitude belt
middle forest
upper forest
subalpine
alpine
subnival and nival
crustose
38,6
45,2
36,2
47,8
46,9
foliose
40,6
36,0
37,1
29,7
27,6
fruticose
20,8
18,8
26,7
22,5
25,5
There are 123 species (31,6%) of foliose lichens
(e.g. Brodoa atrofusca, Melanelia glabra, Physcia
aipolia, Physconia distorta, Umbilicaria cylindrica) in the reserve. This group is more sensitive
to environmental factors than crustose lichens
(Golubkova, 1977) and predominates in the middle forest belt where a milder climate persists.
Fruticose lichens (e.g. Allocetraria madreporiformis, Bryoria nadvornikiana, Cetraria
islandica, Cladonia pyxidata) are the least numerous (77 species or 19,8%) in all altitudinal
belts.
In general, the percentage of crustose
lichens of the total lichen flora increases with
altitude whereas the percentage of foliose lichens
falls (Makryi, 1990), but our studies do not
support this to date (Fig. 2); foliose lichens are
much better known than crustose lichens in the
Teberda Reserve, thus evidently many crustose
taxa have yet to be reported. Further inquiry
is needed to show the real picture. However,
species richness of all growth forms tends to
decrease with altitude. This phenomenon is
probably connected with the smaller diversity
of substrates in the upper belts as well as the
rigorous environmental conditions.
Substrate preferences
Substrate is the most tangible element of all
environmental conditions influencing lichens.
The type and properties of substrate have a great
impact on ecology, distribution and taxonomy
of lichens (Brodo, 1974).
Five substrate groups of lichens are
distinguished in this study: corticolous (on
bark of trunks and branches of trees), saxicolous
(directly attached to the rock), terricolous
(on soil), muscicolous (on bryophytes) and
lignicolous (on lignum) species. Although some
lichens regularly colonize a wide variety of
substrates, most species tend to reveal a distinct
preference for a single substrate type (284
species or 73,1%). This tendency is especially
well developed among corticolous and saxicolous
species of crustose lichens.
Corticolous species (e.g Arthonia radiata,
Arthothelium ruanum, Bacidia subincompta, Biatora vernalis, Bryoria subcana) form the biggest
substrate group in the Teberda Reserve (174
species, 44,8%). Their distribution is controlled
by the distribution of tree species, being absent
above the limit of trees and bushes (Fig.3). Otherwise, crustose species predominate on a wide
variety of deciduous trees (Acer platanoides, A.
trautvetteri, Alnus incana, Berberis vulgaris,
Betula litwinowii, B. pendula, Carpinus betulus,
Corylus avellana, Crataegus monogyna, Fagus
orientalis, Fraxinus excelsior, Malus sylvestris,
Padus avium, Populus alba, P. tremula, Quercus
robur, Rosa spp., Salix spp., Sorbus aucuparia,
Ulmus glabra) and outnumber species on coniferous trees (Abies nordmanniana, Juniperus
communis, Picea orientalis, Pinus sylvestris).
26
Folia Cryptog. Estonica
subnival and nival belts
saxicolous
alpine belt
terricolous
saxicolous
subalpine belt
corticolous
terricolous
saxicolous
corticolous
upper forest belt
middle forest belt
0
10
20
30
40
50
musci
terricolous
saxicolous
corticolous
%
musci
mus
terri mus
ligni
saxi
terri musci
60
70
80
90
ligni
100
Fig. 3. The distribution of substrate groups of lichens in different altitude belts.
However, the greatest numbers of lichens where
found on Abies nordmanniana (67 species), Pinus
sylvestris (51), Acer platanoides (46) and Fagus
orientalis (43), which are the most numerous
trees in the Teberda Reserve (Table 2).
Most of the species (239 species, or 61,6%) are
typical epiphytes (mainly species of Lecanora,
Ramalina and Usnea). However, many facultative epiphytes which often grow on other types
of substrates (e.g. Cladonia fimbriata and C.
pyxidata) also occur on trees.
133 species (34,2%) found on rock (e.g.
Cornicularia normoerica, Lecanora intricata,
Lecanora polytropa) are all acidophilous as only
siliceous rocks occur in the research area (see
above, Study Area). Since this group, mainly
crustose, will prove to be the most biodiverse,
further investigation is obviously necessary. The
percentage of saxicolous lichens increases with
altitude and reaches a maximum in the alpine
belt (Fig. 3).
The 90 terricolous species (23%) found in the
Teberda Reserve (e.g. Cetraria islandica, Peltigera
canina, P. lepidophora, Pertusaria geminipara,
Phaeorrhiza nimbosa, Physconia muscigena) are
mostly fruticose. The percentage of terricolous
lichens increases with altitude (Fig. 3).
Muscicolous lichens (71 species, 18,2%;
e.g. Fuscopannaria praetermissa, Lepraria
cacuminum, Peltigera leucophlebia, Physconia
muscigena) are mainly foliose and maintain
a more-or-less constant percentage along the
altitudinal gradient (Fig. 3).
The lowest number of species are lignicolous
(50 species, 12,8%; e.g. Absconditella lignicola,
Alectoria sarmentosa). As in the case of the corticolous species, the distribution of lignicolous
Table 2. Occurrence of corticolous lichens on
different tree species
Tree species
Number of lichen species
conifers
Abies nordmanniana
67
Juniperus communis
9
Picea orientalis
7
Pinus sylvestris
51
deciduous
Acer platanoides
46
Acer trautvetteri
17
Alnus incana
26
Berberis vulgaris
3
Betula litwinowii
23
Betula pendula
22
Carpinus betulus
6
Corylus avellana
11
Crataegus monogyna
5
Fagus orientalis
43
Fraxinus excelsior
17
Malus sylvestris
10
Padus avium
18
Populus alba
6
Populus tremula
8
Quercus robur
14
Rosa spp.
2
Salix spp.
26
Sorbus aucuparia
14
Ulmus glabra
4
27
species is restricted by the range of forests
(Fig. 3). All growth forms were present among
lignicolous lichens almost in equal numbers.
ACKNOWLEDGEMENTS
The authors would like to thank the Department
of Flora and Vegetation of the Polar-Alpine
Botanical Garden-Institute and especially
Irina Urbanavichene and Tamara Dudoreva
for identifying some of the lichens, and
Denis Davydov for helpful comments on the
manuscript. The authors are also grateful to
Vladimir Onipchenko for valuable information
and the opportunity to do this work. Finally, we
would like to thank all members of MSU Teberda
expedition for their support.
REFERENCES
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(in Russian). Baku. 338 pp.
Brodo, I. M. 1974. Substrate ecology. In: Ahmadjian,
V. & Hale, M. E. (eds). The lichens, pp. 401–441.
Academic Press, New York.
Golubkova, N. S. 1977 Notes about geographical
connections of lichens in Pamir (in Russian). Novitates Systematicae Plantarum non Vascularium
14: 172–185.
Hafellner, J. & Türk, R. 2001. Die lichenisierten
Pilze Osterreichs – eine Checkliste der bisher
nachgewiesenen Arten mit Eerbreitungsangaben.
Stapfia 76: 3–167.
Jatta, A. 1900. Lichenes. In: Sommier, S. & Levier,
E. (eds). Enumeratio plantarum anno 1890 in
Caucaso lectarum. Acta Horto Petropoliatani 16:
523–536.
Kirk, P. M. & Ansell, A. E. 1992. Authors of fungal
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Kirk, P. M., Cannon, P. F., David J. C. & Stalpers, J.A.
(eds) 2001. Dictionary of the Fungi, 9th Edition.
CABI, Wallingford. 624 pp.
Kononov, V. N. 1957. The vegetation of Teberda
Reserve (in Russian). Transactions of Teberda
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phytogeographic provinces in the Caucasus (in
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1–177.
Makryi, T. V. 1990. The lichens of Baikal’s mountain
ridge (in Russian). Nauka, Novosibirsk. 198 pp.
Novruzov, V. S. & Onipchenko, V. G. 1985 Notes to the
lichen flora of alpine belt in the Teberda State Reserve (in Russian). Bot. Journ. 70 (6): 799–802.
Oksner, A.N. 1974. The key of lichens of USSR. Part
2. The morphology, taxonomy and geography (in
Russian). Nauka, Leningrad. 283 pp.
Onipchenko, V. G. 1987. Lichens and higher plants (in
Russian) In: Rabotnov, T. A. (ed), Biogeocenoses of
alpine desert, pp. 19–30. Nauka, Moscow.
Onipchenko, V. G. 2002. Alpine vegetation of the Teberda Reserve, the Northwestern Caucasus. Ver
ff. des Geobot. Inst. der ETH 130: 1–168.
Onipchenko, V. G., Egorov, A. V., Gluchova, E. M.
& Hanina L. G. 1999. The distribution of high
mountain plants on their ecotopes: the analysis
of date bank of geobotanical descriptions (in
Russian). Transactions of Teberda State Reserve
(Stavropol) 15: 166–206.
Pavlov, V. N. 2004. Plant geographical description
of the area. In: Alpine ecosystems in the Northwest Caucasus, pp. 25–54. Kluwer Academic,
Dordrecht.
Takhthadjan, A. L. (ed.) 1988. Red Data Book of RFSR:
Plants (in Russian). Moscow.
Tushinskii, G. K. 1957. Geomorphology of the Teberda
Reserve (in Russian). Transactions of Teberda
State (Stavropol) 1: 3–49.
Urbanavichus, G. P. 1998. The biogeographical
regularities of the lichens flora organization in
the South Baikal areas (in Russian). Thesis of
dissertation of candidate degree.
Vorobyova, F. M. & Kononov, V. N. 1991. Flora of the
Teberda Reserve (vascular plants) (in Russian).
Transactions of Teberda State Reserve (Stavropol)
13: 1–137.
Vorobyova, F. M. & Onipchenko, V. G. 1994. The
species of lichens and mushrooms from the Red
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and fauna of the reserves 99. Moscow. 100 pp.
28
Folia Cryptog. Estonica
APPENDIX
The list of lichens in the Teberda Reserve
Abbreviations of substrate: B – bryophytes, L
– lignum, R – rock, S – soil, Abi – Abies nordmanniana, AceP – Acer platanoides, AceT – Acer
trautvetteri, Aln – Alnus incana, Ber – Berberis
vulgaris, BetL – Betula litwinowii, BetP – Betula
pendula, Car – Carpinus betulus, Cor – Corylus
avellana, Cra – Crataegus monogyna, Fag – Fagus
orientalis, Fra – Fraxinus excelsior, Jun – Juniperus communis, Mal – Malus sylvestris, Pad
– Padus avium, Pic – Picea orientalis, Pin – Pinus
sylvestris, PopA – Populus alba, PopT – Populus
tremula, Que – Quercus robur, Ros – Rosa spp.,
Sal – Salix spp., Sor – Sorbus aucuparia, Ulm
– Ulmus glabra.
Range of altitudes where the species were
collected is given in the brackets. The literature
data are added as well. Species new to Russia are
marked by ***, to Caucasus – by **, and to North
Caucasus – by *. Species in the Red Data Book of
Russia (Takhthadjan, 1988) are given in bold.
**A BROTHALLUS PRODIENS (Harm.) Diederich &
Hafellner – on thallus of Hypogymnia
physodes (1450).
**ABSCONDITELLA LIGNICOLA Vězda & Pišut – on L (2000).
ACAROSPORA ATRATA Hue – on R (2400).
ACAROSPORA MOLYBDINA (Wahlenb.) Trevis. – on R
(2000) (Novrusov & Onipchenko, 1985).
ACAROSPORA RUFA (Vain.) H. Magn. – on R (2000)
(Novrusov & Onipchenko, 1985).
ACAROSPORA SMARAGDULA (Wahlenb.) A. Massal. – on
R (3850).
ACAROSPORA VERONENSIS A. Massal. – on R (2700).
ALECTORIA SARMENTOSA (Ach.) Ach. – on AceP and
Pic, on L (1500–1700).
ALLOCETRARIA MADREPORIFORMIS (Ach.) Kärnefelt & A.
Thell – on S (2750–3151).
**AMANDINEA CACUMINUM (Th. Fr.) H. Mayrhofer &
Sheard – on R (2050–2600).
AMANDINEA PUNCTATA (Hoffm.) Coppins & Scheid. –
on B and R, on BetP, Cra, Cor, Abi, (2700).
**AMYGDALARIA PANAEOLA (Ach.) Hertel & Brodo – on
R (1600).
ANAPTYCHIA CILIARIS (L.) Körb. – on L, on Aln, Fra,
Cor, BetP, Sor, Fag (1300–1600).
ARTHONIA RADIATA (Pers.) Ach. – on AceP and Cor
(1350–1500).
ARTHOTHELIUM RUANUM (A. Massal.) Körb. – on AceP
and Fag (1500–1600).
*ARTHROSPORUM POPULORUM A. Massal. – on Ulm and
AceP (1350).
ASPICILIA CAESIOCINEREA (Nyl. ex Malbr.) Arnold – on
R (1600–3850).
ASPICILIA CINEREA (L.) Körb. – on R (1700–2400).
ASPICILIA DESERTORUM (Kremp.) Mereschk. – on R
(2700).
*ASPICILIA LAEVATA (Ach.) Arnold – on R (2000).
ASPICILIA MACULATA (H. Magn.) Oxner – on R (2000)
(Novruzov & Onipchenko, 1985).
*ASPICILIA SIMOËNSIS Räsänen – on R (2400).
ASPICILIA SPHAEROSPORA (Tomin) Oxner – on R (2000)
(Novruzov & Onipchenko, 1985).
***ASPICILIA SZECHENYI (Vain.) Hue – on R (2000–
3850).
ASPICILIA VERRUCIGERA Hue – on R (2000–2800).
BACIDIA HERBARUM (Stizenb.) Arnold – on S (2500).
**B ACIDIA SUBINCOMPTA (Nyl.) Arnold – on Abi
(2000).
***BACIDINA DELICATA (Larbal. ex Leight.) V. Wirth &
Vězda – on Fra (1350).
BELLEMEREA ALPINA (Sommerf.) Clauzade & Cl. Roux
– on R (1560–2800).
BELLEMEREA CUPREOATRA (Nyl.) Clauzade & Cl. Roux
– on R (2000).
*BIATORA VERNALIS (L.) Fr. – on AceP (1500).
B RODOA ATROFUSCA (Schaer.) Goward – on R
(2000–3850).
BRODOA INTESTINIFORMIS (Vill.) Goward – on R (1800
–3850).
BRYOCAULON DIVERGENS (Ach.) Kärnefelt – on S
(2800–3050).
BRYONORA CASTANEA (Hepp) Poelt – on S (2850).
BRYORIA BICOLOR (Ehrh.) Brodo & D. Hawksw. – on
B and R (1300–2700).
BRYORIA CHALYBEIFORMIS (L.) Brodo & D. Hawksw.
– on AceP, on L and S (1500–2030).
BRYORIA FUSCESCENS (Gyeln.) Brodo & D. Hawksw.
– on L, on Pic, Pin (1550–1800).
BRYORIA IMPLEXA (Hoffm.) Brodo & D. Hawksw. – on
Abi, Pin, AceP (1500–2700).
B RY ORI A NAD V O R NI KI ANA (Gyeln.) Brodo & D.
Hawksw. – on Sal, Fag, Abi, Pin, on R, B, L
(1400–2750).
BRYORIA NITIDULA (Th. Fr.) Brodo & D. Hawksw. – on
R, B, S (2980–3050).
BRYORIA SUBCANA (Nyl. ex Stizenb.) Brodo & D.
Hawksw. – on Fag (1500–1600).
BRYORIA TRICHODES (Michx.) Brodo & D. Hawksw.
– on Fag, Pin (1400–1500).
BUELLIA AETHALEA (Ach.) Th. Fr. – on R (2000).
*BUELLIA CHLOROLEUCA Körb. – on L (1600).
BUELLIA DISCIFORMIS (Fr.) Mudd. – on Fag and Pin
(1600–2000).
BUELLIA ERUBESCENS Arnold – on L, on Abi (1900–
2000).
29
BUELLIA GRISEOVIRENS (Turn. & Borr. ex Sm.) Almb.
– on Pin (1800).
*BUELLIA INSIGNIS (Naegeli ex Hepp) Th. Fr. – on Pin,
S, B (1800, 3000–3050).
**BUELLIA LEPTOCLINE (Flot.) A. Massal. – on R
(2600).
**BUELLIA MIRIQUIDICA Scheid. – on R (2700–3850).
BUELLIA PAPILLATA (Sommerf.) Tuck. – on S (3060).
**BUELLIA PULVERULENTA (Anzi) Jatta – on thallus of
Physconia muscigena (3050).
BUELLIA SCHAERERI De Not. – on Abi (2000).
BUELLIA VILIS Th. Fr. – on S (2700).
CALICIUM ADAEQUATUM Nyl. – on Fag (1400).
CALOPLACA AMMIOSPILA (Wahlenb.) H. Olivier – on S
(3060).
CALOPLACA ARENARIA (Pers.) Müll. Arg. – on R (1300,
3050–3060).
CALOPLACA CERINA (Ehrh. ex Hedw.) Th. Fr. – on
Aln, Ulm, Fra, AceP, AceT, Sal, Pad, BetL
(1300–2000, 3050).
*CALOPLACA CRENULARIA (With.) J.R. Laundon – on
R (2700).
**CALOPLACA EPITHALLINA Lynge – on R, on the tallus
of a lichen (1560–2000).
CALOPLACA FERRUGINEA (Huds.) Th. Fr. – on Jun, on
S (2050–2800).
CALOPLACA HERBIDELLA (Hue) H. Magn. – on AceP,
Fag, Sor, Abi (1500–2000).
CALOPLACA HOLOCARPA (Hoffm.) Wade – on PopT
(1600–2000).
**CALOPLACA HUNGARICA H. Magn. – on Pin (1600).
CALOPLACA JUNGERMANNIAE (Vahl) Th. Fr. – on S, B
(1900, 3050).
**CALOPLACA NIVALIS (Körb.) Th. Fr. – on S (3000).
CALOPLACA SAXICOLA (Hoffm.) Nordin – on R (2400)
(Novruzov & Onipchenko, 1985).
*CALOPLACA TIROLIENSIS Zahlbr. – on S (2600).
C ALVITIMELA ARMENIACA (DC.) Hafellner – on R
(2700).
CANDELARIA CONCOLOR (Dicks.) Stein – on Aln, Ber,
Que, AceP (1300–1350).
CANDELARIELLA AURELLA (Hoffm.) Zahlbr. – on Que,
BetL (1300–2000).
CANDELARIELLA CORALLIZA (Nyl.) H. Magn. – on R
(2400–2600).
**CANDELARIELLA LUTELLA (Vain.) Räsänen – on AceP,
Cor (1350).
CANDELARIELLA REFLEXA (Nyl.) Lettau – on Mal, Cra,
BetL (1300–2000).
CANDELARIELLA VITELLINA (Hoffm.) Müll. Arg. – on Aln,
BetL, on R (1300–3850).
CANDELARIELLA XANTHOSTIGMA (Ach.) Lettau – on AceP,
BetL, Sal, Abi (1650–2000).
CARBONEA VITELLINARIA (Nyl.) Hertel – on thallus of
Candelariella (2700–3100).
CARBONEA VORTICOSA (Flörke) Hertel – on R (3050).
CATAPYRENIUM CINEREUM (Pers.) Körb. – on S (3080–
3850).
CETRARIA ACULEATA (Schreb.) Fr. – on R, S (2400–
3150).
CETRARIA ERICETORUM Opiz – on S (2950).
CETRARIA ISLANDICA (L.) Ach. – on S (1800–3150).
CETRARIA LAEVIGATA Rassad. – on S (2700).
CETRARIA MURICATA (Ach.) Eckfeldt – on S (2800–
2900).
**CETRARIA ODONTELLA (Ach.) Ach. – on R (3000).
CETRELIA CETRARIOIDES (Duby) W.L. Culb. & C.F.
Culb. – on R, S (1550–2000).
CETRELIA OLIVETORUM (Nyl.) W.L. Culb. & C.F. Culb.
– on B (1450).
C HAENOTHECA FUR FURACEA (L.) Tibell – on Abi
(1450).
CHRYSOTHRIX CANDELARIS (L.) J.R. Laundon – on Fag,
Abi (1400–1600).
CLADONIA AMAUROCRAEA (Flörke) Schaer. – on S
(2000–2400).
CLADONIA ARBUSCULA (Wallr.) Flot. – on S (2630).
CLADONIA CARIOSA (Ach.) Spreng. – on S (3050).
CLADONIA CENOTEA (Ach.) Schaer. – on S, L (1400–
2000).
CLADONIA CHLOROPHAEA (Flörke ex Sommerf.) Spreng.
– on S, B (1400–2700).
CLADONIA CONIOCRAEA (Flörke) Spreng. – on S, L
(1450–2630).
CLADONIA CORNUTA (L.) Hoffm – on S (2000–2750).
C LADONIA CRISPATA (Ach.) Flot. – on S (2000,
3000).
CLADONIA DIGITATA (L.) Hoffm. – on Pin (1800).
CLADONIA ECMOCYNA (Ach.) Leigth. – on S (2200,
3050).
CLADONIA FIMBRIATA (L.) Fr. – on S, L, B (1500–
2800).
CLADONIA FURCATA (Huds.) Schrad. – on S (2000–
2900).
CLADONIA GRACILIS (L.) Willd. – on S (1800–2900).
C LADONIA MACILENTA Hoffm. – on S, L (1600–
2050).
CLADONIA MACROCERAS (Delise) Hav. – on S (2750–
2900).
CLADONIA MITIS Sandst. – on S (2000–2800).
CLADONIA OCHROCHLORA Flörke – on S (1300–1600,
2100).
CLADONIA PHYLLOPHORA Hoffm. – on S (1800–2900).
CLADONIA PLEUROTA (Flörke) Schaer. – on S (1600–
2600).
CLADONIA POCILLUM (Ach.) Grognot – on S (2630–
3050).
30
Folia Cryptog. Estonica
CLADONIA PYXIDATA (L.) Hoffm. – on S, B, on Abi,
Jun (1300–3850).
CLADONIA RAMULOSA (With.) J.R. Laundon – on R
(2000).
CLADONIA RANGIFERINA (L.) Weber ex F.H. Wigg. – on
S (1800, 3050).
CLADONIA SQUAMOSA Hoffm. – on L (1850).
C LADONIA STELLARIS (Opiz) Pouzar & Vězda – on S
(2950–3000).
CLADONIA SYMPHYCARPIA (Flörke) Fr. – on S (1450–
2400).
CLIOSTOMUM CORRUGATUM (Ach.: Fr.) Fr. – on Abi
(1600).
**CLIOSTOMUM LEPROSUM (Räsänen) Holien & Tønsberg – on Abi (1500).
COLLEMA AURIFORME (With.) Coppins & J.R. Laundon
– on R (2920).
COLLEMA FLACCIDUM (Ach.) Ach. – on R, B, on Fra,
AceT, Sal (1350–1650).
COLLEMA FURFURACEUM (Arnold) Du Rietz – on Jun,
AceT (1350–1650).
COLLEMA FUSCOVIRENS (With.) J.R. Laundon – on
Fra (1350).
COLLEMA TENAX var. CERANOIDES (Borrer) Degel. – on
S (2100).
COLLEMA UNDULATUM Laurer ex Flot. – Teberda
(Jatta, 1900).
CORNICULARIA NORMOERICA (Gunnerus) Du Rietz – on
R (2050–2700).
DENDRISCOCAULON UMHAUSENSE (Auersw.) Degel. – on
AceP, AceT (1500–1600).
DERMATOCARPON ARNOLDIANUM Degel. – on R (1600–
2800).
DERMATOCARPON INTESTINIFORME (Körb.) Hasse – on
R (2000–3850).
DERMATOCARPON LURIDUM (With.) J.R. Laundon – Teberda (Jatta, 1900).
D ERMATOCARPON MINIATUM (L.) W. Mann – on R
(1300–2620).
DERMATOCARPON RIVULORUM (Arnold) Dalla Torre &
Sarnth. – on R (2800) (Novruzov & Onipchenko, 1985).
DIMELAENA OREINA (Ach.) Norman – on R (2000–
2500).
DIPLOSCHISTES MUSCORUM (Scop.) R. Sant. – on B, S
(2800–3050).
DIPLOSCHISTES SCRUPOSUS (Schreb.) Norman – on
R (2000).
EVERNIA DIVARICATA (L.) Ach. – on AceP, Sal, Abi, Pic
(1400–2000).
EVERNIA PRUNASTRI (L.) Ach. – on PopT, Fra (1400).
FLAVOCETRARIA CUCULLATA (Bellardi) Kärnefelt & A.
Thell – on S (2750–3150).
FLAVOCETRARIA NIVALIS (L.) Kärnefelt & A. Thell – on
S (3150).
FLAVOPARMELIA CAPERATA (L.) Hale – on Que, BetP,
Sal, Mal, B (1250–2200).
FLAVOPUNCTELIA SOREDICA (Nyl.) Hale – on BetP
(1300).
FUSCIDEA CYATHOIDES (Ach.) V. Wirth & Vězda – on R
(2700).
FUSCOPANNARIA LEUCOPHAEA (Vahl) P.M. Jørg. – on B
(2200–3050).
F USCOPANNARIA MEDITERRANEA (Tav.) P.M. Jørg.
– on S, on thallus of Lobaria scrobiculata
(1500–2000).
FUSCOPANNARIA PRAETERMISSA (Nyl.) P.M. Jørg. – on
B (1400).
HETERODERMIA SPECIOSA (Wulfen) Trevis. – on Sal,
B, S (1350–2000).
HYPOCENOMYCE SCALARIS (Ach.) M. Choisy – on Pin
(1500).
HYPOGYMNIA AUSTERODES (Nyl.) Räsänen – on B, L
(1400–2000).
HYPOGYMNIA BITTERI (Lynge) Ahti – on BetL (1900).
HYPOGYMNIA PHYSODES (L.) Nyl. – on BetP, BetL, AceP,
Sor, Fag, Pin, B, L (1300–2200).
HYPOGYMNIA TUBULOSA (Schaer.) Hav. – on Abi, BetL
(2000–2050).
HYPOGYMNIA VITTATA (Ach.) Parrique – on Abi, on B
(1500–1700, 2600).
H YPOTRACHYNA LAEVIGATA (Sm.) Hale – on Abi
(1500).
HYPOTRACHYNA REVOLUTA (Flörke) Hale – on Fag
(1500).
I CMADOPHILA ERICETORUM (L.) Zahlbr. – on L, B
(1800–2400).
IMSHAUGIA ALEURITES (Ach.) S.L.F. Meyer – on Pin
(1800).
LASALLIA PENSYLVANICA (Hoffm.) Llano – on R (1750–
1800).
LECANIA CYRTELLA (Ach.) Th. Fr. – on Fag, PopA
(1300–1350).
**L ECANIA CYRTELLINA (Nyl.) Sandst. – on PopA
(1300).
*LECANIA FUSCELLA (Schaer) Körb. – on Fag (1350).
LECANIA NAEGELII (Hepp) Diederich & van den Boom
– on Pad (1900).
LECANORA ALBELLA (Pers.) Ach. – on Cra (1300).
LECANORA ALLOPHANA Nyl. – on Fag, Aln, PopT,
(1300–2050, 3850).
LECANORA ARGENTATA (Ach.) Malme – on Ros, Fag,
Pin (1300–1600).
**LECANORA CADUBRIAE (A. Massal.) Hedl. – on Pin
(1900).
LECANORA CAMPESTRIS (Schaer.) Hue – on L (1850–
2400).
31
LECANORA CARPINEA (L.) Vain. – on AceP, Fag, Aln,
Pad, Pin, Abi (1500–2000).
LECANORA CENISIA Ach. – on R (2700).
LECANORA CHLAROTERA Nyl. – on Aln, PopT, Fag, Sor,
Pin, Abi, on L (1300–2000).
LECANORA DISPERSOAREOLATA (Schaer.) Lamy – Teberda (Jatta, 1900).
LECANORA FRUSTULOSA (Dicks.) Ach.– on R (2000–
2800) (Novruzov & Onipchenko, 1985).
**LECANORA FUSCESCENS (Sommerf.) Nyl. – on Fag
(1700).
LECANORA GLABRATA (Ach.) Malme – on Carp, Fag,
Pad, Abi (1350–2000).
LECANORA HAGENII (Ach.) Ach. – on B, S (2500).
LECANORA HIEROGLYPHICA Poelt – on R (2000) (Novruzov & Onipchenko, 1985).
L ECANORA INTRICATA (Ach.) Ach. – on R (1700–
3050).
*LECANORA INTUMESCENS (Rebent.) Rabenh. – on Que,
Pop, AceP, AceT, Fra (1300–2000).
*LECANORA LAATOKKAENSIS (Räsänen) Poelt – on R
(2600).
*LECANORA LEPTYRODES (Nyl.) Degel. – on Que, AceP,
Pad, Abi (1300–2000).
L ECANORA MURALIS (Schreb.) Rabenh. – on R
(1300–3000).
LECANORA POLYTROPA (Hoffm.) Rabenh. – on R (a
single finding – on L) (1400–3850).
LECANORA POPULICOLA (DC.) Duby – on Aln (1300).
LECANORA RUGOSELLA Zahlbr. – on Pop, Fag (1550,
3850).
LECANORA RUPICOLA (L.) Zahlbr. – on R (1700–2000,
2800).
*LECANORA SALIGNA (Schrad.) Zahlbr. – on Aln, Pin,
L (1600–2000).
LECANORA SUBCARPINEA Szatala – on Ulm, AceT, Pic,
Abi (1300–1500).
*LECANORA SUBINTRICATA (Nyl.) Th. Fr. – on Pin
(1800).
LECANORA SYMMICTA (Ach.) Ach. – on Abi (1600).
L ECANORA VARIA (Hoffm.) Ach. – on Pin, BetL
(1800–2200).
LECIDEA ARTROBRUNNEA (Ramond ex Lam. & DC.)
Schaer – on R (2400–2500, 3850).
LECIDEA AURICULATA Th. Fr. – on R (2000–2100,
3850).
LECIDEA ECRUSTACEA (Anzi ex Arnold) Arnold – Teberda (Jatta, 1900).
LECIDEA LAPICIDA (Ach.) Ach. – on R (2000, 2700–
3850).
LECIDEA PROMISCENS Nyl. – on R (2400–2500).
LECIDEA TESSELLATA Flörke – on R (2050).
LECIDELLA CARPATHICA Körb. – on R (2100, 2700).
L ECIDELLA ELAEOCHROMA (Ach.) M. Choisy – on
Aln, Ros, AceP, Sor, Pad, Carp, Que, Ber
(1300–1900).
LECIDELLA EUPHOREA (Flörke) Hertel – on BetP, BetL,
PopA, Fag, Pin, Jun (1300–2050).
LECIDELLA LAURERI (Hepp) Körb. – on Carp, Sor, Pad,
Abi, Pin (1350–2000).
LECIDELLA STIGMATEA (Ach.) Hertel & Leuckert – on
R (2800).
LECIDOMA DEMISSUM (Rutstr.) Gutth. Schneid. &
Hertel – on S, B (2000–2900).
*LEMPHOLEMMA POLYANTHES (Bernh.) Malme – on S
(1450).
**LEPRARIA CACUMINUM (A. Massal.) Loht. – on S
(2200).
**LEPRARIA CAESIOALBA (de Lesd.) J.R. Laundon – on
S, B and plant remains (2200–3100).
**LEPRARIA JACKII Tønsberg– on Sal (1400).
LEPRARIA LOBIFICANS Nyl. – on Abi (1450).
LEPTOGIUM BURNETIAE C.W. Dodge – on Fra, Fag,
AceT (1350–1650).
LEPTOGIUM CYANESCENS (Rabenh.) Körb. – on B
(1600).
LEPTOGIUM HILDENBRANDII (Garov.) Nyl. – on Fag
(1550–2000).
LEPTOGIUM SATURNINUM (Dicks.) Nyl. – on BetL, AceP,
Pad, S (1500–2000).
LEPTOGIUM TERETIUSCULUM (Wallr.) Arnold – on Fra,
Sal (1350).
LETHARIA VULPINA (L.) Hue – on Pin (1860–2200).
LOBARIA AMPLISSIMA (Scop.) Forss. – on Fag, AceP
(1400–1600).
LOBARIA PULMONARIA (L.) Hoffm. – on Sal, Fa, AceP,
AceT, Abi (1350–1650).
L OBARIA SCROBICULATA (Scop.) DC. – on Bet, B
(1500).
LOBOTHALLIA ALPHOPLACA (Wahlenb.) Hafellner – on R
(2000) (Novruzov & Onipchenko, 1985).
MEGASPORA VERRUCOSA (Ach.) Hafellner & V. Wirth
– on B, S (2750).
MELANELIA COMMIXTA (Nyl.) A. Thell – on R (2000).
M ELANELIA DISJUNCTA (Erichsen) Essl. – on R
(2700).
MELANELIA ELEGANTULA (Zahlbr.) Essl. – on Fag
(1800–2000).
MELANELIA EXASPERATA (De Not.) Essl. – on AceP,
BetP, BetL, Sal, Pad, Pin, Abi (1350–2000).
MELANELIA EXASPERATULA (Nyl.) Essl. – on Aln, Que,
Sal, AceP, BetP, BetL, Sor, Pin, Abi, B, L
(1300–3050).
MELANELIA GLABRA (Schaer.) Essl. – on Aln, Que, Sal,
AceP, Fra, Ulm, Cor, Fag, BetP, BetL, Sor, Abi
(1250–1900).
32
Folia Cryptog. Estonica
MELANELIA HEPATIZON (Ach.) A. Thell – on R (2000–
3100).
MELANELIA INFUMATA (Nyl.) Essl. – on S (3150).
MELANELIA OLIVACEA Essl. – on Aln (1500).
MELANELIA STYGIA (L.) Essl. – on R, B, S (1800–
2750).
MELANELIA SUBARGENTIFERA (Nyl.) Essl. – on PopA,
BetP, Cor, Jun, B (1300–1350).
MELANELIA SUBAURIFERA (Nyl.) Essl. – on Cra, AceP,
B (1300–3050).
**MICAREA BAUSCHIANA (Körb.) V. Wirth & Vĕzda
– on L (2000).
MICAREA PELIOCARPA (Anzi) Coppins & R. Sant. – on
Pin (1900).
MICAREA PRASINA Fr. – on Abi (1600–2000).
MICAREA SYLVICOLA (Flot.) Vězda & V. Wirth – on R
(2000–2700).
**MIRIQUIDICA DEUSTA (Stenh.) Hertel & Rambold
– on R (3150, 3850).
MIRIQUIDICA GAROVAGLII (Schaer.) Hertel & Rambold
– on R (2700).
**MIRIQUIDICA NIGROLEPROSA (Vain.) Hertel & Rambold
– on R (3150).
MYCOBILIMBIA BERENGERIANA (A. Massal.) Hafellner &
V. Wirth – on Abi (1600).
M YCOCALICIUM SUBTILE (Pers.) Szatala – on L
(1500).
MYXOBILIMBIA LOBULATA (Sommerf.) Hafellner – Teberda (Jatta, 1900).
NAETROCYMBE PUNCTIFORMIS (Pers.) R.C. Harris – on
Abi (2000).
NEOFUSCELIA DELISEI (Duby) Essl. – on R (1400–
2450).
N EOFUSCELIA VERRUCULIFERA (Nyl.) Essl. – on R
(1400).
NEPHROMA BELLUM (Spreng.) Tuck. – on AceP, Abi,
on S (1600–2500).
NEPHROMA PARILE (Ach.) Ach. – on Jun, AceP, BetL,
B, S (1350–2650).
NEPHROMA RESUPINATUM (L.) Ach. – on Fag, AceP,
AceT, B, L (1250–2000).
OCHROLECHIA PALLESCENS (L.) A. Massal. – on AceP
(1300).
OCHROLECHIA SZATALAENSIS Vers. – on Sor (2000).
OPEGRAPHA VARIA Pers. – on AceP (1500).
OPHIOPARMA VENTOSA (L.) Norman – on R (2700–
2800).
PANNARIA CONOPLEA (Ach.) Bory –on AcerT, B, S
(1400–2000).
PARMELIA OMPHALODES (L.) Ach. – on R, B, S (1400,
2750–2800).
PARMELIA SAXATILIS (L.) Ach. – on Fag, Abi, Pin, L,
R, S (1450–2800).
P ARMELIA SUBMONTANA Nádv. ex Hale – on Fag
(1500).
PARMELIA SULCATA Taylor – on AceP, AceT, BetP,
BetL, Fra, Aln, Sal, Pad, Cra, Abi, Pin, Jun
(1300–2300).
PARMELIELLA TRIPTOPHYLLA (Ach.) Müll. Arg. – on BetP,
B (1500).
PARMELIOPSIS AMBIGUA (Wulfen) Nyl. – on L (1500–
2000).
P AR MELIOPSIS HYPEROPTA (Ach.) Arnold – on L
(1500–2000).
PARMOTREMA CHINENSE (Osbeck) Hale & Ahti – on R
(2000) (Novruzov & Onipchenko, 1985).
P AR MOTREMA STUPPEUM (Taylor) Hale – on Sal
(1350).
PELTIGERA APHTHOSA (L.) Willd – on S (1400–2000).
P ELTIGERA CANINA (L.) Willd. – on S, B (1250–
2650).
PELTIGERA DEGENII Gyeln. – on S, B (1900–2000).
PELTIGERA ELISABETHAE Gyeln. – on S (1500–1550).
PELTIGERA HORIZONTALIS (Huds.) Baumg. – on L, S
(1450–2000).
PELTIGERA LEPIDOPHORA (Nyl. ex Vain.) Bitter – on
S (1450).
PELTIGERA LEUCOPHLEBIA (Nyl.) Gyeln. – on S, B
(2800–3150).
PELTIGERA MALACEA (Ach.) Funck – on S, B (1800–
2900).
P ELTIGERA MEMBRANACEA (Ach.) Nyl. – on S, B
(1600–2650).
PELTIGERA POLYDACTYLON (Neck.) Hoffm. – on L, B, S
(1250–2000).
**PELTIGERA PONOJENSIS Gyel. – on S, B (1400–
2050).
PELTIGERA PRAETEXTATA (Flörke ex Sommerf.) Zopf
– on S, B, L (1600–2200).
PELTIGERA RUFESCENS (Weiss) Humb. – on S, B, L
(1300–3050).
**PERIDIOTHELIA FULIGUNCTA (Norman) D. Hawksw.
– on Aln (1300).
PERTUSARIA ALBESCENS (Huds.) M. Choisy & Werner
– on AceP, Fag, Abi, Pic, Pin (1300–2000).
PERTUSARIA AMARA (Ach.) Nyl. – on AceP, Fag, Sal,
Abi (1300–1800).
PERTUSARIA COCCODES (Ach.) Nyl. – on Fag and Abi
(1600–1700).
PERTUSARIA CORONATA (Ach.) Th. Fr. – on Abi (1500–
1700).
**PERTUSARIA GEMINIPARA (Th. Fr.) C. Knight – on S
(2950).
P ERTUSARIA LEIOPLACA DC. – on B, Fag (1300–
1700).
*P ERTUSARIA SERVITIANA Erichsen – on Abi, L
(1500).
33
PHAEOPHYSCIA CILIATA (Hoffm.) Moberg – on AceP,
Sal, Pad, Pin (1350–1900).
PHAEOPHYSCIA ENDOCOCCINA (Körb.) Moberg – on R,
B (1300–2400).
PHAEOPHYSCIA NIGRICANS (Flörke) Moberg – on Aln
(1300).
PHAEOPHYSCIA ORBICULARIS (Neck.) Moberg – on Ulm,
Aln, Cor, Que, Pin, B (1250–1500).
PHAEOPHYSCIA PUSILLOIDES (Zahlbr.) Essl. – on Sal
(1300).
PHAEORRHIZA NIMBOSA (Fr.) H. Mayrhofer & Poelt – on
S, B (1900, 2600–2950).
PHLYCTIS ARGENA (Spreng.) Flot. – on Abi (1450–
1500).
PHYSCIA ADSCENDENS (Fr.) H. Olivier – on Aln, Ma,
Sal, Pad, Abi (1300–2000).
PHYSCIA AIPOLIA (Ehrh. ex Humb.) Fürnr. – on R,
on Ulm, Aln, PopA, PopT, Fra, AceP, Cor, Sal,
BetP, BetL, Pad, Fag, Sor, Abi (1250–3850).
PHYSCIA CAESIA (Hoffm.) Fürnr. – on R (2400).
P HYSCIA DUBIA (Hoffm.) Lettau – on R and S
(2000–2200).
PHYSCIA STELLARIS (L.) Nyl. – on Mal, Cor, Aln, Sor,
BetL, Abi, Pin (1300–2000).
PHYSCIA TENELLA (Scop.) DC. – on Mal, Fag, BetL,
PopT, Abi (1300–2050)
P HYSCIA TRIBACIA (Ach.) Nyl. – on Mal, on R
(1300).
PHYSCONIA DISTORTA (With.) J.R. Laundon – on B,
on Ulm, Aln, PopA, PopT, Fra, AceP, Cor, Sal,
BetP, BetL, Pad, Fag, Sor (1250–1900).
PHYSCONIA ENTEROXANTHA (Nyl.) Poelt – on B (1500).
PHYSCONIA MUSCIGENA (Ach.) Poelt – on R, B (1350–
3850).
PHYSCONIA PERISIDIOSA (Erichsen) Moberg – on B,
Que, Fag, AceP, Pad (1250–1900).
PLACIDIUM LACHNEUM (Ach.) Breuss – on S (3080–
3150).
PLACYNTHIELLA DASAEA (Stirt.) Tønsberg – on Betp,
Abi (1300–1600).
PLACYNTHIELLA ICMALEA (Ach.) Coppins & P. James
– on L (1650–1900).
PLACYNTHIELLA ULIGINOSA (Schrad.) Coppins & P.
James – on B (3150).
PLATISMATIA GLAUCA (L.) W.L. Culb. & C.F. Culb. – on
Abi (1500–2300).
PLEOPSIDIUM FLAVUM (Bellardi) Körb. – on R (2500–
2600).
PLEUROSTICTA ACETABULUM (Neck.) Elix & Lumbsch
– on R (2000).
PORPIDIA CRUSTULATA (Ach.) Hertel & Knoph – on R
(1400–2000).
PORPIDIA MACROCARPA (DC.) Hertel & A.J. Schwab
– on R (2700, 3850).
PROTOPANNARIA PEZIZOIDES (Web.) P.M. Jørg. – on B,
S (1400–3100).
PROTOPARMELIA BADIA (Hoffm.) Hafellner – on R
(2800).
**PSEUDEPHEBE MINUSCULA (Nyl ex Arnold) Brodo &
D. Hawksw. – on R, B, S (2600–3000).
P SEUDEPHEBE PUBESCENS (L.) M. Choisy – on R
(2200–3000).
PSEUDEVERNIA FURFURACEA (L.) Zopf – on BetP, Aln,
Sor, Sal, Pin, Abi (1300–2200).
PSORA DECIPIENS (Hedw.) Hoffm. – on S (2850–
3080).
**P YCNORA PRAESTABILIS (Nyl.) Hafellner – on L
(1600–2000).
RAMALINA DILACERATA (Hoffm.) Hoffm. – on AceT
(1500).
RAMALINA FARINACEA (L.) Ach. – on R, on AceP, Sal,
Abi, Pic (1400–1600).
RAMALINA FASTIGIATA (Pers.) Ach. – on Abi, AceT
(1500).
R AMALINA FRAXINEA (L.) Ach. – on BetL, BetP
(1600–2000).
RAMALINA OBTUSATA (Arnold) Bitter – on Abi (1300–
1500).
RAMALINA POLLINARIA (Westr.) Ach. – on R, B, S, on
AceP, Fag (1250–1900).
RAMALINA POLYMORPHA (Lilj.) Ach. – on R (2300).
RAMALINA SINENSIS Jatta – on AceP, AceT, Sal, Cor,
Pad (1300–1900).
RAMALINA THRAUSTA (Ach.) Nyl. – on Abi (1500).
***RHIZOCARPON CARPATICUM Runem. – on R (2700–
3850).
RHIZOCARPON GEOGRAPHICUM (L.) DC. – on R (1560–
3850).
RHIZOCARPON MACROSPORUM Räsänen – on R (2500).
RHIZOCARPON OBSCURATUM (Ach.) A. Massal. – on R
(3150) (Novruzov & Onipchenko, 1985).
RHIZOCARPON POLYCARPUM (Hepp) Th. Fr. – on R
(1800–2750).
RHIZOCARPON VIRIDIATRUM (Wulfen) Körb. – on R
(2000) (Novruzov & Onipchenko, 1985).
RHIZOPLACA CHRYSOLEUCA (Sm.) Zopf – on R (2000–
2700).
RHIZOPLACA MELANOPHTHALMA (DC.) Leuckert & Poelt
– on R (2500–2700).
R INODINA EXIGUA (Ach.) Gray –on Pin, Pad, L
(1500–1900).
**R INODINA INTERPOLATA (Stirt.) Sheard – on R
(2400).
RINODINA LAEVIGATA (Ach.) Malme – on Abi (2000).
**RINODINA OLIVACEOBRUNNEA C.W. Dodge & G.E.
Baker – on plant remains (3150).
RINODINA PYRINA (Ach.) Arnold –on Abi, L (1900–
2000).
34
Folia Cryptog. Estonica
*RINODINA SEPTENTRIONALIS Malme – on Sal (1500–
2000)
*SCHAERERIA FUSCOCINEREA (Nyl.) Clauzade & Cl.
Roux – on R (2700).
SCOLICIOSPORUM CHLOROCOCCUM (Stenh.) Vĕzda – on
Mal, Sor, Abi, on L (1300–1600).
SCOLICIOSPORUM UMBRINUM (Ach.) Arnold – on Pin
(1600).
SOLORINA CROCEA (L.) Ach. – on S (2850–2950,
3850).
SOLORINA SACCATA (L.) Ach. – on S (2850–3150).
SQUAMARINA LENTIGERA (Weber) Poelt – on R (3150)
(Novrusov & Onipchenko, 1985).
STAUROTHELE FRUSTULENTA Vain. – on R (2050).
STEREOCAULON ALPINUM Laurer – on S, B (2050–
3160).
STEREOCAULON NANODES Tuck. – on R (3850).
STICTA SYLVATICA (Hudson) Ach. – on Abi (2000).
STRIGULA STIGMATELLA (Ach.) R.C. Harris – on Abi,
on L (2000)
T HAMNOLIA VER MICULARIS (Sw.) Schaer. – on S
(2750–2950).
**THELIDIUM PYRENOPHORUM (Ach.) Mudd – on R
(2200).
TONINIA SEDIFOLIA (Scop.) Timdal – Teberda (Jatta,
1900).
T RAPELIOPSIS GRANULOSA (Hoffm.) Lumbsch – on L,
B, S (1400–1900, 3000).
TUCKERMANNOPSIS CHLOROPHYLLA (Willd.) Hale – on
AceP, Abi, L (1450–1700).
UMBILICARIA CYLINDRICA (L.) Delise ex Duby – on R
(1500–3850).
UMBILICARIA DECUSSATA (Vill.) Zahlbr. – on R (2000–
3150).
UMBILICARIA DEUSTA (L.) Baumg. – on R (1600–
3150).
UMBILICARIA HIRSUTA (Sw. ex Westr.) Hoffm. – on R
(2500).
U MBILICARIA HYPERBOREA (Ach.) Hoffm. – on R
(2400–2700).
UMBILICARIA LEIOCARPA DC. – on R (2700).
UMBILICARIA NYLANDERIANA (Zahlbr.) H. Magn. – on
R (2000).
UMBILICARIA POLYPHYLLA (L.) Baumg. – on R (2700–
3000).
UMBILICARIA PROBOSCIDEA (L.) Schrad. – on R (2000–
2700).
UMBILICARIA SUBGLABRA (Nyl.) Frey – on R (2000).
U MBILICARIA VELLEA (L.) Hoffm. – on R (1300–
3000).
USNEA CAUCASICA Vain. – on Sor (1600–1700).
USNEA CAVERNOSA Tuck. – on Abi, Pin, AceP, L, B
(1300–2000).
**USNEA CHAETOPHORA Stirton – on Abi (1600).
USNEA FILIPENDULA Stirton – on AceP, L (1500–
1650).
USNEA FLORIDA (L.) F.H. Wigg. – on Que, BetL, Pin,
Abi, L (1350–2000).
USNEA FULVOREAGENS (Räsänen) Räsänen – on BetP,
Pin (1300–2000).
USNEA GLABRATA (Ach.) Vain. – on Abi, Sor, BetL, L
(1400–2050).
USNEA GLABRESCENS (Nyl. ex Vain.) Vain. – on Aln,
Pin, L (1500–2500).
USNEA LAPPONICA Vain. – on AceP, Sal, Pin (1500–
2200).
USNEA PROSTRATA Vain. – on AceP (1500).
USNEA SCABRATA Nyl. – on AceP, Abi, Pin (1500–
2000).
USNEA SUBFLORIDANA Stirt. – on L, BetL, Abi, Pin
(1400–2000).
VERRUCARIA DOLOSA Hepp – on R (2000).
VERRUCARIA FUNCKII (Spreng.) Zahlbr. – on R
(2050).
VERRUCARIA NIGRESCENS Pers. – on R (1300–3850).
VULPICIDA PINASTRI (Scop.) J.-E. Mattsson & M.J. Lai
– on L, Pin, Abi, BetL (1800–2200).
XANTHOPARMELIA CONSPERSA (Ehrh. ex Ach.) Hale – on
L, S, Pin (1300–2400).
XANTHOPARMELIA SOMLOËNSIS (Gyeln.) Hale – on L, B
(1300–3850).
**X ANTHORIA ECTANEOIDES (Nyl.) Zahlbr. – on R
(2200–2400).
XANTHORIA ELEGANS (Link) Th. Fr. – on R, B, S
(1250–2850).
X ANTHORIA FALLAX (Hepp) Arnold – on B, Que
(1300).
XANTHORIA PARIETINA (L.) Th. Fr. – on Aln (1300).
*XANTHORIA SOREDIATA (Vain.) Poelt – on R (2500–
2700).
XANTHORIA ULOPHYLLODES Räsänen – on R, Aln, Fra,
PopA, Pin (1250–1500).
XYLOGRAPHA PARALLELA (Ach.: Fr.) Behlen & Desberg
– on L (1900–2000).
**XYLOGRAPHA VITILIGO (Ach.) J.R. Laundon – on L
(1500–2000).
Folia Cryptog. Estonica, Fasc. 41: 35–44 (2005)
Lichens from Uummannaq, Qilakitsoq and Qaarsut, Central West
Greenland
Eric Steen Hansen
Botanical Museum, University of Copenhagen, Gothersgade 130, DK-1123 Copenhagen K, Denmark
E-mail: erich@bot.ku.dk
Abstract: A total of 177 taxa of lichens are reported from three localities in the Uummannaq area in Central West Greenland. Aspicilia myrinii is reported as new to Greenland. Northern extensions of 13 species in West Greenland are presented.
Geology, climate and vegetation of the localities are briefly treated.
Kokkuvõte: Kesk-lääne Gröönimaa (Uummannaq, Qilakitsog ja Qaarsut) samblikud.
Esitatakse Kesk-lääne Gröönimaa Uummannaqi piirkonna kolme kogumisala 177 samblikutaksoni leiuandmed. Aspicilia myrinii
on esmasleid Gröönimaalt. Andmetest nähtub 13 Lääne-Gröönimaa liigi ulatuslik levik põhja suunas. Lühidalt peatutakse
piirkonna geoloogial, kliimal ja taimkattel.
INTRODUCTION
The lichen flora of Uummannaq Ø and two additional lowland localities situated at the north
coast of Nuussuaq Peninsula, viz. Qilakitsoq and
Qaarsut (Fig. 1), was investigated by the author in July 1989 and July 2003. Uummannaq
covers an area of c. 12 square kilometres. The
town, Uummannaq, is located at the southern
point of the island. A 1170 m high mountain,
Hjertefjeldet, occupies the central part of the
island (Fig. 2). A small bay, Spraglebugt, occurs
at the western side of the island just north of
Uummannaq. All parts of the island except the
uppermost and steepest slopes of Hjertefjeldet
were studied by the author. Qilakitsoq and
Qaarsut are separated from Uummannaq by
the channel, Sarqarput. The distance between
Uummannaq and the two localities is 8 km and
16 km, respectively. The former, now abandoned
village, Qilakitsoq, is primarily known because
of the rare find of some well-preserved, more
than 500 year old mummies now deposited at
the Greenland national museum in Nuuk. The
author investigated the gneissic foreland near
the graves. Qaarsut is a settlement with c. 250
inhabitants. The author studied its surroundings, in particular the coastal lowland and the
lowest level of the mountains, which rise more
or less steeply to altitudes about 2000 m in the
central part of Nuussuaq.
J. Vahl visited the Uummannaq District in
1834 (Branth & Grønlund, 1888), E. Vanhöffen
in 1892 (Darbishire, 1897). The first publica-
tions list only about twenty species, most of
them very common and widely distributed
Greenland lichens. A more complete survey of
the lichen flora of Uummannaq has not been
published so far, but a number of recent publications contain informations about lichens collected in the area including the surroundings of
the previous mining town, Maarmorilik (Breuss
& Hansen, 1988; Hansen, 1990a, 1990b, 1991,
1995; Hansen & Poelt, 1987; Hansen et al.,
1987a, 1987b; Leuckert et al., 1987; Moberg &
Hansen, 1986; Thomson, 1997).
Localities and geology
The following three localities were investigated
by the author (Fig. 1).
1. Uummannaq. 70°41´N, 52°08´W. Alt.
0–200 m (Fig. 2). 15–19 July 1989, 27 July
1989 & 22–23 July 2003. Archaean gneiss
(”Uummannaq gneiss”) with layers of amphibolite (Escher & Stuart Watt, 1976).
2. Qilakitsoq. 70°36´N, 52°13´W. Alt. 0–100 m.
28 July 1989. Archaean gneiss.
3. Qaarsut. 70°44´N, 52°39´W. Alt. 0–300 m.
24–29 July 2003. Archaean gneiss and
scattered occurrences of amphibolite. The
locality is distincly influenced by Tertiary
basalts and Cretaceous sediments derived
from surrounding rock formations.
36
Folia Cryptog. Estonica
naq, whereas the mean temperature of the
coldest month, March, is –17°C according to
measurements made by Asiaq/Grønlands Forundersøgelser. The annual precipitation is 120
mm (2000).
MATERIAL AND METHODS
Fig. 1. Location of investigation area in Central
West Greenland. 1 – Uummannaq Ø; 2 – Qilakitsoq; 3 – Qaarsut. The small Greenland map
shows the situation of the investigation area.
Climate
The Uummannaq area has a low arctic and
oceanic climate. The mean temperature of
the warmest month, July, is 8°C at Uumman-
Lichens were collected at numerous sample
plots at the three localities situated in the Uummannaq area. The collected material, a total of
630 specimens of lichens, was studied with Zeiss
light microscopes. Selected specimens of Bryoria
and Stereocaulon were identified by means of
TLC. The material is deposited at the Botanical
Museum, University of Copenhagen (C).
RESULTS AND DISCUSSION
About 100 lichens have been reported from the
Uummannaq area prior to the present investigation by Hansen and different co-authors (see
Introduction). Floristically Uummannaq Ø and
Nuussuaq Peninsula are more similar to the
Fig. 2. Rocky hills just north of Uummannaq town. Hjertefjeldet (= Uummannaq Fjeld) with distinct
black bands of amphibolite is seen in the background.
37
northernmost part of Disko and Svartenhuk
Peninsula than to the inland area between the
two big peninsulas. Thus the surroundings of
Maarmorilik are richer in lichens with affinity
to more or less calciferous substrates than,
for example, Uummannaq Ø (Hansen, 1990a,
1991). Selected, mostly rare species of particular
interest reported from the investigation area, but
not found during the trips in 1989 and 2003,
are listed together with the references in the
following.
Caloplaca anchon-phoeniceon Poelt & Clauzade
(Hansen et al., 1987a)
Caloplaca cacuminum Poelt (Hansen et al.,
1987a)
Caloplaca celata Th. Fr. (Hansen et al., 1987a)
Caloplaca fulvolutea (Nyl.) Jatta (Hansen et al.,
1987a)
Caloplaca insularis Poelt (Hansen et al.,
1987a)
Caloplaca paulii Poelt (Hansen et al., 1987a)
Caloplaca saxifragarum Poelt (Hansen et al.,
1987a)
Caloplaca xanthostigmoidea (Räsänen) Zahlbr.
(syn. Caloplaca epiphyta Lynge) (Hansen et
al., 1987a)
Catapyrenium cinereum (Pers.) Körb. (Breuss &
Hansen, 1988)
Catapyrenium daedaleum (Kremp.) Stein.
(Breuss & Hansen, 1988)
Glypholecia scabra (Pers.) Müll. Arg. (Hansen &
Poelt, 1987)
Involucropyrenium waltheri (Kremp.) Breuss
[syn. Catapyrenium waltheri (Kremp.) Körb.]
(Hansen & Poelt, 1987)
Lecidea ileiformis Fr. (Hansen & Poelt, 1987)
Megaspora verrucosa (Ach.) Hafellner & Wirth
[syn. Pachyospora verrucosa (Ach.) A. Massal.] (Hansen, 1991)
Melanolecia transitoria (Arnold) Hertel (Hansen
& Poelt, 1987)
Placidium norvegicum (Breuss) Breuss (syn.
Catapyrenium norvegicum Breuss) (Breuss
& Hansen, 1988)
Phaeophyscia kairamoi (Vain.) Moberg (Moberg
& Hansen, 1986)
Psora globifera (Ach.) A. Massal. (Hansen,
1991)
Psora vallesiaca (Schaer.) Timdal (Hansen,
1991)
Sagiolechia protuberans (Ach.) A. Massal.
(Hansen et al., 1987)
Schadonia fecunda (Th. Fr.) Vězda & Poelt
(Hansen & Poelt, 1987)
General remarks on the lichen vegetation
Uummannaq Ø is characterized by a fairly
diverse terricolous lichen vegetation in spite
of the relatively simple geological conditions
prevailing on the island. Moist heath mosaics with dominating Cassiope tetragona,
Phyllodoce coerulea and Salix herbacea occur
in places with a moderate to prolonged snow
cover during winter. Cetrariella delisei, Cladonia
borealis, C. mitis, C. trassii, Cetraria islandica,
Ochrolechia frigida, Peltigera malacea, Psoroma
tenue and Stereocaulon alpinum are common
lichens in this snow-patch-like heath type. A
related, but somewhat drier heath vegetation
with Cassiope, Empetrum hermaphroditum
and Vaccinium uliginosum covers the thin soil
layer near Qilakitsoq. However, proper snowpatches dominated by Salix herbacea, Solorina
crocea and Pertusaria oculata also occur at this
locality. Bryoria nitidula, Cladonia bellidiflora,
C. cyanipes and C. sulphurina are among the
most typical lichens in this gneissic area.
Empetrum, Vaccinium and Salix glauca form a
lichen-rich community on many, more or less
exposed rocks on Uummannaq Ø. Alectoria
nigricans, Cetraria islandica, C. muricata and
Sphaerophorus globosus are commonly found
at these sites. Wind exposed rocks with a thin
snow cover in winter sometimes hold a dry
Dryas integrifolia-Vaccinium heath with
Alectoria nigricans, A. ochroleuca, Cetraria
muricata, Flavocetraria cucullata, F. nivalis and
Thamnolia vermicularis. These lichens also occur
abundantly in open places in the extensive Betula
nana heath at Spraglebugt and in a steppe-like
Carex rupestris-Artemisia borealis community
occurring on south-facing slopes on the island.
Caloplaca tiroliensis, Candelariella placodizans,
Cetraria muricata, Physconia muscigena and
Psora rubiformis are additional species occurring
in the last-mentioned community. Dry DryasCarex rupestris heaths with well-developed
Alectoria ochroleuca and Flavocetraria nivalis
occur commonly in the vicinity of Qaarsut.
Bryoria nitidula and Flavocetraria cucullata are
the dominant lichen species in mixed VacciniumDryas heaths around this settlement. Cassiope,
Empetrum, Betula nana and Rhododendron
lapponicum are occasional dwarf shrubs in the
heath mosaics occurring at Qaarsut. Their content of lichens is comparable with that of Uum-
38
Folia Cryptog. Estonica
mannaq. However, Protopannaria pezizoides was
not found at Uummannaq. At Qaarsut it grows
on moist soil in these dwarf shrub heaths. Some
moist depressions near Spraglebugt just north
of Uummannaq are covered by, for example,
Arctocetraria nigricascens, Arthrorhaphis alpina
and Lecanora geophila.
A few corticolous lichens, for example, Leptogium saturninum and Melanelia
septentrionalis, have previously been reported
from Maarmorilik (Alstrup, 1982; Hansen, 1991).
Lecanora fuscescens was collected at Qaarsut,
only. The comparatively cold climatic conditions
prevailing in the Uummannaq area evidently do
not favour the development of extensive scrubs
with a corticolous lichen flora of the type known
from more southern parts of Greenland.
Ornithocoprophilous saxicolous lichens
are of great importance at all of the three localities, indicating the very rich bird life of the
area. Guano of ravens and different sea birds,
in particular gulls, manures exposed rocks
and thus favours lichens such as Candelariella
vitellina, Dimelaena oreina, Lecidea atrobrunnea,
Phaeophyscia sciastra, Physcia caesia, P. dubia,
Rhizoplaca melanophthalma, Umbilicaria arctica,
U. decussata and Xanthoria elegans. Umbilicaria
decussata is sometimes the dominant lichen on
north- and east-facing rocks on Uummannaq
Ø, while Xanthoria elegans dominates in the
lichen communities on south-facing rocks. The
basal part of the rocks is occasionally covered
by Xanthoria sorediata. Umbilicaria vellea occurs
abundantly on overhanging rocks and in places
with percolating water. The lowest, west-facing
amphibolite band on Hjertefjeldet holds the following lichens: Dimelaena oreina, Physcia caesia,
Pseudephebe minuscula, Rhizocarpon geminatum,
Rhizoplaca melanophthalma, Umbilicaria
hyperborea, Xanthoria elegans (dominant) and
X. sorediata. A similar sociation (without
Dimelaena oreina and Xanthoria sorediata) occurs at Qilakitsoq. Here Usnea sphacelata was
found growing on a gneissic rock. The nitrophilous sociation is quite common at Qaarsut
(Fig. 3), but differs in the abundant occurrence
of Melanelia infumata. At this locality Xanthoria
elegans binds gravel in the same way as at, for
example, Qeqertarsuaq on Disko (Hansen, 1999).
Somewhat exposed, horizontal gneissic rock faces
Fig. 3. Top of bird rock at Qaarsut. The mouse-grey thalli of Umbilicaria decussata have
distinct reticulate ridges. They are surrounded by pale yellowish white thalli of Rhizoplaca
melanophthalma.
39
at Qaarsut support species such as Calvitimela
armeniaca, Dimelaena oreina, Sporastatia testudinea (infected by Rhizocarpon pusillum),
Umbilicaria decussata and U. lyngei.
Gneissic rocks are much wider distributed in
the Uummannaq area than rocks composed of
amphibolite and accordingly contain more lichen
species than the latter. Acarospora molybdina,
Caloplaca alcarum and Lecanora contractula
form a characteristic sociation on gneissic seashore rocks manured by, for example, gulls. It is
widely distributed in coastal areas in Greenland
(Hansen et al., 1987a). A south-facing gneissic
rock at the east side of Uummannaq Ø is covered
by Lecanora intricata, Rhizocarpon geminatum
(dominant, infected by Caloplaca castellana), Umbilicaria hyperborea, U. torrefacta and U. virginis.
A blackish lichen sociation with Arctoparmelia
incurva, Orphniospora moriopsis, Pseudephebe
minuscula and Umbilicaria hyperborea occurs
on gneissic rocks at a somewhat higher level
(alt. 100 m a.s.l.) in this area. Another dark coloured community occurring more or less commonly at Uummannaq consists of the following
lichens: Allantoparmelia alpicola, Arctoparmelia
incurva, Ophioparma ventosa, Orphniospora
moriopsis, Pseudephebe minuscula (dominant),
Rhizocarpon inarense, Sphaerophorus fragilis,
Umbilicaria hyperborea and U. lyngei. A horizontal rock just north of Uummannaq also holds
a greyish to black coloured association with
Melanelia hepatizon, Parmelia saxalis, Sphaerophorus fragilis, Umbilicaria hyperborea and
U. lyngei.
Annotated list of lichens
The following list of lichens is based on the
author´s collections. The list cannot be considered representative as regards genera such as
Aspicilia, Verrucaria and a number of lecideoid
microlichens, which have been neglected during
the present investigation. Nomenclature follows
Santesson et al. (2004) with some exceptions.
Numbers 1, 2, 3 indicate the three localities
listed above. Annotations are given as regards
the substrate of the lichens and presence of
apothecia (ap.) or perithecia (pe.); ”st.” means
that the specimens is sterile. The asterisk * in
front of the name indicates that the collection
represents a northern range extension of the
taxon in West Greenland. The frequency is
mentioned, where it was possible to estimate
it. Collections which have been distributed previously from herbarium C as part of ”Lichenes
Groenlandici Exsiccati” (LGE) are stated by their
numbers. Selected references are cited.
ACAROSPORA MOLYBDINA (Wahlenb.) A. Massal. – 1.
On gneissic seashore rocks; ap.
* A. PELISCYPHA (Th. Fr.) Arn. – 1. On gneissic rocks
manured by birds; ap.
A. RHIZOBOLA (Nyl.) Alstrup – 1, 3. On soil; ap.
* A. VERONENSIS A. Massal. – 2. On gneissic rocks;
ap.
ALECTORIA NIGRICANS (Ach.) Nyl. – 1, 2, 3. On soil;
st.
A. OCHROLEUCA (Hoffm.) A. Massal. – 1, 2, 3. On
soil; st. LGE 910.
ALLANTOPARMELIA ALPICOLA (Vain.) Essl. – 1, 2. On
gneissic rocks; st.
* AMANDINEA CONIOPS (Wahlenb.) M. Choisy ex
Scheid. & H. Mayrhofer – 1. On gneissic
rocks; ap.
ARCTOCETRARIA NIGRICASCENS (Nyl.) Kärnefelt & A.
Thell – 1. On soil; st. LGE 356.
ARCTOPARMELIA INCURVA (Pers.) Hale – 1, 2, 3. On
gneissic rocks; st.
A RCTOPELTIS THULEANA Poelt – 1. On gneissic
rocks manured by birds; ap. The species is
common on Disko and Nuusuaq Peninsula,
but very rare outside this area (Hansen,
1997).
ARTHRORHAPHIS ALPINA (Schaer.) R. Sant. – 1, 2, 3.
On soil; st.
A. CITRINELLA (Ach.) Poelt – 1. On soil; st.
A SPICILIA MASTRUCATA (Wahlenb.) Th. Fr. – 1,
2, 3. On rocks composed of gneiss and
amphibolite; st.
A. MYRINII (Fr.) Hafellner – 1. On gneissic rocks;
ap. New to Greenland.
* BAEOMYCES CARNEUS Flörke – 1, 3. On soil and
plant debris; st.
B RODOA OROARCTICA (Krog) Goward – 3. On
siliceous rocks; st.
BRYOCAULON DIVERGENS (Ach.) Kärnefelt – 1. On
soil; st.
BRYONORA CASTANEA (Hepp) Poelt – 1, 2, 3. On
mosses and plant debris; st.
BRYORIA CHALYBEIFORMIS (L.) Brodo & D. Hawksw.
– 1. On soil; st.
* B. LANESTRIS (Ach.) Brodo & D. Hawksw. – 3. On
gneissic rocks; st.; rare. Thallus contains
fumarprotocetraric acid (TLC).
B. NITIDULA (Th. Fr.) Brodo & D. Hawksw. – 1, 2,
3. On soil; st. LGE 360, 908.
40
Folia Cryptog. Estonica
BUELLIA PAPILLATA (Sommerf.) Tuck. – 1, 3. On
soil; ap.
CALOPLACA ALCARUM Poelt – 1, 3. On Lecanora
contractula on seashore rocks composed
of gneiss and amphibolite; rarely on old
bones; ap.
C. AMMIOSPILA (Wahlenb.) H. Olivier – 1, 2, 3. On
plant debris, mosses and soil; ap.
C. CASTELLANA (Räsänen) Poelt – 1, 3. On
Placynthium asperellum and Rhizocarpum
geminatum on gneissic rocks; ap.
C. CERINA (Ehrh. ex Hedw.) Th. Fr. – 1, 2, 3. On
mosses, plant debris and old bones; ap.
C. CITRINA (Hoffm.) Th. Fr. – On old bones; ap.
Previously reported from a few localities in
South-, West- and North Greenland (Hansen
et al., 1987a).
C. EPITHALLINA Lynge – 1, 3. On Dimelaena oreina
and Rhizoplaca melanophthalma on gneissic
rocks; ap.
C. FRAUDANS (Th. Fr.) H. Olivier – 2. On gneissic
rocks; ap.
C. JUNGERMANNIAE (Vahl) Th. Fr. – 1, 2. On plant
debris; ap.
C. TETRASPORA (Nyl.) H. Olivier – 1, 3. On plant
debris; ap.
C. TIROLIENSIS Zahlbr. – 1, 2, 3. On mosses and
plant debris; ap.; common.
C. TOMINII L.I. Savicz – 3. On calciferous soil; st.
CALVITIMELA ARMENIACA (DC.) Hafellner – 1, 3. On
gneissic rocks; ap.
CANDELARIELLA AURELLA (Hoffm.) Zahlbr. – 3. On
old bone; ap.
C. DISPERSA (Räsänen) Hakul. – 3. On Placynthium
asperellum on gneissic rocks; st.
C. PLACODIZANS (Nyl.) H. Magn. – 1, 2, 3. On soil;
ap.
C. TERRIGENA Räsänen – 1. On soil rich in humus;
st.
C. VITELLINA (Hoffm.) Müll. Arg. – 1, 3. On gneissic
rocks; ap.
C. XANTHOSTIGMA (Ach.) Lettau – 1. On plant debris;
st.; rare.
CETRARIA ISLANDICA (L.) Ach. – 1, 2, 3. On soil; st.
LGE 359.
C. MURICATA (Ach.) Eckfeldt – 1, 2, 3. On soil;
st.
CETRARIELLA DELISEI (Bory ex Schaer.) Kärnefelt &
A. Thell – 1, 2, 3. ; st.; common. LGE 368.
CHAENOTHECA FURFURACEA (L.) Tibell – 2. On plant
debris and mosses; st.
CLADONIA AMAUROCRAEA (Flörke) Schaer. – 1, 2, 3.
On soil rich in humus; st.; common.
C. BELLIDIFLORA (Ach.) Schaer. – 1, 2. On soil and
plant debris; ap.
C. BOREALIS S. Stenroos – 1, 2, 3. On soil rich in
humus; ap.
C. CARIOSA (Ach.) Spreng. – 1. On soil; ap.
C. CHLOROPHAEA (Flörke ex Sommerf.) Spreng. – 1,
2, 3. On soil and mosses; st.
C. CORNUTA (L.) Hoffm. – 1, 2. On soil rich in
humus and on mosses; st.
C. CYANIPES (Sommerf.) Nyl. – 2. On soil rich in
humus; st.
C. GRACILIS (L.) Willd. – 1, 2. On soil rich in
humus; ap.
C. LUTEOALBA Wheldon & A. Wilson – 2. On plant
debris; st.
C. MACROCERAS (Delise) Hav. – 1, 3. On soil; st.
C. MACROPHYLLA (Schaer.) Stenh. – 2. On soil rich
in humus; ap.
* C. MACROPHYLLODES Nyl. – 1, 2. On soil and
mosses; st.
C. MITIS Sandst. – 1, 2, 3. On soil; st.; common.
C. PHYLLOPHORA Hoffm. – 1. On soil rich in humus;
st.
C. PLEUROTA (Flörke) Schaer. – 1, 2. On soil rich
in humus and on plant debris; st.
C. POCILLUM (Ach.) Grognot – 1, 3. On soil; st.
C. PYXIDATA (L.) Hoffm. – 1, 2, 3. On soil rich in
humus; st.; common.
C. STYGIA (Fr.) Ruoss – 2. Among mosses on soil;
st.
C. SULPHURINA (Michx.) Fr. – 1, 2. On soil rich in
humus; st.
* C. TRASSII Ahti – 1, 2. On soil rich in humus;
ap.
COLLEMA UNDULATUM Laurer ex Flot. var. GRANULOSUM
Degel. – 1. On calciferous soil; st.
DACTYLINA RAMULOSA (Hook.) Tuck. – 3. On soil,
mosses and plant debris; st. LGE 909
DIMELAENA OREINA (Ach.) Norman – 1, 2, 3. On
rocks composed of gneiss and amphibolite;
ap.
* EUOPSIS PULVINATA (Schaer.) Vain. – 1. On gravel;
ap.
FLAVOCETRARIA CUCULLATA (Bellardi) Kärnefelt & A.
Thell – 1, 2, 3. On soil; st.; common. LGE
907.
F. NIVALIS (L.) Kärnefelt & A. Thell – 1, 2, 3. On
soil; st.; common. LGE 906.
FULGENSIA BRACTEATA (Hoffm.) Räsänen – 3. On
plant debris and mosses on calciferous
soil; ap.; rare.
FUSCOPANNARIA PRAETERMISSA (Nyl.) P.M. Jørg. – 1.
On mosses on calciferous soil; st.
41
HYPOGYMNIA AUSTERODES (Nyl.) Räsänen – 1, 3. On
plant debris and mosses; st.
H. SUBOBSCURA (Vain.) Poelt – 3. On plant debris;
st.
LECANORA ARGOPHOLIS (Ach.) Ach. – 1, 3. On rocks
composed of gneiss and amphibolite; ap.
L. CENISIA Ach. – 1. On gneissic rocks; ap.
L. CHLOROLEPROSA (Vain.) H. Magn. – 1, 2. On
gneissic rocks; st.
L. CONTRACTULA Nyl. – 1, 2. On seashore rocks
composed of gneiss and amphibolite; rarely
on old bones; ap.
L. EPIBRYON (Ach.) Ach. – 1, 3. On soil, mosses
and plant debris; ap.
L. FUSCESCENS (Sommerf.) Nyl. – 3. On dead twigs
of Salix glauca; ap.
L. GEOPHILA (Th. Fr.) Poelt – 1. On soil rich in
humus; st.
L. HAGENII (Ach.) Ach. var. FALLAX Hepp. – 1, 2, 3.
On plant debris and old bone; ap.
L. INTRICATA (Ach.) Ach. – 1. On gneissic rocks;
ap.
L. POLYTROPA (Ehrh. ex Hoffm.) Rabenh. – 1,
2, 3. On rocks composed of gneiss and
amphibolite; ap.; common.
LECIDEA ATROBRUNNEA (Ramond ex Lam. & DC.)
Schaer. – 1, 3. On gneissic rocks; ap.
L. LAPICIDA (Ach.) Ach. var. PANTHERINA Ach. – 3. On
gneissic rocks; ap.
L. TESSELLATA Flörke – 1, 3. On gneissic rocks;
ap.
LECIDELLA WULFENII (Hepp) Körb. – 1. On soil rich
in humus; ap.
LEPROCAULON SUBALBICANS (I.M. Lamb) I.M. Lamb &
A.M. Ward – 1, 2, 3. On mosses.
LICHENOMPHALIA HUDSONIANA (H.S. Jenn.) Redhead
et al. – 2. On mosses.
LOPADIUM CORALLOIDEUM (Nyl.) Lynge – 3. On soil;
ap.
MELANELIA DISJUNCTA (Erichsen) Essl. – 1, 2, 3. On
gneissic rocks; st.
M. HEPATIZON (Ach.) A. Thell – 1, 3. On gneissic
rocks; st.
M. INFUMATA (Nyl.) Essl. – 1, 3. On gneissic rocks
manured by birds; rarely on twigs (Salix
glauca); st. LGE 905.
MICAREA ASSIMILATA (Nyl.) Coppins – 1. On soil;
ap.
MIRIQUIDICA ATROFULVA (Sommerf.) A.J. Schwab &
Rambold – 1, 2. On gneissic rocks rich in
iron minerals; st.
M. GAROVAGLII (Schaer.) Hertel & Rambold – 1, 3.
On gneissic rocks; st.
M. NIGROLEPROSA (Vain.) Hertel & Rambold – 1. On
gneissic rocks; st.
MYXOBILIMBIA LOBULATA (Sommerf.) Hafellner – 1,
3. On soil and mosses; ap.
OCHROLECHIA FRIGIDA (Sw.) Lynge – 1, 2, 3. On soil,
plant debris and mosses; ap.; common.
O. GRIMMIAE L ynge – 1, 2. On Racomitrium
lanuginosum; ap.
* O. INAEQUATULA (Nyl.) Zahlbr. – 1. On mosses;
st.
O. LAPUËNSIS (Vain.) Räsänen – 1, 2, 3. On plant
debris; st.
O PHIOPARMA VENTOSA (L.) Norman – 1, 2. On
gneissic rocks; ap.
ORPHNIOSPORA MORIOPSIS (A. Massal.) D. Hawksw.
– 1, 2. On gneissic rocks; ap.
P ARMELIA OMPHALODES (L.) Ach. – 1, 2, 3. On
mosses; st.
P. SAXATILIS (L.) Ach. – 1, 2, 3. On gneissic rocks
and mosses; st.
P. SULCATA Taylor – 2. On gneissic rocks; st.
PARMELIELLA TRIPTOPHYLLA (Ach.) Müll. Arg. – 3. On
mosses; st.
PELTIGERA DIDACTYLA (With.) J.R. Laundon – 1,
2. On soil and mosses; st. The species is
infected by Illosporium carneum.
P. LEPIDOPHORA (Nyl. ex Vain.) Bitter – 3. On soil;
st.; rare.
P. LEUCOPHLEBIA (Nyl.) Gyeln. – 2. On mosses; st.
P. MALACEA (Ach.) Funck – 1, 2, 3. On soil and
mosses; st.
P. RUFESCENS (Weiss) Humb. – 1, 2, 3. On soil and
mosses; st.
P. SCABROSA Th. Fr. – 1, 2, 3. On soil and mosses;
st.
PERTUSARIA CORIACEA (Th. Fr.) Th. Fr. – 1, 3. On
soil, mosses and plant debris; st.
P. DACTYLINA (Ach.) Nyl. – 2. On soil; st.
P. GEMINIPARA (Th. Fr.) C. Knight ex Brodo – 1, 2.
On mosses and plant debris; st.
P. OCULATA (Dicks.) Th. Fr. – 1, 2, 3. On soil and
mosses; ap.
PHAEOPHYSCIA CONSTIPATA (Norrl. & Nyl.) Moberg – 1.
On mosses and soil; st.; rare.
P. ENDOCOCCINA (Körb.) Moberg – 1. On gneissic
rocks and on mosses; ap.
P. SCIASTRA (Ach.) Moberg – 1, 3. On gneissic rocks
and on mosses; ap.
PHYSCIA CAESIA (Hoffm.) Fürnr. – 1, 2, 3. On
gneissic rocks and old bones; st.
P. DUBIA (Hoffm.) Lettau – 1, 3. On gneissic rocks
and old bones; st.
42
Folia Cryptog. Estonica
* P. TENELLA (Scop.) DC. – 1, 3. On gneissic rocks
and dead twigs; st.; rare.
PHYSCONIA MUSCIGENA (Ach.) Poelt – 1, 2, 3. On soil,
mosses, planr debris and gneissic rocks; ap.;
common.
PLACIDIUM LACHNEUM (Ach.) de Lesd. – 1, 3. On
calciferous soil and mosses; pe.
PLACYNTHIUM ASPERELLUM (Ach.) Trevis – 1, 2, 3. On
gneissic rocks; ap.
PLEOPSIDIUM CHLOROPHANUM (Wahlenb.) Zopf – 1, 2,
3. On rocks composed af amphibolite and
gneiss; ap.
* POLYCHIDIUM MUSCICOLA (Sw.) Gray – 1. On plant
debris; st.
PORPIDIA FLAVOCAERULESCENS (Hornem.) Hertel &
A.J. Schwab – 2, 3. On gneissic rocks; ap.
PROTOPANNARIA PEZIZOIDES (Weber) P.M. Jørg. & S.
Ekman – 3. On soil; ap.
PROTOPARMELIA BADIA (Hoffm.) Hafellner – 1, 3. On
gneissic rocksmanured by birds; ap.
PSEUDEPHEBE MINUSCULA (Nyl. ex Arnold) Brodo &
D. Hawksw. – 1, 2, 3. On rocks and stones
composed of amphibolite and gneiss; st.;
common.
P. PUBESCENS (L.) M. Choisy – 2. On gneissic
rocks; rare.
PSORA RUBIFORMIS (Ach.) Hook – 1, 3. On calciferous
soil and mosses; ap.
PSOROMA TENUE Henssen var. BOREALE Henssen
– 1, 2, 3. On soil, mosses and plant debris;
ap. Recently reported as overlooked, but
widespread, arctic-alpine lichen (Jørgensen,
2004). The taxon is common in more
northern parts of Greenland.
RHIZOCARPON BOLANDERI (Tuck.) Herre – 1. On
gneissic rocks; st.; rare.
R. DISPORUM (Nägeli ex Hepp) Müll. Arg. – 1. On
gneissic rocks; ap.
R. EUPETRAEOIDES (Nyl.) Blomb. & Forssell – 1. On
gneissic rocks; ap.
R. GEMINATUM Körb. – 1, 3. On rocks composed of
amphibolite and gneiss; ap. LGE 902.
R. GEOGRAPHICUM (L.) DC. – 1, 3. On gneissic
rocks; ap.
R. GRANDE (Flörke) Arnold – 1. On gneissic rocks;
ap.
R. INARENSE (Vain.) Vain. – 1, 2, 3. On gneissic
rocks; ap.; common.
* R. JEMTLANDICUM (Malme) Malme – 1, 3. On rocks
composed of amphibolite and gneiss; ap.
R. PRAEBADIUM (Nyl.) Zahlbr. – 1. On gneissic
rocks; ap.
R.
Runemark – 1, 3. On Sporastatia
testudinea on gneissic rocks; ap.
R. SUPERFICIALE (Schaer.) Vain. – 1, 3. On gneissic
rocks; ap.
RHIZOPLACA MELANOPHTHALMA (DC.) Leuckert & Poelt
– 1, 3. On gneissic rocks manured by birds;
also on rocks composed of amphibolite;
ap.
RINODINA TURFACEA (Wahlenb.) Körb – 1, 2. On soil
and plant debris; ap.
SOLORINA BISPORA Nyl. – 1. On calciferous soil;
ap.
S. CROCEA (L.) Ach. – 2. On soil; ap.
S. SPONGIOSA (Ach.) Anzi – 3. On calciferous soil;
ap.
S PHAEROPHORUS FRAGILIS (L.) Pers. – 1, 2. On
gravelly soil and gneissic rocks; st.
S. GLOBOSUS (Huds.) Vain. – 1, 2. On soil and
mosses; st.
SPORASTATIA POLYSPORA (Nyl.) Grummann – 1, 3.
On gneissic rocks; ap.
S. TESTUDINEA (Ach.) A. Massal. – 1, 3. On rocks
composed of amphibolite and gneiss; ap.
STEREOCAULON ALPINUM Laurer – 1, 2, 3. Among
mosses on soil; ap.; common.
* S. ARCTICUM Lynge – 1, 2. On soil; st. LGE
355.Thallus contains atranorin and stictic
acid (TLC)
S. ARENARIUM (L.I. Savicz) I.M. Lamb – 1, 2, 3. On
mosses or less gravelly soil; st.; common.
S. BOTRYOSUM Ach. – 2. On gneissic rocks; st.
S. GLAREOSUM (L.I. Savicz) H. Magn. – 1, 2. On
soil; st.
T H A M N O L I A V E R M I C U L A R I S (Sw.) Schaer. var.
SUBULIFORMIS (Ehrh.) Schaer. – 1, 2, 3. On
soil and mosses; common. LGE 911.
T REMOLECIA ATRATA (Ach.) Hertel – 1, 2, 3. On
gneissic rocks; ap.
UMBILICARIA ARCTICA (Ach.) Nyl. – 1, 2, 3. On gneissic
rocks manured by birds; ap.; common.
U. DECUSSATA (Vill.) Zahlbr. – 1, 2. On bird rocks
composed of amphibolite and gneiss; ap.
LGE 354, 904.
U. HYPERBOREA (Ach.) Hoffm. – 1, 2, 3. On rocks
composed of amphibolite and gneiss; ap.;
common.
U. LYNGEI Schol. – 1, 2, 3. On gneissic rocks; ap.;
common.
U. PROBOSCIDEA (L.) Schrad. – 1, 2, 3. On gneissic
rocks; ap.
U. TORREFACTA (Lightf.) Schrad. – 1. On gneissic
rocks; st.
PUSILLUM
43
U.
VELLEA
U.
VIRGINIS
ap.
ap.
(L.) Hoffm. – 1, 2. On gneissic rocks;
Schaer. – 1, 2, 3. On gneissic rocks;
USNEA SPHACELATA R. Br. – 2. On gneissic rocks;
st.; rare.
XANTHORIA BOREALIS R. Sant. & Poelt – 1, 3. On
gneissic rocks manured by birds; rarely on
soil and mosses; ap.
X. ELEGANS (Link) Th. Fr. – 1, 2, 3. On bird rocks
composed of amphibolite and gneiss, rarely
on mosses and old bones; ap.; common. LGE
903.
X. SOREDIATA (Vain.) Poelt – 1, 3. On vertical faces
of gneissic rocks manured by birds; st.
ACKNOWLEDGEMENTS
I wish to thank J. & N. Jakobsen (Nuuk), M.
Pihlsbech (Qaarsut) and A. Stephensen (Uummannaq) for their hospitality during my trips to
Central West Greenland. Thanks are also due
to L. Saag (Tartu) for help with identification of
selected leprarioid microlichens and S. Christensen for assistance with TLC. The investigation
was financially supported by the Commission for
Scientific Research in Greenland (KVUG) and the
Botanical Museum, University of Copenhagen.
REFERENCES
Alstrup, V. 1982. The epiphytic lichens of Greenland.
Bryologist 85: 64–73.
Branth, J. S. D. & Grønlund, C. 1888. Grønlands
lichen-flora. Meddelelser om Grønland 3: 449–
513.
Breuss, O. & Hansen, E. S. 1988. The lichen genera
Catapyrenium and Placidiopsis in Greenland.
Plant Systematics and Evolution 159: 95–105.
Darbishire, O. V. 1897. Flechten aus dem
Umanakdistrikt. Bibliotheca Botanica 42:
53–61.
Escher, A. & Stuart Watt, W. 1976. Geology of
Greenland. The Geological Survey of Greenland.
Copenhagen. 603 pp.
Hansen, E. S. 1990a. Uummannaqs lavflora oplevet
under en naturvandring. Forskning/Tusaat 2/90:
25–28.
Hansen, E. S. 1990b. Qilakitsoq, Maarmorilik
og Upernavik – set med en botanikers øjne.
Tidsskriftet Grønland 4/90: 101–112.
Hansen, E. S. 1991. The lichen flora near a leadzinc mine at Maarmorilik in West Greenland.
Lichenologist 23(4): 381–391.
Hansen, E. S. 1995. Greenland lichens. Atuagkat,
Rhodos and Danish Polar Center. Copenhagen.
124 pp.
Hansen, E. S. 1999. Lichens from Qeqertarsuaq/
Godhavn, Disko, Central West Greenland. Folia
Cryptog. Estonica 39: 3–12.
Hansen, E. S. & Poelt, J. 1987. New or interesting
Greenland lichens IV. Mycotaxon 30: 69–80.
Hansen, E. S., Poelt, J. & Søchting, U. 1987a.
Die Flechtengattung Caloplaca in Grönland.
Meddelelser om Grønland, Bioscience 25: 1–52.
Hansen, E. S., Poelt, J. & Vězda, A. 1987b. The lichen genera Gyalecta, Gyalidea and Sagiolechia
in Greenland. Herzogia 7: 367–374.
Jørgensen, P. M. 2004. Psoroma tenue var. boreale,
an overlooked, widespread, arctic-alpine lichen.
Graphis Scripta 15(1/2): 60–64.
Leuckert, C., Schirrmeister, M. & Hansen, E. S. 1987.
Chemosystematical investigation of lichens from
Greenland. Dimelaena oreina (Ach.) Norm. Bot.
Jahrb. Syst. 108: 373–379.
Moberg, R. & Hansen, E. S. 1986. The lichen genus Physcia and allied genera in Greenland.
Meddelelser om Grønland, Bioscience 22: 1–32.
Santesson, R., Moberg, R., Nordin, A., Tønsberg, T. &
Vitikainen, O. 2004. Lichen-forming and lichenicolous fungi of Fennoscandia. Museum of Evolution, Uppsala University. Uppsala. 359 pp.
Thomson, J.W. 1997. American arctic lichens. II. The
Microlichens. The University of Wisconsin Press.
Wisconsin. 675 pp.
44
Folia Cryptog. Estonica
Folia Cryptog. Estonica, Fasc. 41: 45–54 (2005)
Influence of environmental factors on the local-scale distribution
of cyanobacterial lichens:
case study in the North Urals, Russia
Irina Mikhailova1, Marina Trubina1, Eugene Vorobeichik1 & Christoph Scheidegger2
Institute of Plant and Animal Ecology UD RAS, 8 Marta St. 202, 620144 Ekaterinburg, Russia
E-mail: mikhailova@ipae.uran.ru
2
WSL Swiss Federal Research Institute, CH-8903 Birmensdorf, Switzerland
1
Abstract: Distribution and frequency of 11 cyanobacterial lichen species on Populus tremula trunks were studied in the
mountain taiga of the North Urals. Within an area of 150×200 m, all aspen trunks were registered, and a range of environmental variables (including characteristics of tree stand, herbs, dwarf-shrubs and mosses; 30 variables in total) were noted
within a circle of 6 m diameter around each tree. This study suggests the environmental variables to be of small importance
for spatial distribution of the lichens on a local scale. However, the abundance of the species was shown to depend on size
and vitality of their host trees, reaching maximal values in oldest damaged trees.
Kokkuvõte: Keskkonnategurite mõju tsüanobaktereid sisaldavate samblike levikule lokaalskaalas: pilootprojekt Põhja-Uuralitest, Venemaalt.
Põhja-Uuralite mägitaigas uuriti 11 tsüanobaktereid sisaldava samblikuliigi levikut ja sagedust haava (Populus tremula) tüvedel.
150×200 m alal registreeriti kõik haavatüved, ja 6 m läbimõõduga ringides ümber iga puu märgiti üles rida keskonna muutujaid (sealhulgas puistu karakteristikud, taimed, puhmad ja samblad; kokku 30 muutujat). Uurimusest tuleneb, et samblike
ruumilise leviku jaoks lokaalskaalas on keskkonna muutujad vähese tähtsusega. Ometi sõltus liikide ohtrus peremeespuude
suurusest ja elujõulisusest, saavutades suurima väärtuse vanimatel vigastatud puudel.
INTRODUCTION
Cyanobacterial lichens are a rather specific
group of lichens from both physiological and ecological points of view. Many of them are highly
sensitive to any kind of ecosystem disturbances
and are suggested as being indicators of ecological continuity, or old-growth forest indicators
(Gauslaa, 1994, Kuusinen, 1996, Sillett & McCune, 1998, etc.). Among cyanolichens, there
is a range of rare and endangered species included to Red lists of many European countries.
Use of cyanolichens as biological indicators as
well as necessity of their conservation require
better understanding of environmental factors
influencing their distribution in different spatial
scales, from a forest patch up to the species
distribution area as a whole.
Factors determining distribution of lichen
species at different spatial levels can be very
different, for example, climate parameters, altitude, and regional air quality are responsible
for the lichen distribution at regional scale, and
forest vegetation parameters are of the highest
importance at subregional spatial scale (WillWolf et al., 2004).
Our investigations were carried out in one
of few old-growth forests left in the North Urals.
Continental climate of the Ural Mountains together with high level of industrialization of the
region does not advantage high diversity and
abundance of epiphytic cyanolichens. Hot spots
of cyanolichen diversity can be found in mountain taiga under conditions of higher humidity
and low level of any kind of human activity
though even these habitats are unfavorable for
cyanolichens. Under these conditions, all relationships between lichens and environment are
expected to be more pronounced. Our study was
aimed to assess the importance of environmental factors for the distribution and frequency of
cyanolichens on the local scale (few hectares
within more or less homogeneous tree stand).
MATERIAL AND METHODS
The study was carried out in the mountain taiga
of the North Ural near the village of Kytlym (Sverdlovsk region, 59º28’N, 59º14’E). The climate is
moderately continental, with an annual aver-
46
Folia Cryptog. Estonica
age precipitation of 492 mm. The mean annual
temperature is –0.2°C; mean January and July
temperatures are –16.4°C and 13.9°C, respectively. The duration of the frost-free period is 96
days (Reference book..., 1965, 1968).
The area belongs to the middle taiga
phytogeographical subzone (Kolesnikov et al.,
1973). The sample plot was established in the
lower part (460 m a.s.l.) of the western slope of
Pervyi Bugor mountain (931 m a.s.l.). The tree
stand was composed by Picea obovata, Abies
sibirica, Pinus sibirica, Populus tremula, and
Betula pubescens with an average age of 120
years. The soil was well-drained, deep and rocky,
and was classified as brown forest mountain soil
(Firsova, 1977). In the understorey vegetation,
Vaccinium myrtillus, Oxalis acetosella,
Calamagrostis obtusata, Pleurozium schreberi,
and Hylocomium splendens dominated.
Populus tremula was selected as a phorophyte for studying cyanolichens because of
high water capacity of its bark (Barkman, 1958)
which is advantageous for cyanolichens. Within
an area of 150×200 m, all aspen trees with diameter at breast height (DBH) of at least 30 cm were
registered (182 trees). On all registered trunks,
the frequencies of cyanolichens were recorded
using a sample grid of 10 units adapted to fit
half the circumference of each trunk (Herzig &
Urech, 1991). The grid was placed sequentially
on the basal (0–0.5 m above ground) halves of
the trunks and on those at 1.0–1.5 m above
ground. Thus, species frequency on the trunk
ranged from 0 to 40. For species growing out of
quadrates, only presence was recorded.
The following parameters were noted for
each tree: DBH, class of vitality [1 – healthy, 2
– damaged (parasitic fungi or mechanical damages), 3 – dead], and inclination (measured with
an inclinometer). Age of 45 healthy trees was determined with an increment borer. Obtaining of
accurate age estimates for damaged aspens was
problematic because of decaying of heartwood.
A range of environmental variables was registered within a circle of 6 m diameter around
each tree: 1) tree stand characteristics (basal
area of coniferous and deciduous trees, total
basal area, and crown parameters); 2) density of
young growth in three height classes: < 50 cm,
50–100 cm, and >100 cm; 3) density of shrubs
and trees of undergrowth in three height classes:
< 50 cm, 50–100 cm, and >100 cm; 4) characteristics of understorey vegetation [abundance
and number of herb and dwarfshrub species,
total cover of bryophytes, total cover of herbs
and dwarf-shrubs, average height of lowest and
tallest herb and dwarf-shrub layers, the share of
three dominant species (Calamagrostis obtusata,
Oxalis acetosella, and Vaccinium myrtillus)].
To estimate canopy cover above sample
trees, 8 digital pictures of crowns were taken
around each tree: at 4 cardinal points in circles
of 0.5 m and 1 m radius around the tree (the
camera was placed at the 1 m height from the
ground). The percentage of open sky area (ratio
of number of pixels that fall on the open sky
to the total number of pixels) and the index of
crown density (ratio of number of pixels that
fall on the boundary sky/crown to the number
of pixels that fall on the crown) were calculated
in color images using SIAMS Photolab software
(developed by SIAMS, Ekaterinburg, http://
siams.com).
To determine ecological indices of the sites
(indices of insolation, moistening, soil acidity,
continentality, soil fertility, and thermal regime),
Ellenberg’s indicator values scale (Ellenberg et
al., 1991) was used. Each species of the herb and
dwarf-shrub layer was referred to one of ecological groups. Indices were calculated as weighted
averages of Ellenberg’s indicator values.
To determine bark acidity, samples of upper
layer (1–2 mm) were collected from four cardinal points of 80 model aspen trees at 1–1.5
m height. Model trees were selected to allow
for the balanced model of 2-way ANOVA (tree
vitality and tree DBH as independent factors).
In the laboratory, bark samples were cleaned
from bryophytes and lichens under dissecting
microscope and were ground with a laboratory
mill. One gram of the powder was suspended
in 25 ml of distillated water in stoppered vials,
placed to a shaker for 30 min and then left for 30
min. After this, pH was measured with standard
glass electrode without filtration.
RESULTS AND DISCUSSION
Sample tree and site characteristics
Sample aspens varied significantly in their size
(Table 1). Age of model healthy trees ranged
between 53 and 130 years. Pearson correlation
coefficient between age and DBH was 0.66 at
p<0.05; the mean DBH corresponds to the age of
about 80 years. The age of largest damaged trees
47
Table 1. Some characteristics of the sample plot
Variables
DBH of aspen trees (cm)
Total basal area of tree stand (m2/ha)
% open sky: 0.5 m from the trunk
% open sky: 1 m from the trunk
Index of insolation
Index of moistening
Bryophyte cover (%)
Cover of herb and dwarf-shrub layer (%)
Average bark pH per trunk
Minimal pH value per trunk
Maximal pH value per trunk
Mean
80.57
41.42
0.22
0.24
5.07
5.60
55.57
68.57
6.10
5.69
6.49
can probably exceed 150 years. Healthy trees
prevailed among aspens; 40.7% of aspens had
different signs of damage (fruit bodies of fungi,
mainly Phellinus tremula, mechanical injuries,
frost splits, etc.), and only one tree was dead.
Bark pH was found to vary in great extent
both between and within sample trees. Range
within the same trunk sometimes reached 2.2
units, so we had to treat minimal, maximal and
average pH values for each trunk as separate
variables (Table 1). Two-way ANOVA (SS III, DBH
and vitality as fixed factors) revealed significant
contribution of DBH to the variability of maximal
pH values (F = 5.5, df=4, 63, p<0.01); bark pH
smoothly rose with the increase of tree size (Fig.
SE
1.88
1.47
<0.01
<0.01
0.03
0.01
1.68
0.79
0.05
0.07
0.04
CV (%)
31.54
48.07
26.33
25.34
8.62
3.53
40.99
15.57
7.62
11.49
5.79
Min
32.50
0.00
0.07
0.10
3.60
5.03
10.00
40.00
4.40
4.19
4.82
Max
145.00
108.61
0.48
0.47
5.88
6.05
95.00
86.00
6.77
6.64
7.09
1). Bark of damaged trees tended to be slightly
more acid (difference was not significant, F =
2.57, df=1, 63, p=0.11). For the variability of
minimal pH values, DBH was not found to be an
important factor; tree vitality, on the contrary,
contributed significantly (F = 4.4, df=1, 63,
p<0.05). Acidification of the bark of damaged
trees might probably be explained by activity
of parasitic fungi secreting acidic substances
though this question needs special investigations. High variability of pH values within the
same trunk is most probably due to “dripzone
effect” (Goward & Arsenault, 2000). For example, acid leachates from neighbouring coniferous trees can cause acidification of aspen bark
Fig. 1. Maximal values of bark pH (from 4 measurements) vs aspen DBH (means ± SE). White bars
– healthy trees, dotted bars – damaged trees.
48
Folia Cryptog. Estonica
which in turn leads to the inoculation of aspens
by acidophytic lichens that are generally not
typical for this phorophyte, such as Hypogymnia
physodes and Vulpicida pinastri. Generally, the
range of pH values at the sample plot (4.19–7.09)
corresponds to values found for aspen trunks in
middle and southern boreal forests of Finland
(Kuusinen, 1994).
Mean values of the total basal area and
crown cover indicate rather close tree stand.
Basal area was the most variable among environmental parameters registered; crown characteristics were more stable (Table 1). Cover of bryophytes also varied in a great extent indicating
differences in soil moistening. Though variation
coefficients of ecological indices were low, actual
range of ecological factors can be rather high:
for example, minimal found value of insolation
index (3.6) corresponds to shadowed habitats
while maximal value (5.88) falls between halfshadowed and half-opened habitats (Ellenberg
et al., 1991). Thus, within the more or less homogeneous local tree stand, some heterogeneity
of environmental factors exists which can be of
importance for distribution and abundance of
epiphytic lichens.
Cyanolichen species composition
A total of 11 cyanolichen species were found
on the 182 aspens sampled (Table 2). Most of
them, especially Nephroma species and redlisted species Lobaria pulmonaria, have been
often mentioned as indicators of old-growth
forests in Eurasia and North America (Rose,
1976, Kuusinen, 1996, Kondratyuk et al.,
1998, Sillett & McCune, 1998, etc.). At 60.18%
of investigated aspen trunks, at least one cyanolichen species was found. However, the share of
trunks inoculated by individual species proved
to be very low and in most cases did not exceed
10%. Leptogium intermedium, which was found
on a single aspen tree, was excluded from the
further analysis.
Frequency of cyanolichens vs site characteristics
Factor analysis was applied to reduce the total
variation of measured environmental variables
to three independent factors that were interpreted respectively as site insolation, soil fertility
and canopy openness (Table 3). Interpretation
of factor 1 and factor 3 looks at the first glance
Table 2. List of species under study and their
frequencies
Species
Collema subflaccidum Degel.
Leptogium intermedium
(Arnold) Arnold
L. saturninum (Dicks.) Nyl.
Lobaria pulmonaria (L.)
Hoffm.
Nephroma parile (Ach.) Ach.
N. resupinatum (L.) Ach.
Pannaria pezizoides (Weber)
Trevis.
Peltigera aphthosa (L.) Willd.
P. canina (L.) Willd.
P. polydactylon (Neck.)
Hoffm.
P. praetextata (Sommerf.)
Zopf
No of
findings
5
1
% of
P. tremula
trunks
colonized
2.74
0.55
12
17
6.59
9.34
18
16
10
9.89
8.79
5.49
6
24
6
3.30
13.19
3.30
5
2.76
very similar: both of them reflect light conditions. The difference seems to be in the following: factor 1 includes parameters of grass and
dwarf-shrub layer thus reflecting light regime
over the 6 m diameter sample plot while factor
3 includes mainly the crown parameters which
were registered within 1 m from the trunk and
hence reflects insolation of the sample trunk.
Factor loadings of parameters of the host trees,
undergrowth trees and young growth were negligible. Three extracted factors explained 45% of
the total variance.
At the next step of analysis, Spearman rank
order correlation coefficients were calculated
between lichen frequencies and the factors extracted (Table 4). Peltigera species and Collema
subflaccidum did not show any dependency from
the factors extracted; for the rest of species, significant correlation with at least one factor was
revealed. Frequency of four cyanolichen species
negatively correlated with site insolation (though
correlation was extremely weak). The most probable, this is not the low site insolation which
favors development of cyanolichens, but accompanying higher ambient humidity. Besides,
49
Table 3. Factor loadings of site and phorophyte variables (marked with * loadings are >0.5)
Variables
Factor 1
Site insolation
Factor 2
Soil fertility
Factor 3
Canopy openness
-0.24
-0.23
-0.31
0.00
-0.06
-0.02
-0.33
-0.06
-0.03
Basal area of coniferous -0.29
trees
Basal area of decidious 0.35
trees
Total basal area
0.02
% open sky:
0.5 m from the trunk
0.17
1 m from the trunk
0.10
Crown density:
0.5 m from the trunk
-0.04
1 m from the trunk
-0.04
0.11
-0.40
0.03
-0.41
0.11
-0.62*
0.04
0.05
0.81*
0.80*
-0.23
-0.26
-0.68*
-0.68*
height < 50 cm
height 50–100 cm
height >100 cm
-0.13
-0.28
-0.46
0.35
0.13
0.02
-0.10
0.11
0.30
height < 50 cm
height 50–100 cm
height >100 cm
0.19
0.19
-0.16
-0.22
0.00
-0.03
-0.15
0.28
0.17
0.64*
0.31
0.77*
0.25
-0.58*
0.14
0.33
0.32
-0.02
0.15
Phorophyte data
DBH
Vitality
Inclination
Tree stand characteristics
Density of young growth
Density of shrubs and trees of undergrowth
Understorey vegetation
Abundance of herb and 0.30
dwarf-shrub species
Number of herb and dwarf- 0.24
shrub species
Cover of bryophytes
-0.07
Cover of herb and dwarf- 0.67*
shrub layer
Mean height of the lowest 0.69*
herb layer
Mean height of the tallest 0.72*
herb layer
The share of:
Calamagrostis obtusata
0.66*
Oxalis acetosella
-0.69*
Vaccinium myrtillus
-0.07
0.34
0.11
-0.12
0.52*
-0.89*
-0.07
-0.17
-0.12
Index of insolation
Index of moistening
Index of soil acidity
Index of continentality
Index of soil fertility
Index of thermal regime
% of variation explained
-0.40
0.09
0.66*
-0.76*
0.90*
-0.15
16
0.07
0.30
0.05
0.01
0.14
-0.20
13
Site indices (based on Ellenberg’s indicator values scale)
0.82*
0.73*
0.61*
0.28
0.01
0.06
16
50
Folia Cryptog. Estonica
Table 4. Spearman rank order correlation coefficients for correlation between lichen frequencies
and environmental data (* – p<0.05, ** – p<0.01,
*** – p<0.001)
Species
Collema subflaccidum
Factor 1
Site
insolation
0.04
Factor 2
Soil
fertility
0.09
Factor 3
Canopy
openness
-0.11
Leptogium saturninum
-0.23 **
0.04
-0.16 *
Lobaria pulmonaria
-0.19 **
0.12
-0.10
Nephroma parile
-0.16 *
0.10
-0.06
N.resupinatum
-0.28 *** 0.06
-0.09
Pannaria pezizoides
-0.04
0.15 * -0.10
Peltigera aphthosa
0.106
-0.003
0.074
P. canina
-0.111
0.062
-0.002
P. polydactylon
0.013
0.026
0.008
P. praetextata
-0.021
0.075
-0.090
site insolation has been reflecting history of the
plot: for instance, high insolation means previously happened tree-fall which was followed by
rapid development of light-demanding forbs and
deciduous young growth. Epiphytic lichens inoculated the sample trunk at least tens of years
ago, so looks probable that lichen occurrence
correlates not with present-day but with previous-day environment.
Rather unexpected was the absence of correlation of lichen frequency with canopy openness (excluding for Leptogium saturninum), i.e.
with light regime of the microhabitat. As in the
case of site insolation, the effect of trunk insolation hardly can be separated from that of other
factors, especially, from water relations. Crown
parameters significantly affect amount of water
passing down as stemflow. Epiphytic gelatinous
cyanolichens often grow in rain-tracks; however,
presence and pattern of rain-tracks can not be
predicted from our data on crown cover and
density.
Multiple linear stepwise regression analysis
(ridge regression) showed that environmental
variables accounted for a very low (<20%) part
of variability of cyanolichen species frequency
(Table 5). For different species, from 6 to 15 site
variables were included to the model; so extracting of one or two major factors was not possible.
Integral parameters (number of cyanolichen species per trunk and total frequency of cyanolichens) were found to be “more predictable” on
the base of collected site characteristics.
Discriminant analysis was applied to data
on presence/absence of lichen species (without
quantitative information on frequency). The results confirmed the conclusion of the multiple
regression analysis: frequency of correct predictions of presence/absence of lichen species was
extremely low and ranged from 16 to 66%.
Table 5. Results of ridge step-wise regression analysis (df1 is a number of variables included to
the model, df2 is df Error)
Collema subflaccidum
Leptogium saturninum
Lobaria pulmonaria
Nephroma parile
N. resupinatum
Pannaria pezizoides
Peltigera aphthosa
P. canina
P. polydactylon
P. praetextata
Peltigera spp. (total frequency)
Total frequency of cyanolichens
No of cyanolichen species
Adjusted R2
0.036
0.140
0.151
0.052
0.180
0.119
0.046
0.139
0.033
0.063
0.097
0.214
0.345
df1
7
12
11
6
12
9
6
9
7
7
7
15
13
df2
179
174
175
180
174
177
174
171
173
173
173
171
173
F
1.99
3.52
4.01
2.71
4.40
3.79
2.45
4.24
1.89
2.72
3.77
4.38
8.52
p
0.059
<0.001
<0.001
0.015
<0.001
0.001
0.027
<0.001
0.074
0.010
<0.001
<0.001
<0.001
51
Thus, predictive power of collected environmental data is very low. This means that at the
few hectares spatial scale heterogeneity of site
characteristics seems not to influence significantly the occurrence of cyanolichens.
Relationship between phorophyte parameters and frequency of cyanolichen species
The dependency of lichen community composition on such trunk parameters as size (i.e. tree
age), inclination and vitality is of wide knowledge
(Barkman, 1958; Kalgutkar & Bird, 1968; Bates,
1992, etc.). However, as indicated above, phorophyte parameters showed very weak correlation
with factors extracted, and their influence on
lichen occurrence was not adequately analysed.
Because of this, two-way ANOVA was used to
estimate contribution of tree DBH and vitality
to the overall variability of lichen frequency [5
groups of DBH × 2 vitality classes (healthy and
damaged)]. To assess influence of tree inclina-
tion, one-way ANOVA was used (number of observations did not allow three-way ANOVA).
For Lobaria pulmonaria, Nephroma
resupinatum and Peltigera aphthosa DBH
was found to contribute significantly to the
variability of frequencies (Table 6). Frequency
of L. pulmonaria and N. resupinatum was higher
on larger trunks (Fig. 2). Tree vitality was also of
high importance for these species, especially for
N. resupinatum, which was found on damaged
trees only. Combination of the factors (DBH +
vitality) was highly significant for three species
mentioned above, and also for Nephroma parile
and for the total frequency of Peltigera species.
Most cyanolichen species had a similar pattern of response to phorophyte parameters: their
frequency increased with an increase of tree size
and with lowering of its vitality thus reaching
maximal values on the largest damaged trunks.
The only exception was Peltigera species (Fig. 2)
that showed maximal frequency on small-sized
Table 6. Results of ANOVA (SS III) for the lichen frequencies (df Error= 170 in all cases)
Species
Collema subflaccidum
Lobaria pulmonaria
Leptogium saturninum
Nephroma parile
N. resupinatum
Pannaria pezizoides
Peltigera aphthosa
P. canina
P. polydactylon
P. praetextata
Peltigera spp. (total
frequency)
Sources of variation
DBH
Tree vitality
df=4
df=1
F
F
p
p
1.59
1.91
0.179
0.169
5.68
7.63
<0.001
0.006
1.30
3.86
0.271
0.051
1.86
2.69
0.120
0.103
3.72
8.46
0.006
0.004
1.29
1.59
0.275
0.210
2.57
1.35
0.040
0.246
1.53
0.99
0.197
0.322
0.52
0.61
0.719
0.436
0.44
0.82
0.782
0.366
0.65
0.06
0.626
0.806
DBH + vitality
df=4
F
p
0.12
0.883
4.22
0.016
1.80
0.169
1.46
0.235
7.57
0.001
1.08
0.343
6.52
0.002
0.67
0.511
2.05
0.131
0.97
0.380
1.10
0.334
Tree inclination
df=2
F
p
1.56
0.186
5.65
<0.001
1.01
0.406
2.47
0.046
3.79
0.006
1.34
0.258
3.80
0.005
2.01
0.095
1.45
0.219
0.44
0.782
2.96
0.021
52
Folia Cryptog. Estonica
Fig. 2. Frequencies of cyanolichens vs DBH of host trees (means ± SE). Axes: X – DBH (cm), Y
– lichen frequency. White bars – healthy trees, dotted bars – damaged trees.
53
damaged trunks. Making comparisons with
successional stages, we may suggest different
patterns of epiphytic successions on bases of
healthy and damaged trunks. We suppose that
reason for this difference can be in relationships
between Peltigera species and characteristics of
successively changing epiphytic bryophyte communities. This relationship can include, first,
direct moss-lichen competition and, second,
indirect influence of bryophytes on Peltigera
species through changes in the water regime
of microhabitat.
Frequencies of three species (Lobaria
pulmonaria, Nephroma resupinatum, and
Peltigera aphthosa) were influenced by inclination
of trunks: the highest frequency was found on
trunks with inclination more than 15º.
Thus, preferred habitats for the majority of
studied cyanolichens are large trees that were
shown to have highest values of bark pH. However, we failed to reveal statistically significant
correlation between frequency of cyanolichens
and bark pH. The only exception was Nephroma
resupinatum which showed significant correlation with minimal pH value per trunk (Spearman
rank order correlation coefficient –0.26, p <0.05).
Thus, pH values, at least in the range found in
this study, hardly can be regarded as an important factor for the growth of cyanolichens.
CONCLUSIONS
This study suggests low probability to predict
both presence and frequency of cyanolichens
from the data on the range of environmental
variables within small (few hectares) relatively
homogeneous tree stand. This is in contrast to
results on larger scales where authors usually
manage to reveal major factors influencing lichen species distribution (forest type, macroclimate, air pollution, etc.). Spatial scale of our
study causes relatively low range of variability of
environmental factors, and within this range the
factors seem not to be limiting for cyanolichens
(except for the size of host tree). Impossibility to
extract major factors that are responsible for the
lichen distribution causes low predictive power
of collected variables. Besides, low frequency of
studied cyanolichen species makes the obtaining
of statistically significant results very difficult.
Another reason of failure to find clear dependency between lichen distribution and site
characteristics is in not taking in the account re-
productive potential of lichens studied (number
of propagules produced, distance of propagule
dispersion, success of survival, etc.). However,
biological features like limited dispersion ability,
which was mentioned for most of lichen species
studied (Scheidegger, 1995; Zoller et al., 2000),
can be of high importance in defining patterns
of lichen distribution within a small area and
can mask significance of ecological factors.
Simply stated, lack of propagules can cause
absence of lichens even in the most favorable
environment.
ACKNOWLEDGMENTS
We are grateful to Yu. Paivina, A. Kharin, and
I. Sharounova for their help in the field works,
O. Mezhevikina for the measuring of bark pH,
and A. Ermakov for the analysis of digital images of canopy cover. The study was funded
by Swiss National Science Foundation (Project
7SUPJ62353)
REFERENCES
Barkman, J. J. 1958. Phytosociology and ecology
of cryptogamic epiphytes. Van Gorcum, Assen.
628 pp.
Bates, J. W. 1992. Influence of chemical and physical factors on Quercus and Fraxinus epiphytes at
Loch Sunart, western Scotland: a multivariant
analysis. J. Ecol. 80: 163–179.
Ellenberg, H., Weber, H. E., Dull, R., Wirth, R., Werner,
W. & Paulissen, D. 1991. Zeigerwerte von Pflanzen
in Mitteleuropa. Scripta Geobotanica 9: 1–122.
Firsova, V. P. 1977. Soils of taiga zone of the Urals (in
Russian). Nauka, Moscow. 176 pp.
Gauslaa, Y. 1994. Lungenever, Lobaria pulmonaria,
som indikator på artsrike kontinuitetsskoger.
Blyttia 52: 119–128.
Goward, T. & Arsenault, A. 2000. Cyanolichen distribution in young unmanaged forests: a dripzone
effect? Bryologist 103: 28–37.
Herzig, R.& Urech, M. 1991. Flechten als Bioindikatoren. Integriertes biologisches Messsystemder
Luftverschmutzung für das Schweizer Mittelland.
Biblioth. Lichenol. 43: 1–283.
Kalgutkar, R. M. & Bird, C. D. 1968. Lichens found on
Larix lyallii and Pinus albicaulis in southwestern
Alberta, Canada. Can. J. Bot. 47: 627–648.
Kolesnikov, B. P., Zubareva, R. S. & Smolonogov, E. P.
1973. Conditions for the forest growth and forest
types of Sverdlovsk region (in Russian). UNC AN
SSSR, Sverdlovsk. 175 pp.
Kondratyuk, S. Ya., Coppins, B. J., Zelenko, S. D.,
Khodosovtsev, A. Ye., Coppins, A. M. & Wolseley, P. A. 1998. Lobarion lichens as indicators of
54
Folia Cryptog. Estonica
primeval forests in the Ukrainian part of the proposed trilateral reserve ‘Eastern Carpathians’. In:
Kondratyuk, S. Ya & Coppins, B. J. (eds) Lobarion
lichens as indicators of the primeval forests of the
Eastern Carpathians, pp. 64–79. Phytosociocentre, Kiev.
Kuusinen, M. 1994. Epiphytic lichen flora and diversity on Populus tremula in old-growth and
managed forests of southern and middle boreal
Finland. Ann. Bot. Fenn. 31: 245–260.
Kuusinen, M. 1996. Cyanobacterial macrolichens on
Populus tremula as indicators of forest continuity
in Finland. Biological Conservation 75: 43–49.
Rose, F. 1976. Lichenological indicatrs of age and environmental continuity in woodlands. In: Brown,
D. H., Hawksworth, D. L. & Bailey, R. H. (eds)
Lichenology: Progress and Problems, pp. 279–307.
Academic Press, London & New York.
Scheidegger, C. 1995. Early development of transplanted isidioid soredia of Lobaria pulmonaria
in an endangered population. Lichenologist 27:
361–374.
Sillett, S. C. & McCune, B. 1998. Survival and growth
of cyanolichen transplants in douglas-fir forest
canopies. The Bryologist 101: 20–31.
Reference book on the climate of USSR, 9(2) (in Russian). 1965. Gidrometeoizdat, Leningrad. 362
pp.
Reference book on the climate of USSR 9(4) (in Russian).
1968. Gidrometeoizdat, Leningrad, 373 pp.
Will-Wolf, S., Geiser, L. H. & Neitlich, P. 2004. Comparison of lichen community response to environmental variables at regional and subregional
geographic scales. In: Randlane, T. & Saag, A.
(eds) IAL5 book of abstracts: Lichens in Focus, pp.
60–61. Tartu University Press, Tartu.
Zoller, S., Frey, B. & Scheidegger, C. 2000. Juvenile
development and diaspore survival in the threatened epiphytic lichen species Sticta fuliginosa,
Leptogium saturninum and Menegazzia terebrata: conclusions for in-situ conservation. In: The
Fourth IAL Symposium, Progress and Problems in
Lichenology at the Turn of the Millennium, p. 127.
Universitat de Barcelona, Barcelona.
http://siams.com
Folia Cryptog. Estonica, Fasc. 41: 55–58 (2005)
Lichens from the Khorasan Province, Iran
Mahroo Hadji Moniry1, F. Fallahian2 & A. Maassoumi3
Biology Department, Islamic Azad University, Mashhad, Iran
Biology Department, Science & ResearcheCampus, Islamic Azad University, Tehran, Iran
3
Botany Department, Researches Institute of Forests & Rangelands, Tehran, Iran
1
2
Abstract: Lichens were investigated in 13 localities of the Khorasan province (NE Iran). 39 infrageneric taxa in 22 genera
of 14 families were identified. Two genera and nine species are new to Iran, 21 species are new to the province.
Kokkuvõte: Iraani Khorasani provintsi samblikud.
Khorasani provintsis (Iraani kirde-osas) koguti 13 kohas samblikke. Määrati 39 liiki ja liigisisest taksonit 14 sugukonna 22
perekonnast. 2 perekonda ja 9 liiki on uued Iraani, 21 liiki Khorasani provintsi jaoks.
INTRODUCTION
Iran, with a surface area of ca 1,645,000 km2,
is located in the southwest of Asia between
25˚–40˚N. The knowledge of Iranian lichens is
still very scarce, which also produces a deep gap
in our understanding of Iranian vegetation. In
comparison with 275 published Floras of flowering plants in different parts of Iran, not even an
exhaustive catalogue is available for lichens. So
far, the most comprehensive reference to Iranian
lichens is a survey of 248 species compiled by
Szatala (1957). The present paper, devoted to the
lichens of the Khorasan province, is a contribution to a better knowledge of the lichen flora of
this large and ecologically diverse country.
MATERIALS AND METHODS
Survey area
The Khorasan province is located in the northeast of Iran, between 30˚21’–38˚17’N and 55˚28’–
61˚30’E (Fig. 1). This area is transitional between
the Siberian region and the desert area. Four climate types can be distinguished: cold-montane;
temperate-montane; temperate semi-desert and
hot-dry desert. The survey area is located in two
different geological zones called Alborz and EastSoutheastern, with expansion over 800 miles
from north to south, and over 200 miles from
east to west. Khorasan is distinguished by the
presence of regular and symmetrical mountain
ranges, the fold structures occurring as successive rows of ridges, each row being provided
with local names; altitude ranges between 300
and more than 3000 m (Parsa, 1978).
TURKMENISTAN
Caspian Sea
IRAN
IRAQ
KHORASAN
AFGANISTAN
Persian
Gulf
Fig. 1. The location of the Khorasan province
in Iran.
Materials and identification of lichens
825 samples of lichens were collected by the
first author from different parts of the Khorasan
province in 2002–2003. Due to the inadequacy
of literature and absence of reference material
concerning Iran, only part of them could be
identified. The original samples are stored in
FUMH, duplicates in B.
56
Folia Cryptog. Estonica
Identifications were made using some keybooks, separate papers and websites: Giordani
et al. (2003), Goward et al. (1994), Heidmarsson
(2000), Magnusson (1929), Mc Cune (2002),
Nash et al. (2002), Nimis & Martellos (2004),
Poelt & Vezda (1981), Poelt & Wirth (1968),
Purvis et al. (1992), Szatala (1957), and Wetmore
(2003). Morphology and chemistry were studied
using standard microscopes and the current
reagents (Purvis et al., 1992).
RESULTS
Two varieties and 37 species in 22 genera and
14 families were identified from the Khorasan
province. The genera Farnoldia Hertel and
Placidium A. Massal. are new to Iran. 10 infrageneric taxa are new to Iran, 22 taxa are new to
the Khorasan province.
Annotated list of the lichens found in the
Khorasan province, Iran
Numbers indicate voucher specimens deposited in FUMH. Species marked with * are new
to Iran, and those marked with ** are new to
Khorasan.
ACAROSPORA CERVINA A. Massal. – 1436c.
**A. STAPFIANA (Müll. Arg.) Hue – 1429a.
**A. STRIGATA (Nyl.) Jatta – 1519.
*ANAPTYCHIA ROEMERI Poelt – 1506a.
A. ULOTRICHOIDES (Vain.) Vain. – 1463.
**ASPICILIA CANDIDA (Anzi) Hue – 1408.
**CALOPLACA ALOCIZA (A. Massal.) Mig. – 1497a.
*C. CHALYBEIA (Fr.) Müll. Arg. – 1496a.
**C. TRANSCASPICA (Nyl.) Zahlbr. – 1420.
CANDELARIELLA AURELLA (Hoffm.) Zahlbr. – 1497b.
*COLLEMA AURIFORME (With.) Coppins & J.R. Laundon – 1435a.
**DERMATOCARPON MINIATUM (L.) W. Mann – 1432a.
*D. M I N I A T U M var. C I R S O D E S (Ach.) Zahlbr.
– 1470a
**D. MINIATUM var. COMPLICATUM (Lightf.) Hellb.
– 1470a.
**DIPLOTOMMA VENUSTUM Körb. – 1496a.
**ENDOCARPON PUSILLUM Hedw. – 1433b.
*FARNOLDIA JURANA (Schaer.) Hertel – 1528a.
*FULGENSIA SUBBRACTEATA (Nyl.) Poelt – 1434a.
**GLYPHOLECIA SCABRA (Pers.) Müll. Arg. – 1455.
LECANORA AGARDHIANA Ach. – 1534.
**L. CRENULATA Hook.– 1420.
**L. MARGINATA (Schaer.) Hertel & Rambold
– 1532b.
**L. MURALIS (Schreb.) Rabenh. – 1469a.
*NEOFUSCELIA PERRUGATA (Nyl.) Elix – 1436a.
*PLACIDIUM PILOSELLUM (Breuss) Breuss – 1523.
**PSORA DECIPIENS (Hedw.) Hoffm. – 1433a.
RHIZOCARPON GEMINATUM Körb. – 1436a.
**R. GEOGRAPHICUM (L.) DC. – 1436a.
**R. LECANORINUM Anders – 1663.
**R. VIRIDIATRUM (Wulfen) Körb.; (11); 1436c.
**RHIZOPLACA MELANOPHTALMA (Ramond) Leuckert
& Poelt – 1471.
RINODINA BISCHOFFII (Hepp) A. Massal. – 1522.
**S QUAMARINA CARTILAGINEA (With.) P. James
– 1653.
**S. LENTIGERA (Weber) Poelt – 1537.
**TONINIA CANDIDA (Weber.) Th. Fr. – 1492.
*T. DIFFRACTA (A. Massal.) Zahlbr. – 1453a.
*T. PHYSAROIDES (Opiz) Zahlbr. – 1480a
**T. SEDIFOLIA (Scop.) Timdal. – 1480a.
XANTHORIA ELEGANS (Link.) Th. Fr. – 1491.
ACKNOWLEDGMENTS
We are most grateful to Dr. H. J. M Sipman
and Prof. M. R. D. Seaward for support with
identifications.
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57
Nimis, P. L. & Martellos, S. 2004. Keys to the lichens of
Italy I. Terricolous lichens. Edizioni Goliardiche,
341 pp.
Parsa, A. 1978. Flora of Iran, Vol I. Offset Press Inc.,
Tehran–Iran, 506 pp.
Poelt, J. & Vezda, A. 1981. Bestimmungschluessel
Europaeischer Flechten. Ergaenzungsheft II.
Cramer, 400 pp.
Poelt, J. & Wirth, V. 1968. Flechten aus dem nordoestlichen Afghanistan gesammelt von H. Roemer im Rahmen der deutschen Wakhan Expedition
1964. Mitt. Bot. München 2: 219–261.
Purvis, W. O., Coppins, B. J., Hawksworth, D. L.,
James, P. W., Moore, D. M. 1992. The Lichen
Flora of Great Britain and Ireland. The British
Lichen Society, 710 pp.
Szatala, O., 1957. Prodromus einer Fletchenflora des
Irans. Ann. Historico-Naturales Musei Nationali
Hungarici 8: 101–154.
Wetmore, C. 2003. Key to lobed Caloplaca. URL: http:
//www.tc.umn.edu/~Wetmore/Gasp.html.
58
Folia Cryptog. Estonica
Folia Cryptog. Estonica, Fasc. 41: 59–80 (2005)
Lichens in the Red Data Books in Russia
Eugenia E. Moutchnik1 & Alexei A. Zavarzin2
All-Russia Institute of Nature Protection, “Znamenskie Sadki” estate, M-628, 113628 Moscow, Russia;
E-mail: eugenia@lichenfield.com
2
Department of Botany, St.Petersburg State University, Universitetskaya emb. 7/9, 199034 St.Petersburg, Russia;
E-mail: baltic@teia.org
1
Abstract: Legal aspects of lichen conservation through red-listing in Russia are described in the paper. A compilation of
the list of officially protected lichen taxa in Russia is made for the first time. It includes 29 species included in the Red Data
Book of the Russian Federation and 368 currently accepted taxa (species, subspecies, and varieties) from regional Red Data
Books. Progress and problems of redlisting as a basis for lichen conservation in the Russian regions are discussed.
Kokkuvõte: Samblikud Venemaa Punases Raamatus
Kirjeldatakse samblike kaitse õiguslikke aspekte, mis avalduvad Venemaa punaste nimekirjade koostamise kaudu. Esmakordselt
on kokku liidetud Venemaal ametlikult kaitstavate samblike loendid. Saadud nimestik sisaldab 29 liiki Vene Föderatsiooni
Punasest Raamatust ja 368 taksonit (liiki, alamliiki, varieteeti) regionaalsetest punastest raamatutest. Arutletakse punaste
nimekirjade ja samblike kaitse aluste üle.
INTRODUCTION
Lichen conservation is a complex scientific and
environmental task aimed at preserving the diversity of lichen taxa. Red Lists is an important
political tool and attract people’s attention to
threatened organisms. Red-listed species are
often monitored and inventoried. This is also
true for the Russian Federation (as it was for
the former Soviet Union). Red Data Books of
different levels are compiled here in order to
express the concern of the scientific community
for the fate of certain taxa and propose necessary
actions to preserve these elements of biological
diversity. However, the redlisting process in
Russia differs from that accepted in most other
countries. The reason behind this difference is
that Russian Red Lists normally function as
legal documents naming the species to be protected either in the whole country (in the case
of the Red Data Book of the Russian Federation)
or at the regional level (by means of the Red
lists and/or the Red Data Books of regions of
the Russian Federation). Thus the Red Lists in
Russia are a combination of scientific paper and
legal act. The latter allows to regulate protection
of rare species and to ensure preservation of rare
species’ habitats (Zavarzin et al., 2003).
Redlisting is regulated by the Federal Law
“On the Red Data Book of the Russian Federation” or by the regional laws or regulations on
regional Red Data Books. Usually the authorized
group of good standing researchers, who submit their proposals to the responsible agencies,
performs drafting of national and regional Red
lists. The agencies (Federal Ministry of Natural
Resources or respective regional authorities)
evaluate the proposals before approving the
lists. After the lists are approved they are to
be adopted either by the Government of Russia (in case of Federal Red Data Book) or by
the regional administration (in case of regional
Red Data Book or Red List). When the lists are
adopted they are published in a form of the Red
Data Book with description of species, their biology, distribution in the area, limiting factors
and necessary conservation measures. The laws
usually set the period of up to 10 years for the
Red lists to be revised. It is also required that the
Red list is distributed over to state institution,
private stakeholders and public as soon as the
list is adopted, however, electronic forms of the
Red lists are still not considered to be valid.
In practice the redlisted species are protected by the government, and their populations or
individuals as well as their habitats must not be
directly or indirectly destroyed. If the destruction
happens the responsible person or organization
is a subject for different administrative penalties.
Furthermore, registered populations of redlisted
species are an important requirement for establishing of protected area or introduction of
restrictions on economic use of the certain area.
Redlisted species are always priority objects for
environmental monitoring and important part
of every Environmental Impact Assessment.
60
Folia Cryptog. Estonica
However, generally low knowledge of redlisted
species, absence of readily available guidebooks
for species identification as well as week law
enforcement in Russia do not allow the easy use
of the legal opportunities presented by the Red
lists. This is especially true for such groups of
organisms as invertebrates, bryophytes, algae,
and lichens.
MATERIAL AND METHODS
This paper is based on analysis of literature data
– published Red Data Books and Red Lists of
the whole Russian Federation (Red Data Book
of RSFSR, 1988) as well as of its separate regions. Respective publications containing data
about lichens for the following regions (Fig. 1)
were used:
1. Central Federal District: Belgorod region (Moutchnik, 2003), Lipetsk region
(Moutchnik, 2003), Moscow region (Zubakin & Tikhomirov, 1998), City of Moscow
(Samojlov & Morozov, 2001), Ryazan region
(Kazakova, 2002), Smolensk region (Kruglov,
1997), Tambov region (Usova et al., 2002),
Tver region (Sorokin, 2002).
2. North-Western Federal District: Arkhangelsk region and Nenets autonomous district
(Andreev, 1995), Republic of Karelia (Ivanter & Kuznetsov, 1995), Republic of Komi
(Taskaev, 1998), Leningrad region (Tsvelev,
2000), Murmansk region (Konstantinova,
Koryakin, Makarova, 2003 ).
3. Northern Caucasus Federal District:
Republic of Adygeja (Kozmenko, 2000), Republic of Dagestan (Abdurakhmanov, 1998),
Republic of Kabardino-Balkaria (Ivanov,
2000), Krasnodar region (Nagalevsky, 1994),
Republic of Northern Osetia-Alania (Komzha
et al. 1999).
4. Privolzhsky Federal District: Republic of
Bashkortostan (Solomesha, 2002), Kirov
region (Dobrinsky, Korytin, 2001), Perm
region and Komi-Perm autonomous district
(Bolshakov & Gorchakovsky, 19961), Saratov region (Michurin & Shlyakhtin, 1996),
Republic of Tatarstan (Shepovskikh, 1995),
Republic of Udmurtia (Tuganaev, 2001).
5. Urals Federal District: Khanty-Mansy autonomous district (Vasin, 2003), YamaloNenets autonomous district (Dobrinsky,
1997), Sverdlovsk region (Bolshakov &
Gorchakovsky, 1996).
6. Siberian Federal District: Republic of Altaj
(Krasnoborov & Sidelnikov, 1996), Republic
of Buriat (Bojkov, 2002), Chita region and
Agin’-Buriat autonomous district (Ostrovsky, 2002), Kemerovo region (Krasnoborov,
2000), Republic of Khakassia (Krasnoborov,
2002), Novosibirsk region (Krasnoborov et
al. 1998), Republic of Tyva (Krasnoborov,
1999).
7. Far East Federal District: Khabarovsk
region (Voronov, 2000), Republic of Sakha
(Yakutia) (Isaev, 2000).
RESULTS
LICHENS IN THE RED DATA BOOK OF THE
RUSSIAN FEDERATION
The currently functioning Red Data Book of the
Russian Federation was approved and published
in 1988 and contains 29 species of lichens (Red
Data Book of RSFSR, 1988)2. All taxa in this Red
Data Book are distributed between five Red List
categories (Table 1) with lichen species being
mainly placed in categories 2 and 3. Presently
used categories are not corresponding to those
of IUCN, however, a tendency to find the ways of
adjusting national and international categories
is becoming more and more clear (Zavarzin &
Moutchnik, 2005). The second edition of the Red
Data Book of the Russian Federation devoted to
the threatened plants, fungi and lichens is to be
published in 2005.
Many of the nationally redlisted lichen taxa
are found in the Far East of Russia and just a
few could be found in the rest of the territory
(Table 2). However, due to the extremely uneven
knowledge of lichen diversity in different regions
of Russia it is hard to verify whether populations of these taxa are really threatened and so
narrowly localized or not. This problem could
be solved through analysis of the data gathered
for regional Red Data Books.
LICHENS IN THE REGIONAL RED LISTS AND
RED DATA BOOKS IN RUSSIA
Following the nationwide activities the redlisting
process has started recently in most regions of
the Russian Federation. Analysis of the state
of populations3 in the regions of Russia and
identification of threatened species are essential
for at-the-site preservation of species populations
61
13
10
12
9b
11
9a
8
6 43 5
27
1
20 21a 21b
24
23
19
26
35
25
22
32
30
31
27
33
28
29
34
17
14
161815
Fig.1. Map of the administrative division of the Russian Federation.
– – – Polar circle
Numbers (following those in Table 2) refer to the regions of Russia where Red Data Books contain
lichenised taxa.
I. Central Federal District:
1 – Belgorod region;
2 – Lipetsk region;
3 – Moscow region;
4 – city of Moscow;
5 – Ryazan region;
6 – Smolensk region;
7 – Tambov region;
8 – Tver region.
II. North-Western Federal
District:
9a – Archangel region;
9b – Nenets autonomous
district;
10 – Republic of Karelia;
11 – Republic of Komi;
12 – Leningrad region;
13 – Murmansk region.
III. Northern Caucasus Federal
District:
14 – Republic of Adygeja;
15 – Republic of Dagestan;
16 – Republic of KabardinoBalkaria;
17 – Krasnodar region;
18 – Republic of Northern Osetia
(Alania).
IV. Privolzhsky Federal District:
19 – Republic of Bashkortostan;
20 – Kirov region;
21a – Perm region and Komi-Perm
autonomous district;
22 – Saratov region;
23 – Republic of Tatarstan;
24 – Republic of Udmurtia.
V. Urals Federal District:
21b – Sverdlovsk region
25 – Khanty-Mansy autonomous
district;
26 – Yamalo-Nenets
autonomous district.
VI. Siberian Federal District:
27 – Republic of Altaj;
28 – Republic of Buriat;
29 – Chita region and Agin’-Buriat
autonomous district;
30 – Kemerovo region;
31 – Republic of Khakassia;
32 – Novosibirsk region;
33 – Republic of Tyva.
VII. Far East Federal District:
34 – Khabarovsk region;
35 – Republic of Sakha (Yakutia).
62
Folia Cryptog. Estonica
Table 1. Categories of wild plant taxa used in the Red Data Book of the Russian Federation
(1988)
Categories
Description of the categories used
0 – Possibly extinct
Taxa, previously known from the territory of Russia, but without viable
populations registered for at least 50 years, though the possibility of their
findings is not totally excluded.
1 – Facing extinction
Taxa that have declined to critical numbers (in terms of organisms and/or
localities) to the extent that they can get extinct in the nearest future.
2 – Decreasing in numbers
Taxa with steadily decreasing numbers of individuals or localities so that
under the existing pressures they will be the subjects for Category 1, including:
a) taxa that are declining due to the changes in environmental conditions
and habitat destruction;
b) taxa that are declining because of the overexploitation by humans and
require special conservation measures (e.g. medicinal, decorative plants,
etc.)
3 – Rare
Taxa with naturally low density and small area of distribution or sporadically distributed over large areas and which require special protective
measures, including:
a) endemic taxa with small areas of distribution;
b) taxa with large areals but sporadically distributed and having small
populations;
c) taxa with strong requirements for specific and rare habitats;
d) taxa with large areals but present in Russia at the limits of distribution;
e) taxa with limited areal with the part of it being in Russia.
4 – Of undetermined status
(=Data deficient)
Taxa assumed to be threatened and thus requiring special protective measures but having insufficient data to be assigned to one of the previous
categories.
5 – Restored or restoring
Taxa that are restoring their numbers and/or area of distribution under
natural or artificially maintained conditions and that will soon reach the
point when further protective measures will not be necessary.
including lichens. 62 out of 89 regions of Russia
(so called “Subjects of the Russian Federation”)
have compiled and published their regional Red
Data Books4 by the year 2004. These books cover
redlisted species of animals, vascular plants,
bryophytes, algae, fungi and lichens.
Species of lichens are included in 35 of
these regional Red lists covering 40 regions of
about a billion of hectares (924 370 thousands
ha), i.e. 54,1% of the territory of Russia (Fig. 1).
The number of protected lichen species varies
from 1 in Saratov region to 77 in the Republic of
Karelia. The total number of lichen taxa (species,
subspecies, and varieties) included in the Red
Data Books is 3685 (Table 2).
Most of the lichen taxa are categorized as
3 (Rare) or 4 (Data deficient). Seven lichens
from the Red Data Book of the Russian Federation (1988) are absent in the regional Red
Data Books. These are: Cladonia graciliformis
Zahlbr., C. vulcani Savicz, Icmadophila japonica
(Zahlbr.) Rambold & Hertel, Stereocaulon exutum
Nyl., S. saviczii Du Rietz, Teloschistes flavicans
(Sw.) Norm., and Umbilicaria esculenta (Mioshi)
Minks. These species, with the exception of
S. saviczii, will be included in the Red List of
Primorje region which is still not published. S.
saviczii should be protected in the Kamchatka
region where the Red List contains only animal
taxa so far.
63
DISCUSSION
Despite of certain positive trends in redlisting of lichens (e.g. increasing number of inventories prior
to compilation of Red lists and involvement of experienced lichenologists into revision of proposals
for the Red lists), some serious problems exist in
Russia. The overall low level of knowledge about
the lichen diversity in many regions of Russia is
clearly visible when comparing lists of lichens
and higher plants from the Red Data Books. The
majority of Red lists contain only large foliose and
fruticose lichens (Table 2). Along with the lack
of modern floristic studies obvious mistakes are
not uncommon. For example, Parmelia sulcata
is listed in the Red Data Book of Archangelsk
region or Phaeophyscia nigricans is included into
Red Data Books of Moscow and Riazan regions,
though it is obvious that these species are quite
common in those areas. Nephroma laevigatum is
protected in Kirov region even though it is hard
to imagine the presence of this oceanic species
in the quite continental part of Russia.
Furthermore, the term “rare lichen” in Russia is also subjective, mainly because different
regions of the Russian Federation have been
unevenly explored in terms of lichen diversity.
Taking into account the ongoing process of lichen
diversity inventories it is essential to distinguish
at least three different groups of regionally rare
lichens:
(1) “constitutively rare” meaning that the taxon
is scarcely spread in the whole area of distribution due to its ecological requirements;
(2) taxon that is present in the region at the limit
of its distribution, but which is common in
some neighbouring regions;
(3) taxon associated with rare biotopes especially
sensitive to anthropogenic impact.
A serious problem is arising from the lack of
detailed description of criteria for assigning lichens to different categories in the regional Red
Data Books in Russia. Analysis of lichenological
papers that present more or less reliable data on
lichen density in the regions indicates that most
of the authors use only subjective measures even
for this criterion. Usually species are ranked as
“rare”, “scarce”, “average” or “abundant” [see for
examples all issues of the Handbook of Lichens
of the USSR (Russia), 1971–2003; Shustov,
1989; Makryi, 1990; Dudoreva & Ahti, 1996;
Krivorotov, 1997; Hermansson, Kudriavtseva,
1997; Poriadina, 1998; Urbanavichene & Urbanavichus, 1998; Zhurbenko, 2000; Pystina,
2000; Vedeneev, 2001 and others]. Only a few
researchers indicate exact number of localities
where the species was found as a criterion for a
specific rank. For example, Budaeva (2001) ranks
as rare only taxa encountered at 3 to 5 locations
in the region. Kataeva (2002) considers species
with less than 4 localities as demanding regional
conservation efforts.
Possible solutions could be found in the enhancement of regional floristic studies especially
in protected areas and adoption of the system
of IUCN criteria (2001) for analyzing the state
of lichen populations prior to compiling regional
Red Data Books in Russia (Zavarzin & Moutchnik, 2005).
Due to their biology lichens are special objects for conservation. The relatively small size,
the ability for vegetative propagation, and the
low pressure from direct use by people help to
reduce the potential danger of decrease of their
populations. On the other hand, often narrow
substrate and biotope affiliation coupled with
high sensitivity to environmental quality make
the task of lichen conservation very difficult. As
there is no possibility to preserve lichens in botanical gardens or arboretums, the only way left
is to maintain viable populations through protecting biotopes and developing networks of protected
areas. In our opinion this is the most effective approach to be introduced in Russia and therefore
the redlisting efforts in the Russian regions have
to be always focused on ensuring the establishment and maintenance of the effective network
of protected areas. The Red Data Books can be
used as a powerful legal tool in this case.
Remarks
1
This Red Data Book covers two federal districts of Russia:
Perm region and Komi-Perm autonomous district belonging to the Privolzhsky Federal District, and Sverdlovsk
region being within Urals Federal District.
2
One of the species – Parmelia borisorum – is not
considered to be an independent taxon and is currently
included in Melanelia tominii.
3
The term “population” is used in a geographic sense
meaning all organisms of a certain taxon in a certain
region.
4
Regional Red Data Books follow the same categories
that are used in the Red Data Book of Russian Federation
(Table 1). However, they may contain additional categories
depending upon the needs and necessities of the
region.
5
Except for 85 species belonging the region-specific
category “demanding biological monitoring” in the Red
Data Book of Murmansk region, and for 7 taxa currently
treated as synonyms (these 7 taxa are also included in
Table 2 but are marked by “!”).
VI
VII
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
V
Acarospora sinopica (Wahlenb.) Körb.
+
Acrocordia gemmata (Ach.) A.Massal.
Adelolecia kolaënsis (Nyl.) Hertel & Rambold (as
Lecidea conferenda Nyl.)
Alectoria sarmentosa (Ach.) Ach.
+
+
+
A. sarmentosa ssp. vexillifera (Nyl.) D.Hawksw.
+
+
Amygdalaria pelobotryon (Wahlenb.) Norman
Anaptychia ciliaris (L.) Körb.
A. runcinata (With.) J.R.Laundon
Arctocetraria andrejevii (Oxner) Kärnefelt &
A.Thell
A. nigricascens (Nyl.) Kärnefelt & A.Thell
+
+
+ +
+
+
+
Arctoparmelia centrifuga (L.) Hale
+
A. incurva (Pers.) Hale
+
A. subcentrifuga (Oxner) Hale
+
Arthonia arthonioides (Ach.) A.L.Sm.
+
A. incarnata Th.Fr. ex Almq.
+
A. leucopellaea (Ach.) Almq.
+
A. ligniariella Coppins
A. ruana A.Massal. [as Arthothelium ruanum
(A.Massal.) Körb.]
Asahinea chrysantha (Tuck.) W.L.Culb. & C.F.Culb.
+
+
+
A. scholanderi (Llano) W.L.Culb. & C.F.Culb.
+
Aspicilia canina Räsänen
A. laevata (Ach.) Arnold
+
+
+
+ +
+ +
Folia Cryptog. Estonica
Lichen species
Red Data Books of the Subjects of the Russian Federation
I
II
III
IV
64
Table 2. Lichen species included into the Red Data Books of the regions of the Russian Federation (see Fig 1 for explanation of numbers). Lichen species protected at the national level (included in the Red Data Book of Russian Federation, 1988) are marked in bold.
Species that have been synonymised but are treated here still as separate taxa are marked with !.
A. protuberans Räsänen
Aspilidea myrinii (Fr.) Hafellner [as Aspicilia myrinii
(Fr.) Stein.]
Bacidia incompta (Borrer ex Hook.) Anzi
B. polychroa (Th.Fr.) Körb. [as B. acerina (Ach.)
Arnold]
Belonia russula Körb. & Nyl.
+
+
+
+
Brodoa intestiniformis (Vill.) Goward
+
Bryoria bicolor (Ehrh.) Brodo & D.Hawksw.
+
B. capillaris (Ach.) Brodo & D.Hawksw.
B. chalybeiformis (L.) Brodo & D.Hawksw. (as B.
intricans (Vain.) Brodo & D.Hawksw.)
B. fremontii (Tuck.) Brodo & D.Hawksw.
+
+
+
+
+
+ +
B. fuscescens (Gyeln.) Brodo & D. Hawksw.6
+
+
+ + +
+
+
B. implexa (Hoffm.) Brodo & D.Hawksw.
+
B. nadvornikiana (Gyeln.) Brodo & D.Hawksw.
+ +
B. nitidula (Th.Fr.) Brodo & D.Hawksw.
+
B. smithii (Du Rietz) Brodo & D.Hawksw.
+ +
B. subcana (Nyl. ex Stizenb.) Brodo & D.Hawksw.
Buellia geophila (Flörke ex Sommerf.) Lynge (as B.
triphragmia (Nyl.) Arnold)
Calicium abietinum Pers.
+
+
+
C. adaequatum Nyl.
+
C. lenticulare Ach.
+ +
C. quercinum Pers.
+
C. viride Pers.
+
Caloplaca decipiens (Arnold) Blomb. & Forssell
+
Cetraria aculeata (Schreb.) Fr.
+
C. ericetorum Opiz
+
+
+
+
+
+
65
C. odontella (Ach.) Ach.
+
+
C. sinapisperma (Lam. & DC.) Maheu & Gillet
C. islandica (L.) Ach.
+
Red Data Books of the Subjects of the Russian Federation
I
II
III
IV
VI
VII
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
V
C. sepincola (Ehrh.) Ach. (as Tuckermanopsis
sepincola (Ehrh.) Hale)
C. steppae (Savicz) Kärnefelt (as Cornicularia
steppae Savicz)
Cetrariella delisei (Bory ex Schaer.) Kärnefelt &
A.Thell
C. fastigiata (Delise ex Nyl.) Kärnefelt & A.Thell
Cetrelia alaskana (C.F.Culb. & W.L.Culb.)
W.L.Culb. & C.F.Culb.
! C. cetrarioides (Del. ex Duby) W.L. Culb. &
C.F.Culb.7
C. olivetorum (Nyl.) W.L.Culb. & C.F.Culb.
+
+
+
+
+
+
+
+
+ +
+
+
C. cinerea (Pers.) Tibell
+
C. chlorella (Ach.) Müll.Arg.
C. gracilenta (Ach.) J.Mattsson & Middelb. [as Cybebe gracilenta (Ach.) Tibell]
C. gracillima (Vain.) Tibell
+
+ +
+ +
C. hispidula (Ach.) Zahlbr.
+
C. laevigata Nádv.
+
C. phaeocephala (Turner) Th.Fr.
+
C. stemonea (Ach.) Müll.Arg.
+
+
+
C. subroscida (Eitner) Zahlbr.
+
Chaenothecopsis consociata (Nádv.) A.F.W.Schmidt
+
C. haematopus Tibell
+
C. vainioana (Nádv.) Tibell
+
C. viridialba (Krempelh.) A.F.W.Schmidt
C. amaurocraea (Flörke) Schaer.
+
+
Chaenotheca brachypoda (Ach.) Tibell
Cladonia acuminata (Ach.) Norrl.
+
+
+
+
+
Folia Cryptog. Estonica
Lichen species
66
Table 2 (continued)
C. arbuscula (Wallr.) Flot. ssp. arbuscula
+
+
+
+
+
+
C. bellidiflora (Ach.) Schaer.
C. botrytes (K.G.Hagen.) Willd.
+
C. caespiticia (Pers.) Flörke
C. cariosa (Ach.) Spreng.
+
+
+
C. cenotea (Ach.) Schaer.
+
C. crispata (Ach.) Flot.
+
C. deformis (L.) Hoffm.
+ +
+
+
+
+
+
C. digitata (L.) Hoffm.
C. ecmocyna Leight.
+
C. floerkeana (Fr.) Flörke
+
C. foliacea (Huds.) Willd.
+ + +
C. furcata (Huds.) Schrad.
+
C. gracilis (L.) Willd.
+
C. incrassata Flörke
+
+
+
+
+
C. kanewskii Oxner
+
C. luteoalba Wheldon & A. Wilson
C. macilenta Hoffm. ssp. macilenta
+
C. macroceras (Delise) Hav.
+
+
C. macrophylla (Schaer.) Stenh.
C. magyarica Vain.
+
+ +
C. nipponica Asahina
+
C. phyllophora Hoffm.
+
C. pleurota (Flörke) Schaer.
+
C. polycarpoides Nyl.
+
C. portentosa (Dufour) Coem.
C.rangiferina (L.) F.H.Wigg.
C. rangiformis Hoffm.
+
+
+
+
C. scabriuscula (Delise) Nyl.
+
+
+
+
67
C. stellaris (Opiz) Pouzar & Vĕzda
+
+
Red Data Books of the Subjects of the Russian Federation
I
II
III
IV
VI
VII
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
V
C. subrangiformis Sandst.
+
C. subulata (L.) Weber ex F.H.Wigg.
+
C. symphycarpia (Flörke) Fr.
+ +
C. turgida Hoffm.
C. verticillata (Hoffm.) Schaer. [as C. cervicornis
(Ach.) Flot. ssp. verticillata (Hoffm.) Ahti]
Cliostomum corrugatum (Ach.:Fr.) Fr.
+
+ +
+
+
C. leprosum (Räsänen) Holien & Tønsberg
Coccocarpia palmicola (Spreng.) Arv. &
D.J.Galloway [as C. cronia (Tuck.) Vain.]
C. erythroxyli (Spreng.) Swinscow & Krog
+
+ + +
+ +
Collema auriforme (With.) Coppins & J.R.Laundon
+
C. bachmanianum (Fink) Degel.
+
C. crispum (Huds.) Weber ex F.H.Wigg.
+
C. cristatum (L.) Weber ex F.H.Wigg.
+
+
C. curtisporum Degel.
C. fragrans (Sm.) Ach.
C. fuscovirens (With.) J.R.Laundon [as C. tunaeforme (Ach.) Ach.]
C. furfuraceum (Arnold) Du Rietz
C. limosum (Ach.) Ach.
C. nigrescens (Huds.) DC.
+
+
+
+ +
C. subflaccidum Degel.
+
+ +
+
+
+
+
+
C. subnigrescens Degel.
Cyphelium inquinans (Sm.) Trevis.
+
+
C. occultatum var. populinum (Th.Fr.) Degel.
C. polycarpon Hoffm.
+
+
+
+
+
Folia Cryptog. Estonica
Lichen species
68
Table 2 (continued)
C. karelicum (Vain.) Räsänen
+ +
! Dendriscocaulon umhausense (Auersw.) Degel.
+ +
8
Dermatocarpon deminuens Vain.
+
D. meiophyllizum Vain.
+
D. miniatum (L.) W.Mann.
+
Dimerella lutea (Dicks.) Trevis.
+
Endocarpon adscendens (Anzi) Müll.Arg.
+
E. psorodeum (Nyl.) Blomb. & Forssell
+
E. pusillum Hedw.
+
Evernia divaricata (L.) Ach.
+ + + +
E. mesomorpha Nyl.
+
+
+ +
+
E. prunastri (L.) Ach.
+
+
Flavocetraria nivalis (L.) Kärnefelt & A.Thell
+
Flavoparmelia caperata (L.) Hale
+ +
+
Flavopunctelia soredica (Nyl.) Hale
+
+
Fuscopannaria ahlneri (P.M.Jørg.) P.M.Jørg.
+
F.confusa (P.M. Jørg.) P.M.Jørg.
Gonohymenia nigritella (Lettau) Henssen [as
Lichinella nigritella (Lettau) Moreno]
Graphis scripta (L.) Ach.
+
+
Gyalecta flotowii Körb.
+
G. kukriensis (Räsänen) Räsänen
+
+
G. ulmi (Sw.) Zahlbr.
+
Heppia solorinoides (Nyl.) Nyl.
+
Heterodermia japonica (M.Sato) Swinscow & Krog
+
H. speciosa (Wulfen) Trevis.
+ +
Hypogymnia austerodes (Nyl.) Räsänen
+ +
+
+
+
+
69
H. bitteri (Lynge) Ahti
H. hypotrypa (Nyl.) Rass. [as H. hypotrypella
(Asahina) Rass.]
H. pseudophysodes (Asahina) Rass.
+
Red Data Books of the Subjects of the Russian Federation
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16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
V
H. pulverata (Nyl. ex Cromb.) Elix
+
H. submundata (Oxner) Rass.
+
H. tubulosa (Schaer.) Hav.
+
+
H. vittata (Ach.) Parrique
+
Hypotrachyna revoluta (Flörke) Hale
+
H. sinuosa (Sm.) Hale
Icmadophila ericetorum (L.) Zahlbr.
+
+
Imshaugia aleurites (Ach.) S.L.F.Meyer
Lecanora albescens (Hoffm.) Branth & Rostr.
+
+
+
+
L. cenisea Ach.
+
L. hypopta (Ach.) Vain.
+
L. mughicola Nyl.
+
L. subcarnea (Lilj.) Ach.
+
L. sulphurea (Hoffm.) Ach.
+
Lecidea albofuscescens Nyl.
+
L. lapicida (Ach.) Ach.
+
L. leprarioides Tønsberg
L. plana (J.Lahm.) Nyl.
Leptochidium albociliatum (Desm.) M. Choisy
+
+
+
Leptogium asiaticum P.M.Jørg.
+ +
L. brebissonii Mont.
+
L. burnetiae C.W.Dodge
L. cyanescens (Rabh.) Körb.
L. issatschenkoi Elenk.
L. hildenbrandii (Garov.) Nyl.
+
+
+
+ + +
+
+ +
+ + +
+
+
+ +
+
Folia Cryptog. Estonica
Lichen species
70
Table 2 (continued)
L. lichenoides (L.) Zahlbr.
+
+
+
L. rivulare (Ach.) Mont.
+
+
+
L. saturninum (Dicks.) Nyl.
L. schraderi (Bern.) Nyl.
+
+
L. subtile (Schrad.) Torss.
+
L. tenuissimum (Dicks.) Körb.
+
L. teretiusculum (Wallr.) Arnold
+
Letharia vulpina (L.) Hue
Lichenomphalia hudsoniana (H.S.Jenn) Redhead et al. [as Omphalina hudsoniana (H.S.Jenn)
H.E.Bigelow]
Lobaria amplissima (Scop.) Forssell
+
+
+
+ +
+ +
L. hallii (Tuck.) Zahlbr.
+ +
+ +
+
L. isidiosa (Müll.Arg.) Vain.
+ +
L. linita (Ach.) Rabenh.
+
+
L. meridionalis Vain.
+
L. orientalis (Asahina) Yoshim.
+
+ + + + +
L. pulmonaria (L.) Hoffm.
+
+
+
L. scrobicalata (Scop.) DC.
+ + + + + + + + + + + +
+
+ + +
+
L.virens (With.) J.R. Laundon
+
+ +
+ +
+
+ +
+
Lobothallia radiosa (Hoffm) Hafellner
+
Masonhalea richardsonii (Hook.) Kärnefelt
+ +
Melanelia commixta (Nyl.) A.Thell
+
M. exasperata (De Not.) Essl.
+
M. exasperatula (Nyl.) Essl.
+
+
M. fuliginosa (Fr. ex Duby) Essl.
+
+
+
+
+
+
M. hepatizon (Ach.) A.Thell
M. olivacea (L.) Essl.
+
+
L. retigera (Bory) Trevis.
M. glabra (Schaer.) Essl.
+
+
+
+
+
71
72
Table 2 (continued)
V
VI
VII
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
Lichen species
M. septentrionalis (Lynge) Essl.
+
M. sorediata (Ach.) Goward & Ahti
+
M. stygia (L.) Essl.
+
M. subargentifera (Nyl.) Essl.
+
M. subaurifera (Nyl.) Essl.
M. tominii (Oxner) Essl. (as Parmelia borisorum
Oxner)
Menegazzia terebrata (Hoffm.) A.Massal.
+
+
+
+
! Micarea kivakkensis Vain.
+
+
M. tuberculata (Sommerf.) R.A.Anderson
+
+
+
+
+
+
+
+
N. ryssolea (Ach.) Essl.
+
Nephroma arcticum (L.) Torss.
+
N. bellum (Spreng.) Tuck.
+
N. helveticum Ach.
+
N. isidiosum (Nyl.) Gyeln.
+
+
+
+
+
N. parile (Ach.) Ach.
N. resupinatum (L.) Ach.
+ +
+
Neofuscelia loxodes (Nyl.) Essl.
N. laevigatum Ach. non auct.
+
+
Miriquidica deusta (Stenh.) Hertel & Rambold
Mycobilimbia lurida (Ach.) Hafellner &Türk [as
Lecidea lurida (Ach.) DC.]
Myelochroa metarevoluta (Asahina) Elix & Hale
N. pulla var. delisei Duby.
+
+
Microcalicium ahlneri Tibell
N. pulla (Ach.) Essl.
+
+
9
+
+
+
+
+
+
Folia Cryptog. Estonica
Red Data Books of the Subjects of the Russian Federation
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II
III
IV
Nephromopsis komarovii (Elenk.) Wei
(as Cetraria komarovii Elenk.)
Normandina pulchella (Borrer) Nyl.
+ + +
+ +
Ochrolechia pallescens (L.) A.Massal.
+
Opegrapha atra Pers.
Ophioparma lapponica (Räsänen) Hafellner &
R.W.Rogers [as O. ventosa (L.) Norman var. lapponica Räsänen]
Pannaria conoplea (Ach.) Bory
+
+
+
+
P. rubiginosa (Ach.) Bory
+
Parmelia fraudans (Nyl.) Nyl.
+
P. saxatilis (L.) Ach.
+
P. sulcata Taylor
+
Parmeliella triptophylla (Ach.) Müll.Arg.
+
Parmelina quercina (Willd.) Hale
+
+
P. tiliacea (Hoffm.) Hale
+
Parmeliopsis ambigua (Wulfen) Nyl.
+
+
Peltigera aphthosa (L.) Willd.
+
P. canina (L.) Willd.
+
+
+
P. collina (Ach.) Schrad.
+ +
P. degenii Gyeln.
+
P. elisabethae Gyeln.
+
+
P. horizontalis (Huds.) Baumg.
+
+
P. leucophlebia (Nyl.) Gyeln.
+
P. malacea (Ach.) Funck
+
+
P. ponojensis Gyeln.
+
P. praetextata (Flörke ex Sommerf.) Fr.
+
P. rufescens (Weiss) Humb.
+
+
73
P. scabrosa Th.Fr.
+
+ +
+
P. hyperopta (Ach.) Arnold
P. neckeri Hepp ex Müll.Arg.
+ +
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24
25
26
27
28
29
30
31
32
33
34
35
V
P. venosa (L.) Hoffm.
+
+ +
Peltula radicata Nyl.
+
P. zabolotnoji (Elenk.) Golubk.
+
Pertusaria hemisphaerica (Flörke) Erichsen
+ +
P. pseudophlyctis Erichs.
+
P. raesaenenii Erichsen
+
P. sommerfeltii Fr.
Phaeocalicium compressulum (Szatala)
A.F.W.Schmidt
P. populneum (Brond ex Duby) A. F.W.Schmidt
+
+
+
+ +
Phaeophyscia endophoenicea (Harm.) Moberg
+
P. hirsuta (Mereschk.) Moberg
+
P. hispidula (Ach.) Moberg
+
P. kairamoi (Vain.) Moberg
+ +
P. nigricans (Flörke) Moberg
+
+
+
+
P. pyrrophora (Poelt) D.D.Awasthi & M.Joshi
+
Phylliscum demangeonii (Moug. & Mont.) Nyl.
+
Physcia caesia (Hoffm.) Fürnr.
Physconia detersa (Nyl.) Poelt
+
+
+
P. perisidiosa (Erichsen) Moberg
+
Placolecis opaca (Fr.) Hafellner
+
P. cribellans (Nyl.) Räsänen
Platismatia glauca (L.) W.L.Culb. & C.F.Culb.
Pleopsidium flavum (Bellardi) Körb.
+
+ +
+
Pleurosticta acetabulum (Neck.) Elix & Lumbsch
Polychidium muscicola (Sw.) Gray
+
+
+
+
+
Folia Cryptog. Estonica
Lichen species
74
Table 2 (continued)
Protopannaria pezizoides (Weber) P.M.Jørg. [as
Pannaria pezizoides (Web.) Trev.]
Protoparmelia badia (Hoffm.) Hafellner
P. nephaea (Sommerf.) R.Sant.
Protoparmeliopsis muralis (Schreb.) M.Choisy [as
Lecanora muralis (Schreb.) Rabenh.]
Pseudevernia furfuracea (L.) Zopf
+
+
+
+
+ +
+
+
+
Psilolechia clavulifera (Nyl.) Coppins
+
+
Psora globifera (Ach.) A.Massal.
+
P. rubiformis (Ach.) Hook.
+
Psoroma hypnorum (Vahl.) Gray
+
Punctelia subrudecta (Nyl.) Krog
+
Pycnothelia papillaria (Ehrh.) Dufour
+
Pyrenocarpon flotowianum (Hepp.) Trevis.
+
Pyrrhospora elabens (Fr.) Hafellner
Pyxine endochrysoides (Nyl.) Degel. [as Pyxine
sorediata (Ach.) Mont.]
! Ramalina asahinana Zahlbr.10
+
+ + + + +
+
R. baltica Lettau
+ + +
+
R. calicaris (L.) Fr.
+
+ +
R. dilacerata (Hoffm.) Hoffm.
+
+
+
+
R. elegans (Bagl. & Carestia) Jatta
+
R. evernioides Nyl.
R. farinacea (L.) Ach.
+
+
+
+
+
+
R. fastigiata (Pers.) Ach.
R. fraxinea (L.) Ach.
+
+
+
R. obtusata (Arnold) Bitter
R. pollinaria (Westr.) Ach.
R. roesleri (Hochst. ex Schaer.) Hue
+
+
+
+ +
+
+
+ +
R. sinensis Jatta
R. subfarinacea (Nyl. ex Cromb.) Nyl.
+
+
+ +
+
75
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24
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28
29
30
31
32
33
34
35
V
R. thrausta (Ach.) Nyl.
+ + + +
Rhizocarpon grande (Flörke) Arnold
+
R. hochstetteri (Körb.) Vain.
R. petraeum (Wulfen) A.Massal. (as R. concentricum auct.)
R. postumum (Nyl.) Arnold
Sclerophora coniophaea (Norman) J. Mattsson &
Middelb.
Sphinctrina anglica Nyl.
+
+
+
+
+
+
S. turbinata (Pers.:Fr.) De Not.
+
Stenocybe major Nyl.
+
Stereocaulon arenarium (Savicz) I.M.Lamb
+
S. capitellatum H.Magn.
+
S. dactylophyllum Flörke
+
S. symphycheilum I.M.Lamb
+
+
+ +
+
Sticta arctica Degel.
+
S. fuliginosa (Hoffm.) Ach.
+
S. limbata (Sm.) Ach.
+
S. nylanderiana Zahlbr.
+
S. sylvatica (Huds.) Ach.
S. wrightii Tuck.
+
Synalissa symphorea (Ach.) Nyl.
+
Tholurna dissimilis (Norman) Norman
+
Tornabenia atlantica (Ach.) Kurok.
Trapeliopsis viridescens (Schrad.) Coppins &
P.James
Tuckermannopsis chlorophylla (Willd.) Hale
+
+ +
+
+
+
+
+
+
+
+ +
+
+
Folia Cryptog. Estonica
Lichen species
76
Table 2 (continued)
T. ciliaris (Ach.) Gyeln.
Tuckneraria laureri (Kremp.) Randlane &
A.Thell [as Nephromopsis laureri (Krempelh.)
Kurok. or Cetraria laureri Krempelh.]
Umbilicaria deusta (L.) Baumg.
+
+
+
+
+
+
+
+ + + + +
+
U. hirsuta (Sw. ex Westr.) Hoffm.
+
U. hyperborea (Ach.) Hoffm.
+
+
+
U. krascheninnikovii (Savicz) Zahlbr.
+
U. polyphylla (L.) Baumg.
+
+
U. polyrrhiza (L.) Ach.
+
U. proboscidea (L.) Schrad.
+
U. pulvinaria (Savicz) Frey11
+
Usnea articulata (L.) Hoffm.
+
+
U. barbata (L.) Weber ex F.H.Wigg
+
+
U. cavernosa Tuck.
+
! U. distincta Motyka
+
12
! U. extensa Vain.13
+
U. filipendula Stirt.
+
U. florida (L.) Weber ex F.H.Wigg.
+
+
+
+ + + + + + +
! U. foveata Vain.
U. fulvoreagens (Räsänen) Räsänen
+
U. glabrata (Ach.) Vain.
+
U. glabrescens (Nyl. ex. Vain.) Vain.
+
+
+
U. hapalotera (Harm.) Mot.
U. hirta (L.) Weber ex F.H.Wigg.
U. lapponica Vain.
+
+
+ + +
+
+
U. longissima Ach.
U. subfloridana Stirt.
+ +
+ +
+
+
U. wasmuthii Räsänen
Varicellaria rhodocarpa (Körb.) Th.Fr.
+
+
14
+
+
77
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17
18
19
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22
23
24
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27
28
29
30
31
32
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35
V
Verrucaria margacea (Wahlenb.) Wahlenb.
+
V. onegensis Vain.
+
Vulpicida juniperinus (L.) J.-E. Mattsson & M.J.Lai
+
+
Xanthoparmelia camschadalis (Ach.) Hale
+
X. conspersa (Ach.) Hale
+
X. somloënsis (Gyeln.) Hale
+
Xanthoria calcicola Oxner
+
X. candelaria (L.) Th.Fr.
X. fallax (Hepp) Arnold
Remarks
+
+
+
6
In two regional Red Data Books (of Republic of Adygeja and of Krasnodar region) this species is listed under Bryoria jubatus (L.) Brodo & D.
Hawksw.
7
Cetrelia cetrarioides is currently treated as a synonym to C. olivetorum (see e.g. Santesson et al., 2004). However, we prefer to maintain the chemical
species here as separate taxa.
8
Dendriscocaulon umhausense is currently treated as Lobaria amplissima (see e.g. Santesson et al., 2004). However, the taxon is redlisted in two regions
of Russia under this name which is used also here.
9
Micarea kivakkensis Vain. is considered to be conspecific with Catillaria contristans (Nyl.) Zahlbr. (see e.g. Kotlov, 2004) which is a rather widespread
lichen. This taxon was included into the officially adopted list of threatened species of the Republic of Karelia presumably due to its original description
from the Karelian locality.
10
Ramalina asahinana is often synonymized with R. sinensis; however, both taxa are included in the Red Data Book of Novosibirsk Region and Red
Data Book of the Republic of Khakassia. We also prefer to keep these two taxa separate. Furthermore, R. asahinana is misspelled in the Red Data Book
of Republic of Adygeja and is included there as “Ramalina asahina Zahlbr.” while R.sinensis is mentioned in the latter publication as a synonym.
11
Umbilicaria pulvinaria was originally mentioned as Gyrophora pulvinaria by Savicz (1911: 256) and was described later under the same name (Savicz,
1914). The taxon needs revision.
12
Usnea distincta is not reported for Russia in the recent treatment of the genus (Handbook of the Lichens of Russia, Vol. 6, 1996), but is mentioned
there as a potential synonym of U. glabrescens (p. 86). However, the taxon is included into the Red Data Book of the Republic of Adygeja and therefore
is kept in the list of officially protected lichen species. Presumably should be treated as U. glabrescens.
13
Usnea exstensa is treated as a separate species in the recent treatment of the genus in Russia (Handbook of the Lichens of Russia, Vol. 6, 1996:
78–79) and is redlisted in the Republic of Karelia. According to other authors (e.g. Santesson et al., 2004) the taxon is a synonym of U. glabrescens.
14
Usnea foveata is treated as a separate species in the recent treatment of the genus in Russia (Handbook of the Lichens of Russia, Vol. 6, 1996:
82–83) and is redlisted in Khanty-Mansy autonomous district. According to other authors (e.g. Santesson et al., 2004 ) the taxon is a synonym of U.
barbata.
Folia Cryptog. Estonica
Lichen species
78
Table 2 (continued)
79
REFERENCES
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Republic of Dagestan. Rare, threatened species of
animals and plants (in Russian). Makhachkala.
338 pp.
Andreev, V. (ed.) 1995. Red Data Book of Arkhangelsk
region. Rare and protected species of plants and
animals (in Russian). Arkhangelsk. 330 pp.
Bojkov, T. (ed.) 2002. Red Data Book of the Republic of
Buriat. Rare and decreasing species of plants and
fungi (in Russian). Novosibirsk. 340 pp.
Bolshakov, V. & Gorchakovsky, P. (eds) 1996. Red
Data Book of Middle Urals (Sverdlovsk and Perm
regions): rare and threatened species of animals
and plants (in Russian). Ekaterinburg. 279 pp.
Budaeva, S. 2001. Rare species of lichens of Buriatia
(in Russian). Novitates Systematicae Plantarum
non Vascularium 35: 96–101.
Dobrinsky, L. (ed.) 1997. Red Data Book of YamaloNenets autonomous district: Animals, plants, fungi
(in Russian). Ekaterinburg. 240 pp.
Dobrinsky, L. & Korytin, N. (eds) 2001. Red Data Book
of Kirov region: Animals, plants, fungi (in Russian).
Ekaterinburg. 288 pp.
Dudoreva, T. & Ahti, T. 1996. Rare species of
macrolichans in Murmansk region (in Russian).
Novitates Systematicae Plantarum non Vascularium 31: 109–113.
Isaev, A. (ed.) 2000. Red Data Book of the Republic
of Sakha (Yakutia). Vol. 1. Rare and threatened
species of plants and fungi (in Russian). Yakutsk.
256 pp.
Ivanov, I. (ed.) 2000. Red Data Book of the Republic
of Kabardino-Balkaria. Rare, threatened species
of animals and plants (in Russian). Nalchik. 308
pp.
Ivanter, E. & Kuznetsov, V. (eds) 1995. Red Data Book
of Karelia (in Russian). Petrozavodsk, Karelia.
186 p.
Handbook of the Lichens of the USSR (Russia)
Vol. 1–9 (in Russian). 1971–2004. Leningrad
(St.Petersburg).
Hermansson, J. & Kudryatseva, D. 1995. Notes on
the lichens of the Pechoro-Ilych Zapovednik, Komi
Republic, Russia. Graphis Scripta 7: 67–78.
IUCN Red List Categories and Criteria: Version 3.1.
2001. Gland. 30 pp.
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Folia Cryptog. Estonica, Fasc. 41: 81–88 (2005)
Lithophilous lichens of Middle Urals
Alexander G. Paukov & Svetlana N. Trapeznikova
Biological Faculty, Urals State University, Lenin av. 51, 620083, Ekaterinburg, Russia
E-mail: alexander_paukov@mail.ru
Abstract: The list of 194 lichen species spatially connected with rocky outcrops at Middle Urals is presented. The differences
in species composition and specificity, taxonomic spectra are shown for the rock types under consideration. The groups of
rarity and pollution sensitivity are segregated within epilithic lichens.
Kokkuvõte: Kesk-Uurali litofiilsed samblikud.
Esitatakse Kesk-Uurali kaljupaljandite samblike nimekiri, mis sisaldab 194 liiki. Käsitletakse erinevat tüüpi kivisubstraatide
lihhenofloora liigilise koosseisu iseärasusi. Epiliitsete samblike hulgas on eristatud erineva haruldus- ja saastetundlikkusastmega rühmad.
INTRODUCTION
The lithophilous lichen flora of Middle Urals has
never been thorougly studied and only a few data
on lichen biodiversity on rocks are available for
this territory (Ryabkova, 1998; Kotlov, 2003;
Paukov & Trapeznikova, 2003).
Middle Urals lay between latitudes 55º54’
and 59º15’N (Fig. 1) (Urals and ..., 1968). It is a
less elevated part of Ural mountains but it bears
a high complexity in the aspects of geology and
vegetation. The outcrops of limestone, ultramafic, basic and acidic rocks situated mainly at the
river banks are equally ubiqiutous here. Limestone is a sedimental rock type that consists
of about 95% of calcium carbonate and 5% of
calcium-magnesium carbonate. The acidity rate
of silicate rocks is often estimated indirectly by
comparison of content of SiO2 (weak acid) and
some additional components which may influence on pH. Serpentines and pyroxenites are the
ultramafic rocks which contain about 41% of
SiO2 and 36–42% of MgO. Basalts are the basic
rocks which contain up to 50% of SiO2, 16% of
Al2O3, 9% of CaO and 6% of MgO. Acidic rocks
– granites – consist of 72% of SiO2 and 14% of
Al2O3 (Voitkevich et al., 1990).
The highest relative altitude is 878 metres
(Katchkanar mountain) that is almost twice as
low as neighbouring mountains of Northern
Urals so there is no altitudinal zonation seen
within Middle Urals. Climatic conditions on the
outcrops vary from rather cool and wet from a
seepage water on the northern slopes to dry and
hot with temperatures rising above 60 ºC at the
southern slopes. Coniferous forests with pine
(Pinus sylvestris L.) and fir (Picea obovata Ledeb.)
is the dominant vegetation with birch forests and
steppe patches are on the south.
Middle Urals is a territory with high concentration of metallurgical factories. Main pollut-
Fig. 1. The location of Middle Urals within the
Ural mountains.
82
Folia Cryptog. Estonica
ants are heavy metals (Cu, Pb, Fe and others)
and sulphur dioxide. Car exhaust gases are the
major pollutants in Ekaterinburg that result in
high concentration of nitrogen oxides, formaldehyde, phenol and lead in the air.
The aims of this study were to reveal the
lichen biodiversity of the territory, to compare
species richness and composition on different
rocks and to evaluate the possibility to delimit
pollution sensitivity groups within the lithophilous lichens.
MATERIAL AND METHODS
Lichens spatially connected with rocky outcrops
and growing directly on rock, plant debris, soil
and mosses have been collected at the banks
of rivers Bagaryak, Iset, Sysert, Pyshma, Serga,
Chusovaya, Ufa, Rhezh, Neiva, Tagil, Tura and
on mountains such as Uktus, Volchikha, Shunut and others. Different types of rocks, e.g.
limestones, basalts, serpentines, pyroxenites,
dunites, gabbro, granites and diorites have been
studied. Attention has also been payed to artificial stones and concrete in towns and villages.
The term “lithophilous lichens” was interpreted
here as the species not growing directly on rock
only but as all species occuring in these habitats. Many of them are not obligately epilithic
(e.g. Caloplaca cerina var. chloroleuca (Sm.) Th.
Fr., Endocarpon pusillum Hedwig etc.).
List of lichens was compiled for every rock
type separately. Sørensen coefficient was calculated to evaluate the similarity of such species
lists. Delimitation of categories of rare species
was made using the following criteria: 1) lichens
with a single known locality and rare within it
(category rr1); 2) lichens with a single locality
but abundant there (category rr2); 3) lichens
known from two or three distant localities and
rare within them (category rr3).
The affinity groups of lichen families to
substrate type were derived from the ranks
of families counted on the number of species
recorded on the certain rock type (e.g.
Parmeliaceae with 15 species on pyroxenite
was graded to the rank 1, Lecanoraceae with
10 species was graded to the rank 2, Physciaceae
with 8 species was graded to the rank 3). Using
the highest ranks for particular family the
segregation of affinity groups was made. The
term “acidofilic” is used for the families which
have highest ranks on granite, "neutrofilic" are
the families with highest ranks on ultramafics
and basalt as the intermediate rocks in SiO2
content. "Basifilic" family is the one with highest
ranks on limestone, and species of which had
not been found on granite. Some additional
groups were also segregated.
Species on granite were used to delimit
groups of sensitivity to air pollution. Six points
with granite outcrops in different parts of Ekaterinburg and 50 km NW from the town were
selected. The location of inner points of species distribution in the town together with the
quantity of specimens found were the criteria for
delimitation of sensitivity groups of lithophilous
lichens. Three groups were delimited: tolerant
– species abundant in the centre of the town;
moderately tolerant – species growing only in the
town periphery (the park zone) and non-polluted
zone; sensitive – species collected only in the
non-polluted territory.
RESULTS AND DISCUSSION
194 species of lithophilous lichens are known
from the Middle Urals. Total species number on
different rocky outcrops vary from 82 species on
limestone and serpentine to 69 species on granite (Table 1). Only 11 species (6% of total) are
amphitolerant growing on all rock types while
Table 1. Lichen species quantity on different rock types
Total number of species
Number of species specific to the
rock type
Specificity index = (Specific/Total)
Limestone
no SiO2
82
Serpentine
41% SiO2
82
Pyroxenite
41% SiO2
70
Basalt
50% SiO2
78
Granite
72% SiO2
69
47
14
6
15
22
0,57
0,17
0,09
0,19
0,32
83
the others appear to be selective. Limestone is a
substrate with the highest percentage of specific
lichens – 57% of all taxa recorded on limestone
have not been recorded elsewhere. Granites (incl.
diorites) have the second highest percentage of
specific species – 32%. Ultramafics and basalt
are rather poor in specific lichens (less than
20% of recorded species). The most chemically
deviating rocks bear the most specific flora while
the substrates with the intermediate values of
SiO2 concentration have a lot of species which
are able to migrate there either from granite or
from limestone. The data on the serpentine flora
of higher plants (Kruckeberg, 1954; Proctor &
Woodell, 1971) are different and imply that the
pH rather than availability of metal ions is a key
factor determining lichen species distribution
on different rocks.
A dendrogram based on the Sørensen’s
index shows the similarity of lichen floras on
different types of rocks (Fig. 2). Ultramafics (serpentine and pyroxenite) and basalt are rather
Fig. 2. Similarity dendrogram of the species
composition of lichens on different rocks.
Numbers are the Sørensen similarity indexes
of the linkage.
similar in their species composition while flora
of limestone is the most deviating.
Taxonomic analysis of lichen biodiversity on
different rocks revealed the affinity of different
lichen families to a particular substrate group
(Table 2). Physciaceae and Lecanoraceae are
the amphitolerant families with a high number
of species on every substrate. Porpidiaceae
and Rhizocarpaceae are acidophilic families
with the biggest number of species on granite.
Parmeliaceae has the affinity to neutral and acidic
rocks. This is different from Teloschistaceae
which is widespread on limestone, basalt and
ultramafics. Collemataceae is a basifilic family
that is very usual on limestone and have not
been found on granite. According to our data
for the same territory, these affinities of the
families including also epiphytic lichens are
not necessarily concordant with the affinities
for different bark types segregated by the acidity.
Teloschistaceae, Lecanoraceae and Parmeliaceae
are concordant while Physciaceae is more
acidophilic on rocks.
50 species are rare in the region. Acarospora
oligospora (Nyl.) Arnold, Melanelia panniformis
(Nyl.) Essl., Micarea sylvicola (Flotow) Vězda
& V. Wirth, M. tuberculata (Sommerf.) R. A.
Anderson, Miriqiudica leucophaea (Rabenh.)
Hertel & Rambold, Toninia opuntioides (Vill.)
Timdal and others belong to the first category of
rare species. Bacidia herbarum (Stizenb.) Arnold,
Heterodermia speciosa (Wulfen) Trevis., Trapelia
placodioides Coppins & P. James are the species
of the second category. Caloplaca chrysodeta
(Vainio et Räsänen) Dombr., Cetrelia olivetorum
(Nyl.) W.L. Culb. & C.F. Culb., Flavoparmelia
caperata (L.) Hale, Phlyctis argena (Sprengel)
Flot., Sarcogyne privigna (Ach.) Anzi are the
examples of the third category of rare species.
Granite is a substrate with a biggest percentage of rare species (28% of total) (Table 3). The
discrepancy between the poorest lichen flora and
the biggest number of rare species can probably
be explained by microclimatic conditions on this
substrate. Granite outcrops are situated under
the forest canopy within the studied territory.
The surface of this stone is permanently cold so
arctic and hypoarctic species are common on
this substrate. Rare steppe lichens (Cladonia
pocillum (Ach.) Grognot, Fulgensia bracteata
(Hoffm.) Räsänen) occur mainly on limestone
(the most xeric substrate) or on the southern
slopes of other rock types.
84
Folia Cryptog. Estonica
Table 2. Taxonomic spectrum of lichen flora on different rocks. Numbers (1–26) are the ranks of
families counted on the number of species recorded on the certain rock type. Highest ranks (1–5)
for the particular family are in bold.
Family
Physciaceae
Lecanoraceae
Porpidiaceae
Rhizocarpaceae
Parmeliaceae
Hymeneliaceae
Teloschistaceae
Collemataceae
Limestone
3
4
12–16
9–11
12–16
1
2
Serpentine
1
2
12–13
14–26
3–4
3–4
5
9–11
Substrate
Pyroxenite
3
2
6–8
6–8
1
5
4
9–10
Granite outcrops are common within
Ekaterinburg and suburbs. It gives a good
possibility to estimate changes of biodiversity
of epilithic lichens in the town and delimit
sensitivity groups of species on this substrate.
Granite of the central part of the town is
characterized by the poorest lichen biodiversity
(17 species only, some of them are included into
the tolerant group). The most usual species are:
Acarospora fuscata (Schrad.) Arnold, Aspicilia
cinerea (L.) Körb., Caloplaca crenularia (With.)
J.R. Laundon., Candelariella vitellina (Hoffm.)
Müll. Arg., Lecanora dispersa (Pers.) Sommerf.,
Lecanora muralis (Schreb.) Rabenh., L. polytropa
(Hoffm.) Rabenh., Physcia caesia (Hoffm.)
Fürnr., Porpidia crustulata (Ach.) Hertel &
Knoph, Scoliciosporum umbrinum (Ach.) Arnold.
Some lichens known from the centre of the city
with the one specimen only are not included
into this group. 25–29 lichen species are
recorded on different outcrops in the periphery
of Ekaterinburg. Moderately tolerant lichens
with an inner distribution border in a town
periphery are: Dibaeis baeomyces (L.) Rambold
& Hertel, Diploschistes scruposus (Schreb.)
Norman, Phaeophyscia sciastra (Ach.) Moberg,
Basalt
2–4
5
9–13
9–13
1
2–4
2–4
7
Granite
5–6
2
3–4
3–4
1
7–10
13–22
-
Affinity
Amphitolerant
Amphitolerant
Acidophilic
Acidophilic
Calciphobous
Neutrophilic
Basi-neutrophil.
Basifilic
Polysporina simplex (Davies) Vězda, Rhizocarpon
grande (Flörke) Arnold, R. obscuratum (Ach.)
A. Massal., Stereocaulon tomentosum Fr. and
Umbilicaria deusta (L.) Baumg. The latter was
recorded in the town on granite in 1995 but
has not been found later. The group of sensitive
lichens include: Arctoparmelia centrifuga (L.)
Hale, Bryoria simplicior (Vain.) Brodo & D.
Hawksw., Cetrelia olivetorum (Nyl.) W.L. Culb. &
C.F. Culb., Lepraria rigidula (de Lesd.) Tønsberg,
Melanelia panniformis (Nyl.) Essl., Parmelia
fraudans (Nyl.) Nyl., Porpidia cinereoatra (Ach.)
Hertel & Knoph. Granites outside the town bear
the most diverse lichen biota that includes 67
species. Caloplaca crenularia and Lecanora
dispersa have not yet been found on granite in
the non-polluted territory. Melanelia disjuncta
(Erichsen) Essl. and M. infumata (Nyl.) Essl.
were found outside the town only on granites,
and regarded as sensitive. Inside the town they
were found on different substrate (pyroxenite).
It may reflect the possible changes of the rock
chemistry as have been previously reported in
Sweden (Arup et al., 1989).
Some rare fruticose and foliose lichens as the
most susceptible group to anthropogenic stress
Table 3. Number of rare species on different rock types
Number of rare species
Percentage of rare species of total
number of species on the rock type
Limestone
18
Serpentine
8
Pyroxenite
3
Basalt
8
Granite
19
22
10
4
10
28
85
are in need of protection. These species are:
Bryoria simplicior, Cetrelia olivetorum, Collema
crispum (Hudson) F.H. Wigg., C. flaccidum
(Ach.) Ach., C. glebulentum (Nyl. ex Crombie)
Degel., Flavoparmelia caperata, Heterodermia
speciosa, Leptogium lichenoides (L.) Zahlbr.,
Melanelia panniformis, Peccania coralloides
(A. Massal.) A. Massal., Solorina saccata (L.)
Ach. and Umbilicaria rossica (Dombr.) Golubk.
Some of them grow on the territory of “Olenyi
ruchyi” national park and Visim state biospheric
reserve both situated within the Middle Urals.
Nevertheless, the forest fires and tourist
activities still leave the possible threat to these
species.
CONCLUSIONS
The biodiversity of lithophilous lichens of the
Middle Urals is rather high. This is caused
mainly by the substrate and microclimatic
diversity that gives suitable sites for acidofilic
and basifilic lichens and a possibility for both
steppe and arctic lichens to occur. Limestone
and granite which bear the most specific flora
are very valuable substrates for the conservation of rare species. Atmospheric pollution and
recreation are the most deleterious factors for
lithophilous lichens in the studied area. The
segregated sensitivity groups can be used as
an alternative way for monitoring lichens especially in polluted sites where epiphytic lichens
are poorly developed.
ACKNOWLEDGMENTS
We are very grateful to Mikhail Andreev, Irina
Makarova, Einar Timdal, Ulrik Søchting, Alexei
Zavarzin and Mikhail Zhurbenko for the check of
lichens, and to a reviewer for the patient reading
and checking of the manuscript.
The study has been financially supported by
grants of Russian fund of basic research (0404-96131) and Russian Ministry of education
(A.03-2.12-493).
REFERENCES
Arup, U., Ekman, S., Fröberg, L., Knutsson, T. &
Mattsson, J. E. 1989. Changes in the lichen flora
on Romeleklint, S. Sweden, over a 50-year period.
Graphis Scripta 2(4): 148–155.
Kotlov, Yu. V. 2003. Lichen species from Sverdlovsk
region new to the Urals (in Russian). Botanicheskii
Zhurnal 88(4): 145–146.
Kruckeberg, A. R. 1954. The ecology of serpentine
soils. III. Plant species in relation to serpentine
soils. Ecology 35: 267–274.
Proctor, J. & Woodell, S. R. J. 1971. The plant ecology
of serpentine. J. Ecol. 59(2): 375–411.
Paukov, A. G., & Trapeznikova, S. N. 2003. Lithophilous lichens new for the Urals flora (in Russian).
Botanicheskii Zhurnal 88(2): 104–109.
Ryabkova, K. A. 1998. A systematic list of Ural’s
lichens (in Russian). Novitates Systematicae
Plantarum non Vascularium 32: 81–87.
Urals and surrounding areas (in Russian). 1968.
Moscow. 459 pp.
Voitkevich, G. V., Kokin, A. V., Miroshnikov, A. E.,
Prokhorov, V. G. 1990. Guide on geochemistry (in
Russian). Moscow. 480 pp.
APPENDIX
The list of lichens of Middle Urals
Abbreviations of substrates: bas. – basalt, concr.
– concrete, gr. – granite, limest. – limestone, pyr.
– pyroxenite, serp. – serpentine.
Rr1, rr2, rr3 are the rarity groups described
in Material and Methods.
ACAROSPORA BADIOFUSCA (Nyl.) Th. Fr. – bas.
A. FUSCATA (Schrad.) Arnold – serp., pyr., bas.,
gr.
A. GLAUCOCARPA (Wahlenb.) Körb. – limest.
A. MACROSPORA (Hepp) Bagl. – limest.
A. NITROPHILA H. Magn. – serp., pyr., bas.
A. OLIGOSPORA (Nyl.) Arnold – serp.; rr1.
AMANDINEA PUNCTATA (Hoffm.) Coppins & Scheid.
– serp., bas.
ANEMA JENISEJENSIS H. Magn. – limest.
ARCTOPARMELIA CENTRIFUGA (L.) Hale – gr.
A RTHONIA LAPIDICOLA (Taylor) Branth & Rostr.
– concr.; rr1.
ASPICILIA CINEREA (L.) Körb. – serp., pyr., bas., gr.
A. CONTORTA (Hoffm.) Krempelh. – limest., serp.,
pyr.
A. CONTORTA SSP. HOFFMANNIANA S. Ekman & Fröberg
– serp., pyr., bas.
86
A.
Folia Cryptog. Estonica
(Krempelh.) Mereschk. – serp.,
pyr.; rr1.
A. LAPPONICA (Zahlbr.) Oxner – serp., pyr., bas.
A. MOENIUM (Vain.) Thor &Timdal – concr.,
limest.
A. CF. PERRADIATA (Nyl.) Hue – bas.
A. SIMOËNSIS Räsänen – serp., bas.
A. CF. SUPERTEGENS Arnold – bas.
A. CF. VERRUCIGERA (Hue) Zahlbr. – serp., pyr.,
bas., gr.
A. TRANSBAICALICA Oxner – bas.
BACIDIA HERBARUM (Stizenb.) Arnold – mosses on
serp.; rr2.
B. INUNDATA (Fr.) Körb. – serp.; rr1.
BACIDINA ARNOLDIANA (Körb.) V. Wirth & Vězda
– limest.; rr1.
BAEOMYCES RUFUS (Huds.) Rebent. – gr.
BELLEMEREA CUPREOATRA (Nyl.) Clauzade & Cl. Roux
– serp., pyr., bas., gr.
BRYORIA SIMPLICIOR (Vain.) Brodo & D. Hawksw.
– gr.; rr1.
BUELLIA LEPTOCLINE (Flot.) A. Massal. – bas.; rr1.
CALOPLACA ARENARIA (Pers.) Müll. Arg. – serp.,
bas.
C. CERINA VAR. CHLOROLEUCA (Sm.) Th. Fr. – debris
on limest.
C. C HR Y S ODE T A (Vain. & Räsänen) Dombr.
– limest.; rr3.
C. CIRROCHROA (Ach.) Th. Fr. – limest.
C. CRENULARIA (With.) J.R. Laundon – gr.
C. DIPHYODES (Nyl.) Jatta – bas.; rr1.
C. DOLOMITICOLA (Hue) Zahlbr. – limest.
C. FLAVOVIRESCENS (Wulfen) Dalla Torre & Sarnth.
– serp., pyr., bas., limest.
C. GRIMMIAE (Nyl.) H. Olivier – On Candelariella
vitellina.
C. HOLOCARPA (Ach.) A.E. Wade – serp., pyr.,
bas.
C. JUNGERMANNIAE (Vahl.) Th. Fr. – debris on
limest.; rr1.
C. LACTEA (Massal.) Zahlbr. – limest.
C. SAXICOLA (Hoffm.) Nordin – limest., bas.
C. SUBPALLIDA H. Magn. – bas., serp., pyr.
C. VARIABILIS (Pers.) Müll. Arg. – limest.
C. VITELLINULA (Nyl.) H. Olivier – serp., bas.,
limest.
CANDELARIELLA MEDIANS (Nyl.) A.L. Sm. – limest.;
rr1.
C. VITELLINA (Ehrh.) Müll. Arg. – gr., serp., pyr.,
bas., limest.
CATAPYRENIUM SP. – limest.
CETRELIA OLIVETORUM (Nyl.) W.L. Culb. & C.F. Culb.
– gr.; rr3.
DESERTORUM
CHROMATOCHLAMYS MUSCORUM (Fr.) H. Mayrhofer &
Poelt – moss on pyr.; rr1.
CHRYSOTHRIX CHLORINA (Ach.) J.R. Laundon – gr.,
serp.
CLADONIA ACUMINATA (Ach.) Norrl. – soil on limest.
C. AMAUROCRAEA (Flörke) Schaer. – soil on serp.
C. CHLOROPHAEA (Flörke) Spreng. – soil on serp.,
pyr., gr., bas.
C. CONIOCRAEA (Flörke) Spreng. – soil on serp.,
gr.
C. POCILLUM (Ach.) Grognot – soil, moss on
limest.
C. PYXIDATA (L.) Hoffm. – soil, moss on gr., serp.,
pyr., limest., bas.
C. RAMULOSA (With.) J.R. Laundon – gr.
CLAUZADEA MONTICOLA (Schaer.) Hafellner & Bellem.
– limest.; rr1.
COLLEMA AURIFORME (With.) Coppins & J.R. Laundon – moss on limest.
C. CRISPUM (Huds.) F.H. Wigg. – limest.; rr2.
C. CRISTATUM (L.) F.H. Wigg. – serp., bas.,
limest.
C. FLACCIDUM (Ach.) Ach. – serp., bas., limest.;
rr2.
C. FUSCOVIRENS (With.) J.R. Laundon – limest.
C. GLEBULENTUM (Nyl. ex Cromb.) Degel. – limest.;
rr1.
C. POLYCARPON Hoffm. – serp., pyr., limest.
C. TENAX (Sw.) Ach. – soil on serp., pyr., limest.,
bas.
DERMATOCARPON MINIATUM (L.) W. Mann. – serp.,
bas.
DIBAEIS BAEOMYCES (L.) Rambold & Hertel – serp.,
pyr., gr.
DIMELAENA OREINA (Ach.) Norman – bas., gr.
DIPLOSCHISTES MUSCORUM (Scop.) R. Sant. – moss
on limest., serp.
D. SCRUPOSUS (Schreb.) Norman – gr., pyr., bas.
DIPLOTOMMA ALBOATRUM (Hoffm.) Flot. – limest.;
rr1.
ENDOCARPON PUSILLUM Hedw. – soil on limest.,
serp.
EVERNIA MESOMORPHA Nyl. – serp., pyr., bas.
FLAVOPARMELIA CAPERATA (L.) Hale – pyr.; rr3.
FLAVOPUNCTELIA SOREDICA (Nyl.) Hale – bas.
FULGENSIA BRACTEATA (Hoffm.) Räsänen – limest.;
rr1.
FUSCOPANNARIA LEUCOPHAEA (Vahl) P.M. Jørg. – serp.;
rr1.
F. PRAETERMISSA (Nyl.) P.M. Jørg. – serp.; rr1.
GYALIDEA FRITZEI (B. Stein) Vězda – gr.; rr1.
HETERODERMIA SPECIOSA (Wulfen) Trevis. – moss on
serp., pyr.; rr2.
87
HYPOGYMNIA PHYSODES (L.) Nyl. – gr., serp., pyr.,
bas.
LECANIA ERYSIBE (Ach.) Mudd – limest.
L. TURICENSIS (Hepp) Müll. Arg. – limest.
LECANORA CAMPESTRIS (Schaer.) Hue – serp., pyr.,
gr., bas.
L. CRENULATA Hook. – limest.
L. DISPERSA (Pers.) Sommerf. – serp., pyr., gr.,
bas.
L. FRUSTULOSA (Dicks.) Ach. – serp., pyr., bas.,
limest.
L. INTRICATA (Ach.) Ach. – pyr., gabbro.
L. MURALIS (Schreb.) Rabenh. – gr., serp., pyr.,
bas., limest.
L. POLYTROPA (Hoffm.) Rabenh. – serp., pyr., gr.,
bas.
L. C F . X A N T H O S T O M A Cl. Roux ex Fröberg
– limest.
LECIDEA ATOMARIA Th. Fr. – serp., pyr.; rr1.
LECIDELLA CARPATHICA Körb. – serp., pyr., bas.,
limest.
L. STIGMATEA (Ach.) Hertel & Leuckert – gr., serp.,
pyr., bas., limest.
LEPRARIA INCANA (L.) Ach. – serp., pyr., bas.
L. MEMBRANACEA (Dicks.) Vain. – serp., pyr., bas.,
gr., limest.
L. RIGIDULA (de Lesd.) Tønsberg – gr.
LEPTOGIUM LICHENOIDES (L.) Zahlbr. – limest.; rr1.
LEPTOGIUM SP. – limest.
L. TENUISSIMUM (Dicks.) Körb. – soil on serp.,
bas.
LICHINELLA STIPATULA Nyl. – serp., pyr., bas.
MELANALIA OLIVACEA (L.) Essl. – pyr.
M. DISJUNCTA (Erichsen) Essl. – gr., pyr., bas.
M. EXASPERATULA (Nyl.) Essl. – serp., pyr.
M. INFUMATA (Nyl.) Essl. – pyr., gr., serp., bas.
M. PANNIFORMIS (Nyl.) Essl. – gr.; rr1.
M. SUBARGENTIFERA (Nyl.) Essl. – serp.
M. TOMINII (Oxner) Essl. – pyr., bas.
MICAREA ERRATICA (Körb.) Hertel, Rambold & Pietschm. – serp., gr.
M. LAPILLICOLA (Vain.) Coppins & Muhr – pyr.
M. SYLVICOLA (Flot.) Vězda & V. Wirth – gr.; rr1.
M. TUBERCULATA (Sommerf.) R.A. Anderson – gr.;
rr1.
MIRIQIUDICA LEUCOPHAEA (Rabenh.) Hertel & Rambold – gr.; rr1.
MYCOBILIMBIA CARNEOALBIDA (Müll. Arg.) Printzen
– moss on limest.
M. MICROCARPA (Th. Fr.) Brunnb. – moss on
limest.
M. SABULETORUM (Schreb.) Hafellner – moss on
limest.
M.
TETRAMERA
limest.
(De Not.) Vitik. et al. – moss on
PARMELIA FRAUDANS Nyl. – pyr., bas., gr.
P. SAXATILIS (L.) Ach. – gr.; rr1.
P. SULCATA Tayl. – gr., serp., pyr., bas., limest.
PECCANIA CORALLOIDES (A. Massal.) A. Massal.
– limest.; rr1.
PELTIGERA LEPIDOPHORA (Vain.) Bitter – soil on
serp., bas.
P. RUFESCENS (Weiss) Humb. – soil on limest.,
bas.
PELTULA CF. EUPLOCA (Ach.) Poelt – serp.
PERTUSARIA ALBESCENS (Huds.) M. Choisy & Werner
– serp., gr.
PHAEOPHYSCIA CONSTIPATA (Norrl. & Nyl.) Moberg
– moss on limest., serp.
P. NIGRICANS (Flörke) Moberg – serp.
P. SCIASTRA (Ach.) Moberg – gr., serp., pyr., bas.,
limest.
PHLYCTIS ARGENA (Spreng.) Flot. – limest.; rr3.
PHYSCIA ADSCENDENS (Fr.) H. Olivier – pyr.
PH. CAESIA (Hoffm.) Fürnr. – gr., serp., pyr., bas.,
limest.
PH. DUBIA (Hoffm.) Lettau – gr., serp., pyr., bas.
PH. SP. – bas.; rr1.
PHYSCONIA DETERSA (Nyl.) Poelt – serp., pyr., bas.
PH. MUSCIGENA ( Ach.) Poelt – serp., pyr., bas.
P H . PERISIDIOSA (Erichsen) Moberg – limest.,
serp.
PLACYNTHIELLA ICMALEA (Ach.) Coppins & P. James
– gr.
P. ULIGINOSA (Schrad.) Coppins & P. James – gr.
PLACYNTHIUM NIGRUM S. Gray – limest.
POLYSPORINA SIMPLEX (Davies) Vězda – gr., bas.
PORPIDIA CINEREOATRA (Ach.) Hertel & Knoph – gr.,
serp.
P. CRUSTULATA (Ach.) Hertel & Knoph – serp., pyr.,
gr., bas., limest.
P. AFF. GLAUCOPHAEA (Körb.) Hertel & Knoph – gabbro; rr2.
P. MACROCARPA (DC.) Hertel & A.J. Schwab – gr.
P. CF. SOREDIZODES (Lamy ex Nyl.) J.R. Laundon
– gr.; rr1.
P. TUBERCULOSA (Sm.) Hertel & Knoph – bas.;
rr1.
P ROTOBLASTENIA RUPESTRIS (Scop.) J. Steiner
– limest.
PROTOPANNARIA PEZIZOIDES (Weber) P.M. Jørg. & S.
Ekman – soil on gr.
PSORA ELENKINII Rass. – limest.
RAMALINA POLLINARIA (Westr.) Ach. – serp.
RHIZOCARPON BADIOATRUM (Spreng.) Th. Fr. – gr.,
pyr., bas.
88
Folia Cryptog. Estonica
R. GRANDE (Flörke) Arnold – gr., serp., pyr., bas.
R. OBSCURATUM (Ach.) A. Massal. – gr., pyr.
R. CF. PLICATILE (Leighton) A. L. Sm. – gr.; rr1.
R. SUBGEMINATUM Eitner – gr.; rr1.
RHIZOPLACA CHRYSOLEUCA (Sm.) Zopf – bas.
RINODINA BISCHOFII (Hepp) A. Massal. – limest.
R. CALCAREA (Hepp) Arnold – limest.
R. CONFRAGOSA (Ach.) Körb. – serp., pyr., bas.
R. MILVINA (Wahlenb.) Th. Fr. – serp., pyr.,
limest.
SARCOGYNE PRIVIGNA (Ach.) Anzi VAR. CALCICOLA H.
Magn. – limest.; rr3.
S. REGULARIS Körb em. Oxner – limest.
SCOLICIOSPORUM UMBRINUM (Ach.) Arnold – serp.,
pyr., gr., bas.
SOLORINA SACCATA (L.) Ach. – soil on limest.; rr1.
STAUROTHELE FUSCOCUPREA (Nyl.) Zschacke – bas.;
rr1.
STEREOCAULON SAXATILE H. Magn. – gabbro.
S. TOMENTOSUM Fr. – gr., pyr.
THYREA CF. CONFUSA Henssen – limest.
TONINIA OPUNTIOIDES (Vill.) Timdal – limest.; rr1.
T. CINEREOVIRENS (Schaer.) A. Massal. – serp.;
rr2.
T RAPELIA COARCTATA (Sm.) M. Choisy – gr.
T. CF. INVOLUTA (Taylor) Hertel – bas.; rr1.
T. CF. OBTEGENS (Th. Fr.) Hertel – gr.; rr1.
T.
PLACODIOIDES
rr2.
Coppins & P. James – gr, bas.;
TUCKERMANNOPSIS SEPINCOLA (Ehrh.) Hale – pyr.,
gr.
UMBILICARIA DEUSTA (L.) Baumg. – gr., pyr.
U. ROSSICA (Dombr.) Golubk. – bas.; rr1.
VERRUCARIA CAERULEA DC. – serp.
V. CALCISEDA DC. – limest.
V. HYDRELA Ach. – pyr. in springs.
V. CF. LEIGHTONII A. Massal. – concr.
V. MURALIS Ach. – serp., pyr.
V. CF. MUTABILIS Borr. – limest.
V. NIGRESCENS Pers. – limest., concr.
V. CF. VIRIDULA (Schrad.) Ach. – serp.
VULPICIDA PINASTRI (Scop.) J.-E. Mattsson & M.J.
Lai – gr., pyr.
XANTHOPARMELIA CAMTSCHADALIS (Ach.) Hale – soil
on limest.; rr1.
X. CONSPERSA (Ach.) Hale – pyr., gr., serp., bas.
X. SOMLOËNSIS (Gyeln.) Hale – pyr., gr., serp.,
bas., limest.
X. TINCTINA (Maheu & A. Gillet) Hale – gr., pyr.
XANTHORIA ELEGANS (Link) Th. Fr. – bas., serp.,
pyr., limest.
X. FALLAX (Hepp) Arnold – pyr., bas., limest.
X. SOREDIATA (Vain.) Poelt – serp., pyr., bas.,
limest.
Folia Cryptog. Estonica, Fasc. 41: 89–96 (2005)
Distribution patterns of primary and secondary species in the
genus Vulpicida
Tiina Randlane & Andres Saag
Institute of Botany and Ecology, University of Tartu, Lai Street 38, 51005, Tartu, Estonia
E-mail: Tiina.Randlane@ut.ee
Abstract: World-wide distribution maps are presented for all six Vulpicida species as well as for the groups of primary
and secondary species separately. Taxa which usually bear apothecia (V. canadensis, V. juniperinus and V. viridis) are treated
as primary species; the status of secondary species is assigned in this paper to the sorediate taxon (V. pinastri) as well as to
these two species (V. tilesii and V. tubulosus) that probably reproduce mainly vegetatively by the fragments of thalli. There are
two north-american endemics and one eurasian taxon among primary species, and two circumboreal taxa plus one european
taxon among secondary species. The idea that the ancestor of Vulpicida may have had a circumpolar distribution, and later
speciation processes took place in Eurasia as well as in North America may explain the contemporary distribution patterns of
Vulpicida species. The shape of pycnidiospores (differing in two north-american endemics and other four species) supports
the speculation that the former taxa have evolved apart from the latter, which probably developed somewhere in Eurasia,
and some of them (V. pinastri, V. tilesii) have distributed secondarily to North America.
Kokkuvõte: Rebasesamblike (Vulpicida) primaarsete ja sekundaarsete liikide levikumustrid.
Esitatakse kõigi kuue Vulpicida liigi levikukaardid, samuti eraldi koondkaardid primaar- ja sekundaarliikide leviku kohta. Liike,
mis tavaliselt moodustavad apoteetsiume, käsitletakse primaarliikidena (V. canadensis, V. juniperinus ja V. viridis); sekundaarliigi
staatus on selles töös antud soredioossele taksonile (V. pinastri) ning neile kahele liigile (V. tilesii ja V. tubulosus), mis tõenäoliselt
paljunevad vegetatiivselt talluse tükkide abil. Primaarsete liikide hulgas on kaks põhja-ameerika endeemi ja üks euraasia takson
ning sekundaarsete liikide hulgas kaks tsirkumboreaalset ning üks euroopa takson . Idee, et rebasesamblike eellane võis olla
levinud tsirkumpolaarselt ja edasine liigiteke toimus nii Euraasias kui ka Põhja-Ameerikas, seletaks Vulpicida liikide tänapäevast
levikumustrit. Püknidiospooride kuju (erinev kahel põhja-ameerika endeemil ja ülejäänud neljal liigil) toetab seisukohta, mille
kohaselt põhja-ameerika liigid arenesid eraldi ülejäänutest; viimaste areng toimus tõenäoliselt kusagil Euraasias ning osa neist
(V. pinastri, V. tilesii) võisid uuesti levida Põhja-Ameerikasse.
INTRODUCTION
The heterogeneous group of cetrarioid lichens
(fam. Parmeliaceae, Ascomycota) including
about 150 species from more than 20 genera
comprises eight genera in which both primary
and secondary species are represented –
Allocetraria, Cetrelia, Cetreliopsis, Platismatia,
Tuckermanella, Tuckneraria, Tuckermannopsis
and Vulpicida (Randlane & Saag, 2004).
World-wide distribution maps for four of them,
mainly Asian genera (Allocetraria, Cetrelia,
Cetreliopsis and Tuckneraria), were presented
lately (Randlane & Saag, 2004) while the
distribution of three further, mainly North
American genera, Platismatia, Tuckermanella
and Tuckermannopsis, will be analysed in
another paper.
Here we focus on the distribution of species
from the genus Vulpicida: V. canadensis
(Räsänen) J.-E. Mattsson & M. J. Lai, V.
juniperinus (L.) J.-E. Mattsson & M. J. Lai,
V. pinastri (Scop) J.-E. Mattsson & M. J. Lai,
V. tilesii (Ach.) J.-E. Mattsson & M. J. Lai, V.
tubulosus (Schaerer) J.-E. Mattsson & M. J.
Lai, and V. viridis (Schwein.) J.-E. Mattsson &
M. J. Lai.
Genus Vulpicida was described and thoroughly
characterized more than ten years ago (Mattsson & Lai, 1993; Mattsson, 1993). Six cetrarioid
taxa with intensely yellow coloured foliose or
subfruticose thallus were incorporated in this
unit. The main diagnostic character of the genus
is the presence of pinastric and vulpinic acids in
the medulla. Broadly clavate asci and broadly
ellipsoid to subglobose ascospores are also of
importance. Still, phylogenetic analyses based
on morphological (s. lat.) characters (Mattsson,
1993; Saag et al., 2002) reveal that Vulpicida is
not quite homogenous: it includes two separate
clades which anatomically differ in the shape
of pycnidiospores (citriform in North-American
endemics V. canadensis and V. viridis and
90
Folia Cryptog. Estonica
sublageniform in other four species). Analyses
using molecular characters (Saag et al., 2002;
Mattsson & Articus, 2004) group the species
inside the genus differently. As only four species of the six have been sequenced till now,
the topography of cladograms can be changed
in future analyses.
Both primary and secondary species are
represented in the genus.
The concept of species pairs was first and clearly
formulated by Du Rietz in 1924 who used the
term “Artenpaaren” for indicating the phenomenon that pairs of lichen species exist which
differ from each other in just one character
– having or lacking soredia (or isidia). He also
drew attention to the fact that sorediate and
usually sterile taxa have a different ecology and
distribution from their nonsorediate fertile allies
(Mattsson & Lumbsch, 1989). Hale (1965) pointed out that sorediate species often have a wider
distribution than their fertile counterparts. Poelt
(1970, 1972) expanded the species pair concept
and suggested that to every sorediate taxon corresponds a nonsorediate one, considering that
sorediate taxa represent evolutionary dead ends.
It was also pointed out that all taxa with sexual
reproduction appear to be quite rare with very
restricted distribution. Still, this is not the absolute rule. For example, two primary Allocetraria
species (A. madreporiformis, A. stracheyi) have
spread from the probable speciation centre of the
genus in Asia to North America. Furthermore,
if several sorediate cetrarioid lichens occur extensively on two, three or even four continents,
then isidiate species are clearly less successful
in their distribution. For example, of cetrarioid
isidiate taxa which evidently have evolved in Asia
(Allocetraria isidiigera, Cetrelia braunsiana, C.
isidiata and Tuckneraria togashii), all four have
“stayed” in this continent (Randlane & Saag,
2004).
In Vulpicida one sorediate species (V. pinastri)
is known, and three taxa which usually bear
apothecia (V. canadensis, V. juniperinus and V.
viridis). They can be treated as secondary and
primary species, respectively. However, it is not
as simple as it seems to distinguish between
primary and secondary species using the presence of apothecia versus soredia or isidia only.
In Vulpicida there are some species which lack
vegetative propagules of reproduction but for
which apothecia have been observed occasionally
only. Both such taxa (V. tilesii and V. tubulosus)
grow on the ground – differently from the other
species in the genus which are epiphytic – and
presumably reproduce through the fragmentation of thalli. In this paper we assign the status
of secondary species to the sorediate taxon (V.
pinastri) as well as to these two species (V. tilesii
and V. tubulosus) that probably reproduce mainly
vegetatively by the fragments of thalli.
MATERIAL AND METHODS
Distribution maps were compiled using the
computer program DMAP. Both herbarium
and literature data were used as sources for
the localities. The materials from the following
herbaria were taken into consideration: B, GZU,
H, LD, LE, TU, UPS, and the collections of Irina
Skirina, Vladivostok (from the Far East of Russia). Data of the checklists published on the
internet (Feuerer, 2004) were accounted only
in exceptional cases, when no other data were
available. The literature sources for distribution
data of each lichen species are listed below.
Vulpicida canadensis: Brodo et al., 2001; Mattsson, 1993; Nash III et al., 2002.
V. juniperinus: Azuaga & Gomez-Bolea, 1996,
2000; Barkhalov, 1983; Biazrov et al.,
1989; Davydov, 2001; Fadeyeva et al.,
1997; Feuerer, 2004; Hermansson et al.,
1998; Himelbrant et al., 2001; Kotlov, 1995;
Kurokawa, 2003; Martellos & Nimis, 2001;
Mattsson, 1993; Randlane & Saag, 1999;
Randlane et al., 2001; Santesson et al.,
2004; Scholz, 2000; Urbanavichene, 1998;
Wei, 1991; Zavarzin et al., 1999.
V. pinastri: Azuaga & Gomez-Bolea, 2000;
Barkhalov, 1983; Biazrov et al., 1989; Brodo
et al., 2001; Davydov, 2001; Diederich &
Sérusiaux, 2000; Fadeyeva et al., 1997;
Feuerer, 2004; Gorbatch, 1973; Hafellner
& Türk, 2001; Hermansson et al., 1998;
Himelbrant et al., 2001; Kopatchevskaya,
1986; Kotlov, 1995; Kurokawa, 2003;
Makarova, 1998; Makarova et al., 2002;
Mattsson, 1993; Motiejunaite, 1999; Piterans, 2001; Pisút et al., 1996; Poryadina,
2001; Randlane & Saag, 1999; Santesson
et al., 2004; Sorokina, 2001; Suppan et al.,
2000; Urbanavichene, 1998; Vězda & Liska,
1999; Wei, 1991; Zhuravleva & Zhigunov,
2002; Zhurbenko & Vechov, 2001.
91
V. tilesii: Biazrov et al., 1989; Brodo et al., 2001;
Davydov, 2001; Glew, 2004; Hermansson
et al., 1998; Kotlov, 1994, 1995; Mattsson,
1993; Urbanavichene, 1998.
V. tubulosus: Fadeyeva et al., 1997; Hafellner &
Türk, 2001; Martellos & Nimis, 2001; Mattsson, 1993; Pisút et al., 1996; Randlane &
Saag, 1999; Santesson et al., 2004; Suppan
et al., 2000; Vězda & Liska, 1999.
V. viridis: Brodo et al., 2001; Mattsson, 1993.
RESULTS
Vulpicida canadensis (Fig.
1) is endemic to western
North America and is distributed in Canada (British
Columbia, Alberta), Mexico
(Baja California) and USA
(Washington, Idaho, Montana, Oregon, California)
only. The lichen grows on
bark and wood of mainly
conifers (Pinus, Pseudotsuga, Picea, Abies) in open
and relatively dry sites.
V. juniperinus (Fig. 2)
is distributed mainly in
northern Europe (Estonia,
Finland, Norway, Russia,
Sweden), but known also
from a few isolated sites
in subalpine areas of central or southern Europe Fig. 1. World distri(Andorra, Germany, Italy, bution of Vulpicida
Spain, United Kingdom), canadensis – on
and sparsely in Asia (Chi- the western coast
na, Georgia, India, Japan, of North America
Mongolia, Russia, South ( – after Mattsson,
Korea). The species has 1993; – complebecome today very rare in mented).
southern Fennoscandia, in
one of its main distribution regions (Thell et al.,
2004). It is an epiphyte that is mainly restricted
to Juniperus communis in Europe and to Pinus
pumila in Asia; occasionally grows also on Betula
or on calcareous soil (Randlane et al., 2001).
V. pinastri (Fig. 3) is a circumboreal lichen which
occurs in subalpine, subarctic and boreal zones
of North America (Canada, Greenland, USA),
Europe (Andorra, Austria, Belgium, Bulgaria,
Byelorussia, Czech Republic, Denmark, Estonia, Finland, France, Germany, Hungary,
Iceland, Italy, Latvia, Lithuania, Luxembourg,
Netherlands, Norway, Poland, Portugal, Romenia, Russia, Slovakia, Slovenia, Spain, Sweden,
Switzerland, Ukraine, United Kingdom) and Asia
(Armenia, China, Georgia, Japan, Kazakhstan,
Mongolia, Russia, Turkey). It grows mainly on
branches and trunks of coniferous trees (e.g.
Abies, Juniperus, Larix, Picea, Pinus, in Europe
preferably on Pinus sylvestris) and on deciduous
trees with acidic bark (e.g. Alnus, Betula, Populus etc.), occasionally also on lignum, siliceous
rocks and soil.
V. tilesii (Fig. 4) is an arctic-alpine species distributed mainly in the north-western part of
North America (Canada, Greenland, USA) and
northern part of Asia (Kazakhstan, Mongolia,
Russia); in Europe only a few localities are
known from its very eastern areas (Russia). It is
a terricolous lichen occurring on exposed, periodically wet calciferous soil (Mattsson, 1993).
V. tubulosus (Fig. 5) has a disjunct distribution
area, being found abundantly on the Baltic
Isles and adjacent territories (Estonia, Finland,
Norway, Sweden, Russia), and in the alpine regions of Central Europe (Austria, Czech Republic, France, Germany, Italy, Poland, Romania,
Slovenia, Slovakia, Spain, Switzerland). The
northernmost localities are known from Torne
Lappmark in Sweden (≈ 68°N), where it is partly
extinct (Santesson et al., 2004).
V. viridis (Fig. 6) occurs in the south-eastern
part of North America (USA), close to the Atlantic coast. The species is endemic to this
area; recent findings of this taxon in Russian
Fig. 2. World distribution of Vulpicida juniperinus
( – after Mattsson, 1993; – complemented).
92
Folia Cryptog. Estonica
Fig. 3. World distribution of Vulpicida pinastri ( – after Mattsson, 1993; – complemented).
Fig. 4. World distribution of Vulpicida tilesii ( – after Mattsson, 1993; – complemented).
Far East, Primorye region, close to the Pacific
coast (Tchabanenko, 2002; Randlane et al.,
2004) turned out to be misidentifications. It
is an epiphytic lichen which mainly grows in
humid conditions on Chamaecyparis thyoides
(in bogs along the north-eastern coastal plain)
or mature trees of Quercus in the south (Brodo
et al., 2001).
DISCUSSION
Fig. 5. World distribution of Vulpicida tubulosus
( – after Mattsson, 1993; – complemented).
Fig. 6. World distribution
of Vulpicida viridis – on
the eastern coast of North
America (after Mattsson,
1993).
None of the primary species (V. canadensis, V.
juniperinus and V. viridis) occurs simultaneously in Europe, Asia and North America. V.
juniperinus has the widest distribution area of
the three, occurring abundantly in Europe and
scarcely in wide territories of Asia. Two other
primary taxa, V. canadensis and V. viridis have
rather restricted distribution areas; although
they are both endemic to North America, their
distributions do not overlap.
93
All primary species are predominantly epiphytic, however, V. juniperinus can occasionally
grow also on soil.
epiphytic V. pinastri shows the widest ecological amplitude growing on various tree species,
lignum, and rarely on rocks or soil.
Among the secondary species, the distribution
area of the sorediate V. pinastri is the widest,
covering North America, Europe and a major
part of Asia. V. tilesii has also rather a wide
distribution in North America and northern Asia
(with only a few localities in the very eastern
part of Europe). The third secondary species, V.
tubulosus, has a restricted and disjunct distribution area, with both parts of its distribution
only in Europe.
Epiphytic and epigeic lichens are present
among secondary species of Vulpicida. The
The distribution patterns of primary (Fig. 7) and
secondary species of Vulpicida (Fig. 8) do not
refer unambiguously to the biodiversity or speciation centres of the genus. If secondary taxa
and especially sorediate lichens usually occur
on different continents, then primary species
are either too specialized (to the substratum,
ecological conditions etc.) or unable to cross the
wide distances between the continents. In this
genus there are two north-american endemics
and one eurasian taxon among primary species,
and two circumboreal taxa plus one european
Fig. 7. Distribution patterns of primary Vulpicida species.
Fig. 8. Distribution patterns of secondary Vulpicida species.
94
Folia Cryptog. Estonica
taxon among secondary species. The idea that
the ancestor of Vulpicida may have had a circumpolar distribution (Mattsson 1993), and
later speciation processes took place in Eurasia
as well as in North America may explain the contemporary distribution patterns of Vulpicida species. The shape of pycnidiospores which differs
in two north-american endemics (citriform) and
other four species (sublageniform) supports the
speculation that the former taxa have evolved
apart from the latter, probably somewhere in
Eurasia, and some of them (V. pinastri, V. tilesii)
have distributed secondarily to North America.
ACKNOWLEDGEMENTS
The authors are grateful to the keepers of herbaria mentioned in the text for kindly sending
the lichen specimens. Special thanks are due
to Maria Angeles Herrera-Campos who provided
us with the distribution data of Vulpicida species from Mexico. This study was financially
supported by the Estonian Science Foundation
(grants 5505 and 5823).
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Folia Cryptog. Estonica, Fasc. 41: 97–104 (2005)
Some lichens have incomplete distribution ranges in the Aegean
(Greece)
Harrie J. M. Sipman¹, Th. Raus¹ & A. Scharlau²
¹Botanic Garden & Botanical Museum Berlin-Dahlem, Free University of Berlin, Berlin, Germany
²P. O. Box 81, GR-84300 Naxos, Greece
Abstract: An investigation of the epiphytic lichen flora on two comparable islands in the Aegean Sea (Greece) showed that
some of the common species are present on one of the islands but absent from the other. This suggests that they failed to
colonize all suitable habitats in the region and that insular isolation has a similar reducing effect on lichen diversity as it has
for other organism groups. Further observations, which support dispersal restrictions, are that vegetative diaspore frequency
deviates from that in continental areas, and that chemical strains of Pseudevernia furfuracea occupy separate ranges. A list of
208 epiphytic lichens encountered on the islands Ikaria and Naxos is appended.
Kokkuvõte: Mõnede samblike levik on Egeuse piirkonnas (Kreeka) lünklik.
Kahe Egeuse meres asuva Kreeka saare samblike uurimine näitas, et mõned tavalised liigid ühel saartest esinevad, teisel mitte.
See tekitab arvamuse, et nad pole suutnud asustada kõiki selle regiooni sobivaid substraate ja et saareline eraldatus vähendab
samblike mitmekesisust samuti nagu teistelgi organismirühmadel. Edasised, leviku piiranguid toetavad uuringud näitavad
vegetatiivsete diaspooride sageduse erinevust sellest, mis mandriosas on tavaline, ja et hariliku karesambliku (Pseudevernia
furfuracea) erinevad keemilised rassid on erineva levikuga. Lisatud on Ikaria ja Naxose saarte epifüütsete samblike nimestik,
mis koosneb 208 liigist.
INTRODUCTION
For a long time it is known that islands have
a reduced flora and fauna. This is commonly
explained by difficult access and increased risk
for extinction because of the small population
sizes. Lichens, however, as spore-producing organisms, are supposed to be easily dispersed,
and their presence or absence to be determined
by the availability of suitable habitats rather
than geographical connections or other forms
of easy access. Indeed, Hayward & Hayward
(1986) conclude in an investigation of island
lichen biota that the number of species present
is correlated with the availability of different
habitats. However, there are also indications
that spatial isolation might play a role in lichen
diversity. Warren (2003) reports that increasing
distance reduces genetic exchange, and Degelius
(1986) concludes that the remote island of Anholt is mainly colonized by late immigrants. He
gives no further interpretation of his observation
but it bears the suggestion that other species
have been unable to reach the island.
An ongoing project to inventory the lichen flora
of individual islands in the Aegean See, Greece,
by the first two authors (Sipman & Raus, 1995,
1999, 2002) provided an opportunity to test the
effect of insular isolation on lichen diversity. The
Aegean Sea, situated between the mainland of
Greece and Turkey, contains many islands, remains of a submersed mountain system (Higgins
& Higgins, 1996). These share a Mediterranean,
rather arid climate, but differ considerably as to
the availability of bedrock. Some are exclusively
composed of limestone, others have large intrusions of granite or metamorphic rocks, while a
few have volcanic rocks. Consequently the total
lichen diversity of the islands is much influenced
by the available rock substrate. The epiphytic
substrates, however, show much less variation,
because the number of available tree species is
very restricted (Appendix 1). Moreover, epiphytic
lichens tend to be unspecialized and grow on a
wide range of tree species. Thus the epiphytic
diversity seems to be less influenced by geological differences between the islands and more
suitable for a study of insular effects.
For an estimation whether islands have an
impoverished lichen flora, it would seem most
suitable to compare them with an equally sized
continental area. However, the adjacent continental areas of the Aegean are very mountainous
and exceed the islands considerably in altitu-
98
Folia Cryptog. Estonica
dinal range. Also the climate is different from
the islands, which have a pronounced maritime
climate. This makes considerable differences in
lichen flora probable. Therefore a comparison
between two islands with comparable size and
elevation, situated close enough to assure a
climatic similarity, seemed more appropriate
to investigate isolation effects. When there is
an impoverishment, it is unlikely that it has
exactly the same effect in two islands, but it
is more probable that some species are absent
from one island and other species absent from
the other island.
MATERIAL AND METHODS
Epiphytic lichen inventories were available for
two islands, Naxos, coordinates 37°03’ N, 25°28’
E and area 428 km², and Ikaria, coordinates
37°35’ N, 26°10’ E and area 255 km² (Fig. 1),
situated in the central part of the Aegean sea
and about 60 km apart from each other.
The epiphytic lichens of Naxos were studied
in detail by the third author during three months
in 1995 (Scharlau, 1996). The island is largely
covered by mountains reaching to about 900 m
elevation and composed of various rock types,
including limestone, granite and gneiss. Trees
are available mainly in cultivated areas in the
higher valleys and on the mountain slopes. The
author investigated 304 individual trees belonging to 22 phorophytes (Appendix 1) in 55 localities. In total some 4000 observations of 148 taxa
were made (Appendix 2).
Ikaria is very similar to Naxos, and differs
mainly in shape. It is elongated rather than
rounded and W-E-oriented, so that clouds
from the sea driven by the prevailing northerly
winds (called “meltémia”) have a stronger effect.
Fig. 1. The Aegean sea with the investigated islands. 1 – Naxos; 2 – Ikaria.
99
It was surveyed by the first two authors, who
made a lichen inventory of all available habitats in 44 localities during two weeks in 2002,
including 18 phorophytes (Appendix 1). Some
650 epiphytic records of 147 taxa were gathered
(Appendix 2).
Vouchers of the encountered lichens are preserved in B, some in ATHU. A small number of
records was omitted because of incomplete identification, usually caused by poor state of the
material: 37 from Ikaria and 38 from Naxos.
RESULTS AND DISCUSSION
As Appendix 2 shows, there are 87 taxa shared
by both islands. Naxos has in addition 61 species not observed on Ikaria, and Ikaria has 60
species not observed on Naxos. This suggests
considerable floristic differences. However, the
differentiating species are usually uncommon
and may have been missed by accident on one
of the islands. When taking into account only
the 70 commonest species (3 or more records
from Ikaria or over 40 from Naxos), and leaving
aside taxa where identification problems might
have caused observation differences, 8 taxa
remain, which are evidently present on one of
the islands and absent from the other (Table 1).
It concerns common and widespread species
in the Mediterranean, which must have had
sufficient nearby diaspore sources to reach the
islands. Therefore the only explanation for their
uneven distribution seems to be insular isolation. They evidently reached one of the islands,
but failed to reach the other, although suitable
habitats are surely present.
Another way by which an effect of insular
isolation might become visible, is the frequency
of means for vegetative reproduction. For longrange dispersal they may be disadvantageous,
because the propagules are larger and probably
less easily dispersed over wider distances than
most ascospores.
An evaluation of the 208 listed taxa demonstrates that 71 or about 34% are vegetatively
reproducing by means of soredia, blastidia,
granules or isidia. A comparison with a study
of a mainland area (Giralt, 1996) shows that only
40 out of 180 species, or 22 %, produce vegetative diaspores. A possible explanation is that
the vegetatively reproducing species disperse
themselves over long distances by ascospores,
and that the vegetative diaspores serve for
short-distance dispersal, enabling the species
to build a strong population quickly once the
first individual has become established. This is
indicated by the high number of sorediate species in Table 1.
A remarkable distribution was found for two
chemical strains of Pseudevernia furfuracea on
Ikaria, var. ceracea and var. olivetorina (Fig. 2).
The species is found only above 700 m, and the
island has two areas reaching above this level,
separated by a pass of about 500 m of altitude.
The physodic acid-containing race was found
only on the eastern part, and the olivetoric acidcontaining race only in the west. Allopatric distribution for the races is unusual, as they have
been shown always to be indistinguishable by
ecology or distribution so far (e.g. Culberson et
al., 1977). It suggests that both areas have been
colonized in two separate events and that the
populations have been unable to spread across
the pass.
Table 1. List of species common on one of the islands Ikaria and Naxos, but absent from the other.
Numbers of observations and presence of vegetative reproduction (sor = sorediate) in brackets.
Only observed on Ikaria
5 out of 59 frequent species:
Bryoria capillaris (3; sor)
Hyperphyscia adglutinata (3; sor)
Hypogymnia tubulosa (5; sor)
Lecanora expallens (5; sor)
Pyrrhospora quernea (3; sor)
Only observed on Naxos
3 out of 49 frequent species:
Caloplaca aegatica (79)
Collema multipunctatum (43)
Melanelia glabra (88)
100 Folia Cryptog. Estonica
Fig. 2. Distribution of the chemotypes of Pseudevernia furfuracea on Ikaria. Dots – physodic
acid-strain; triangles – olivetoric acid-strain.
Thin line: 500 m contour.
ACKNOWLEDGEMENTS
The German Research Society (DFG) is gratefully acknowledged for financial support. Prof.
F. Daniels (Münster, Germany) supportively
enabled one of us (AS) to investigate the island
of Naxos.
REFERENCES
Culberson, W.L., Culberson, C.F. & Johnson, A. 1977.
Pseudevernia furfuracea-olivetorina relationships:
chemistry and ecology. Mycologia 69: 604614.
Degelius, G. 1986. The lichen flora of the island of Anholt, Denmark. Acta Regiae Soc. Sci. Litt. Gothob.,
Bot. 3: 1–60.
APPENDIX 1
The investigated phorophytes (27 species); IK =
Ikaria; NA = Naxos.
Acer sempervirens – NA; Alnus glutinosa – NA;
Arbutus unedo – IK; Castanea sativa – IK, NA;
Ceratonia siliqua – IK, NA; Crataegus monogyna
– IK, NA; Cupressus sempervirens – IK, NA; Erica
arborea – NA; Erica manipuliflora – IK, NA; Ficus
carica – IK, NA; Juglans regia – NA; Juniperus
oxycedrus subsp. macrocarpa – IK, NA; Malus
domestica – IK; Morus nigra – IK, NA; Nerium
oleander – NA; Olea europaea – IK, NA; Phillyrea
latifolia – IK; Pinus brutia – IK; Pinus halepensis
– NA; Pistacia lentiscus – NA; Platanus orientalis
– NA; Prunus cerasifera – IK; Pyrus spinosa – IK,
NA; Quercus coccifera – IK, NA; Quercus ilex – IK,
NA; Quercus ithaburensis subsp. macrolepis NA; Quercus pubescens – IK, NA.
Giralt, M. 1996. Líquens Epífits i Contaminació
Atmosfèrica a la Plan i les Serralades Litorals
Tarragonines. Institut d’Estudis Catalans, Secció
de Ciències Biològiques, Barcelona. 525 pp.
Hayward, B.W. & Hayward, G.C. 1986. Lichen flora
of the offshore islands of northern New Zealand.
The Offshore Islands of Northern New Zealand.
New Zealand Department of Lands and Survey
Information Series 16: 153–160.
Higgins, M.D. & Higgins, R. 1996. A geological companion to Greece and the Aegean. Cornell University
Press, Ithaca, New York. 240 pp.
Scharlau, A. 1996. Untersuchungen zur Flechtenflora
der Insel Naxos (Kykladen, Griechenland). Unpubl.
thesis, University of Münster, Germany. 184 pp.
with appendices.
Sipman, H. J. M. & Raus, Th. 1995. Lichen observations from Santorini (Greece). In: Knoph, J.-G.,
Schrüfer, K. & Sipman, H. J. M. (eds). Studies in
Lichenology with Emphasis on Chemotaxonomy,
Geography ad Phytochemistry. Festschrift Christian Leuckert. Biblioth. Lichenol. 57: 409–428.
Sipman, H. & Raus, Th. 1999. A lichenological comparison of the Paros and Santorini island groups
(Aegean, Greece), with annotated checklist. Willdenowia 29: 239–297.
Sipman, H. & Raus, Th. 2002. An inventory of the
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Warren, J. 2003. Isolation by distance in the crustose
lichens Candelariella vitellina and Placynthium nigrum colonising gravestones in northeast Scotland. Biodivers. & Conservation 12(2): 217–224.
APPENDIX 2
The epiphytic lichen taxa (208), with indication
of island (IK = Ikaria, NA = Naxos), number of
records in brackets and collecting numbers of
selected vouchers.
ACROCORDIA GEMMATA (Ach.) A. Massal. – IK (3):
48924, 49152; NA (9): 270.
AGONIMIA OCTOSPORA Coppins & P. James – IK (5):
48943.
ANAPTYCHIA CILIARIS (L.) Körb. – IK (7): 48728,
48854; NA (61): 96.
ANISOMERIDIUM BIFORME (Borrer) R.C. Harris – NA
(1): 362.
A RTHONIA CINNABARINA (DC.) Wallr. – IK (1):
49197.
ARTHONIA LIGNIARIA Hellb. – IK (1): 48942.
101
ARTHONIA PATELLULATA Nyl. – IK (1): 48939a (cf.);
NA (1).
BACIDIA AUERSWALDII (Stizenb.) Mig. – NA (4).
BACIDIA CIRCUMSPECTA (Vain.) Malme – NA (12):
258, 238a.
BACIDIA CROZALSIANA (H. Olivier) Zahlbr. – NA (1):
381.
BACIDIA ROSELLA (Pers.) De Not. – IK (2): 49226a;
NA (89): 215.
BACIDIA RUBELLA (Hoffm.) A. Massal. – IK (5):
48910; NA (26): 250.
B ACIDIA SUBINCOMPTA (Nyl.) Arnold – NA (12):
238.
BACIDINA PHACODES (Körb.) Vězda – NA (13): 251.
BACTROSPORA PATELLARIOIDES (Nyl.) Almq. – IK (2):
48921; NA (3).
BIATORELLA OCHROPHORA (Nyl.) Arnold – NA (9):
283.
BRYORIA CAPILLARIS (Ach.) Brodo & D. Hawksw. – IK
(5): 48746, 49179.
BRYORIA FUSCESCENS (Gyeln.) Brodo & D. Hawksw.
– IK (1): 49006.
BUELLIA ERUBESCENS Arnold – IK (1): 48971a.
BUELLIA GRISEOVIRENS (Sm.) Almb. – IK (3): in
48994, in 48810.
BUELLIA SCHAERERI De Not. – IK (1): 48912.
CALICIUM GLAUCELLUM Ach. – IK (1): 48974.
CALICIUM SALICINUM Pers. – NA (1): 372.
CALICIUM VIRIDE Pers. – IK (1): 48987.
CALOPLACA AEGATICA Giralt, Nimis & Poelt – NA
(79): 330.
CALOPLACA ALNETORUM Giralt, Nimis & Poelt – NA
(112): 227.
CALOPLACA CERINA (Hedw.) Th. Fr. – IK (2): 49055,
49625; NA (49): 230.
CALOPLACA CERINELLA (Nyl.) Flagey – NA voucher
nr. 301.
CALOPLACA CERINELLOIDES (Erichsen) Poelt – IK (2):
in 48468, in 49015; NA (67): 283.
CALOPLACA FERRUGINEA (Huds.) Th. Fr. – IK (2):
49015. 49587; NA (42): 235.
CALOPLACA FLAVOCITRINA (Nyl.) H. Olivier – IK (2):
49227, in 48910; NA (15): 210.
CALOPLACA FLAVORUBESCENS (Huds.) J.R. Laundon
var. QUERCINA (Flagey) Giralt, Nimis & Poelt
– IK (10): 47791, 48434.
CALOPLACA HAEMATITES (St.-Amans) Zwackh – IK
(4): 48466, 48471; NA (215): 34.
CALOPLACA HERBIDELLA (Hue) H. Magn. – IK (4):
48717, 48864; NA (3): 307.
CALOPLACA HOLOCARPA (Ach.) A.E. Wade – NA (3).
CALOPLACA HUNGARICA H. Magn. – IK (3): 48972,
48997.
CALOPLACA LOBULATA (Flörke) Hellb. – NA (2): 295.
CALOPLACA OBSCURELLA (Körb.) Th. Fr. – IK (4):
48469; NA (9): 289.
CANDELARIELLA AURELLA (Hoffm.) Zahlbr. – NA (1).
C ANDELARIELLA REFLEXA (Nyl.) Lettau – IK (1):
49127.
CANDELARIELLA VIAE-LACTAE G. Thor & V. Wirth
– NA (1).
CANDELARIELLA VITELLINA (Hoffm.) Müll. Arg. – IK (1):
49017; NA (59): 32.
CANDELARIELLA XANTHOSTIGMA (Ach.) Lettau – NA
(6): 308.
CATAPYRENIUM PSOROMOIDES (Borrer) R. Sant. – NA
(4): 354.
CATILLARIA CHALYBEIA (Borrer) A. Massal. – IK (5):
49231a, 48398; NA (44): 105.
CATILLARIA MEDITERRANEA Hafellner – NA (1): in
291.
CATILLARIA NIGROCLAVATA (Nyl.) Schuler – IK (3):
49204.
CATILLARIA PRAEDICTA Tretiach & Hafellner – IK (2):
47790, in 49313; NA (24): 314.
CATINARIA ATROPURPUREA (Schaer.) Vězda & Poelt
– IK (1): 48939; NA (6): 297.
CETRARIA ACULEATA (Schreb.) Fr. – NA (1).
CETRARIA CHLOROPHYLLA (Willd.) Vain. – IK (4):
48744, 48999.
CLADONIA CHLOROPHAEA (Sommerf.) Spreng. – IK (2):
48806, 48857a; NA (6): 394.
CLADONIA FIMBRIATA (L.) Fr. – IK (1): 49121.
CLADONIA FOLIACEA (Huds.) Willd. – NA (2).
CLADONIA GLAUCA Flörke – NA (2).
CLADONIA POCILLUM (Ach.) O.J. Richter – NA (9).
CLADONIA PSEUDOPITYREA Vain. – IK (1): 48979; NA
(4): 365.
CLADONIA PYXIDATA (L.) Hoffm. – IK (9): 48718,
48805; NA (16): 12.
C LADONIA RANGIFOR MIS Hoffm. – NA (5): 107,
107a.
COLLEMA FLACCIDUM (Ach.) Ach. – NA (10): 306.
COLLEMA FURFURACEUM (Arnold) Du Rietz – IK (2):
48927; NA (5): 392.
COLLEMA MULTIPUNCTATUM Degel. – NA (43): 202.
COLLEMA NIGRESCENS (Huds.) DC. – IK (6): 48928,
49030; NA (157): 204.
DEGELIA ATLANTICA (Degel.) P.M. Jørg. & P. James
– IK (1): 49027.
DEGELIA PLUMBEA (Lightf.) P.M. Jørg. & P. James
– IK (2): 49014; NA (2): 159.
DIMERELLA PINETI (Ach.) Vězda – IK (1): 48926.
D IPLOICIA CANESCENS (Dicks.) A. Massal. var.
CANESCENS – IK (2): in 48922, in 48900; NA
(5): 169.
102 Folia Cryptog. Estonica
DIPLOSCHISTES MUSCORUM (Scop.) R. Sant. – IK (1):
49133.
DIPLOTOMMA ALBOATRUM (Hoffm.) Flot. – IK (1): in
48467; NA (55): 256.
DIPLOTOMMA PULVERULENTUM (Anzi) D. Hawksw.
– NA (1).
DIRINA CERATONIAE (Ach.) Fr. – IK (1): in 47792.
EVERNIA PRUNASTRI (L.) Ach. – IK (16): 48729,
48743; NA (71): 249.
FUSCOPANNARIA OLIVACEA (P.M. Jørg.) P.M. Jørg. – IK
(6): 48865, 48930; NA (45): 35.
GYALECTA TRUNCIGENA (Ach.) Hepp – IK (1): 49232b;
NA (1).
GYALIDEOPSIS ANASTOMOSANS P. James & Vězda
– NA (1).
HAFELLIA DISCIFORMIS (Fr.) Marbach & H. Mayrhofer
– IK (1): 49190a.
HYPERPHYSCIA ADGLUTINATA (Flörke) H. Mayrhofer &
Poelt – IK (4): 48426, 48436.
HYPOGYMNIA FARINACEA Zopf – IK (1): 49003; NA
(1): 390.
HYPOGYMNIA PHYSODES (L.) Nyl. – IK (5): 48730,
48966; NA (3): 369.
HYPOGYMNIA TUBULOSA (Schaer.) Hav. – IK (18):
48715, 48722.
LECANIA CYRTELLA (Ach.) Th. Fr. – NA (4): 250.
LECANIA KOERBERIANA J. Lahm – NA (12): 293.
LECANIA NAEGELII (Hepp) Diederich & van den Boom
– IK (4): 49132a, 49312; NA (39): 255.
LECANORA ALLOPHANA Nyl. – NA (1): 225.
LECANORA ARGENTATA (Ach.) Malme – NA (399):
33.
LECANORA CARPINEA (L.) Vain. – NA (12): 261.
LECANORA CHLAROTERA Nyl. – IK (15): 48467, 48473
(4 not chemically tested); NA (400): 207.
L ECANORA DISPERSA (Pers.) Sommerf. – IK (5):
49311.
LECANORA EXPALLENS Ach. – IK (14): 48913, 49194
(7 uncertain).
LECANORA HAGENI (Ach.) Ach. – IK (1): in 49194;
NA (104): 268, 245.
LECANORA HORIZA (Ach.) Linds. – IK (10): 47788,
48433.
LECANORA HYBOCARPA (Tuck.) Brodo – IK (6): 48429,
48872.
LECANORA LEUCKERTIANA Zedda – IK (1): 48906.
LECANORA SAMBUCI (Pers.) Nyl. – NA (1).
LECANORA STROBILINA (Spreng.) Kieff. – NA (34):
210.
LECANORA SYMMICTA (Ach.) Ach. – IK (4): 48973,
49191; NA (12): 226.
LECIDELLA ELAEOCHROMA (Ach.) M. Choisy – IK (26):
48428, 48470; NA (592): 34-1, 34-2, 219.
LEPRARIA NIVALIS J.R. Laundon – IK (7): 48719,
48808; NA (49): 213.
LEPRARIA VOUAUXII (Hue) Kukwa – IK (1): 48925.
LEPROCAULON MICROSCOPICUM (Vill.) Gams – IK (5):
48944, 49131; NA (15): 56.
LEPTOGIUM BREBISSONII Mont. – IK (1): 48852.
LEPTOGIUM CYANESCENS (Rabenh.) Körb. – NA (2):
305.
LEPTOGIUM GELATINOSUM (With.) J.R. Laundon – NA
(5): 378.
LEPTOGIUM TENUISSIMUM (Dicks.) Körb. – NA (2):
386.
LEPTOGIUM TERETIUSCULUM (Wallr.) Arnold – IK (2):
48875, in 49146; NA (27): 269, 388.
LOBARIA AMPLISSIMA (Scop.) Forssell – IK (2): 49035,
49154.
LOBARIA PULMONARIA (L.) Hoffm. – IK (4): 48848,
49153; NA (2): 364.
L OBARIA SCR OBICULATA (Scop.) Nyl. – IK (1):
48853.
MELANELIA GLABRA (Schaer.) Essl. – NA (88): 4.
MELANELIA GLABRATULA (Lamy) Essl. – NA (24):
205.
M ELANELIA SUBAURIFERA (Nyl.) Essl. – IK (1):
48742.
MICAREA LITHINELLA (Nyl.) Nyl. – IK (1): 49159a.
MICAREA PRASINA Fr. – IK (2): 48953, in 48949.
MYCOCALICIUM SUBTILE (Pers.) Szatala – NA (3):
340.
NEPHROMA LAEVIGATUM Ach. – IK (5): 48860, 48946;
NA (3): 155.
NORMANDINA PULCHELLA (Borrer) Nyl. – IK (2): 48850,
in 49131; NA (1): 368.
OCHROLECHIA ANDROGYNA (Hoffm.) Arnold – IK (6):
48868, 48983; NA (97): 216, 277.
OCHROLECHIA BALCANICA Verseghy – NA (19): 252.
OCHROLECHIA PARELLA (L.) A. Massal. – IK (10):
48724, 48881; NA (140): 229.
OCHROLECHIA SUBVIRIDIS (Høeg) Erichsen – IK (9):
48726, 48809.
OCHROLECHIA TURNERI (Sm.) Hasselrot – IK (1):
49602a.
OPEGRAPHA ATRA Pers. – NA (9): 350, 352.
OPEGRAPHA CELTIDICOLA (Jatta) Jatta – NA (10):
346.
OPEGRAPHA CULMIGENA Lib. – NA (10).
OPEGRAPHA NIVEOATRA (Borrer) J.R. Laundon – NA
(1): 273.
OPEGRAPHA OCHROCINCTA Werner – NA (3): 347.
OPEGRAPHA VARIA Pers. – IK (1): 49229; NA (41):
231.
PACHYPHIALE CARNEOLA (Ach.) Arnold – IK (3): 48871,
48940.
103
PANNARIA RUBIGINOSA (Ach.) Bory – IK (1): 48851.
PARMELIA SAXATILIS (L.) Ach. – IK (15): 48716,
48721; NA (16): 234.
PARMELIA SULCATA Taylor – IK (3): 48803, 48948;
NA (28): 214.
P ARMELINA PASTILLIFERA (Harm.) Hale – IK (3):
48720, 48862; NA (12).
PARMELINA QUERCINA (Willd.) Hale – IK (1): 49591;
NA (41): 235.
PARMELINA TILIACEA (Hoffm.) Hale – IK (8): 49592;
NA (334): 2, in 325.
PARMOTREMA CHINENSE (Osbeck) Hale & Ahti – IK
(13): 48733, 48739; NA (16): 235.
PELTIGERA COLLINA (Ach.) Schrad. – IK (1): 49124.
PELTIGERA HYMENINA (Ach.) Delise – NA: 362.
PELTIGERA NECKERI Müll. Arg. – NA (2): 362.
PERTUSARIA ALBESCENS (Huds.) M. Choisy & Werner
– IK (16): 48727, 48735; NA (175): 10, 162,
300.
PERTUSARIA AMARA (Ach.) Nyl. – IK (1): 48810.
PERTUSARIA COCCODES (Ach.) Nyl. – IK (6): 48879,
48932; NA (6): 260.
PERTUSARIA DALMATICA Erichsen – NA (16): 240.
PERTUSARIA GRAECA Erichsen – IK (12): 48723,
48807; NA (23): 124.
PERTUSARIA HEMISPHAERICA (Flörke) Erichsen – IK
(1): 49434; NA voucher nr. 216.
PERTUSARIA HETEROCHROA (Müll. Arg.) Erichsen – IK
(3): in 48736, 48917.
PERTUSARIA HYMENEA (Ach.) Schaer. – IK (9): 48423,
48878; NA (238).
PERTUSARIA LEIOPLACA DC. – NA (32).
P ERTUSARIA OPHTHALMIZA (Nyl.) Nyl. – IK (1):
49436.
PERTUSARIA PERTUSA (Weigel) Tuck. – IK (18): 48725,
48736; NA (187): 93.
PERTUSARIA RHODIENSIS Erichsen – NA (22): 303.
PERTUSARIA WERNERIANA Boqueras – IK (2): 48427,
49201.
PHAEOPHYSCIA ORBICULARIS (Neck.) Moberg – IK (5):
49316, 49601; NA (83): 271.
PHLYCTIS AGELAEA (Ach.) Flot. – IK (3): 48957,
49202, in 49197; NA (28): 496.
PHLYCTIS ARGENA (Spreng.) Flot. – IK (15): 48424,
48880; NA (96): 92.
PHYSCIA ADSCENDENS (Fr.) H. Olivier – IK (6): 48465,
48919; NA (309): 97.
PHYSCIA AIPOLEA (Humb.) Fürnr. – IK (1): 49599;
NA (150): 215.
PHYSCIA BIZIANA (A. Massal.) Zahlbr. var. BIZIANA – IK
(8): 48464, 48908; NA (260): 7.
PHYSCIA BIZIANA (A. Massal.) Zahlbr. var. LEPTOPHYLLA
Vězda – IK (1): 48903; NA (10): 37.
PHYSCIA LEPTALEA (Ach.) DC. – IK (6): 49040, 49119;
NA (334): 122, 278, 133.
PHYSCIA TENELLA (Scop.) DC. – IK (11): 48425,
48902; NA (101): 158.
P HYSCONIA ENTEROXANTHA (Nyl.) Poelt – IK (2):
49237a, in 48425; NA (32): 36.
PHYSCONIA GRISEA (Lam.) Poelt – IK (1): 49237; NA
(18): 332.
PHYSCONIA PERISIDIOSA (Erichsen) Moberg – IK (4):
48421, 49012; NA (31): 212.
PHYSCONIA SERVITII (Nádv.) Poelt – IK (9): 48422,
48437; NA voucher nr. in 215, in 224.
PHYSCONIA SUBPULVERULENTA (Szatala) Poelt – IK (6):
48861, 48963; NA voucher nr. 224.
PHYSCONIA VENUSTA (Ach.) Poelt – IK (1): 49011; NA
(83): 3, in 224.
PLACYNTHIELLA ICMALEA (Ach.) Coppins & P. James
– IK (2): 48975.
PLACYNTHIELLA ULIGINOSA (Schrad.) Coppins & P.
James – IK (1): 48991.
PLATISMATIA GLAUCA (L.) W.L. Culb. & C.F. Culb.
– IK (1) obs. Pirintsos; NA (2): 229.
PLEUROSTICTA ACETABULUM (Neck.) Elix & Lumbsch
– IK (2): 48968, in 48950; NA (33): 228.
PORINA AENEA (Wallr.) Zahlbr. – NA (3): 263.
PSEUDEVERNIA FURFURACEA (L.) Zopf var. CERATEA
(Ach.) D. Hawksw. – IK (3): 48745.
PSEUDEVERNIA FURFURACEA (L.) Zopf var. FURFURACEA
– IK (5): 49000; NA (2): 231.
PYRENULA CHLOROSPILA Arnold – IK (1): 49232a;
NA (1): 364.
PYRRHOSPORA LUSITANICA (Räsänen) Hafellner – IK
(1): in 49188.
PYRRHOSPORA QUERNEA (Dicks.) Körb. – IK (8):
48984, 49165.
RAMALINA CANARIENSIS J. Steiner – IK (2): 48431, in
48912; NA (76): 101, 73.
RAMALINA FARINACEA (L.) Ach. – IK (3): 48430,
48858; NA (51): 121.
RAMALINA FASTIGIATA (Pers.) Ach. – IK (18): 48432,
48438; NA (173): 100.
RAMALINA FRAXINEA (L.) Ach. var. CALICARIFORMIS (Nyl.)
Hue – IK (1): 48945; NA (37): 99.
RAMALINA FRAXINEA (L.) Ach. var. FRAXINEA – NA
(31): 98.
RINODINA BOLEANA Giralt & H. Mayrhofer – NA (4):
341.
RINODINA CAPENSIS Hampe – IK (7): 49308, 49584;
NA (13): 289.
RINODINA COLOBINA (Ach.) Th. Fr. – NA (2): 393.
RINODINA NIMISII Giralt & H. Mayrhofer – NA (1):
333.
104 Folia Cryptog. Estonica
RINODINA OLEAE Bagl. – IK (1): in 48716; NA (126):
348.
RINODINA PYRINA (Ach.) Arnold – NA (8): 275.
RINODINA SEPTENTRIONALIS Malme – NA (46): 281.
SCHISMATOMMA DECOLORANS (Sm.) Clauzade & Vězda
– IK (2): 48920.
SCOLICIOSPORUM CHLOROCOCCUM (Stenh.) Vězda – IK
(2): 48996, 49203.
SCOLICIOSPORUM SAROTHAMNI (Vain.) Vězda – IK (2):
49167, in 49190; NA (29): in 333.
STAUROLEMMA OMPHALARIOIDES (Anzi) P.M. Jørg. &
Henssen – IK (2): 49031, 49236; NA (2):
329.
STRIGULA MEDITERRANEA Etayo – NA (4): 321.
T ELOSCHISTES FLAVICANS (L.) Norman – IK (1):
48855.
TEPHROMELA ATRA (Huds.) Hafellner – IK (9): 48955,
49143; NA (149): 8.
THELENELLA MODESTA (Nyl.) Nyl. – NA (4): 385.
THELOPSIS ISIACA Stizenb. – IK (1): 47792; NA (1):
343.
TORNABEA SCUTELLIFERA (With.) J.R. Laundon – NA
(8): 228.
T RAPELIOPSIS GRANULOSA (Hoffm.) Lumbsch – IK (1):
in 49188.
USNEA ARTICULATA (L.) Hoffm. – IK (3): 48747,
48845; NA (3): 230-1.
USNEA CORNUTA Körb. – IK (5): 49177, 49186,
49430; NA (5): 230-2.
USNEA ESPERANTINA P. Clerc – IK (2): 49432a, in
48730.
USNEA FLAMMEA Stirt. – IK (1): 48748; NA (2).
USNEA GLABRESCENS (Vain.) Vain. – IK (2): 48749,
49185.
USNEA SCABRATA Nyl. – IK (1): 48849.
USNEA WIRTHII P. Clerc – IK (2): 49428, in 49186.
WAYNEA STOECHADIANA (Abbassi Maaf & Cl. Roux)
Cl. Roux & P. Clerc – IK (1): 48905.
XANTHORIA PARIETINA (L.) Th. Fr. – IK (13): 48435,
49232; NA (488): 1.
Folia Cryptog. Estonica, Fasc. 41: 105–108 (2005)
Lichens from Golestan National Park (Iran)
Mohammad Sohrabi
Department of Biology, Faculty of Science, Gorgan University of Agricultural Science and Natural Resources,
Gorgan, Iran
E-mail: sohrabi_lichen@yahoo.com
Abstract: 41 lichen species belonging to 28 genera are reported from Golestan National Park (Iran). 3 genera and 11 species
are new to the lichen flora of Iran.
Kokkuvõte: Golestani Rahvuspargi (Iraan) samblikud.
Esitatakse Golestani Rahvuspargi (Iraan) samblike nimekiri, mis sisaldab 41 liiki 28 perekonnast, koos leiuandmetega. 3
perekonda ja 11 liiki on uued Iraani samblikeflooras.
INTRODUCTION
This paper is a contribution to a better knowledge of the lichen flora of Iran. It reports several species which were found in the Golestan
National Park (province of Golestan). Golestan
National Park is one of the UNESCO’s designated reserves and the first Iranian national
park. The area, with 91.895 hectares, is located in the northeast of Iran between three
provinces; Golestan, Khorassan, and Semnan.
The most outstanding feature of the area lies in
its extremely different climatic conditions, which
vary from temperate subhumid in the west to
cold-arid and cold-semiarid in the south and
the east of the park, with an annual precipitation ranging from ca 150 to ca 1000 mm. The
complex of geomorphologic, geologic, hydrologic,
and climatic conditions provide a wide range of
biotopes leading to high biodiversity.
Although, there are some reports on the
lichen flora of northern Iran (Buhse, 1860;
Müller, 1892; Steiner, 1896, 1910, 1916; Szatala, 1940, 1957; Oxner, 1946; Weber, 1965;
Riedl, 1979), the only records from Golestan are
those of a few species cited by Hertel (2001) and
Seaward et al (2004). The present paper is the
first devoted to a better knowledge of the poorly
known lichen flora of the Park.
MATERIAL AND METHODS
The samples were collected between 20–21 June
2003 in three different zones in the southeast
of Golestan National Park: Almeh, Golzar, and
Sharleg (Fig. 1).
The samples were numbered, and stored
in paper envelopes. A stereomicroscope, light
microscope, and usual colour tests were used in
the identification. The main reference texts were:
Clauzade & Roux (1985), Magnusson (1929,
1940), Nimis (1993), Orange et al (2001), Timdal (1991), Wirth (1987). The samples are kept
in the private herbarium of Mohammad Sohrabi
(hb. M. Sohrabi), all duplicates kept in the TSB
herbarium. New records are marked with *.
LIST OF TAXA
ACAROSPORA CERVINA A. Massal. – Golestan National
Park, Almeh zone, 850 m; saxicolous. 20
June 2003, Sohrabi 2507.
*ACAROSPORA LAQUEATA Stizenb. – Golestan National
Park, Sharleg zone, 750 m; saxicolous. 21
June 2003, Sohrabi 2592.
A NAPTYCHIA CILIARIS (L.) Körb. ex A. Massal.
– Golestan National Park, Golzar camping,
600 m; corticolous on Quercus sp. 21 June
2003, Sohrabi 2479.
ANAPTYCHIA CRINALIS (Schrad.) Vězda – Golestan
National Park, Golzar camping, 600 m;
corticolous on Quercus sp. 21 June 2003,
Sohrabi 2481.
ANAPTYCHIA ULOTHRICHOIDES (Vain.) Vain. – Golestan
National Park, Sharleg zone, 750 m;
corticolous on Quercus sp. 21 June 2003,
Sohrabi 2596.
*ASPICILIA EMILIAE Aggr. – Golestan National Park,
Golzar camping, 600 m; saxicolous. 21 June
2003, Sohrabi 2497.
106 Folia Cryptog. Estonica
Fig. 1. Location of Golestan National Park.
*BIATORELLA OCHROPHORA (Nyl.) Arnold – Golestan
National Park, Golzar camping, 600 m;
corticolous on Pyrus sp. 21 June 2003,
Sohrabi 2484.
C ALOPLACA CERINA (Ehrh. ex Hedw.) Th. Fr.
– Golestan National Park, Golzar camping,
600 m; lignicolous, on Quercus sp. 21 June
2003, Sohrabi 2478.
CALOPLACA PERSICA (J. Steiner) M. Steiner & Poelt
– Golestan National Park, Almeh zone, 850
m; lignicolous, on Pyrus sp. 20 June 2003,
Sohrabi 2500.
C ANDELARIELLA VITELLINA (Hoffm.) Müll. Arg.
– Golestan National Park, Almeh zone, 850
m.; saxicolous, lignicolous on Juniperus. 20
June 2003, Sohrabi 2507.
CLADONIA POCILLUM (Ach.) Grognot – Golestan
National Park, Sharleg zone, 750 m;
terricolous. 21 June 2003, Sohrabi 2584.
C OLLEMA AURIFOR ME (With.) Coppins & J.R.
Laundon – Golestan National Park, Almeh
zone, 850 m; saxicolous, or muscicolous. 20
June 2003, Sohrabi 2525.
C OLLEMA FUSCOVIRENS (With.) J.R. Laundon
– Golestan National Park, Almeh zone, 850
m; saxicolous or on calcareous rock under
trees. 20 June 2003, Sohrabi 2525
C OLLEMA FUSCOVIRENS (With.) J.R. Laundon –
Golestan National Park, Sharleg zone, 750
m; saxicolous or on calcareous rock under
trees. 21 June 2003, Sohrabi 2578.
DIPLOSCHISTES DIACAPSIS (Ach.) Lumbsch – Golestan
National Park, Sharleg zone, 750 m;
terricolous. 21 June 2003, Sohrabi 2582.
DIPLOTOMMA VENUSTUM Körb. – Golestan National
Park, Sharleg zone, 750 m; saxicolous. 21
June 2003, Sohrabi 2583.
EVERNIA PRUNASTRI (L.) Ach. – Golestan National
Park, Golzar camping, 600 m; corticolous,
on Quercus sp. 21 June 2003, Sohrabi
2495.
LECANORA FLOTOWIANA Körb. – Golestan National
Park, Almeh zone, 850 m; lignicolous, on
Quercus sp. 20 June 2003, Sohrabi 2499.
LECANORA HAGENII (Ach.) Ach. – Golestan National
Park, Golzar camping, 600 m; corticolous,
on Quercus sp. 21 June 2003, Sohrabi
2486.
LECANORA MURALIS (Schreb.) Rabenh. – Golestan
National Park, Sharleg zone, 750 m;
saxicolous, on calcareous rock. 21 June
2003, Sohrabi 2589.
M ELANELIA GLABRA (Schaer.) Nyl. – Golestan
National Park, Golzar camping, 600 m.
corticolous, on Quercus sp. 21 June 2003,
Sohrabi 2483.
MELANELIA SUBARGENTIFERA (Nyl.) Essl. – Golestan
National Park, Golzar camping, 600 m;
corticolous, on Quercus sp. 21 June 2003,
Sohrabi 2487.
P AR MELINA TILIACEA (Hoffm.) Hale – Golestan
National Park, Golzar camping, 600 m;
107
corticolous, on Quercus sp. 21 June 2003,
Sohrabi 2493.
*PECCANIA TERRICOLA H. Magn. – Golestan National
Park, Almeh zone, 850 m; terricolous, on
soil and in crevices of rocks. 20 June 2003,
Sohrabi 2506, 2529.
PELTIGERA RUFESCENS (Weiss) Humb. – Golestan
National Park, Sharleg zone, 750 m;
terricolous. 21 June 2003, Sohrabi 2586.
*PERTUSARIA ALBESCENS (Huds.) M.Choisy & Werner
– Golestan National Park, Golzar camping,
600 m; corticolous, on Quercus sp. 21 June
2003, Sohrabi 2482, 2491.
*P HLYCTIS ARGENA (Spreng.) Flot. – Golestan
National Park, Golzar camping, 600 m.
corticolous, on Acer sp. 21 June 2003,
Sohrabi 2492.
PHYSCIA ADSCENDENS (Fr.) H. Olivier – Golestan
National Park, Almeh zone, 850 m;
corticolous, on Quercus sp. 20 June 2003,
Sohrabi 2500.
PHYSCIA TENELLA (Scop.) DC. – Golestan National
Park, Golzar camping, 600 m; corticolous,
on Quercus sp. 21 June 2003, Sohrabi
2485.
PHYSCIA TRIBACIA (Ach.) Nyl. – Golestan National
Park, Almeh zone, 850 m; corticolous, on
Quercus sp. 20 June 2003, Sohrabi 2504.
PHYSCONIA GRISEA (Lam.) Poelt – Golestan National
Park, Golzar camping, 600 m; corticolous,
on Quercus sp. 21 June 2003, Sohrabi
2480.
PHYSCONIA PERISIDIOSA (Erichsen) Moberg – Golestan
National Park, Golzar camping, 600 m;
corticolous, on Quercus sp. 21 June 2003,
Sohrabi 2481.
*PLEUROSTICTA ACETABULUM (Neck.) Elix & Lumbsch
– Golestan National Park, Golzar camping,
600 m; corticolous, on Quercus sp. 21 June
2003, Sohrabi 2484.
*RAMALINA FRAXINEA (L.) Ach. – Golestan National
Park, Golzar camping, 600 m; corticolous,
on Quercus sp. 21 June 2003, Sohrabi 2478,
2490.
R AMALINA POLLINARIA (Westr.) Ach. – Golestan
National Park, Golzar camping, 600 m;
corticolous, on Quercus sp. 21 June 2003,
Sohrabi 2494.
RHIZOCARPON GEOGRAPHICUM (L.) DC. – Golestan
National Park, Sharleg zone, 750 m;
saxicolous. 21 June 2003, Sohrabi 2579.
RHIZOPLACA CHRYSOLEUCA (Sm.) Zopf – Golestan
National Park, Almeh zone, 850 m;
saxicolous. 20 June 2003, Sohrabi 2521.
T ONINIA CINEREOVIRENS (Schaerer) A. Massal.
– Golestan National Park, Sharleg zone,
750 m. Saxicolous, in fissures of rocks. 21
June 2003, Sohrabi 2580.
*T ONINIA TAURICA (Szatala) Oxner – Golestan
National Park, Sharleg zone, 750 m;
saxicolous, in fissures of rocks. 21 June
2003, Sohrabi 2594.
*VERRUCARIA MARMOREA (Scop.) Arnold – Golestan
National Park, Sharleg zone, 750 m;
saxicolous. 21 June 2003, Sohrabi 2595.
*XANTHORIA FULVA (Hoffm.) Poelt & Petutschnig
– Golestan National Park, Golzar camping,
600 m; corticolous, on Acer sp. 21 June
2003, Sohrabi 2486, 2488.
ACKNOWLEDGEMENTS
I am most grateful to Prof. P. L. Nimis, (Trieste,
Italy) for confirmation of my identifications. I
especially want to thank Prof. Mansouri and
Mis. Shimi (Center of International Research
and Collaboration – ISMO) and Dr. Ali Tavili
(Teheran University) and all people who helped
me in various ways during my research in different parts of Iran. This study was financially
supported by ISMO.
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Nimis, P. L. 1993. The Lichens of Italy. Museo Regionale
di Scienze Naturali, Torino. 897 pp.
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Microchemical methods for the identification of
lichens. British Lichen Society, London. 101 pp.
Oxner, A. N. 1946. Lichens of northern Iran collected
by A. B. Shelkovnikov (in Ukrainian). Botanical
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Halbwüsten. Plant Systematics and Evolution
131: 217–233.
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Maassoumi, A. A., Hadjmoniry, M. & Sohrabi,
M. 2004. A lichen checklist of Iran. Willdenowia
(accepted)
Steiner, J. 1896. Beitrag zur Flechtenflora Südpersiens.
Sitzungsberichte der kaiserlichen Akademie
der Wissenschaften in Wien, mathematischnaturwissenschaftliche Classe 105(1): 436–446.
Steiner, J. 1910. Lichenes Persici coll. a cl. Consule
Th. Strauss. Annales Mycologici 8: 212–245.
Steiner, J. 1916. Aufzählung der von J. Bornmüller
im Oriente gesammelten Flechten. Annalen
des Naturhistorischen Hofmuseums in Wien 30:
24–39.
Szatala, O. 1940 (1939). Lichenes. In: K. H. Rechinger,
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einer botanischen Reise nach dem Iran, 1937.
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des Irans. Annales Historico-Naturales Musei
Nationalis Hungarici, ser. nov. 8: 101–154.
Timdal, E. 1991. A monograph of the genus Toninia.
Opera Botanica 110: 1–137.
Weber, W. A. 1964. Iranian plants collected by Per
Wendelbo in 1959. VIII. Lichenes. Årbok for
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Eugen Ulmer, GmbH & Co., Stuttgart. 528 pp.
Folia Cryptog. Estonica, Fasc. 41: 109–114 (2005)
Chemical constituents from lichens for pharmaceutical and
industrial uses
Kunio Takahashi1, Kaoru Kinoshita1, Yoshikazu Yamamoto2, Kiyotaka Koyama1 &
Isao Yoshimura3
Department of Pharmacognosy and Phytochemistry, Meiji Pharmaceutical University, Noshio 2-522-1, Kiyose-shi,
Tokyo 204-8588, Japan
E-mail: diamonds@my-pharm.ac.jp
2
Department of Biological Production Science, Faculty of Bioresource Sciences, Akita Prefectural University, 241-7,
Kaidobata-nishi, Shimoshinjo-nakano, Akita 010-0195, Japan
3
Hattori Botanical Laboratory, Kochi Branch, Edagawa 2576-27, Ino-cho, Agawagun, Kochi, Japan
1
Abstract: Some pharmaceutical and industrial uses of lichen metabolites from both cultured isolates and the thalli of lichens
are reported. 5-propylresorcinol and bis(2,4-dihydroxy-5-propylphenyl)methane are the anti-tyrosinase active constituents
of mixture of Protousnea spp. Since tyrosinase is an enzyme producing melanin pigments, the anti-tyrosinase activity might
be effective in whitening the facial skin. Both the activity and cytotoxicity of some synthetic resorcinol derivatives were
tested. Furthermore, the inhibitory activities of confluentic acid and its synthetic analogues on liver monoamine oxidases
were screened. New compounds, arthoniafurone A and B, and cinnabarinol, were obtained from the cultured isolate of
Arthonia cinnabarina. These novel metabolites showed cytotoxicity against U937 human leukemia cells. Other novel substances
panaefluoroline A, B, and C were obtained from the medium where cultured isolate of Amygdalaria panaeola was grown.
These substances are significant regarding their structures and also their fluorescent characters, which were stable both in
water and in organic solvents over one year. The metabolites of cultured mycobionts may be one of new natural resources
for developing of fluorescent agents.
Kokkuvõte: Samblikuained farmaatsia ja tööstuse tarbeks.
Tutvustatakse nii samblikutallustest eraldatud samblikuainete kui söötmel kasvatatud mükobiontidest saadud ainete kasutusvõimalusi farmaatsias ja tööstuses. 5-propüülresorsinool ja bis(2,4-dihüdroksü-5-propüülfenüül)metaan on türosinaasivastase
toimega ained perekonna Protousnea liikides. Kuna türosinaas on ensüüm, mis katalüüsib melaniinsete pigmentide teket, siis
türosinaasivastase toimega ained võiksid olla kasulikud näonaha pleegitamisel. Kontrolliti veel mõnede sünteetiliste resorsinooli derivaatide bioaktiivsust ja tsütotoksilisust. Uuriti konfluenthappe ja selle sünteetiliste analoogide inhibeerivaid omadusi
maksa monoamiini oksidaaside suhtes. Arthonia cinnabarina kultuurist eraldati uued ühendid arthoniafuroon A ja B ning tsinnabarinool. Need ained ilmutasid tsütotoksilisust inimese leukeemia U937 rakkude suhtes. Söötmest, millel oli kasvatatud
Amygdalaria panaeola kultuuri, eraldati veel kolm uut ühendit – panaefluoroliin A, B, ja C. Need ained on tähelepanuväärsed nii
oma struktuuri ja kui ka fluorestseeruvate omaduste poolest, mis säilusid nii vees kui orgaanilistes lahustites üle aasta. Kultuuris
kasvatatud mükobiontidest eraldatavad ained võivad olla üheks uute looduslike fluorestseeruvate ühendite allikaks.
INTRODUCTION
Lichens, symbiotic associations of algal and
fungal partners, produce many characteristic phenolic compounds such as depsides,
depsidones, benzoquinones, naphthquinones,
anthraquinones, xanthones, biphenyls and
dibenzofuranes (Huneck & Yoshimura, 1996),
which are considered to be biosynthesized by
the mycobionts, fungal partners (Culberson,
1969). We are interested in primary and secondary metabolites of lichens and for this reason we performed both structural elucidation
of novel metabolites and screened their possible
applications in the pharmaceutical field (Takahashi et al., 1974, 1979, 1981; Watanabe et
al., 1986; Kinoshita et al., 1993, 1994a, 1994b,
2002, 2003; Matubara et al., 1994, 1997, 1998,
1999; Yamamoto et al., 2002b). As a result of
our search of secondary metabolites in cultured
lichen mycobionts, we surprisingly isolated
some novel metabolites, which had never been
found in intact lichen thalli from nature. New
secondary compounds were recently found in
cultured isolates of lichens under stress conditions, for example under osmotic stress – if
10% sucrose was supplemented to the nutrient
medium (Kon et al., 1997; Miyagawa et al., 1993,
1994, 1997; Tanahashi et al., 1997; Yamamoto
et al., 1996).
110 Folia Cryptog. Estonica
A major objective of the present study is to
determine the chemical structures of hitherto unknown secondary lichen compounds
isolated from aposymbiotically grown lichen
fungi (Yamamoto et al., 1996, 2002a, 2002b;
Kinoshita et al., 2003). A further aim of these
investigations is to compare past and new reports about pharmaceutical and industrial uses
of lichen metabolites from both cultures and
the thalli.
MATERIAL AND METHODS
Plant materials
Protousnea spp. mixture [P. dusenii (DR.) Krog,
P. malacea (Stirt.) Krog and P. magellanica
(Mont.) Krog] was collected in Tierra del Fuego
in 1990; the specimens are kept in Department
of Biological Production Science, Faculty of Bioresource Sciences, Akita Prefectural University,
Japan. Arthonia cinnabarina (DC.) Wallr. was collected in 1995 in Japan; the specimen is in the
harbarium of the Natural History Museum and
Institute, Chiba, Japan. Amygdalaria panaeola
(Ach.) Hertel & Brodo was collected in Finland
in 1990; the specimen is in Hattori Botanical
Laboratory, Kochi Branch, Kochi, Japan.
All lichens were identified by Dr I. Yoshimura, Hattori Botanical Laboratory, Kochi Branch,
Kochi, Japan.
Chemical material
Isolated confluentic acid used by the authors
of this paper is from Asahina Collection which
is stored in the Department of Pharmacognosy
and Phytochemistry, Meiji Pharmaceutical University, Japan.
Methods
The inhibition assay on MAO-A and MAO-B was
performed according to the method reported by
Kagaya et al. 1996.
RESULTS
In search of new biologically active compounds,
we found some secondary metabolites of lichens
that could be potential sources for pharmaceutical and industrial uses.
PHARMACEUTICAL USES
Anti-tyrosinase active substances
Natives in the island of Tierra del Fuego have
used the extract of lichen mixtures of Protousnea
spp. as a skin conditioner to whiten the facial
skin. We reported earlier that the anti-tyrosinase
active constituents of those are 5-propylresorcinol (Fig. 1, substance 1) and bis(2,4- dihydroxy5-propylphenyl)methane (Fig. 1, substance 2)
(Kinoshita et al., 1994a). Since tyrosinase is an
enzyme producing melanin pigments, the antityrosinase activity might be effective in whitening the facial skin.
In an additional study to find out more
details about the structure-activity interrelationship, some synthetic resorcinol derivatives
were prepared. The elongation of the alkyl chain
of resorcinol from methyl to nonyl showed an
increase of the activity. Moreover, 4-alkylresorcinols, non-natural and synthetic analogues,
and also arbutin (Fig. 1, substance 3), which
have been used as a component of cosmetics to
cause whitening effects of the skin, were prepared and tested for their biological activities.
Our experiments demonstrated that the activity
of 4-alkylresorcinols, for example 4-methylresorcinol (substance 4), was stronger than those
of 5-alkylresorcinols (5-methylresorcinol, substance 5) and also of a natural diphenylmethane, bis(2,4-dihydroxy-5-methylphenyl)methane
(substance 6) (Matsubara et al., 1997) (Figs
1–2).
In a new test series we screened both the activity and cytotoxicity of those compounds. As a
conclusion, 4-alkylresorcinols showed inhibition
of melanin production and cytotoxicity effects
were visible in cultured B-16 mouse melanoma
cells at concentrations from 10 mM to 1.2 mM.
Among the tested compounds, substance 4
showed a comparatively stronger activity with a
slightly weaker cytotoxicity at 80 μM (Matsubara
et al., 1998). It was noticed that the difference
in the position and the length of the alkyl group
in resorcinol derivatives affected both the antityrosinase activity and the cytotoxicity. Such
findings may be useful for understanding the
chemical background of recorcinol derived compounds and their applications in skin care.
Monoamine oxidase (MAO) inhibitors
MAO is located in the outer mitochondrial membrane and exists in two forms, type A and B
111
(MAO-A and MAO-B). Endo et al. (1994) reported
that confluentic acid, isolated from a Brazilian
plant, showed a selective inhibitory effect on
MAO-B. Confluentic acid (Fig. 3, substance 7)
was first isolated form the lichen, Lecidea confluens (Weber) Ach. and is a depside comprising
two alkylresorcinolic acid units. We examined
the inhibitory activities of some lichen compounds and their synthetic analogues on liver
MAO to determine the role of confluentic acid
(Kinoshita et al., 2002). We have since found
that the synthetic 6-alkyl-α-resorcinolic acids
had no activity, but 5-heptyl-3-methoxyphenol
showed activity (56%) at a concentration of 2.5
X 10-5 g/ml. These findings indicated that the
presence of a carboxylic acid group was not essential for the biological activity.
Mouse liver MAO is a mixture of MAO-A and
B. We have now tested confluentic acid and the
synthetic compounds 1-(2,4-dihydroxyphenyl)1-heptanone (substance 8) and 1-(4-hydroxy-2methoxyphenyl)-1-heptanone (substance 9) for
their activity on MAO-A and MAO-B, respectively
(Fig. 3). It has previously been reported that
MAO-B inhibitors can be used effectively in
combination with L-DOPA (L-dihydroxyphenylalanine) for the treatment of Parkinson’s disease.
The inhibitory activity of compound 8 on MAOB was found to be much stronger than that of
compound 9. Thus methylation of ortho-phenolic
group of 8 decreased its activity. Among various
synthetic analogues the heptanone 8 showed
the strongest and most selective activity against
MAO-A and B, and it had almost the same potency as natural confluentic acid (Table 1).
Cytotoxicity against cancer cells
Some aposymbiotically grown lichen mycobionts (cultured without the algal partner) can
synthesize novel and extraordinary chemical
constituents. We successfully cultured the
isolate of Arthonia cinnabarina derived from
its spores. New compounds, arthoniafurone A
(substance 10) and B (substance 11), and cinnabarinol (substance 12), were obtained from the
culture (Fig. 4) (Yamamoto et al., 2002b). These
novel metabolites showed cytotoxicity against
U937 human leukemia cells. The concentration of 50% effective dose of substance 10 was
<30μM, that of substance 11 was <30μM, and
of 12 was 25μM.
INDUSTRIAL USES
Cultured isolate of Amygdalaria panaeola was
obtained from the thallus fragment. The medium, on which the mycobiont grew, showed
yellowish green fluorescence under a fluorescent lamp. Novel fluorescent substances, panaefluoroline A (13), B (14), and C (15), were
first isolated to determine their structures (Fig.
5) (Kinoshita et al., 2003).
As mentioned already, these fluorescent
pigments could not be detected in the natural
lichen thallus by HPLC analysis. Moreover,
polyketides like depsides or depsidones were
not isolated and could not be detected in the
culture medium by TLC. However, fluorescent
compounds were isolated from the medium of
cultured isolate. These results suggest that the
biosynthetic pathway of the cultured mycobiont, Amygdalaria panaeola, may be changed by
stress under variable culture conditions.
Anyway, the obtained novel compounds are
significant regarding their structures and also
their fluorescent characters, which were stable
both in water and in organic solvents over one
year. The metabolites of cultured mycobionts
may be one of new natural resources for developing fluorescent agents.
AKNOWLEDGEMENTS
We wish to thank Emeritus Professor of Tokyo
University, Shoji Shibata, for his encouragement
and suggestion.
Table 1 Inhibition of monoamine oxidases (MAO-A and MAO-B) by confluentic acid and its analogues.
Confluentic acid (7)
MAO-A(IC50, µM)
MAO-B(IC50, µM)
Ratio (A/B)
25
1.4
18
1-(2,4-dihydroxy-phenyl)1-heptanone (8)
70
2.4
29
1-(4-hydroxy-2-methoxyphenyl)-1-heptanone (9)
>100
16
>6.3
112 Folia Cryptog. Estonica
Fig. 1. Structures of 5-propylresorcinol (1), bis(2,4-dihydroxy-6-propylphenyl)-methane (2), arbutin (3),
4-methylresorcinol (4), 5-methylresorcinol (5) and bis(2,4-dihydroxy-6-methylphenyl)methane (6).
Fig. 2. Anti-tyrosinase activities of resorcinol derivatives and arbutin.
113
Fig. 3. Structures of inhibitors of monoamine oxidases – confluentic acid (7), and its analogues,
1-(2,4-dihydroxyphenyl)-1-heptanone (8) and 1-(4-hydroxy-2-methoxyphenyl)-1-heptanone (9).
Fig. 4. Structures of new compounds arthoniafurone A (10), B (11) and cinnabarinal (12).
Fig. 5. Structures of other novel substances panaefluoroline A (13), B (14) and C (15).
114 Folia Cryptog. Estonica
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Folia Cryptog. Estonica, Fasc. 41: 115–122 (2005)
A preliminary phylogeographic study of Flavopunctelia and Punctelia
inferred from rDNA ITS-sequences
Arne Thell1, B. Herber2, A. Aptroot3, M. T. Adler4, T. Feuerer2 & E. I. Kärnefelt1
Lund University, the Biological Museums, Botanical Museum, Östra Vallgatan 18, 223 61 Lund, Sweden.
E-mail: Arne.Thell@sysbot.lu.se
2
Biozentrum Klein Flotbek und Botanischer Garten, Universität Hamburg, Ohnhorststrasse 18, D-22609 Hamburg,
Germany
3
Centraalbureau voor Schimmelcultures, P. O. Box 85167, NL-3508 AD Utrecht, Netherlands
4
Departemento de Biodiversidad y Biologica Experimental, Facultad de Ciencias Exactas y Naturales, Universidad de
Buenos Aires, Piso 4, Pabellon II, Ciudad Universitaria, 1428 Ciudad Autonoma de Buenos Aires, Argentina
1
Abstract: A preliminary phylogeny of the genera Flavopunctelia and Punctelia is presented. Genus and species delimitations
have been investigated using ITS rDNA-sequencing of populations from different continents. Current genus delimitations of
Flavopunctelia, Punctelia and Parmelia are confirmed and the species status of recently resurrected Punctelia ulophylla is confirmed.
The status of three cryptic species, Flavopunctelia soredica, Punctelia perreticulata and P. stictica is discussed. Flavopunctelia borrerioides
and Punctelia perreticulata are reported from China for the first time.
Kokkuvõte: Esialgne ülevaade perekondade Flavopunctelia ja Punctelia fülogeograafiast rDNA ITSsekventside põhjal.
Esitatakse perekondade Flavopunctelia ja Punctelia esialgne fülogeneesi rekonstruktsioon. Perekondade ja liikide eraldamist on
uuritud erinevatelt kontinentidelt pärinevate populatsioonide ITS rDNA sekventside alusel. Senine perekondade Flavopunctelia,
Punctelia ja Parmelia piiritlemine on leidnud kinnitust, samuti liigi Punctelia ulophylla staatus. Arutletakse kolme krüptilise
liigi, Flavopunctelia soredica, Punctelia perreticulata ja P. stictica staatuse üle. Teatatakse liikide Flavopunctelia borrerioides ja Punctelia
perreticulata esmasleidudest Hiinas.
INTRODUCTION
The genus Punctelia Krog was segregated from
Parmelia Ach. on differences in pseudocyphellae ontogeny, secondary chemistry and phytogeography (Krog, 1982). The genus, originally
including 22 species, was subdivided into two
distinct subgenera: Punctelia subgenus Punctelia, characterized by unciform spermatia and
atranorin as a a major cortical substance, and
Punctelia subgenus Flavopunctelia Krog characterized by bifusiform spermatia and usnic acid
as a major cortical substance. Spermatial shape
has been considered to be of great importance
in genus delimitations (Kärnefelt, 1998). Based
on spermatial and additional chemical characters, Flavopunctelia (Krog) Hale was recognized
as a separate genus composed of four species
(Hale, 1984); two additional species have been
discovered in Flavopunctelia (Elix & Adler, 1987;
Kurokawa, 1999) as compared with 30 species
which constitute Punctelia today (Crespo et
al., 2004; Egan, 2003; Elix & Johnston, 1988;
Galloway & Elix, 1994; Kurokawa, 1999; Sérusiaux, 1983, 1984; Wilhelm & Ladd, 1992). Both
Flavopunctelia and Punctelia have a temperate
to subtropical distribution and reach their
highest diversity in South- and North-America
and in Africa (Krog, 1982). DNA-investigations
support that Flavopunctelia and Punctelia are
sister groups and that Parmelia Ach. may be
the sister group of the two genera (Blanco et
al., 2004, Thell et al., 2004). Populations of
Flavopunctelia and Punctelia species, of which
some are represented by collections from different continents, are analysed here, together
with some Parmelia species, using nuclear ITS
rDNA-sequences. Genus and species delimitations are studied and discussed.
MATERIALS AND METHODS
The material was collected by the authors and
collegues during recent travels, resulting in 20
new sequences from the ITS1-5.8S-ITS2 rDNA
region that were submitted to the NCBI GenBank (http://www.ncbi.nlm.nih.gov, Table 1).
Eleven sequences were downloaded from the
116 Folia Cryptog. Estonica
Table 1. Specimens used in the study, extraction numbers (LD), sample-IDs and GenBank accession numbers
Species
Flavoparmelia caperata
Flavopunctelia borrerioides
Flavopunctelia flaventior
Flavopunctelia flaventior
Flavopunctelia flaventior
Flavopunctelia soredica
Parmelia ernstiae
Parmelia ernstiae
Parmelia saxatilis
Parmelia saxatilis
Parmelia submontana
Parmotrema crinitum
Punctelia borreri
Punctelia borreri
Punctelia borreri
Punctelia borreri
Punctelia borreri
Punctelia borreri
Punctelia perreticulata
Punctelia perreticulata
Punctelia perreticulata
Punctelia stictica
Punctelia stictica
Punctelia stictica
Punctelia subpraesignis
Punctelia subrudecta
Punctelia subrudecta
Punctelia subrudecta
Punctelia ulophylla
Punctelia ulophylla
Punctelia ulophylla
Extr.
555
1521
1285
1517
1520
1518
858
965
518
534
1273
945
959
960
1338
1339
1506
1286
1331
1505
1020
1340
1609
1310
944
958
1509
956
957
1507
Sample-ID
Estonia, Tartumaa, Ahunapalu,Thell 9906 (TUR)
China, Yunnan Prov., 29 Oct 2002, van Herk (ABL)
Germany, Bavaria, Dachau, Feuerer s. n. (HBG)
China, Yunnan Prov., Aptroot 56024 (ABL)
China. Yunnan Prov., Aptroot 560101 (ABL)
U. S. A., New York, Aptroot 50612 (ABL)
Germany, Schleswig-Holstein, Feuerer & Thell (HBG)
Sweden, Scania, Eslöv, Thell 0101 (HBG)
Chile, Magallanes, Feuerer 29542 (HBG)
Finland, Regio aboënsis, Ruissalo, Thell 9926 (TUR)
Spain, Hoya Redonda (MAF 3729)
Yemen, Socotra, Schultz 14297c (HBG)
Italy, Trentino-Alto Adige, Feuerer & Thell s. n. (HBG)
Italy, Abruzzo, Tretiach 34124 (HBG)
Italy, Friuli-Venezia-Giulia, Gambera 34126 (HBG)
Kenya, Kakamega Forest Nat. R., Killmann (priv. herb.)
Kenya, Kakamega Forest Nat. R., Killmann (priv. herb.)
China, Yunnan Prov., Aptroot 56028 (ABL)
U. S. A., Missouri, Osage Co., Ladd 23798 (HBG)
China, Yunnan Prov., Yunlong, Aptroot 56005 (ABL)
China, Yunnan Prov., Aptroot 56094 (ABL)
Venezuela, La Culata, 31 Oct 1995, Feuerer s. n. (HBG)
Kenya, Kakamega Forest Nat. R., Killmann (priv. herb.)
Peru, Pisac, 20 Sept 2003, Thell & Feuerer s. n. (HBG)
Argentina, B. Aires, Adler & Protomastro s. n (BAFC)
Germany, Schleswig-Holstein, Feuerer & Thell (HBG)
Italy, Venezia-Giulia, Gambera 34128 (dupl. HBG)
Germany, Eifel, Aptroot 55416 (ABL)
The Netherlands, Gelderland, Aptroot 44450 (ABL)
The Netherlands, Prov. Utrecht, Sipman 43579 (HBG)
Belgium, Liege, Aptroot 57873 (ABL).
*Sequences downloaded from the GenBank
same GenBank, of which 10 have appeared in
earlier publications (Adler et al., 2004; Molina et
al., 2004; Thell et al., 2002, 2004). The laboratory work was performed at the Department of
GenB. acc.
AF451750*
AY773129
AF251420*
AY773126
AY773127
AY773128
AF410834*
AF247007*
AF410672*
AF410835*
AY037000*
AY251442*
AY773113
AF451769*
AY773114
AY773110
AY773111
AY773115
AY773123
AY773124
AY773122
AY773125
AY773112
AY773119
AY267010*
AY773116
AY773117
AY773118
AY773120
AY251726*
AY773121
General Botany and Botanical Garden, University of Hamburg.
Minute fragments of the fresh collections
were ground with sterile plastic pestels. Total
117
DNA was extracted using the DNEasy Plant
Mini Kit from Qiagen as described in Thell et
al. (2004). ITS standard primers, ITS 4 and ITS
5, were used (White et al., 1990)
Ready To Go PCR beads (in 0.2 ml tubes) from
Pharmacia Biotech Inc. were dissolved in 11.8 µl
distilled water, 0.35 µl of a 16µM concentration
of each of the primers ITS5 and ITS4 (White et
al., 1990). The ITS fragments were amplified with
a Perkin-Elmer Gene Amp PCR System 9700
thermal cycler. 12.5 µl of the concentrated DNA
extractions were added to the solution, resulting in final reaction volumes of ca. 25 µl. The
PCR started with 2 minutes at 95°C, followed
by a 30–35 cycle schedule using a denaturation
temperature of 95°C for 1 min., an annealing
temperature of 60°C for 1 min., and an extension temperature of 72°C for 1 min.
The PCR products were purified with QIAquick
PCR purification kit, and diluted in 30 µl of the
enclosed elution buffer. A 25 cycle sequencing
PCR, with a denaturation temperature of 96°
C for 10 seconds, an annealing temperature of
50° C for 5 seconds, and an extension time of
60° C for 4 minutes, was performed to amplify
the DNA-fragments prior to the sequencing procedure. 12 µl deionized water including 30–90
ng of the purified PCR-product and 3.2 pmol
of the primers ITS1LM (Myllys, 1999) and ITS4
were added to 8 µl BigDye Terminator Cycle
Sequencing Ready Reaction Kit with AmpliTaq
Polymerase FS from Perkin Elmer according to
the accompanying protocol. The sequences were
produced using an automatic sequencer, ABI
Prism 377 from Perkin-Elmer.
The phylogenetic analyses of the manually
aligned ITS sequences were done with PAUP version 4.0b (Swofford, 1998). Trees were searched
by using the heuristic option, with TBR branch
swapping, 1000 replicates of random addition
sequence order, and branches collapsed if the
maximum length is zero. Gaps in the alligned
sequences were treated as missing characters.
Bootstrap analyses with 1000 replicates were
done, using the same settings as in the heuristic
search. Bootstrap support values of 60 or above
are marked in the cladogram above the branches
(Fig. 1). Large surveys of Parmeliaceae phylogeny were consulted when selecting the outgroup
(Crespo et al., 2001; Thell et al., 2004)
RESULTS AND DISCUSSION
Results from the phylogeny analysis
The aligned matrix was composed of 520 nucleotide long sequences, including the gaps. Of the
163 variable characters, 114 were parsimony
informative. The phylogeny was based on parsimony analysis using PAUP 4.0b. The analysis
resulted in 12 shortest trees of length 328 (RI =
0,834; CI = 0,668). Bootstrap consensus (Fig. I)
was identical to strict consensus of the 12 most
parsimonious trees, except for some branching
orders of the Punctelia subclades (Fig. 1. I–V).
Genus and species delimitations
Present genus delimitations are supported in the
analysis, where Parmelia and Flavopunctelia are
strongly supported, with bootstrap values of 94
and 100 respectively. Punctelia has a more moderate support, 75, of the three ingroup genera,
and the clade is divided into five subclades (Fig.
1, I–V). Four of these subclades, however, constitute single species, and two of these species constitute monophyletic clades, P. subrudecta (Nyl.)
Krog and P. ulophylla (Ach.) van Herk & Aptroot
(Fig. 1, subclades II–III), both having a bootstrap
support value at least 99. Flavopunctelia soredica (Nyl.) Hale, Punctelia perrecticulata (Räs.) G.
Wilh. & Ladd and P. stictica (Duby) Krog appear
as cryptic species (Fig. 1, subclades IV–V). The
bootstrap support for the largest subclade (Fig.
1, I) is rather strong, 75. At the adjacent node of
the tree, however, an even stronger monophyletic
clade is identified, composed of P. borreri (Sm.)
Krog, the P. perreticulata-samples collected in
China, P. subpraesignis (Nyl.) Krog. African P.
borreri and the Chinese P. perreticulata are not
supported as separate species by ITS-sequences
(Wilhelm & Ladd, 1987).
The number of cryptic species have increased
rapidly in recent years becuse they are revealed
by DNA-techniques. How to taxonomically treat
these morphologically and chemically more or
less identical but genetically different species
is currently under constant review. Cryptic
species are most frequently discovered by ITSsequences, but the results should preferrably
be confirmed by a second or third gene, such
as mitochondrial SSU (Crespo et al., 2001) or
GAPDH (Myllys et al., 2002). On the contrary,
118 Folia Cryptog. Estonica
Flavoparmelia caperata 555 Estonia
Parmotrema crinitum 1273 Yemen
Punctelia borreri 1338 Kenya
55
84
69
80
Punctelia borreri 945 Italy
Punctelia borreri 959 Italy
Punctelia borreri 960 Italy
Punctelia perreticulata 1505 China
86
Punctelia perreticulata 1331 China
91
Punctelia borreri 1339 Kenya
95
Punctelia borreri 1506 China
75
I
Punctelia subpraesignis 1310 Argentina
Punctelia stictica 1340 Kenya
Punctelia stictica 1609 Peru
72
100
75
II
Punctelia subrudecta 944 Germany
Punctelia subrudecta 958 Italy
Punctelia subrudecta 1509 Germany
Punctelia ulophylla 956 The Netherlands
99
Punctelia ulophylla 957 The Netherlands
III
Punctelia ulophylla 1507 Belgium
Punctelia perreticulata 1286 U. S. A.
IV
Punctelia stictica 1020 Venezuela
V
Flavopunctelia flaventior 1285 Germany
74
98
94
Flavopunctelia flaventior 1517 China
Flavopunctelia soredica 1520 China
Flavopunctelia borrerioides 1521 China
Flavopunctelia soredica 1518 U. S. A.
97
Parmelia ernstiae 965 Sweden
100
Parmelia saxatilis 518 Chile
69
100
Parmelia ernstiae 858 Germany
Parmelia saxatilis 534 Finland
Parmelia submontana Spain
Fig. 1. Phylogeny of Flavopunctelia, Parmelia and Punctelia based on ITS rDNA-sequences. Bootstrap consensus, identical to strict consensus from 12 most parsimonious trees. Bootstrap support
values of 60 or above are indicated above the branches, and the five subclades (I–V) of Punctelia
below the branches.
ITS-sequences do not always separate formerly
accepted morphological species (Grube & Kroken, 2000).
Punctelia ulophylla – a successfully resurrected taxon
Punctelia ulophylla was resurrected only recently
(van Herk & Aptroot, 2000), based on detailed
morphological observations and co-occurrence
in Western Europe with P. subrudecta. Although
the species was formerly described as a variety,
it was rarely mentioned, and was not thought to
merit much attention between the many other
varieties and forms described in Parmelia s.
lat. However, it has now been widely reported,
mainly from Western Europe, but also from
Northern Europe (Gauslaa, 2000) and Central
Europe (Truong & Clerc, 2003). Crespo et al.
(2004) confirmed its status as a separate species
and even found it to be only distantly related to
119
P. subrudecta, which is confirmed here by our
results. It is surprising how such a common
and well-recognizable macrolichen has remained
unnoticed for so many years. Even more surprising is the current reluctence of some authors to
accept this fact, even including the unexplained
citation of P. ulophylla as a synonym of P. subrudecta in Santesson et al. (2004).
Flavopunctelia borrerioides and Punctelia
perreticulata – new to China
Among the material cited are two species which
are new to China, namely Flavopunctelia borrerioides Kurok. and Punctelia perreticulata.
Flavopunctelia borrerioides was described
by Kurokawa (1999) from Peru and Mexico and
more recently has been reported from India by
Divakar et al. (2003). It differs from the common
F. flaventior (Stirton) Hale, with which it occurs,
by the conspicuous rounded laminal pseudocyphellae developing into soredia, giving it the
aspect of Punctelia borreri (hence the name). Our
specimen from China and an additional one from
South Africa (leg. C.M. van Herk, in ABL) suggest
that this is a widespread subtropical species.
Punctelia perreticulata was recently reported
(Aptroot, 2003) to be the most common and
widespread sorediate Punctelia in North- and
Central America (Egan & Aptroot, 2004). It is
therefore not surprising that it is also present
in Asia, as our specimen from China shows.
Notes on the cryptic species
Punctelia perreticulata is a species mainly distinguished by atranorin in the upper cortex
and sorediate pseudocyphellae. The lower side
is ivory to tan towards the centre and lecanoric
acid in the medulla is the major secondary metabolite. It has been treated as a synonym of P.
subrudecta by Krog (1982) and Nimis (1993),
a view not supported in the cladogram, which
contradicts a conspecific habit of the two species
(Fig. 1). Alternatively, Whilhelm & Ladd (1987),
who studied corticolous populations from the
interior highlands of the USA, considered P. perreticulata to be a distinct species, distinguished
by a strongly fovelate-reticulate upper surface
compared with the occasionally somewhat
ridged upper surface of P. subrudecta. Adler
& Ahti (1996) also considered P. perreticulata
as a distinct species, mainly differring from P.
subrudecta in shape and length of spermatia:
European studied specimens of P. perreticulata
were all saxicolous, had a strongly foveolate upper surface, and 7 µm long, filiform spermatia.
Specimens form Argentina were characterized
by longer, 9–11 µm, spermatia, whereas those
from the USA were mostly corticolous and supplied with an even upper surface (Adler & Ahti,
1996; Aptroot, 2003; Egan & Aptroot, 2004).
The Chinese specimens of P. perreticulata are
corticolous and have 9–12 µm long spermatia
similar to the North-American material. Adler
& Ahti (1996) concluded that P. perreticulata is
a species with an unusually wide infraspecific
geographic variation. On the contrary, Longán
et al. (2000) prefer a narrow species concept
for samples with long spermatia. The present
analysis, comparing samples from China and
the USA only, supports the view that several
species may be accumulated in P. perreticulata,
compared with the circumscription by Adler &
Ahti (1996).
Punctelia stictica (Duby) Krog is characterized by
a light to dark brown upper surface with secondarily sorediate pseudocyphellae, with granular
to isidioid soredia, the underside brown to black
towards the centre, with gyrophoric acid as the
major medullary metabolite and long filiform
spermatia (Adler, 1996). It is also a widely distributed, cosmopolitan species, reported from
North- and South-America, Africa and Europe,
growing mostly on rocks in very different climates and altitudes, but found also on soil
and trunks (Adler, 1996). Both P. perrecticulata
and P. stictica are very widely distributed and
have wide ecological amplitudes. Apart from P.
perreticulata, the cryptic habit of P. stictica is,
however, not supported by any hitherto known
morphological, chemical or anatomical traits,
which, on the other hand, is the definition
of being cryptic. Furthermore, the P. stictica
samples do not group according to geographic
origin (Fig. 1)
Finally, the Flavopunctelia soredica sample from
the USA (New York) constitutes a sister group to
the single F. borrerioides sample, the two F. flaventior samples and the F. soredica sample from
China. Interestingly, in this preliminary study
based on a single DNA-fragment, populations of
the three possibly cryptic species correlate with
DNA according to their geographical origins.
120 Folia Cryptog. Estonica
ACKOWLEDGMENTS
Prof. Mark Seaward is thanked for reviewing the
manuscript. M. T. Adler is a Research Member
of The National Research Council of Argentina
(CONICET)and their support is highly appreciated. Jack Elix is thanked for determining the
African specimens.
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White T. J., Burns T., Lee S. & Taylor J. 1990. Amplification and direct sequencing of fungal ribosomal DNA genes for phylogenetics. In: Innis M.,
Gelfand, J., Sninsky, J. & White, T. (eds) PCR
protocols: a guide to methods and applications, pp.
315–322. Academic Press, Orlando, Florida.
Wilhelm, G & Ladd, D. 1987. Punctelia perreticulata, a
distinct lichen species. Mycotaxon 28: 249–250.
Wilhelm, G & Ladd, D. 1992. A new species of the lichen genus Punctelia from the midwestern United
States. Mycotaxon 44: 495–504.
122 Folia Cryptog. Estonica
Folia Cryptog. Estonica, Fasc. 41: 123–134 (2005)
Secondary chemistry of lichen-forming ascomycetes: culture
experiments with isolates of Xanthoparmelia species from Australia
Bettina Zocher & Elfie Stocker-Wörgötter
Institute of Organismic Biology, University of Salzburg, Hellbrunner Strasse 34, 5020 Salzburg, Austria
E-mails: bettina.zocher@sbg.ac.at
elfriede.stocker@sbg.ac.at
Abstract: Sterile cultured isolates of different Xanthoparmelia species from Australia were analyzed and their chemistry
compared with their voucher specimens. Isolates produced the same major and satellite substances as typically formed in
lichen thalli, such as usnic, salazinic and consalazinic acids in Xanthoparmelia cheelii, X. tasmanica and X. antleriformis. Norlobaridone, loxodin, usnic acid and, surprisingly, divaricatic acid, which was never before reported for that lichen were found
in voucher specimens and cultures of X. flavescentireagens. Many of the cultured fungi produced additional compounds and
traces of various other secondary metabolites, which are either closely related to the major substances and/or parts of a
chemosyndrome and precursors. X. flavescentireagens produced different secondary compounds, depending on the composition of the culturing media.
Kokkuvõte: Samblikke moodustavate kottseente sekundaarsete ainete keemia: Austraaliast pärit koldsamblike
(Xanthoparmelia) isolaatide kultuuris kasvatamise katsed.
Analüüsiti Austraaliast pärit Xanthoparmelia liikide steriilseid kultuuri viidud isolaate ning võrreldi nende samblikuaineid samade
liikide lihheniseerunud eksemplaride ainetega. Isolaadid tootsid samu põhi- ja kõrvalaineid, mis tavaliselt moodustuvad nende
samblike talluses, nt usniin-, salatsiin- ja konsalatsiinhappeid liikides Xanthoparmelia cheelii, X. tasmanica ja X. antleriformis. Sambliku
X. flavescentireagens tõendeksemplarides ja kultiveeritud isolaatides leiti norlobaridooni, loksodiini, usniinhapet ja üllatuslikult ka
divarikaathapet, mida sellest samblikust varem määratud pole. Paljudest kultuuri viidud seentest leiti lisaaineid ning erinevate
metaboliitide jälgi, mis on tihedalt seotud põhiainetega ja/või kemosündroomi osade ja eelkäijatega. X. flavescentireagens tootis
kultuuri koostisest sõltuvalt erinevaid sekundaaraineid.
INTRODUCTION
Lichen forming ascomycetes produce a wide
range of secondary metabolites such as depsides, depsidones and dibenzofurans (Culberson, 1969), some of which are potentially useful,
biologically active compounds. Antiviral, antibiotic, antitumor, photoprotective, but also allergenic and enzyme inhibitory effects have been
recognized (e.g. Rancan et al., 2002; Yamamoto
et al., 1993). Lichen substances can also be
produced by cultured mycobionts, depending
on the culture conditions, composition of the
nutrient media and stress parameters such as
osmotic and temperature stress or exposure to
high light intensities (Tanahashi et al., 1997;
Stocker-Wörgötter & Elix, 2002; Yamamoto et
al., 1987). The biosynthesis of secondary metabolites in the cultured mycobionts may differ
from the production in intact thalli. Possible
causes for these differences can be osmotic
conditions, nutrient supply, culture age, culture conditions or the presence of a photobiont
(Molina et al., 2003).
Lichen thalli are known to be frequently
inhabited by parasitic, endo- and epiphytic
fungi. Therefore, chemical or genetic analyses
are often required for identification of sterile
cultured isolates (Yamamoto, 1990; StockerWörgötter, 2001).
The genus Xanthoparmelia (Vain.) Hale, a segregate of Parmelia Ach., occurs mainly in southern
Africa, Australia and New Zealand. In comparison, only few species are known from the northern hemisphere. Xanthoparmelia is one of the
most dominant groups of saxicolous lichens in
very warm, semi-arid regions, where its representatives frequently grow in open habitats.
Xanthoparmelia thalli are foliose to subcrustose, loosely to very tightly adnate, and
their upper surface is pale yellow-green to
grey-green. Common secondary compounds in
the genus Xanthoparmelia are usnic, salazinic,
consalazinic, norstictic and protocetraric acid,
loxodin, norlobaridone and chemically closely
related minor substances (Elix, 1994).
124 Folia Cryptog. Estonica
In general, representatives of Parmeliaceae
reveal highly variable and complex chemical
profiles (including many biologically active
compounds) and thus are exciting subjects for
culturing and chemical investigations.
The objective of this investigation was to
screen culture conditions and their influence on
the formation of secondary compounds in sterile
cultured isolates of Australian Xanthoparmelia
species, and to compare them with lichen substances contained in the original thalli.
MATERIAL AND METHODS
Lichen specimens
Xanthoparmelia antleriformis (Elix) Elix & Johnst.
was collected S of Morton National Park near
Nerriga in New South Wales, Australia, 35°05’S,
150°09’E, 700 m; X. cheelii (Gyeln.) Hale and
X. flavescentireagens (Gyeln.) D.J. Galloway:
Brindabella Range, Mt Majura, Australian Capital Territory (ACT), 35°14’S, 149°11’E (Fig. 1); X.
tasmanica (Hook.f. & Taylor) Hale: Brindabella
Range, summit of Mt Aggie, ACT, 43 km WSW
of Canberra, 35°28’S, 148°48’E, 1490 m.
List of culture media
Different culturing media were used. LBM+B
[Lilly-Barnett-Medium with bark extract (LillyBarnett, 1951)] contains glucose as a carbohydrate source, agar, asparagine, thiamine, biotin
and different mineral nutrients (Mg, K, Fe, Zn,
Mn) and additionally bark extract prepared by
the method of Esser (1976).
MIX [MIX-Medium (Stocker -Wörgötter,
2002)] contains glucose, malt, yeast extract,
agar, peptone and NaCl.
MS [Murashige-Skoog-Medium modified
(Stocker-Wörgötter, 2001)] consists of sucrose,
mannitol, malt, caseine, agar and the Murashige
mineral salts.
MY [Malt-Yeast-Medium (Yamamoto, 1990)]
is composed of malt, yeast extract and agar.
PDA [Potato-Dextrose-Agar (Sigma P-2182)]
is an extract of potatoes, glucose and agar.
S2% [Sabouraud-2%-Glucose-Agar (Fluka
84086)] and S4% [Sabouraud-4%-Glucose-Agar
modified (Stocker-Wörgötter, 2001)] contain
polypeptone, glucose and agar.
Mycobiont isolations
The Yamamoto-method (modified after
Yamamoto, 1990) was used for the isolation of
the symbionts from selected Xanthoparmelia
species. Small thallus fragments were washed
in bidistilled water containing a drop of Tween
80, then homogenized in sterile water with a
mortar and pestle. The suspension was filtered
through two sieves (500 and 150µm mesh size).
Fragments (160–200 µm size) of the washed and
homogenized lichen material were transferred
with bamboo sticks into tubes containing MSmedium. The tubes were kept in darkness (covered by aluminium foil) for about 3 months until
small colonies had formed. The fungal colonies
were homogenized for subculturing on the different media as described above.
Isolates were kept in a culture chamber with
a 14 h/20°C:10 h/10°C day-night cycle and a
light intensity of 50-100 µE m-2s-1.
Chemical analyses
For the chemical analyses, sterile cultures of
the mycobionts (c. 1.0–1.5 cm) were cut out of
the agar plates and freeze-dried at –42°C for
at least 12 h. Dried cultures and small pieces
of the original lichen thalli were extracted in
methanol for 4 h; the extracts were transferred
to HPLC vials. The secondary compounds of the
thalli and mycobiont cultures were observed
by HPLC using a Merck-Hitachi system with
two pumps, a DAD (photodiode array detector; 190–800 nm wavelength) and a computer
system. Two solvent systems were used: (A) 1%
aqueous orthophosphoric acid and methanol in
the ratio 7:3, and (B) methanol. The run started
with 100% A and was raised to 58% B within
15 min, then to 100% B after a further 15 min,
followed by isocratic elution in 100% B for a
further 10 min. The spectra were identified by
means of a spectrum library (comparisons with
reference substances), and chemical data of
Huneck and Yoshimura (1996). TLC was performed using the standardized method with 3
solvent systems (Culberson & Ammann, 1979;
Culberson & Kristinsson, 1969).
RESULTS
Cultures
Isolates of the different Xanthoparmelia spp.,
when grown on the same media (Fig. 2–4), differ in color, growth rate and growth form. They
all produce brown pigments. Protruding hyphae
cause a velvet appearance. The Xanthoparmelia
cheelii isolate (Fig. 3) forms more hyphae that dip
into the medium than the other species.
125
Depending on the composition of the nutrient media, isolates of the same species vary in
their growth rates, colouration and also in their
developmental patterns. Figures 5–10 show isolates of X. flavescentireagens growing on various
test media. Pigments leak into the medium and
darkly stain it, particularly on MIX and S2%.
In general, the Xanthoparmelia isolates show
a high increase of biomass on all tested media,
except on S4% where slow growth was recorded
(Fig. 10).
Chemistry of the voucher specimens and
their cultured mycobionts
The chemical profiles of the selected lichens and
their response to different media were compared.
Isolates of Xanthoparmelia tasmanica, X. cheelii
and X. antleriformis showed best growth on MS,
whereas the isolate of X. flavescentireagens revealed no preference for a particular nutrient
medium. Qualitatively, the chemical profile
(types of secondary metabolites) did not change,
but quantitative differences (major, minor, traces) were found on different media. Table 1 gives
an overview of the chemical analyses.
XANTHOPARMELIA TASMANICA
The HPLC and TLC data revealed that the thallus contained salazinic acid (major), usnic acid
(minor) and a small amount of consalazinic acid
(Fig. 11).
The mycobiont grown on MS produces
salazinic acid (major), usnic acid (minor) and has
additional minor compound conorlobaridone.
Tab. 1. Results of the chemical analyses of Xanthoparmelia spp. and its cultured isolates
X. antleriformis
X. cheelii
X. tasmanica
Species
Source of substances
Quantity of substances
minor
low quantities/traces
usnic acid
consalazinic acid
usnic acid, conor- consalazinic acid,
lobaridone
unknown
usnic acid
consalazinic acid, norstictid acid, protocetraric
acid
conorlobaridone
unknowns
thallus (Fig. 11)
isolate (MS) (Fig. 12)
major
salazinic acid
salazinic acid
thallus (Fig. 13)
salazinic acid
isolate (MS) (Fig. 14)
thallus (Fig. 15)
salazinic acid,
usnic acid
salazinic acid
isolate (MS) (Fig. 16)
salazinic acid
usnic acid
thallus (Fig. 17)
usnic acid, norlobaridone
loxodin, divaricatic
acid
usnic acid
X. flavescentireagens
isolate (MS)
isolate (MIX)
isolate (PDA)
isolate (MY)
isolate (LBM+B) (Fig. 18)
isolate (S2%)
isolate (S4%)
consalazinic acid, norstictic acid, protocetraric acid
consalazinic acid, conorlobaridone,
unknowns
usnic acid, nor-lobaridone, loxodin, divaricatic
acid
norlobaridone
2´O-methylsubdivaricatic acid
norlobaridone
norlobaridone
norlobaridone
usnic acid
loxodin, divaricatic acid
conorlobaridone
norlobaridone,
loxodin, divaricatic acid,
usnic acid
conorlobaridone
loxodin, usnic acid conorlobaridone
loxodin, usnic acid conorlobaridone
126 Folia Cryptog. Estonica
baridone (minor), usnic acid (minor), a small
amount of loxodin and divaricatic acid, and a
trace of conorlobaridone. On MY, the mycobiont
did not produce any major lichen substances,
except a trace of conorlobaridone. On S2% and
Fig. 1. Thallus of Xanthoparmelia
flavescentireagens.
There are also traces of consalazinic acid and
another unknown (UK) substance (Fig. 12).
XANTHOPARMELIA CHEELII
The voucher specimen contains salazinic acid
(major), usnic acid (minor) and traces of consalazinic acid, norstictic acid and protocetraric
acid (Fig. 13).
The cultured mycobiont produces the two
major substances (salazinic and usnic acid), and
conorlobaridone as another minor compound
(Fig. 14). There are also several peaks of unidentified trace substances.
XANTHOPARMELIA ANTLERIFORMIS
Salazinic acid (major) and usnic acid (minor)
and a small quantity of consalazinic acid can
be found both in the thallus and the cultured
mycobiont. Additionally, the lichen thallus forms
traces of norstictic acid and protocetraric acid;
in contrast the mycobiont in culture produces
traces of conorlobaridone and different additional unknown compounds (Fig. 15, 16).
XANTHOPARMELIA FLAVESCENTIREAGENS
The thallus contains usnic acid and norlobaridone as major metabolites, and loxodin and
divaricatic acid as minor compounds (Fig. 17).
The mycobiont grown on MS produces the same
substances, but only a trace quantity of them.
However, when grown on MIX, the mycobiont
only forms norlobaridone. When cultured on
PDA the mycobiont produces this substance
as a major compound, usnic acid as a minor
compound, and there are traces of loxodin and
divaricatic acid. The extract of the mycobiont
on LBM+H (Fig. 18) shows a depside, probably
2`-O-methylsubdivaricatic acid (major), norlo-
Fig. 2. Isolate of X. tasmanica cultured on MS
medium.
Fig. 3. Isolate of X. cheelii cultured on MS
medium.
Fig. 4. Isolate of X. antleriformis cultured on
MS medium.
127
S4% the mycobiont produced norlobaridone
(major), loxodin (minor), divaricatic acid (minor), usnic acid (minor), and also a trace of
conorlobaridone.
Figs 19–22 show UV-spectra of prominent
substances, as found in Xanthoparmelia spp.
(salazinic acid, usnic acid, norlobaridone and
divaricatic acid).
Extracts of the culture media show that the
mycobionts had released only a small quantity
Figures 5–10: Sterile cultured isolate of X. flavescentireagens grown on different media:
Fig. 5. MIX medium;
Fig. 6. PDA medium;
Fig. 7. LBM+B medium;
Fig. 8. MY medium;
Fig. 9. S2% medium;
Fig. 10. S4% medium.
128 Folia Cryptog. Estonica
Figs 11–13. HPLC chromatograms of secondary compounds produced by thalli (T) and sterile
cultured isolates (I) of Xanthoparmelia spp.:
Fig. 11. X. tasmanica (T); Fig. 12. X. tasmanica (I) grown on MS medium; Fig. 13. X. cheelii (T).
129
Figs 14–18. HPLC chromatograms of secondary compounds produced by thalli (T) and sterile
cultured isolates (I) of Xanthoparmelia spp.: Fig. 14. X. cheelii (I) grown on MS medium; Fig. 15.
X. antleriformis (T); Fig. 16. X. antleriformis (I) grown on MS medium.
130 Folia Cryptog. Estonica
Figs 17–18. HPLC chromatograms of secondary compounds produced by thalli (T) and sterile
cultured isolates (I) of Xanthoparmelia spp.:
Fig. 17. X. flavescentireagens (T); Fig. 18. X. flavescentireagens (I) grown on LBM+B medium.
of substances into the medium, probably chemically related metabolites of norlobaridone and
traces of several unidentifiable compounds. Depending on the type of medium and/or the species (e.g. X. flavescentireagens, X. antleriformis),
pigments were released into the medium. Obviously, these pigments were not solubilized in
methanol and therefore could not be detected
in HPLC analyses.
DISCUSSION
Most of the cultured mycobionts produced the
same major substances as the voucher speci-
mens, but cultured isolates often showed considerable chemosyndromic variation in the composition of satellite compounds and precursors
(e.g. conorlobaridone, 2`-O-methylsubdivaricatic
acid). Due to the very low contents, these traces were often difficult to identify, because the
spectra showed a high background noise. One
explanation for the differences in the production
of secondary compounds in voucher specimens
and cultures could be that the aposymbiotically
grown mycobionts represent juvenile and deviate
developmental stages that have been grown under “artificial” lab conditions considerably different to the ones parent thalli experience in their
natural environment (Molina et al., 2003).
131
The cultured isolates of the different Xanthoparmelia species exhibited similar growth forms,
probably because they are representatives of the
same genus and closely related to each other.
Mostly, the cultured isolates showed increased
growth rates on nutrient rich media. The only
exception is S4%, obviously a less than optimal
substrate, since all Xanthoparmelia isolates grew
much more slowly on this medium than on S2%.
In general, the production of lichen substances
by cultured isolates also depended on the composition of the nutrient media.
However, high growth rates on one medium
do not necessarily mean that the culture also
produces high quantities of secondary compounds, as can be seen in the cultured isolates
of X. flavescentireagens grown on MY or MIX.
A negative correlation was found between fast
growth on nutrient-rich media and production
of secondary compounds. Our results indicate
that mycobionts exhibiting low growth rates
favour the polymalonate pathway. Further
essential parameters include environmental
factors such as temperature shifts, increased
light intensities and various other stress factors
(Tanahashi et al., 1997; Stocker-Wörgötter &
Elix, 2002; Yamamoto et al., 1987). The impact
of these factors could be tested in further investigations. Our study showed that the cultured
fungal isolates responded to physiological stress,
such as reduced and altered nutrient content
of the artificial media. Seasonal changes and
alternating day-night cycles in the natural environment can only be partly simulated in the
Figs 19–20: UV-spectra: Fig. 19. salazinic acid; Fig. 20. usnic acid.
132 Folia Cryptog. Estonica
Figs 21–22: UV-spectra: Fig. 21. norlobaridone; Fig. 22. divaricatic acid.
culture chamber (i.e. by exposing the cultures
to a 14 h/20°C:10 h/10°C day-night cycle).
The detection of divaricatic acid in the original
thallus and the culture of X. flavescentireagens
was very surprising, as this compound has not
been previously described from this species
(Elix, 1994; Elix, pers. comm.).
ACKNOWLEDGEMENTS
We are very grateful to the Austrian Science
Foundation (grant 15378 B06) for support. Prof.
J. A. Elix is very much thanked for his help with
the collection and determination of the investigated lichens during several stays of EST-W
in Australia and his advice in confirming the
identification of the lichen substances. Alfredo
Passo (Centro Regional Universitario Bariloche,
Universidad Nacional del Comahue, Argentina),
Armin Hager and Georg Brunauer (Institute of
Plant Physiology, University of Salzburg, Austria)
are also thanked for valuable discussions.
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Biochemistry 41: 175–180.
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against UVB irradiation by natural filters extracted from lichens. Journal of Photochemistry
and Photobiology B: Biology 68: 133–139.
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the lichen symbiosis: DNA-analyses, differentiation and secondary chemistry of selected mycobionts, artificial resynthesis of two- and tripartite
symbioses. Symbiosis 30: 207–227.
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A. K. (eds), pp. 47–60. Protocols in Lichenology:
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in Biomonitoring. Springer-Verlag, Berlin, Heidelberg, New York.
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chemistry of cultured mycobionts: formation of a
complete chemosyndrome by the lichen fungus of
Lobaria spathulata. Lichenologist 34: 351–359.
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N. & Hamada, N. 1997. Four phenolics from the
cultured lichen mycobiont of Graphis scripta var.
pulverulenta. Chemical and Pharmaceutical Bulletin 45: 1183–1185.
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134 Folia Cryptog. Estonica
Folia Cryptog. Estonica, Fasc. 41: 135–136 (2005)
NEW ESTONIAN RECORDS
Lichens and lichenicolous fungi
Ave Suija1, Maarja Nõmm1 & Steffen
Boch2
Institute of Botany and Ecology, University of Tartu, 38
Lai St., 51005 Tartu
2
Universität Lüneburg, Institut für Ökologie und
Umweltchemie, Scharnhorststrasse 1, D-21335 Lüneburg,
Germany
1
Eleven new species of lichens and lichenicolous
fungi and one new variety are reported here. Abbreviations of distribution regions and frequency
classes follow Randlane & Saag (1999). All cited
material is deposited in the lichenological herbarium of University of Tartu (TU).
AGONIMIA GLOBULIFERA Brand & Diederich – WIs:
Saaremaa, Tagamõisa, Tammese, alvar
(58°25,5’N, 21°59,7’E), on calcareous pebbles
and soil over mosses in Sedo-Scleranthenea,
24 July 2004, leg. S. Boch T31/B11.5. Freq.:
rr. – This only recently described species
(Sérusiaux et al., 1999) is known from
northern and western Europe in lowland
areas. It has been confused with Agonimia
gelatinosa, which is a mountainous-boreal
species. A. globulifera differs from this
species by the thallus possessing numerous
shiny, black sterile globules (young fruiting
bodies). The common A. tristicula has a
distinct squamulose thallus, no globules
and larger ascospores, 2 per ascus.
# ARTHONIA FARINACEA (H. Olivier) R. Sant. – NW:
Raplamaa, Märjamaa comm., Niidiaia wooded meadow with oaks (58°53’N 24°28’E), on
soralia of Ramalina farinacea on Quercus
robur, 14 July 2004, leg. & det. A. Suija No.
45. Freq.: rr.
# DIDYMELLOPSIS COLLEMATUM (J. Steiner) Grube &
Hafellner – WIs: Hiiumaa, Kadakalaid islet
(58°59’N 23°00’E), limestone bank in the
northern coast, on Collema sp. on ground
mosses, 6 July 2004, leg. A. Suija & I. Jüriado No. 609, det. A. Suija. Freq.: rr.
# ENDOCOCCUS PROTOBLASTENIAE Diederich – WIs:
Hiiumaa, Vohilaid islet (58°55’N 23°02’E),
eastern coast, on Protoblastenia rupestris on
limestone shingle, 7 July 2004, leg. A. Suija
& I. Jüriado No. 657, det. A. Suija. Freq.: rr.
– This species has been previously reported
from Luxemburg (Sérusiaux et al., 1999) and
Sweden (Santesson et al. 2004).
FELLAHANEROPSIS VEZDAE (Coppins & P. James)
Sérus. & Coppins – SW: Pärnumaa, Surju
comm., Sigaste (58°13’2’’N 24°48’18’’E),
spruce and ash dominating eutrophic
paludified forest, on Alnus glutinosa, 10 July
2003, leg. & det. M. Nõmm. Freq.: rr. – The
species is recorded in Norway, Sweden and
Lithuania and is not found in Finland and
Latvia (Motiejúnaité, 1999; Piteráns, 2001;
Santesson et al., 2004).
# MONODICTYS FULIGINOSA Etayo – NE: Ida-Virumaa,
Oonurme forestry, q. 82/8, (59°08’45’’N
26°57’17’’E), Oxalis myrtillus-Vaccinium
uliginosum type aspen forest with some
Picea abies and Betula pendula, on thallus
of Lobaria pulmonaria on Populus tremula,
23 Jan 2004, leg. I. Jüriado, det. A. Suija.
Freq.: rr. – This inconspicuous hyphomycete, restricted to L. pulmonaria has been
recorded from Spain and Scotland (Etayo
& Diederich, 1996), Canary islands (Etayo,
1996) and Russia (Hermansson et al., 1998;
Zhurbenko, 2001).
NAETEROCYMBE FRAXINI (A. Massal) R.C. Harris
– NE: Jõgevamaa, Tabivere comm., Sortsi
(58°34’27’’N 26°25’12’’E), birch and spruce
dominating eutrophic paludified forest, on
Alnus incana, 16 Sept 2003, leg. & det. M.
Nõmm. Freq.: rr. – The species is recorded in
Norway and Sweden, not found in Finland,
Latvia and Lithuania (Motiejúnaité, 1999;
Piteráns, 2001; Santesson et al., 2004).
# PHACOPIS FUSCA (Triebel & Rambold) Diederich
– WIs: Saaremaa, Muhu island, at the
edge of Raugi wooded meadow (58°39’N
23°17’E), on Xanthoparmelia sp. on granite,
29 May 2004, leg. & det. A. Suija; Hiiumaa,
Vohilaid islet, alvar (58°55’N 23°02’E), on
Xanthoparmelia sp. (infected also with
Lichenostigma cosmopolites) on granite, 7
July 2004, leg. I. Jüriado & A. Suija No.
632, det. A. Suija. Freq.: rr.
# PRONECTRIA ROBERGEI (Mont. & Desm.) Lowen
– NW: Raplamaa, Kõnnumaa Landscape
136 Folia Cryptog. Estonica
Reserve, near Lalli village, former gravel
quarry (58°53’N 25°00’E), on Peltigera sp.
(infected also with Graphium aphthosae and
Corticifraga fuckelii) on soil, 1 Oct 2003, leg.
& det. A. Suija No. 13. Freq.: rr.
SARCOSAGIUM CAMPESTRE (Fr.) Poetsch & Schied.
– SE: Võrumaa, Karula National Park, near
to the Pautsjärve reservation, former gravel
quarry (57°41’N 26°29’E), on Peltigera sp.
(infected also with Graphium aphthosae) and
on soil mosses, 24 Aug 2003, leg. & det. A.
Suija (TU-26822). Freq.: rr. – Sarcosagium
campestre is one of a few ephemeral lichens.
The life-cycle of this species has been described by Gilbert (2004) in detail. According
to his field observations, the fruit-bodies of
S. campestre are detected in rather short
period in a year, from the late summer to
the beginning of spring.
# SPHAERELLOTHECIUM ARANEOSUM (Arnold) Zopf
var. CLADONIAE Alstrup & Zhurb. – WIs:
Hiiumaa, Vohilaid islet (58°55’N 23°02’E),
alvar in central part of the islet, on Cladonia
symphycarpia on soil, 7 July 2004, leg. I.
Jüriado & A. Suija No. 626, det. A. Suija.
Freq.: rr. – This a while ago described
taxon (Zhurbenko & Alstrup, 2004) is
rather easily noticed through the dark
net-like mycelium bound with fruit-bodies
on primary squamules of various Cladonia
species growing on soil.
THELIDIUM MINUTULUM Körb. – NW: Raplamaa,
Kädva, Kädva stream (58°54’N 25°05’E), on
inundated stones, 15 July 2004, leg. A. Suija
No. 56, det. H. Thüs. Freq.: rr. – Thelidium
minutulum was repeatedly reported from
European watercourses, e.g. by Gilbert
(1996) from British limestone creeks, by
Keller (2000) on gneiss from the fluvial
mesic Zone in the river Teigitsch in Austria
and also on schist in the fluvial mesic zone
of the small river Wisper in Germany (Thüs,
2002: p. 191). Certainly it is a species with
a very wide ecological amplitude and will be
found in other habitats in Estonia also.
ACKNOWLEDGEMENTS
Maarja Nõmm thanks Dr. Christine Keller (Switzerland) for verification of Fellhaneropsis vezdae
and Naeterocymbe fraxini; Ave Suija thanks Dr.
Holger Thüs (Germany) for determining and for
comments on Thelidium minutulum.
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