Folia Cryptog. Estonica, Fasc. 57: 73–84 (2020)
https://doi.org/10.12697/fce.2020.57.09
Where the interesting species grow – remarkable records of lichens
and lichenicolous fungi found during a Nordic Lichen Society
meeting in Estonia
Ave Suija1, Inga Jüriado1, Piret Lõhmus1, Rolands Moisejevs2, Jurga Motiejūnaitė3,
Andrei Tsurykau4,5, Martin Kukwa6
Institute of Ecology and Earth Sciences, University of Tartu, Lai 40, EE-51005 Tartu, Estonia.
E-mails: ave.suija@ut.ee; inga.juriado@ut.ee; piret.lohmus@ut.ee
2
Institute of Life Sciences and Technology, Daugavpils University, Parades 1A, LV-5401 Daugavpils, Latvia. E-mail:
rolands.moisejevs@biology.lv
3
Institute of Botany, Nature Research Centre, Žaliųjų Ežerų 49, LT-08406 Vilnius, Lithuania.
E-mail: jurga.motiejunaite@gamtc.lt
4
Department of Biology, Francisk Skorina Gomel State University, Sovetskaja 104, BY-246019 Gomel, Belarus.
E-mail: tsurykau@gmail.com
5
Department of Ecology, Botany and Nature Protection, Institute of Natural Sciences, Samara National Research
University, Moskovskoye road 34, RU-443086 Samara, Russia
6
Department of Plant Taxonomy and Nature Conservation, Faculty of Biology, University of Gdańsk, Wita Stwosza 59,
PL-80–308 Gdańsk, Poland. E-mail: martin.kukwa@ug.edu.pl
1
Abstract: In August 2019, the Nordic Lichen Society held its bi-annual meeting and excursion in south-western Estonia. The
most remarkable findings of lichenized and lichenicolous fungi are recorded herewith, including nine new species (of them
two lichenicolous), and one new intraspecific taxon for the country. Full species lists are provided for two notable locations,
sandstone outcrop at the river Pärnu and an oak woodland in the Naissoo Nature Reserve, for which no previous data were
available, to illustrate the importance of collective survey effort.
Kokkuvõte: 2019. a augustis toimus Eestis korraline, iga kahe aasta järel korraldatav Põhjamaade Lihhenoloogide Ühingu
kokkutulek ja ekskursioon. Artiklis anname ülevaate huvitavatest lihheniseerunud ja lihhenikoolsete liikide leidudest, sealhulgas
üheksa (neist kahe lihhenikoolse) liigi esmaleiust Eestis ja ühe liigisisese taksoni esmaleiust. Lisaks esitatakse lihheniseerunud,
lihhenikoolsete ja neile lähedaste seente täisnimekirjad, mis koostati kahe elupaiga – Tori Põrgu Maastikukaitseala
liivakivipaljandi ning Naissoo Looduskaitseala koosseisu jääva Naissoo tammiku – kohta.
Keywords: new species, red-listed species, sandstone lichens, lichens in oakwood
INTRODUCTION
Nordic Lichen Society (NLS) is an organization
of Nordic and Baltic lichenology aiming to share
knowledge about lichen diversity between professionals and amateurs, and to educate next
generation of researchers and lichen enthusiasts. The society holds bi-annual meetings and
excursions mainly within member countries,
and this time, the opportunity to organize the
meeting was given to Estonia. During 6–10
August 2019, the 23rd NLS meeting was held
in south-western Estonia, consisting of four
full-days field excursions to different habitats
including coastal meadows and dry calcareous
grasslands (alvars), sandstone and limestone
cliffs, and different wooded habitats. In addi-
tion to the field activities, Martin Kukwa from
the University of Gdansk gave a lecture and
held a workshop about sterile sorediate lichens
containing usnic acid and xanthones.
The meetings bring together professionals and
amateurs, and it is not a surprise that such
gatherings notably raise the knowledge about
biodiversity of the region (see, e.g., Thell et
al., 2014; Holien et al., 2016). Several locally
new lichenized or lichenicolous taxa have been
recorded in recent international lichenological
gatherings in Estonia: e.g., 17 (of them 10 lichenicolous) species in 14th symposium of Baltic
Mycologists and Lichenologists (BLS) (Halonen
74
Folia Cryptog. Estonica
et al., 2000), 30 (13 lichenicolous) species in
5th symposium of International Association of
Lichenologists (IAL) (Aptroot et al., 2005) and 11
(6 lichenicolous) species in 17th BLS symposium
(Suija et al., 2009). To follow the idea, we asked
the participants to survey and record as many
lichenized and lichenicolous species as possible
in two locations that represented little-studied
habitat types in Estonia and from where previous data were missing. These habitats were
middle-Devonian sandstone outcrops at the
bank of the Pärnu river in the Tori Põrgu Landscape Reserve and oak woodland in the Naissoo Nature Reserve. In this paper, we present
findings that (1) represent records of new, rare
(with less than ten known localities), protected,
red-listed or otherwise interesting species in Estonia from 12 locations that were visited during
the meeting; and (2) provide full lists of lichens
and lichenicolous fungi found in two study sites.
MATERIALS AND METHODS
Study area
South-West of Estonia and islands were chosen
as the site for NLS meeting because of the richness of habitats valuable for lichens. During
the 4-days meeting, we visited 12 locations (Fig.
1), covering oak-dominated woodlands, alvars,
sandstone and limestone outcrops, a park
around manor house, and old-growth forests. We
made detailed surveys within limited time frame
and compiled lists of lichenized and lichenicolous species for two locations (4 and 10 in the
list below). Short descriptions of these habitats
are given before their species lists.
List of localities
6 August 2019 (all localities in Pärnu County,
Kihnu island)
1. Rootsiküla, north-western coast of Kihnu
island, juniper shrubland in pasture and
the granite stones at the sea, 58.10007ºN,
23.9764ºE
2. Rootsiküla, surroundings of Kihnu lighthouse, 58.09906ºN, 23.96923ºE
3. Linaküla, wooded and coastal meadows with
erratic boulders, 58.137699ºN, 23.966962ºE
7 August 2019 (all localities in Pärnu County)
4. Tori community, Tori borough, Tori Põrgu
(Tori Hell), sandstone outcrop at the river
Pärnu, 58.483497ºN, 24.816716ºE
5. Tori community, Tori borough, Tori cemetery,
58.483139ºN, 24.81857ºE
6. Häädemeeste community, Nigula Nature
Reserve, Nigula bog study trail, Salupeaksi
bog island with old-growth broad-leaved
deciduous forest, 58.018707ºN, 24.68077ºE
8 August 2019
7. Saare County, Muhu island, Muhu community, Nõmmküla alvar, 58.66775ºN,
23.20594ºE
8. Saare County, Muhu island, Üügu Nature
Park, Üügu cliffs and alvar, 58.67169ºN,
23.2373ºE
9. Pärnu County, Lääneranna community,
Puhtu-Laelatu Nature Reserve, Puhtu peninsula, old-growth broad-leaved deciduous
forest, 58.55744ºN, 23.55018ºE
9 August 2019 (all localities in Pärnu County)
10. Lääneranna community, Naissoo Nature
Reserve, Naissoo oak forest mixed with some
birches, 58.609033ºN, 24.187376ºE
11. Lääneranna community, Nedrema Nature Reserve, Nedrema wooded meadow,
58.538175ºN, 24.071151ºE
12. Pärnu, Tõstamaa borough, Tõstamaa park
around Tõstamaa manor house, park
with Acer platanoides, Larix europaea and
Quercus robur, 58.343499ºN, 23.997766ºE
Species identification
To confirm field identifications of some specimens, lichen substances were detected using
thin layer chromatography method described
in Orange et al. (2001) using solvent system A
(AS, AT) and C (AT, MK), and DNA sequences
were analyzed. DNA was extracted with a lysis
procedure, amplified, purified and sequenced
following the protocols in Voitk et al. (2020). The
internal transcribed spacer (nuITS) region was
amplified using the primer pair ITS0F / LA-W
(Tedersoo et al., 2008). The voucher specimens
are deposited in BILAS, DAU, GSU, TU, and
UGDA, and the DNA sequences are accessible
under UDB-codes through public Web output
UNITE (http://unite.ut.ee; Kõljalg et al., 2013).
The distribution and rarity data were extracted
from Data management and Publishing Platform
PlutoF (https://plutof.ut.ee).
RESULTS
During the 4-days field excursions, ten new taxa
for Estonia were recorded, among them seven
75
Fig. 1. The excursion sites of the 23rd Nordic Lichen Society meeting in Estonia
lichenized species, one lichenized intraspecific
taxon and two lichenicolous species. New localities were found for two threatened and three
near-threatened species, according to the recent
assessment of the threat status of Estonian lichens (Lõhmus et al., 2019). These species are:
Enchylium limosum (red-listed category Vulnerable) and Lobaria pulmonaria (Vulnerable and III
protection category), and Cladonia portentosa,
Peltigera ponojensis and Sclerophora pallida
(red-listed category Near Threatened, the latter
also III protection category). New localities were
found for 14 rare species, i.e species with up to
10 localities in Estonia. However, the frequency
class sensu Randlane & Saag (1999) changed
for one lichenized – Ochrolechia bahusiensis –
and for two lichenicolous species – Abrothallus
caerulescens (from rare to rather rare) and Trichonectria rubefaciens (from very rare to rare).
During the excursions several protected and
/or red-listed species were re-found in previously known localities, e.g. Lobaria pulmonaria
in Nigula Nature Reserve and in Puhtu-Laelatu
Nature Reserve, Gyalolechia bracteata (Critically
Endangered and III category), Placidium squamulosum (Near Threatened), Solorina saccata (Endangered and II category), Vulpicida juniperinus
(Near Threatened) in Üügu Landscape Reserve
on Muhu island.
THE SPECIES LISTS
Abbreviations of the names of collectors and determiners: AS – Ave Suija, AT – Andrei Tsurykau,
IJ – Inga Jüriado, JM – Jurga Motiejūnaitė, MK –
Martin Kukwa, PL – Piret Lõhmus, RM – Rolands
Moisejevs. Species that are new to Estonia are
marked in bold. Lichenicolous / algicolous species are marked with #, non-lichenized species
with + and indicator species of Woodland Key
Habitat (WKH) (Anonymous, 2017) with !
New, rare, protected, threatened and otherwise interesting species
# AbrothAllus cAerulescens I. Kotte – 7: on Xanthoparmelia conspersa on granite, AS (TU86821).
76
Folia Cryptog. Estonica
This is the sixth locality for the species, and the
first record in Muhu island.
Botryolepraria lesdainii (Hue) Canals, Hern.Mar., Gómez-Bolea & Llimona – 8: cave, 3 m
from entrance, on limestone, MK 20494 (UGDA),
TLC: lesdainin. The species usually occurs on
strongly shaded calcareous rocks, where it
grows directly on rocks or on calcareous bryophytes in rain-sheltered habitats, but sometimes
it is found also on other substrates (Laundon,
1992; Baruffo et al., 2006; Kukwa & Czarnota,
2008). So far B. lesdainii has been reported from
Europe (e.g., Austria, Belgium, France, Germany, Italy, Finland, the British Isles, Poland,
Portugal, Spain, Sweden) and North America
(e.g., Laundon, 1992; Kümmerling & Leuckert,
1993; Kukwa, 2000; Baruffo et al., 2006; Kukwa
& Czarnota, 2008).
Candelariella effloresCens R. C. Harris & W.
R. Buck (aggregate) – 10: on Quercus robur,
leg. AT, det. MK (GSU). It is the single taxon
in Candelariella which produces soredia and
has polysporous asci. Sterile specimens are
similar to C. reflexa which differs by its much
larger, indistinctly effigurate thallus, crateriform
soralia arising in the center of the thallus and
larger soredia (Westberg, 2007). Candelariella
efflorescens is a common species in Europe,
North America, reported also from the Caucasus
(Westberg, 2007; Gasparyan & Sipman, 2016;
Ismailov et al., 2017). The specimen is sterile
but well-developed and has delimited punctiform soralia. However, according to Westberg &
Clerc (2012), when sterile, it is usually impossible to separate between C. efforescens and C.
xanthostigmoides and therefore the specimen is
referred here as C. efforescens aggregate.
Catillaria CroatiCa Zahlbr. – 6: on Corylus avellana, MK 20475 and 20476 (UGDA), TLC: no
lichen substances (traces of terpenoids are the
same as in the analyzed bark). This is a rarely
reported species as it is almost always found
sterile, but can be distinguished by light greengrey, superficial, well or poorly developed to immersed thallus consisting of scattered or almost
contiguous areoles and numerous soralia, which
are rounded, flat, convex or weakly capitate,
discrete or partly fused and forming a leprose
crust. Soredia are green, but externally pale
brown pigmented in some specimens (Kukwa et
al., 2012). The species has been reported from
Europe (Austria, Belgium, Croatia, France, Lux-
embourg, Poland, Romania, Slovakia, Slovenia
and Ukraine) and North America (Kukwa et al.,
2012 and literature cited therein).
# chAenothecopsis vAinioAnA (Nádv.) Tibell – 10:
on Quercus robur, together with Anisomeridium
polypori, leg. AS, det. PL (TU87247). Rather rare
species in Estonia, all records are from wooded
meadows and oak-dominated woodlands.
Cladonia monomorpha Aptroot, Sipman & Herk
– 4: on bryophytes growing on sandstone, AT
(GSU), TLC: fumarprotocetraric acid; 10: on
saxicolous bryophytes, MK 20531 (UGDA), TLC:
fumarprotocetraric and protocetraric acids; AT
(GSU). This is a member of a taxonomically difficult Cladonia pyxidata group and has not always been distinguished from C. pyxidata (Kowalewska et al., 2008 and literature cited therein),
but recent molecular studies treat this species
as a distinct taxon (Stenroos et al., 2019). The
species is most similar to C. pyxidata and C.
pocillum, from which it differs by the presence of
bullate plates (commonly with whitish margins)
inside and outside scyphi, and glomerulose apothecia (Aptroot et al., 2001; Kowalewska et al.,
2008; Tsurykau & Golubkov, 2015). Chemically
similar C. chlorophaea and C. fimbriata differ
by producing soredia. Cladonia monomorpha is
widely distributed in Europe, being also known
from North America, including Greenland
(Kowalewska et al., 2008), and Asia (Mongolia,
Turkey and Russian Arctic) (Kowalewska et
al., 2008; Osyczka et al., 2011; Golubkov &
Tsurykau, 2017). Cladonia monomorpha has
been reported previously from Estonia (Aptroot
et al., 2005), but the specimens from a single
locality (TU28247, TU30991, TU30992a) were
redetermined as Cladonia pyxidata.
# didymoCyrtis ramalinae (Roberge ex Desm.)
Ertz, Diederich & Hafellner – 3: on the thallus
of Ramalina fastigiata growing on Padus avium,
MK 20447 (UGDA). The finding represents the
asexual stage of the fungus. The dimensions of
hyaline, ellipsoid conidia are 5.5–6.5 × 3.5 μm.
The fungus, mainly its asexual stage, is recorded
from many countries in Europe but the species
is also known in Africa and Australasia (Ertz et
al., 2015).
enchylium limosum (Ach.) Otálora, P. M. Jørg.
& Wedin – 4: on sandstone, AS (TU86765,
TU86767). The species is rather rare (this is the
seventh record), and it belongs to the red-listed
77
category Vulnerable according to the the latest
version of the Red List of Estonia (Lõhmus et
al., 2019).
h aleCania viridesCens Coppins & P. James
– 6: on wood of log, MK 20474 (UGDA), TLC:
argopsin (major), norargopsin (minor), ‘gracilenta unknown 1’. This is the second, but the
first published record in Estonia. The species
was previously found from Heinlaid islet in
Hiiu County as growing on twig of Rhamnus
cathartica (TU45760.a.), but because of the
scarce material remained unreported. Halecania
viridescens is usually sterile and characterized
by small, fragile, usually punctiform or rarely
confluent soralia (often with a brown pigment
in the external soredia), and the presence of
argopsin and ‘gracilenta unknown 1’ (Tønsberg,
1992; Kukwa & Jabłońska, 2009). The species
is rarely reported, but widespread and has been
reported from several countries in Europe, North
America, central America (Guatemala) and Africa
(the Canary Islands) (Kukwa & Jabłońska, 2009
and literature cited therein).
lecAnorA persimilis (Th. Fr.) Arnold – 2: roadside
trees, on Fraxinus excelsior, MK 20442a (UGDA).
This is the second record of this species in Estonia, known previously from northern Estonia
(Martin et al., 2011).
lecAnorA subcArpineA Szatala – 10: on Quercus
robur, MK 20536 (UGDA), TLC: atranorin (major), psoromic acid (minor), unknown (trace). The
disc of ascomata is slightly pruinose, C+ yellow,
apothecial margin Pd+ yellow. The species is rare
in Estonia (this is the fifth locality), the closest
locality in Nedrema wooded meadow is ca 10
kilometres (Aptroot et al., 2005).
leCidella elaeoChroma f. soralifera (Erichsen)
D. Hawksw. – 3: on Fraxinus excelsior, MK 20449
(UGDA); leg. AS, det. MK (TU86789). This is a
rarely reported sorediate form of a very common
Lecidella elaeochroma (Tønsberg, 1992).
lepraria rigidula (B. de Lesd.) Tønsberg – 1: on
Juniperus communis, MK 20435 (UGDA), TLC:
atranorin, nephrosteranic acid. The presence of
a rare fatty acid, nephrosteranic acid, and soredia with long projecting hyphae are diagnostic
characters of this taxon. It is a rather common
epiphytic species in open habitats in Europe
but found also on other substrates and outside
Europe (Tønsberg, 1992; Baruffo et al., 2006;
Kukwa, 2006; Saag et al., 2009).
leprAriA vouAuxii (Hue) R. C. Harris – 4: on
sandstone, MK 20456, MK 20458 (UGDA); 9:
together with L. finkii on Ulmus scabra, MK
20515 (UGDA); 12: on Acer platanoides, AS
(TU87653). All specimens contain pannaric acid
6-methylester detected with TLC. The species is
rather rare, known from less than ten localities
in Estonia. Lepraria vouauxii grows on various substrates, but often on tree bark in open
habitats (Tønsberg, 1992; Baruffo et al. 2006;
Kukwa, 2006; Saag et al., 2009). Many records
from South America were recently segregated
as L. cryptovouauxii (Guzow-Krzemińska et al.,
2019).
lobAriA pulmonAriA (L.) Hoffm. – 12: on Quercus
robur, AS (not collected). The species is protected
in Estonia and was recently evaluated according
to IUCN criteria as Vulnerable (Lõhmus et al.,
2019). It was recorded growing on a single tree
in Tõstamaa park.
ochrolechiA bAhusiensis H. Magn. – 10: together
with Lepra albescens (KC negative) on Quercus
robur, MK 20532 (UGDA); 12: on Pinus sylvestris, MK 20550 (UGDA), TLC: gyrophoric acid,
murolic acid complex, unidentified pigments.
The species had ten localities in Estonia but
is probably under-recorded due to similarities
with several other Ochrolechia species. This is
a widespread species in Europe which is mostly
confined to lower altitudes, and found mainly
on bark of deciduous trees in various types of
ecosystems (Kukwa, 2011).
o chrolechiA szAtAlAensis Verseghy – 7: on
Juniperus communis, MK 20505 (UGDA;
UDB0779091). The cortex of apothecial margin
in the Estonian specimen is not glassy at the
base in sectioned apothecia, but white and thus
more similar to O. upsaliensis (Kukwa, 2011).
However, the nuITS sequences are one-to-one
identical to the sequences of O. szatalaensis
(MK811817, MK811903, MK811865, FR799244)
deposited in GenBank. This is a frequent but
scattered species in Estonia, however, as morphological characters of the specimen deviate
from the known description, we still report it
here.
parmelia serrana A. Crespo, M. C. Molina & D.
Hawksw. – 6: on Betula pendula, leg. AS, det.
AT & AS (TU86826; UDB0778439); AT (GSU;
verified by A. Thell), TLC of both collections:
salazinic acid, atranorin and fatty acids. The
78
Folia Cryptog. Estonica
population in Salupeaksi is a mixture of two
cryptic species, P. ernstiae and P. serrana,
which are realistically identifiable only by DNA
sequences (Haugan & Timdal, 2019; Tsurykau
et al., 2019). We compared nuITS sequences
and composition of lichen substances of three
specimens, two of which corresponded to P.
serrana and one to P. ernstiae (E. Ossowska,
pers. comm.). The latter specimen is deposited
in UGDA.
# pronectriA erythrinellA (Nyl.) Lowen. – 5: all
records on Peltigera praetextata growing on
mossy stone fence, AS (TU86773), AT (GSU), JM
(BILAS). The species has less than ten scattered
localities in Estonia.
# pronectriA sAntessonii (Lowen & D. Hawksw.)
Lowen – 3: on Anaptychia ciliaris on Fraxinus
excelsior, AS (TU87916), AT (GSU), both det.
JM. The species has been found previously
only once in Estonia (Martin et al., 2012). The
characteristics of the specimens are in concordance with the protologue of the species (Lowen &
Hawksworth, 1986). The dimensions of the ascospores of TU87916 are 12.5–(13.96±1.05)–16
× 5–(5.8±0.59)–7 μm (n=13).
# sclerococcum homoclinellum (Nyl.) Ertz &
Diederich (syn. Dactylospora homoclinella (Nyl.)
Hafellner) – 7: on Protoparmeliopsis muralis on
granite, AS (TU86806; UDB0778820). This is
the second record of this species from Estonia,
previously reported as growing on Buellia griseovirens (Suija, 2005).
sclerophorA pAllidA (Pers.) Y. J. Yao & Spooner
– 5: on Fraxinus excelsior, leg. AS, det. PL
(TU86820); 6: RM (DAU600001114); PL (TU).
This species is protected in Estonia and has
recently been evaluated as Near Threatened in
the latest version of the national Red List (Lõhmus et al., 2019).
# s pirographa triCupulata (F. Berger & E.
Zimm.) Flakus, Etayo & Miadlikowska (syn. Cornutispora tricupulata F. Berger & E. Zimm) – 11:
on Physcia cf. tenella on willow twig, MK 20547
(UGDA). Spirographa tricupulata is characterized
by having conidia composed of three equal arms,
4–5.2 × 2.4–3 μm, which are strongly swollen at
the base and develop 1.2–1.5 μm long cilia at the
ends (Zimmermann & Berger, 2018). The conidia
of the Estonian specimen have swollen, almost
equal arms, 4 × 2 μm and with cilia c. 1.5 μm
long. So far, the species has been known only
from Austria (Zimmermann & Berger, 2018).
This is the second Spirographa species besides S.
lichenicola (D. Hawksw. & Sutton) Flakus, Etayo
& Miadlikowska (syn. Cornutispora lichenicola
D. Hawksw. & B. Sutton) that is reported from
Estonia (Aptroot et al., 2005).
# stigmidium solorinArium (Vain.) D. Hawksw.
– 8: on Solorina sp. on limestone cliffs, AS
(TU86805). This is the second record of this
Solorina-specialized lichenicolous species in
Estonia (Suija et al., 2009).
# trichonectriA rubefAciens (Ellis & Everh.)
Diederich & Schoers – 10: on Parmelia sulcata,
AT (not collected). This is the third locality for the
species in Estonia. Trichonectria rubefaciens is
one of those lichenicolous species that is easily
recognizable in field because of characteristic
reddish orange perithecia on the discoloured
thalli of Parmelia species (Sérusiaux et al.,
1999).
# xAnthoriicolA physciAe (Kalchbr.) D. Hawksw.
– 3: on Xanthoria parietina, AS (TU86790). The
dematiaceous lichenicolous species has less
than ten localities in Estonia but is probably
under-recorded although it is one of the most
easily recognizable species among lichenicolous
fungi as the host apothecia turn black due to
the infection. Microscopically, each blackened
apothecium contains mycelium that forms conidiogenous area in the host hymenium near the
surface of the apothecium. The conidiogenous
area is characteristic for this fungus as it is
formed by penicillate, short and brown, monophialidic conidiogenous cells. Brown, globose
and warted conidia in dimensions of 3.5–6 μm
that develop terminally on conidiogenous cells
lie on the surface of the host apothecium (Hawksworth & Punithalingam, 1973) giving apothecia
a black appearance.
xylogrAphA pArAllelA (Ach.) Fr. – 10: on wood,
together with Buellia griseovirens, Lecanora symmicta and Placynthiella icmalea, IJ (TU88412).
This is the tenth record of the lignicolous species in Estonia.
List of species on Tori sandstone outcrop
The Tori Põrgu Landscape Reserve was created
in 1959 to protect middle-Devonian sandstone
bank and caves at the Pärnu river in south-western Estonia. The length of the bank is ca 400 m
and its maximum height is 8.25 m being thus
79
the most representative outcrop of this epoch
in the East Baltic area (Fig. 2). There are three
caves created by the stream waters eroding the
sandstone bank and expanded by human activities. The largest cave is called “Tori Põrgu” (“Tori
Hell” in translation) as in the Estonian folklore,
this was the lodging of horned family, namely
Vanapagan (“The Old Heathen”, also known as
“The Old Devil” or „The Old Empty One”) and
others (Laugaste & Liiv, 1970).
Systematic studies of lichens on Devonian
sandstone cliffs are scarce in Estonia. There is
one diploma work (Tenson, 1970), and one species inventory (Ingerpuu & Suija, 2010) dealing
with sandstone lichens in central and eastern
parts of Estonia. The list here is the first from
the southwestern part of Estonia, including
26 species, all recorded as growing directly on
sandstone. The number of species is close to
that of Kallaste sandstone outcrop in eastern
Estonia (28 species; Ingerpuu & Suija, 2010).
The list includes three notable species, Cladonia
monomorpha, Enchylium limosum and Lepraria
vouauxii (see the list above).
AthAlliA holocArpA (Hoffm.) Arup, Frödén &
Søchting – AS (TU86768).
bAcidiA bAgliettoAnA (A. Massal. & De Not.) Jatta
– AS (TU86772).
bAeomyces rufus (Huds.) Rebent. – AS (TU89609).
In some spots, the specimens of Baeomyces
were surrounded by the black mycelial
network. Sphaerellothecium conioides is the
only species that has such dark-brown vegetative hyphae and grows on B. rufus (Roux
& Triebel, 1994). However, no ascomata were
developed and therefore the identity of the
fungus remained unclear.
bilimbiA sAbuletorum (Schreb.) Arnold – AS
(TU86766).
chAenothecA furfurAceA (L.) Tibell – AS.
c lAdoniA chlorophAeA (Flörke ex Sommerf.)
Spreng. – AS (TU86825; UDB0778437); MK
20457 (UGDA); AT (GSU).
Cladonia monomorpha Aptroot, Sipman & Herk
– AT (GSU).
enchylium limosum (Ach.) Otálora, P. M. Jørg. &
Wedin – AS (TU86767; TU86765).
everniA prunAstri (L.) Ach. – AS.
f lAvoplAcA citrinA (Hoffm.) Arup, Frödén &
Søchting – AS (TU86764).
hAemAtommA ochroleucum (Neck.) J. R. Laundon
– MK 20463 (UGDA).
lecidellA cArpAthicA Körb. – RM (DAU 600001111).
leprAriA finkii (B. de Lesd.) R. C. Harris (syn. L.
lobificans auct.) – MK 20459 (UGDA).
leprAriA incAnA (L.) Ach. – MK 20467 (UGDA).
leprAriA vouAuxii (Hue) R. C. Harris – MK 20456;
MK 20458 (UGDA).
leproplAcA chrysodetA (Vain. ex Räsänen) J.
R. Laundon – AS (TU86800); MK 20460
(UGDA).
pArmeliA sulcAtA Taylor – AS.
peltigerA cAninA (L.) Willd. – MK 20464 (UGDA).
peltigerA didActylA (With.) J. R. Laundon – AS.
peltigerA membrAnAceA (Ach.) Nyl. – MK 20465
(UGDA).
p eltigerA neckeri Hepp ex Müll. Arg. – AS
(TU86770); MK 20466 (UGDA).
p eltigerA prAetextAtA (Flörke ex Sommerf.)
Zopf – AS.
p el tigerA rufescens (Weiss) Humb. – AS
(TU86769).
phlyctis ArgenA (Ach.) Flot. – MK 20461 (UGDA).
rAmAlinA fArinAceA (L.) Ach. – AS (TU86771); MK
2045 (UGDA).
rAmAlinA pollinAriA (Westr.) Ach. – MK 20455
(UGDA).
List of species in the oak wood in Naissoo
Nature Reserve
Fig. 2. The sandstone outcrop of Tori Põrgu
Landscape Reserve. Photo: Jurga Motiejūnaitė
The Naissoo Nature Reserve (115.71 ha) is located in south-western part of Estonia and was
created in 1964 to protect old oak woods and
spruce forests. The oldest oaks (Quercus robur)
80
Folia Cryptog. Estonica
in the area are 230 years old. The calciferous
meadows (alvars) are less represented in the
area. The oakwoods are probably former wooded
pastures and wooded meadows that are now
overgrown due to the cessation of mowing and
grazing (Naissoo looduskaitseala ...). In some
places, the remnants of limestone fences and
piles of stones are visible under the oak trees.
The list is based on the collective survey effort made during 1.5 hours in a limited area
in eastern part of the reserve. The species list
includes 95 species, of which 83 are lichenized,
11 lichenicolous-algicolous fungi, and one is
a non-lichenized calicioid fungus. During the
limited timeframe, the participants found four
rare species (Lecanora subcarpinea, Ochrolechia
bahusiensis, Trichonectria rubefaciens and
Xylographa parallela), and two species new to
Estonia (Candelariella efflorescens and Cladonia monomopha). In addition, participants
recorded several Woodland Key Habitat (WKH)
species that grow mainly on oaks in the area.
No collector’s initials are indicated for the common species that were registered by most of the
participants.
! AcrocordiA gemmAtA (Ach.) A. Massal. – on
Quercus robur, PL (TU87927).
! AlyxoriA vAriA (Pers.) Ertz & Tehler – on Q.
robur, MK 20527 (UGDA); AS (TU86801).
AnAptychiA ciliAris (L.) Körb. – on Q. robur.
Anisomeridium polypori (Ellis & Everh.) M. E.
Barr – on Q. robur, AS (TU87247.b).
ArthoniA mediellA Nyl. – on Q. robur, MK 20520
(UGDA); on Picea abies, MK 20526 (UGDA);
PL (TU87926).
ArthoniA rAdiAtA (Pers.) Ach. – on twig of Q.
robur, MK 20525 (UGDA).
ArthoniA spAdiceA Leight. – on Q. robur, PL.
ArthoniA vinosA Leight. – on Q. robur.
AspiciliA contortA subsp. hoffmAnniAnA S. Ekman & Fröberg – on limestone, IJ.
# AtheliA ArAchnoideA (Berk.) Jülich – on Xanthoria parietina.
bAcidiA ArceutinA (Ach.) Arnold – on Q. robur, AS.
biAtorA efflorescens (Hedl.) Räsänen – on Q.
robur, MK 20533 (UGDA).
b ilimbiA sAbuletorum (Schreb.) Arnold – on
mosses on limestone, IJ (TU88410).
buelliA griseovirens (Turner & Borrer ex Sm.)
Almb. – on lignum, IJ (TU88409.c); on lignum, IJ (TU88412.b); on Q. robur, PL; on
Juniperus communis, AS.
cAlicium glAucellum Ach. – on lignum of Q. robur
log, PL (TU87929).
cAlicium sAlicinum Pers. – on Q. robur, PL.
cAlicium viride Pers. – on Q. robur, PL.
Candelariella effloresCens R. C. Harris & W.
R. Buck (aggregate) – on Q. robur, leg. AT,
det. MK (GSU).
cAndelAriellA xAnthostigmA (Ach.) Lettau – on
Q. robur, MK 20535 (UGDA); PL.
! chAenothecA brAchypodA (Ach.) Tibell – on lignum of Q. robur log, PL.
chAenothecA chrysocephAlA (Turner ex Ach.) Th.
Fr. – on Pinus sylvestris, IJ.
chAenothecA trichiAlis (Ach.) Th. Fr. – on P.
sylvestris, IJ; on Q. robur, PL.
# chAenothecopsis vAinioAnA (Nádv.) Tibell – on
Q. robur, leg. AS, det. PL (TU87247.a); PL
(TU87928).
c lAdoniA chlorophAeA (Flörke ex Sommerf.)
Spreng. (aggregate) – IJ.
clAdoniA coniocrAeA (Flörke) Spreng. – on lignum, IJ (TU88408.a); on mosses on limestone, IJ (TU88411.b).
clAdoniA fimbriAtA (L.) Fr. – on Q. robur, IJ.
Cladonia monomorhpa Aptroot, Sipman & Herk
– on saxicolous bryophytes, MK 20531
(UGDA); AT (GSU).
clAdoniA ochrochlorA Flörke – on mosses on
limestone, AS (TU87246.a); on base of
Betula, IJ (TU88405); on Juniperus communis, AS.
clAdoniA pocillum (Ach.) Grognot – on mosses
on limestone, IJ (TU88407).
coenogonium pineti (Schrad. ex Ach.) Lücking &
Lumbsch – on Q. robur AS.
everniA prunAstri (L.) Ach. – on Q. robur.
# heterocephAlAcriA physciAceArum (Diederich)
Millanes & Wedin – on thallus of Physcia
adscendens, AS.
hypocenomyce scAlAris (Ach.) M. Choisy – on old
Betula, IJ (TU88406.b).
hypogymniA physodes (L.) Nyl. – on Q. robur; B.
pendula.
hypogymniA tubulosA (Schaer.) Hav. – on Q. robur.
# intrAlichen sp. – in apothecia of Lecanora, AS
(TU86802).
lAthAgrium fuscovirens (With.) Otálora, P. M.
Jørg. & Wedin – on mosses on limestone,
JM (TU86797.a); IJ.
lecAnorA cArpineA (L.) Vain. – on twig of Q. robur,
MK 20525a (UGDA); on Q. robur, MK 20536a
(UGDA).
lecAnorA chlAroterA Nyl. – on Q. robur.
81
lecAnorA expAllens Ach. – on Q. robur, PL.
lecAnorA pulicAris (Pers.) Ach. – on B. pendula,
IJ.
lecAnorA strobilinA (Spreng.) Kieff. – on twig of
Q. robur, MK 20522 (UGDA).
lecAnorA subcArpineA Szatala – on Q. robur, MK
20536 (UGDA).
lecAnorA symmictA (Ach.) Ach. – on lignum, IJ
(TU88412.c).
lecidellA elAeochromA (Ach.) M. Choisy – all on
Q. robur, MK 20521 (UGDA); MK 20530a
(UGDA); MK 20537 (UGDA).
lecidellA flAvosorediAtA (Vězda) Hertel & Leuckert – on Q. robur, MK.
leprA Albescens (Huds.) Hafellner (syn. Pertusaria albescens (Huds.) M. Choisy & Werner)
– on Q. robur, MK 20532 (UGDA).
leprA AmArA (Ach.) Hafellner (syn. Pertusaria amara (Ach.) Nyl.) – on lignum, IJ (TU88408.b).
leprAriA eburneA J. R. Laundon – both on Q.
robur, MK 20539 (UGDA); MK 20541a
(UGDA).
leprAriA finkii (B. de Lesd.) R. C. Harris – on Q.
robur, MK 20540 (UGDA).
leprAriA incAnA (L.) Ach. – both on Q. robur, MK
20541 (UGDA); MK 20542 (UGDA).
# lichenoconium xAnthoriAe M. S. Christ. – on
Xanthoria parietina, AT (GSU).
# mArchAndiomyces AurAntiAcus (Lasch) Diederich & Etayo – on Physcia tenella, AS.
melAnelixiA glAbrAtulA (Lamy) Sandler & Arup
– on Q. robur.
melAnelixiA subAuriferA (Nyl.) O. Blanco, A. Crespo, Divakar, Essl., D. Hawksw. & Lumbsch
– on Q. robur.
m elAnohAleA exAsperAtulA (Nyl.) O. Blanco,
A. Crespo, Divakar, Essl., D. Hawksw. &
Lumbsch – on Q. robur.
micAreA misellA (Nyl.) Hedl. – on lignum, IJ
(TU88409.a).
# microcAlicium disseminAtum (Ach.) Vain. – on
Q. robur, PL (TU87930).
+ m ycocAlicium subtile (Pers.) Szatala – on
lignum, IJ (TU88409.b); on lignum of Q.
robur, PL.
# nesolechiA oxysporA (Tul.) A. Massal. – all on
thalli of Parmelia sulcata, AT; growing on
twig of Salix, MK 20523 (UGDA), on twig of
Q. robur, AS (TU86774).
ochrolechiA ArboreA (Kreyer) Almb. – on twig of
Q. robur, MK 20524 (UGDA).
ochrolechiA bAhusiensis H. Magn. – on Q. robur,
MK 20532 (UGDA).
opegrAphA vulgAtA (Ach.) Ach. – on Q. robur, PL
(TU87931).
pAchyphiAle fAgicolA (Arnold) Zwackh – on Q.
robur, MK 20535a (UGDA).
pArmeliA ernstiAe Feuerer & A. Thell – on Q.
robur, MK 20529 (UGDA).
pArmeliA sulcAtA Taylor – on B. pendula.
peltigerA cAninA (L.) Willd. – on Q. robur, PL, det.
IJ (TU87932).
peltigerA neckeri Hepp ex Müll. Arg. – on ground
over mosses, IJ (TU89562; UDB0779902).
peltigerA prAetextAtA (Flörke ex Sommerf.) Zopf
– on mosses in stone pile, AS.
pertusAriA coccodes (Ach.) Nyl. – both on Q.
robur, MK 20532a (UGDA); MK 20534
(UGDA).
pertusAriA coronAtA (Ach.) Th. Fr. – on Q. robur,
PL. In the field UV + orange, K+ yellow, then
orange.
pertusAriA leioplAcA DC. – on Q. robur.
phlyctis ArgenA (Spreng.) Flot. – all on Q. robur,
MK 20528 (UGDA); MK 20530 (UGDA); MK
20538 (UGDA); AS (TU86808).
physciA Adscendens (Fr.) H. Olivier – on Q. robur.
physciA AipoliA (Ehrh. ex Humb.) Fürnr. – on
Q. robur.
physciA stellAris (L.) Nyl. – on Q. robur.
physciA tenellA (Scop.) DC. – on Q. robur.
physconiA distortA (With.) J. R. Laundon – on
Q. robur.
physconiA perisidiosA (Erichsen) Moberg – on
Q. robur.
plAcynthiellA icmAleA (Ach.) Coppins & P. James
– on lignum, IJ (TU88412.d); on log of Q.
robur, on wood, PL.
plAtismAtiA glAucA (L.) W. L. Culb. & C. F. Culb.
– on Q. robur.
polycAulionA polycArpA (Hoffm.) Frödén, Arup &
Søchting – on Q. robur.
pseudeverniA furfurAceA (L.) Zopf – on Q. robur.
rAmAlinA fAstigiAtA (Pers.) Ach. – on Q. robur.
scytinium lichenoides (L.) Otálora, P. M. Jørg.
& Wedin – all records on mosses on limestone, JM (TU86797.b); AS (TU87246.b); IJ
(TU88411.a).
toniniopsis subincomptA (Nyl.) Kistenich, Timdal,
Bendiksby & S. Ekman (syn. Bacidia subincompta (Nyl.) Arnold) – on Q. robur, PL.
trApeliopsis flexuosA (Fr.) Coppins & P. James
– on base of Betula, IJ (TU88406.a); on J.
communis.
# tremellA hypogymniAe Diederich & M. S. Christ.
– on Hypogymnia physodes, JM (TU86796).
82
Folia Cryptog. Estonica
# trichonectriA rubefAciens (Ellis & Everh.)
Diederich & Schroers – on thallus of Parmelia sulcata, AT.
verrucAriA dolosA Hepp – on limestone, AS
(TU86823; UDB0778442).
# vouAuxiellA lichenicolA (Linds.) Petr. & Syd.
– on apothecia of Lecanora chlarotera, AS.
vulpicidA pinAstri (Scop.) J.-E. Mattsson & M. J.
Lai – on Q. robur.
xAnthopArmeliA conspersA (Ach.) Hale – on granite stone, AS.
xAnthoriA pArietinA (L.) Th. Fr. – on Q. robur.
xylogrAphA pArAllelA (Ach.) Fr. – on lignum, IJ
(TU88412.a).
ACKNOWLEDGEMENTS
Arne Thell (Lund) and Emilia Ossowska (Gdansk) are thanked for confirming identity of
Parmelia serrana and P. ernstiae respectively.
Rasmus Puussepp (Tartu) is thanked for DNA
work. Ede Oja, Tiina Randlane, Andres Saag,
Maarja Nõmm, Magdalena Kosecka, Sigrid
Maasen and Indrek Tammekänd are thanked for
participation. The financial support for the meeting was provided by the Gambling Tax Council
and by bilateral exchange programme between
Academies of Sciences (enabling participation of
JM and MK). The financial support of AS was
provided by the European Regional Development
Fund (Centre of Excellence EcolChange).
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