New Records of Pannariaceae (Lichenized Ascomycota) from Sakhalin and the Kuril Islands, Russian Far East

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New Records of Pannariaceae (Lichenized Ascomycota) from
Sakhalin and the Kuril Islands, Russian Far East
Author(s): Alexander K. Ezhkin and Per M. Jørgensen
Source: Evansia, 35(2):43-52.
Published By: The American Bryological and Lichenological Society, Inc.
URL: http://www.bioone.org/doi/full/10.1639/0747-9859-35.2.043
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Evansia 35(2): 2018
43
New records of Pannariaceae (lichenized Ascomycota) from Sakhalin and the
Kuril Islands, Russian Far East
Alexander K. Ezhkin
Laboratory of Plant Ecology and Geoecology, Institute of Marine Geology and Geophysics, Far
Eastern Branch of the Russian Academy of Sciences, Nauki, 1B, Yuzhno-Sakhalinsk, 693022,
Russia
E-mail: ezhkin@yandex.ru
Per M. Jørgensen
Department of Natural History, Bergen University Museum, Box 7800, Allégt.41, N’5012
Bergen, Norway
E-mail: per.jorgensen@um.uib.no
Abstract. Eleven taxa in the lichen family Pannariaceae connected with boreal coniferous or
nemoral broad-leaved forests are documented. Notes on their taxonomy, ecology and
geography are provided. Fuscopannaria leprosa is new to Eurasia; Fuscopannaria poeltii,
Pannaria insularis, Pannaria lurida ssp. russellii and Parmeliella miradorensis are new to
Russia.
Keywords. Biodiversity, plant geography, broad-leaved forests, Asia, rare species.
INTRODUCTION
Sakhalin Island and the Kuril Islands are located in the northwestern part of the Pacific Ocean
(Fig. 1). The climate is temperate, with monsoons, cold winters and cool, rainy summers. According
to Takhtadzhyan (1978) Northern Sakhalin belongs to the Okhotsk-Kamchatka Province of the
Circumboreal floristic Region, while Southern Sakhalin along with the southern Kuril Islands
belong to the Sakhalin-Hokkaido Province of the East Asian floristic Region. Most of Sakhalin and
the Kuriles are covered by coniferous forests. In the 20th century, major forests in Sakhalin were
destroyed by wood harvesting and fires. At present time, large territories are occupied with
managed, secondary forests except some places with difficult access.
The first lichenological investigations of Sakhalin and the Kuriles were performed by
Japanese specialists as late as in the beginning of the twentieth century. The first list of lichens of
Sakhalin Island numbered 105 species (Sato 1936a) and 44 species for the Kuriles (Sato 1936b),
among them only one in the family Pannariaceae. In 2002 the list of lichens included 322 species
for Sakhalin and 260 species for the Kuriles (Tchabanenko 2002), and since then, new records were
published in several papers (Taran and Tchabanenko 2003; Joneson and al. 2004; Tchabanenko
2004; Tchabanenko and Taran 2004; Tchabanenko 2006; Galanina 2013, Ezhkin and Galanina
2014, 2016; Kondratyuk et al. 2013; Skirina et al. 2016; Ezhkin 2016; Galanina et al. 2017).
Regardless of the period of Sakhalin lichen flora research—almost 100 years—many areas of the
region are being actively investigated.
Recent papers list just two species of Pannariaceae for Sakhalin—Protopannaria pezizoides
(Weber) Trevis. (Sato 1936a) and Psoroma hypnorum (Vahl.) S. Gray. (Tchabanenko 2002). A
third species, however—Pannaria lurida (Mont.) Nyl.—is also known for Kunashir, Sakhalin, as
well as for the Primorskiy Region of Sakhalin. In the latter region, upwards of ten species of
Pannariaceae may occur (Urbanavichus 2010); some are included in the present paper, while others
are doubtful records in need of confirmation, e.g. Degelia plumbea (Lightf.) P. M. Jørg. and P.
James, which is likely a species of Coccocarpia. In all, 11 taxa in the Pannariaceae are reported in
the present paper as either new to Russia (5) or new to the Russian Far East and/or Sakhalin/Kuriles
(6). This number could grow if further species from neighboring regions in Asia and North America

Evansia 35(2): 2018
44
are found in the future in Sakhalin and the Kuriles. Both of these two neighboring regions are known
to be very rich in species of Pannariaceae (especially in the genera Fuscopannaria and Parmeliella;
Jørgensen 2000b, 2002; 2004, Jørgensen and Kashiwadani 2001a; Tchabanenko 2002; Liu et al.,
2016).
Figure 1. Map of Pannariaceae collecting sites in the Russian Far East (Sakhalin and South
Kuriles).
MATERIAL AND METHODS
Lichen samples were collected by the first author from the Sakhalin Island and the
Kuriles—the Kunashir Islands, the Iturup Islands and the Shikotan Islands—from 2012 through
2017. Collecting localities are specified on the map (Fig. 1). Examined specimens are deposited in
the herbarium of Institute of Marine Geology and Geophysics (SAK). All the material was
examined using standard microscopic techniques with a Wild M3Z Plan stereomicroscope,
Olympus BH-A research microscope and a Carl Zeiss A1 Axio Scope. Photographs were taken
with a Canon EOS 600D digital camera and an Olympus BX51 light microscope. Secondary
metabolites were characterized by high performance thin-layer chromatography (HPTLC)
according to the methods described in Schumm and Elix (2015). Identifications were done with the
help of published keys such as Jørgensen (1978, 2000b) and Jørgensen and Kashiwadani (2001a,
2001b).

Evansia 35(2): 2018
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THE SPECIES
Records new to Russia
1. Fuscopannaria leprosa P.M. Jørg. & Tønsberg (Fig. 2 D)
This species was previously only known from a few collections in Pacific North America
(Jørgensen 2000b) and is here reported for the first time from Eurasia. It has some resemblance to
Fuscopannaria mediterranea (C. Tav.) P.M. Jørg. but has a more crustose thallus which dissolves
completely into soredia (not marginal soralia on upturned lobe-ends). It has a rather special ecology,
preferring to grow up on tree stems in warm, south-facing valley-sides. It has a disjunct
amphibehringian distribution.
Specimens Examined. — Sakhalin, Noglikskiy District, The Dagi River valley, N 52°06'38,43'' E
142°57'27,82'', alt 5m, floodplain forest, on bark of Salix udensis Trautv., 12.10.2012, A.K. Ezhkin
(SAK 1271). Kunashir, Golovnino Volcano surroundings, N 43°52'20.9208'' E 145°36'00.9251'',
alt 89 m, mixed forest, on bark of old Quersus crispula Blume, 23.08.2017, A.K. Ezhkin (SAK
1272).
2. Fuscopannaria poeltii (P.M. Jørg.) P.M. Jørg. (Fig. 2 C)
The species has only been reported once from Nepal (Jørgensen 1978), and is here recorded for the
first time from Russia. Fuscopannaria poeltii belongs to the F. leucosticta complex. Superficially
it looks rather similar to F. leucostica (Tuck.) P.M. Jørg., but is more uniformily brown. The spores
(Fig. 2 F) are distinctly different, being smaller 17‒20 (25) × 7‒9 µm without any apical
enlargement of the exospore. The apparent disjunction between Nepal and Sakhalin is hardly real
since it certainly has been overlooked in the intervening area. While F. leucosticta is a warm-
temperate species, mainly found in the broad-leaved deciduous forests of the lowland hills in SE
Asia (Jørgensen and Sipman 2006), F. poeltii appears to prefer cooler and damper conditions.
Specimens examined. — Sakhalin, Noglikskiy District, Nogliki town surroundings, N 51°49'14,4''
E 142°56'36,17'', alt 83m, dark coniferous forest, on bark of old Betula ermanii Cham., 12.10.2012,
A.K. Ezhkin (SAK 1273).
3. Pannaria insularis P.M. Jørg. & Kash. (Fig. 2 A)
Thallus squamulose-foliose, tightly adnate, 2–3 cm diam., grey-greenish to brown, with numerous
teeth-like secondary lobes with pruinose margins, apothecia rare, scattered on thallus to 0.5 cm with
brown disc and squamulose thalline margin. Photobiont: Nostoc. Hymenium I + blue. Ascospores
simple, colorless, ovate to ellipsoid 10‒14 (17) × 7‒9 µm, with indistinct warty epispore, 8 in ascus.
One unknown trace was detected by HPTLC (fatty acid) which is unique to taxa in the Pannaria
lurida group, most of which contain pannarin, a substance absent from this specimen. P. insularis
is a very rare corticolous lichen and previously known only from southern Japanese Ryuku Islands
(Jørgensen and Kashiwadani 2001a). The species is reported here or the first time from Russia.
Specimens examined. — Sakhalin, Slepikovskogo Cape, N 47°17'26.3'' E 141°59'10.0'', alt 12m,
oak forest, on bark of Quercus crispula Blume, 18.05.2013, A.K. Ezhkin (SAK 133).
4. Pannaria lurida ssp. russellii (Tuck.) P.M. Jorg. (Fig. 2 E)
Pannaria lurida has been reported from the Russian Far East by Tschabanenko (2002), but without
specifying the subspecies or further details. It is unlikely to be ssp. lurida since that is a subtropical,
mainly Pacific species (described from Hawaii). The majority of specimens collected on Sakhalin
appear to belong in the pannarin-deficient, small-spored ssp. russellii which is also present in
Japan, Honshu and Hokkaido (Jørgensen and Kashiwadani 2001a), as well as eastern North
America (Jørgensen 2000b), and accordingly is one of those strongly disjunctive lichen taxa
believed to reflect events in the Tertiary (Culberson 1972). This subspecies is reported here as new
to Russia. It grows in broad leaved and mixed coniferous-broad leaved forests in low elevations on
deciduous trees. It should also be noted that there is a further, as yet not identified, pannarin-

Evansia 35(2): 2018
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Figure 2. Pictures of new lichens for Russia from Sakhalin. A. Pannaria insularis, Bar = 2 mm; B.
Parmeliella miradorensis, Bar = 0,5 mm; C. Fuscopannaria poeltii, Bar = 1 mm; D. F. leprosa,
Bar = 2 mm; E. P. lurida ssp. russellii, Bar = 5 mm; F. Spore of F. poeltii, Bar = 10 µm.
containing taxon in spruce forests in Sakhalin, further adding to the complex taxonomy of this
species (Jørgensen and Kashiwadani 2001a).
Specimens examined. — Kunashir, Dubovoye village surroundins, N 43°47'46.9787'' E
145°30'15.2819'', alt 107m, oak forest, on bark of Quercus crispula Blume, 23.08.2017, A.K. Ezhkin
(SAK 1327, 1328); same place, N 43°46'47.6040'' E 145°29'48.6995'', alt 69m, oak forest, on bark
of Quercus crispula Blume, 23.08.2017, A.K. Ezhkin (SAK 1329, 1330, 1331); Lagunnoye village
surroundins, N44°2'7,044" E145°45'8,524", alt 123m, mixed coniferous-broad leaved forest, on
bark of old Betula ermanii Cham., 16.08.2013, A.K. Ezhkin (SAK 1357); same place, N 44°02'50.2''
E 145°46'01.6'', alt 79m, mixed coniferous-broad leaved forest, on bark of Kalopanax septemlobus
(Thunb. ex A. Murray) Koidz., 14.08.2015, A.K. Ezhkin (SAK 134); Aliger Lake surroundins, N
44°01'45.8328'' E 145°43'12.6732'', alt 79m, mixed coniferous-broad leaved forest, on bark of
Ulmus laciniata Trantv., 20.06.2014, A.K. Ezhkin (SAK 1359); Golovnino Volcano surroundins, N
43°52'20.9208'' E145°36'00.9251'', alt 89m, mixed forest, on bark of Quercus crispula Blume,

Evansia 35(2): 2018
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23.08.2017, A.K. Ezhkin (SAK 1360); Zmeinyi Creek valley, N 44°0'27,66" E 145°40'57,30", alt
39m, broad leaved forest, on bark of Ulmus laciniata Trantv., 31.07.2013, A.K. Ezhkin (SAK 488).
Shikotan, Mt. Notoro surroundins, N 43°46'50.6603'' E 146°42'16.3872'', alt 50m, mixed forest, on
bark of Acer pictum Thunb., 17.06.2017, A.K. Ezhkin (SAK 1332); same place, N 43°46'10.7652''
E 146°41'56.0255'', alt 238m, birch forest, on bark of old Betula ermanii Cham., 14.06.2017, A.K.
Ezhkin (SAK 1351); Mt. Brysova surroundins, N 43°50'54.6'' E 146°51'03.8'', alt 56m, mixed forest,
on bark of Acer pictum Thunb., 19.06.2017, A.K. Ezhkin (SAK 1333); same place, on bark of Salix
caprea L., 19.06.2017, A.K. Ezhkin (SAK 1345); Mt. Shikotan surroundins, N 43°52'15.9024'' E
146°50'42.5472'', alt 157m, mixed forest, on bark of Acer pictum Thunb., 11.06.2017, A.K. Ezhkin
(SAK 1348).
5. Parmeliella miradorensis Vain. (Fig. 2 B)
Thallus repeatedly areolate-sqamulose, tightly adnate, 1‒2 cm diam., with grey-blue to brownish,
thin, leafy, imbricating squamules up to 2 mm wide without a visible blackish prothallus.
Photobiont: Nostoc. Apothecia brown, convex, 0.3‒0.4 mm wide, with a distinct white proper
exciple, clearly visible since the thalline margin is absent, no photobiont in subhymenial areas.
Hymenium I + blue. Ascospores simple, ellipsoid, hyaline, 8 in ascus, (9) 12‒15 x (4) 6‒7 µm. No
lichen substances detected with HPTLC. Parmeliella miradorensis is a corticolous species found
mostly in humid forests of Central and South America, as well as the West Indies, Macaronesian
islands and Japan (Jørgensen 2000b; Jørgensen and Kashiwadani 2001b). P. miradorensis is
reported here for the first time from Russia. Its closest known locality is in Hokkaido, Japan
(Jørgensen and Kashiwadani 2001b) and it is thus showing a disjunct distribution.
Specimens examined. — Kunashir, Mendeleev Volcano surroundings, Lesnaya River valley, N
44°00'04.8'', E 145°42'26.9'', alt 135m, mixed coniferous-broad leaved forest, on bark of Salix
sakhalinensis Sekka, 26.07.2013, A.K. Ezhkin (SAK 131); same place, N 43°59'54,65" E
145°42'25,75", alt 160 m, on bark of Ulmus laciniata Trantv., 04.08.2013, A.K. Ezhkin (SAK 1281).
Iturup, Kurilka River valley, oak forest, N 45°07'49,84'' E 147°47'21,44'', alt 225 m, on bark of
Quercus crispula Blume, 25.08.2012, A.K. Ezhkin (SAK 1282).
Records new to Russia Far East, Sakhalin and Kuriles
6. Fuscopannaria ahlneri (P.M. Jorg.) P.M. Jorg.
This species is characterized by a foliose, rosette-like thallus with a scabrous surface and
hemisphaeric soralia. Fuscopannaria ahlneri is mainly confined to moist, boreal, conifer forests in
N. America, Europe and Asia (the Vladivostok Region and Japan, Jørgensen 1978, 2000b). It was
also recorded in the continental part of Far East of Russia (Primorskiy District) by Tchabanenko
(2002), as well as from the Kamtchatka Territory (Neshatayeva and al. 2003). F. ahlneri is reported
here for the first time from Sakhalin and the Kuriles.
Specimens examined. — Kunashir, Baranskogo Volcano surroundings, near the Kipyashaya River,
N 45°4'43,31" E 147°59'6,968", alt 225 m, mixed forest, on bark of Quercus crispula Blume,
21.08.2015, A.K. Ezhkin (SAK 1278). Sakhalin, Yuzhno-Sakhalinsk city surroundings, Mt. Mitsul,
N 47°02'13,4'' E 142°31'04,9'', alt 712 m, dark conifer forest with Betula ermanii Cham., on bark
of Sorbus commixta Hedl., 26.08.2016, A.K. Ezhkin (SAK 1279); Nevelskiy District, Lovetskiy
pass, N 46°44'35,91" E 142°6'21,02", alt 424 m, mixed forest, on bark of Sorbus commixta Hedl.,
11.06.2013, A.K. Ezhkin (SAK 1280).
7. Fuscopannaria confusa (P.M. Jorg.) P.M. Jørg.
This species possesses a squamulous, deeply incised thallus, often resting on a thin blackish
prothallus, and soralia which are marginal and contain blue-gray, coarse-grained, and often isidioid
soredia. Fuscopannaria confusa is a species of very humid localities in boreal forests, most often
occurring at waterfalls in the places it is known from N. America and Asia (Jørgensen 2000a,

Evansia 35(2): 2018
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2000b). In Russia, it is known from several regions, including the continental part of the Far East
(Urbanavichus 2010). F. confusa is reported here for the first time from Sakhalin and the Kuriles.
Specimens examined. — Sakhalin, Yuzhno-Sakhalinsk city surroundings, Susunaiskiy Range, Mt.
Vorobyinaya, N 46°58'44,24" E 142°48'18,46", alt 511 m, mixed forest, on mossy rock, 12.11.2015,
A.K. Ezhkin (SAK 1277); Mt. Notoro surroundings, N 43°46'40.9296'' E 146°42'08.0495''E, alt 92
m, floodplain forest, on bark of Padus ssiorii (Fr. Schmidt) C.K. Schneid, 17.06.2017, A.K. Ezhkin
(SAK 1274); Tserkovnaya Bay, near the steam, N 43°43'31.1807'' E 146°40'25.4316'', alt 34 m,
mixed forest, on rocks, 13.06.2017, A.K. Ezhkin (SAK 1275).
8. Parmeliella parvula P.M. Jorg.
This species is characterized by its small size and isidioid thallus with discrete soralia but without
terpenoid crystals. Parmeliella parvula is a mostly corticolous species and is widespread in cool-
temperate regions of both hemispheres (Jørgensen 2000b). In Russia, it is known only from the
Caucasus (Urbanavichus 2010). This species is reported here for the first time from the Far East of
Russia.
Specimens examined. — Kunashir, Zmeyinyi Creek valley, N 44°0'25,57" E 145°41'5,686", alt 58
m, broad leaved forest, on bark of Ulmus laciniata Trautv., 31.07.2013, A.K. Ezhkin (SAK 1284).
Shikotan, Tserkovnaya Bay, N 43°44'16.4904'' E 146°40'57.9216'', alt 47 m, mixed forest, on bark
of Salix udensis Trautv., 15.06.2017, A.K. Ezhkin (SAK 1283).
9. Protopannaria pezizoides (Weber) P.M. Jørg. & S. Ekman
This species is characterized by a small-squamulose thallus, often forming compact mats with
prominent apothecia, up to 2 mm in diameter, with a crenulate margin. Protopannaria pezizoides
is a northern, terricolous species widespread in N. America, Europe, Asia, Africa (Jørgensen 1978).
It is also known from the Far Eastern Russia from Sakhalin (Sato 1936a). The species is reported
here for the first time from the Kuriles. It is worth noting that the species in this region prefers to
grow on seaside meadows or in the aerohaline zone of beaches. This unusual habit is previously
known only from the Pacific Northwest coast of North America (Vancouver Island region).
Specimens examined. — Shikotan, the Krai Sveta Cape, N 43°50'28.9140'' E 146°54'35.4996'', alt
59 m, seaside meadow, on soil on old Japanese millitary bunker, 19.06.2017, A.K. Ezhkin (SAK
1268).
10. Santessoniella grisea (Hue) Henssen
Thallus grayish brown, forming a crust over bryophytes and soil, partly subfruticose with 0.1 mm
broad branchlets covered by a single to double cortical layer. Apothecia flat, to 2 mm broad,
brownish with pale proper exciple. Photobiont: Nostoc. Hymenium J+ red-brown. Asci with
indistinct amyloid apical ring-structure. Ascospores simple, colorless, rugulose, ellipsoid (15) 20–
30 (33) x (5) 8–10 (11) µm. All reactions in the thallus negative (colour test); no lichen substances
detected by HPTLC. The species was previously known from the Russian Far East, specifically the
Primorskiy Region (Knyazheva 1973). The species occurs in cool, humid habitats in North America
(Washington) and Japan (Tønsberg and Henssen 1999; Henssen 1997). Santessoniella grisea is
reported here for the first time from Sakhalin and the Kuriles.
Specimens examined. — Sakhalin, Yuzhno-Sakhalinsk city surrounding, Susunaiskiy Range, Mt.
Vorobyinaya, near the Rogatka River, N 46°58'44,24" E 142°48'18,46", alt. 511 m, dark coniferous
forest, on mossy rock, 12.11.2015, A.K. Ezhkin (SAK 490); same place, N 46°58'34.3811'' E
142°48'25.1063'', alt 448 m, 03.09.2016, A.K. Ezhkin (SAK 1261, 1262);
Korsakovskiy District, Tunaicha Lake surroundings, N 46°48'30.3227'' E 143°04'16.1904'', alt 20
m, dark coniferous forest, on soil, 20.09.2017, A.K. Ezhkin (SAK 1263). Kunashir, Mendeleev
Volcano surroundings, Lesnaya River valley, alt 50 m, mixed broad-leaved coniferous forest, on
mossy trunk of old Kalopanax septemlobus (Thunb. Ex. A. Murray) Koidz., 27.08.2017, A.K.
Ezhkin (SAK 67); Tretyakova village surroundings, N 43°58'38.6579'' E 145°39'36.4823'', alt 163

Evansia 35(2): 2018
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m, old mixed broad-leaved coniferous forest, on mossy trunk of old Kalopanax septemlobus
(Thunb. ex. A. Murray) Koidz. and Ulmus laciniata (Trautv.), 15.08.2017, A.K. Ezhkin (SAK 1264,
1265); Lagunnoye Lake surroundings, N 44°02'50.2'' E 145°46'01.6'', alt 79 m, old mixed broad-
leaved coniferous forest, on mossy trunk of Kalopanax septemlobus (Thunb. erx. A. Murray)
Koidz., 22.08.2017, A.K. Ezhkin (SAK 1267).
11. *Vahliella leucophaea (Vahl) P.M. Jørg.
The species is characterized by a squamulose thallus forming a crust-like cover on a thin blue-black
hypothallus. Vahliella leucophaea is a saxicolous species of wet rocks in cold-temperate regions of
the Northern Hemisphere (Jørgensen 1978). It is widespread in Russia including the Far East,
specifically the Primorskiy Region (Urbanavichus 2010; Knyazheva 1978). It was previously
reported for Sakhalin as an epiphyte by mistake (Ezhkin and Galanina 2014); those samples were
redetermined by Jørgensen as F. poeltii. A new finding of V. leucophaea is here confirmed from
one specimen collected from rocks in Sakhalin.
Specimens examined. — Sakhalin, Yuzhno-Sakhalinsk city surroundings, Susunaiskiy Range, Mt.
Vorobyinaya, N 46°58'44,24" E 142°48'18,46", alt 511 m, mixed forest, on rocks, 12.11.2015, A.K.
Ezhkin (SAK 97).
This species has recently been placed in a family of its own, Vahliellaceae (Wedin et al. 2012), but
is still easily mistaken for a member of the Pannariaceae and accordingly included here.
CONCLUSIONS
The marked increase in the number of species known from the Sakhalin Region shows that
the lichen flora has been poorly known, and that in spite of human activities (destruction of primary
forests) it is still possible to find interesting additions to the flora. It is also noteworthy that the
number of known species now exceeds that of the neighbouring region to the south. This is not
surprising since Sakhalin is partly in one plant geographical zone (a southern with broad leaved
deciduous trees dominating) and partly in another (a northern, where coniferous forests are
dominant). This results in a multitude of different habitats. There is one species which cannot easily
be placed in one group or the other, Fuscopannaria poeltii. This lichen represents a Himalayan
element, possibly at its easternmost locality in Sakhalin. The northern group consists of the
following; Fuscopannaria confusa, Pannaria ahlneri, Partmeliella parvula, Santessoniella grisea,
and Protopannaria pezizoides. The first four of these are species confined to moist, boreal
coniferous forests, and represent remnants of the flora present in the large coniferous forests
covering the Northern Hemisphere before the last Ice Age. Today they occur in disjunct areas of
forests in Asia, North America and Europe. The two latter species are more evenly dispersed in the
Northern Hemisphere, Protopannaria pezizoides having a quite different ecology, normally
growing on the ground among mosses or on turf. On the Shikotan Is. it has found a rather special
niche: maritime meadows. The southern group is a rather more mixed one: Fuscopannaria leprosa,
Pannaria insularis, Pannaria lurida ssp. russellii, and Parmeliella miradorensis. They can all be
characterized as warm-temperate, but each with a different origin and distribution. Parmeliella
miradorensis has a wide distribution from Macaronesia, South and Central America to South East
Asia (Jørgensen and Kashiwadani 2001b). The others show more restricted distributions. The most
interesting one is Pannaria insularis, which appears to be an endemic around the Japan Sea,
described from the Riyuku Islands (Jørgensen and Kashiwadani 2001a) and now surprisingly
turning up much further north in the Sakhalin Island. It is a characteristic member of the tropical
Pannaria lurida complex, which has at least one other member in the region: Pannaria lurida ssp.
russelli. This taxon is described from eastern North America and is an example of an
American/Asian disjunction related to geological events in the Tertiary. In addition, the two
Fuscopannaria species are disjunct between North America and Asia, but in a different way. They
are both incompletely amphiberingian in distribution, absent from the coldest part in the North.
They well may have had a more continuous distribution in times when the climate here was milder

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in the northern parts. In contrast, the southern group could remain from one of the warm climatic
periods in the Pleistocene or Holocene when the Sakhalin Island and the Kuriles were connected to
Japan and the Asian continent (Bogatov et al., 2006; Razjigaeva et al. 2013). Most of the species
discovered and reported in this paper are considered rare, occurring mostly in low, disturbed areas
that are difficult to access; because of this, they could be recommended for the Russian Red List.
We conclude that the pannariaceous lichens reflect the varying historical events that the Sakhalin
Island has been part of, as they are mostly remnants of the flora of different, previous geological
periods. It is necessary to intensify the search for lichens in this region to further understand these
distributional patterns.
ACKNOWLEDGEMENTS
We dedicate this paper to Professor Emeritus William A. Weber, Boulder, Colorado on the
occasion of his 100th birthday. We are indebted to the following persons for assistance: Felix
Schumm for performing HPTLC and making micrographs for some specimens; and Alexander
Kordyukov for preparing the map. This study was partly financially supported by the Russian
Foundation for Basic Research (Grants No. 18-04-00098\18).
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