Bull. Natl. Mus. Nat. Sci., Ser. B, 43(1), pp. 1–10, February 22, 2017

Contribution to the Knowledge of the Genus Usnea(Parmeliaceae, Ascomycota) in Southern Far East Russia
Yoshihito Ohmura1,*, Irina Skirina2 and Fedor Skirin2
1

Department of Botany, National Museum of Nature and Science,
Amakubo 4–1–1, Tsukuba, Ibaraki 305–0005, Japan2
Paciﬁc Geographical Institute, Far East Branch of the Russian Academy of Sciences,
Radio street 7, Vladivostok 690041, Russia* E-mail: ohmura-y@kahaku.go.jp(Received 18 November 2016; accepted 21 December 2016)

Abstract Specimens of the lichen genus Usnea from the southern part of Far East Russia were
examined. As the result of our modern taxonomic studies, following 19 species were conﬁrmed:
Usnea cornuta, U. dasaea, U. dasopoga, U. diffracta, U. diplotypus, U. fragilescens, U. fulvoreagens, U. glabrata, U. glabrescens, U. intumescens, U. lapponica, U. longissima, U. pangiana, U.
praetervisa, U. rubicunda, U. rubrotincta, U. subﬂoridana, U. trichodeoides and U. wasmuthii, of
which U. cornuta, U. dasaea, U. intumescens, U. pangiana, U. praetervisa and U. rubrotincta are
new to Russia. Although U. aciculifera, U. articulata, U. barbata, U. bismolliuscula, U. cavernosaand U. mutabilis have been reported from the southern part of Far East Russia, they were notfound in our examined specimens, but they were included here in an artiﬁcial key to the 25 species
recorded for this region.
Key words: Eastern Asia, ﬂora, Jewish Autonomous Region, Khabarovsk, lichen, Primorsky,
Sakhalin, taxonomy.

Introduction
The genus Usnea (Parmeliaceae, Ascomycota)
is characterized by a fruticose thallus with cortex, medulla and a cartilaginous central axis, andthe presence of usnic acid in the cortex. It consists of c. 350 species which are widely distributed throughout the world from polar to tropicalregions (Kirk et al., 2008; Divakar et al., 2015).
In 2015, the ﬁrst author had an opportunity to
examine 96 specimens of the genus collectedfrom the southern part of Far East Russia (Fig. 1)
which were housed in the herbarium of the
Paciﬁc Geographical Institute, Vladivostok.
Although 20 species were listed for this region(Urbanavichus, 2010), they have not beenrevised in the light of modern taxonomic studies
conducted in related regions (e.g. Ohmura andKashiwadani, 2000; Ohmura et al., 2000, 2010;

Ohmura, 2001, 2002, 2008, 2011, 2012, 2014;
Ohmura and Kanda, 2004; Ohmura andOnimaru, 2010; Clerc, 2011). Therefore, the purpose of this study is to contribute to our knowledge of the genus Usnea in this region based on
herbarium material.
Materials and Methods
This study is based on a total of 110 specimens, 96 of which were mainly collected by thesecond author between 1959 and 2014 andhoused in the Paciﬁc Geographical Institute, Far
East Branch of the Russian Academy of Sciences(abbreviated as PGI in the present paper) and 14
specimens housed in the National Museum of
Nature and Science, Tsukuba (TNS).
Morphological observations were made using
dissecting and light microscopes. Ratio measure-Yoshihito Ohmura et al.

2

Fig. 1.

Map showing the area of southern Far East Russia (black-painted). Modiﬁed after Urbanavichus (2010).

ments of the thicknesses of the cortex, medulla
and axis of branches according to the method of
Clerc (1984), and lichen substances were determined by thin layer chromatography (TLC)
according to Culberson and Johnson (1982). Solvent B system (hexane : methyl tert.-butyl ether :formic acid, 140 : 72 : 18) was used for all TLC
analyses. Morphological terminology follows
that of Ohmura (2001).

The 19 species listed below were conﬁrmed by
the results of a taxonomic study based on 110
examined specimens.

ictic acids; and (3) usnic acid.
New to Russia.
Specimens examined. RUSSIA. Primorsky
Krai: Terneiskiy District, upper part of Bolʼshaya
Ussurka River, north side of the valley, ﬁr-spruce
forest, on Abies nephrolepis, 1981, I. F. Skirina
19147 (PGI); Chuguevskiy District, Golets
mountain, ﬁr-spruce forest, (1300 m elev.), on
Abies nephrolepis, Betula lanata, and Picea ajanensis, 10 July 1991, I. F. Skirina 13464 (PGI);
Kedrovaya Pad Nature Reserve, right side of
Kedrovaya River, near the upper part of watershed, 652 m elev., black ﬁr forest, on Tilia amurensis, 11 July 1999, collector unknown 5455
(PGI).

1. U. cornuta Körb.
For a detailed description and synonyms, see
Clerc (2011).
Chemistry. Three chemotypes were detected in
the examined specimens: (1) usnic, norstictic,
protocetraric (trace), and salazinic acids; (2)usnic, norstictic, menegazziaic, stictic, and const-

2. U. dasaea Stirt.
For detailed descriptions and synonyms, see
Clerc (2011) and Ohmura (2001).
Chemistry. All examined specimens containusnic, norstictic, protocetraric (trace) and salazinic acids.
New to Russia.

Results and Discussion

Usnea in Southern Far East Russia

Specimens examined. RUSSIA. Primorsky
Krai: Terneiskiy District, Krivoy creek, pine forest, on Pinus koraiensis, 30 June 1977, I. F. Skirina 14649 (PGI); Chuguevskiy District, Upper
Ussurian Stationary, test area no. 12, pine forest,
on Pinus koraiensis, 11 Aug. 1976, V. A. Eﬁmova& I. F. Skirina 14672 (PGI); Dalnegorskiy District, Medvezhʼya mountain, N slope, ﬁr-spruce
forest, on deadwood of Picea ajanensis, 28 July
1982, I. F. Skirina 14667 (PGI); Dalnegorskiy
District, upper part of Krivaya River, ﬁr-spruce
forest, on Pinus koraiensis, 1 Aug. 1992, I. F.
Skirina 9707 (PGI); Krasnoarmeiskiy District,
Serokamenka River, ﬁr-spruce forest, on deadwood, 1980, I. F. Skirina 14808 (PGI); Lazovskiy District, Lazovskiy Pass, Suchanskiy creek,
N side of valley, ﬁr-spruce forest, on Abies
nephrolepis, Picea ajanensis, Betula lanata andAcer barbinerva, April 1985, I. F. Skirina 19139(PGI); Michailovskiy District, Ivanovskiy forestry, ﬁr-spruce forest, on Abies nephrolepis andPicea ajanensis, 1994, I. F. Skirina 30495 (PGI);
Terneyskiy District, Zacharovskiy creek, ﬁrspruce forest, on Abies nephrolepis, 10 Oct.
1976, I. F. Skirina 14913 (PGI); Terneiskiy District, Sichote-Alinskiy Nature Reserve, plot of
land Blagodatnoe, oak forest, on Quercus mongolica, 13 July 1977, I. F. Skirina 14241 (PGI);
Shkotovskiy District, near Shkotovo village,pine-broadleaved forest, on Pinus koraiensis,
1976, collector unknown (PGI); Sichote-Alinskiy
Nature Reserve, Krivoy creek, pine forest, on
Pinus koraiensis, 30 June 1977, I. F. Skirina
14630 (PGI); Sichote-Alinskiy Nature Reserve,
Tayozjnaya River, ﬁr-spruce forest, on Picea ajanensis and Abies nephrolepis, 21 July 1980, I. F.
Skirina 28772 (PGI). Jewish Autonomous
Region: Bastak Nature Reserve, Kirga River valley, near the Polkovnikova apiary, ﬁr-spruce forest, on Pinus koraiensis, 7 Aug. 2002, I. F. Skirina & V. I. Skirin 14635 (PGI); Bastak Nature
Reserve, Kirga River valley, near the
Polkovnikova apiary, ﬁr-spruce forest, on Picea
ajanensis, 7 Aug. 2002, I. F. Skirina & V. I. Skirin 18946 (PGI); Bastak Nature Reserve, quarter
140, hill near bee-yard, oak forest, tree no. 3, on

3

Quercus mongolica, 22 Sept. 2004, I. F. Skirina
16542 (PGI); Bastak Nature Reserve, valley of
Kirga River, pine-broadleaved forest, on Abiessp., 2003, I. F. Skirina 34211 (PGI); Bastak
Nature Reserve, right conﬂuence of Bolshoy
Sorennak River, near road, swamp with larch, on
Rhododendron davurica, 13 Aug. 2002, I. F. Skirina & V. I. Skirin 14640 (PGI).
3. U. dasopoga (Ach.) Nyl.
For a detailed description and synonyms, see
Clerc (2011).
Chemistry. All examined specimens containusnic and salazinic acids.
Specimens examined. RUSSIA. Khabarovsk
Krai: Miao-Chan ridge, 10 km N of Gorniy village, left side of Amur River, ﬁr forest, on Picea
sp. 1983, P. V. Elpatyevskiy s.n. (PGI). Sakhalin
Oblast: Timovskiy District, Pilinga River valley,ﬁr-spruce forest, on Abies sachalinensis, 17 July
2014, I. F. Skirina s.n. (PGI, TNS).
4. U. diffracta Vain.
For a detailed description and synonyms, see
Ohmura (2001).
Chemistry. All specimens examined containusnic, barbatic, diffractaic, baeomycesic (trace)and squamatic acids.
Specimens examined. RUSSIA. Sakhalin
Oblast: Mt. Suzuya, 19 July 1932, Y. Asahina s.n.
(TNS); Toyohara, 12 Aug. 1920, A. Yasuda s.n.
(TNS); Sakaehama, 21 July 1932, Y. Asahina s.n.
(TNS); Takinosawa, 31 July 1932, Y. Asahina
s.n. (TNS); Mt. Tosso, 24 July 1932, M. Sato s.n.
(TNS); Tinnai, 14 Aug. 1933, M. Sato s.n.
(TNS); Toyohara, 12 Aug. 1920, M. Sato s.n.
(TNS).
5. U. diplotypus Vain.
For a detailed description, see Halonen et al.
(1999).
Chemistry. All specimens examined containusnic and salazinic acids.
Specimens examined. RUSSIA. Primorsky
Krai: Partizanskiy District, Olhovaya mountain,
540 m elev., ﬁr-spruce forest, on Picea ajanensis

4

Yoshihito Ohmura et al.

and Abies nephrolepis, 2010, I. F. Skirina 29423(PGI). Sakhalin Oblast: Timovskiy District, Pilinga River valley, ﬁr-spruce forest, on Abiessachalinensis, 18 July 2014, I. F. Skirina s.n.
(PGI).
6. U. fragilescens Hav. ex Lynge
For a detailed description and synonyms, see
Ohmura (2001).
Chemistry. Two chemotypes were detected in
the examined specimens: (1) usnic, norstictic,
salazinic and ± protocetraric (trace) acids; and(2) usnic, norstictic and stictic (trace) acids.
Specimens examined. RUSSIA. Primorsky
Krai: Chuguevskiy District, valley of Ussury
River, Berezoviy Pass, Berezoviy creek, ﬁrspruce forest, on Abies nephrolepis and Picea
ajanensis, 20 Dec. 2001, F. V. Skirin s.n. (PGI);
Dalnegorskiy District, headwater of Krivaya
River, mixed pine-broadleaved forest, “Chortovi
vorota” (Hellʼs gate), on Pinus koraiensis, 6 July
1976, I. F. Skirina14563 (PGI); Lazovskiy District, Elamovskiy creek, pine-broadleaved forest,
on Abies nephrolepis, 20 July 2000, I. F. Skirina
14964 (PGI).
7. U. fulvoreagens (Räsänen) Räsänen
For a detailed description, see Ohmura (2001).
Chemistry. Two chemotypes were detected in
the examined specimens: (1) usnic, diffractaic,
norstictic and squamatic (trace) acids, and zeorin(trace); and (2) usnic and barbatic (trace) acids,
and zeorin (trace).
Although Clerc (2011) treated this taxon under
the synonym of U. glabrescens, U. fulvoreagens
has deeply excavated soralia (as seen in U. lapponica) and constantly contains zeorin. In contrast, U. glabrescens has discrete soralia withslightly excavated surface and lacking zeorin.
Therefore, U. fulvoreagens is treated as a distinct
taxon in the current paper.
Specimens examined. RUSSIA. Primorsky
Krai: Krasnoarmeiskiy District, Dalʼniaya River,
Porojistiy creek, ﬁr-spruce forest, on Abies
nephrolepis, 8 Aug. 2014, I. F. Skirina & F. V.
Skirin s.n. (PGI). Sakhalin Oblast: Timovskiy

District, Pilinga River valley, ﬁr-spruce forest, on
Abies nephrolepis, 18 July 2014, I. F. Skirina s.n.
(PGI).
8. U. glabrata (Ach.) Vain.
For detailed descriptions, see Ohmura (2001)
and Clerc (2011).
Chemistry. Two chemotypes were detected in
the examined specimens: (1) usnic and salazinic
acids; and (2) usnic, barbatic, 4-O-demethylbarbatic (trace), ± norstictic and ± protocetraric(trace) acids.
Specimens examined. RUSSIA. Primorsky
Krai: Dalnegorskiy District, Eldorado mountain(1300 m elev.), ﬁr-spruce forest, on deadwood of
Picea ajanensis, 1982, I. F. Skirina 14323 (PGI);
Terneiskiy District, Dalʼniy mountain ridge, ﬁrspruce forest, on Abies nephrolepis, 22 June
1977, I. F. Skirina 14314 (PGI); Terneiskiy District, Solontsoviy creek, pine forest, on Abies
nephrolepis, 7 July 1977, I. F. Skirina 14665(PGI); Terneiskiy District, estuary of Serebrianka
River, Chosenia forest, on Chosenia arbutifolia,
11 July 1977, I. F. Skirina 28605 (PGI); Terneiskiy District, Sichote-Alinskiy Nature Reserve,
Tayozhnaya River, ﬁr-spruce forest, on Abies
nephrolepis and Picea ajanensis, 21 July 1980, I.
F. Skirina 28773 (PGI); Partizanskiy District,near Olhovaya mountain, Vetvistiy creek, 700 m
elev., ﬁr-spruce forest, on Picea ajanensis andBetula lanata, 19 Aug. 2006, I. F. Skirina 29587(PGI); Partizanskiy District, 25 km N of Sergeevka village, Sergeevskoe (Suchano-Malazskoe) plateau, 875 m elev., swamp with larch, on
Larix cajanderi, 14 July 2002, E. P. Kudryavceva
14354 (PGI); Partizanskiy District, spur of Skalistaya mountain, 1000 m elev., pine-broadleaved
forest, area no. 10, tree no. 1 (N43°07′82.2″,
E132°59′18.9″), on Pinus koraiensis, 25 Aug.
2012, I. F. Skirina & F. V. Skirin 31979 (PGI).
9. U. glabrescens (Nyl. ex Vain.) Vain.
For descriptions and synonyms, see Halonen
et al. (1999) and Ohmura (2001).
Chemistry. Usnic and norstictic acids weredetected in the single specimen examined.

Usnea in Southern Far East Russia

5Fig. 2. Usnea intumescens (Demenshina 14995, fragment in TNS) showing punctiform soralia and the broadenedbase of branches. Scale ＝ 0.5 mm.

See also U. fulvoreagens above for the differences.
Specimen examined. RUSSIA. Sakhalin
Oblast: Timovskiy District, Pilinga River valley,ﬁr-spruce forest, on Abies nephrolepis, 20 July
2014, I. F. Skirina s.n. (PGI).
10. U. intumescens Asahina
For a description and synonyms, see Ohmura(2001).
Chemistry. Usnic, psoromic and 2′-O-demethylprosormic acids were detected.
New to Russia. This species was previously
known only from central Japan (Ohmura, 2001).
However, the features of the Russian material[i.e. broadened base of branches, punctiform
soralia (Fig. 2), and the presence of psoromic orsalazinic acid as the major substance] coincide
well with those seen in U. intumescens (seeOhmura, 2001).
Specimen examined. RUSSIA. Primorsky
Krai: Khasanskiy District, Posʼet Bay, Sosnovaya
Cove, pine forest, on Pinus densiﬂora, 5 Aug.
1980, Demenshina 14995 (PGI).
11. U. lapponica Vain.
For descriptions and synonyms, see Clerc(2011) and Mark et al. (2016).
Chemistry. All specimens examined containusnic and salazinic acids; a trace amount of zeorin was also detected in one specimen.
Usnea lapponica is morphologically difﬁcult
to distinguish from U. fulvoreagens.
Specimens examined. RUSSIA. Primorsky
Krai: Partizanskiy District, Olhovaya mountain,
1500-1600 m elev. and Olchoviy creek (540 m
elev.), ﬁr-spruce forest, on Abies nephrolepis,
Picea ajanensis and Betula costata, 27 July
2007, I. F. Skirina & F. V. Skirin 29636 (PGI);
Partizanskiy District, Postishevka River valley,
353 m elev., pine-broadleaved forest, area no. 2,
tree no. 1 (N43°09′53.0″, E132°59′28.9″), on
Salix caprea, 24 Aug. 2012, I. F. Skirina & F. V.
Skirin 31976 (PGI); Partizanskiy District, Olhovaya mountain, 540 m elev., ﬁr-spruce forest, on
Picea ajanensis and Abies nephrolepis, 2 July
2007, I. F. Skirina & F. V. Skirin 29643 (PGI);
Partizanskiy District, Olhovaya mountain, fallen
ﬁr, on Abies nephrolepis, 4 July 2007, I. F. Skirina & F. V. Skirin 31121 (PGI). Sakhalin Oblast:
Timovskiy District, Pilinga River valley, ﬁrspruce forest, on Abies sachalinensis, 18 July
2014, I. F. Skirina, s.n. (PGI, TNS); Timovskiy
District, Timovsk city, on Populus sp., 18 July

6

Yoshihito Ohmura et al.

2014, O. E. Chrapko s.n. (PGI); Timovskiy District, Pilinga River valley, ﬁr-spruce forest, on
Abies sachalinensis, 18 July 2014, I. F. Skirina
s.n. (PGI).
12. U. longissima Ach.
For a description and synonyms, see Ohmura(2001).
Chemistry. Two chemotypes were detected in
the examined specimens: (1) usnic, barbatic(trace), diffractaic, baeomycesic and squamatic
acids; and (2) usnic and evernic acids. Ohmura(2001) also reported a further chemotype (usnic
and barbatic acids) found in northern Hokkaido
of Japan, and further investigation would no
doubt ﬁnd the chemotype in Far East Russia.
Specimens examined. Sakhalin Oblast:
Timovskiy district, Pilinga River valley, ﬁrspruce forest, on Abies sachalinensis, 20 July
2014, I. F. Skirina s.n. (PGI); Takinosawa. 30
July 1932, Y. Asahina s.n. (TNS); ibid., 31 July
1932, Y. Asahina 2503 (TNS); Sakaehama, 21
July 1932, Y. Asahina s.n. (TNS); loc. non cit.,
1923, O. Kikuchi s.n. (herb. Y. Asahina 2506)(TNS); Aikawa (Picecharomai), 28 Feb. 1930, O.
Kikuchi s.n. (TNS).
13. U. pangiana Stirt.
For a description and synonyms, see Ohmura(2001).
Chemistry. Two chemotypes were detected in
the examined specimens: (1) usnic and salazinic
acids; and (2) usnic acid. Although barbatic acidis also commonly detected in eastern Asian material (Ohmura, 2001), it was not detected in theexamined specimens in this study.
New to Russia.
Specimens examined. RUSSIA. Primorsky
Krai: Ussuriyskiy Reserve, on slope of watershedof Mironov creek, pine-broadleaved forest, on
Betula platyphylla, 9 June 1974, E. Semenova
32888 (PGI); Khasanskiy District, Kedrovaya
Pad Reserve, right side of Kedrovaya river,
652 m elev., black ﬁr-spruce forest, on Tilia amurensis, 11 July 1999, I. F. Skirina 18651 (PGI);
Khasanskiy District, Kedrovaya Pad Nature

Reserve, Kedrovaya River, right side, 562 m
elev., near the watershed, black ﬁr forest, on Tilia
amurensis, 11 July 1999, I. F. Skirina 19120(PGI); Khasanskiy District, Riazanovka railway
station, near FEFU Biological Station, on hillslope, oak forest, on Tilia amurensis, 18 July
1984, E. Shparko 14312 (PGI).
14. U. praetervisa (Asahina) P. Clerc
For a detail description, see Clerc (2004).
Chemistry. All examined specimens containusnic and norstictic acids.
New to Russia.
Specimens examined. RUSSIA. Primorsky
Krai: Sichote-Alinskiy Nature Reserve, Krivoy
creek, east side of the valley, larch forest, scree,
on Rhododendron sp., 1 July 1977, I. F. Skirina
s.n. (PGI); Sichote-Alinskiy Nature Reserve,
Listvenichniy island, estuary of Serebrianka
River, birch forest, on Betula platyphylla, 14
June 1985, I. F. Skirina 5419 (PGI). Sakhalin
Oblast: Timovskiy District, Pilinga River valley,ﬁr-spruce forest, on Abies sachalinensis, 18 July
2014, I. F. Skirina s.n. (PGI). Jewish Autonomous Region: Bastak Nature Reserve, mixedpine-broadleaved forest, on Tilia amurensis, 14Aug. 2002, I. F. Skirina & V. I. Skirin 14637(PGI).
15. U. rubicunda Stirt.
For a detail description and synonyms, see
Ohmura (2001).
Chemistry. Usnic, norstictic, menegazziaic,
stictic and constictic acids were detected.
Specimen examined. RUSSIA. Primorsky
Krai: Muraviev-Amurskiy Peninsula, Bogataya
griva ridge, on Fraxinus rinchophilla, 14 Sept.
1959, Vokueva 28901 (PGI).
16. U. rubrotincta Stirt.
For a detail description and synonyms, see
Ohmura (2001).
Chemistry. All specimens examined containusnic, norstictic, protocetraric (trace) and salazinic acids.
New to Russia.

Usnea in Southern Far East Russia

Specimens examined. RUSSIA. Primorsky
Krai: Shkotovskiy District, near Novaya Moskva
village, the upper reaches of the Tigroviy creek,ﬁr-spruce forest, on Abies nephrolepis, Taxus
kuspidata, Picea ajanensis and Acer sp., 1 Sept.
2002, I. F. Skirina 14865 (PGI); Shkotovskiy
District, Livadiyskaya mountain, Pryamoi creek,ﬁr-spruce forest, on Abies nephrolepis, 12 June
2003, I. F. Skirina 30257 (PGI); Shkotovskiy
District, Livadiyskaya mountain, Pryamoi creek,ﬁr-spruce forest, on Picea ajanensis, 12 June
2003, I. F. Skirina 34207 (PGI); Kedrovaya Pad
Nature Reserve, near small hut, on Pinus
koraiensis, 13 Sept. 1959, Vasilʼeva 34229 (PGI);
Khasanskiy District, Gamova Peninsula, nearMarine Experimental Station, oak forest with
Rhododendron, on Quercus mongolica, 24 May
1988, A. Mikulin 10687 (PGI); Khasanskiy District, Kedrovaya Pad Nature Reserve, near small
hut, pine forest, on deadwood of Pinus koraiensis, 16 Aug. 1975, I. F. Skirina & L. A. Knyazheva
18306 (PGI).
17. U. subﬂoridana Stirt.
For a detail description and synonyms, see
Ohmura (2001).
Chemistry. Two chemotypes were detected in
the examined specimens: (1) usnic and squamatic
acids; and (2) usnic and thamnolic acids.
Specimens examined. RUSSIA. Khabarovsk
Krai: Lazo District, Nelta River valley, rightinﬂow of Muchen River, upper and middle ﬂowing, 400-500 m elev., ﬁr-spruce forest, on Abies
nephrolepis, 29 March 2003, I. F. Skirina s.n.
(PGI); Lazo district, Nelta River valley, rightinﬂow of Muchen River, upper and middle ﬂowing, 400-500 m elev., ﬁr-spruce forest, on Abies
nephrolepis, 29 March 2003, I. F. Skirina s.n.
(PGI). Primorsky Krai: Ussuri Nature Reserve,pine-broadleaved forest, on fallen Pinus koraiensis, 7 Aug. 1974, E. Semenova s.n. (PGI); Partizanskiy District, Olhovaya mountain, ﬁr-spruce
forest, on Picea ajanensis, 4 July 2007, I. F. Skirina & F. V. Skirin 31126 (PGI); Partizanskiy
District, Postishevka River valley, 353 m elev.,pine-broadleaved forest, area no. 2, tree no. 1

7(N43°09′53.0″, E132°59′28.9″), on Salix caprea,
24 Aug. 2012, I. F. Skirina & F. V. Skirin 31977(PGI); Terneyskiy District, Djigitovka River,
Zolotaya polyana village, north slope, oak forest,on Quercus mongolica, 1982, I. F. Skirina 14809(PGI); Terneyskiy District, Sichote-Alinskiy
Nature Reserve, Blagodatnoe, Sukhoy creek,pine-broadleaved forest, on stones, 1982, I. F.
Skirina s.n. (PGI); Terneyskiy District, SichoteAlinskiy Nature Reserve, Blagodatnoe, oak forest with larch, on Larix cajanderi, 15 June 1985,
I. F. Skirina 5417 (PGI); Terneyskiy District,near Terney village, watershed, oak forest withlarch, on Larix cajanderi, 15 June 1985, I. F. Skirina 5437 (PGI); Partizanskiy District, Olhovaya
mountain, on fallen Picea ajanensis, 4 July 2007,
I. F. Skirina & F. V. Skirin 31120 (PGI); Dalnegorskiy District, Rudnaya River valley, upper
part of Novitskogo, watershed, on Rhododendron
davurica, Betula davurica and B. platyphylla,
1981, I. F. Skirina 10903 (PGI). Sakhalin Oblast:
Timovskiy District, Pilinga River valley, ﬁrspruce forest, on Abies sachalinensis, 18 July
2014, I. F. Skirina s.n. (PGI, TNS); 10 km N of
Buyukhli village, larch forest, on Larix cajanderi, 11 July 2014, I. F. Skirina s.n. (PGI);
Timovskiy District, near Uskovo village, larch
forest, on Larix cajanderi, 17 July 2014, I. F. Skirina s.n. (PGI); Timovskiy District, Pilinga Rivervalley, ﬁr-spruce forest, on Abies sachalinensis,
18 July 2014, I. F. Skirina s.n. (PGI).
18. U. trichodeoides Vain.
For a detail description and synonyms, see
Ohmura (2001).
Chemistry. Atranorin, and fumarprotocetraricand usnic (trace) acids were detected. Ohmura(2001) also reported another chemotype (usnic
and salazinic acids); since it was found in northern Hokkaido of Japan, further investigation no
doubt detect the chemotype in Far East Russia.
Specimen examined. RUSSIA. Primorsky
Krai: Khasanskiy District, Kedrovaya Pad
Reserve, test area no. 6, behind small hut, pine
forest, on Pinus koraiensis, 16 Aug. 1980, L. A.
Kniazsheva 14787 (PGI).

8

Yoshihito Ohmura et al.

19. U. wasmuthii Räsänen
For a detail description and synonyms, see
Ohmura (2001).
Chemistry. All specimens examined containusnic and salazinic acids.
Specimens examined. RUSSIA. Primorsky
Krai: Partizanskiy District, Olhovaya mountain,
Olchoviy creek, 540 m elev., ﬁr-spruce forest, on
Picea ajanensis and Abies nephrolepis, 2 July
2007, I. F. Skirina 29615 (PGI); Partizanskiy
District, Olhovaya mountain, 540 m elev., ﬁrspruce forest, on Picea ajanensis and Abies

nephrolepis, 2 July 2007, I. F. Skirina 29404(PGI); Partizanskiy District, Skalistaya mountain, watershed, 430–475 m elev., pine-broadleaved forest, on Picea ajanensis and Abies
nephrolepis, 24 Aug. 2012, I. F. Skirina & F. V.
Skirin 31978 (PGI); Terneiskiy District, Zakharovskiy creek, ﬁr-spruce forest, on Abies nephrolepis, 10 Oct. 1976, I. F. Skirina 15014 (PGI);
Terneiskiy District, Zakharovskiy creek, ﬁrspruce forest, on Abies nephrolepis, 10 Oct.
1976, I. F. Skirina 15021 (PGI).

Key to Usnea taxa in southern Far East Russia
The taxa in squared parentheses [ ] have been reported from southern Far East Russia, but notfound in our examined specimens. Therefore this key includes such species possibly occurring in thisregion.
1. Red pigment present in the cortex ................................................................................................. 2
1. Red pigment absent in the cortex ................................................................................................... 3
2. Terminal and subterminal branches elongated; sorediate ﬁbrils abundant on branches; soralia
present both on ﬁbrils and thick branches; salazinic acid present (or very rarely stictic acid could
be present) .................................................................................................... U. rubrotincta Stirt.
2. Terminal and subterminal branches not elongated; ﬁbrils sparse and sorediate ﬁbrils very rare on
branches; soralia present mainly on thick branches; stictic acid present ........ U. rubicunda Stirt.
3. Cortex fragile on main branches, decorticated or areolate-corticated; thallus “ﬁsh-bone” like inappearance with numerous perpendicular ﬁbrils on elongated branches ..................................... 4
3. Cortex stable, consistent on the branches; thallus not as above ................................................... 5
4. Branches terete, without longitudinal-furrows on the surface; diffractaic, barbatic (or possibly
evernic) acids present .................................................................................... U. longissima Ach.
4. Branches ﬂattened in well-developed thallus with longitudinal-furrows on the surface; fumarprotocetraric or salazinic acid present ....................................................... U. trichodeoides Motyka
5. Annular-pseudocyphellae distinct on the segments of branches (bamboo-like appearance); thallus isotomic-dichotomous branching ................................................................. U. diffracta Vain.
5. Annular-pseudocyphellae absent; thallus anisotomic- or isotomic-dichotomous branching ........ 6
6. Medulla wine-red ........................................................................................... [U. mutabilis Stirt.]
6. Medulla white ................................................................................................................................ 7
7. Branches foveolate and/or ridged ................................................................................................. 8
7. Branches not foveolate nor ridged ................................................................................................ 9
8. Soralia absent ............................................................................................... [U. cavernosa Tuck.]
8. Soralia present, punctiform, ± plane to slightly stipitate ............... [U. barbata (L.) F.H. Wigg.]
9. Main branches inﬂated and segmented (sausage-like) ........................ [U. articulata (L.) Hoffm.]
9. Branches do not form sausage-like segments ............................................................................. 10
10. Thallus with jet black base ......................................................................................................... 11
10. Thallus with concolorous to dark brown base ............................................................................ 19
11. Soralia convex ............................................................................................................................. 12

Usnea in Southern Far East Russia

9

11.
12.
12.
13.
13.
14.
14.
15.
15.
16.
16.
17.
17.
18.
18.
19.
19.
20.
20.
21.
21.
22.
22.
23.
23.
24.

Soralia concave ........................................................................................................................... 15
Squamatic or thamnolic acid present .......................................................... U. subﬂoridana Stirt.
Salazinic or norstictic acid present ............................................................................................. 13
Norstictic acid present .............................................................. U. praetervisa (Asahina) P. Clerc
Salazinic acid present .................................................................................................................. 14
Thallus usually pendulous; terminal branches elongated ....................... U. dasopoga (Ach.) Nyl.
Thallus shrubby; apical parts often sinuous ................................................... U. diplotypus Vain.
Soralia deeply excavated nearly into central axis ....................................................................... 16
Soralia slightly excavated and never reaching to central axis .................................................... 17
Salazinic acid present; norstictic acid absent ................................................... U. lapponica Vain.
Norstictic acid present; salazinic acid absent ....................... U. fulvoreagens (Räsänen) Räsänen
Branches glossy on the surface, constricted at the ramiﬁcation points .......... U. fragilescens Hav.
Branches mat on the surface, not constricted at the ramiﬁcation points ..................................... 18
Soralia discrete, rounded; norstictic acid present ................ U. glabrescens (Nyl. ex Vain.) Vain.
Soralia conﬂuent, elliptic; norstictic acid absent ....................................... U. wasmuthii Räsänen
Branches inﬂated, constricted at the ramiﬁcation points ............................................................ 20
Branches not inﬂated, not constricted at the ramiﬁcation points ................................................ 25
Soralia concave, cortical margin reﬂexed ............................................... U. glabrata (Ach.) Vain.
Soralia convex to ﬂat .................................................................................................................. 21
Stictic acid present ...................................................................................................................... 22
Stictic acid absent ....................................................................................................................... 23
Perforations present on thicker branches ........................................... [U. bismolliuscula Zahlbr.]
Perforations absent on thicker branches ............................................................ U. cornuta Körb.
Lateral branches broadened at the base (Fig. 2) ..................................... U. intumescens Asahina
Lateral branches constricted at the base ...................................................................................... 24
Main branches inﬂated and segmented as sausage-like; soralia (usually cited as pseudocyphellae)punctiform and sparse; galbinic acid absent, fumarprotocetraric acid present or absent ................
............................................................................................................. [U. articulata (L.) Hoffm.]
24. Branches do not form sausage-like segments; soralia irregular in shape which are developed
from scars of detached ﬁbrils; galbinic acid present or absent, fumarprotocetraric acid absent .....
............................................................................................................................... U. dasaea Stirt.
25. Base of thallus continuous or irregularly cracked; stictic acid present ........ [U. aciculifera Vain.]
25. Base of thallus distinctly annularly cracked; salazinic acid present ................. U. pangiana Stirt.

Acknowledgements
We are grateful to Prof. Mark Seaward (Bradford University, UK) for help in the preparationof this manuscript, and to an anonymous reviewer
for a critical appraisal of this manuscript. This
study was partly supported by JSPS KAKENHI(Grant no. 24300314) for the ﬁrst author.

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