Arctoa (2021) 30: 79–90
doi: 10.15298/arctoa.30.09
CONTRIBUTION TO THE MOSS FLORA OF AYAN
(WEST COAST OF THE SEA OF OKHOTSK, THE RUSSIAN FAR EAST)
К ФЛОРЕ МХОВ ОКРЕСТНОСТЕЙ ПОСЕЛКА АЯН
(ЗАПАДНОЕ ПОБЕРЕЖЬЕ ОХОТСКОГО МОРЯ, РОССИЙСКИЙ ДАЛЬНИЙ ВОСТОК)
ELENA A. IGNATOVA1, MICHAEL S. IGNATOV1,2, KSENIA G. KLIMOVA3* & VADIM A. BAKALIN3
ЕЛЕНА А. ИГНАТОВА1, МИХАИЛ С. ИГНАТОВ1,2, КСЕНИЯ Г. КЛИМОВА3*, ВАДИМ А. БАКАЛИН3
Abstract
The exploration of the area near Ayan Settlement and adjacent part of Dzhugdzhur Range on the
Western coast of the Sea of Okhotsk revealed 164 species and 1 variety of mosses; their annotated list
is provided. The cold and humid climate restricts the occurrence of the East Asian species that are
more numerous in a few hundred kilometers to the south, and xerophytic plants that are characteristic
to continental areas, at less than 300 km from the seacoast. Species with the North Pacific distribution
are well represented in the area: Coscinodon yukonensis, Trachycystis ussuriensis, Myuroclada maximowiczii, Echinophyllum sachalinense, Sciuro-hypnum uncinifolium, Bartramiopsis lescurii, Oligotrichum falcatum, O. parallelum, and Rhizomnium nudum. The frequent occurrence of Aquilonium
plicatulum, Brachythecium baicalense, Echinophyllum sachalinense, and Dicranum majus, and finding of Oedipodium griffithianum are other features of oceanic environment here.
Резюме
По итогам изучения флоры мохообразных в окрестностях поселка Аян и прилежащей части
хребта Джугджур на западном побережье Охотского моря выявлено 164 вида и 1 разновидность
мхов; приводится аннотированный список. Распространение южных восточноазиатских видов,
более многочисленных в нескольких сотнях километров к югу, а также ксерофитных видов,
характерных для континентальных районов, расположенных не менее чем в 300 км от морского
побережья, ограничено холодным и влажным климатом района исследований. Хорошо представлены виды, распространенные в северной части побережья Тихого океана: Coscinodon
yukonensis, Trachycystis ussuriensis, Myuroclada maximowiczii, Echinophyllum sachalinense, Sciurohypnum uncinifolium, Bartramiopsis lescurii, Oligotrichum falcatum, O. parallelum и Rhizomnium
nudum. Океаничность климата подчеркивается частой встречаемостью Aquilonium plicatulum,
Brachythecium baicalense, Echinophyllum sachalinense, Dicranum majus, а также обнаружением
Oedipodium griffithianum.
KEYWORDS: biodiversity, phytogeography, bryophytes, Khabarovsk Territory, Dzhugdzhur Range
INTRODUCTION
The flora of the Russian Far East was in a focus of
extensive studies in the recent decades, and numerous
discoveries in this area continue (Cherdansteva et al.,
2018). At the same time, the coverage of exploration remains fairly uneven. Pisarenko & Bakalin (2017) summarized all published materials for the regional moss
diversity of the Russian Far East by 39 spatial units of
5×5 degrees latitude/longitude. Most of such units include records for more than 200 moss species, with maximal diversity of 416 species in the Primorsky Territory.
1
Eight regions are least known, with less than 100 moss
species reported, and the region of the present study on
the western coast of the Sea of Okhotsk is one of these:
the total number of known species is 52 cf. Pisarenko &
Bakalin (2017). This number is based only on Lazarenko (1940, 1941a, b, 1945) catalogue that refers to the
unpublished data of Victor N. Vassiliev, who studied vegetation in Ayan region in 1936. These records include
mostly common species of the main type of vegetation,
and lack exact locality data.
– Lomonosov Moscow State University, Faculty of Biology, Plant Ecology and Geography Dept., Leninskie Gory Str. 1–12,
Moscow 119234 Russia; ORCID (EA): 0000-0001-6287-5660, (MI): 0000-0001-6096-6315
2 – Tsitsin Main Botanical Garden, Russian Academy of Sciences, Botanicheskaya Str., 4, Moscow 127276 Russia
3 – Botanical Garden-Institute, Far Eastern Branch of the Russian Academy of Sciences, Makovskogo Street, 142, Vladivostok,
690024, Russia ORCID (KK): 0000-0002-3229-1880; (VB): 0000-0001-7897-4305
* – Author for correspondence: ksenia.g.klimova@mail.ru
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E.A. IGNATOVA, M.S. IGNATOV, K.G. KLIMOVA & V.A. BAKALIN
Fig. 1. Collecting localities. See also Table 1. More precise locality data are shown in the map in supplementary material.
#
Table 1. Collecting localities, with brief notes on vegetation and substrates
Locality
1
Southward of Ayan Settl., Sarafanovka River valley, Betula lanata and Duschekia fruticosa
crooked forest communities on the edge with Pinus pumila thickets at the bottom of NWfacing slope.
2 Small hill southward of Ayan Settl., dwarf-shrub-lichen tundra intermingled with spots of
bare ground and Pinus pumila clumps on S-SW-facing slope.
3 Small hill southward of Ayan Settl., edge of dwarf-shrub-lichen tundra bordered with Pinus pumila
thickets and Betula lanata and Duschekia fruticosa crooked forest communities.
4 Eastern spur of Dzhugdzhur Range, left tributary of Aldoma River, wide valley of small stream
with flood plain vegetation (Populus suaveolens, Salix udensis), surrounded by Larix forest
with some clumps of Pinus pumila in understory.
5 Eastern spur of Dzhugdzhur Range, left tributary of Aldoma River, Larix forest with Picea and
Betula admixture in stream valley with some clumps of Pinus pumila in understory.
6 Eastern spur of Dzhugdzhur Range, left tributary of Aldoma River, rock field on steep slope to
stream valley surrounded by Larix forest with Picea and Betula admixture and Pinus pumila
thickets.
7 Upper reaches of Tugorma River near the main ridge of Dzhugdzhur Range, stream valley in
subalpine belt with many rocks and rocky outcrops along streambed and thickets of subalpine
vegetation (Duschekia fruticosa, Pinus pumila, Betula lanata s.l.) with dwarf-shrub-mosslichen tundra spots on slope to stream.
8 Upper reaches of Tugorma River near the main ridge of Dzhugdzhur Range, NE-facing slope of
stream valley in alpine belt: mostly dwarf-shrub-moss-lichen tundra with rocky outcrops, spots of
bare ground and clumps of dwarf subalpine shrubs (Pinus pumila, Duschekia fruticosa).
9 Upper reaches of Tugorma River near the main ridge of Dzhugdzhur Range, rock fields with
small spots covered by tundra vegetation and sparse rocky outliers on NE-facing slope to
stream valley and ridgeline in alpine belt.
10 Upper reaches of Tugorma River near the main ridge of Dzhugdzhur Range, rocky ridge.
11 Upper reaches of Tugorma River near the main ridge of Dzhugdzhur Range, alpine vegetation
moistened by neutral to basic reaction percolate water with many rocky outcrops and rock
fields on steep N-NW-facing slopes.
Latitude, longitude, altitute
56°27'24"N,
138°10'16"E, 11 m
56°27'13"–56°27'10"N,
138°10'27"–138°10'25"E, 140 m
56°27'08"N,
138°11'01"E, 65 m
56°36'39" – 56°36'48"N,
137°34'15"–137°34'31"E,
294–330 m
56°36'53"N,
137°34'46"E, 377 m
56°36'58"N,
137°34'52"E, 418 m
56°39'18"–56°39'06"N,
137°15'34"–137°15'37"E,
904–935 m
56°38'52"N,
137°15'47"E, 1066 m
56°38'44"–56°38'48"N,
137°15'26"–137°15'31"E,
1261–1328 m
56°38'35"–56°38'39"N,
137°15'20"–137°15'19"E,
1393–1436 m
56°38'29"–56°38'24"N,
137°15'24"–137°15'10"E,
1473–1484 m
81
Contribution to the moss flora of Ayan
12 Upper reaches of Tugorma River near the main ridge of Dzhugdzhur Range, small rock
field at the bottom of a slope.
13 Area near the pass over main ridge of Dzhugdzhur Range, its western macro-slope (Arctic Ocean Basin), upper reaches of Birandya River, alpine belt with sparse vegetation
represented by dwarf shrub-lichen tundra and Sphagnum lenense mats over percolate
water openings, surrounded by rock fields.
14 Area near the pass over main ridge of Dzhugdzhur Range, its western macro-slope (Arctic Ocean Basin), upper reaches of Birandya River, moist mossy tundra near stream.
15 Upper reaches of Sarafanovka River northward of Ayan Settl., mostly Picea ajanensis
forest with admixture of Betula lanata and grass cover in river valley with some Salix
udensis near watercourse turned into crooked Betula lanata-Duschekia fruticosa-Pinus
pumila forest along river.
16 Upper reaches of Sarafanovka River northward of Ayan Settl., Picea forest on SW-facing
slope to the river valley and on a ridge.
17 Upper reaches of Sarafanovka River northward of Ayan Settl., Betula lanata forest on Wfacing slope.
18 Saddle in the hilly landscape near sea coast northward of Ayan Settl., windy community
with low grasses and shrubs.
19 Pribrezhnyi Range, Unych'ya River valley, large riverside SE-S-facing cliffs.
20 Pribrezhnyi Range, its northern part, Unych'ya River middle course, large N-facing conglomerate cliffs with scattered Betula lanata along riverside.
21 Pribrezhnyi Range, its northern part, eastern spur of Unych'ya Mt., small mountain with
many rocky outcrops, rock fields and Pinus pumila clumps (virtually crooked forest belt).
22 Pribrezhnyi Range, its northern part, eastern spur of Unych'ya Mt., SE-facing slope to
Unych'ya River valley, crooked Betula forest.
23 Pribrezhnyi Range, its northern part, upper course of Unych'ya River, small waterfall in
deep canyon and rocks along river bank in mixed forest.
24 Pribrezhnyi Range, its northern part, middle course of Unych'ya River, Picea forest with
some Betula lanata trees in the valley.
25 Westward of Ayan Settl., Sivakchan River valley, boggy community covered in winter by
aufeis glade not far from slightly salted estuary occasionally filling with salted water.
26 Northwestward of Ayan Settl., steep N-NE-facing slope to sea with many limestone outcrops
surrounded by tundra-like communities (developed mostly due to severe wind conditions).
27 Northwestward of Ayan Settl., lighted Larix forest in small stream valley with many
limestone boulders in the streambed.
28 Northwestward of Ayan Settl., Ayanka River valley, eutrophic hypnaceous swamp developed over limestone subbase with sluggishly flowing waters.
29 Ayan Settl. surroundings, northern coast of Ayanskaya Bay, basic sedimentary N-facing
rock outcrops along seacoast.
30 Ayan Settl. surroundings, western coast of Ayanskaya Bay, basic sedimentary N-facing
rock outcrops along seacoast.
Deficiency of data is mainly because the area is very
difficult to access. In 19th Century the Ayan Settlement
was founded in 1843 on the main road for the RussianAmerican Company trade. The road connected Europe to
Pacific seas via Yakutsk Town and Ayan (Lightfoot, 2003),
but severe climate and possible usage only in winter resulted in that after 30 years of the road exploitation it was
abandoned.
In the Soviet Union period few botanical explorations
were conducted in the area, but publications included only
vascular plants of this land (Khokhrjakov & Berkutenko,
1979; Kharkevich et al., 1983; Schlothgauer, 1990; Schlothgauer & Kryukova, 2005). Therefore, present account is
a first step to fill this ‘blank spot’ on the map of exploration of the moss flora of the Russian Far East.
Fieldworks were conducted by KK and VB over a
total of three weeks in late June – early July of 2019,
56°38'31"N,
137°14'33"E, 1119 m
56°37'47"N,
137°14'26"E, 1148 m
56°37'48"N,
137°14'03"E, 1109 m
56°27'44"–56°27'33"N,
138°12'04"–138°13'02"E,
54–143 m
56°27'35.7"–56°27'40"N, 138°13'06"–
138°13'10"E,182–273 m
56°27'51"N,
138°12'07"E, 125 m
56°27'59"N,
138°12'01"E, 105 m
56°25'26"–56°25'21"N,
137°58'16"–137°57'56"E, 82–93 m
56°25'07"N,
137°56'55", 129–154 m
56°24'36"N,
137°54'34"E, 780 m
56°24’25"N,
137°54'57"E, 517 m
56°24'15"–56°23'44"N,
137°55'19"–137°55'50"E, 179–235 m
56°25'10"N,
137°56'59"E, 117 m
56°26'49"N, 138°03'40"E, -2 m
56°29'37"N,
138°07'59"E, 34 m
56°29’38"N,
138°07’53"E, 22 m
56°29’26"N,
138°07’24"E, 91 m
56°27’33"N, 138°09’49"E, 5 m
56°27’07"N, 138°09’09"E, 5 m
addressed specifically to liverwort exploration (Bakalin
et al., 2021); however, mosses where gathered whereever possible, and 350 moss specimens were collected.
STUDY AREA
Liverwort flora analysis for the area was recently published (Bakailn et al., 2021), with the expanded overview of the area climate and vegetation. Thus here only a
brief data are provided.
The Ayan mean annual temperature at sea level is
–3.3°C, with +13.0 in August and –19.6°C in January.
The mean annual precipitation is 856 mm, with a vast
majority of them in July to September, and quite scanty
precipitation in winter (https://en.climate-data.org/asia/
russian-federation/khabarovsk-krai/ayan-718644/).
The vegetation in the study area is highly intermingled mosaic of Picea ajanensis and Larix cajanderi forest, Pinus pumila thickets, crooked forests (krummholz)
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E.A. IGNATOVA, M.S. IGNATOV, K.G. KLIMOVA & V.A. BAKALIN
A
B
C
D
E
F
G
H
Fig. 2. Vegeration types in the studied area. А, B: Dzhugdzhur Range vegetation (loc. 14). С: Dzhugdzhur Range, stream valley
(loc. 7). D: SE-facing slope to Unych’ya River, Betula lanata crooked forest and Pinus pumila thickets (loc. 22). E: Seacoast area,
wind-faced vegetation with Picea ajanensis and Betula lanata (loc. 18). F: ‘Flag formed’ trees of Larix cajanderi among tundralike community with grasses and Caragana jubata in the seacoast area (loc. 18). G: Dwarf Pinus pumila and tundra vegetation on
coastal hills along Ayanskaya Bay (loc. 2). H: Cliffy shore of the Sea of Okhotsk, with lighted Picea ajanensis stands in ‘wind
shade’ and Pinus pumila in wind exposed places (loc. 2).
Contribution to the moss flora of Ayan
83
A
B
C
D
E
F
G
H
Fig. 3. Vegeration types in the studied area and habitats of rare moss species. A: Larix cajanderi forest in the stream valley, eastern
spurs of Dzhugdzhur Range, 300 m alt., habitats of Rhizomnium nudum and Sphagnum tundrae. B: Rock field on a slope to stream valley
surrounded by Larix forest with Picea, Betula and Pinus pumila, eastern spurs of Dzhugdzhur Range, 400 m alt., habitat of Grimmia
jacutica. C: Ridgeline in alpine belt with rock fields, small spots covered by tundra vegetation and sparse rocky outliers, axial part of
Dzhugdzhur Range, 1300 m alt., habitat of Grimmia jacutica. D: Large N-facing conglomerate cliffs, Pribrezhnyi Range, Unych’ya River
valley, 129–154 m alt., with scattered Betula lanata along riverside, habitat of Hedwigia kuzenevae. E: Riverside SE-S-facing cliffs,
Pribrezhnyi Range, Unych’ya River valley, 80 m alt., habitats of Coscinodon yukonensis, Echinophyllum sachalinense and Oedipodium
griffithianum. F: Oedipodium griffithianum from ‘E’. G: Boggy community covered in winter by aufeis glade not far from slightly salted
estuary occasionally filling with salted water, -2 m alt. (sic!), westward of Ayan Settl., Sivakchan River valley, habitat of Cinclidium
stygium, Distichium inclinatum, Meesia triquetra. H: Rich fen over limestone subbase, with sluggishly flowing waters, NW of Ayan
Settlement, 90 m alt., habitats of Catoscopium nigritum, Cinclidium arcticum, Paludella squarrosa, and Tortella spitsbergensis.
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E.A. IGNATOVA, M.S. IGNATOV, K.G. KLIMOVA & V.A. BAKALIN
of Alnus fruticosa, and mountain tundra. Rock fields are
numerous. Late snow beds have willow dominated or
mossy communities. Mires are few, and one with thick
peat deposit was seen only once. Rock types are many,
ranging from acid, e.g. granites, to presumable alkaline
metamorphic rocks and limestone.
Specimens were collected in the three main areas: 1)
Ayan Settlement surroundings within a radius of 3–7 km
from the settlement, 2) northernmost part of Pribrezhnyi
Range (low and middle altitudinal belts) and Unych’ya
River valley, 3) sources of Tugorma and Birandya Rivers
in the axial part of Dzhugdzhur Range (the watershed of
Pacific and Arctic Oceans). All collected localities are
shown in Table 1 and Fig. 1.
SPECIES LIST
The names of taxa are arranged alphabetically, with
nomenclature following Cherdantseva et al. (2018), with
some recent taxonomic changes. Species is annotated by:
1) altitudinal range, in meters (given in square brackets); 2) collection localities according Table 1 and Fig.
1.; 3) accompanying moss species, if available; 4) description of habitats.
Abietinella abietina (Hedw.) M. Fleisch. – [1066–1484 m], 8,
11, 13. Humus and plant debris and those covering rocks in
mountain tundra communities with clumps of dwarf subalpine shrubs.
Amphidium cf. lapponicum (Hedw.) Schimp. – [54–935 m], 7,
15, 19, 21. Covering rocks and fine soil covering rocks in
niches in subalpine belt, fine soil filling crevice in riverside
SE-S-facing cliffs in forest belt, fallen decaying barked branch
in Picea ajanensis-Betula lanata forest in stream valley.
Andreaea alpina Hedw. (=A. obovata Thed.) – [1066 m], 8.
Covering rocks on NE-facing slope to stream valley in alpine belt.
A. rupestris Hedw. – [101–1328 m], 4, 7, 9, 15, 21. Covering
rocks and boulders in subalpine and alpine belts and in stream
and river valleys in forest belts.
Anoectangium thomsonii Mitt. – [129–154 m], 20. Covering
rocks in a big crevice of N-facing conglomerate cliffs with
scattered Betula lanata along riverside.
Anomobryum concinnatum (Spruce) Lindb. – [82 m], 19. Soil
covering rocks in niche in SE-S-facing riverside cliffs in forest belt.
Aquilonium plicatulum (Lindb.) Hedenäs, Schlesak & D. Quandt – [54–1148 m], 4, 5, 7, 13, 14, 15, 21, 24. In pure mats or
with Sanionia uncinata, Sciuro-hypnum reflexum. Picea and
Betula trunk bases and decaying wood in Picea ajanensisBetula lanata forests, covering boulders in stream valley with
flood plain vegetation surrounded by Larix forest and in
Duschekia fruticosa thickets in subalpine belt, soil in tundra community in alpine belt.
Arctoa fulvella (Dicks.) Bruch, Schimp. & W. Gümbel – [935–
1328 m], 7, 9. Sandy soil covering rocks in crevice in subalpine belt, humus in niche under rock in alpine belt.
Aulacomnium palustre (Hedw.) Schwägr. – [91–1109 m], 4,
14, 28. Plant debris covering soil in stream valley with flood
plain vegetation surrounded by Larix forest, soil in eutrophic
hypnaceous swamp developed over limestone subbase, soil
in moist mossy tundra in alpine belt.
A. turgidum (Wahlenb.) Schwägr. – [129–1148 m], 7, 13, 20.
Ground cover in thickets of subalpine vegetation in subalpine belt and in tundra communities in alpine belt, soil covering N-facing conglomerate cliffs with scattered Betula lanata along riverside.
Bartramia deciduaefolia Broth. & Yasuda – [54 m], 15. Soil
covering vertical surface of rocks in Picea ajanensis-Betula
lanata forest in the valley.
Bartramiopsis lescurii (James) Kindb. – [101 m], 15. Soil covering rocks in Picea ajanensis dominating forest with admixture of Betula lanata in river valley.
Brachythecium baicalense Ignatov – [54 m], 15. Fallen decaying barked branch in Picea ajanensis-Betula lanata forest in
the valley.
B. cirrosum (Schwägr.) Schimp. – [34–200 m], 20, 23, 26.
Humus covering rocks along river bank in mixed forest, humus on steep N-NE-facing slope to sea with many limestone
outcrops surrounded by tundra-like communities, covering
bottom of N-facing conglomerate cliffs with scattered Betula lanata along riverside.
B. erythrorrhizon Bruch, Schimp. & W. Gümbel – [129–154
m], 20. Humus covering vertical surface of N-facing conglomerate cliffs with scattered Betula lanata along riverside.
B. rotaeanum De Not. – [117–330 m], 4, 24. Decaying stump
in stream valley with flood plain vegetation surrounded by
Larix forest, Populus suaveolens trunk in Picea forest with
some Betula lanata trees in the valley.
Bryoerythrophyllum recurvirostrum (Hedw.) P.C. Chen – [5
m], 29. Soil in crevice in basic sedimentary N-facing rock
outcrops along seacoast.
Bryum amblyodon Müll. Hal. – [-2–140 m], 2, 25. Humus in
dwarf shrub-lichen tundra intermingled with spots of bare
ground and Pinus pumila clumps on a ridge near seacoast,
humus and plant debris in boggy community covered in winter by aufeis glade not far from slightly salted estuary occasionally filling with salted water.
B. argenteum Hedw. – [140 m], 2. Humus in dwarf shrub-lichen tundra intermingled with spots of bare ground and Pinus pumila clumps on a ridge near seacoast.
B. caespiticium Hedw. – [93 m], 19. Humus covering riverside
S-facing cliffs in wide river valley in forest belt.
B. creberrimum Taylor – [11 m], 1. Clayish soil covering
rocks near road surrounded by Betula lanata and Duschekia fruticosa crooked forest on NW-facing slope in
settlement area.
B. pseudotriquetrum (Hedw.) P. Gaertn., B. Mey. & Scherb. –
[11–1109 m], 1, 4, 14, 18, 26, 27. Soil and plant debris covering boulders in Betula lanata and Duschekia fruticosa
crooked forest communities, soil and soil covering rocks on
a stream banks in forest, tundra belt and in tundra-like communities on slopes to sea.
Buckia vaucheri (Lesq.) D. Ríos, M.T. Gallego & J. Guerra –
[140–1478 m], 2, 11. Clayish soil covering boulder in dwarf
shrub-lichen tundra intermingled with spots of bare ground
and Pinus pumila clumps on a ridge near seacoast, fine soil
covering rocks in niche on steep slope with scattered alpine
vegetation moistened by neutral to basic reaction percolate
water and many rocky outcrops and talus.
Bucklandiella microcarpa (Hedw.) Bedn.-Ochyra & Ochyra –
[904–1066 m], 7, 8. Covering rocks in stream valley in subalpine belt, humus and granitic sediments covering rocks in
alpine belt.
Contribution to the moss flora of Ayan
B. sudetica (Funck) Bedn.-Ochyra & Ochyra – [904 m], 7.
Covering rocks on N-facing slope to stream in subalpine belt.
Calliergonella lindbergii (Mitt.) Hedenäs – [129–154 m], 20.
Clayish soil covering vertical surface of N-facing conglomerate cliffs with scattered Betula lanata along riverside.
Campylium stellatum (Hedw.) C.E.O. Jensen – [1478 m], 11.
Clayish soil moistened by neutral to basic reaction percolate
water on steep W-NW-facing slope with scattered alpine
vegetation with many rocky outcrops and talus.
Catoscopium nigritum (Hedw.) Brid. – [91 m], 28. Soil in
eutrophic hypnaceous swamp developed over limestone subbase with sluggishly flowing waters.
Ceratodon purpureus (Hedw.) Brid. – [65–780 m], 2, 3, 19,
21. Humus in open sites in alpine and subalpine belts, on
rocks along river in forest belt, clayish soil on a roadside in
settlement area, old animal excrements in dwarf shrub-lichen tundra on a ridge near seacoast.
Cinclidium arcticum (Bruch & Schimp.) Schimp. – [91 m], 28.
Soil in eutrophic hypnaceous swamp developed over limestone subbase, with sluggishly flowing waters.
C. stygium Sw. – [-2 m], 25. Soil in boggy community covered
in winter by aufeis glade not far from slightly salted estuary
occasionally filling with salted water.
Climacium dendroides (Hedw.) F. Weber & D. Mohr – [54–
200 m], 3, 15, 23. Humus in Picea ajanensis-Betula lanata
forests in the valley and in Betula lanata crooked forest.
Cnestrum alpestre (Wahlenb.) Nyholm – [1478 m], 11. Humus
covering rocks on steep W-NW-facing slope with scattered
alpine vegetation moistened by neutral to basic reaction percolate water with many rocky outcrops and talus.
Conostomum tetragonum (Hedw.) Lindb. – [935 m], 7. Sandyhumus soil covering rocks in crevice in stream valley in subalpine belt.
Coscinodon yukonensis Hastings – [82–780 m], 19, 21. Covering rocks in subalpine belt, covering vertical surface of SES-facing riverside cliffs in wide river valley in forest belt.
Cratoneuron filicinum (Hedw.) Spruce – [22–105 m], 18, 27.
Soil in windy community with low grasses and shrubs close
to the seacoast, clayish soil covering rocks on a stream bank
in lighted Larix forest.
Cynodontium asperifolium (Lindb. & Arnell) Paris – [140 m],
2. Humus and that covering rocks in dwarf shrub-lichen tundra intermingled with spots of bare ground and Pinus pumila
clumps on a ridge near seacoast.
C. strumiferum (Hedw.) Lindb. – [418–1484 m], 6, 7, 11, 13,
21. Humus covering boulder and fine soil in niche in alpine
belt, humus with granitic sediments in subalpine belt, humus covering boulder in niche on a rock field on steep slope
to stream valley surrounded by Larix forest.
Cyrtomnium hymenophylloides (Huebener) T.J. Kop. – [82 m],
19. Sandy-clayish soil in deep niche at the bottom of SE-Sfacing riverside cliffs in wide river valley in forest belt.
Dicranella curvipes (Lindb.) Ignatov – [935 m], 7. Humus filling
crevice in rocky outcrops along streambed in subalpine belt.
D. subulata (Hedw.) Schimp. – [11 m], 1. With Ditrichum heteromallum. Clayish alluvial soil near road surrounded by
Betula lanata and Duschekia fruticosa crooked forest communities on NW-facing slope in settlement area.
Dicranum elongatum Schleich. ex Schwägr. – [294–1393 m],
4, 7, 9, 10, 13. Soil in subalpine and alpine belts, rotten
wood in stream valley in forest belt.
D. flexicaule Brid. – [904 m], 7. Soil in Duschekia fruticosa
thickets in stream valley in subalpine belt.
85
D. fuscescens Turner – [101–294 m], 4, 15, 24. Humus, rotten
wood in Picea forest with some admixture of Betula lanata
in river valley, humus with plant debris covering boulder in
stream valley with flood plain vegetation surrounded by Larix
forest.
D. laevidens R.S. Williams – [140–1148 m], 2, 13. In pure
mats or with Dicranum majus. Soil in alpine belt with dwarf
shrub-lichen tundra and Sphagnum mats over percolate water openings surrounded by rock fields, soil shaded by Pinus
pumila in dwarf shrub-lichen tundra intermingled with spots
of bare ground and Pinus pumila clumps on S-SW-facing
slope near seacoast.
D. majus Turner – [11–294 m], 1, 2, 4, 7, 15, 23, 24. Humus,
plant debris and those covering rocks in alpine and subalpine belts, Picea ajanensis-Betula lanata forest, decaying
wood in Picea forest with some admixture of Betula lanata.
D. montanum Hedw. – [418 m], 6. Humus covering boulder in
niche between tree roots on rock field on steep slope surrounded by Larix forest with Picea and Betula admixture
and Pinus pumila thickets.
D. schljakovii Ignatova & Tubanova – [935 m], 7. Sandy-clayish soil covering rocks in stream valley in subalpine belt.
Didymodon ferrugineus (Schimp. ex Besch.) M.O. Hill – [5
m], 29. Soil filling crevice in basic sedimentary N-facing
rock outcrops along seacoast.
D. rigidulus Hedw. – [179 m], 23. Covering boulder on a stream
bank in mostly inundated forest along river.
D. cf. validus Limpr. – [34 m], 26. Fine soil covering rocks on
steep N-NE-facing slope to sea with many limestone outcrops surrounded by tundra-like communities.
Distichium capillaceum (Hedw.) Bruch, Schimp. & W. Gümbel – [34–154 m], 18, 20, 26. Soil in tundra-like communities on slopes and saddle adjacent to the seacoast, humus
covering N-facing conglomerate cliffs with scattered Betula
lanata along riverside.
D. inclinatum (Hedw.) Bruch, Schimp. & W. Gümbel – [-2 m],
25. Soil in boggy community covered in winter by aufeis
glade not far from slightly salted estuary occasionally filling
with salted water.
Ditrichum heteromallum (Hedw.) E. Britton – [11 m], 1. With
Dicranella subulata. Clayish alluvial soil near road surrounded by Betula lanata and Duschekia fruticosa crooked forest communities on NW-facing slope in settlement area.
Echinophyllum sachalinense (Lindb.) O’Brien – [54–179 m],
15, 23, 24. Picea trunk bases in Picea ajanensis-Betula lanata valley forests, forest litter in mostly inundated forest
along river bank.
Encalypta brevicolla (Bruch & Schimp.) Ångstr. – [1436 m],
10. Humus covering rocks on rocky ridge in alpine belt.
E. ciliata Hedw. – [93–1478 m], 11, 19. Humus covering rocks
on steep W-NW-facing slope with scattered alpine vegetation with many rocky outcrops, on large riverside S-facing
cliffs in wide river valley in forest belt.
E. pilifera Funck – [5 m], 29. Soil filling crevice in basic sedimentary N-facing rock outcrops along seacoast.
Fissidens dubius P. Beauv. – [34–154 m], 20, 26. Soil covering
rocks in crevice in N-facing conglomerate cliffs with scattered Betula lanata along riverside and on steep N-NE-facing slope to sea with many limestone outcrops surrounded
by tundra-like communities.
Flexitrichum flexicaule (Schwägr.) Ignatov & Fedosov – [235
m], 23. Cliff crevice near small waterfall in deep canyon in
Duschekia crooked forest in subalpine belt.
86
E.A. IGNATOVA, M.S. IGNATOV, K.G. KLIMOVA & V.A. BAKALIN
F. gracile (Mitt.) Ignatov & Fedosov – [34 m], 26. Soil covering rocks on steep N-NE-facing slope to sea with many limestone outcrops surrounded by tundra-like communities.
Funaria hygrometrica Hedw. – [5 m], 29. Soil filling crevice
in basic sedimentary N-facing rock outcrops along seacoast.
Grimmia donniana Sm. – [935 m], 7. Crevice in upper surface
of rocks in stream valley in subalpine belt.
G. jacutica Ignatova, Bedn.-Ochyra, Afonina & J. Muñoz –
[418–1328 m], 6, 9. Soil with admixture of sand covering
rocks in alpine belt, humus covering boulder in niche on
rock field on steep slope surrounded by Larix forest and Pinus pumila thickets.
G. longirostris Hook. – [780 m], 21. Humus covering rocks in
subalpine belt.
G. reflexidens Müll. Hal. – [1478 m], 11. Covering rocks on
steep W-NW-facing slope with scattered alpine vegetation
with many rocky outcrops.
Hedwigia kuzenevae Ignatova & Ignatov – [129–154 m], 20.
Covering vertical surface of N-facing conglomerate cliffs with
scattered Betula lanata along riverside.
Herzogiella turfacea (Lindb.) Z. Iwats. – [82 m], 19. Rotten
log near riverside in wide river valley in forest belt.
Hygrohypnella ochracea (Turner ex Wilson) Ignatov & Ignatova – [54–179 m], 15, 23. Covering boulders and sand over
boulders submerged in running water in stream and river
valleys in forest belt.
Hylocomiadelphus triquetrus (Hedw.) Ochyra & Stebel – [101–
154 m], 15, 20. Forest litter in Picea ajanensis dominating
forest with admixture of Betula lanata in river valley, soil
covering N-facing conglomerate cliffs with scattered Betula
lanata along riverside.
Hylocomiastrum pyrenaicum (Spruce) M. Fleisch. – [101 m],
15. Soil in Picea ajanensis dominating forest with admixture of Betula lanata in river valley.
Hylocomium splendens (Hedw.) Bruch, Schimp. & W. Gümbel
– [54–904 m], 2, 5, 7, 13, 15, 24. Soil in alpine and subalpine belts and Picea ajanensis-Betula lanata forests, covering
boulder in Larix forest with Picea and Betula admixture in
stream valley.
Hymenostylium recurvirostrum (Hedw.) Dixon – [129–154 m],
20. Covering N-facing conglomerate cliffs with scattered
Betula lanata along riverside.
Hypnum cupressiforme Hedw. – [117 m], 24. Populus trunk in
Picea forest with some Betula lanata trees in the valley.
H. cupressiforme var. subjulaceum Molendo – [780 m], 21.
Fine soil covering rock in niche in subalpine belt.
Isopterygiella alpicola (Lindb.) Ignatov & Ignatova – [93 m],
19. Humus covering riverside S-facing cliffs in forest belt.
I. pulchella (Hedw.) Ignatov & Ignatova – [1484 m], 11. With
Pohlia beringiensis. Humus covering boulder on steep Nfacing slope in alpine belt.
Isopterygiopsis catagonioides (Broth.) Ignatov & Ignatova –
[129–935 m], 7, 20, 21. Soil covering rocks in niche and soil
between boulders in subalpine belt, covering N-facing conglomerate cliffs with scattered Betula lanata along riverside.
Iwatsukiella leucotricha (Mitt.) W.R. Buck & H.A. Crum –
[101–780 m], 5, 15, 16, 17, 20, 21, 24. Betula trunks in Picea forests and in subalpine belt, Picea trunk in Larix forest
with Picea and Betula admixture, decaying barked Betula
and Pinus pumila branches in Picea forest, covering vertical
surface of N-facing conglomerate cliffs with scattered Betula lanata along riverside.
Lewinskya sordida (Sull. & Lesq.) F. Lara, Garilleti & Goffinet – [54–330 m], 4, 15, 23. Salix udensis and Populus suaveolens trunks near streams in different type forests.
Loeskypnum badium (Hartm.) H.K.G. Paul – [91 m], 28. With
Paludella squarrosa, Tomentypnum nitens. Soil in eutrophic
hypnaceous swamp developed over limestone subbase with
sluggishly flowing waters.
Meesia triquetra (Jolycl.) Ångstr. – [-2 m], 25. Soil partly submerged in water in boggy community covered in winter by
aufeis glade not far from slightly salted estuary occasionally
filling with salted water.
Mnium lycopodioides Schwägr. – [101–1484 m], 4, 7, 11, 15.
Small soil niche in tundra in alpine belt, humus covering
rocks in niche in subalpine belt, Larix needles stratum covering soil in stream valley with flood plain vegetation surrounded by Larix forest, Salix udensis trunk base in Picea
ajanensis dominating forest in stream valley.
M. marginatum (Dicks.) P. Beauv. – [330 m], 4. Tree trunk in
stream valley with flood plain vegetation surrounded by Larix forest.
M. spinosum (Voit) Schwägr. – [117 m], 24. Populus trunk base
in Picea forest with some Betula lanata trees in the valley.
M. thomsonii Schimp. – [780 m], 21. Humus covering rock in
crevice in subalpine belt.
Myurella julacea (Schwägr.) Bruch, Schimp. & W. Gümbel –
[5–1478 m], 11, 19, 26, 29. Clayish soil covering rocks in
alpine belt, humus covering riverside S-facing cliffs in wide
river valley in forest belt, soil and that covering coastal basic
sedimentary N-facing rock outcrops and in tundra-like communities near seacoast.
Myuroclada maximowiczii (G.G. Borshch.) Steere & W.B.
Schofield – [5 m], 29. Soil covering basic sedimentary Nfacing rock outcrops along seacoast.
Niphotrichum canescens (Hedw.) Bedn.-Ochyra & Ochyra – [1119–
1478 m], 11, 12. Sandy-clayish and sandy soil moistened by
neutral to basic reaction percolate water on steep N-facing slope
with alpine vegetation, many rocky outcrops and rock fields.
Oedipodium griffithianum (Dicks.) Schwägr. – [82 m], 19. Soil
covering riverside SE-S-facing cliffs in wide river valley in
forest belt.
Oligotrichum falcatum Steere – [935–1066 m], 7, 8. Sandyhumus soil between boulders in alpine belt, humus filling
rock crevice in stream valley in subalpine belt.
O. parallelum (Mitt.) Kindb. – [101 m], 15. Soil wall on a
stream bank and covering vertical surface of rocks in narrow
crevice in Picea ajanensis dominating forest with admixture
of Betula lanata in river valley.
Oncophorus virens (Hedw.) Brid. – [54–154 m], 15, 20. Clayish-sandy soil covering boulder on a stream bank in Picea
ajanensis-Betula lanata forest, sandy soil covering boulder
partly submerged in water on a riverbank near large N-facing conglomerate cliffs with scattered Betula lanata.
Paludella squarrosa (Hedw.) Brid. – [91 m], 28. With Loeskypnum badium, Tomentypnum nitens. Soil in eutrophic
hypnaceous swamp developed over limestone subbase with
sluggishly flowing waters.
Plagiomnium ellipticum (Brid.) T.J. Kop. – [54 m], 15. Humus
and plant debris in wet hollow in Picea ajanensis-Betula
lanata forest in stream valley.
Plagiopus oederianus (Sw.) H.A. Crum & L.E. Anderson –
[93–154 m], 19, 20. Humus covering rocks in niche in riverside S-facing cliffs, covering vertical surface of rocks N-fa-
Contribution to the moss flora of Ayan
cing conglomerate cliffs with scattered Betula lanata along
riverside.
Plagiothecium cavifolium (Brid.) Z. Iwats. – [82 m], 19. Soil
covering riverside SE-S-facing in wide river valley cliffs.
P. denticulatum (Hedw.) Bruch, Schimp. & W. Gümbel – [101–
418 m], 6, 15, 23. Soil near small pool and covering rocks in
Picea ajanensis dominating forest in river valley, clayish soil
on a stream bank in Larix forest, humus covering rocks along
river bank in mixed forest.
P. svalbardense Frisvoll – [377 m], 5. Picea trunk base in Larix forest with Picea and Betula admixture in stream valley.
Platyhypnum alpestre (Hedw.) Ochyra – [235 m], 23. Sandy
soil covering boulder in streambed near small waterfall in
deep canyon.
P. duriusculum (De Not.) Ochyra – [179–330 m], 4, 23. Sandy
soil covering boulder on a stream bank near water’s edge
and covering boulder in a streambed in Larix forest with
flood plain vegetation along stream, sand covering boulder
submerged in water in mostly inundated forest along river
bank.
P. norvegicum (Bruch, Schimp. & W. Gümbel) Ochyra – [182
m], 16. Covering boulder in small temporary watercourse in
Picea forest on SW-facing slope of the river valley.
Pleurozium schreberi (Brid.) Mitt. – [11–904 m], 1, 4, 7, 15,
21, 24. Soil in subalpine vegetation thickets in subalpine
belt, in Picea ajanensis forests with Betula lanata admixture and in Larix forest with flood plain vegetation along
stream.
Pogonatum dentatum (Brid.) Brid. – [1148 m], 13. Humus filling crevice between boulders in alpine belt with sparse vegetation surrounded by rock fields.
P. urnigerum (Hedw.) P. Beauv. – [11–1066 m], 1, 2, 8. Clayish soil and sandy-clayish soil along streambed in alpine belt,
clayish alluvial soil on a roadside surrounded by Betula lanata and Duschekia fruticosa crooked forest communities
on NW-facing slope in settlement area.
Pohlia beringiensis A.J. Shaw – [1484 m], 11. With Isopterygiella pulchella. Humus covering boulder on steep N-facing
slope in alpine belt.
P. cruda (Hedw.) Lindb. – [54–294 m], 4, 15. Humus covering
rocks in small niche, soil wall in stream valley with flood
plain vegetation surrounded by Larix forest, humus covering
vertical surface of rocks in Picea ajanensis-Betula lanata
forest in the valley.
P. crudoides (Sull. & Lesq.) Broth. – [101–1261 m], 9, 15.
Coarse grained sand with small admixture of humus in niche
on NE-facing slope in alpine belt, covering vertical surface
of rocks in Picea ajanensis dominating forest with admixture of Betula lanata in river valley.
P. drummondii (Müll. Hal.) A.L. Andrews – [1484 m], 11.
Sandy-clayish soil on steep N-facing slope in alpine belt.
P. longicolla (Hedw.) Lindb. – [82 m], 19. Soil covering riverside SE-S-facing cliffs in wide river valley in forest belt.
P. ludwigii (Spreng. ex Schwägr.) Broth. – [1109 m], 14. Sandy
soil near stream in moist mossy tundra.
P. nutans (Hedw.) Lindb. – [117–1066 m], 4, 7, 8, 24. Sandyhumus soil covering rocks in alpine belt, humus and granitic
sediments covering rocks in stream valley in subalpine belt,
humus covering vertical surface of rocks in stream valley
with flood plain vegetation surrounded by Larix forest, rotten wood in Picea forest with some Betula lanata trees in
the valley.
87
Polytrichastrum alpinum (Hedw.) G.L. Sm. – [54–1109 m],
14, 15. Soil partly submerged in water near stream in moist
mossy tundra, humus in Picea ajanensis-Betula lanata forest in the valley.
Polytrichum commune Hedw. – [294 m], 4. Larix needles stratum covering soil in stream valley with flood plain vegetation surrounded by Larix forest.
P. juniperinum Hedw. – [11–780 m], 1, 21. Clayish soil covering rocks along road in Betula lanata and Duschekia fruticosa crooked forest communities in settlement area, humus
covering rocks in subalpine belt.
P. piliferum Hedw. – [140–1393 m], 2, 4, 10, 21. Humus and
that filling crevice in alpine belt, clayish-sandy soil covering
rock in subalpine belt, humus covering decaying wood in stream
valley with flood plain vegetation surrounded by Larix forest.
P. strictum Brid. – [1109 m], 14. Soil in moist mossy tundra
near stream.
Pseudohygrohypnum fauriei (Cardot) Jan Kučera & Ignatov –
[54 m], 15. Picea trunk base in Picea ajanensis-Betula lanata forest in the valley.
Ptilium crista-castrensis (Hedw.) De Not. – [117–904 m], 7,
24. Soil in Duschekia fruticosa thickets in subalpine belt,
decaying wood in Picea forest with some Betula lanata trees
in the valley.
Pylaisia polyantha (Hedw.) Bruch, Schimp. & W. Gümbel –
[101–330 m], 4, 15, 24. Salix udensis and Populus suaveolens trunks in Picea ajanensis forests with admixture of Betula lanata in valleys, in stream valley with flood plain vegetation surrounded by Larix forest.
P. condensata (Mitt.) A. Jaeger (P. selwynii auct. non Kindb.)
– [117 m], 24. Populus suaveolens trunk in Picea forest with
some Betula lanata trees in the valley.
Racomitrium lanuginosum (Hedw.) Brid. – [780–1148 m], 7,
13, 21. Humus covering boulder in alpine belt with sparse
vegetation surrounded by rock fields, covering rocks and
sandy-humus soil covering rocks in stream valley in subalpine belt.
Rhabdoweisia crispata (Dicks. ex With.) Lindb. – [82 m], 19.
Soil covering riverside SE-S-facing cliffs in wide river valley in forest belt.
Rhizomnium andrewsianum (Steere) T.J. Kop. – [1484 m], 11.
Humus and plant debris moistened by neutral to basic reaction percolate water on steep N-facing slope with alpine vegetation with many rocky outcrops and rock fields.
R. magnifolium (Horik.) T.J. Kop. – [179 m], 23. Humus in
moist hollow in mostly inundated forest along river bank.
R. nudum (E. Britton & R.S. Williams) T.J. Kop. – [294 m], 4.
Sandy-clayish soil in stream valley with flood plain vegetation surrounded by Larix forest.
Rhytidium rugosum (Hedw.) Kindb. – [140–1066 m], 2, 8, 20,
21. Soil and plant debris covering rocks in alpine belt, covering upper surface of N-facing conglomerate cliffs with scattered Betula lanata along riverside.
Saelania glaucescens (Hedw.) Broth. – [1478 m], 11. Clayish
soil covering rocks moistened by neutral to basic reaction
percolate water on steep W-NW-facing slope with scattered
alpine vegetation with many rocky outcrops and talus.
Sanionia uncinata (Hedw.) Loeske – [11–1148 m], 1, 4, 7, 8,
13, 15, 20, 23, 24. Soil, humus and plant debris covering
rocks in alpine belt, covering rocks and soil covering boulders in crooked Betula lanata, Duschekia fruticosa forests,
decaying wood and soil with plant debris in Picea ajanen-
e
88
E.A. IGNATOVA, M.S. IGNATOV, K.G. KLIMOVA & V.A. BAKALIN
sis-Betula lanata valley forests, Larix needles stratum covering boulder in stream valley with flood plain vegetation
surrounded by Larix forest.
Sarmentypnum exannulatum (Bruch, Schimp. & W. Gümbel)
Hedenäs – [904 –1109 m], 7, 14. Soil, submerged in water
in moist mossy tundra near stream in alpine belt, covering
rocks on N-facing slope of stream valley in subalpine belt.
Schistidium agassizii Sull. & Lesq. – [129–154 m], 20. Covering big boulder in riverbed, near large N-facing conglomerate cliffs with scattered Betula lanata along riverside.
S. papillosum Culm. – [93–1478 m], 11, 19. Humus covering
rocks on steep W-NW-facing slope in alpine belt, covering
riverside S-facing cliffs in wide river valley in forest belt.
S. pulchrum H.H. Blom – [82 m], 19. Covering vertical surface of
riverside SE-S-facing cliffs in wide river valley in forest belt.
S. rivulare (Brid.) Podp. – [101–330 m], 4, 15. Covering boulders in streambeds in stream valley with flood plain vegetation surrounded by Larix forest and in Picea ajanensis dominating forest with admixture of Betula lanata in river valley.
S. sibiricum Ignatova & H.H. Blom – [179 m], 23. Fine soil
covering boulder in a stream bank in mostly inundated forest along river bank.
Sciuro-hypnum latifolium (Kindb.) Ignatov & Huttunen – [65–
101 m], 3, 15. Humus near stream with sluggishly flowing
water in Betula lanata crooked forest in settlement area,
partly submerged in small stream in Picea ajanensis dominating forest with admixture of Betula lanata in river valley.
S. plumosum (Hedw.) Ignatov & Huttunen – [129–154 m], 20.
Covering N-facing conglomerate cliffs with scattered Betula
lanata along riverside.
S. reflexum (Starke) Ignatov & Huttunen – [11–517 m], 1, 15,
22, 24. Soil and soil covering boulder in crooked Betula lanata forest, decaying wood and humus covering boulder in
Picea ajanensis-Betula lanata forests in valleys.
S. uncinifolium (Broth. & Paris) Ochyra & Żarnowiec – [54
m], 15. Humus in Picea ajanensis-Betula lanata forest in
the valley.
Scorpidium cossonii (Schimp.) Hedenäs – [91 m], 28. Soil in
eutrophic hypnaceous swamp developed over limestone subbase with sluggishly flowing waters.
S. revolvens (Sw. ex anon.) Rubers – [91 m], 28. Submerged
and partly submerged in water in hollows in eutrophic hypnaceous swamp developed over limestone subbase with sluggishly flowing waters.
S. scorpioides (Hedw.) Limpr. – [-2 m], 25. Soil on the bottom
of hollows and pools in boggy community covered in winter
by aufeis glade not far from slightly salted estuary occasionally filling with salted water.
Seligeria diversifolia Lindb. – [129–154 m], 20. Covering Nfacing conglomerate cliffs with scattered Betula lanata along
riverside.
Sphagnum alaskense R.E. Andrus & Janssens – [904 m], 7.
Soil on N-facing slope to stream with dwarf shrub-moss-lichen tundra spots to stream in subalpine belt.
S. angustifolium (C.E.O. Jensen ex Russow) C.E.O. Jensen –
[904 m], 7. Soil on N-facing slope to stream with dwarf shrubmoss-lichen tundra spots to stream in subalpine belt.
S. aongstroemii C. Hartm. – [1148 m], 13. Soil in alpine belt
with sparse vegetation represented by dwarf shrub-lichen
tundra and Sphagnum mats over percolate water openings,
surrounded by rock fields.
S. balticum (Russow) C.E.O. Jensen – [1148 m], 13. Soil in
alpine belt with sparse vegetation represented by dwarf shrub-
lichen tundra and Sphagnum mats over percolate water openings, surrounded by rock fields.
S. capillifolium (Ehrh.) Hedw. – [91 m], 28. Soil in eutrophic
hypnaceous swamp developed over limestone subbase with
sluggishly flowing waters.
S. compactum DC. – [1109 m], 14. Soil in moist mossy tundra
near stream.
S. girgensohnii Russow – [11–294 m], 1, 4, 15. Soil in Duschekia fruticosa thickets, in Picea ajanensis dominating forest with
admixture of Betula lanata in river valley and in stream valley with flood plain vegetation surrounded by Larix forest.
S. lenense H. Lindb. ex L.I. Savicz – [904–1148 m], 7, 13. Soil
in alpine belt with sparse vegetation represented by dwarf
shrub-lichen tundra and Sphagnum mats over percolate water
openings surrounded by rock fields, in dwarf-shrub-moss-lichen tundra spots on N-facing slope to stream in subalpine
belt.
S. tundrae Flatberg – [294 m], 4. Soil in stream valley with
flood plain vegetation surrounded by Larix forest.
S. warnstorfii Russow – [1148 m], 13. In pure mats or with
Sphagnum lenense. Soil in alpine belt with sparse vegetation represented by dwarf shrub-lichen tundra and Sphagnum mats over percolate water openings, surrounded by rock
fields.
Tetraphis pellucida Hedw. – [101 m], 15. Rotten stump in Picea ajanensis dominating forest with admixture of Betula
lanata in river valley.
Tetraplodon angustatus (Hedw.) Bruch, Schimp. & W. Gümbel – [140 m], 2. Old animal excrements in dwarf shrublichen tundra intermingled with spots of bare ground and
Pinus pumila clumps on a ridge near seacoast.
T. mnioides (Hedw.) Bruch, Schimp. & W. Gümbel – [140 m],
2. In pure mats or with Tetraplodon angustatus. Old animal
excrements in dwarf shrub-lichen tundra intermingled with
spots of bare ground and Pinus pumila clumps on a ridge
near seacoast.
Tomentypnum nitens (Hedw.) Loeske – [91 m], 28. In pure mats
or with Loeskypnum badium, Paludella squarrosa. Soil in
eutrophic hypnaceous swamp developed over limestone subbase with sluggishly flowing waters.
Tortella fragilis (Hook.& Wilson) Limpr. – [5 m], 30. Soil on
basic sedimentary N-facing rock outcrops along seacoast.
T. spitsbergensis (Bizot & Thér.) O. Werner, Köckinger &
Ros – [91 m], 28. Soil in eutrophic hypnaceous swamp developed over limestone subbase with sluggishly flowing
waters.
T. tortuosa (Hedw.) Limpr. – [34–105m], 18, 26. Humus and
plant debris shaded by Caragana jubata, soil covering rocks
in tundra-like communities near seacoast.
Tortula hoppeana (Schultz) Ochyra – [93 m], 19. Humus covering riverside S-facing cliffs in forest belt.
Trachycystis ussuriensis (Maack & Regel) T.J. Kop. – [129–
154 m], 20. Humus covering conglomerate cliffs with scattered Betula lanata along riverside.
Ulota rehmannii Jur. – [54 m], 15. Picea trunk in Picea ajanensis-Betula lanata forest in the valley.
Warnstorfia pseudostraminea (Müll. Hal.) Tuom. & T.J. Kop.
– [-2 m], 25. Soil partly submerged in water in boggy community covered in winter by aufeis glade not far from slightly salted estuary occasionally filling with salted water.
Zygodon sibiricus Ignatov, Ignatova, Z. Iwats. & B.C. Tan –
[117 m], 24. Populus suaveolens trunk in Picea forest with
some Betula lanata trees in the valley.
Contribution to the moss flora of Ayan
DISCUSSION
The short time of field work revealed 164 species and
one variety, which indicates a rather high moss diversity
for this area with such harsh environmens. Of course, the
revealed diversity is far from comprehensive, and its features can be discussed only preliminary. Nevertheless, some
of them are interesting and worthy mentioning. The closest
for comparison are: (1) the local floras of Yudoma River
(tributary of Maya River, which is a tributary of
Aldan)(Ignatov et al., 2001), ca. 350 km NNW; and (2) the
Upper Bureya River (Ignatov et al., 2000), ca. 550 km SSW.
1. Iwatsuki (1972) classified mosses that occur both
in East Asia and North America in two groups. Temperate species common with North America occur in eastern North America, comprising classical ‘Arcto-Tertiary’ disjunction, between East Asia and eastern North
America. The second group occurs in North-Pacific regions of North America. There is only one species with
‘Arcto-Tertiary’ disjunction among Ayan mosses,
Pseudohygrohypnum fauriei, while North Pacific flora
elements (or Beringian sensu lato) are rather many: Coscinodon yukonensis, Trachycystis ussuriensis, Myuroclada maximowiczii, Echinophyllum sachalinense, Sciurohypnum uncinifolium, Bartramiopsis lescurii, Oligotrichum falcatum, O. parallelum, and Rhizomnium nudum.
All of them were found in the studied area at lower elevations. In Upper Bureya moss flora about the same number of North-Pacific species (9) are known (Ignatov et
al., 2000), but species with ‘Arcto-Tertiary’ disjunction
are also many (8).
Second flora taken for comparison, from Yudoma
River in South Yakutia lacks species of both aforementioned groups, except of North-Pacific Pylaisia steerei.
Here we can find a contrast in distribution patterns
between mosses and hepatics. The analysis of Bakalin et
al. (2021) found Ayan liverwort flora to be especially similar to Northern Sikhote-Alin, then Southern SikhoteAlin, and then Northern Hokkaido. For mosses, the Northern Sikhote-Alin is enriched by temperate elements
(Fedosov et al., 2016) even more than the Upper Bureya
moss flora, and in Southern Sikchote-Alin and then
Northern Hokkaido ‘Arcto-Tertiary disjuncts’ their number still increased.
2. The moss species composition characterizes the
area as cold and humid. The abundance of Sciuro-hypnum reflexum and Sanionia uncinata at most elevations
is similar to Kola Peninsula (NW Europe), Kamchatka,
and also Botchi State Nature Reserve in the south of Khabarovsk Territory, also a very humid territory (Ignatova
et al., 2013). In the latter area Aquilonium plicatulum,
Brachythecium baicalense, Echinophyllum sachalinense,
Dicranum majus, Hylocomiadelphus triquetrus, and Tetraphis pellucida are common, and Herzogiella turfacea
also is not rare. One of the most interesting mosses in
Ayan is Oedipodium griffithianum. This species was only
recently found in Russia (Ignatov et al., 2006) at about a
89
timberline in Middle Sikhote-Alin. In Ayan this species
occurs at about sea level, similar to the that in other regions with hyperoceanic climate (Størmer, 1969). The
Yudoma moss flora, being only 200 km from the Sea of
Okhotsk, includes such strict xerophytic species as Indusiella thisnschanica, Grimmia poecilostoma, and Entosthodon pulchellus. Ayan moss flora has no such elements. However, a somewhat more xeric flora in the studied area exists at upper elevation, in high mountains,
above 1000 m alt. Such species as Abietinella abietina
seems absent or at least very rare at lower elevations: all
its collections were done in the interval 1066–1484 m.
Only or mostly at the same level occur Pogonatum dentatum, Niphotrichum canescens, Racomitrium lanuginosum, Buckia vaucheri, and Conostomum tetragonum.
Some species from these elevations represent northern
flora element, close to the southern limit of general distribution: Cnestrum alpestre, Encalypta brevicollis,
Sphagnum aongstroemii, S. lenense, Rhizomnium andrewsianum.
3. Wetland and peat land moss composition is unusual. There is no Calliergon and Drepanocladus, and
Campylium was found just once at ca. 1500 m. Sarmentypnum exannulatum was found only twice, and Wanstorfia pseudostraminea – once. Rich fen species were
collected in two places, one calcareous (Catoscopium nigritum, Cinclidium arcticum, Loeskypnum badium, Paludella squarrosa, Scorpidium cossonii, S. revolvens, Tomentypnum nitens), and one salty (Cinclidium stygium,
Meesia triquetra, Scorpidium scorpioides). Interestingly, there were no one species in common in these places.
Campylium stellatum, that commonly co-occurs with
these species, was found in third place.
Although the short field trip conditions makes exploration fairly uneven, the following data on species diversity may show at least roughly a pattern of moss distribution along the altitude:
Altitude, m
studied locs.
Species
0–400
20
125
400–800
3
20
800–1200
5
44
1200–1484
3
22
As expected, low elevations comprise the maximal
diversity in this area.
ACKNOWLEDGMENTS
We are deeply indebted to Mr. Andrey Karamzin, Ayan
Settlement resident, for excellent organization of the field
research and guiding in the difficult environmental conditions. The geographic basis of the map (Fig. 1) was kindly
prepared by Mrs. Ekaterina A. Petrunenko to whom authors
are also sincerely grateful. The work of VB and KK was partially supported by the Russian Foundation for Basic Research
(grant no. 20-04-00278). We thank Ministry of Higher Education and Science of Russian Federation for the support the Center of Collective Use “Herbarium MBG
RAS”, grant 075-15-2021-678.
90
E.A. IGNATOVA, M.S. IGNATOV, K.G. KLIMOVA & V.A. BAKALIN
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Received 24 May 2021
Accepted 6 August 2021
Supplementary materials: Detailed map of collecting localities: https://kmkjournals.com/upload/PDF/Arctoa/29/
Arctoa30_79_90_supplement.pdf
Vadim Bakalin