Bull. Natl. Mus. Nat. Sci., Ser. B, 43(1), pp. 11–18, February 22, 2017

Mosses of Kedrovaya Pad Nature Reserve,
Primorskiy Territory, Far East Russia
Masanobu Higuchi1,* and Vadim Bakalin2
1

Department of Botany, National Museum of Nature and Science,
Amakubo 4–1–1, Tsukuba, Ibaraki 305-0005, Japan
2
Botanical Garden-Institute, Far Eastern Branch, Russian Academy of Sciences,
Makovskogo Street 142, Vladivostok, 690024, Russia* E-mail: higuchi@kahaku.go.jp(Received 6 December 2016; accepted 21 December 2016)

Abstract The moss ﬂora of Kedrovaya Pad Nature Reserve, Far East Russia, was investigated in2015. The mosses recognized comprise 26 families, 65 genera and 93 species. Three species, Hypnum oldhamii, H. tristo-viride and Palamocladium leskeoides, are new to Russia. The moss ﬂora
of Kedrovaya Pad Nature Reserve mainly consists of East Asian and circumboreal species.
Eocronartium muscicola, parasitic bryophilous basidiomycete, was found on four species, Myuroclada maximoviczii, Oncophorus crispifolius, Pylaisia stereodontoides and Taxiphyllum
aomoriense. For each species, locality, substrate, specimen number and element of distribution
were given.
Key words: bryophytes, Eocronartium muscicola, Far East Russia, ﬂora, Kedrovaya Pad Nature
Reserve, mosses, Primorskiy Territory.

Introduction
This study deals with the moss ﬂora of Kedrovaya Pad Nature Reserve, Primorskiy Territory,
Far East Russia. Although the outline of the moss
ﬂora in Russia is becoming clear (cf. Ignatov etal., 2006), there has never been enough knowledge on the local ﬂora. In 2015 we made a ﬁeld
research and collected bryophytes mainly from
Kedrovaya River basin.
Kedrovaya Pad Nature Reserve is situated at
ca. 20 km west of Vladivostok, 43°01′ to
43°09′N; 131°24′ to 131°36′E, and located in thesouthernmost of Far East Russia, that is, on the
Khasansky District in the southern part of Primorsky territory, and it occupies an area of ca.
180 square kilometers. The native deciduous
broad-leaved and the coniferous forests are dominated by Quercus mongolica, Acer mono, etc.
and Abies holophyla, Pinus koraiensis, etc.,

respectively. This area is also famous forendemic animals such as Amur tiger, Amur leopard and Asian black bear.
Materials and Methods
Field study was carried out in 2015, and a totalof ca. 160 bryophyte specimens were collected.
The complete set of the specimens are preservedin the herbarium of the Department of Botany,
National Museum of Nature and Science (TNS)and some duplicates in the herbarium of the
Botanical Garden-Institute, Far Eastern Branch,
Russian Academy of Sciences (VBGI). The mainsites investigated are as follows.
A. Kedrovaya River Valley, the ﬂood plain,broad-leaved deciduous temperate forest,
43°06′12N, 131°32′28E, ca. 45–60 m alt.,
July 8, 2015.
B. Kedrovaya River system, Vtoroj Zolotoj

12Masanobu Higuchi and Vadim Bakalin

Stream middle course, broad-leaved deciduous temperate forest with admixture ofAbies holophylla and Pinus koraiensis,
43°05′52N, 131°32′30E, ca. 150–500 m alt.,
July 9, 2015.
C. Kedrovaya River system, ﬂood plain in middle course, broad-leaved deciduous temperate forest with admixture of Abies holophylla and Pinus koraiensis, 43°06′12N,
131°32′58E, ca. 60–80 m alt., July 10, 2015.
D. Kedrovaya River system, upper course of
Pervyj Zolotoj Stream, broad-leaved BetulaAbies holophylla-Pinus koraiensis forest on
slope, 43°05′19N, 131°32′02E, ca. 65–350 m
alt., July 11, 2015.
Results and Discussion
The mosses recognized in this study comprise
26 families, 65 genera and 93 species. Three species, Hypnum oldhamii, H. tristo-viride and Palamocladium leskeoides, are new to Russia (cf.
Ignatov et al., 2006).
1.

Phytogeographical analysis of the moss ﬂora
To make clear the characteristics of the moss
ﬂora of Kedrovaya Pad Nature Reserve, the elements of distribution was studied. A total of 93
species recognized from this area include several
elements of distribution. They are classiﬁedroughly into the following ﬂoristic groups: East
Asian species (41 spp., 44%), circumboreal species (31 spp., 33%), cosmopolitan species (8spp., 9%), or others (13 spp., 14%). There are noendemic species to this area. The East Asian andcircumboreal species constitute about 77% of thespecies and represent the moss ﬂora of Kedrovaya Pad Nature Reserve. The East Asian species
occur in East Asia including China, Korea andJapan, but exceptionally extend to westwards
Himalayan region and northeastwards Alaska
region. Twenty one species of 41 East Asian species are known only from the “South Far East” in
Russia (cf. Ignatov et al., 2006). The circumboreal species, widely occurring in the Northern
Hemisphere, include Anomodon rugelii, Bartra-mia pomiformis, Calliergonella lindbergii, etc.
The cosmopolitan species, occurring widely
without relation to any continent or vegetation
types, are represented by Anoectangium thomsonii, Fissidens taxifolium, Haplocladium angstifolium, etc. Among others, Thuidium cymbifolium
is only species known as palaeotripic species
which is distributed mainly in tropical Asia.
Infection of Eocronartium muscicola
Eocronartium muscicola is a parasitic bryophilous basidiomycete and is known as an obligate
parasite recorded on 21 moss species of different
genera and families (Boehm and McLaughlin,1988). We found fruiting bodies of Eocronartium
muscicola at young sporophytes of Myuroclada
maximoviczii (Fig. 3), Oncophorus crispifolius,
Pylaisia stereodontoides (Fig. 9) and Taxiphyllum aomoriense (Fig. 10) in different sites of thisarea. Although they belong to the different taxonomic groups, they could have the similar developmental stage of the sporophytes in the ﬁeld.
2.

3.

Enumeration of species
The species recognized from Kedrovaya Padʼ
Strict Nature Reserve are listed below. The scientiﬁc names follow Iwatsuki (2004) and Gofﬁnet
et al. (2009). The families, genera and species
are arranged alphabetically. Each species isreferred by collecting sites (A–D), substrates,
collectorʼs numbers of Higuchi and distributional
elements. An asterisk (*) preceding a species
indicates a species to be known only from the“South Far East” in Russia (cf. Ignatov et al.,2006).
Amblystegiaceae
Campylium hispidulum (Brid.) Mitt. B: on
basal part of rock-cliff, mh-53510 (＋Oxystegus tenuirostris), 53512. C: on decaying log,
mh-53573. Circumboreal.
*Pseudohygrohypnum eugyrium (Bruch &Schimp.) Kanda. B: on boulder, mh-53540(＋Brachythecium rivulare). Circumboreal.
Sanionia uncinata (Hedw.) Loeske B: on rock-Mosses of Kedrovaya Pad Nature Reserve, Far East Russia

13

Figs. 1–8. 1. A broad-leaved deciduous forest, along a branch of Kedrovaya River, ca. 60 m alt. 2. A mixed Betula sp, Abies holophylla and Pinus koraiensis forest, at upper course of Pervyj Zolotoj Stream, Kedrovaya
River system, ca. 350 m alt. 3. Myuroclada maximoviczii growing on boulder. Eocronartium muscicola, parasitic bryophilous basidiomycete, infected the species. 4. Dicranum mayrii growing on decaying log. 5. Physcomitrium sphaericum growing on soil. 6. Gollania ruginosa growing on basal part of trunk. 7. Hypnum oldhamii growing on boulder. 8. Hypnum tristo-viride growing on boulder.

14

Masanobu Higuchi and Vadim Bakalin

Figs. 9–16. 9. Pylaisia stereodontoides growing on decaying log. Eocronartium muscicola infected the species.
10. Taxiphyllum aomoriense growing on boulder. Eocronartium muscicola infected the species. 11. Trachycystis ussuriensis growing on boulder. 12. Macromitrium japonicum growing on trunk of Juglans sp. 13. Ulota
japonica growing on tree-trunk. 14. Pogonatum japonicum growing on soil. 15. Glyphomitrium humillimum
growing on decaying branch. 16. Bryonoguchia molkenboeri growing on boulder.

Mosses of Kedrovaya Pad Nature Reserve, Far East Russia

cliff, mh-53511. Circumboreal.
Anomodontaceae*Anomodon abbreviatus Mitt. D: on trunk of
Quercus mongolica, mh-53579. East Asian.
Anomodon rugelii (Müll.Hal.) Keissl. D: onroot, mh-53582 (＋Rauiella fujisana). Circumboreal.
Anomodon thraustus Müll.Hal. A: on trunk of
Quercus dentata, mh-53470 (＋Haplohymenium triste); on tree-trunk, mh-53487. Others.
Haplohymenium triste (Ces.) Kindb. A: on treetrunk of Quercus dentata, mh-53470 (＋Anomodon thraustus); on trunk of Acer mono,
mh-53493 (＋Pylaisia stereodontoides). Circumbreal.
Herpetineuron toccoae (Sull. & Lesq.) Cardot
A: on trunk of Alnus sp., mh-53498). Cosmopolitan.
*Schwetschkeopsis fabronia (Schwägr.) Broth.
C: on rock-cliff, mh-53565. Others.
Bartramiaceae
Bartramia pomiformis Hedw. B: on rock-cliff,
mh-53529. Circimboreal.
Brachytheciaceae
Brachythecium rivulare Schimp. A: on boulder,
mh-53474. B: on boulder, mh-53540(＋Hygrohypnum eugyrium). D: on rock-cliff,
mh-53586. Circumboreal.
Brachythecium salebrosum (F.Weber & Mohr)
Schimp. D: on soil bank, mh-53589. Others.
Bryhnia hultenii E.B.Bartram A: on boulder
covered with humus, mh-53479. C: on rockcliff, mh-53570. East Asian (incl. west N.
America).
Bryhnia novae-angliae (Sull. & Lesq.) Grout A:
on humus, mh-53500 (＋Plagiomnium acutum). C: on boulder covered with thin layer ofsoil, mh-53576. Circumboreal.
Myuroclada maximoviczii (Borcz.) Steere & W.
B.Schoﬁeld A: on boulder, mh-53491. East

15

Asian (incl. west N. America). (Fig. 3)*Okamuraea hakoniensis (Mitt.) Broth. C: on
decaying log, mh-53572. East Asian.
Palamocladium leskeoides (Hook.) E.G.Britton
D: on rock-cliff, mh-53603 (＋Thamnobryum
plicatulum). East Asian.
Notes. This species is new to Russia.
Rhynchostegium riparioides (Hedw.) Cardot C:
on boulder, mh-53563. Circumboreal.
Bryaceae
Bryum cyclophyllum (Schwägr.) Bruch &Schimp. B: on boulder, mh-53546. Circumboreal.
Climaciaceae
Climacium dendroides (Hedw.) F.Weber &Mohr A: on boulder covered with humus,
mh-53482. Others.
Dicranaceae
Dicranella heteromalla (Hedw.) Schimp. B: onrock-cliff, mh-53535 (＋Rhabdoweisia crispata). D: on rock-cliff, mh-53590, 53593. East
Asian.
Dicranum ﬂagellare Hedw. B: on decaying log,
mh-53549. Others.
Dicranum japonicum Mitt. B: on boulder,
mh-53561. D: on boulder, mh-53606, 53626(＋Polytrichum formosum), 53627. East Asian.
*Dicranum mayrii Broth. A: on decaying log,
mh-53477 (＋Haplocladium microphyllum),
53489, 53497 (＋Haplocladium microphyllum). East Asian.
*Dicranum nipponense Besch. B: on decaying
log, mh-53517 (＋Rhizomnium parvulum).
East Asian.
Entodontaceae
Entodon ﬂavescens (Hook.) A.Jaeger A: on
decaying log, mh-53478. East Asian (incl. S.
India).

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Masanobu Higuchi and Vadim Bakalin

Entodon scabridens Lindb. C: on tree-trunk,mh-53578. East Asian.
*Entodon sullivantii (Müll.Hal.) Lindb. C: onboulder, mh-53562 (＋Haplocladium angstifolium), 53566. East Asian.
Fissidentaceae
Fissidens dubius P.Beauv. D: on rock-cliff,
mh-53602. Circumboreal.
*Fissidens gymnogynus Besch. A: on base oftree-trunk, mh-53472. East Asian.
Fissidens taxifolius Hedw. D: on soil at creviceof rock-cliff, mh-53584. Cosmopolitan.
Funariaceae
Physcomitrium sphaericum (Ludw.) Fuernr. A:
on soil, mh-53469. Others.
Grimmiaceae
Grimmia pilifera P.Beauv. B: on boulder,
mh-53548. Others.
Hylocomiaceae
Hylocomiastrum pyrenaicum (Spruce) Broth.
D: on boulder, mh-53614 (＋Thuidium cymbifolium). Circumboreal.
Hylocomium splendens (Hedw.) Schimp. D: onboulder covered with humus, mh-53617,
53629. Circumboreal (incl. New Zealand).
Pleurozium schreberi (Brid.) Mitt. D: on boulder, mh-53625, 53631. Circumboreal.
*Rhytidiadelphus japonicus (Reimers) T.J.
Kop. B: on humus, mh-53530. D: on boulder,
mh-53615. East Asian.
Rhytidiadelphus triquetrus (Hedw.) Warnst. B:
on boulder, mh-53554. D: on boulder,
mh-53623. Circumboreal.
Hypnaceae
Callicladium haldanianum (Grev.) H.A.Crum
A: on decaying log, mh-53496. Circumboreal.

Calliergonella lindbergii (Mitt.) Hedenäs B: on
boulder, mh-53528. C: on decaying log,
mh-53571; on boulder, mh-53580. Circumboreal.
Gollania ruginosa (Mitt.) Broth. A: growing on
basal part of trunk, mh-53483. B: on decaying
log, mh-Higuchi 53526. East Asian.
Note. Higuchi (2016) reported this species
from Kedrovaya Padʼ Strict Nature Reserve.
Herzogiella turfacea (Lindb.) Z.Iwats. C: on
decaying log, mh-53568. Circumboreal.
Hypnum cupressiforme Hedw. B: on rock-cliff,
mh-53536. Cosmopoliatan.
Hypnum oldhamii (Mitt.) A.Jaeger & Sauerb.
D: on boulder, mh-53633. East Asian.
Note. This species is new to Russia.
Hypnum plicatulum (Lindb.) A.Jaeger & Sauerb. D: on boulder, mh-53619. Circumboreal.
Hypnum tristo-viride (Broth.) Paris D: on boulder, mh-53636. East Asian.
Note. This species is new to Russia.
Isopterygiopsis muelleriana (Schimp.) Z.Iwats.
B: on rock-cliff, mh-53508 (＋Taxiphyllum
taxirameum). D: on soil at crevice of rockcliff, mh-53604; on boulder, mh-53632. Circumboreal.
*Pylaisia brotheri Besch. A: on trunk of Alnussp., mh-53495. East Asian.
*Pylaisia stereodontoides Broth. & Yasuda ex
Iisiba A: on decaying log, mh-53480; on fallen
tree-trunk, mh-53481; on trunk of Acer mono,
mh-53493 (＋Haplohymenium triste). B: on
dead tree-trunk, mh-53560. C: on decaying log,
mh-53574; on trunk of Acer mono, mh-53577;
on fallen tree-trunk, mh-53601. East Asian.
*Taxiphyllum aomoriense (Besch.) Z.Iwats. A:
on base of tree-trunk, mh-53471; on boulder,
mh-53476. C: on rock-cliff, mh-53564. East
Asian.
*Taxiphyllum taxirameum (Mitt.) M.Fleisch.
B: on rock-cliff, mh-53508 (＋Isopterygiopsis
muelleriana). Others.
Lembophyllaceae
Dolichomitriopsis diversiformis (Mitt.) Nog. B:

Mosses of Kedrovaya Pad Nature Reserve, Far East Russia

17

Forsstroemia japonica (Besch.) Paris B: on
tree-trunk, mh-53544 (＋Forsstroemia trichomitria). East Asian.
*Forsstroemia trichomitria (Hedw.) Lindb. B:
on tree-trunk, mh-53544 (＋Forsstroemia
japonica). Others.

*Rhizomnium parvulum (Mitt.) T.J.Kop. B: on
decaying log, mh-53516, 53517 (＋Dicranum
nipponense); on humus, mh-53531. D: on soil
at crevice of rock-cliff, mh-53591. East Asian.
Trachycystis ﬂagellaris (Sull. & Lesq.) Lindb.
B: on humus, mh-53532. D: on boulder,
mh-53611. East Asian (incl. west N. America).
Trachycystis ussuriensis (Maack & Regel) T.J.
Kop. B: on boulder, mh-53524. D: on boulder,
mh-53608. East Asian.

Leskeaceae

Neckeraceae

Haplocladium angstifolium (Hampe & Müll.
Hal.) Broth. C: on boulder, mh-53562(＋Entodon sullivantii). Cosmopolitan.
Haplocladium microphyllum (Hedw.) Broth. A:
on decaying log, mh-53477 (＋Dicranum
mayrii), 53492, 53497 (＋Dicranum mayrii).
Cosmopolitan.
*Hylocomiopsis ovicarpa (Besch.) Cardot D: onboulder, mh-53620. East Asian.
*Miyabea fruticella (Mitt.) Broth. B: on trunkof Acer mono, mh-53547. East Asian.

Homalia trichomanoides (Hedw.) Schimp. B:
on rock-cliff, mh-53509; on root, mh-53543.
Circumboreal.
*Thamnobryum coreanum (Cardot) Nog. & Z.
Iwats. B: on boulder, mh-53520, 53541. East
Asian.
*Thamnobryum plicatulum (Sande Lac.) Z.
Iwats. B: on rock-cliff, mh-53519. D: on rockcliff, mh-53603 (＋Palamocladium leskeoides). East Asian.

on base of tree-trunk, mh-53552. East Asian.
Leptodontaceae

Leucobryaceae
Brothera leana (Sull.) Müll.Hal. B: on basal
part of tree, mh-53555; on decaying stump,
mh-53556. Others.
Mniaceae
Mnium lycopodioides (Hook.) Schwägr. B: onrock-cliff, mh-53507; on boulder, mh-53545.
Circumboreal.
Plagiomnium acutum (Lindb.) T.J.Kop. A: onbase of tree-trunk, mh-53485. B: on boulder,
mh-53515. D: on boulder, mh-53596. East
Asian.
Plagiomnium medium (Bruch & Schimp.) T.J.
Kop. A: on humus, mh-53499. C: on humus,
mh-53567. Circumboreal.
Pohlia wahlenbergii (F.Weber & Mohr) A.L.
Andrews B: on rock-cliff, mh-53537. Cosmopolitan.

Orthotrichaceae*Macromitrium japonicum Dozy & Molk. A:
on trunk of Juglans sp., mh-53486. East Asian.
*Orthotrichum consobrinum Cardot D: on
tree-trunk, mh-53598. East Asian.
Ulota crispa (Hedw.) Brid. B: on decaying log,
mh-53550 (＋Glyphomitrium humillimum). D:
on branch, mh-53610. Circumpolitan.
*Ulota japonica (Sull. & Lesq.) Mitt. D: on treetrunk, mh-53609. East Asian (incl. west N.
America).
Plagiotheciaceae
Plagiothecium cavifolium (Brid.) Z.Iwats. D: on
rock-cliff, mh-53592. Circumboreal.
Plagiothecium laetum Schimp. B: on rock-cliff,
mh-53514. C: on boulder, mh-53569. D: on
rock-cliff, mh-53583. Circumboreal.
Plagiothecium nemorale (Mitt.) A.Jaeger B:
on rock-cliff, mh-53533. D: on rock-cliff,

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Masanobu Higuchi and Vadim Bakalin

mh-53600. Circumboreal.
Polytrichaceae*Atrichum rhystophyllum (Müll.Hal.) Paris A:
on soil, mh-53494. Circumboreal.
Pogonatum contortum (Brid.) Lesq. D: on soil,
mh-53634. East Asian (incl. west N. America).
Pogonatum japonicum Sull. & Lesq. D: on soil,
mh-53622, 53635. East Asian.
Polytrichastrum alpinum (Hedw.) G.L.Sm. D:
on boulder, mh-53612. Circumboreal.
Polytrichastrum formosum (Hedw.) G.L.Sm.
D: on boulder, mh-53624, 53626 (＋Dicranum
japonicum). Cosmopolitan.
Pottiaceae
Anoectangium thomsonii Mitt. B: on rock-cliff,
mh-53506. Cosmopolitan.
Oxystegus tenuirostris (Hook. & Taylor)
A.J.E.Smith A: on boulder, mh-53490. B: on
basal part of rock-cliff, mh-53510 (＋Campylium hispidulum); on boulder, mh-53523. D:on boulder, mh-53597. Circumboreal.
Pylaisiadelphaceae
Platygyrium repens (Brid.) Bruch & Schimp. B:
on decaying log, mh-53502; on decaying
branch, mh-53504. Circumboreal.
Pylaisiadelpha tenuirostris (Bruch & Schimp.)
W.R.Buck A: on fallen tree-trunk, mh-53475;
on decaying log, mh-53501. B: on decaying
log, mh-53505; on trunk of Quercus mongolica, mh-53559. D: on trunk of Betula sp.,
mh-53628. East Asian.
Rhabdoweisiaceae*Glyphomitrium humillimum (Mitt.) Cardot
B: on decaying branch, mh-53539, 53550(＋Ulota crispa); on trunk of Quercus mongolica, mh-53558. D: on fallen branch,
mh-53581. East Asian.
Oncophorus crispifolium (Mitt.) Lindb. D: on

rock-cliff, mh-53595. East Asian.
Rhabdoweisia crispata (With.) Lindb. B: onrock-cliff, mh-53534, 53535 (＋Dicranella
heteromalla). D: on rock-cliff, mh-53594. Others.
Thuidiaceae*Boulaya mittenii (Broth.) Cardot B: on trunkof Quercus mongolica, mh-53557. East Asian.
*Bryonoguchia molkenboeri (Sande Lac.) Z.
Iwats. & Inoue D: on boulder, mh-53621. East
Asian.
Rauiella fujisana (Paris) Reimers D: on root,
mh-53582 (＋Anomodon rugelii). East Asian.
*Thuidium cymbifolium (Dozy & Molk.)
Broth. B: on boulder, mh-53542. D: on boulder, mh-53614 (＋Hylocomiastrum pyrenaicum). Others.
*Thuidium subglaucinum Cardot B: on rockcliff, mh-53538. East Asian.
Acknowledgments
We wish to thank Dr. Y. Ohmura and Ms. Olga
Deryugina for their help in the ﬁeld research andMiss A. Watanabe for her help in the identiﬁcation of Eocronartium muscicola.
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