Arctoa (2021) 30: 407–416
doi: 10.15298/arctoa.30.26
LIVERWORT FLORA OF THE JIRISAN NATIONAL PARK IN KOREA
ФЛОРА ПЕЧЁНОЧНИКОВ НАЦИОНАЛЬНОГО ПАРКА ЧИРИСАН В КОРЕЕ
HYUN MIN BUM1, VADIM A. BAKALIN2, SEUNG JIN PARK3, SEUNG SE CHOI4*,
CHUL HWAN KIM1 & SOONKU SO5
ХЁН МИН БОМ1, ВАДИМ А. БАКАЛИН2, СЫНГ ЧИН ПАК3, СЫНГ СЭ ЧЕ4*,
ЧУЛ ХВОН КИМ1, СУН КУ СО5
Abstract
The liverwort flora of the Jirisan National Park was studied and analyzed to create a list of taxa,
that includes 177 species. Of these, 14 species are recorded only in literature and 78 species are newly
recorded for the Jirisan Mountains. One species (Lopholejeunea subfusca) was excluded from the list
because its occurrence in the Jirisan mountains is doubtful. The flora is characterized by the dominance of the East Asian temperate elements with great participation of northern boreal and subalpine
taxa. The rather noticeable diversity of the flora is explained by a wide altitudinal range in the study
area, which results in a high diversity of communities and habitat types.
Резюме
Изучена и проанализирована флора печеночников национального парка Чирисан. По имеющимся данным, она включает 177 видов. Из них 14 видов известны только по литературным
данным, а 78 видов приводятся для гор Чирисан впервые. Один вид (Lopholejeunea subfusca)
исключен из списка, поскольку его присутствие в горах Чирисан вызывает сомнения. Для флоры
характерно преобладание восточноазиатских неморальных видов с большим участием бореальных
и субальпийских таксонов. Значительное разнообразие флоры объясняется большим высотным
диапазоном исследуемой территории, что обуславливает большое разнообразие сообществ и типов
местообитаний.
KEYWORDS: liverworts, flora, checklist, Jirisan mountains, South Korea, East Asia
INTRODUCTION
The Jirisan National Park designated South Korea’s
first national park in 1967, is located on the boundaries
of Jeollabuk-do, Jeollanam-do and Gyeongsangnam-do
provinces. It comprises differing plant communities, from
warm-temperate broad-leaved deciduous forests and dispersed evergreen shrubs in the lowlands to extensive gravelly barrens with scattered vegetation, subalpine shrubs
and crooked forests at upper elevations. The vascular
plants of the Jirisan Mountains have been well studied
with 2042 species recorded in the literature (Nakai, 1915;
Jang et al., 2007; Hyun, 2019). In contrast, only fragmentary information on the region’s liverwort flora is
available in a scattering of publications (Uno & Takahasi, 1940; Hong & Yoo, 1961; Hattori et al., 1962; Song
1
& Yamada, 2009; Bakalin et al., 2009). There is a similar paucity of data for the park’s mosses with only 133
species recorded in the literature (Uno & Takahasi, 1940;
Hong & Ando, 1961; Choe, 1962, 1980). However, considering the high-altitude landscape and plant community variations of the study area, we proposed that the
diversity of liverworts in the park is likely greater than
previously recorded. Moreover, comprehensive data on
liverwort taxonomic diversity is urgently needed to inform conservation effort within the Korean Peninsula,
and to improve understanding of liverwort distribution
patterns and ecology in a broad, East Asian context. Thus,
the aim of the present study is to provide new and comprehensive records of liverwort diversity in the Jirisan
National Park.
– Department of Life Science, Jeonbuk National University, Jeonju, 54896, South Korea. E-mail: know8318@jbnu.ac.kr,
plantaxa@hanmail.net (Chul Hwan Kim is deceased); ORCID (HB): 0000-0001-7518-0702, (CK): 0000-0003-4894-0651
2 – Botanical Garden-Institute, Far Eastern Branch of the Russian Academy of Sciences, Makovskogo Street, 142, Vladivostok,
690024, Russia; E-mail: vabakalin@gmail.com; ORCID: 0000-0001-7897-4305
3 – Division of botany, Honam National Institute of Biological Resources, Mokposi, 58762, South Korea. E-mail: moss89@hnibr.re.kr
ORCID: 0000-0002-7967-8361
4 – Team of National Ecosystem Survey, National Institute of Ecology, Seocheon, 33657, South Korea. E-mail: hepaticae@nie.re.kr
ORCID: 0000-0002-3332-5544
5 – Korea National Park Research Institute, Korea National Park Service, Wonju, 26441, South Korea. E-mail: ssk822@knps.or.kr
ORCID: 0000-0001-7044-5441
* – Corresponding author: hepaticae@jbnu.ac.kr
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H.M. BUM, V.A. BAKALIN, S.J. PARK, S.S. CHOI, C.H. KIM & S. SO
Fig. 1. The position of the Jirisan National Park within Korea and the surveyed routes in this study. For details of collections
see supplementary materials.
STUDY AREA
The Jirisan National Park lies between longitudes
127°17'09"E – 127°49'29"E and latitueds 35°12'42"N –
35°26'43"N. Altitudes range from ca. 600 m a.s.l. in the
lowlands to 1915 m a.s.l. at Cheonwangbong Mt., the
highest point of mainland South Korea (Figs. 1–2). Other relatively high mountains in the park are Nogodan
Mt. (1507 m a.s.l., western sector) and Banyabong Mt.
(1732 m a.s.l., central sector). The total area of the national park is 472 km2. The lowland area of the park is
populated by broad-leaved deciduous forest that gradually changes into coniferous mountain forest near ridgelines intermixed with thickets of Magnolia sieboldii K.
Kochand and Weigela florida (Bunge) A. DC. sometimes
forming a kind of crooked forest at high elevations (Park,
2019). The dominant species of the deciduous forest are
Quercus mongolica Fisch. ex Ledeb., Carpinus laxiflora
(Siebold & Zucc.) Blume, Fraxinus rhynchophylla Hance,
and Acer pictum Thunb. var. mono (Maxim.) Franch.
The dominant species of the coniferous mountain forest
are Abies koreana E.H. Wilson, Taxus cuspidata Siebold
& Zucc., Pinus koraiensis Siebold & Zucc., and Abies
nephrolepis (Trautv. ex Maxim.). The ridgeline area is
also rich in rocky outcrops and occasional gravelly barrens on steep slopes.
The mean annual temperature in the lowland area is
12.8°C, with the coldest month being January (average
temperature varying from 8.4–1.4°C) and the warmest
month is August with an average temperate between 23.3
and 31.4°C. These parameters vary greatly depending
on elevation and exposition. Mean annual precipitation
in the lowlands is 2136 mm, with the summer maximum
1444.4 mm (https://data.kma.go.kr). At higher altitudes
the amount of precipitation noticeably increases due to
interception of wet air masses from the Pacific Ocean.
The area of the park features southern inland climate.
The dominant mineral rocks in the park are granitic
gneiss (Shin, 2019).
Exploration of the hepatic flora of the park
The liverwort flora of the park has been studied for
over 70 years, although there have been some lengthy
breaks between reports in the literature. The first report
on liverworts from the Jirisan Mountians was published
by Uno and Takahashi (1940) who recorded four species
(Brachiolejeunea sandvicensis (Gottsche) A. Evans (=
Acrolejeunea sandvicensis), Frullania fauriana, Frullania moniliata (Reinw., Blume & Nees) Mont. subsp. obscura Verd. (= Frullania appendiculata) and Madotheca setigera (Steph.) S. Hatt. (= Porella caespitans var.
cordifolia)) in the current territory of the park. Then Hong
& Yoo (1961) recorded four species (Frullania diversitexta, F. schensiana, F. taradakensis, and F. usamiensis) as
new for Korean flora. Hattori et al. (1962) recorded 50
taxa, also including 15 species new for the liverwort flora of the Korean Peninsula. After a long gap, Song &
Liverwort Flora of the Jirisan National Park in Korea
409
Fig. 2. Landscapes of the Jirisan National Park in Korea. A. Main ridge of Cheonwangbong (main peak, 1915 m) – Nogodan
(1507 m). B. Abies koreana forest near peak of Jeseokbong (1808 m). C. Abies koreana forest near peak of Banyabong (1732 m).
D. Hansin valley (1350 m). E. Chilseon valley. F. Baemsagol valley (1100 m). Photos: S.S.Choi.
Yamada (2009) published a checklist that included 79
species known in the park, 16 of which were first records.
During the same year, the authors of the present account
started a floristic exploration of the area. Bakalin et al.
(2009) described Tritomaria koreana as new to science;
the species occupies an isolate position in Lophozia
(Bakalin et al., 2021a). Choi et al. (2012) reported five
taxa (Neotrichocolea bissetii, Calypogeia angusta, Cephaloziella massalogi, Harpanthus scutatus, and Plectocolea torticalyx) from the Jirisan National Park that were
new records for the Korean Peninsula. Recently, Bakalin
et al. (2019) described Marsupella koreana and Bakalin
et al. (2020) described Solenostoma jirisanense as new
to science. In total, 99 species were recorded from the
park, 19 of which were recorded by the authors’ team.
METHODS
Field surveys
Our purposeful studies began in 2009 when Choi and
Bakalin first visited the park. Subsequently, Choi visited
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H.M. BUM, V.A. BAKALIN, S.J. PARK, S.S. CHOI, C.H. KIM & S. SO
many localities in the park, as shown in Fig. 1 (see also
Supplementary Materials). In total, over 2500 specimens
were collected and identified. Most of them are stored in
JNU, with some duplicates and Bakalin’s originals in
VBGI. The specimens were identified by Choi and Bakalin during a special collaborative project.
Latitudinal and longitudinal fractional composition
To analyze of the distribution characteristics of bryophytes, we obtained global distribution data from Bakalin
(2010), and Korean distribution information from Choe
(1980), Kim & Hwang (1991), and Choi et al. (2021).
LIST OF TAXA
The nomenclature applied Söderström et al. (2016)
with some updates from the recent literature. Each species is annotated by: 1) presence of sexual and asexual
reproductive structures in the studied specimens, using
the abbreviations spor. – sporogonia, arch. – archegonia,
ant. – antheridia, per. – perianthia, gemm. – gemmae; 2)
habitat; 3) altitudinal range in the studied area; 4) accompanying taxa; 5) selected voucher specimen numbers (Choi’s collections are prefixed by CS; Bakalin’s
collections are prefixed by VB). New records from the
Jirisan Mountains are marked with asterisks. Unconfirmed records are marked with a degree symbol (°).
Acanthocoleus yoshinaganus (S. Hatt.) Kruijt. – on shaded wet
rocks; 647–848 m alt.; CS 3646, CS 3651, CS 8122, CS
110662 (JNU).
*Acrobolbus ciliatus (Mitt.) Schiffn. – on wet rock near the
waterfall; 840–1241 m alt.; CS 8144, CS 8294 (JNU).
°Acrolejeunea pusilla (Steph.) Grolle et Gradst. – This species
was reported by Song & Yamamda (2009) but is absent the
collections available for us.
A. sandvicensis (Gottsche) Steph. – per.; on shaded dry rocks;
647–1667 m alt.; CS 3607, CS 9550, CS 110667 (JNU).
Anastrophyllum assimile (Mitt.) Steph. – on shaded cliffs; with
Bazzania parabidentula, Cheilolejeunea obtusifolia, Scapania ampliata; 1805–1820 m alt.; CS 3779, CS 3783, CS
111164 (JNU).
*Aneura pinguis (L.) Dumort. – on wet rocks; 685–1005 m
alt.; CS 3932, CS 7368, CS 8187 (JNU).
Apopellia endiviifolia (Dicks.) Nebel et D. Quandt – on wet
soil; with Calypogeia tosana, Heteroscyphus coalitus; 685–
1667 m alt.; CS 3432, CS 9131 (JNU).
*Asterella leptophylla (Mont.) Grolle – arch.; on shaded wet
cliffs; 860 m alt.; CS 3675, CS 3676 (JNU).
Bazzania denudata (Lindenb. et Gottsche) Trevis. – on humus
and decaying wood; with Fuscocephaloziopsis lunulifolia,
Cololejeunea macounii, Scapania ampliata; 840–1840 m alt.;
CS 3750, CS 7382, CS 111096 (JNU).
*B. imbricata(Mitt.) S. Hatt. – on humus and decaying wood;
1000–1840 m alt.; CS 1911001,CS 111096 (JNU).
*B. parabidentula Bakalin – on humus and bark of Quercus
mongolica; with Bazzania denudata, Scapania ciliata, Tetralophozia filiformis; 840–1820 m alt.; CS 3550, CS 8282,
CS 111158 (JNU).
B. tricrenata (Wahlenb.) Lindb. – on humus and decaying wood;
with Diplophyllum taxifolium, Scapania ciliata; 1421–1841
m alt.; CS 3571, CS 8267, CS 111081 (JNU).
*B. trilobata (L.) Gray – on humus and decaying wood; with
Douinia plicata; 840 m alt.; CS 8159 (JNU).
Blepharostoma minor Horik. – on wet rocks; with Herbertus
aduncus; 840–1421 m alt.; CS 3559, CS 3686, CS 6023, CS
8168, CS 8193 (JNU).
B. trichophyllum (L.) Dumort. – on humus and decaying wood;
with Fuscocephaloziopsis lunulifolia, Syzygiella autumnalis, Tritomaria exsecta; 840–1841 m alt.; CS 3798, CS 3833,
CS 7269, CS 8164, CS 8263 (JNU).
Calycularia laxa Lindb. et Arnell – on humus and shaded rocks
covered with soil; 848–1840 m alt.; CS 3534, CS 8226, CS
111165 (JNU).
Calypogeia angusta Steph. – on decaying wood; 1134 m alt.;
CS 3689 (JNU).
C. arguta Nees et Mont. – on shaded soil near roadsides; with
Cephalozia otaruensis; 749–1300 m alt.; CS 3968, CS 6047
(JNU).
C. orientalis Buczkowska et Bakalin – on humus and decaying
wood; 1134–1820 m alt.; CS 3685, CS 3801 (JNU).
*C. tosana (Steph.) Steph. – on shaded soil; with Cephalozia
otaruensis; 840–1667 m alt.; CS 3560, CS 9135 (JNU).
*Cephalozia bicuspidata (L.) Dumort. – on shaded soil; 1202–
1805 m alt.; CS 4230, CS 111074 (JNU).
C. otaruensis Steph. – on shaded rocks and humus ; 860–1840
m alt.; CS 3677, CS 8237, CS 111120 (JNU).
*Cephaloziella divaricata (Sm.) Schiffn. – on shaded rocks;
1820 m alt.; CS 3786 (JNU).
*C. hampeana (Nees) Schiffn. ex Loeske – ant., per.; on shaded rocks; with Syzygiella autumnalis; 840–1711 m alt.; CS
4236, CS 8167, CS 8168 (JNU).
C. massalogi (Spruce) Müll. Frib. – on shaded rocks; 1620 m
alt.; CS 3748 (JNU).
*C. microphylla (Steph.) Douin – spor.; on shaded soil near
roadside; 511–1540 m alt.; CS 3726, CS 6002 (JNU).
C. spinicaulis Douin – on soil and wet rocks; 144–1835 m alt.;
CS 3601, CS 111087, CS 111157 (JNU).
*C. spinigera (Lindb.) Jørg. – on shaded rocks; with Syzygiella autumnalis, Microlejeunea ulicina; 1840 m alt.; CS 8232
(JNU).
*C. varians (Gottsche) Steph. – on shaded rocks; 1713 m alt.;
CS 111062b, CS 111070b (JNU).
Cheilolejeunea obtusifolia (Steph.) S. Hatt. – on wet rocks and
shaded rocks; 685–1667 m alt.; CS 3629, CS 7372, CS 111046
(JNU).
Chiastocaulon mayebarae (S. Hatt.) S.D.F.Patzak, M.A.M.
Renner, Schäf.-Verw. & Heinrichs – on rocks covered thin
soil near the stream; 1667 m alt.; CS 9132 (JNU).
*Chiloscyphus pallescens (Ehrh.) Dumort. – on wet rocks; 969–
1667 m alt.; CS 3553, CS 9322 (JNU).
C. polyanthos (L.) Corda – on wet rocks; 1427 m alt.; CS 3533,
CS 8273, CS 111126 (JNU).
*Cololejeunea longifolia (Mitt.) Benedix ex Mizut.– on shaded rocks; 685–1667 m alt.; CS 3696, CS 7357, CS 9127 (JNU).
*C. macounii (Spruce) A. Evans – on shaded rocks; 860–1808
m alt.; CS 3681, CS 8248 (JNU).
*Conocephalum japonicum (Thunb.) Grolle – on shaded soil
and rocks; 749–781 m alt.; CS 3971, CS 3991 (JNU).
*C. salebrosum Szweyk., Buczk. et Odrzyk. – ant.; on shaded
rocks; 511–1427 m alt.; CS 3613, CS 8152 (JNU).
°Cryptolophocolea compacta (Mitt.) L. Söderstr. – This species was reported by Hattori et al. (1962) but is absent the
collections available for us.
*Cylindrocolea kiaeri (Austin) Váňa – on shaded rocks; with
Liverwort Flora of the Jirisan National Park in Korea
Herbertus aduncus; 848–1200 m alt.; CS 1910844 (JNU).
*C. recurvifolia (Steph.) Inoue – on shaded wet rocks; with
Cheilolejeunea obtusifolia, Plicanthus birmensis; 848 m alt.;
CS 3634, CS 3635 (JNU).
Diplophyllum albicans (L.) Dumort. – on humus layer over
rocks; with Anastrophyllum assimile, Blepharostoma trichophyllum, Cephaloziella hampeana, Odontoschisma grosseverrucosum, Scapania irrigua; 840–1841 m alt.; CS 3521,
CS 8271, CS 111148 (JNU).
*D. taxifolium (Wahlenb.) Dumort. – on humus and shaded
rocks; 1667–1820 m alt.; CS 3836, CS 9126 (JNU).
Douinia plicata (Lindb.) Konstant. et Vilnet – per., spor.; on
shaded rocks and shaded humus; 904–1820 m alt.; CS 3791,
CS 8249, CS 111166 (JNU).
*Drepanolejeunea angustifolia (Mitt.) Grolle – on shaded rocks;
with Frullania appendiculata; 1793 m alt.; CS 8214 (JNU).
Frullania appendiculata Steph. – on bark of tree, decaying
wood, dry rocks and shaded rocks; with Bazzania denudata,
Metzgeria lindbergii, Metzgeria temperata, Nipponolejeunea pilifera, Scapania ampliata; 658–1840 m alt.; CS 6026,
CS 9890, CS 111159 (JNU).
F. davurica Hampe ex Gottsche, Lindenb. et Nees – ant., per.;
on bark of Quercus mongolicaand decaying wood; with Frullania appendiculata; 658–820 m alt.; CS 4219, CS 7374, CS
7537, CS 8121 (JNU).
F. densiloba Steph. ex A. Evans – on dry rock; with Microlejeunea ulicina; 578–1585 m; CS 3640, CS 3825, CS 6011
(JNU).
F. diversitexta Steph. – on dry rocks and shaded rocks; 658–
860 m; CS 3608, CS 3625, CS 3639, CS 3643, CS 7267, CS
8114 (JNU).
°F. ericoides (Nees) Mont. – This species was reported by
Song & Yamada (2009) but is absent the collections available for us.
°F. fauriana Steph. – This species was reported by Uno and
Takahashi (1940), Hong (1960), Song and Yamada (2009)
but is absent the collections available for us.
F. hamatiloba Steph. – on bark of Malus sieboldii and bark of
Fraxinus sieboldiana; 1337 m alt.; CS 7545, CS 7546 (JNU).
*F. inflata Gottsch – on bark of tree; 700–800 m alt.; CS
1910127 (JNU).
F. kagoshimensis Steph. – per.; on bark of tree; with Frullania
appendiculata, Microlejeunea ulicina; 842–1820 m alt.; CS
3796, CS 7561 (JNU).
F. muscicola Steph. – per.; on dry rock and bark of Carpinus
laxiflora; 578–1585 m alt.; CS 3603, CS 111145 (JNU).
°F. parvistipula Steph. – This species was reported by Song &
Yamada (2009) but is absent the collections available for us.
*F. pedicellata Steph. – per.; on bark of Quercus mongolica;
with Frullania kagoshimensis; 1202 m alt.; CS 4208 (JNU).
*F. polyptera Taylor – on bark of Quercus mongolica; with
Acrolejeunea sandvicensis; 658–1202 m alt.; CS 4205, CS
8108 (JNU).
F. schensiana C. Massal. – on bark of Quercus mongolica; with
Frullania muscicola; 1000–1320 m alt.; CS 1910651 (JNU).
F. taradakensis Steph. – on bark of trees; 685–1337 m; CS
7375, CS 7536,CS 7548 (JNU).
F. usamiensis Steph. – per.; on bark of Quercus mongolica;
with Frullania muscicola; 685–1820 m alt.; CS 3767, CS
7543, CS 7544(JNU).
*Fuscocephaloziopsis catenulata (Huebener) Váňa & L. Söderstr. subsp. nipponica (S. Hatt.) Váňa et L. Söderstr. – decay-
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ing wood; with Metacalypogeia cordifolia, Schistochilopsis
cornuta; 969–1841 m alt.; CS 7380, CS 8265 (JNU).
F. leucantha (Spruce) Váňa et L.Söderstr. – on decaying wood;
with Fuscocephaloziopsis lunulifolia; 1808–1841 m alt.; CS
8260, CS 8266 (JNU).
*F. lunulifolia (Dumort.) Váňa et L.Söderstr. – on decaying
wood; with Blepharostoma trichophyllum, Calypogeia arguta, Cephalozia bicuspidata, Kurzia makinoana; 969–1835
m alt.; CS 3838, CS 4230, CS 8188, CS 111069, CS 111090
(JNU).
*Gymnomitrion commutatum (Limpr.) Schiffn. – on shaded
rocks; with Anastrophyllum assimile, Cephaloziella spinicaulis, Diplophyllum albicans, Microlejeunea ulicina, Tetralophozia filiformis; 1820 m alt.; CS 3778, CS 3827 (JNU).
G. parvitextum (Steph.) Mamontov, Konstant. et Potemkin. –
on shaded rocks; 1820 m alt.; CS 3816 (JNU).
°Harpanthus flotovianus (Nees) Nees – This species was reported by Choe (1980) but is absent the collections available for us.
H. scutatus (F. Weber et D. Mohr) Spruce – on decaying wood
in broad-leaved forest with admixture of conifers and shrubs;
with Tritomaria exsecta; 1720 m alt.; CS 3763a (JNU).
*Hattorianthus erimonus (Steph.) R.M.Schust. et Inoue – on
wet humus near stream; 685 m alt.; CS 7352, CS 7360 (JNU).
Herbertus aduncus (Dicks.) Gray – on shaded rocks; with Nipponolejeunea pilifera; 658–1840 m alt.; CS 6015, CS 111088,
CS 111146 (JNU).
*H. buchii Juslén – on shaded rocks; 1317 m alt.; CS 7556
(JNU).
*H. dicranus (Gottsche, Lindenb. et Nees) Trevis. – on shaded
rocks; 1500–1820 m alt.; CS 3728, CS 3813, CS 3850 (JNU).
*Heteroscyphus coalitus (Hook.) Schiffn. – on wet soil; 842 m
alt.; CS 6019 (JNU).
*H. planus (Mitt.) Schiffn. – on wet rocks covered soil; with
Calypogeia tosana, Solenostoma pyriflorum, Trichocoleopsi ssacculata; 840 m alt.; CS 8148 (JNU).
*Jubula hutchinsiae (Hook.) Dumort. subsp. javanica (Steph.)
Verd. – on wet rocks; 860–1421 m alt.; CS 3550, CS 3665
(JNU).
*J. hutchinsiae (Hook.) Dumort. subsp. japonica (Steph.) Horik.
et Ando – on wet rocks near the stream; 1241 m alt.; CS
8276 (JNU).
*Jungermannia atrovirens Dumort. – on wet rocks; 1118–1440
m alt.; CS 3679, CS 3716, CS 110665 (JNU).
*J. exsertifolia Steph. – on wet rocks; 578 m alt.; CS 3900
(JNU).
*Kurzia makinoana (Steph.) Grolle – on rocks covered with
soil; with Blepharostoma minus, Cephaloziella hampeana,
Herbertus aduncus; 820–1820 m alt.; CS 3782, CS 8125
(JNU).
Lejeunea compacta (Steph.) Steph. – on humus; with Lejeunea
parva, Lophocolea minor; 658 m alt.; CS 8112 (JNU).
L. japonica Mitt. – per., spor.; on wet rocks; with Chiloscyphus polyanthus, Dicranolejeunea yoshinagana; 647–1667
m alt.; CS 3621, CS 9948, CS 110660 (JNU).
*L. parva (S. Hatt.) Mizut. – on dry rocks; with Cephaloziella
spinicaulis; 658–1427 m alt.; CS 3604, CS 6007, CS 8117
(JNU).
*Lepidozia reptans (L.) Dumort. – on decaying wood; with
Fuscocephaloziopsis leucantha, Fuscocephaloziopsis lunulifolia, Cephalozia otaruensis, Metacalypogeia cordifolia, Mylia
verrucosa, Riccardia multifida subsp. decrescens, Xenochila
integrifolia; 1118–1841 m alt.; CS 8270, CS 111167 (JNU).
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H.M. BUM, V.A. BAKALIN, S.J. PARK, S.S. CHOI, C.H. KIM & S. SO
*L. subtransversa Steph. – on rocks covered with humus; 840–
1808 m alt.; CS 6077, CS 8138, CS 8255 (JNU).
*L. vitrea Steph. – on decaying wood; 1713 m alt.; CS 111066
(JNU).
°Liochlaena subulata (A. Evans) Schljakov – This species was
reported by Hattori et al. (1962) but is absent the collections
available for us.
*Lophocolea heterophylla (Schrad.) Dumort. – on decaying
wood and wet rocks; 685–1720 m alt.; CS 3552, CS 7371
(JNU).
*L. horikawana S. Hatt. – on shaded rocks; 842–1720 m alt.;
CS 3749, CS 8183, CS 111064 (JNU).
L. minor Nees – gemm.; on shaded rocks; 144–1134 m alt.; CS
3623, CS 8172 (JNU).
*Lophozia guttulata (Lindb. et Arnell) A. Evans – on decaying
wood; with Calypogeia tosana, Tritomaria exsecta; 1820–
1841 m alt.; CS 3798, CS 8265 (JNU).
L. koreana (Bakalin, S.S. Choi et B.-Y. Sun) Maltseva, Vilnet
et Bakalin – on shaded cliffs; with Bazzania denudata, Fuscocephaloziopsis lunulifolia, Douinia plicatum, Scapania
ampliata; 1808–1840 m alt.; CS 8225, CS8254 (JNU).
*Lophoziopsis excisa (Dicks.) Konstant. et Vilnet – on decaying wood; 1000–1134 m alt.; CS 1910198 (JNU).
*Makinoa crispata (Steph.) Miyake – on wet humus; 685–1421
m alt.; CS 3552, CS 7350, CS 7359 (JNU).
°Marchantia paleacea Bertol. – This species was reported by
Song & Yamada (2009), but is absent in our collections.
*M. paleacea Bertol. subsp. diptera (Nees et Mont.) Inoue –
on shaded rocks; 1427 m alt.; CS 5035, CS 5040 (JNU).
M. polymorpha L. subsp. ruderalis Bischl. et Boissel.-Dub. –
gemm.; on shaded soil; 1427 m alt.; CS 5041 (JNU).
Marsupella apertifolia Steph. – per.; on shaded rocks; 1300–
1720 m alt.; CS 3766, CS 6090 (JNU).
M. koreana Bakalin et Fedosov – on cliffs; 1440–1500 m alt.;
CS3058 (JNU).
M. pseudofunckii S.Hatt. – ant.; on shaded rocks; 1440–1720
m alt.; CS 3717, CS 40866 (JNU).
M. tubulosa Steph. – on wet rocks; with Cephaloziella hampeana, Lejeunea parva, Trichocoleopsis sacculata, Tritomaria exsecta; 840–1820 m alt.; CS 3641, CS 8172, CS 111147
(JNU).
M. yakushimensis (Horik.) S.Hatt. – on shaded rocks; 840–
1805 m alt.; CS 3628, CS 7278, CS 111166 (JNU).
*Metacalypogeia cordifolia (Steph.) Inoue – on soil and wet
rocks; with Bazzania denudata, Blepharostoma trichophyllum, Fuscocephaloziopsis catenulata subsp. nipponica, Lepidozia reptans, Tritomaria exsecta; 658–1840 m alt.; CS 3602,
CS 8245, CS 111107 (JNU).
Metzgeria furcata (L.) Corda – on rocks, bark of trees and wet
rocks; with Lejeunea parva; 860–1808 m alt.; CS 3682, CS
8277 (JNU).
M. lindbergii Schiffn. – on bark of trees; 685 m alt.; CS 7354,
CS 7370 (JNU).
M. pubescens (Schrank) Raddi – on shaded humus; 578–1835
m alt.; CS 3917, CS 111091 (JNU).
M. temperata Kuwah. – on rocks; with Cheilolejeunea obtusifolia; 848–1835 m alt.; CS 8197, CS 111085 (JNU).
*Microlejeunea ulicina (Taylor) Steph. – on base of Abies koreana; with Syzygiella autumnalis; 720–1700 m alt.; CS 3605,
CS 7547 (JNU).
*Mylia taylorii (Hook.) Gray – on humus; 1840–1841 m alt.;
CS 8233, CS 8238, CS 8264 (JNU).
*M. verrucosa Lindb. – on decaying wood; with Bazzania denudata, Bazzania trilobata, Fuscocephaloziopsis catenulata
subsp. nipponica; 840–1841 m alt.; CS 6049, CS 6051, CS
8269 (JNU).
Nardia assamica (Mitt.) Amakawa – on shaded soil near road;
625–1711 m alt.; CS 4217, CS 7571 (JNU).
Neotrichocolea bissetii (Mitt.) S.Hatt. – on wet rocks; with
Scapania undulata, Trichocoleopsis sacculata; 996–1793 m
alt.; CS 6059, CS 8295 (JNU).
Nipponolejeunea pilifera (Steph.) S.Hatt. – on bark of Abies
holophylla and shaded dry rocks; with Frullania appendiculata, Frullania schensiana, Plagiochila porelloides, Radula
cavifolia; 1202–1915 m alt.; CS 3573, CS 8231, CS 111086
(JNU).
Nowellia curvifolia (Dicks.) Mitt. – on decaying wood near
the stream; with Cephalozia otaruensis, Odontoschisma denudatum; 820–1241 m alt.; CS 8123, CS 8288 (JNU).
*Odontoschisma denudatum (Nees) Dumort. – on decaying
wood; 658 m alt.; CS 8115 (JNU).
*O. pseudogrosseverrucosum Gradst., S.C.Aranda et Vanderp.
– on decaying wood and shaded rocks; with Fuscocephaloziopsis lunulifolia, Syzygiella autumnalis, Diplophyllum taxifolium, Marsupella pseudofunkii, Solenostoma pyriflorum;
1317–1915 m alt.; CS 3831, CS 8219 (JNU).
*Pedinophyllum truncatum (Steph.) Inoue – per.; on humus
and wet rocks; 511–842 m alt.; CS 3608, CS 7338 (JNU).
*Pellia neesiana (Gottsche) Limpr. – on wet rocks; 842–1327
m alt.; CS 3694, CS 111132 (JNU).
Plagiochila gracilis Lindenb. et Gottsche – on shaded rocks;
1840 m alt.; CS 8234 (JNU).
*P. hakkodensis Steph. – on shaded rocks; 1820 m alt.; CS
3818 (JNU).
P. ovalifolia Mitt. – ant., per.; on shaded wet rocks; with Cololejeunea macounii; 647–1667 m alt.; CS 3615, CS 8153, CS
111130 (JNU).
°P. parvifolia Lindenb. – This species was reported by Song &
Yamada (2009) but is absent the collections available for us.
P. porelloides (Torr. ex Nees) Lindenb. – on wet rocks; 685–
1319 m alt.; CS 3680, CS 8202 (JNU).
P. sciophila Nees – on shaded rocks; 860–1134 m alt.; CS 3660,
CS 110664(JNU).
P. semidecurrens (Lehm. et Lindenb.) Lindenb. – on shaded
rocks; 1700–1800 m alt.; VB Kor-75-2-19 (VBGI).
*P. trabeculata Steph. – on shaded rocks; 1700–1820 m alt.;
CS 3811, CS 3831 (JNU).
Plectocolea erecta Amakawa – per.; on shaded rocks near the
stream; 904–1300 m alt.; CS 6066, CS 8192 (JNU).
P. infusca Mitt. – per.; on wet rocks covered thin soil; 689–
1222 m alt.; CS 3515, CS 3919 (JNU).
*P. infusca Mitt. var. recondita Bakalin – on wet rocks covered
with thin soil; 1202–1300 m alt.; CS 3855, CS 4247 (JNU).
P. ovalifolia (Amakawa) Bakalin et Vilnet – on wet rocks covered with thin soil; 1134–1222 m alt.; CS 3702a, CS 3522
(JNU).
P. rosulans (Steph.) S.Hatt. – on wet rocks covered with thin
soil layer; wih Radula constricta, Scapania undulata; 685–
1319 m alt.; CS 3618, CS 8289 (JNU).
P. torticalyx (Steph.) S.Hatt. – on wet rocks near the stream in
broad-leaved forest; 1300–1900 m alt.; CS 3093, CS 3856 (JNU).
P. virgata Mitt. – on wet rocks covered with thin soil layer;
860–1134 m alt.; CS 3659, CS 3697 (JNU).
Plicanthus birmensis (Steph.) R.M.Schust. – on shaded rocks;
Liverwort Flora of the Jirisan National Park in Korea
with Cephaloziella spinigera; 658–1840 m alt.; CS 3636,
CS 3644, CS 8235 (JNU).
*Porella acutifolia (Lehm. & Lindenb.) Trevis. subsp. tosana
(Steph.) S. Hatt. – on shaded rocks; 860 m alt.; CS 7270
(JNU).
P. caespitans (Steph.) S. Hatt. var. cordifolia (Steph.) S. Hatt.
ex T.Katag. et T.Yamag. – per.; on shaded rocks; 511–1100
m alt.; CS 3876, CS 3876, CS 7393, CS 8136 (JNU).
*P. chinensis (Steph.) S. Hatt. – per.; on shaded rocks; 1000–
1100 m alt.; CS 1910624 (JNU).
P. faurieri (Steph.) S. Hatt. – on shaded rocks; 1700 m alt.; CS
3841 (JNU).
P. grandiloba Lindb. – on shaded rocks; 511–860 m alt.; CS
3884, CS 7369 (JNU).
P. ulophylla (Steph.) S. Hatt. – on bark of tree; 647–1700 m
alt.; CS 3840, CS 7563, CS 110666 (JNU).
°P. japonica (Sande Lac.) Mitt. – This species was reported by
Song & Yamada (2009) but is absent the collections available for us.
P. vernicosa Lindb. – on shaded rocks and bark of trees; 860–
1711 m alt.; CS 3553, CS 7271, CS 7562 (JNU).
Protosolenostoma fusiforme (Steph.) Vilnet et Bakalin – 1421
m alt.; CS5042 (JNU).
*Ptilidium pulcherrimum (Weber) Vain. – spor.; on bark of tree;
with Cephaloziella spinigera, Scapania ampliata; 1820–1840
m alt.; CS 3802, CS 8242 (JNU).
Radula auriculata Steph. – on shaded rocks; 685–1820 m alt.;
CS 3800, CS 7358 (JNU).
R. constricta Steph. – on bark of trees; 1317–1421 m alt.; CS
3551, CS 7551 (JNU).
R. japonica Gottsche – on wet rocks; 647–1134 m alt.; CS 3703,
CS 110661 (JNU).
°R. obtusiloba Steph. – This species was reported by Song &
Yamada (2009) but is absent the collections available for us.
°R. oyamensis Steph. – This species was reported by Song &
Yamada (2009) but is absent the collections available for us.
*R. tokiensis Steph. – on shaded rocks; with Lejeunea japonica, Metzgeria lindbergii; 685 m alt.; CS 7340 (JNU).
Reboulia hemisphaerica (L.) Raddi – on shaded rocks; 749–
1320 m alt.; CS 3709, CS 3967, CS 3973 (JNU).
°Riccardia chamedryfolia (With.) Grolle – This species was
reported by Choe (1980) but is absent the collections available for us.
*R. multifida subsp. decrescens (Steph.) Furuki – on decaying
wood; 1300 m alt.; CS 6070 (JNU).
*R. palmata (Hedw.) Carruth. – on decaying wood; with Cephalozia sp.; 1241–1820 m alt.; CS 3810, CS 8287 (JNU).
*Scapania ampliata Steph. – on wet rocks; with Bazzania denudata, Cephaloziella spinigera, Solenostoma pyriflorum;
860–1840 m alt.; CS 3688, CS 8234, CS 111152 (JNU).
*S. apiculata Spruce – on humus; 1427 m alt.; CS 5038 (JNU).
*S. ciliata Sande Lac. – on decaying wood; 749–1667 m alt.;
CS 3963, CS 9134 (JNU).
°S. curta (Mart.) Dumort. – This species was reported by Hattori et al. (1962) but is absent the collections available for us.
S. integerrima Steph. – on wet rocks; with Plectocolea rosulans; 842–1540 m alt.; CS 3741, CS 8192 (JNU).
*S. irrigua (Nees) Nees – on wet rocks near the stream; with
Metacalypogeia cordifolia; 1134–1805 m alt.; CS 3690, CS
111162 (JNU).
*S. paludosa (Müll. Frib.) Müll. Frib. – on wet soil; with Makinoa crispata; 720–1319 m alt.; CS 3614, CS 8209 (JNU).
*S. parvitexta Steph. – on shaded rocks; with Scapania irri-
413
gua, Scapania integerrima; 685–1840 m alt.; CS 3609, CS
8231 (JNU).
*S. subalpina (Nees ex Lindenb.) Dumort. – gemm.; on wet
rocks; 1000–1385 m alt.; CS 1910732 (JNU).
S. undulata (L.) Dumort. – gemm.; on wet rocks; 658–1585 m
alt.; CS 3555, CS 8292, CS 111144 (JNU).
*Schistochilopsis cornuta (Steph.) Konstant. – on decaying
wood; with Bazzania parabidentula, Bazzania tricrenata,
Blepharostoma trichophyllum, Fuscocephaloziopsis catenulata ssp. nipponica, Fuscocephaloziopsis lunulifolia, Douinia
plicata; 1100–1840 m alt.; CS 3812, CS 8256, CS 111151
(JNU).
Solenostoma bilobum (S. Hatt. ex Amakawa) Potemkin et Nyushko – 1729 m alt.; CS3762 (JNU).
S. cyclops (S. Hatt.) R.M. Schust. – on shaded wet rocks along
stream; 860–1319 m alt.; CS 7279, CS 8204 (JNU).
S. jirisanense Bakalin et S.S. Choi – on shaded rocks covered
thin soil; 1500–1800 m alt.; CS 3747-3(JNU), VB Kor-25-515 (VBGI).
S. pyriflorum Steph. – on cliff, wet rocks and shaded rocks;
with Scapania integerrima; 1440–1915 m alt.; CS 3710, CS
6085, CS 8280 (JNU).
S. rotundatum Amakawa – on wet rocks covered thin soil; 848
m alt.; CS 3658 (JNU).
S. sunii Bakalin et Vilnet – on shaded rocks covered with thin
soil layer; 848–1720 m alt.; CS 3653, CS3759 (JNU).
*Sphenolobus saxicola (Schrad.) Steph. – on shaded rocks in
stony field; 904 m alt.; CS 8170 (JNU).
Syzygiella autumnalis (DC.) K. Feldberg, Váňa, Hentschel et
Heinrichs – spor.; on wet rocks and dry rocks; with Bazzania
parabidentula, Cephaloziella spinigera, Diplophyllum taxifolium, Metacalypogeia cordifolia, Plagiochila gracilis, Ptilidium pulcherrimum, Tritomaria exsecta; 685–1915 m alt.;
CS 3561, CS 9129, CS 111114 (JNU).
*Tetralophozia filiformis (Steph.) Urmi – on shaded cliffs; with
Cephaloziella spinicaulis, Diplophyllum taxifolium, Herbertus aduncus, Odontoschisma pseudogrosseverrucosum; 840–
1835m alt.; CS 3684, CS 8171, CS 111058 (JNU).
*Trichocolea tomentella (Ehrh.) Dumort. – on shaded humus;
with Calypogeia tosana, Cephalozia otaruensis, Metacalypogeia cordifolia, Schistochilopsis cornuta; 1118 m alt.; CS
6036, CS 6037, CS 6039 (JNU).
Trichocoleopsis sacculata (Mitt.) S. Okamura – on rocks; 511–
1805 m alt.; CS 3524, CS 7344, CS 111160 (JNU).
*Trilophozia quinquedentata (Huds.) Bakalin – on shaded
rocks; with Lophozia koreana, Scapania ampliata, Schistochilopsis cornuta; 1808 m alt.; CS 6561 (JNU).
Tritomaria exsecta (Schmidel) Schiffn. ex Loeske – on wet
rocks; with Bazzania denudata, Blepharostoma minus, Fuscocephaloziopsis lunulifolia, Scapania ampliata; 781–1808
m alt.; CS 3751, CS 8251 (JNU).
*Xenochila integrifolia (Mitt.) Inoue – on shaded wet rocks
along stream; 685 m alt.; CS 7342, CS 7362, CS 7364 (JNU).
DISCUSSION
Based on the analysis of 2500 herbarium specimens
and a review of data in the literature, we found that the
bryophyte flora of the Jirisan Mountains includes 177
taxa, of which 78 species are newly records for the park.
There were 14 species recorded in the literature that had
no corresponding herbarium specimen. The species Lopholejeunea subfusca was excluded from the list of taxa
because, although it was reported on the Jirisan Moun-
414
H.M. BUM, V.A. BAKALIN, S.J. PARK, S.S. CHOI, C.H. KIM & S. SO
Table 1. The latitude fractional composition in the hepatics
of the Jirisan National Park in Korea; The treatment of floristic
elements (latitudinal types) follows to Konstantinova (2000)
and Bakalin (2010).
Floristic elements
(latitudinal types)
Arctic
Arctic-boreal
Boreal
Boreal-temperate
Temperate
Temperate-subtropical
Subtropical
Subtropical-tropical
Multizone
Total
Taxa
Ratio (%)
12
25
18
19
48
31
13
4
7
177
6.8
14.1
10.2
10.7
27.1
17.5
7.3
2.3
4.0
100
tains by Hong (1962), its distribution is limited to tropical and subtropical areas, so its occurrence on the Jirisan
mountains is unlikey (Choi et al, 2021). The plants observed by Hong (1962) most likely belonged to Acanthocoleus yoshinaganus. In recent research on the Jirisan
Mountain’s liverworts, Lophozia koreana (Fig. 3D),
Marsupella koreana (Fig. 3E), and Solenostoma jirisanense (Fig. 3F) were recorded as being new to science
(Bakalin et al., 2009, 2019, 2020, 2021b). Lophozia koreana has subsequently been observed in Japan (Bakalin, 2016), Marsupella koreana has been discovered in
Daedunsan, Gyaysan, Namdeogyusan, Dureunsan, and
Hallasan Mountains in Korea (Bakalin et al., 2021b).
In the analysis of longitudinal distribution characteristics of liverwors, 89 taxa (50.3%) were East Asian, 49
taxa (27.7%) were circumpolar, nine taxa (5.1%) were
widespread in Asia, eight taxa (4.5%) were amphioceanic and seven taxa (4.0%) were amphipacific (Table 1).
This result is consistent with the Korean Peninsula belonging to the Eastern Asia floristic region defined by
Takhtajan (1986). A latitudinal analysis of floristic elements found, 48 taxa (27.1%) were temperate, 31 taxa
(17.5%) were temperate-subtropical, 25 taxa (14.1%)
were arctic-boreal, 19 taxa (10.7%) were boreal-temperate, 18 taxa (10.2%) were boreal, 13 taxa (7.3%) were
Table 2. The longitudinal fractional composition in the
Jirisan National Park in Korea; floristic elements (longitudinal
types) follows Konstantinova (2000) and Bakalin (2010).
Area types
(longitudinal types)
Amphioceanic
Amphipacific
Asia
Asian-American
Asian-Euro
Circumpolar
Disjunctive
Eastern Asia
Eastern Asia-American
Multiarea
Total
Taxa
Ratio (%)
8
7
9
4
2
49
1
89
1
7
177
4.5
4.0
5.1
2.3
1.1
27.7
0.6
50.6
0.6
4.0
100
Table 3. Altitudinal distribution of all taxa in the Jirisan
National Park.
Altitude, m
0–500
500–1000
1000–1500
1500–1917
Species
58
114
131
94
Ratio (%)
32.7
64.4
74.0
53.1
subtropical and 12 (6.8%) were arctic species (Table 2).
The following data on species diversity may show at
least roughly a pattern of liverwort distribution along the
altitude. As expected, high elevations of over 1000 m
a.s.l. comprise the maximal diversity in this area (Table
3). This result is because the altitudes of the Jirisan Mountains ridge are between 1000 and 1500 m a.s.l.
Neotrichocolea bissetii, Lepidozia subtransversa and
Nipponolejeunea pilifera (Fig. 3A, B, G) are rare Korean species recorded from the Jirisan Mountains. Neotrichocolea bissetii is endemic to East Asia, and is distributed in China (Anhui, Fujian, Yunnan, Zhejing Provinces) and Japan (Honshu, Shikoku, and Kyushu Islands).
In Korea, it is recorded only in the Jirisan Mountains
(Choi et al., 2012). We discovered this species on wet
rock near a waterfall on humic soil near N-facing slope
near the top of the mountain in broad-leaved forest with
Oplopanax elatus (Nakai) Nakai, Acer sp., Quercus sp.,
and a Abies koreana forest. Lepidozia subtransversa is
restricted to temperate regions of East Asia, including
the Korean Peninsula, Japan, China and the southernmost of the Russian Far East (Yamada & Iwatsuki, 2006;
Piippo, 1990; Choi & Bakalin, 2012). In Korea, it was
previously recorded only in the Jirisan and Seolaksan
Mountains. We recorded the species on humus covered
rocks, decaying wood near northwestern slopes of a top
area and Hansin Stream valley. Nipponolejeunea pilifera
represents mainly Japanese-Korean oro-hemiboreal to
oro-temperate endemic, known from Hokkaido to Kyushu in Japan, Korea, Taiwan, and northwardly known
in southern Kurils (Piippo 1990; Yamada & Iwatsuki,
2006; Bakalin, 2019). In Korea, it was previously recorded from Deogyusan, Gayasan, Ullengdo Island
(Seonginbong), and the Jirisan Mountains. This species
grows in coniferous forests (Abies koreana, A. nephrolepis), along with Frullania appendiculata, F. usamienesis, Radula constricta and on shaded cliffs together with
Scapania ampliata, Anastrophyllum assimile, and Marsupella yakushimensis.
We found the most significant area for liverwort diversity in the Jirisan Mountains – the plant communities
of Abies koreana, A. nephrolepis, and Oplopanax elatus
located near the northern slope of Cheonwangbong Peak
(1915 m a.s.l.), the highest peak of the Jirisan Mountain
(Fig. 2A–B). In this area, we found Lophozia koreana,
Anastrophyllum assimile, Nipponolejeunea pilifera, Neotrichocolea bissetii, Douinia plicata, and Schistochilopsis cornuta, and some species that are typically found in
several subalpine regions of the Korean Peninsula. An-
Liverwort Flora of the Jirisan National Park in Korea
415
Fig. 3. Some main liverworts of the Jirisan National Park in Korea. A. Neotrichocolea bissetii. B. Lepidozia subtransversa. C.
Schistochilopsis cornuta. D. Lophozia koreana. E. Marsupella koreana. F. Solenostoma jirisanense. G. Nipponolejeunea pilifera.
H. Makinoa crispata. I. Hattorianthus erimonus. Photos: S.S. Choi.
416
H.M. BUM, V.A. BAKALIN, S.J. PARK, S.S. CHOI, C.H. KIM & S. SO
other area is the upper half of the Hansin Valley, which
runs from Seseok Ridge to Hanshin Valley (Fig. 2D).
Waterfalls and rock fields on the north slopes of the
Seseok Ridge are common. There we found Lepidozia
subtransversa, Plectocolea rosulans, Scapania ciliata,
Bazzania imbricata, Neotrichocolea bissetii, Scapania
undulata, and some additional species recorded from
subalpine areas of the Korean Peninsula.
ACKNOWLEDGMENTS
This work was supported by a grant from the 5th National Ecosystem Survey of National Institute of Ecology
(NIE), funded by the Ministry of Environment (MOE) of
the Republic of Korea (NIE-A-2021-01). Also, this work
was partially supported by a grant from “Natural resource
survey in the Jirisan National Park in 2019” of Korea
National Park Research Institute (no NPRI 2019-40). Bakalin also thanks Russian Foundation for Basic Research
(grant 20-04-00278) and Botanical Garden-Institute research project (no. AAAA-A-20-120031990012-4).
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Received 23 September 2021
Accepted 10 November 2021