Alpine Vegetation on the Paekdu-San (Changbaishan) Summit of the North-east China

Abstract

We aim to provide a syntaxonomical scheme for the alpine tundra vegetation of volcano Mount Paekdu (Changbaishan) bordering China and North Korea. From the field investigations in 1991, 2004, and 2009 a total of 89 phytosociological relevés were sampled by the Braun-Blanquet method. Obtained data sets were analyzed by tabular hand-sorting technique. Syntaxonomic position and nomenclature of units were determined by comparison with mainly Japanese relevant communities. Ten rankless units in four physiognomic vegetation types were classified: alpine desert vegetation (Papaver radicatum var. pseudoradicatum community, Minuartia arctica community, Polytrichastrum alpinum-Minuartia macrocarpa var. koreana community), alpine windswept dwarf vegetation (Chrysanthemum zawadskii var. alpinum-Festuca ovina var. koreano-alpina community, Scirpus maximowiczii community, Oxytropis anertii-Salix rotundifolia community, Carex longirostrata-Dryas octopetala var. asiatica community), snow-bed vegetation (Phyllodoce caerulea-Tofieldia coccinea community, Viola biflora-Calamagrostis langsdorfii community), and colluvial drainage channel vegetation (Carex atrata-Sanguisorba stipulata var. riishiriensis community). The variation of the vegetation resulted from primarily the rock and soil particle size and soil mobility, and slope inclination. Some of the plant communities were regarded as local and endemic types. Yet we firmly believe that further researches are requested by much richer materials and syntaxonomical comprehensive evaluation in between elsewhere in the Pan-East Sea (Sea of Japan).

Full text

J. Plant Biol. (2019) 62:436-450
DOI 10.1007/s12374-019-0305-x
Alpine Vegetation on the Paekdu-San (Changbaishan) Summit of the
North-east China
Jong Won Kim
1
, Sung Yeol Kim
2
and
Byoung Ki Choi
3,
*
1
Unit of Vegetation Science, School of Biological Sciences, University of Keimyung, Daegu 42601, Korea
2
Research Division, National Science Museum, 481 Daedeokdaero Yuseong-gu Daejeon 34143, Korea
3
Warm Temperate and Subtropical Forest Research Center, National Institute of Forest Science, 22 Donnaeko Rd. Seogwipo-
si, Jeju Special Self-governing Province, 63582, Korea
Received: July 30, 2019 / Accepted: October 14, 2019
© Korean Society of Plant Biologists 2019
Abstract We aim to provide a syntaxonomical scheme for
the alpine tundra vegetation of volcano Mount Paekdu
(Changbaishan) bordering China and North Korea. From the
field investigations in 1991, 2004, and 2009 a total of 89
phytosociological relevés were sampled by the Braun-Blanquet
method. Obtained data sets were analyzed by tabular hand-
sorting technique. Syntaxonomic position and nomenclature of
units were determined by comparison with mainly Japanese
relevant communities. Ten rankless units in four physiognomic
vegetation types were classified: alpine desert vegetation
(Papaver radicatum var. pseudoradicatum community,
Minuartia arctica community, Polytrichastrum alpinum-
Minuartia macrocarpa var. koreana community), alpine
windswept dwarf vegetation (Chrysanthemum zawadskii var.
alpinum-Festuca ovina var. koreano-alpina community, Scirpus
maximowiczii community, Oxytropis anertii-Salix rotundifolia
community, Carex longirostrata-Dryas octopetala var. asiatica
community), snow-bed vegetation (Phyllodoce caerulea-
Tofieldia coccinea community, Viola biflora-Calamagrostis
langsdorfii community), and colluvial drainage channel
vegetation (Carex atrata-Sanguisorba stipulata var. riishiriensis
community). The variation of the vegetation resulted from
primarily the rock and soil particle size and soil mobility, and
slope inclination. Some of the plant communities were regarded
as local and endemic types. Yet we firmly believe that further
researches are requested by much richer materials and
syntaxonomical comprehensive evaluation in between
elsewhere in the Pan-East Sea (Sea of Japan).
Keywords: Alpine tundra grassland, Paekdu-san, Rankless
unit, Zurich-Montpellier
Introduction
Mount Paekdu (Paekdu-San; Changbaishan in Chinese) is
the highest mountain in continental East Asia and a major
border between North Korea and northeastern China. As the
largest single volcano in the region the alpine belt of Paekdu-
San is an important refuge for glacial relict species (Qian et
al. 1999). It provides a habitat for 336 plant species, and its
bryophyte and lichen floras are remarkably rich (CNRI
1982). Due to their small population size, 48 phanerogams,
including the endemic Papaver radicatum var. pseudoradicatum,
have been protected by the Chinese Jiling Province (Yang
2009).
Although Paekdu-San has long been acknowledged as a
main research site on the biogeography of alpine plant
communities, its phytosociological composition has been little
investigated. An earlier research by Qian (1990) revealed three
TWINSPAN groups and 59 associations, and that the alpine
tundra plant communities depend primarily on the soil-water
gradient; however, this study did not investigate in terms of
phytosociological species composition. Although some
syntaxonomical names such as Caricetalia tenuiformis and
Festucetum ovinae koreanoalpinae from the alpine vegetation
on Paekdu-San of North Korea (Kim et al. 1991) were found,
they did not comply with the phytosociological standards.
Similarly, phytosociological studies established on the Chinese
slopes of Changbaishan did not use the conventional
syntaxonomic system (Qian 1992; Qian et al. 1999). To the
best of our knowledge, the first valid syntaxonomic unit
was the Dryado tschonoskii-Rhododendretum described by
Dostálek et al. (1988) from Paekdu-San of North Korea.
Despite the limited studies and poor understanding of the
plant community structure, Chang et al. (1992) hypothesized
that wind and soil moisture are major factors that determine
species composition of plant communities around the caldera
RESEARCH ARTICLE
*Corresponding author; Byoung Ki Choi
Tel : +82-64-730-7260
E-mail : vegetation01@korea.kr

J. Plant Biol. (2019) 62:436-450 437
lake Cheonji located atop the mountain. The altitudinal
changes in the local flora have been described for the slopes
below the summit of Cheungseuk-bong on Paekdu-San (Lee
2002), and another survey documented the altitudinal
distribution patterns of dominant species mainly along the
main tourist trail above 2,000 m elevation on the Chinese
side (Kil et al. 1998; Lee et al. 1998). The scarcity of
vegetation surveys seems to be due to the restrictions
imposed by Chinese local authorities. Field surveys by
Korean scientists particularly were bound by spatio-
temporal constraints. Before diplomatic relations between
China and South Korea, in 1991 we had investigated the
Chinese side of northern Paekdu-San without any guarantee of
personal security.
The present study aims to describe the phytosociological
characteristics of the alpine tundra vegetation mainly along
Tabl e 1 . Synoptic table of the alpine open tundra-like formation of Paekdu-san. A synoptic value is the net contribution degree (NCD)
combining a frequency and a coverage of species in a relevant unit. Syntaxa A: Alpine desert vegetation, A1: Papaver radicatum var.
pseudoradicatum community, A2: Minuartia arctica community, A3: Polytrichastrum alpinum-Minuartia macrocarpa var. koreana
community, B: Alpine windswept grassland, B1: Chrysanthemum zawadskii var. alpinum-Festuca ovina var. koreano-alpina community,
B2: Scirpus maximowiczii community, B3: Carex longirostrata-Dryas octopetala var. asiatica community, B4: Oxytropis anertii-Salix
rotundifolia community, C: Snowbed vegetation, C1: Phyllodoce caerulea-Tofieldia coccinea community, C2: Viola biflora-Calamagrostis
langsdorfii community, D: Colluvial drainage channel vegetation, Carex atrata-Sanguisorba stipulate var. riishiriensis community, D1: Luzula
oligantha subcommuniy, D2: typical subcommunity, D3: Ligularia jamesii subcommunity, r-NCD: Relative net contribution degree (%)
Vegetation type A B C D r-NCD
Syntaxa A1A2A3 B1B2B3B4 C1C2 D1D2D3
Number of relevés 224167249 45 646
Diagnostic species group:
Papaver radicatum var. pseudoradicatum 7.50 0.50 . . . . . . . . . . 0.39
Minuartia arctica . 8.50 . 0.07 . 0.04 0.04 . . 0.06 . . 2.83
Polytrichastrum alpinum . . 6.50 . . 0.13 0.79 0.19 0.12 . . 0.08 8.04
Minuartia macrocarpa var. koreana ..5.75.0.39 .. .. ...1.38
Cardamine bellidifolia . . 1.50 0.06 . . 0.02 . . . . . 1.00
Phyllodoce caerulea . . . . . 0.01 . 7.75 3.40 0.28 0.31 0.06 6.86
Tofieldia coccinea agg. . 0.50 1.13 0.01 0.75 0.16 0.02 1.31 1.76 . . . 9.51
Calamagrostis langsdorfii agg. . . . . . . . 0.19 8.40 ...2.07
Aquilegia flabellate var. pumila ... .... .2.20
2.33 . 0.11 2.69
Sibbaldia procumbens ... .... .1.80
0.06 . 0.03 0.64
Sedum roseum . . . 0.01 0.06 0.06 0.10 0.19 3.20 3.17 2.75 3.50 20.22
Viola biflora ... .... .1.28
. . 0.11 0.46
Festuca ovina var. koreanoalpina 1.25 . 1.13 5.59 0.67 1.83 0.80 . . . . . 66.59
Chrysanthemum zawadskii var. alpinum 2.00 2.50 0.13 1.95 . 0.16 0.80 0.88 . 0.08 0.06 0.03 24.77
Scirpus maximowiczii . . 1.31 0.64 8.33 0.29 1.19 0.19 . . . . 29.20
Dryas octopetala var. asiatica 0.25 1.50 . 0.10 3.67 8.08 0.59 1.88 . . . . 83.92
Gagea serotina . . 0.19 0.12 0.22 2.86 0.69 . . . . . 26.54
Carex longirostrata . 0.25 0.06 0.19 0.44 2.57 0.02 . . . . . 22.29
Saussurea alpicola . 0.75 . 0.21 0.22 1.42 0.04 0.06 0.08 . 0.50 0.03 18.61
Salix rotundifolia . 0.75 1.50 0.15 0.67 0.23 5.56 1.69 0.08 . 0.19 . 28.19
Oxytropis anertii 2.00 0.50 0.25 0.59 0.50 1.80 5.04 1.31 . 0.22 . 0.03 57.43
Sanguisorba stipulate var. riishiriensis ... .... ..5.33
6.75 7.50 12.75
Carex atrata ...0.01 ... ..1.39
2.75 0.92 3.39
Erigeron thunbergii var. glabratus 0.50 . 0.06 0.01 . 0.01 . 0.75 . 1.94 . . 2.79
Hierochloe alpina . . . 0.12 0.08 0.03 . . . 1.83 . 0.06 3.65
Luzula oligantha 0.75 . . . . . . . 0.12 2.17 0.25 . 1.61
Viola sacchalinensis var. alpicola ... .... ..1.00
. . 0.28
Poa malacantha var. shinanoana ... .... ..2.36
0.13 . 0.87
Phleum alpinum ... .... ..1.11
. . 0.31
Ligularia jamesii ... . ... .. ..4.50
1.50
Veratrum maackii s.l. ... .... .. ..2.92
0.80
Saussurea triangulate var. alpina ... . .. . .. ..1.11
0.60

438 J. Plant Biol. (2019) 62:436-450
the Seo-Pa route over the western ridge closed strictly by
China until 2003. We also compared with the Dicentro-
Stellarietea nipponicae, Asplenietea rupestria, Phyllodoco-
Harrimanelletea, Cetrario-Loiseleurietea, and other alpine
windswept grassland vegetation reported for Japan (Nakamura
1987, 1997; Miyawaki et al. 1994). This phytosociological
study is clearly a pioneer work on the very remote area of
Changbaishan and we expect further research.
Results and Discussions
The open grass-like formation of the Paekdu-San summit is
the most predominant vegetation in the alpine tundra
landscape. Four vegetation types were recognized: alpine
desert, windswept grassland, snowbed vegetation, and
colluvial drainage channel vegetation. Each type comprised
several plant communities based on floristic affinities
corresponding to habitat conditions (Table 1). A total of 10
rankless units were classified and comprised 80 species of 68
vascular and 12 non-vascular plants. This number of vascular
plants is about 52.3% of the total 129 species of Chinese
Paekdu-San’s alpine tundra zone compiled by Qian et al.
(1999). Polygonum ochotense, Dryas octopetala var. asiatica,
Festuca ovina var. koreanoalpina, Oxytropis anertii, Gentiana
algida, Scirpus maximowiczii, Rhodiola angusta, Salix
rotundifolia, Gagea serotina, Carex longirostrata, and Sedum
roseum were highly influential species as indicated by r-
NCD >20%. Of 12 bryophytes recorded, the mat-forming,
widespread moss Racomitrium lanuginosum was the most
frequent and abundant. Only three low-growing rhododendrons
were recorded; Rhododendron aureum, R. lapponicum subsp.
parvifolium, and R. redowskianum. The evergreen subshrub
and yellow-flowered rosebay (R. aureum) made the most
Tabl e 1 . Continued
Vegetation type A B C D r-NCD
Syntaxa A1A2A3 B1B2B3B4 C1C2 D1D2D3
Number of relevés 224167249 45 646
Companions:
Polygonum ochotense 1.00 0.50 2.75 1.44 2.00 1.62 3.67 2.50 3.20 1.53 1.00 . 100
Rhododendron aureum . . . . 0.06 0.06 0.19 0.75 0.24 0.11 0.63 0.06 4.43
Niphotrichum canescens . . 0.19 0.01 0.28 0.06 0.25 0.44 . . . 0.06 3.49
Gentiana algida . . 1.13 1.94 0.50 1.16 1.81 0.13 . . . . 39.46
Rhodiola angusta . . 1.88 0.72 0.89 1.12 0.68 0.50 . . . . 28.81
Dicranum spadiceum . . . 0.07 1.08 0.14 0.06 0.13 0.28 . . . 5.28
Brachythecium brotheri . . 0.31 0.01 0.08 0.08 1.30 . 0.40 . . . 5.15
Persicaria vivipara . . . 0.02 0.06 0.03 0.07 . . . 0.38 0.22 1.93
Saxifraga laciniata 0.50 . 0.19 0.22 . 0.01 0.15 . . . . . 2.36
Bupleurum euphorbioides . . . . 0.06 . 0.02 0.13 . 0.42 0.13 . 0.70
Racomitrium lanuginosum . . 2.25 . 0.39 0.61 1.04 . . . . . 11.03
Festuca vivipara . . . . . . . . . 0.89 0.38 0.75 1.38
Gentiana jamesii . . . . . . . . 0.04 . 0.38 0.03 0.15
Luzula arcuata var. unalaschkensis . . . 0.01 . . 0.02 . . . 0.13 . 0.14
Saxifraga punctata . . 0.06 . . . . 0.06 . . 0.13 . 0.09
Cetraria s.l. 1 . . . . 0.28 0.01 0.74 0.38 . . . . 0.87
Trollius japonicus ... .... . . . 0.50 0.33 0.32
Saussurea triangulata . . . . . . . . . 0.44 0.13 . 0.23
Anthoxanthum nipponicum var. furumii ... ..0.00. .. ..0.44.0.21
Rhytidium rugosum . . 0.13 . . . . 0.31 . . . . 0.21
Cladonia rangiferina (lichen) . . . . 0.06 . 0.01 . . . . . 0.11
Veronica stelleri var. longistyla . . . . . . . . . 0.03 0.13 . 0.05
Salix divaricata var. metaformosa . . . . . . . . . . 0.13 0.03 0.05
Ligusticum tachiroei .0.25. .... .. ..0.03 0.05
Cetraria s.l. 2 . . . . . 0.02 . 0.38 . . . . 0.03
Single occurrence: Syntaxa
A3
–
Cerastras sp. (Keimling) 0.75, Dicranum scoparium 0.19,
B1 –
Sphagnum magellanicum 0.01,
B2 –
Vaccinium
uliginosum 0.33, Rhododendron lapponicum subsp. parvifolium 0.06, Rhododendron redowskianum 0.03,
C2 –
Abietinella sp. 0.12, Arctous rubra
0.04,
C3 –
Thalictrum aquilegiifolium var. sibiricum 0.02, Angelica nakaiana, 0.01, D1
–
Silene repens 0.22, Crepis coreana 0.22, Saussurea erio-
phylla 0.17, D2
–
Calamagrostis angustifolia 0.19, Saussurea sp. 0.19, Cerastium baischanense 0.13, D3
–
Eranthis stellata 0.08, Petasites saxati-
lis 0.06, Dryopteris sp. 0.03, Oxyria digyna 0.03

J. Plant Biol. (2019) 62:436-450 439
flourish community. On the other hand, a representative class
of the alpine open grass-like tundra formation in North
Korea, Dryado-Vaccinietea uliginosi, was described by Lee
et al. (1992), but the class is invalid and ineffective based on
the International Code of Phytosociological Nomenclature
(Weber et al. 2000).
Alpine Desert Vegetation
The harshest environments on Paekdu-San summit range are
inhabited only by the plant community with a very high floristic
homogeneity (H
opt
= 0.89) which reflects the distributional
limitations imposed on plants by extreme conditions.
Polygonum ochotense, the greatest contributor to the whole
alpine tundra formation in the study area, is the only notable
companion species characterized by high constancy [V] and
low coverage (1–4).
Another circumpolar bistort (Persicaria vivipara) occurs
very rarely in the Scirpus maximowiczii community of
windswept vegetation, which also distributes in North America
and arctic northern regions of Europe. The evergreen
Rhododendron species, Gagea serotina, Sedum roseum, and
Hierochloe alpina, which are the principal elements of the
dwarf subshrub formation just above the timberline, were
almost absent. Only 29 species were identified in 8
phytosociological relevés, and three rankless units were
classified by differential species responding to different
habitat types.
Papaver radicatum var. pseudoradicatum community
The community shows a small, very distinctive patch-like
physiognomy growing over a very unstable surface with
Tabl e 2 . Phytosociological table of alpine desert vegetation. Syntaxa A1: Papaver radicatum var. pseudoradicatum community, A2:
Minuartia arctica community, A3: Polytrichastrum alpinum-Minuartia macrocarpa var. koreana community
Syntaxa A1 A2 A3
Relevé no. 12 34 5678
No. of taxa 6 6 8 6 10 13 12 13
Differential species
Papaver radicatum var. pseudoradicatum 87 2 · · ·· ·
Minuartia arctica · · 89· · · ·
Polytrichastrum alpinum · · · · 5588
Minuartia macrocarpa var. koreana · · · · 7466
Cardamine bellidifolia · · · · 2211
Companions:
Bistorta ochotensis 11 2· 2234
Chrysanthemum zawadskii var. alpinum 31 14· · · 2
Festuca ovina var. koreanoalpina 5· · · · 222
Saxifraga laciniata 2· · · · · 3·
Erigeron thunbergii var. glabratus · 2· · · 1·
Oxytropis anertii · 8· 2· · · 4
Dryas octopetala var. asiatica · 112· · · ·
Carex longirostrata · · 1· · · · 1
Tofieldia coccinea agg. · · · 222· 2
Salix rotundifolia · · · 3· · 75
Rhodiola angusta · · · · 334·
Cerastra sp. (Keimling) · · · · 33· ·
Racomitrium lanuginosum · · · · 99· ·
Scirpus maximowiczii · · · · · 133
Gentiana algida · · · · · 123
Single occurrence: Relevé No., Coverage (r-NCD). No 1, Luzula oligantha 3 (2.9); No. 3, Saussurea alpicola 3 (2.9), Ligusticum tachiroei 1,
(1.0); No. 5, Gagea serotina 3 (2.9), Rhytidium rugosum 2 (1.9); No. 6, Saxifraga punctata 1 (1.0); No. 7, Dicranum scoparium 3 (2.9), Nipho-
trichum canescens 3 (2.9); No. 8, Brachythecium brotheri 5 (4.8).
Site environment: Relevé No., Latitude (° ′ ″ N) & longitude (° ′ ″ E), Area (m ), Altitude (m), Aspect, Slope (°), Height (cm) & coverage (%) of
herb, Coverage of moss (%). No. 1, no data, 0.3, 2127, SW, 40, 20, 70; No. 2, 42 00 23, 128 01 52, 1, 2623, N, 30, 15, 95, No. 3, no data, 0.8,
2274, SW, 15, 20, 70, No. 4, 41 59 51, 128 01 38, 0.08, 2572, N, 2, 4, 80; No. 5, 42 01 02, 128 01 52, 1, 2600, N, 20, 5, 80; No. 6, 42 00 11, 128
01 35, 1, 2565, N, 40, 5, 95; No. 7, 42 00 50, 128 01 40, 0.3, 2489, N, 25, 3, 60; No. 8, 42 01 01, 128 01 24, 1, 2410, N, 3, 3, 20.
Investigation date: 03.07.2009 (No. 4~8), 07.10.2004 (No. 1~3). Investigators: JWKIM, YMJEONG (No. 1, 3), JWKIM, SUHAN (No. 2),
JWKIM, SYKIM (No. 4, 5, 7, 8), JWKIM, MSKIM (No. 6).

440 J. Plant Biol. (2019) 62:436-450
scoria, scree, and debris in the uppermost part of the alpine
belt. Occurring at the margins of the belt, this community
develops in the transition zone between inhabitable and non-
inhabitable sites on the Paekdu-san summits. The community is
found sporadically on microhabitats of the Cheonji caldera
ridges, in areas with rock sediments of weathered, rough-
edged, dark grey volcanic materials, such as basaltic scoria,
rhyolitic pumice, and trachytic and tuffaceous breccia. Fine
grained soil particles in the sites are mostly blown and
washed away by strong winds and snowdrifts and heavy
rainfalls. Habitat conditions are similar to those of pioneer
communities dominated by Papaver species found in polar
deserts with comparatively little snowfall and aeolian erosion
(Kienast et al. 2005). Species composition of the community
is very simple, and only P. radicatum var. pseudoradicatum
is diagnostic (Table 2).
The present community obviously is geographically vicariant
with the Papaveretum fauriei described in Rishiri Island
(45°10′N) off the west coast of Hokkaido (Ohba 1977).
Luzula oligantha is the only common species for both units.
The community on Rishiri comprises richness in diagnostic
species such as P. fauriei, Stellaria ruscifolia, Oxyria digyna,
and Arabis lyrata var. kamchatica. In contrast, the community
on Paekdu-San includes the widely distributed alpine desert
species associated with stony steppe: Polygonum ochotense,
Chrysanthemum zawadskii var. alpinum, Oxytropis anertii,
and Festuca ovina var. koreanoalpina. Papaveretum fauriei
of Japan is the most southerly distributed component of
poppy alpine desert vegetation (Nakamura and Krestov
2012), but the present community of P. radicatum var.
pseudoradicatum on Paekdu-San (42°00′N) is regarded as an
endemic type occurring in the southernmost area of the
continental Northeast Asia. The Hokkaido Papaveretum
fauriei belongs to the class Dicentro-Stellarietea nipponicae
(Ohba 1968), but the syntaxonomy of Paekdu-San’s poppy
unit remains unresolved due to insufficient phytosociological
data. The subassociation Dryado tschonoskii-Rhododendretum
aurei papaveretosum radicatae, described by Kolbek and
Jarolímek (2007) on the North Korean side of Paekdu-San,
was assigned to the subshrub tundra vegetation in the alpine
and subalpine belt by the presence of various differential
species, such as Potentilla nivea, Androsace capitata, Ligusticum
tachiroei, Pedicularis verticillata, and other species with
higher constancy (Kolbek and Jarolímek 2007). However,
Papaver radicatum has been found chiefly on unstable and
fully open sites, as the Papaver radicatum var. pseudoradicatum
community is likewise. Somewhat similar poppy associations
occur in the Carpathian Mountains at the border between
Poland and Slovakia (Valachovič et al. 1997; Biţă-Nicolae
2009), which may be considered vicarious to the northeastern
Asian syntaxa. Another poppy class, Drabo corymbosae-
Papaveretea dahliani, existed in the High Arctic polar
deserts (Daniëls et al. 2016).
Minuartia arctica community
This is another threshold community type dominated by
Minuartia arctica, which is found sporadically on a very
unstable habitat with stone crumbs at higher altitudes on
Paekdu-San. M. arctica is a pioneer encouraging clayey
lithosol deposition, thereby gradually increasing its patch
size. The syntaxonomic status remains undefined due to
insufficient data. M. arctica occurs in northern Siberia
(Matveyeva 1994) and northwest Canada and Alaska’s
northern polar areas with barren landscapes of exposed soil
and low phytomass (Zhu and Rowe 1987; Victoria and
Hilary 2001). In the arctic coastal plains of Beringian Alaska, its
habitats are dry, exposed sites with small-patterned ground
feature (Walker et al. 2011). The Minuartia community
differs from the Papaver community by the presence of Salix
rotundifolia, a dwarf subshrub and pioneer on barren sites.
Cerastio-Minuartietum vernae japonicae (Ohba 1968; Okuda
1985) on ultramafic stony-slumped sites in central Honshu of
Japan is quite similar to the physiognomy of the Minuartia
arctica community, but both syntaxa do not share any
common species (Table 2).
Polytrichastrum alpinum-minuartia macrocarpa var. koreana
community
Present community is found in humid habitats with frequent
clouds and long-lasting fog. Its differential species are M.
macrocarpa var. koreana, Cardamine bellidifolia, and
Polytrichastrum alpinum (Table 2). A low-lying, 10 cm thick
cushion mat of mosses dominated by P. alpinum and partly
Racomitrium lanuginosum forms a unique matrix and develops
on stabilized and flattened tuffaceous block streams. Such
edaphic situation of community site is a very favorable home
to the Korean Pika (Ochotona coreana), a small lagomorph
mammal restricted to the Changbai region (Fig. 3). The
vegetation is less than 5 cm high. Fine-grained soils weathered
from the tuff are rarely deposited on the block. The site
occurs sporadically and rarely, and it is found only on the
block stream of northern slopes of Baekwoon-bong. The
species composition is relatively diverse (12 species per
relevé) in comparison with the above phytocoenons. Although
the soils are very shallow and poor, moist conditions of the
habitat prevent thermophilic or xerophilic plants such as
Dryas octopetala var. asiatica, a very common component
of alpine tundra. Gagea serotina and Oxytropis anertii,
which are common in alpine tundra, are very rare and
frequently absent from this community.
Alpine Windswept Dwarf Vegetation
The landscape around the ridges of the Paekdu-San mountain

J. Plant Biol. (2019) 62:436-450 441
top is predominant of wind-exposed dwarf-herb communities.
The habitats are noticeably cold and windy throughout the
year. A total of 56 phytosociological relevés with 44 species
were analyzed, and 4 rankless units were classified:
Chrysanthemum zawadskii var. alpinum-Festuca ovina var.
koreano-alpina community, Scirpus maximowiczii community,
Carex longirostrata-Dryas octopetala asiatica community,
and Oxytropis anertii-Salix rotundifolia community. The
Tabl e 3 . Phytosociological table of alpine windswept grassland. Syntaxa B1: Chrysanthemum zawadskii var. alpinum-Festuca ovina var.
koreano-alpina community, B2: Scirpus maximowiczii community
Syntaxa B1 B2
Relevé no. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23
No. of taxa 5 7 5 6124 9118 9 5 7 812510121091491212
Character and differential species:
Festuca ovina var. koreanoalpina 7896747632338648 42 · · 3 · 3
Chrysanthemum zawadskii var. alpinum · 2222 · 247868.?2.?2 ·······
Scirpus maximowiczii 1····5····3·22325998888
Dryas octopetala var. asiatica ·······2·····2 12343723
Companions:
Polygonum ochotense 122 · 2 · 2 · 121365 · 2 321222 3
Gentiana algida 54323423 · 3 · · 2222 321 · · · 3
Oxytropis anertii ·2·22·1·3· ·3·2· 22· ·1·23
Rhodiola angusta ····2·42···24322222··22
Salix rotundifolia 3···2······3···· 31·2·21
Carex longirostrata ··2····2···12···4····35
Saussurea alpicola ···2······2··4·222··2··
Gagea serotina ····2·22········322····
Tofieldia coccinea agg. ····1·1········· ···333·
Saxifraga laciniata ·····3······3142 ·······
Dicranum spadiceum ········73······ ···5·44
Hierochloe alpina ····2···3····7·· ··3····
Cardamine bellidifolia ·······111······1····· ·
Niphotrichum canescens ·2·············· ···2·3·
Minuartia arctica ···32··2········ ·······
Persicaria vivipara ······2··1······ ···2···
Erigeron thunbergii var. glabratus ········11······ ······ ·
Sedum roseum ·········3······ ·2···· ·
Brachythecium brotheri ·············2·· ····3· ·
Minuartia macrocarpa var. koreana ················ ···5·2·
Cetraria sp.1················ ···2··3
Racomitrium lanuginosum ················ ····52·
Vaccinium uliginosum ················ ····4·2
Single occurrence: Relevé no., Coverage (r-NCD). No. 2, Luzula arcuata var. unalaschkensis, 2 (0.1); No. 5, Sphagnum magellanicum, 3 (0.1);
No. 8, Carex atrata, 2 (0.1), Ligularia jamesii, 1 (0.0); No. 13, Saussurea triangulate var. alpina, 1 (0.0); No. 19, Rhododendron lapponicum
subsp. parvifolium, 2, (0.1), Rhododendron redowskianum, 1 (0.1); No. 20, Rhododendron aureum, 2 (0.1), Bupleurum euphorbioides, 2 (0.1),
Cladonia rangiferina (lichen), 2 (0.1).
Site environment: Relevé no., Latitude (° ′ ″ N) & longitude (° ′ ″ E), Area (m ), Altitude (m), Aspect, Slope (°), Height (cm) & Cover (%) of
herb. Coverage of moss (%). No. 1, 42 00 00, 128 01 28, 4, 2601, S, 5, 5, 60; No. 2, 41 59 58, 128 01 30, 1, 2566, S, 5, 5, 70; No. 3, 41 59 59,
128 01 29, 1, 2583, SE, 40, 10, 90; No. 4, 128 01 40, 42 00 51, 1.5, 2593, W, 30, 5, 30; No. 5, 41 58 37, 128 53 31, 0.1, 2538, N, 3, 10, 50; No. 6,
42 00 15, 128 01 46, 4, 2653, W, 5, 5, 20; No. 7, 41 58 37, 128 53 31, 0.2, 2538, S, 10, 20, 50; No. 8, 41 59 39, 128 01 24, 4, 2332, W, 40, 5, 40;
No. 9, no data, 1.05, 2538, W, 5, 10, 50; No. 10, no data, 1, 2538, W, 10, 10, 70; No. 11, 41 58 44, 128 03 23, 1, 2540, SE, 5, 5, 50; No. 12, 41 58
45, 128 03 21, 0.09, 2552, SSW, 2, 5, 80; No. 13, 41 59 41, 128 01 51, 1, 2462, SW, 3, 15, 90; No. 14, 41 59 44, 128 01 36, 1, 2468, NW, 4, 5, 80;
No. 15, 41 59 58, 128 01 30, 1, 2566, SSW, 20, 2, 20; No. 16, 41 58 40, 128 03 26, 1, 2528, SSE, 5, 3, 80; No. 17, 42 00 09, 128 01 33, 4, 2575,
N, 10, 5, 90; No. 18, 41 59 51, 128 01 38, 0.5, 2572, S, 15, 10, 90; No. 19, no data, 1, 2410, W, 20, 10, 95; No. 20, 41 59 58, 128 01 38, 0.5,
2662, NW, 2, 5, 95; No. 21, 42 00 27, 128 01 54, 0.5, 2413, W, 7, 4, 95; No. 22, 41 59 51, 128 01 38, 0.5, 2572, W, 3, 5, 80; No. 23, 42 00 03,
128 01 46, 0.15, 2568, N, 15, 5, 95.
Investigation date: 07.02.2009 (No. 5, 7, 9~12, 16), 07.03.2009 (No. 1~4, 6, 8, 13~15, 17~23). Investigators: JWKIM, BKCHOI (No. 1, 2, 3,
11, 12, 15~17), JWKIM, BKCHOI, JCLIM (No. 4, 6, 8), JWKIM, SYKIM, JCLIM (No. 5, 7, 9, 10), JWKIM. SYKIM (No. 13, 18~23), JWKIM,
MSKIM (No. 14).

442 J. Plant Biol. (2019) 62:436-450
average homotoneity (H
opt
0.20; max. 0.46, min. 0.26) of
present vegetation type was three-fold lower than other
vegetation types (H
opt
0.68 for snowbed vegetation and H
opt
0.89 for alpine desert vegetation). The reed grass Calamagrostis
langsdorfii agg. was almost exclusive in present vegetation
type; on the North Korean side of Paekdu-San, reed grass
is confined to subshrub rhododendron (R. aureum) stands,
where it shows the high constant degree (92%) in the light
taiga forests (Rhododendro aurei-Laricetum olgensis; Dostalek
et al. 1988) between 1,740-1,950 m altitude (Kolbek and
Jarolímek 2007). This rhododendron has also been considered
as a differential species of Viola biflora-Calamagrostis
langsdorfii community of the snowbed vegetation.
Chrysanthemum zawadskii var. alpinum-Festuca ovina var.
koreano-alpina community
This community was observed mainly on windward habitats
along the ridges, on level terrains with somewhat stronger
gravitational disturbance, but also on and around hiking trails
in the alpine zone. The coarse fraction in the soil is high, and
eroded soil surfaces are extensive. Most of the sites are
exposed to a day-long solar radiation either westward or
southward. In such habitat conditions, fescues occur sporadically
in bundles on semicircular, 2–3 cm high soil mounds, resulting
in an average vegetation cover of 58%. The differential
species Festuca ovina var. koreano-alpina and Chrysanthemum
zawadskii var. alpinum were dominant alternately (Table 3);
the former is regionally more common to the Paekdu-San
range (Kim et al. 1991). Species richness of the community
was very poor, with an average of 7.9 species per relevé.
Gentiana algida, a common species of cold alpine meadows
in Asian Eurasia and North America, emerged with a high
contribution degree to the unit. Another fescue community,
Festuceta ovinae koreanoalpinae ascribed to the Oxitropion
anertii, has been described by Kim et al. (1991) on
southeastern slopes of Paekdu-san in North Korea. However,
these syntaxa could not be compared here because of not
only the missing nomenclatural type, but also especially the
community table of the almost incomplete flora-list.
Scirpus maximowiczii community
This community is classified by two differential species,
Scirpus maximowiczii and Dryas octopetala var. asiatica
(Table 3). An open mat-like bulrush community of the high
moor element S. maximowiczii is found on such microhabitat
conditions caused by late-melting snow. The presence of
various mosses and lichens (Dicranum scoparium, Racomitrium
lanuginosum, R. canescens, Cetraria species, Brachythecium
brotheri, Cladonia rangiferina, and Rhytidium rugosum) is
characteristic for the community. The habitat is strongly
influenced by windy conditions and very wet fine-grained
soil formed by frost boil (Peterson et al. 2003). Present
community is highly homogeneous, with H
opt
0.41. Due to
insufficient data for comparison, however, its syntaxonomical
ranked unit was unresolved. Most stands were located at the
area free from human interference, along the ridge, and on
piles of rocks 3–5 cm in diameter and structured soils. The
community is bordered by Oxytropis anertii-Salix rotundifolia
community. Less frequently, dwarf subshrubs Vaccinium
uliginosum and S. rotundifolia intrude into this bulrush bed,
depending on a protruding microhabitat. Polygonum ochotense
occurs constantly with coverage of 1–3, whereas P. vivipara
is very rare.
Carex longirostrata-Dryas octopetala var. asiatica community
Among the alpine windward plant communities, this community
was predominant in the sward-like landscape on the
northwestern part of Paekdu-San. Only 47 species were
listed in a total of 24 relevés. D. octopetala var. asiatica, C.
longirostrata, Gagea serotina, and Saussurea alpicola are
differential species of the structured community. D. octopetala
var. asiatica forms a dense, mat-like vegetation of prostrate
evergreen subshrub, and C. longirostrata and G. serotina are
intermixed with relatively high frequency. Particularly, the
high contribution of D. octopetala var. asiatica and R.
lanuginosum reflects the habitat conditions (Natalia 2001;
Pearce and van der Wal 2002). Protruding or flattened
microhabitats of the community are oligotrophic with less
organic matter, small rock fragments, and soil blown away
due to heavy winds and early-melting snow. D. octopetala
var. asiatica, which is a diagnostic species to the Oxytropidion
japonicae and Caricetalia tenuiformis in Japan (Miyawaki et
al. 1994), is one of the most abundant components of the
East Asian tundra and alpine windswept heathland. In contrast,
a representative alpine landscape on the North Korean side
of Paekdu-San belongs to the Asian mountain avens biome,
i.e. Dryeta octopetalae (Paekdu-San Series Editorial Committee
1992). The Oxytropido-Dryadetum octopetalae asiaticae,
described on the southeastern slope (Lee et al. 1992), is a
vicarious syntaxon of this Carex longirostrata-Dryas octopetala
var. asiatica community. The Oxytropido-Dryadetum is
characterized by high richness of dwarf shrubs such as
Vaccinium uliginosum, Rhododendron parviflorum var.
alpinum, R. aureum, and Arctous rubra, which replace
windswept herbaceous plants such as C. longirostrata,
Rhodiola angusta, Festuca ovina var. koreanoalpina, and
Gentiana algida.
Present Carex-Dryas community, like a carpet on the
ground, forms a monolayer structure about 8 cm in height.
This community shows the most similar species composition
with the Oxytropis anertii-Salix rotundifolia community
described below. Very common species at the wind-exposed
alpine stands, Phyllodoce caerulea, Anthoxanthum nipponicum
var. furumii, Saussurea triangulata, Luzula arcuata var.

J. Plant Biol. (2019) 62:436-450 443
Tabl e 4 . Phytosociological table of aven and willow subshrub vegetation. Syntaxa B3: Carex longirostrata-Dryas octopetala var. asiatica
community, B4: Oxytropis anertii-Salix rotundifolia community
Syntaxa B3 B4
Relevé no. 123456789101112131415161718192021222324252627282930313233
No. of taxa 6 12 10 13 15 12 9 11 9 12 9 12 10 11 9 8 9 12 10 9 11 11 10 11 11 9 9 8 12 12 10 14 14
Character and differential species:
Dryas octopetala var. asiatica 787998898998799889776979 · · 2 · 3 · · 52
Carex longirostrata 54 · 342 · 36224 · 2546 · 372334 ·······2·
Gagea serotina · 3153424324453345246 · 242 ····33·35
Saussurea alpicola 63 · 22235 · 1 · 2 · · 257 · 423 · 2 · ········3
Salix rotundifolia ··3·2···42·······3··3··2 798375524
Oxytropis anertii · · · 3 · 55463255 · 44322 · · 224 3 · 4968885
Companions:
Polygonum ochotense 32223234453 · · 34 · · 25 · · 44 · 525233328
Festuca ovina var. koreanoalpina 534 · 3 · 62 · 5 · 4 · 3 · 43165 · 422 2 · 3 · 21 · · 5
Gentiana algida · 13142223222 · 2 · · · 1 · 21223 33 · 24 · 432
Rhodiola angusta · 52434433 ········2324112 13 · 22 · 3 · ·
Scirpus maximowiczii ·2··54·····5·····4··2·2· ·2·2·4233
Chrysanthemum zawadskii var.
alpinum ····333·····1···4·1····· 32·2·33··
Polytrichastrum alpinum ····5··2·········9···3·· 5·3·3·5··
Tofieldia coccinea agg. ···412··4··13··········· ········2
Rhododendron aureum ···3·········21··3······ ····21·2·
Sedum roseum ·········4··221········· ·······22
Persicaria vivipara ·····2····2··2·········· ····12···
Minuartia arctica ·3··················4··1 ·····3···
Saxifraga laciniata ··2·2··················· ··33·····
Hierochloe alpina ·········2··········22·· ·········
Cardamine bellidifolia ···········1············ ·2·······
Erigeron thunbergii var. glabratus ··············1···2····· ·········
Thalictrum aquilegiifolium var.
sibiricum ···············32······· ·········
Niphotrichum canescens ·232···············2···· 3·7······
Dicranum spadiceum ·263················2··3 ········5
Racomitrium lanuginosum · · · 74 · · · · ·6673 · · · · · · 8· · 3 · 7 · · ·383 ·
Brachythecium brotheri ·······3·2·2·········5·· ····62355
Cetraria s.l.1 ············3····1······ 54·····33
Cetraria s.l.2 ··········4·3··········· ·········
Single occurrence: Relevé no., Coverage (r-NCD), No. 1, Phyllodoce caerulea, 4 (0.1), No. 13, Anthoxanthum nipponicum var. furumii, 1 (0.0),
No. 14, Saussurea triangulate var. alpina, 2 (0.0), No. 20, Angelica nakaiana, 2 (0.0), No. 25, Luzula arcuata var. unalaschkensis, 2 (0.0), No. 27,
Cladonia rangiferina (lichen), 1 (0.0), No. 32, Bupleurum euphorbioides, 2 (0.0)
Site environment: Relevé no., Latitude (° ′ ″ N) & longitude (° ′ ″ E), Area (m ), Altitude (m), Aspect, Slope (°), Height (cm) & Cover (%) of herb, Cov-
erage of moss (%). No. 1, 41 59 47, 128 01 45, 0.6, 2463, W, 5, 3, 75, No. 2, 42 01 29, 126 01 47, 1, 2606, W, 20, 10, 75, No. 3, 42 00 02, 128 01 28, 1,
2604, W, 3, 3, 70, No. 4, 42 01 44, 128 02 23, 1, 2500, N, 10, 5, 90, No. 5, 42 00 34, 128 01 55, 1, 2476, W, 10, 20, 85, No. 6, 42 00 51, 128 01 42, 0.25,
2301, N, 15, 10, 95, No. 7, 41 58 46, 128 03 25, 1, 2524, SEE, 2, 3, 80, No. 8, 41 59 59, 128 01 30, 1, 2568, W, 3, 5, 90, No. 9, 42 00 30, 128 01 54, 1,
2618, N, 30, 10, 95, No. 10, 42 00 21, 128 01 48, 1, 2635, N, 2, 20, 95, No. 11, 42 00 24, 128 01 53, 0.25, 2596, SEE, 10, 5, 90, No. 12, 42 00 31, 128 01
57, 1, 2503, NNW, 50, 5, 90, No. 13, 42 01 40, 128 02 36, 0.25, 2477, N, 3, 7, 80, No. 14, 41 59 40, 128 01 46, 1, 2446, S, 5, 7, 95, No. 15, 42 00 21, 128
01 50, 0.12, 2584, N, 6, 6, 90, No. 16, 41 59 56, 128 01 34, 0.06, 2422, E, 30, 8, 90, No. 17, 41 59 57, 128 01 35, 0.35, 2424, E, 10, 7, 95, No. 18, 42 00
17, 128 01 46, 1, 2650, SW, 5, 15, 85, No. 19, 41 59 50, 128 01 30, 1, 2454, SE, 10, 20, 90, No. 20, 41 59 50, 128 01 30, 2, 2469, S, 30, 5, 85, No. 21, 42
01 01, 128 01 29, 1, 2463, NW, 5, 5, 30, No. 22, 42 00 13, 128 01 42, 1, 2630, W, 3, 5, 95, No. 23, 41 59 58, 128 01 30, 4, 2564, S, 30, 5, 85, No. 24, 42
01 21, 128 01 52, 1, 2607, S, 10, 2, 95, No. 25, 42 00 18, 128 01 47, 0.25, 2661, W, 50, 5, 50, No. 26, 42 00 23, 128 01 52, 0.25, 2621, NW, 20, 3, 90, No.
27, 41 59 58, 128 01 30, 0.35, 2570, NNE, 2, 3, 80, No. 28, 42 00 21, 128 01 48, 0.25, 2635, ·, 0, 5, 90, No. 29, 42 00 23, 128 01 52, 1, 2621, N, 30, 10,
70, No. 30, 42 00 50, 128 02 13, 0.12, 2466, N, 3, 7, 90, No. 31, 42 00 20, 128 01 49, 1, 2653, W, 30, 10, 90, No. 32, 41 59 48, 128 01 39, 0.5, 2548, NW,
10, 10, 90, No. 33, 42 00 04, 128 01 50, 0.25, 2557, N, 7, 7, 95.
Investigation date: 07.10.2004 (No. 1, 15~17), 07.02.2009 (No. 7), 07.03.2009 (No. 2~6, 8~14, 18~33). Investigators: No. WKIM, JSKIM (No.
1, 16, 17), JWKIM, SYKIM (No. 2, 4, 6, 13, 14, 21, 24, 30, 32, 33), No. 3, JWKIM, BKCHOI (No. 3, 7, 10, 11, 20, 23, 26~28), No. 5, JWKIM,
JCLIM (No. 5, 9, 18, 22, 29, 31), No. 8, JWKIM, MSKIM (No. 8, 19), JWKIM, BKCHOI, JCLIM (No. 12, 25), JWKIM, YMJEONG (No. 15).

444 J. Plant Biol. (2019) 62:436-450
unalaschkensis, and Angelica nakaiana, occur very rarely in
this Carex-Dryas community.
Oxytropis anertii-Salix rotundifolia community
Present community is a type of pioneer vegetation different
from the above Carex longirostrata-Dryas octopetala var.
asiatica community. Its microhabitat shows much more
unstable situation of the volcanic sand- and scoria-alluvial
microtopography with relatively greater inclination range
(min. 3°, max. 50°). Domination of Salix rotundifolia and near
exclusion of Saussurea alpicola and Carex longirostrata are
tied to such edaphic conditions. S. rotundifolia appears as a
loose mat-forming or densely caespitose summer-green
subshrub on Paekdu-San. This round and small-leaved willow
is an amphi-Beringian species of the alpine tundra that
occurs typically at the edge of sites with patterned ground
(Kojima 1991; George 2004). As windswept dwarf subshrub
stands, both communities contain conspicuous, large- and
small-sized cushion-forming mosses, Racomitrium lanuginosum,
R. canescens, Dicranum spadiceum, and Brachythecium
brotheri. Syngeographic vicariant vegetation types, both
Salici-Oxytropidetum yesoensis and Saxifraga oppositifolia-
Salix arctica community have been described on Mt.
Daisetsu in Japan (Nakamura 1988) and Cornwallis Land in
Canada (Kojima 1991), respectively. Habitat conditions of
these syntaxa are all similar, which are a prevailingly
windward site and moderate slopes with sandy soil around
patterned ground. In contrast, the species composition of the
Salicetum rotundifoliae described in a snowbed on the
southeastern slope of the North Korean side of Paekdu-San
(Lee et al. 1992) is rather unusual. Alpine windswept
components such as D. octopetala var. asiatica, Gagea
Tabl e 5 . Phytosociological table of snowbed vegetation. Syntaxa C1: Phyllodoce caerulea-Tofieldia coccinea community, C2: Vio l a
biflora-Calamagrostis langsdorfii community.
Syntaxa C1 C2
Relevé no. 1234 56789
No. of taxa 8 13 7 13 9 8 8 10 14
Differential species
Phyllodoce caerulea 9886 32255
Tofieldia coccinea agg. 23· 233· 23
Calamagrostis langsdorfii agg. · · 3 · 99978
Aquilegia flabellate var. pumila · · · · 32222
Sibbaldia procumbens · · · · 22212
Sedum roseum · 3 · · 52333
Viola biflora ···· 213 ·2
Companions:
Bistorta ochotensis 1333 22354
Salix rotundifolia 3·33 ····2
Rhododendron aureum 1· ·5 · · ·12
Erigeron thunbergii var. glabratus 3··3 ·····
Rhodiola angusta 22·· ·····
Oxytropis anertii ·322 ·····
Dryas octopetala var. asiatica ·235 ·····
Saussurea alpicola ·1· · · · ·2·
Chrysanthemum zawadskii var. alpinum ··34 ·····
Dicranum spadiceum ···2 ····7
Polytrichastrum alpinum ···3 ····3
Brachythecium brotheri · · · · 3·2· ·
Single occurrence: Relevé no., Coverage (r-NCD). No. 1, Saxifraga punctata, 1 (0.2); No. 2, Cetraria sp.-1, 6 (1.4), Niphotrichum canescens, 7
(1.6), Gentiana algida, 2 (0.5), Bupleurum euphorbioides, 2 (0.5), Cetraria sp.-2, 6 (1.4); No. 4, Scirpus maximowiczii, 3 (0.7), Rhytidium rugo-
sum, 5 (1.2); No. 5, Abietinella sp., 3 (0.7); No. 6, Arctous rubra, 1 (0.2), Gentiana jamesii, 1 (0.2); No. 7, Luzula oligantha, 3 (0.7).
Site environment: Relevé no., Latitude (° ′ ″ N) & longitude (° ′ ″ E), Area (m ), Altitude (m), Aspect, Slope (°), Height (cm) & Cover (%) of
herb, Coverage of moss (%). No. 1, no data, 0.3, 2127, SW, 40, 20, 70; No. 2, 42 00 23, 128 01 52, 1, 2623, N, 30, 15, 95; No. 3, no data, 0.8,
2274, SW, 15, 20, 70; No. 4, no data, 0.08, 2572, N, 2, 4, 80; No. 5, 42 01 02, 128 01 52, 1, 2600, N, 20, 5, 80; No. 6, 42 00 11, 128 01 35, 1,
2565, N, 40, 5, 95; No. 7, 42 00 50, 128 01 40, 0.3, 2489, N, 25, 3, 60; No. 8, 42 01 39, 128 02 03, 1, 2410, N, 3, 3, 20; No. 9, 42 01 40, 128 02
36, 0.5, 2476, N, 1, 4, 95.
Investigation date: 07.03.2009 (No, 4~9), 07.10.2004 (No, 1~3). Investigators: JWKIM, YMJEONG (No. 1, 3), JWKIM, SUHAN (No. 2),
JWKIM, SYKIM (No. 4~6, 8, 9), JWKIM, JCLIM, CWLEE (No. 7).

J. Plant Biol. (2019) 62:436-450 445
Tabl e 6 . Phytosociological table of Carex atrata-Sanguisorba stipulate var. riishiriensis community of the colluvial drainage channel vegetation
in the alpine zone. Syntaxa D1: Luzula oligantha subcommunity, D2: typical subcommunity, D3: Ligularia jamesii subcommunity
Syntaxa D1D2D3
Relevé no. 123456 78910111213141516
No. of taxa 18121514149 91795 7879918
Differential species:
Sanguisorba stipulate var. riishiriensis 378734 8568 896958
Carex atrata 22222 · 2243 · 42 · · 5
Festuca vivipara 2222 · · 2 · 1 · 2 · 3 · 4 ·
Luzula oligantha 212224 · · 11 ······
Viola sacchalinensis var. alpicola 1·332· ···· ······
Poa malacantha var. shinanoana 23255 · · 2 · · ······
Phleum alpinum 3322· · · · · · ······
Aquilegia flabellate var. pumila 222224 · · · · ····11
Hierochloe alpina 223121 · · · · ·····2
Erigeron thunbergii var. glabratus 2 · 2226 · · · · ······
Ligularia jamesii ······ ···· 355563
Veratrum maackii s.l. ······ ···· 5455·2
Saussurea triangulata var. alpina ······ ···· 5·2·12
Companions:
Sedum roseum 532333 1334 243345
Polygonum ochotense 22223 · · · 35 ······
Rhododendron aureum 1····132·· 2·····
Persicaria vivipara ······ 12·· ····22
Saussurea triangulata 3··5·· 2··· ······
Saussurea alpicola ······ 22·· ····1·
Gentiana jamesii ······ 12·· ···1··
Veronica stelleri var. longistyla ··1··· ·2·· ······
Chrysanthemum zawadskii var. alpinum ····3· ·1·· ·····1
Phyllodoce caerulea ····32 ·5·· ·····2
Salix divaricata var. metaformosa ······ ·2·· ·····1
Bupleurum euphorbioides 1 · 2 · 2 · · 2 · · ······
Trollius japonicus ······ ··8· ·3·3··
Saxifraga punctata ······ ··2· ······
Oxytropis anertii 22···· ···· ·····1
Silene repens 22···· ···· ······
Sibbaldia procumbens ··2··· ···· ····1·
Crepis coreana 2··2·· ···· ······
Anthoxanthum
nipponicum var. furumii ······ ···· ·5···3
Single occurrence: Relevé no., Coverage (r-NCD), No. 5, Minuartia arctica, 2 (0.1), No. 6, Saussurea eriophylla, 6 (0.4), No. 8, Luzula arcuata
var. unalaschkensis, 2 (0.1), Salix rotundifolia, 3 (0.2), Calamagrostis angustifolia, 3 (0.2), Cerastium baischanense, 2 (0.1), No. 9, Saussurea sp.,
3 (0.2), No. 12, Eranthis stellata, 3 (0.2), No. 14, Viola biflora, 4 (0.2), Petasites saxatilis, 2 (0.1), Dryopteris sp., 1 (0.1), No. 16, Polytrichastrum
alpinum, 3 (0.2), Niphotrichum canescens, 2 (0.1), Oxyria digyna, 1 (0.1), Ligusticum tachiroei, 1 (0.1).
Site environment: Relevé no., Latitude (° ′ ″ N) & longitude (° ′ ″ E), Area (m ), Altitude (m), Aspect, Slope (°), Height (cm) & Cover (%) of
herb. Coverage of moss (%). No. 1, no data, 1, 2260, S, 10, 45, 98, No. 2, no data, 0.8, 2260, S, 10, 25, 98, No. 3, no data, 1.5, 2260, S, 15, 35, 95,
No. 4, no data, 1.5, 2260, S, 15, 40, 90, No. 5, no data, 1, 2270, S, 15, 35, 90, No. 6, 41 59 41, 128 01 26, 6, 2386, S, 25, 12, 90, No. 7, no data, 1,
2100, N, 15, 22, 80, No. 8, no data, 1, 2110, N, 10, 25, 80, No. 9, 42 00 50, 128 01 36, 2, 2513, N, 5, 18, 98, No. 10, 42 00 59, 128 01 41, 0.25,
2548, NWW, 5, 15, 75, No. 11, 41 59 45, 128 01 38, 2, 2435, W, 50, 5, 80, No. 12, 42 00 53, 128 01 09, 4, 2205, N, 3, 20, 95, No. 13, 42 00 35,
128 01 45, 3, 2399, NWW, 3, 50, 80, No. 14, 42 00 45, 128 01 17, 3, 2181, NE, 35, 40, 100, No. 15, 42 00 55, 128 01 14, 1, 2190, N, 40, 20, 50,
No. 16, 42 00 44, 128 01 22, 1.5, 2255, E, 20, 25, 90.
Investigation date: 08.31.1991 (No. 1~5), 09.01.1991 (No. 7, 8), 07.10.2004 (No. 6, 9, 10, 14), 07.03.2009 (No. 11~13, 15, 16). Investigators:
JWKIM, JEKIM (No. 1~5, 7, 8), JWKIM, YMJEONG (No. 6), JWKIM, SUHAN, GJCHO (No. 9, 10, 14), JWKIM, BKCHOI, JCLIM (No.
11~13), JWKIM, JCLIM (No. 15), JWKIM, SYKIM (No. 16).

446 J. Plant Biol. (2019) 62:436-450
serotina, and Gentiana algida are absent. Instead the alpine
snowbed component Phyllodoce caerulea appears high
frequency.
Snowbed Vegetation
The more chionophilous Phyllodoce caerulea-Tofieldia coccinea
community and Viola biflora-Calamagrostis langsdorfii
community were described by using nine phytosociological
relevés containing 31 species (Table 5). Plots were located
mainly away from hiking trails on a leeward slope in the
study area. The species richness was slightly higher than in
the alpine desert vegetation. The average species number per
relevé was 10.1 species (min. 7 spp., max. 14 spp.). Most stands
were found on a very gentle microtopography, with an average
slope of 5.2° (max. 15°, min. 1°). The optimal homotoneity of
this vegetation (average 0.68) was comparatively very high,
nonetheless floristic and structural variation was observed in
between these chionophilous communities.
Phyllodoce caerulea-Tofieldia coccinea community
This community is characterized by evergreen species such
as the subshrub Phyllodoce caerulea and Tofieldia coccinea
agg. Some plots were covered by Salix rotundifolia, a
species that is regarded a stenobasiophyte in the Chukotka-
Alaska province of the subarctic tundra (Razzhivin 1994).
Rhododendron aureum was adopted as a weak differential
species. These dwarf subshrubs were not found in a plot
dominated by the moss Niphotrichum canescens and two
different lichen species of the genus Cetraria (Table 5).
Habitat conditions of the community were considerably
uniform at the bottom of well-like depressions mainly with
lingering snow cover and a layer of 0.5–1 cm wide pebble on
leeward slopes and ridges. The already stabilized and wetter
crevices provided a major adaptive condition for the well-
growing population of T. coccinea. Characteristically, Dryas
octopetala var. asiatica and Oxytropis anertii, representative
diagnostic species of the alpine windswept vegetation, also
appear high frequency. The present community occurred in
somewhat drier conditions in comparison with next Viola
biflora-Calamagrostis langsdorfii community. A geographical
vicariant unit to the present community is Phyllodocetum
yezoensi-aleuticae of the alliance Phyllodocion aleuticae
(Ohba 1967) described in Hokkaido, Japan. The association
Phyllodocetum is characterized by Phyllodoce aleutica, P.
caerulea var. yezoensis, and Harrimanella stelleriana
(Nakamura 1988). Each syntaxa from Paekdu-San and
Hokkaido are markedly different in species composition but
analogous about their habitat conditions. The syntaxonomical
status of the Paekdu-San’s community remains unresolved
due to insufficient materials.
Viola biflora-Calamagrostis langsdorfii community
Calamagrostis langsdorfii grassland was linked to the habitats
with most favorable conditions, with almost complete absence
of gravitational disturbance and abnormal dryness. The plant
community spatially showed a linear distribution pattern
dependent on microtopography and more hospitable slopes
(2–5°) of a U-shaped valley resembling a mini cirque (Kar)
formed by glacial erosion. The ecological and floristic
constancy of the community is somewhat unique. C. langsdorfii,
Viola biflora, Sibbaldia procumbens, and Aquilegia flabellata
var. pumila are nominated as differential species. C. langsdorfii
is the dominant species and other four species are present in
higher frequency. This community is representative of the
snowbed vegetation in Paekdu-San, particularly on its northern
slope. Its geographically vicariant, Calamagrostio-Phyllodocetum
aleuticae has been reported from Mt. Tsukiyama, Yamakata
Prefecture, Japan (Nakamura 1987); it is classified within
Phyllodocion aleuticae of the unresolved order Harimanelletalia
(Fujiwara 1996). The higher syntaxonomic units of the Viola
biflora-Calamagrostis langsdorfii community are undefined,
due to not only insufficient phytosociological data from Paekdu-
San, but also to a very different species composition from that
of the Phyllodocion aleuticae in Japan. However, present
rankless unit is considered as a local vegetation type
according to diagnostic species combination. On the other
hand, a meadow community dominated by C. langsdorfii has
been described on well-drained but permanently moist
floodplain habitats in the Bol’šoj Čivyrkuj River Valley near
Lake Baikal (Chytrý et al. 1995). A circumpolar temperate
species, Sibbaldia procumbens, is typical for such humid
conditions and even frost pockets (Daniëls 1994; Franziska
et al. 2005). This creeping sibbaldia occurs only on Halla-san
summit of Jeju-do, South Korea, where their habitat is
briefly submerged during torrential rains, but it drains fast
due to medium structure of the subsurface.
Colluvial Drainage Channel Vegetation
Carex atrata-Sanguisorba stipulata var. riishiriensis community
There are small and large sizes of sedimentary colluvium on
steep and fairly gentle slope, which shows a perennial drainage
system accompanying mobile and/or stabilized soil and
scree. A total of 16 phytosociological relevés composed of
47 species were analyzed. The relevés represent much of the
variation among the phytocoena, depending on north or
south aspect. Present community occurs often in colluvial
fans and aprons of moist, gently leaning slopes around the
Cheonji caldera lake (Fig. 4). The soil layer is somewhat
developed and well drained, and often contains a significant
fraction of volcanic sandy gravel. Most sites are located
around a periodical gully directly affected by underground
and surface water. Sanguisorba stipulata var. riishirensis is

J. Plant Biol. (2019) 62:436-450 447
tied to such habitat conditions. The differential species of the
community are S. stipulata var. riishirensis, Carex atrata,
and Festuca vivipara (Table 6). The species richness, with an
average of 11.3 species per relevé, is relatively high. Some
species that favor drier conditions, such as Dryas octopetala
var. asiatica, Carex longirostrata, and Tofieldia coccinea,
were absent. Except the diagnostic species group, Sedum
roseum and Polygonum ochotense are influential to the
community showing the higher r-NCD value (3.17~1.53),
reflecting habitat situation of the rocky colluvial deposit.
The community was subdivided into three subunits:
typical subcommunity, Luzula oligantha subcommunity,
and Ligularia jamesii subcommunity. The L. oligantha
subcommunity mainly occurs on south-facing sites in the
inner caldera zone around the lake Cheonji at 2,260–2,380
m. Among differential species (listed in Table 1), various
graminoids including L. oligantha, Poa malacantha var.
shinanoana, Phleum alpinum, and Hierochloe alpina were
noticeable. Other differential species are Erigeron thunbergii
var. glabratus, Aquilegia flabellata var. pumila, and Viola
sacchalinensis var. alpicola; these are diagnostic herbs in
areas with high air moisture levels. Species richness is
relatively high. The L. jamesii subcommunity is identified by
differential species of L. jamesii, Veratrum maackii s.l., and
Saussurea triangulata. Edifiers of the community are
Ligularia and Veratrum, reflecting the highly humid soil
conditions in the valley microhabitat. L. jamesii is an
element mainly found in the ecotone between alpine tundra
and subalpine coniferous forest (Šrůtek et al. 2003). Rainwater
and snowmelt flow into the site from the neighboring sloping
areas, and vegetation is physically disturbed by occasionally
moving rock fragments of 3–5 cm in diameter. The optimal
homotoneity (H
opt
) of the present community was calculated
as 0.21, and those of its subcommunities ranged from 0.51 to
0.54. This results in a fairly different species composition
between subcommunities. In this context, we specified the
present community as a rankless unit, even if regarded as a
local vegetation type.
Materials and Methods
Study Area
Mount Paekdu (Paekdu-San) is a dormant volcano located on the
North Korea-China border region (Fig. 1). The latest volcanic eruption
had occurred in 1903 (Yun and Cui 1996) and it is expected to
explode again in the near future. It is the tallest mountain in the East
Asian continent, reaching 2,744 m above sea level. Its phytogeographical
position is separated and marginal within the Northeast Province in
Takhtajan’s phytogeographical concept (Takhtajan 1986) and contributes
as a refuge for many narrowly distributed or widespread plant taxa.
The high elevations of peaks and ridges are characterized by a
large-volume pumice deposit large and small-scale erosions and rock
Fig. 1. Location and climate of Mt. Paekdu-San (Changbaishan),
and mainly investigated areas along the ridges of the Cheonji
caldera.
Fig. 2. A generalized altitudinal scheme on the zonal vegetation in the Mt. Paekdu-San of Chinese side.

448 J. Plant Biol. (2019) 62:436-450
collapse. The surface of the Cheonji caldera lake is 2,257 m above sea
level, about 3 × 4 km across, and its depth is 375 m. The caldera slope
is very steep in 850 m deep, 5 km wide, and consists of abundant
large and small colluvial sites (Whitford-Stark 1987; Ji 2003). The
substrate is mostly composed of unstable fragile alkaline rocks such
as trachyte, trachyte-andesites, and rhyolitic pumice originated from
the stratovolcano (Ri 1996).
A recognizable altitudinal vegetation zonation of the Paekdu-San
from the cool-temperate to the alpine belt (Zhu and Rowe 1987; Yang
and Xu 2003) is given in Fig. 2. The bottom belt hosts the cool-
temperate altimontane admixed forests, which comprise Tilio-Pinetalia
of Querceneo mongolicae, characterized by deciduous broad-leaved
and evergreen needle-leaved trees. Quercus mongolica, Tilia amurensis,
and Pinus koraiensis are the canopy trees (Kim 1992). Abieti-
Piceetalia jezoensis coniferous forests develop in the subalpine
belt, abundant in conifers such as Picea jezoensis, Abies
nephrolepis, and A. holophylla. In the upper end of the subalpine
belt, Larix olgensis var. koreana and Betula ermanii form tree
islands. The distribution of Pinus pumila krummholz is hitherto
known in the south at 1800–2100 m a.s.l. on China’s Paekdu-San
area (Qian et al. 2003).
The present study was conducted in the alpine belt above this
Larix-Betula timberline (Qian et al. 1999), in wind-exposed vegetation
and dwarf-shrub vegetation, alpine heath, and snowpatch meadows.
This alpine landscape above 1,950 m a.s.l., is characterized by low-
growing plants and a mean annual temperature of about -3.8°C
(Zhang and Dai 2001). The cold winter and winds of all year round,
combined with more than 10 cm snow depth for eight months,
strongly influence vegetation development. Low-growing sub-shrubs
occur in more sheltered sites, whereas herbaceous plants and
bryophytes more in exposed sites.
Annual mean temperature and precipitation at the summit of Mt.
Paekdu (based on data collected from 1982 to 1988 by the Tiānchí
Meteorological Observatory; 2,623 m a.s.l., 42°01′N, 128°05′E) are -
7.39°C and 1,532 mm, respectively. Mean temperature of the
warmest August is 9.9°C and that of the coldest January is -24.9°C.
The temperature lapse rate was estimated roughly at -0.536°C/100 m
by reference to thermal data of the stations in Erdaobaihe (740 m
a.s.l.) and Tiānchí (2,623 m a.s.l.).
Fig. 3. Polytrichastrum alpinum-Minuartia macrocarpa var. koreana community (42°01′01″N, 128°01′24″E, altitude 2,387). The site is a
keystone habitat for the northern pika, Ochotona hyperborean group.
Fig. 4. Carex atrata-Sanguisorba stipulate var. riishiriensis
community (true green colored) at full damp sites in the slope
fringing the Cheonji Lake.

J. Plant Biol. (2019) 62:436-450 449
Field Work, Data Analysis and Tabular Work
Three times expeditions were accomplished July 1991, August 2004,
and September 2009 in the northwestern area of Paekdu-San centered
around the caldera lake Cheonji (Tiānchí in Chinese). Field investigations
were carried out at the crater rim around the Cheonji caldera. On the
Seo-pa route lined by several summits, we started the on-site
investigation from Border Milestone No. 5 (altitude 2,450 m) at 4:00
am and barely arrived at 9:00 pm, after a walking distance of 18 km,
at the Erdaobaihe village. A total of 89 relevés were collected along
the alpine ridge of the Seo-Pa route, and at colluvial sites around the
caldera lake in different days (Fig. 2). All surveyed plots are located
within a strictly protected core zone of the Changbai Mountain
Nature Reserve designated by China since 1960.
Phytosociological relevés (Braun-Blanquet 1964) were collected
with respect to the homogeneousness of species combinations and
habitat conditions. All plant species occurring in a relevé plot were
listed and their performance was determined using the 1 to 9
combined cover scale (Westhoff and van der Maarel 1978), modified
from the Braun-Blanquet cover estimation scale. Latitude and
longitude, altitude, administrative districts, micro-topography, area,
slope direction, soil, inclination, and rock type, were recorded in
every plot. The relevé sizes varied from 0.12 m to 6 m , depending
on the vegetation types. Collected relevés were classified by conventional
tabular hand-sorting method. Important diagnostic species were
chosen in order to describe a phytosociological character of plant
community. The name-giving species of a relevant community has a
high cover value, but they may also occur in other plant communities.
All communities were described without the rank, due to insufficient
materials for establishing the hierarchical syntaxa. In the structured
table, the species are arranged mainly according to synoptic value
size, i.e. relative net contribution degree (r-NCD). The net contribution
degree (0 < NCD ≤ 9), and the relative net contribution degree (0 < r-
NCD ≤ 100) were followed by Kim and Manyko (1994). A relative
homogeneousness of species composition of each community, i.e. the
optimal homotoneity (0 < H ≤ 100) were determined by Kim and
Eom (2017). The optimal homotoneity is based on the numbers of
valid species and other accidental species on the given set of relevés
structured as a syntaxon. The only single occurrence species on a
synthesized table is nominated as an accidental species, and all other
species are considered the valid species. Those formulas were
described as following.
and ,
ΣC, sum of coverage of species i
N, total number of relevés,
n number of relevés with species i,
NCD , maximum NCD value of a certain species for a
synthesized vegetation unit.
,
n, number of relevés,
VS , valid species number of relevé k, VS = S − AS ,
S, total species number of relevé k,
AS , accidental species number of relevé k.
mVS, mean valid species number per relevé,
,
tS, total species number,
tAS, total accidental species number of the given set of relevés.
Owing to the Insufficiency of Existing Phytosociological and
Synecological Knowledge of Mt. Paekdu, the classified communities
here were compared with a few syngeographical vicariants, mainly those
of Japan. We basically adopted the scientific names in the following local
literatures such as Changbaishan Plant and Animal List (CNRI 1982)
and Korean Plant Names Index (KFS 2014) for vascular plant, Choi
(1980) and Yoshimura (1994) for lichens, and Iwatsuki and Mizutani
(1984) for bryophytes, although some of them have been still unresolved
according to the www.theplantlist.org. The study area is even now a very
sensitive and non-free political area, so it is virtually impossible to study
the local flora of the entire mountain. Systematic taxonomic studies on
alpine zone of Chinese Changbaishan and Korean Paekdu-San are still in
their early stages. The collected specimens were stored in the ecological
laboratory of Keimyung University in Daegu, S. Korea.
Acknowledgements
We present special thanks to Dr. KH Ahn, Dr. GJ Cho, Dr. JC Lim,
Mr. SU Han, JS Kim, CW Lee, Mrs. YM Jeong, MS Kim, Mrs. JA Lee,
and Mrs. JE Kim for long-lasting cooperation in very tough fieldworks.
Author’s Contributions
JWK designed and accomplished the whole work from the beginning;
SYK performed the identification of nonvascular plants and
participated in the initial version of manuscript; BKC and SYK
analyzed data, tested statistics, coordinated the figures. All the authors
declare that they have no conflict of interest.
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