Professional Documents
Culture Documents
Phytosociology
of the Beech
(Fagus) Forests
in East Asia
Geobotany Studies
Editor
Franco Pedrotti
University of Camerino
Via Pontoni 5
62032 Camerino
Italy
Editorial Board:
S. Bartha, Vacratot, Ungarn
F. Bioret, University of Brest, France
E.O. Box, University of Georgia, Athens, SA
A. Carni, Slovenian Academy of Sciences, Ljubljana (Slovenia)
K. Fujiwara, University of Yokohama, Japan
D. Gafta, University “Babes-Bolyai” of Cluj-Napoca (Romania)
J.-M. Géhu, Inter-Phyto, Nouvion sur Ponthieux, France
J. Loidi, University of Bilbao, Spain
L. Mucina, University of Perth, Australia
S. Pignatti, University of Rome, Italy
R. Pott, University of Hannover, Germany
A. Velasquez, Centro de Investigacion en Sciencias Ambientales,
Morelia, Mexico
R. Venanzoni, University of Perugia, Italy
Phytosociology of the
Beech (Fagus) Forests
in East Asia
Tukasa Hukusima Tetsuya Matsui
Masato Yoshikawa Plant Ecology and Diversity Group
Hidekazu Honma Forestry and Forest Products Research Institute
Faculty of Agriculture (FFPRI)
Tokyo University of Agriculture Sapporo, Hokkaido, Japan
and Technology
Fuchu-shi, Tokyo, Japan Sandro Pignatti
Department of Environmental Biology
Takayoshi Nishio University of Rome “La Sapienza”
Weed Science Center (WSC) Rome, Italy
Utsunomiya University
Utsunomiya, Tochigi, Japan Sheng-You Lu
Taipei Botanical Gardens
Division of Forest Biology
Liang Yang
Taiwan Forestry Research Institute
Yunnan Environmental Monitoring Center
Taipei, Taiwan
Kunming, Yunnan
Republic of China
People’s Republic of China
Moon-Hong Kim Yuehua Wang
Department of Biology School of Life Sciences
College of Natura Yunnan University
Cheju National University Kunming, Yunnan
Jeju, Republic of South Korea People’s Republic of China
The beeches (genus Fagus, with a dozen of species) are widespread elements in
the woody flora of the northern hemisphere. In the mountain vegetation, beech
woodlands occupy the upper belt of compact deciduous forests, so that, at more
elevated altitudes only the evergreen needle-lived forests of boreal conifers occur.
At the boundary between deciduous and evergreen forest, the beech forests can
be observed on Mt. Fuji, in the mountains of China, in Caucasus, and in the Alps.
The only exception are the north American species of beech (F. grandifolia)
growing in the plains, under cold climate, together with other deciduous trees,
mainly maple. These mountain forests of the northern hemisphere have a counter-
part in the Nothofagus forests of Patagonia and New Zealand, which look very
similar (as to aspect and ecology) although recent results demonstrate that Fagus
and Nothofagus evolved independently from one another.
The beech forest was clearly perceived from botanists and foresters as a distinct
vegetation form, because of the compact structure of the canopy. This vegetation
was clearly described still in the nineteen century, from the most relevant phytogeo-
graphers, in Germany, Switzerland and Austria. The first concept of the beech
forest, as an essential biological and ecological unit of plant life in the mountains,
is expressed in several publications of B. Pawlowski (prof. in Krakow, Poland),
who in 1928 described the alliance Fagion sylvaticae and order Fagetalia based
on the beech forests of the Tatra mountains (Carpathians): the presence of these
coenological units was successively confirmed on all mountain systems of Central
and Southern Europe, from the Pyrenées to the Alps, Balkans and Apennines, until
the Mediterranean islands Corsica and Sicily. Independently from these develop-
ments, in the years 1949–1954, Tokio Suzuki described in Japan the beech forests
with Fagus crenata. The successive travels and field investigations by R. Tüxen
in Japan (in the 1960s) allowed the possibility to compare the parallel adaptations
(and the differences) between these important forest systems in Europe and Japan.
During the following years, several Japanese scientists visited Europe (in particular
A. Miyawaki) and scientists from Europe had the possibility to be introduced to the
study of the vegetation of Japan (for the writer of these lines, the first experience
was during the memorable International Excursion of 1974). The following
discussions and exchange of experiences (in many papers, meetings and in the
v
vi Preface
field) led to the persuasion that the beech forests on both extremities of Eurasia had
similar composition and ecology. From this background, develops the personal
experience of prof. T. Hukusima, the first Author of this book, who in the period
1980–1990 elaborated a synthesis of beech forests in Japan and successively carried
out many research periods investigating directly in the field the beech forests of
Europe, SW-Asia and N-America, and in particular with excursions in different
countries of E-Asia (Korea, continental China, Taiwan). These field investigations
had a focal point when both Hukusima and myself had the possibility, at the
beginning of November 2003, to investigate the habitat of the rare Fagus mexicana,
growing in a remote chain of the Sierra Madre Oriental.
It has been necessary as shown above to briefly summarize the long historical
development, which lead to the origin of this book. It was realized with the
collaboration of leading scientists from different East-Asiatic countries, the follow-
ing elaboration of data and discussion of the results were carried out successively,
during repeated stages of prof. T. Hukusima in the Botanical Garden of the Rome
University. This study gives, for the first time, the possibility to have a general
outlook on the different aspects of the beech forests in East Asia, from Hokkaido to
Taiwan and from the coasts of the Pacific Ocean to the easternmost chains of the
Himalayan system. This large synthesis gives a general information on over 50
different types of beech forests, distributed in two vegetation classes with 22 types
of forest communities. Over 1,500 species of the forest flora of East Asia (trees,
shrubs, herbs, mosses) are reported. In this way, it is shown clearly the extraordi-
nary biodiversity concentrated in the deciduous forests of the East-Asiatic
mountains. From this synthesis, it is also possibly to propose a hypothesis on the
evolution of the beech forests, based on the central role played by the mountain
systems of SW China (Yunnan) as an ancient centre of origin for botanical groups:
many of them in the following eras spread over most of the boreal hemisphere and
presently are an essential component of the flora in the temperate and cold zone of
Eurasia.
Sandro Pignatti
Acknowledgements
This study was possible because it is based on the previous studies performed
by many senior researchers and associate researchers. Especially, it would not
have been possible for us to understand Chinese beech forests, without the
previous pioneer work performed by Wang, Z.X. of College of Resources and
Environmental Science, Hubei University, former Prof. Dr. Kazue Fujiwara of
Yokohama National University and her colleagues. We here express our respect
and gratitude for their precise research results. We are also grateful to our
colleagues, Hiroyuki Takasuna, Yutaka Tsunetomi and Yutaka Kyan. This study
was partly funded by the Global Environmental Research of Japan (S-8) program,
the Ministry of the Environment. Lastly we wish to dedicate this work to our
mutual masters, the late Prof. Dr. Tokio Suzuki and the late Prof. Dr. Hyoji Suzuki,
both of them contributed greatly to the development of vegetation science in Japan.
This research was partly supported by the Environment Research and Technology
Development Fund (S-8) of the Ministry of the Environment, Japan.
vii
.
Contents
ix
x Contents
Appendix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
Appendix 1. Alphabetic List—Species of the Beech Forests
in East Asia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
Appendix 2. Taxonomical Outlook on the Flora of the East Asiatic
Beech Forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 255
List of the Tables
xi
Phytosociology of the Beech (Fagus)
Forests in East Asia 1
1.1 Introduction
The beech (genus Fagus) is often the dominant species in forests in many locations
and is an extremely important species to local ecosystems. In East Asia, forests
including species of Fagus form, in a phytogeografical vision, an extended forest
belt in the zone with temperate climate, that lies between the evergreen broad-
leaved forests spreading out across the lower latitudes of the tropics and the
evergreen coniferous trees (boreal forests) of the northern latitudes. This can be
also considered, in an altitudinal vision, as a horizontal forest belt that also lies
between those two forest types in a vertical sense. In general, beech forests occur in
areas with oceanic climate (at least moderately) and avoid conditions of elevate
continentality. Because of their location, East Asian forests containing species of
the genus Fagus can help clarify, through a comparative study of East Asian
vegetation, the characteristics of beech forests in the Northern Hemisphere (Eurasia
and North America) as well as play a crucial role in providing an understanding of
East Asian vegetation, thus making the beech a key vegetation type.
In addition, although the distribution of the species is wide, beech forests share
their temperate zone habitat with man and in many locations suffer a good deal of
disturbance from human activity. At present, they continue to represent a valuable
resource, providing both material for human habitations and pasturage for stock.
However, severe over-usage has caused a rapid reduction in beech forests in many
areas. East Asia is no exception.
In such an environment, it is extremely important to precisely record the
characteristics of beech forests. Since the 1950s, vegetation studies using phytoso-
ciological procedures have been carried out in each country in East Asia, and the
floristic compositional characteristics of the beech forests distributed over each
country have been clarified. However, in terms of study range, these studies were
focused mainly within each individual country, and until now only limited compar-
ative research has been undertaken. Therefore, there has been little overall study of
the homogeneity or heterogeneity of Asian beech forests as a whole. The present
advance in the accumulation of data for the study of Fagus dominated forests in
each country provides a more comprehensive range from which the compositional
characteristics of the beech forests in East Asia, as a whole, can now be determined.
In this study, we compared the known data on East Asian forests that include
species of the genus Fagus with new data that we obtained from the areas where
little previous research had been conducted. Through this comparison, we aimed to
define the phytosociological characteristics of the beech forests in E-Asia, assess
the forests phytosociologically, and systematically classify the plant communities
therein.
East Asia forms the core area of the worldwide distribution of species of the genus
Fagus, Many species of Fagus are distributed across this region (Fig. 1.1). Two
species of Fagus, (Fagus crenata Blume and F. japonica Max.) are distributed over
Japan, although, generally, when the ranges of these species overlap, F. japonica
tends to be found at lower elevations than F. crenata. Differences can be seen in
both their range and ecological adaptations. F. crenata is single-trunked, and
seldom coppice; however, the F. japonica is sparse-leaved, multi-trunked and
coppice vigorously, but is short-lived. F. crenata ranges from about 31 300 N to
42 450 N, covering an area that stretches from Mt. Takakuma in Kagoshima
Prefecture in southern Kyushu, across Kyushu, Shikoku and Honshu, to the south-
ern area of Hokkaido. F. japonica ranges from approximately 32 N in Miyazaki
Prefecture in Kyushu, over Kyushu and Shikoku, to 40 N of Honshu (Ishizuka et al.
1992), but with the exception of the Chugoku region, it is not found along the Japan
Sea side of the country. This species is seldom dominant and it tends to be
distributed along the Pacific Ocean side of Japan, where it often forms forests
with Abies firma, Tsuga sieboldii, F. crenata and Quercus mongolica var.
grosseserrata.
Fagus multinervis Nakai is found in Korea. However, this species is not
distributed over the Korean Peninsula, but is restricted to Ulleungdo, an island
140 km to the east of the Korean Pensinsula in the Japan Sea (130 500 E, 37 300 N).
This is a small volcanic island, with a land area of 73 km2, known to have erupted
around 9,300 years ago. The highest point on the island is Seonginbong Peak
(983 m), which occupies one corner of the outer rim of the volcano. The beech
trees coppice, and resemble F. japonica in shape. The form of the leaf is also similar
to that of F. japonica, but the cupules and form of the seeds are different from those
of F. multinervis, being generally larger. The trees are distributed above an eleva-
tion of 350 m, and are dominant across large areas. Because steep slopes dominate
the island’s topography, many valleys are formed, and beech forests range almost to
the bottom of the gorges, although no ravine forests, such as found in Japan, are
present. The island is located at the meeting point of warm and cold ocean currents,
and there are foggy conditions throughout the year. The yearly average humidity is
1.2 Fagus Species and Associated Forests in East Asia 3
Fig. 1.1 Study site locations (A to Q) and geographical features of the study area. Note that the
site B is in the areas enclosed by the dashed-dotted line in Japan
likely to exist also between the various types of forest communities, each of them
with the dominance of one among the four Fagus species (a – b – c – d), because of
the high diversity in the phytogeographical elements distributed over the different
regions of East Asia.
forests classes such as the coniferous Abies firma forests or to the deciduous broad-
leaved Quercus forests. Therefore, we extracted relevés with strong relations to F.
crenata forests for comparison of species composition with the other beech forests.
The results showed that in mixed stands, where F. crenata and F. japonica grow
together, many lower units can be distinguished, with species differing quantita-
tively; however, these do not represent compositionally autonomous plant
communities. In consequence, these mixed communities were considered as
lower units and placed at the subassociation level of the F. crenata forest
association.
In the Korean Peninsula or Jeju Island, both of which are in close proximity to
Japan, no species belonging to the genus Fagus are present, although in mountain
areas favourable ecological conditions would exist, at least in terms of elevation. In
Korea, F. multinervis Nakiai is distributed only over the isolated Ulleungdo Island,
located in the Sea of Japan, but under the Korean souverainity. Kim et al. (1986)
and Kim (1988) previously undertook studies of the island and reported on its plant
community with Fagus dominance. For the present analysis, we added the data
collected by the authors to that of Kim et al. (1986) and Kim (1988) to obtain a total
of 54 relevès for the association table.
As to the Fagus woodlands in China, Wang and Fujiwara (2003) published a
synoptic table based on data collected in the beech forests such as Fagus
longipetiolata Seem., F. engleriana Seem., F. lucida Rehd. et Wils. and Fagus
hayatae ssp. pashanica C.C. Yang. The relevés were obtained from forest
communities in southwest Hunan Province and the border of Hunan and Guangxi
Province (Nanzan), Fanjingshan Nature Reserve and Kuankuoshui Nature Reserve
in Guizhou Province, northwest Hunan Province, the Badagongshan Nature
Reserve on the border of Hunan and Hubei Province; the Dalaoling forest station,
Longmenhe forest station, and Houhe Nature Reserve in Hubei Province; the
Baotianman Nature Reserve in Hunan Province, and the Sihaishan Nature Reserve
and Qingliangfeng Nature Reserve in Zhejiang Province. Thereafter, Wang et al.
(2005) reported on the plant communities and classification of forests with F. lucida
and F. engleriana as dominant tree. An original vegetation survey table with a list
of all species appearing, except for data from the Sihaishan Nature Reserve and
Qingliangfeng Nature Reserve in Zhejiang Province, was included. In the present
analysis, data for the two nature reserves in Zhejiang Province were taken directly
from synoptic tables, while we prefer to use the original data from Wang et al.
(2005) for all other areas. Where the study area of Wang et al. (2005) overlapped
with that of the authors for the Fanjingshan Nature Reserve in Guizhou Province,
however, we consolidated the data to create a larger association table. Relevés from
the Sanjiangkou Nature reserve in the north of Yunnan Province, Wenshan and
Xichou counties in the south of Yunnan Province, and Nanjiang country in the north
of Sichuan Province, were all collected by the authors. Raw data from Yunnan
Province are included in Wu et al. (1987), and although these data do not represent a
complete species list, it does provide representative members of the species in each
forest level. Consequently, we decided to include it in this analysis, because these
data were very valuable for these areas, in which no other vegetation survey data
1.4 Classification of the Plant Communities 7
sets were available. We then reached a gross total of 108 Chinese relevès in the
association table.
Because there were no data available for comparison, some of the authors visited
Taiwan to undertake a study in cooperation with researchers from the Taiwan
Forestry Research Institute. In this study we were able to obtain 23 relevès,
distributed on all stands of the Taiwanese beech forests at Mt. Chulu, Mt.
Peichatien, Mt. Lala and Mt. Tonshan. The results were published in Hukusima
et al. (2005).
The present study, is based on the total of 657 relevés, partly collected in the
field by the authors and partly from the literature. These relevés were then used to
produce the general synoptic table, association tables and to select the character
species at all syntaxonomical levels, from the vegetation classes to orders, alliances
and associations.
All published data, as well as our original data have been assembled in association
tables, and successively organized into synoptic tables. In the first elaboration, we
closely examined the data contained in published or unpublished association tables,
and corrected the data for plant communities presented therein to create synoptic
tables for comparison. After this resorting, we created a general synoptic table with
the synthesis for the entire plant communities including all types of the beech
forests until now described for East Asia (Table 3.1a, 3.1b, 3.1c [Online]). Synoptic
tables are the ideal vehicle to obtain the complete overview of all species connected
with a given vegetation unit. However, it has a sense to give a complete list of all
species only in the case that data derive from a small number of study units. When
there are a large number of study units to consider, as in this case, the large number
of species appearing only a few times (sporadic species) can become a problem. In
this case, we listed only species with an incidence of 2 % or greater, whereas the
merely occasional species were not more taken into consideration.
With regard to the identification of associations, we examined the units previ-
ously reported at the level of associations, and, respecting these previous results as
far as possible, we created a new ordered list of the associations described in the
literature. However, with regard to plant communities with different compositions,
we closely examined the character species and consequently, we arrived to an
improved definition for a number of associations. This method was successively
used for the superior hierarchical levels of alliance, order and class. With regard to
orders and above, we basically used the same method as that used to study
associations and alliances; however, we also undertook a comparison with adjacent
orders and classes reported from each study area.
From the analysis of the climatic conditions (Table 3.5), it results that, in the
beech forests the differences in temperature conditions are quite significant between
the different classes of vegetation, whereas differences in precipitation are more
8 1 Phytosociology of the Beech (Fagus) Forests in East Asia
important in the definition of the growth activity of the forests, at the order and
alliance level.
The analysis of the species composition, genus composition, life form composi-
tion and the examination of the relationships between the classified vegetation units
and climatic conditions overall elucidate the clear differences in characters of the
proposed vegetation classes of beech forests in East Asia. Moreover, the classified
orders and alliances show, for the most of the case, clear differences and unique
characters in terms of species, genus and life form compositions. On the other hand,
associations and subassociations show characters which are distinctive only within
a much smaller context. The two last associations (No. 21–22) are differing in many
characters and belong to a still incompletely defined vegetation class. These results
are all suggesting that, when vegetation units are compared, the evaluation of
species composition (raw data) is the most important procedure at the association
and subassociation level, whereas the analysis of genus and life form composition,
such as similarity and dissimilarity (meta-data), are important for the classification
at the order or class level.
The comparisons at genus-level show that the combinations of genera are most
informative feature for a distinction among the classes. The differences are most
accentuated between Ulleungdo Island in Korea and Japan, which have a prevailing
proportion of deciduous genera; the same can be observed in a comparison between
the beech forests of China and Taiwan, which have higher proportions of evergreen
genera. The same tendency was also found at the level of the five orders. However,
at alliance level, distinct genera became less obvious, and furthermore, at associa-
tion and subassociation level, no important differences at the genus level were
found.
For the comparison of the floristic composition of the different phytosociological
units, Table 3.1 is fundamental. Indeed, the consultation of this table is very
difficult, because of his unusually large dimensions (over 1,500 rows by more
than 50 columns). For this reason, Table 3.1 was divided into Table 3.1a (Korea
and Japan) and 3.1b (China and Taiwan); a general outlook is possible in Table 3.1c
(On line) where the presences III, IV and V (40–100 %) are indicated with a black
square and presences 1–39 % with a point. Alphabetical lists of the species as well
as a taxonomical outlook of the flora of the East Asiatic beech forests are given in
the Appendix 1 and Appendix 2.
Syntaxonomy of the East Asiatic Fagus
Forests 2
Table 3.1 is a synoptic table of the forests dominated by the genus Fagus in East
Asia. According to Table 3.1, the phytosociological system of beech forests in East
Asia was classified as in Table 3.9 and the distribution map of the classified classes,
orders, alliances and associations was depicted as Fig. 2.1. In Table 3.1, 68 species
categorized as the species group 49 are the common species occurring in the beech
forests in East Asia, although there is a regional bias. Many of the 68 species are
character species of beech forests in each region.
Species constantly found in the beech forests in Ulleungdo Island are:
Dryopteris crassirhizoma, D. fortunei var. radicans, Disporum sessile, and Viola
selkirkii.
Commonly found in the beech forests both in Taiwan and Japan are: Viburnum
furcataum, Rhus ambigua, Ardisia japonica, Parathelypteris japonica.
Species distributed in China, Taiwan and Japan, but not in Ulleungdo Island are:
Hydrangea paniculata, Sorbus alnifolia, Oxalis griffithii, Acer mono, Athyrium
wardii, Clethra barbinervis, Ilex macropoda, Lindera umbellata, Cornus kousa,
Vaccicinium japonicum. Moreover, these species are highly frequent in the F.
crenata forests in Japan.
Common species in the beech forests in China and Taiwan are: Daphniphyllum
macropodum, Ardisia crenata, Cleyera japonica, Maesea japonica, Plagiogyria
euphlebia, Skimmia reevesiana, Peracarpa carnosa, Symplocos sumuntia, Smilax
lanceofolia var. opaca. There are many evergreen, broad-leaved species. Moreover,
Daphniphyllum macropodum and Ardisia crenata in Japan are growing in the
evergreen, broad-leaved forests rather than in the beech forests.
In the synoptic table (Table 3.1c), it is possible to have an outlook on the
classification of the beech forests in East Asia into 51 vegetation units. Each of
the units was compared with each of the vegetation unit that have been proposed in
the published literature, with regards to the similarities of the species composition.
As a result of this comparison, we were able to organize the phytosociological
Fig. 2.1 Distributions of classes, orders, alliances and associations for the beech forests in
East Asia
classification, including new vegetation units (Table 3.9, Fig. 2.1). For the East
Asian forests with dominance in the tree layer of species of the genus Fagus, it was
possible to organize the classification of the plant communities in the following
syntaxonomical treatment.
The first vegetation class (Fagetea crenatae) includes all beech forests
recognized in Japan and on the Korean Ulleungdo Island, associations no. 1–6,
distributed among 2 orders and 3 alliances.
The second vegetation class (Litseo elongatae-Fagetea) includes the beech
forests with prevalence of deciduous species, recognized in China: associations
7–20, distributed among 2 orders and 4 alliances.
The third vegetation class is still incompletely known (and consequently still
unnamed); it includes the beech forests with prevalence of evergreen species,
recognized in South China: associations 21–22, belonging to 1 order and 1 alliance.
In the following chapters, all these vegetation units are described and discussed.
F. crenata forests and F. multinervis forests on Ulleungdo Island, Korea, fit into this
class. In addition, the composition of the forests distributed over Jeju Island, the
Korean Peninsula and northeastern China, where species of Fagus are not present,
was reported instead to be with Quercus mongolica as dominant tree. A comparison
of the forest composition of the Q. mongolica forests of Jeju Island (Yun et al.
2008), reveals the presence of many common species, suggesting those forests
could be included in this class. Song (1988) defined this class as Quercetea
mongolicae (Song 1988) from a study of the Korean conifer-broad-leaved mixed
forests, and this was supported by Takeda et al. (1994). On the other hand, in the
similar Quercus mongolica forests in North Korea, Kolbek et al. (2003) revised the
concept of the Querco-Fagetea crenatae (Miyawaki et al. 1968) proposed by
Miyawaki et al. (1968). In northeast China, Wang et al. (2006) recognizes
Quercetea mongolicae (Song 1988). Wang et al. (2006) also defined Querco
mongolicae-Betuletea davuricae (Wang et al. 2006) as a new independent class.
A comparison of the reports by Kolbek et al. (2003) and Wang et al. (2006) with the
association tables for beech forests presented in this study shows that many species
not found in Fagus classes, such as Quercus mongolica, Vitis amurensis, Carex
nanella, Lespedeza bicolor, Pinus koraiensis, Philadelphus schrenkii, Tilia
amurensis, Maackia amurensis, Athyrium crenatum, Artemisia keiskeana, Deutzia
glabrata, Rhododendron mucronulatum, and Polygonatum involucratum, are com-
mon and are present at characteristically high constancies. Furthermore, there are
many species that are not distributed in beech forests, indicating that the composi-
tional differences are large. Hence, this class can be judged to be a distinct class.
However, an interesting point to note is that where Quercus mongolica var.
grosseserrata forests (Hoshino 1998; Suzuki 2002) come into contact with beech
forests at lower elevations in Japan, many species common to the Quercus
mongolica forests are found. To explain these relations, it is necessary to examine
not only current climatic factors but also historical factors dating from the Quater-
nary period.
As to the floristic composition, it has to be pointed out that the proportion of
deciduous flora is high, and that of evergreen broad-leaved flora is extremely low in
this class. This represents a major difference between the characteristics of this
class and the Litseo elongatae-Fagetea sp. div.cl. nov. class seen in beech forests in
China and Taiwan.
This class includes two orders: Fagetalia multinervis (Kim et al. 1986) (F.
multinervis order) defined by Kim et al. (1986) on Ulleungdo Island (Korea) and
Saso Fagetalia crenatae (Suzuki 1966) (Fagus-Sasa order) in Japan defined by
Suzuki (1966). Many of the character species of this order are common to the
two abovementioned orders; however, Tripterospermum japonicum, Euonymus
alatus fo. ciliato-dentatus, Smilax nipponica, Styrax obassia, Mitchella undulata,
Maianthemum dilatatum, Asperula odorata, Polystichum retroso-paleaceum and
Viola kusanoana are limited to the beech forests distributed on Ulleungdo Island
and in Japan on the Japan Sea side of Honshu Island, and are extremely rare on the
Pacific Ocean side of Japan, suggesting that Ulleungdo Island and the Japan Sea
side of Japan were historically closely connected.
12 2 Syntaxonomy of the East Asiatic Fagus Forests
Type relevé: Kim et al. (1986), Tab. 1, Relevé reference number 17 (Elevation
820 m, Ulleungdo).
This association was recorded and named by Kim et al. (1986) on Ulleungdo
Island (Figs. 2.3, 2.4 and 2.5). Even the relevés published in Kim (1988) were
carried out on this island. The climate of Ulleungdo is warm and humid. The annual
mean temperature of 11.5 C at 357 m elevation (corresponding to 8 C at 1,000 m,
but the island has not such elevate mountains) is relatively high in comparison with
temperatures of the beech forests in Japan (Table 3.4); rainfall is relatively low
(1,371 mm), indeed the elevate atmospheric humidity is maintained by the isolated
location in the middle of the sea. The beech forests distributed on the highlands
above an elevation of 350 m have been little impacted by humans, and remain in a
predominantly natural state.
The Hepatico-Fagetum multinervis is characterized by the abovementioned
character species with Fagus multinervis, Hepatica maxima and Tilia insularis,
all endemic to the island. In addition, although they are not character species,
Dystaenia takesimana, Acer okamotoana, Ligustrum foliosum and Prunus
takesimensis are also endemic to Ulleungdo, indicating the high endemicity of the
flora in this island.
Pteridophytes, such as Rumohra standishii, Polystichum tripteron and
Polystichum retroso-paleaceum, which are character species in Japanese ravine
forests, are predominant on the forest floors in this association. This confirms that
this island experiences a warm, wet, typical oceanic climate. Kim et al. (1986)
sorted this association into the Rumohra standishii subass. (Running number. 1),
Sasa kurilensis subass. (Run. no. 2), typical subass. (Run. no. 3) and Rhododendron
brachycarpum subass. (Run. no. 4), with Kim (1988) later recognizing the same
subassociations. Our addition of new data obtained by the authors to the previous
data published by Kim et al. (1986) and Kim (1988) resulted in the recognition of an
additional subassociation: Celtis jessoensis subass. (Run. no. 5).
The structural characteristics of this association include the dominance of
deciduous species such as Fagus multinervis, Acer okamotoana and A. takesimense
14 2 Syntaxonomy of the East Asiatic Fagus Forests
in the canopy layer, with herbaceous species such as Allium victorialis var.
platyphyllum, Hepatica maxima and Maianthemum dilatatum and pteridophytes
such as Rumohra standishii, Polystichum tripteron and Polystichum retroso-
paleaceum forming the forest floor. Evergreen trees and shrubs have a reduced
presence (1–3 %). Very significant is the high presence of geophytes with rhizomes
(30 %) or with bulbs (4 %), the highest values in the beech forests of East Asia.
Very diffused are also the perennial herbs (hemicryptophytes) with a total of over
33 % and climbers (12.5 %).
Fig. 2.6 Distribution map of the five beech forest associations in Japan
these species originate from areas on the Pacific Ocean side of Japan and have
adapted to the climate on the Japan Sea side, where the snowfall is much greater. In
this region, the climate is cold-temperate, with annual mean temperatures of 6 to
9 ; heavy snow falls in winter are frequent with up to 2 m and more.
The associations included in this alliance are characterized by the prevalence of
broad leaved deciduous trees (ca. 26 %) and shrubs (23 %), together with perennial
herbs (Hemicryptophytes 27 %, Geophytes 20 %). The alliance Fagion crenatae
includes two associations: Saso kurilensis-Fagetum crenatae (Suzuki 1949) and
Lindero umbellatae—Fagetum crenatae (Horikawa et Sasaki 1959).
2. Saso kurilensis-Fagetum crenatae (Suzuki 1949) (Run. no. 6–10)
Character species: Rhododendron albrechtii, Acer mono var. mairii; Acer
tchonoskii, Streptopus streptopoides var. japonicus.
Type relevé: Hukusima et al. (1984), Tab. 2, Relevé reference number HB 40, in
Mt. Yulap (Elevation 490 m, Hokkaido).
This association (Figs. 2.7, 2.8, 2.9, 2.10, 2.11, 2.12, 2.13, and 2.14) is distributed
across the Hokuriku and Tohoku regions, and on the Oshima Peninsula in Hokkaido
from 36 to 43 N and approx. 136 to 142 E. The association, described by Suzuki
(1949a) for the Hokkaido area, occurs in the central and northern regions of Honshu
and in southern Hokkaido, mainly on the Japan Sea side, from 200 to 1,700 m in the
south and close to sea level in the northernmost areas. This climax community is the
most typical beech forest in Japan, very uniform in species composition and occur-
ring over large areas.
In this association, Hukusima et al. (1995) distinguished five subassociations:
typicum (Run. no. 6), carpinetosum (Run. no. 7), aesculetosum (Run. no. 8),
abietetosum (Run. no. 9) and sasetosum (Run. no. 10).
Canopy layer is always dominated by Fagus crenata, and sub-canopy layer with
Acer japonicum and A. tchonoskii. Upper shrub layer always with dominance of
Sasa kurilensis and other species including Lindera umbellata and Viburnum
furcatum. Lower shrub layer with evergreen broad-leaved shrubs, such as Ilex
leucoclada, Ilex crenata var. paludosa, Cephalotaxus harringtonia and Aucuba
japonica var. borealis, together with Carex dolichostachya and Smilacina japonica.
In the sasetosum subassociation, which grows on the flat mountain ridges with
volcanic ash soil in the Kitakami Mountains, Tohoku region, the shrub layer
occasionally lacks scrub bamboo grasses and is dominated by forbs.
The ecological conditions for this association are characterized by very intense
rain (1,700–3,000 mm yearly rainfall) and in the cold season heavy snowfalls: up to
3 m.
3. Lindero umbellatae—Fagetum crenatae (Horikawa et Sasaki 1959) (Run.
no. 11–15)
Character species: Euonymus lanceolatus, Torreya nucifera var. radicans,
Cryptomeria japonica, Menziesia ciliicalyx.
Type relevé: Sasaki (1964), Table 6, Relevé reference number Mt. Daisen, no. 4
(Elevation 1,100 m).
This association is distributed in the Chugoku region on the Japan Sea side of
Japan and on the lowlands in the Hokuriku area of Chubu, in a range from 34 200 to
18 2 Syntaxonomy of the East Asiatic Fagus Forests
37 400 N and from 131 800 to 137 100 E. It was recorded by Horikawa and Sasaki
(1959) in the Geihoku area in Hiroshima prefecture, located in the Chugoku region,
situated in western Honshu. As few of the mountains in this area have sufficient
elevation for F. crenata forests to grow, the associations are observed only sporadi-
cally. In addition to the character species, this association contains character species
of the Fagion crenatae (Suzuki 1952) alliance, although fewer of these character
species are observed towards the west. On the other hand, this association on the
Pacific Ocean side blends with an increasing number of species from the
Sasamorpho-Fagion crenatae (Miyawaki et al. 1968) alliance. Four subassociations
are classified under this association: typicum (Run. no. 11), polystichetosum (Run.
no. 12), rhododendretosum (Run. no. 13) and torpetaleietosum (Run. no. 14). The
F. japonica forests (Figs. 2.15, 2.16, 2.17, and 2.18) have been classified as the
association, Torreyo-Fagetum japonicae (Nakanishi et al. 1970). However, because
of the similarity in their composition with the Lindero umbellatae-Fagetum
crenatae, we now classify it as Lindero umbellatae-Fagetum crenatae torreyetosum
(Run. no. 15), a subassociation of this association. Although the composition is
basically similar to Saso kurilensis-Fagetum crenatae (Suzuki 1949), Cryptomeria
japonica is occasionally seen in the canopy layer in this association.
The ecological conditions for this association correspond to a moderately cool,
rainy climate, with yearly mean temperatures of ca. 8 C at 1,000 m elevation and
average rainfall 2,300–2,700 mm (Table 3.4). Snowfall during the cold season is
lesser intensive than in Saso kurilensis-Fagetum crenatae, and mostly remains in the
range of 1 m to about 2 m.
B-b. Sasamorpho-Fagion crenatae (Miyawaki et al. 1964) (Run. no. 16–27)
Character species: Sasamorpha borealis, Abies homolepis, Acer palmatum var.
amoenum, Rhododendron quinquefolium, Abelia spathulata, Acer tenuifolium,
Stewartia monadelpha, Carex fernaldiana.
This alliance (Miyawaki et al. 1964) is distributed across all areas of Kyushu and
Shikoku, and the Pacific Ocean side of Honshu, in a range from 31 300 to 39 500
2.2 I. Fagetea crenatae (Miyawaki et al. 1964) (Run. No. 1–27) 21
and 130 200 to 142 E. Although the distribution area of this alliance extends over a
wide area, the mountain environment suitable for beech forest growth does not
continue throughout the region; therefore, the associations of this alliance appear
sporadically and the species composing each plant community reflect the flora of
each region. The alliance is characterized by the character species noted above;
however, Stewartia monadelpha and Carex fernaldiana are not observed in the
Sasamorpho-Fagetum crenatae (Suzuki 1949).
The associations of this alliance develop under conditions of cool, rainy climate,
but, as a difference to the previous alliance (Fagion crenatae), during the cold
season snowfall is relatively limited and in average not reaching 1 m. Average
yearly temperatures (Table 3.4) in general are ca. 8 C at 1,000 m elevation,
whereas rainfall is variable (see the different associations).
In the canopy layer, evergreen broad-leaved species such as Quercus acuta, Q.
salicina, Illicium religiosum and Eurya japonica appear among F. crenata, as well
as evergreen conifer species such as Abies firma, Tsuga sieboldii and Torreya
nucifera. Although the forest floor is often dominated by Sasamorpha borealis,
2.2 I. Fagetea crenatae (Miyawaki et al. 1964) (Run. No. 1–27) 23
Sasa nipponica prevails on the forest floor in the Kanto region. As these scrub
bamboo species have a high degree of coverage, the herb layer is often not well
developed.
4. Sasamorpho-Fagetum crenatae (Suzuki 1949) (Run. no. 16–19)
Character species: This is the typical association without character species.
Type relevé: Suzuki (1949b), Table 2, Relevé number. M5, Tonakayama/
Shirokura National forest, Shizuoka Prefecture (Elevation 1,490 m).
This association is known from the inland areas of central Honshu to the inland
Kanto regions and the Pacific Ocean side of the Tohoku region, in a range from 35
100 to 39 500 and 137 200 to 142 E. This association, recorded by Suzuki (1949b),
is the typical association showing the basic characteristics of the alliance,
Sasamorpho-Fagion crenatae. As this association occasionally adjoins Saso
kurilensis-Fagetum crenatae (Suzuki 1949) on the Japan Sea side, it often contains
the main species of that association, such as Acer japonicum, Dryopteris sabaei,
Prunus grayana, Vaccinium japonicum, Lindera umbellata var. membranacea and
Acer palmatum var. matsumurae. However, Stewartia monadelpha and Carex
fernaldiana, which are character species of the alliance, are seldom recorded.
Hukusima et al. (1995) classified this association into three subassociations;
typicum (Run. no. 16), aucubetosum (Run. no. 17), and enkianthetosum (Run. no.
18). In addition, the floristic composition of the F. japonica forests distributed over
Mt. Oomuro at the foot of Mt. Fuji, suggests that these forests are to be included in
the Sasamorpho-Quercetum crenatae association, as an additional subassociation
oxalietosum (Run. no. 19). Species richness in the canopy layer is high, with
evergreen conifer species Abies homolepis, Tsuga sieboldii and Abies firma,
along with Quercus mongolica var. grosseserrata. However, the sub-canopy layer
is not well developed. The shrub layer is made up of scrub bamboo such as
Sasamorpha borealis and Sasa nipponica. Therefore, the development of the herb
layer is extremely poor.
The inland areas where Sasamorpho-Fagetum develops are humid, but the total
yearly rainfall (1,300–2,000 mm) is inferior than in the areas of the following
associations (3,000 mm and more).
5. Corno-Fagetum crenatae (Miyawaki et al. 1964) (Run. no. 20–22)
Character species: Prunus incisa, Parabenzoin praecox, Plectranthus
umbrosus, Aster ageratoides var. harae f. leucanthus, Aster dimorphophyllus,
Enkianthus campanulatus.
Type relevé: Miyawaki et al. (1964), Table 2-13, Relevé reference number 52,
Mt. Shiragatake in Tanzawa, Kanagawa Prefecture (Elevation 1,380 m).
This association occurs in a limited area in central Honshu on the Pacific Ocean
side of Japan, in a range from 34 500 to 36 N and from 137 500 to 139 100 E. The
association, named by Miyawaki et al. (1964), is distributed across an area referred
to as the Fossa Magna, where relatively recent volcanic mountains, such as Mt.
Fuji, are situated. Reflecting the moist oceanic climate of the area and elevate
rainfall, this association contains species that form ravine forests (Fraxino-
Ulmetalia Suz.-Tok. 1967), such as Viola bissetii, Aster ageratoides var. harae f.
leucanthus and Plectranthus umbrosus. Under this association, Hukusima et al.
24 2 Syntaxonomy of the East Asiatic Fagus Forests
(1995) classified three subassociations; typicum (Run. no. 20), sasetosum (Run. no.
21), and cacalietosum (Run. no. 22). Although F. crenata is dominant in the canopy
layer, Cornus kousa often grows in the sub-canopy layer. Sasamorpha and Sasa
hayatae are dominant in the shrub layer. However, in the subassociations of
sasetosum and cacalietosum, which develop on flat ridges with thick volcanic ash
soil, the shrub layer lacks scrub bamboo and is dominated by forbs.
6. Sapio japonici-Fagetum crenatae (Sasaki 1970) (Run. no. 23–27)
Character species: Parabenzoin trilobum, Enkianthus cernuus f. rubens, Styrax
shiraiana, Lindera sericea var. glabrata, Hydrangea luteo-venosa.
Type relevé: Miyawaki (1981), Table 132, Relevé reference number C80, Mt.
Kunimi, Shiiba Village, Miyazaki Prefecture (Elevation 1,370 m).
This association (Figs. 2.19, 2.20, 2.21, and 2.22) occurs in Kyushu, Shikoku,
the Kii Peninsula, and the Tokai region on the Pacific Ocean side of Japan, in a
range from 31 300 to 35 100 N and 130 200 to 137 500 E. The climate (Table 3.4) is
relatively warmer than in the previous associations (mean temperature 9–10 C at
1,000 m elevation), with abundant rainfall (up to 3,000–3,600 mm yearly). This
association contains evergreen species, more frequently as usual among the F.
crenata forest associations in Japan. Evergreen broad-leaved species such as
Quercus acuta, Q. salicina and Illicium religiosum, as well as evergreen conifer
species such as Abies firma and Tsuga sieboldii normally occur in the canopy and
sub-canopy layers of this association; in the shrub layer, evergreen species such as
Pieris japonica, Skimmia japonica, Eurya japonica, Neolitsea sericea and Camellia
japonica are often present. Another characteristic of this association is that some
species are closely related to species widespread in beech forests of southwestern
China, such as those seen in Yunnan Province, Sichuan Province, and Guizhou
Province. Hukusima et al. (1995) in this association distinguish four different
subassociations: typicum (Run. no. 23), occurring throughout all the area of distri-
bution, leucosceptretosum (Run. no. 24), which is diffused only in Kyushu,
cacalietosum (Run. no. 25) in Shikoku, and aceretsum (Run. no. 26) observed in
Shikoku and the Kii Peninsula. Although further evaluation is still required, in this
2.2 I. Fagetea crenatae (Miyawaki et al. 1964) (Run. No. 1–27) 25
2.3 II. Litseo elongatae-Fagetea sp. div. cl. Nov. (Run. No. 28–48)
Sinarundinaria nitida (¼Fargesia nitida), are the preferred food of the Giant
Panda, and consequently the communities bearing a dense thicket of this bamboo,
often included in natural reserves or National Parks, are among the most important
factors for the survival of the remaining populations of this mammal, worldwide
considered an icon for species highly endangered of extinction.
This order contains four alliances (see Fig. 2.1):
C-a. Abelio englerianae—Fagion all. nov. (associations 7–8) in the Shaanxi
and Sichuan Provinces, North-Western China.
C-b. Aceri davidii—Fagion lucidae (Wang et al. 2005) (associations 9–14) in
the Guizhou, Guangxi, Henan, Hubei Provinces in the central area of China.
C-c. Qiongzheo tumidinodae—Fagion all. nov. (associations 15–17) in Hunan
Province and in the northern part of Yunnan, South-Western China.
C-d. Indocalamo latifolii—Fagion hayatae var. zhejiangensis all. nov.
(associations 18–19) in the Zhejiang Province, Eastern China.
C-a. Abelio englerianae—Fagion all. nov. (Run. no. 28–30)
Character species: Rhododendron bricranthum, Prunus pilosiuscula, Euony-
mus giraldii, Calanthe fimbriata, Abelia engleriana, Quercus glandulifera, Acer
laxiflorum, Holboellia fargesii, Epimedium sagittatum, Lonicera pseudopro-
terantha, Ainsliaea triflora, Berberis dielsiana.
Located at about 32–33 N and 106–107 E, Nanjiang is situated in northern
Sichuan Province, and to the south of the mountains of Shaanxi Province. This is
a relatively restricted territory at the north-western edge of China. In this area,
forests dominated by Fagus engleriana, Quercus glandulifera and Q. spinosa grow
in a mosaic pattern on the lower slopes of the mountains and along the rivers.
Forests dominated by Fagus lucida grow mainly on the upper slopes, and Fagus
hayatae ssp. pashanica occurs on the steep hills and mountain ridges. It is a
characteristic of this area that these three species of beech inhabit different
environments.
This area is situated in a relatively warm part of the Sichuan basin, but with
relatively low rainfall (about 1,000 mm yearly). Deciduous species are the principal
component of the tree layer (30 %) and in the shrub layer (20 %), whereas evergreen
species occur with a lower presence (15 % and about 8 % respectively, cfr.
Table 3.9); in the herb layer, hemicryptophytes and geophytes are prevailing.
This alliance contains the following two associations.
7. Euonymo porphyrei-Fagetum englerianae ass. nov. (Run. no. 28–29)
Character species: Euonymus pourphyreus, Quercus spinosa, Acer ginnola,
Rubus pungens
Type relevé: Relevé reference, page 153–158: number SHI 9, Nanjan in Sichuan
(Elevation 1,640 m).
This association (Figs. 2.23, 2.24, and 2.25) was observed in Nanjan in Sichuan
Province between 31 500 and 32 450 N, and 106 270 and 107 100 E. The associa-
tion, when growing around rivers, is often made up of Fagus engleriana. Under the
particular ecological conditions of this community, Fagus engleriana grows, sprout
from the root with multi-trunked individuals, with in general not over 40 cm in
2.3 II. Litseo elongatae-Fagetea sp. div. cl. Nov. (Run. No. 28–48) 29
each layer: the shrub layer is dominated by Symplocos botryanta and the herb layer
is dominated by Elatostema stewardii. As the elevation increases, Fagus
longipetiolata in the canopy layer of this association is progressively replaced by
Fagus lucida. The floristic composition of the association also changes, and the
association is often occurring in aspects of transition to the Sinarundinario
chungii—Fagetum lucidae (Wang et al. 2005). On the other hand, deciduous trees
disappear at lower elevations, and evergreen flora such as Castanopsis platyacantha
and Lithocarpus cleistocarpus var. omeiensis, expand as dominant species, with the
transition to an evergreen broad-leaved forest.
11. Sinarundinario bashersuto-Fagetum lucidae (Wang et al. 2005) (Run. no.
35–36)
Character species: Eurya loquaiana, Clethra fabri, Cinnamomum bodinieri,
Callicarpa brevipes, Ardisia affinis, Erythroxylum kunthianum, Fordiophyton
faberi, Indosasa shibataeoides, Dioscorea batatas, Sinarundinaria bashersuta,
Rubus alceaefolius, Viburnum foetidum var. rectangulatum, Rhaphiolepis indica,
Quercus bambusaefolia, Manglietia fordiana, Rhododendron haofui, Castanopsis
eryei, Disporum cantoniense, Oreocharis benthamii var. reticulata, Ternstroemia
kwangtungensis, Phellodendron chinense.
Type relevé: Wang et al. (2005), Table 2, Relevé reference number 7, Nanshan
Nature Reserve, Hunan Province (Elevation 1,760 m)
This association was recorded from the data collected by Wang et al. (2005) in
Nanshan (26 70 N, 110 80 E) in Hunan Province, located on the border of Guangxi
Province. The authors designated the following species as character species:
Sinarundinaria bashersuta, Clethra faberi, Erythroxylum kunthianum, Indosasa
shibataeoides, Manglietia fordiana, Dioscorea batatas, Fordiophyton faberi,
Rubus alceaefolius and Callicarpa brevipes. In our study, we classified all of
these species as character species of the association except Actinidia kolomikta.
In addition, we added a wide variety of species as character species, such as Eurya
loquaiana and Cinnamomum bodinieri. In Nanshan, according to Wang et al.
(2005), forests of Fagus lucida develop on the summit of the mountains and the
upper parts of the northeastern slopes at elevations of around of 1,690–1,830 m. The
association includes two subassociations: symplocetosum lancifoliae (Run. no. 35),
and torricellietosum tiliifoliae (Run. no. 36).
In this association, also according to Wang et al. (2005), Fagus lucida dominates
the canopy layer. Mixed with deciduous species such as Liquidambar formosana,
Acer sinense and Sorbus folgneri and evergreen trees such as Quercus multinervis,
Q. bambusaefolia, Castanopsis eyrei, Manglietia fordiana and Rhododendron
indica, it forms a mixed evergreen-deciduous forest. Evergreen trees prevail in
the sub-canopy layer, and the shrub layer is densely covered by Sinarundinaria
bashersuta and Indosasa shibataeoides. Many F. lucida saplings are also observed.
In the herb layer, many evergreen ferns such as Athyrium strigillosum and
Allantodia wichurae are present.
12. Fagetum engleriano-lucidae (Wang et al. 2005) (Run. no. 37–38)
Character species: Abelia macrotera, Euonymus alatus, Aster ageratoides,
Carex capilliformis, Carex sendaica, Quercus aliena var. acutidentata, Quercus
36 2 Syntaxonomy of the East Asiatic Fagus Forests
Based on Wang et al. (2005), this association is distributed over a small area at
the northern limit of the Chinese beech forests, forming a small island stretching
from the northeast to the northwest slopes of the Houhe Nature Reserve. Together
with Fagus engleriana, the tree canopy consists mainly of deciduous species
Quercus glandulifera var. brevipetiolata, Q. aliena var. acutidentata and Carpinus
turczaninowii, with occasional conifers Pinus tabulaeformis and Tsuga chinensis.
The sub-canopy layer is not well developed. The development of the shrub layer
varies, but Lindera obtusiloba and Cercis chinensis are conspicuous. The herb layer
has high frequency of Carex capilliformis, Maianthemum bifolium Ophiorrhiza
japonica, Elatostema sessile, Saxifraga stolonifera, Disporopsis pernyi and
Zingiber mioga.
14. Sinarundinario nitidae-Fagetum lucidae (Wang et al. 2005) (Run. no.
40–41)
Character species: Hedera nepalensis.
Type relevé: Wang et al. (2005), Table 2, Relevé reference number 32,
Badagongshan Nature Reserve, Hunan Province (Elevation 1,460 m).
According to Wang et al. (2005), Sinarundinaria nitida, Lithocarpus sp. Rubus
buergeri and Sarcopyramis bodinieri were proposed as the character species for this
association in the northern region of Hunan province and Badagongshan Nature
Reserve on the border of Hunan and Hubei Province (29 460 N, 110 40 E). Its
distribution is centered on slopes of 30–48 at an elevation of 1,430–1,600 m. Of
the species indicated by Wang et al. (2005), in this study we demonstrate that Rubus
buergeri has to be considered as character species for the class, Sinarundinaria
nitida is a character species for the order, and Sarcopyramis bodinieri is indicated
as the differential species for the subassociation. In the present study, we found that
the only character species, Hedera nepalensis, had low constancy. From this, it can
be said that the association demonstrates the properties of a typical association,
showing the characteristics of the Abelio englerianae—Fagion all. nov. alliance.
We consider this association to be essentially a symplocetosum crassifoliae
subassociation, and Wang et al. (2005) divided this association into two
subassociations; typicum and symplocetosum crassifoliae.
Again according to Wang et al. (2005), the dominance of F. lucida in the canopy
layer in this association is high, with a mixture of the deciduous Sorbus folgneri and
Betula insignis, and the evergreen Quercus gracilis and Q. multinervis. The sub-
canopy layer, shows little dominance of the deciduous species, with the evergreen
species Q. multinervis, Symplocos anomala, Eurya muricata, Camellia caudata and
C. pitardii frequently appearing. Sinarundinaria nitida are scattered among the
shrub layer with little dominance, while deciduous species are dominant in the herb
layer.
C-c. Qiongzheo tumidinodae—Fagion all. nov. (Run. no. 42–44)
Character species: Castanopsis platyacantha, Qiongzhea tumidinoda,
Allantodia hirtipes, Camellia grijsii, Ilex intermedia var. fangli, Rubus
chroosepalus, Rhododendron hypoglaucum, Acanthopanax evodiaefolius var.
gracilis, Viburnum willeanum, Stranvaesia amphidoxa.
38 2 Syntaxonomy of the East Asiatic Fagus Forests
not indicated the Latin association name or type relevé. Consequently, Indocalamus
latifolius-Fagus hayatae community can not be considered a validly published
association. As a matter of fact, the two subunits are described for different
localities (Qingliangfeng Nature Reserve and Sihaishan Nature Reserve, respec-
tively). In addition, they differ from one another in more than 20 differential
species. In our opinion, considering the differences among beech associations in
other areas of China, these two subunits should be considered different associations.
As to the nomenclatural problems, we propose to save the original formulation by
Wang and Fujiwara (2003) for one of these new associations (19), because this
corresponds to the subunit, where Indocalamus latifolius occurs with higher fre-
quency. For the other association (18) the name is derived from Carex lanceolata
(which occurs only in this type of beech forest) instead of the widespread Pieris
japonica.
The two associations included in this alliance are described for mountain ranges
not far away from the coast of the Chinese Sea. The climate is relatively fresh
(11–13 C in 1,000 m elevation) and humid, with 1,700–1,800 mm rainfall
(Table 3.5). In contrast with the previous alliances, here the deciduous species are
prevailing (32 % and 23 %) over the evergreen (16 % and 17 %) in the tree layer as
well as in the shrub layer (Table 3.9).
18. Carici lanceolatae—Fagetum hayatae var. zhejiangensis ass. nov. lim.
(Hukusima et al. 2001) (Run. no. 45).
Character species: Carex lanceolata, Schisandra henryi, Viola rossii,
Callicarpa giraldii, Meliosma myriantha var. discolor, Viburnum hengshenicum,
Aster procerus, Picrasma quassioides, Liquidambar acalycina, Eurya hebeclados.
Type relevé: Wang and Fujiwara (2003), Table 2, Reference number 6. We were
not able to indicate the type relevé, because in the literature only a synoptic table
exists.
Wang and Fujiwara (2003) reported data for this plant community from three
study districts on the steep upper slopes, between 970 and 1,040 m, of the
Qingliangfeng Nature Reserve in Zhejiang Province (30 60 N, 118 530 E). The
Pieris japonica subunit of the Indocalamus latifolius-Fagus hayatae community
was recorded by Wang and Fujiwara (2003) in these districts. The results of the
present study revealed that this community possessed the abovementioned charac-
ter species as well as species peculiar to it, allowing it to be differentiated from
other beech forests. However, we cannot indicate a type relevé as the original data
provided in Wang and Fujiwara (2003) consist only of a synoptic table with
frequency results, without the detailed cover values for the single relevés.
Many of the character species in this association are the differential species for
the subassociation in Wang and Fujiwara (2003). They described the canopy layer
to be dominated by the deciduous Fagus hayatae var. zhejiangensis, although
Schima superba is sometimes observed in that layer. Development of the sub-
canopy layer below the canopy is poor, with Pieris japonica dominant in the sub-
canopy and shrub layers. The scrub bamboo Indocalamus latifolius grows in the
shrub layer with low coverage. Many species are present in the herb layer, though
none is clearly dominant. Unlike other beech forests, Fagus hayatae var.
2.3 II. Litseo elongatae-Fagetea sp. div. cl. Nov. (Run. No. 28–48) 41
zhejiangensis saplings and seedlings are often observed, indicating steady regener-
ation of the community.
19. Indocalamo latifolii-Fagetum hayatae var. zhejiangensis (Wang and
Fujiwara 2003 lim. Hukusima et al. 2001) (Run. no. 46)
Character species: Lyonia ovalifolia, Abelia dielsii, Carex chinensis,
Polygonatum sibiricum, Smilax austro-zhejiangensis, Calamagrostis arundinacea
var. ciliata, Photinia paniculata, Lithocarpus harlandii, Prunus serrulata,
Indigofera nigrescens, Bredia amoena, Ilex ficoidea, Ilex triflora, Spiraea
chinensis, Itea chinensis var. oblonga, Rubus corchorifolius, Quercus fabri,
Diplazium pinfaense.
Type relevé: Wang and Fujiwara (2003), Table 2, Reference number 7. Like in
the previous association, we are not able to indicate the type relevé, because in the
original description only a synoptic table is given.
This association was studied by Wang and Fujiwara (2003) in the Sihaisan
nature reserve in Zhejiang Province (28 300 N, 120 430 E). They made four relevés
at elevations between 900 and 960 m, and the Lyonia ovalifolia subunit of the
Indocalamus latifolius-Fagus hayatae community was recorded. In the present
study, we took the differential species of the lower units presented in Wang and
Fujiwara (2003) as the abovementioned character species. Basing on the species
composition, this unit appear quite independent from one another and consequently
we concluded that the association level is more adequate for this plant communities
so that we propose this one as a distinct association. However, we cannot indicate a
type relevé as only frequency results and not the original data are provided in Wang
and Fujiwara (2003).
A distinctive characteristic of this association is the large number of species,
such as Fagus hayatae var. zhejiangensis and Toxicodendron trichocarpa, in the
canopy layer, and dominated by deciduous species. Evergreen species are dominant
in the shrub and herb layers. Indocalamus latifolius is dominant on the shrub layer.
Unlike other plant communities, Fagus hayatae var. zhejiangensis saplings and
seedlings are plentiful, indicating healthy regeneration of the community.
Within the wide-ranging beech forests of East Asia, differences in flora can be seen
to reflect differences not only in the capacity of compose plant communities,
depending from the climatic and ecological conditions, as it was described in the
syntaxonomical section of this study, but also in the evolutionary history of each
plant group and the migrations following the last glacial maximum in each area. For
these reasons there are very few species that are uniformly diffused across the whole
range of the study area. However, on the other hand, in many cases, even though
species were different, similar structural properties and functional adaptations can be
observed at the genus or family levels.
Consequently a discussion of similarities and differences among plant communities
only based on the presence or absence of character species is not sufficient. In the
following sections, we will try to develop a further analysis at the species, genus and
family levels, as well as on the periodicity of life processes and relationships between
plant and climate.
First of all, a general outlook on the flora of the East Asiatic Fagus forests. The
species listed in the synoptic table (Tables 3.1a and 3.1b) are 1,535, belonging to
the three groups of the Vascular Plants:
Pteridophyta 145
Gymnospermae 20
Angiospermae 1,370
Running number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
Class I
Order A B
Alliance A-a B-a B-b
Association 1 2 3 4 5 6
No. of releves 20 11 9 5 9 67 18 45 15 10 18 14 9 9 20 16 6 10 13 21 7 23 56 13 9 15 11
Char. sp. of Fagetea crenatae Miyawaki, Ohba et Murase 1964
Hydrangea petiolaris V V V II V II II IV IV V II III I + II I V III III III II IV V III II
Schizophragma hydrangeoides III II III IV III III III IV III II IV IV IV III IV III II II IV IV IV III III II III III IV
Sorbus commixta V V V V V III I II IV III III II II II I I I II + I + II
Disporum smilacinum I I I III I II I I II II II III III II + IV II II + II I II
Kalopanax pictus I I II II I II II I II II II II + II + I + I III II I II
Viburnum wrightii I I IV I I I II II III III V III II III IV III I III II V III II III
Trillium smallii I + II II II II + + I II + II
Tripterospermum japonicum I I III II II II II III I II I II II II + I + + + +
Smilax nipponica V V V V III + I II II I + + I I + +
Styrax obassia III I V IV V III + + I I II I II I II
Mitchella undulata I II II I II II III I III II I II IV II I
Maianthemum dilatatum IV II V V IV II II II IV III + I + + I
Char. sp. of Fagion multinervis Kim, Kimura,Yim 1986 and Fagetalia multinervis Kim, Kimura etYim 1986
3
Acer okamotoana V V V V V
Acer takesimense III V V V III
Prunus takesimensis III IV IV III IV
Asperula odorata V II V I IV + III +
Arisaema amurense IV IV III I III
Polystichum retroso-paleaceum IV II IV III I I + I II +
Synthetic Remarks
3.1
Running number 1 2 3 4 5 6 7 8 9
10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
Class I
Order A B
Alliance A-a B-a B-b
Association 1 2 3 4 5 6
No. of releves 20 11 9 5 9 67 18 45 15 10 18 14 9 9 20 16 6 10 13 21 7 23 56 13 9 15 11
Rhododendron brachycarpum I I V
var. roseum
Celtis jessoensis I IV
Hedera rhombea I IV II
Ophiopogon japonicus III +
Camellia japonica II I
Char. sp. of Saso-Fagetalia Suz.-Tok. 1966
Fagus crenata V V V V V V V V V II V V V III V V V V V V V V
Acanthopanax sciadophylloides IV III IV IV III IV III IV IV III III I IV III II II II I II IV
Magnolia obovata II III III II III II III II I I II II II II I I + I + + + III
Fraxinus lanuginosa III IV IV III V II III II II II IV II III V IV III V II IV V III II
Quercus mongolica var. grosseserrata II IV I I III III II IV IV IV III II III + II I II III + III III
Skimmia japonica var. intermedia III II III III I III II III II III + V II III II + II II II
f. repens
Acer rufinerve II II II I IV III III III II II
III II IV V II I III I III III + I
3
Dryopteris sabaei I II II I I II I I II I I I II II + + I
Carex dolichostachya var. glaberrima II II III IV II II III II I II + I I I + + I I
Prunus grayana III III III III II II I I III II I II II + +
Corylus sieboldiana I II II II III II IV III I III I + + I + I I
Sasa senanensis I + II II II + + II I
Betula grossa + I + I + II II II + + II II V
Symplocos coreana III V V II V + I IV IV III IV V V IV II
Acer shirasawanum II III + + II II I III + III
Carpinus japonica + + I + II I II II + II + II II III III
Stewartia pseudo-camellia I II II I III II II I + + I
Char. sp. of Fagion crenatae Suz.-Tok. 1952
Cephalotaxus harringtonia var. nana II III III I II III V III III I
Plagiogyria matsumureana III II III V IV IV III II III
The Flora of the East Asiatic Fagus Forests
(continued)
Table 3.1a (continued)
54
(continued)
Table 3.1a (continued)
56
Running number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
Class I
Order A B
Alliance A-a B-a B-b
Association 1 2 3 4 5 6
No. of releves 20 11 9 5 9 67 18 45 15 10 18 14 9 9 20 16 6 10 13 21 7 23 56 13 9 15 11
Prunus incisa var. kinkiensis II
Pertya rigidula II
Rhododendron reticulatum + III I
var. ciliatum
Sasa veitchii var. tyugokuensis II
Char. sp. of Sasamorpha-Fagion crenatae Miyawaki, Ohba et Murase 1964
Sasamorpha borealis III IV III IV III IV IV III III IV
Abies homolepis IV II I II III IV
Acer palmatum var. amoenum I I I III I II III V II + II
Rhododendron quinquefolium I + IV I I + II
Abelia spathulata + + + III II IV IV II + IV II
Acer tenuifolium + I V + I + III III II
Stewartia monadelpha IV IV + III V II II IV
Carex fernaldiana II IV II II I II I
3
Dryopteris bissetiana I I II
Ainsliaea apiculata + II +
Torreya nucifera II + I
Pertya scandens II
Dioscorea quinqueloba + II
Prunus verecunda I + + II
Enkianthus subsessilis IV
Pertya triloba III
Rhamnus costata + IV +
Alangium platanifolium var. trilobum I I + + IV
Caulophyllum robustum II III I
Smilax sieboldii I I II + III
Pternopetalum tanakae III +
The Flora of the East Asiatic Fagus Forests
(continued)
Table 3.1a (continued)
58
Running number 1 2 3 4 5 6 7 8 9
10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
Class I
Order A B
Alliance A-a B-a B-b
Association 1 2 3 4 5 6
No. of releves 20 11 9 5 9 67 18 45 15 10 18 14 9 9 20 16 6 10 13 21 7 23 56 13 9 15 11
Enkianthus campanulatus + + + I II III +
Diff. sp. of subassociations
Elatostema umbellatum IV I +
Acer diabolicum + I IV
Sasa hayatae III
Galium kikumugura III
Chamaele decumbens III
Viola eizanensis I III + +
Cirsium microspicatum + II
Clinopodium gracile var. multicaule I II +
Stellaria sessiliflora II I
Viburnum sieboldii II
Lamium humile II
Arisaema angustatum II
3
Cimicifuga acerina II IV II
Cacalia hastata var. farfaraefolia II + III
Cacalia yatabei II I + + III
Calamagrostis hakonensis + II + III
Aconitum japonicum var. montanum + I III
Anemonopsis macrophylla II
Synthetic Remarks
3.1
Potentilla yokusaiana + II
Filipendula multijuga II
Cirsium effusum II
Miricacalia makineana II
Aster ageratoides ssp. amplexifolius + II
Stewartia serrata + II I
Synurus pungens II
Char. sp. of Sapio japonici-Fagetum crenatae Sasaki 1970
Parabenzoin trilobum + + II IV V V II IV
Enkianthus cernuus f. rubens + II + I + II
Styrax shiraiana + + II I I III
Lindera sericea var. glabrata I + V II II V + III
Hydrangea luteo-venosa I II II III
The Flora of the East Asiatic Fagus Forests
(continued)
Table 3.1a (continued)
60
Running number 1 2 3 4 5 6 7 8 9
10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
Class I
Order A B
Alliance A-a B-a B-b
Association 1 2 3 4 5 6
No. of releves 20 11 9 5 9 67 18 45 15 10 18 14 9 9 20 16 6 10 13 21 7 23 56 13 9 15 11
Ligularia stenocephala II
Carex grallatoria + + II
Cacalia yatabei var. occidentalis IV
Acanthopanax hypoleucus IV
Cacalia tebakoensis II + II + IV
Arisaema ternatipartitum III +
Galium kinuta + III
Deparia albosquamata I + III
Cirsium nipponicum var. shikokianum II
Carex sachalinensis var. sikokiana II
Euphorbia sieboldiana II
Cirsium buergeri II
Deinanthe bifida II
Rubia chinensis var. glabrescens II II
3
Philadelphus satsumi + II
Carex filipes II
Elatostema umbellatum var. majus + + II
Paeonia japonica I + + II
Leucosceptrum japonicum + II II
Actaea asiatica + I + I II
Synthetic Remarks
3.1
Sasa ishizuchiana II
Hydrangea sikokiana III
Elements of Quercetalia serratae-grosseserratae Miyawaki et al. 1971
Euonymus oxyphyllus II + I II + III I I II I II II III II II II II
III II V II
II
Struthiopteris nipponica III IV II II I IV III IV V V I II + I + III I I II
IV
Callicarpa japonica V I + I + + + I II + II II IV + I + I II
Euonymus alatus f. ciliato-dentatus I I I III I II II III I III II II II + + I +
Rhus trichocarpa IV III II III I III II IV II I II IV + II + II
Schisandra repanda I + II II + I + II I V I I + I + III
Pourthiaea villosa var.laevis + I + II I II + II III III III II I IV III II
Hydrangea hirta + I II III III IV I III II III II II I I
Rhododendron kaempferi + III + + III II I V III II I I II
Hamamelis japonica I + + + II I I III + + II I
The Flora of the East Asiatic Fagus Forests
(continued)
Table 3.1a (continued)
62
Running number 1 2 3 4 5 6 7 8 9
10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
Class I
Order A B
Alliance A-a B-a B-b
Association 1 2 3 4 5 6
No. of releves 20 11 9 5 9 67 18 45 15 10 18 14 9 9 20 16 6 10 13 21 7 23 56 13 9 15 11
Fagus japonica V III II IV V V
Illicium religiosum III I I II + III
Quercus serrata I II II I +
Evodiopanax innovans II III + IV
Buckleya lanceolata + II II I
Elements of Fraxino-Ulmetalia Suz.-Tok. 1966
Ligustrum tschonoskii I + I IV II + + + + V III V III III IV +
Panax japonicus I + II + II I + + IV + II I + II I
Peracarpa carnosa var. circaeoides + III II IV + III I + I I III + III + +
Hydrangea macrophylla I III I + + + I II III II V V + +
var. acuminata
Leptogramma mollissima I I II II + II II + II I + + II
Euonymus melananthus I + IV + + + + IV I I II II
Polystichum tripteron III I II I I III II + I +
3
(continued)
Table 3.1a (continued)
64
Running number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
Class I
Order A B
Alliance A-a B-a B-b
Association 1 2 3 4 5 6
No. of releves 20 11 9 5 9 67 18 45 15 10 18 14 9 9 20 16 6 10 13 21 7 23 56 13 9 15 11
Lindera umbellata IV V V I V II IV I V III II III II + II
Carpinus tschonoskii + I + II + III + II II II I II II II
Athyrium wardii + I II II III + II + I I III II +
Vaccinium japonicum III III I II III II III IV III + I +
Carpinus cordata I + + I I + V I I II II II
Helwingia japonica + + I + + I V I II I III +
Styrax japonica + I + II II + II II II II II
Akebia trifoliata + + + II I III I III + +
Pieris japonica I + IV + + I III II II II
Oxalis griffithii + II II I IV II I + +
Lyonia ovalifolia var. elliptica + I I II II III II + +
Sapium japonicum I III III + + II I I II
Viburnum erosum I I III IV + I I I II
Carex siderosticta + I + II I I +
3
Asarum sieboldii + + I I I I
Rubus palmatus + + I I IV I
Celastrus orbiculatus I I + + I I
Meliosma myriantha II + II II + III
Ilex pedunculosa I II III I I II
Viola grypoceras I + + + II
Synthetic Remarks
3.1
Pertya glabrescens + + I II II
Viburnum plicatum var.tomentosum + II + + I
Acer palmatum II I + + +
Rubus pectinellus IV II + +
Lindera obtusiloba + I III +
Viburnum urceolatum + II I +
Actinidia kolomikta + II II
Oxalis acetosella + II I
Eurya japonica II II II
Lindera erythrocarpa + I +
Lespedeza buergeri + +
Athyrium otophorum I +
Dryopteris erythrosora I +
The Flora of the East Asiatic Fagus Forests
Lepisorus thunbergianus II I
Schisandra chinensis II
Pachysandra terminalis II
Cayratia japonica II
Symplocos paniculata I
Companions
Pyrola japonica III III V II I + II + IV I
Athyrium yokoscense I II II I I + IV + II II + + II II IV I
Polygonatum falcatum II + + + + I II + II
Actinidia arguta II + + I IV + + + + + I
Lepisorus ussuriensis var. distans I + + I II II +
Phryma leptostachya v .asiatica V I IV I III II + II +
Osmunda japonica I I III I + I I
Desmodium oxyphyllum III I IV I IV + I +
65
(continued)
Table 3.1a (continued)
66
Running number 1 2 3 4 5 6 7 8 9
10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
Class I
Order A B
Alliance A-a B-a B-b
Association 1 2 3 4 5 6
No. of releves 20 11 9 5 9 67 18 45 15 10 18 14 9 9 20 16 6 10 13 21 7 23 56 13 9 15 11
Trillium tschonoskii II II II + I
Fraxinus rhynchophylla II I
Goodyera maximowicziana I II
Dryopteris expansa I III III II II + II I
Solidago virga-aurea var. asiatica + II I III + I + + I + + +
Polygonatum lasianthum I + + + + I I + II + I
Fagus hayatae var. hayatae I III I II + III II II
Carex reinii I III I I I I II II III
Shortia uniflora I I + II + III
Goodyera foliosa + II I + + I + +
Lycopodium serratum var. serratum II + I + I + III I
Tricyrtis affinis + I + I + III + I
Carex conica + I I I + II I +
Sambucus sieboldiana + I II II + II I II
3
Vaccinium hirtum II + I I + II + +
Monotropastrum globosum + I + + + + + II
Stephanandra incisa + II I I II II + I
Phegopteris connectilis + II I
Euonymus macropterus I + II
Carex morrowii var. temnolepis I + + + I
Synthetic Remarks
3.1
Vaccinium smallii I + II
Lycopodium serratum II I I II + I + +
Ilex sugerokii var. brevipedunculata + + I + +
Carex sachalinensis + + II
Osmunda cinnamomea var. fokiensis + + I II +
Vaccinium oldhamii I I II
Cacalia adenostyloides + II I
Astilbe thunbergii var. congesta I I III +
Rubus palmatus var. coptophyllus II I IV II III II
Cacalia delphiniifolia I + + + II II I
Smilax china II II II + + I I
Cephalotaxus harringtonia III II + I + II
Viburnum urceolatum II + I + +
The Flora of the East Asiatic Fagus Forests
var. procumbens
Plectranthus longitubus II + II II
Carex morrowii II II + + + V + II
Ilex geniculata I II I I I I
Sasa nipponica II + + + I I
Acer mono f. dissectum II + II + I + I
Lepisorus onoei III I II +
Rhododendron wadanum II III IV II I III
Deutzia crenata I II I II +
Viola bissetii I + II II
Astilbe thunbergii + I + + II +
Alnus firma II II
Salvia nipponica I II
Tricyrtis macropoda II II
67
(continued)
Table 3.1a (continued)
68
Running number 1 2 3 4 5 6 7 8 9
10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
Class I
Order A B
Alliance A-a B-a B-b
Association 1 2 3 4 5 6
No. of releves 20 11 9 5 9 67 18 45 15 10 18 14 9 9 20 16 6 10 13 21 7 23 56 13 9 15 11
Zanthoxylum piperitum II + I
Lilium cordatum I I II
Viola boissieuana + I II
Omphalodes japonica +
Symplocos chinensis var. leucocarpa + + + + + + + +
f. pilosa
Cacalia nikomontana + + + I +
Menziesia multiflora I I + I +
Tripterygium regelii + + + + +
Acer argutum I + I + +
Athyrium niponicum + + + I +
Vitis coignetiae I + I I
Wisteria floribunda I I I I
Leucothoe grayana var. oblongifolia I I + +
3
Euonymus planipes + + + +
Brachypodium sylvaticum + I I I
Deparia pycnosorum I + I I
Acer nipponicum + I + +
Callicarpa mollis I I + I
Vaccinium smallii var. glabrum + + + I
Lonicera gracilipes I + I +
Synthetic Remarks
3.1
Berberis thunbergii + II + +
Oreorchis patens + + +
Ostrya japonica + I +
Gentiana zollingeri I I +
Geum japonicum I + +
Zelkova serrata I I +
Actinidia polygama I + I
Circaea alpina I + +
Stellaria diversiflora I + I
Polygonatum odoratum + + +
var. pluriflorum
Epigaea asiatica I + +
Polygonum filiforme I + I
The Flora of the East Asiatic Fagus Forests
Hydrangea involucrata + I I
Oplismenus undulatifolius I + I
var. japonicus
Chamaecyparis obtusa + + I
Shortia soldanelloides var. magnus + I I
Staphylea bumalda + I I
Astilbe thunbergii var. fujisanensis + I +
Acer mono var. connivens + + +
Davallia mariesii II + +
Rhamnus japonica var. decipiens + I I
Acanthopanax trichodon + + I
Prenanthes acerifolia II + +
Polygonum cuspidatum + II I
Sanicula chinensis + + +
69
(continued)
Table 3.1a (continued)
70
Running number 1 2 3 4 5 6 7 8 9
10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
Class I
Order A B
Alliance A-a B-a B-b
Association 1 2 3 4 5 6
No. of releves 20 11 9 5 9 67 18 45 15 10 18 14 9 9 20 16 6 10 13 21 7 23 56 13 9 15 11
Patrinia triloba var. palmata + + +
Rhododendron metternichii I + +
Valeriana degeletiana I I I
Abelia serrata + II
Chimaphila japonica + I
Aristolochia kaempferi + I
Viscum aibum var. coloratum I +
Luzula plumosa var. macrocarpa I I
Morus bombycis + +
Pinus parviflora + I
Symplocos myrtacea I II
Stachyurus praecox + +
Trachelospermum asiaticum I II
var. intermedium
3
Quercus salicina + +
Acer mono var. savatieri I I
Asarum savatieri + I
Dichocarpum stoloniferum I I
Dioscorea septemloba + +
Dryopteris monticola + +
Arisaema japonicum I I
Synthetic Remarks
3.1
Tylophora aristolochioides + I
Sorbus gracilis I I
Shortia soldanelloides + +
Galium pogonanthum + +
Weigela hortensis + +
Aster scaber + I
Prunus apetala I +
Abelia spathulata var.stenophylla I +
Eupatorium chinense + +
var. simplicifolium
Hosta montana + I
Sasa tsuboiana + +
Magnolia kobus I II
The Flora of the East Asiatic Fagus Forests
Calanthe tricarinata + +
Cacalia farfaraefolia var. bulbifera I +
Polygonatum macranthum + +
Thujopsis dolabrata var.hondae + I
Ligustrum obtusifolium I +
Carex stenostachys var. cuneata I I
Viola violacea I +
Magnolia kobus var. borealis + +
Acanthopanax spinosus + +
Diphylleia grayi I +
Dioscorea tokoro + +
Carex blepharicarpa + +
Rhododendron reticulatum + I
Boehmeria spicata + +
71
(continued)
Table 3.1a (continued)
72
Running number 1 2 3 4 5 6 7 8 9
10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
Class I
Order A B
Alliance A-a B-a B-b
Association 1 2 3 4 5 6
No. of releves 20 11 9 5 9 67 18 45 15 10 18 14 9 9 20 16 6 10 13 21 7 23 56 13 9 15 11
Picea polita + +
Fraxinus apertisquamifera I I
Epipactis papillosa + I
Anemone pesudo-altaica + I
Streptopus amplexifolius + +
var. papillatus
Asarum heterotropoides + +
Cynanchum sublanceolatum I +
Liparis krameri I +
Tsuga diversifolia I I
Pourthiaea villosa I II
Carpesium triste + +
Abelia spathulata var. sanguinea I I
Scrophularia duplicato-serrata I I
3
Hosiea japonica I I
Galium trachyspermum + +
Lindera sericea I II
Veronica cana var. miqueliana + II
Spuriopimpinella nikoensis + +
Smilacina hondoensis + I
Vitis amurensis f. glabrescens I I
Synthetic Remarks
Athyrium henryi I I
3.1
Poa takeshimana II I
Deparia japonica I +
Carex sp.2 I
Carex sp.1 I
Cornus macrophylla +
Viola acuminata +
Dioscorea nipponica +
Goodyera schlechtendalina I
Smilax riparia var. ussuriensis I
Rhododendron dilatatum +
Neolitsea sericea +
Enkianthus campanulatus +
var. ikokianus
The Flora of the East Asiatic Fagus Forests
Scutellaria laeteviolacea I
Polystichum polyblepharum +
Plectranthus shikokianus I
var. intermedius
Anemone nikoensis I
Astilbe thunbergii var. shikokiana I
Orixa japonica +
Plectranthus kameba I
Lycopodium obscurum I
Phellodendron amurense I
Athyrium clivicola +
Ainsliaea acerifolia I
Cardiandra alternifolia I
Celastrus orbiculatus var. papillosus I
73
Dryopteris lacera +
Dumasia truncata +
(continued)
Table 3.1a (continued)
74
Running number 1 2 3 4 5 6 7 8 9
10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
Class I
Order A B
Alliance A-a B-a B-b
Association 1 2 3 4 5 6
No. of releves 20 11 9 5 9 67 18 45 15 10 18 14 9 9 20 16 6 10 13 21 7 23 56 13 9 15 11
Acer mono var. glabrum +
Lycopodium clavatum +
Lilium medeoloides I
Glaucidium palmatum +
Plectranthus inflexus +
Alnus hirsuta var.sibirica +
Syneilesis palmata +
Platanthera florentii +
Tilia maximowicziana +
Rhododendron metternichii +
var. hondoense
Betula maximowicziana +
Dryopteris saxifraga +
Galium kamtschaticum var. +
acutifolium
3
Carex floribunda +
Calanthe discolor +
Euptelea polyandra +
Sasa megalophylla +
Daphne miyabeana +
Carex lasiolepis I
Synthetic Remarks
Viola dissecta I
3.1
Asplenium incisum I
Boehmeria tricuspis I
Circaea mollis I
Isodon excisus I
Saxifraga fortunei var. incisolobata I
Achyranthes japonica I
Cephalanthera falcata I
Chrysosplenium grayanum I
Eutrema japonica I
Elaeagnus macrophylla I
Angelica polymorha I
Polygonum thunbergii II
Lonicera gracilipes var. glandulosa +
The Flora of the East Asiatic Fagus Forests
Berchemia racemosa +
Smilacina yesoensis I
Malus tschonoskii +
Pyrola renifolia I
Alnus hirsuta +
Acer mono var. ambiguum +
Magnolia sieboldii +
Chelonopsis moschata +
Rubus microphyllus +
Liriope platyphylla I
Monotropa hypopitys I
Arisaema peninsulae I
Angelica edulis +
Thujopsis dolabrata +
75
Pterostyrax corymbosa +
(continued)
Table 3.1a (continued)
76
Running number 1 2 3 4 5 6 7 8 9
10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
Class I
Order A B
Alliance A-a B-a B-b
Association 1 2 3 4 5 6
No. of releves 20 11 9 5 9 67 18 45 15 10 18 14 9 9 20 16 6 10 13 21 7 23 56 13 9 15 11
Prunus buergeriana +
Chamaecyparis pisifera +
Acer ukurunduense I
Pinus densiflora +
Betula schmidtii I
Ephippianthus schmidtii +
Aconitum japonicum +
Erythronium japonicum +
Struthiopteris castanea +
Rubus illecebrosus +
Woodsia manchuriensis +
Dryopteris chinensis +
Coptis trifolia +
Carex breviculmis +
3
Elaeagnus montana I
Rhododendron decandrum +
Rhododendron weyrichii +
Epimedium sempervirens I
Dioscorea gracillima +
Lilium auratum +
Solidago virga-aurea var. leiocarpa +
Synthetic Remarks
3.1
Ophiopogon planiscapus +
Cercidiphyllum japonicum +
Sasa ramosa +
Galium paradoxum I
Tilingia holopetala I
Ligularia fischeri I
Arisaema limbatum +
Ulmus japonica +
Betula corylifolia I
Epimedium grandiflorum I
var. thunberganum
Streptopus streptoides I
Galium kamtschaticum +
The Flora of the East Asiatic Fagus Forests
Tricyrtis latifolia +
Pleioblastus chino +
Elaeagnus montana var. ovata +
Lastrea quelpaertensis +
Euonymus sieboldianus +
var. sanguineus
Leucosceptrum sp. I
Carex insaniae +
Pteridophyllum racemosum +
Anemone debilis +
Rhododendron pentaphyllum +
Agrostis scouleri I
Polypodium vulgare I
Cephalanthera erecta I
77
Table 3.1b Synoptic table of the beech forests in East Asia (China and Taiwan)
78
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Char.sp of Litseo elongatae-Fagetea sp. div. cl. nov.
Fagus longipetiolata II 3 3 I 3 V 4
Litsea elongata I I I 2 V V I II 2 IV 1
Smilax discotis 1 1 I I II
Parathelypteris glanduligera 1 I I 2 I
Ardisia crispa I II 1 IV
Dendropanax chevalieri I I III 2 3 II
Rubus buergeri 1 1 IV II II
Symplocos lancifolia I IV III III
Dennstaedtia scabra I II 1 II
Char.sp. of Sinarundinario nitidae—Fagetalia sp. div.
Fraxinus chinensis 2 II II 3 3 4
Pyrola decorata 1 I III 2 I 1
Stewartia sinensis 2 1 3 II 2
3
Sorbus folgneri 2 2 2 II IV I IV 1 IV
Acer sinense 3 1 I I III 3 1 2 III
Viburnum betulifolium 2 3 3 I III
Polystichum neolobatum 1 I II IV I
Fagus lucida 2 V V V V 4 3 V V
Smilax stans 2 1 1 II IV I 1 III 1 3 II I
Viola schneideri II III III I III II
Ophiopogon bodinieri 1 2 1 I I II 1 II I
Camellia pitardii I II 1 I III III
Quercus oxyodon 3 II II 3 II
Lithocarpus cleistocarpus III 2 V
Symplocos botryantha I 2 IV V
Pinus armandii 1 3 2 II 3
The Flora of the East Asiatic Fagus Forests
Dryopteris labordei 2 V 2 I
Acer oliverianum 1 2 1 IV I III 2 II I 2 V
Fagus engleriana 3 3 1 IV 4 1 2 V
Eurya brevistyla 2 III III 2 V
Lithocarpus hancei II II II 2 III I I 1 V
Symplocos caudata I II I I 1 I I 2 IV
Ophiopogon mairei III III II I
Machilus ichangensis I 3 I 2 V
Symplocos anomala II I 3 IV IV
Reineckia carnea III II 1 II
Plagiogyria stenoptera III 3 III 2 III
Rubus malifolius II II IV 2 I
Hydrangea anomala I II III IV 1 II I III
Quercus stewardiana I III 2 I 3
79
(continued)
Table 3.1b (continued)
80
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Eurya muricata II 3 IV 2 4
Rhododendron mariesii II 1 3 3 3
Polygonatum odoratum I I 3 2 3
Schima superba I I 1 3 4
Rhododendron latoucheae I I 1 1 4
Carpinus viminea I I 1 II 2 3
Lindera reflexa II I I 2 3 4
Symplocos stellaris I I 1 3 3
Quercus gracilis II 2 3 II 3 4
Arthraxon hispidus I 2 2 1 IV 2
Parthenocissus heterophylla II III 2 I 1 IV 1
Dryopteris fuscipes I V III 1 1 I 3
Ophiorrhiza japonica III II 3 II 1
Camellia cuspidata III III I 3 II 3 3
3
Ilex wilsonii 1 IV 1 4
Smilax glabra III I II 1 I 3 1
Rhododendron simsii I III 1 III 2 3 4
Char. sp. of Abelio englerianae—Fagion all. nov.
Rhododendron bricranthum 1 1 2
Prunus pilosiuscula 1 1 2 I
Synthetic Remarks
3.1
Euonymus giraldii 1 3 1
Calanthe fimbriata 2 1 1
Abelia engleriana 1 1 1
Quercus glandulifera 3 2 1
Acer laxiflorum 1 1 1 I
Holboellia fargesii 3 2 II I
Epimedium sagittatum 2 1
Lonicera pseudoproterantha 2 1
Ainsliaea triflora 1 2 III
Berberis dielsiana 1 2
Char.sp. of Euonymo porphyrei-Fagetum englerianae ass. nov.
Euonymus pourphyreus 3 2
Quercus spinosa 2 2 II
The Flora of the East Asiatic Fagus Forests
Acer ginnola 1 2
Rubus pungens 1 2
Diff. sp. of subassociations
Pedicularis nasturtifolia 3
Cacalia roborowskii 2
Meliosma veitchiorum 2
Char.sp. of Vaccinio henryi -Fagetum pashanicae ass. nov.
Fagus hayatae subsp. pashanica 3
Rubus bambusarum 3 I
Hugeria vaccinioides 2
Vaccinium henryi 2
Daphniphyllum angustifolium 2
(continued)
81
Table 3.1b (continued)
82
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Char.sp. of Aceri davidii—Fagion lucidae Wang, Fujiwara et Lei 2005
Quercus multinervis V V V 2 III 3 V
Acer davidii I II II III 1 IV 4 3
Lonicera japonica II I 2 I 2 II
Lithocarpus henryi II II II 2 III I 1
Enkianthus serrulatus II III II 2 2 II
Carex filicina I II 2 I 3 2 IV
Polygonatum cyrtonema I I 1 III 1 I
Ainsliaea henryi II I II III
Acer flabellatum I III I 1
Litsea pungens I II 1 IV 2 II
Pterygocalyx volubilis II IV 1 I
Rubus trianthus II II 2 I
Viola davidii I III 1 I I
3
Mahonia japonica II II 1
Viburnum ichangense I 3 3 III
Char.sp. of Sinarundinario chungii-Fagetum luchidae Wang, Fujiwara et Lei 2005
Sinarundinaria chungii IV IV V
Illicium simonsii I III II II
Rubus pacificus I II I 1
Synthetic Remarks
3.1
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Char.sp. of Polypodio argutum—Fagetum longipetiolatae ass. nov.
Quercus engleriana I I 3 II I
Castanopsis carlesii 2
Cymbidium sinense 2
Polypodium argutum 2
Lophatherum gracile 2
Leptogramma scallani 2
Char.sp. of Sinarundinario bashersuto-Fagetum lucidae Wang, Fujiwara et Lei 2005
Eurya loquaiana II V 3
Clethra fabri V 3
Cinnamomum bodinieri I 2
Callicarpa brevipes III 2
Ardisia affinis III 1
Erythroxylum kunthianum V 2 V
3
Rhaphiolepis indica I 3
Quercus bambusaefolia IV 1
Manglietia fordiana IV 2
Rhododendron haofui II IV 1
Castanopsis eryei I II V 1
Disporum cantoniense 1 II 2
Oreocharis benthamii var. reticulata II 1
Ternstroemia kwangtungensis II 1
Phellodendron chinense II 1
Diff. sp. of subassociations
Lithocarpus glober III
Eupatorium shinense IV
Arthromeris lehmannii IV
The Flora of the East Asiatic Fagus Forests
(continued)
Table 3.1b (continued)
86
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Quercus aliena var. acutidentata II 4
Quercus serrata var. brevipetiolata II 2 I 1
Platycarya strobilacea II 1 3
Adenophora hunanensis I 2
Philadelphus incanus I 2
Diff. sp. of subassociations
Carex subpediformis IV
Castanea henryi IV
Carpinus cordata var. chinensis 1 IV
Hamamelis mollis III 2
Rhododendron augustinii 1 III
Rhododendron hypoglancum 2 III
Luzula plumosa III
Parathelypteris nipponica III I
3
Stellaria palustris II
Polygonum suffultum II
Smilax polycorea II
Viburnum rhytidophyllum II
Pieris formosa 1 II
Cerasus conradinae II
Ligularia veitchiana II
Lindera floribunda II
Goodyera repens II
Carpinus turczaninowii 2 1 4
Deyeuxia sinelatior 4
Saussurea cordifolia 4
Smilax trachypoda 4
The Flora of the East Asiatic Fagus Forests
Rosa setipoda 3
Euonymus praewarskii 3
Buckleya henryi 3
Lespedeza formosa 3
Patrinia scabiosaefolia 3
Forsythia suspensa 3
Styrax hemsleyana 2
Spiraea dasyantha 2
Spiraea prunifolia 2
Rhododendron micranthum 2
Morus mongolica 2
Tsuga chinensis 2
Melampyrum roseum 2
Adenophora cordifolia 2
87
(continued)
Table 3.1b (continued)
88
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Adenophora polymorpha 2
Adenophora trachelioides 2
Pertya cordifolia 2
Char.sp. of Elatostemo sessile—Fagetum lucidae ass. nov.
Phoebe sheareri 1 3
Cercis chinensis 2
Elatostema sessile 2
Saxifraga stolonifera 2
Aesculus wilsonii 2
Bletilla striata 2
Disporopsis pernyi 2
Primula ovalifolia 2
Zingiber mioga 2
Toona ciliata 2
3
Trachelospermum jasminoides 2
Phoebe neurantha 2
Char.sp of Sinarundinario nitidoi-Fagetum lucidae Wang et al. 2005
Hedera nepalensis 1 II
Diff. sp. of subassociation
Dryopteris mariformis II
Litsea suberosa II
Synthetic Remarks
3.1
Aristolochia mollissima II
Asarum ichangense II
Sarcopyramis bodinieri V 1 I
Plagiogyria atenoptera V
Symplocos crassifolia V
Betula insignis IV
Ilex latifrons III
Photinia villosa I III
Acanthopanax fargersii II III
Symplocos stapfiana II
Phanerophlebiopsis blinii II
Magnolia biondii II
Lactuca graciliflora II
The Flora of the East Asiatic Fagus Forests
(continued)
Table 3.1b (continued)
90
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Eurya semisenullata II
Carpinus fangiana II
Symplocos ramosissima II
Allantodia squamigera II
Microlepia marginata II
Ligustrum delavayanum II
Eurya graffi II
Hydrangea xanthoneura II
Arachniodes pseudo-aristata II
Spatholirion longifolium II
Acer prolificum II
Char.sp. of Viburno flavescentis—Fagetum englerianae ass. nov.
Sorbus sargentiana 2
Toxicodendron radicans var. hispidus 2
3
Lepisorus bicolor 2
Neolitsea chuii 2
Dichroa febrifuga 2
Athyrium delavayi 2
Polypodiastrum dielsianum 2
Davidia involucrata var. vilmoriniana 1
Synthetic Remarks
3.1
Dendrobenthamia melanotricha 1
Viburnum flavescens 1
Phyllagathis longipes 1
Smilacina yunnanensis 1
Schizophragma hypoleuca 1
Smilax opaca 1
Char.sp. of Tripterospermo cordifolium – Fagetum englerianae ass. nov.
Schima crenata V
Lindera subcaudata var. hemsleyana V
Neolitsea chinensis V
Panicum brevifolium IV
Manglietia duclouxii IV 1
Tripterospermum cordifolium IV
The Flora of the East Asiatic Fagus Forests
Actinidia vitifolia II
(continued)
Table 3.1b (continued)
92
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Char.sp. of Indocalamo latifolii—Fagion hayatae var. zhejiangensis all. nov.
Fagus hayatae var.zhejiangensis 3 4
Indocalamus latifolius 1 3
Toxicodendron trichocarpa 3 4
Quercus nubium 2 4
Tripterospermum chinense 3 2
Ainsliaea macroclinidioides II 2 4
Eurya ribiginosa var. attenuata 1 4
Magnolia cylindrica 3 2
Rhododendron ovatum 2 2
Dioscorea bulbifera I 3 3
Albizia kalkora 1 3
Liriope graminifolia 1 3
Acer elegantulum 1 2
3
Photinia parvifolia 1 1 2
Smilax nervo-marginata 1 2
Char.sp. of Carici lanceolatae—Fagetum hayatae var. zhejiangensis ass. nov.
Carex lanceolata 3
Schisandra henryi 3
Viola rossii 3
Synthetic Remarks
3.1
Callicarpa giraldii 2
Meliosma myriantha var. discolor 2
Viburnum hengshenicum 2
Aster procerus 2
Picrasma quassioides 2
Liquidamber acalycina 2
Eurya hebeclados I 2
Char.sp. of Indocalamo latifolii-Fagetum hayatae Wang et Fujiwara 2003
Lyonia ovalifolia 4
Abelia dielsii 3 4
Carex chinensis 4
Polygonatum sibiricum 4
Smilax austro-zhejiangensis 4
The Flora of the East Asiatic Fagus Forests
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Char.sp. of Yushania-Fagetum hayatae Suz.-Tok. ex Hukusima et al. 2005
Fagus hayatae V V
Neolitsea acuminatissima V V
Yushania nitakayamensis V V
Stauntonia purpurea V V
Elatostema trilobulatum III II
Crypsinus echinosporus II III
Rubus shinkoensis III V
Schizophragma integrifolium var. formosana III III
Camellia tenuifolia II IV
Dryopteris formosana IV II
Acrophorus stipellatus IV III
Plagiogyria formosana III V
Quercus sessilifolia II IV
3
Illicium tashiroi II V
Smilax elongato-reticullata II III
Osmanthus heterophyllus III V
Pourthiaea villosa var. parvifolia IV I
Viburnum luzonicum V I
Tripterospermum lanceolata III I
Synthetic Remarks
3.1
Coptis quinquefolia I V
Eurya leptophylla I IV
Rhododendron formosanum I III
Smilax elongato-umbelata I III
Loxogramme remote-frondigera II I
Damnacanthus angustifolius I II
Ardisia brevicaulis I II
Trochodendron aralioides I II
Polypodium amoenum II I
Pieris taiwanensis I II
Selaginella remotifolia I II
Viola formosana var. formosana I II
Euonymus spraguei II I
The Flora of the East Asiatic Fagus Forests
Cremastra appendiculata I II
Diff. sp. of subassociations
Eurya crenatifolia IV
Camellia brevistyla III
Araiostegia parripinnata II
Lepisorus obscure-venulosus II
Acer kawakamii II
Arachniodes rhomboides II
Elaeocarpus japonicus IV
Enkianthus perlata IV
Lasianthus japonicus III
Symplocos macrostroma III
Neolitsea aciculata III
Barthea formosana III
95
(continued)
Table 3.1b (continued)
96
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Pellionia arisanensis III
Damnacanthus angustifolius var. stenophyllus III
Helicia cochinchinensis III
Litsea acuminata III
Symplocos cochinchinensis subsp. laurina III
Machilus thunbergii II
Hedera rhombera var. formosana II
Myrica rubra var. acuminata II
Rhododendron pseudo-chrysenthnum II
Crypsinus quasioivaricata II
Lycopodium serratum var. longipetiolatum II
Damnacanthus indica II
Berberis mingetsuensys II
Sarcandra glabra II
3
(continued)
Table 3.1b (continued)
98
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Lindera metcalfiana 3 V
Meliosma sichourensis 3 IV
Beilschmiedia robusta II 2 IV
Osmanthus corymbosus 3 V
Eurya trichocarpa 2 III
Camellia forrestii 2 IV
Lithocarpus dealbatus 2 II
Gomphandra tetrandra 1 III
Illicium yunnanensis 2 III
Sloanea elegans 3 I
Lasianthus longicaudus 2 V
Schefflera producta 1 IV
Euonymus forbesianus 1 III
Parathelypteris hirsutipes 1 V
3
Asplenium normale 1 IV
Ophiopogon clavatus 1 III
Athyrium malipoense 2 III
Sarcandra hainanensis 2 III
Kadsura heteroclita 2 IV
Fissistigma acuminatissimum 2 III
Synthetic Remarks
3.1
Neolitsea levinei 1 2 V
Alpinia chinensis II 2 IV
Pyrrosia lingua 2 III
Piper cft. flaviiflorum 2 III
Vittaria yunnanensis 1 III
Diff. sp. of subassociation
Plagiogyria maxima V
Castanopsis pachyrachis V
Allantodia metteniana I IV
Phyllagathis ovalifolia IV
Beccarinda tonkinensis IV
Diacalpe aspidioides IV
Symplocos groffii IV
The Flora of the East Asiatic Fagus Forests
(continued)
Table 3.1b (continued)
100
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Lithocarpus carolinae 3
Liparis japonica 2
Dendropanax macrocarpus 2
Carex perakensis 2
Canthium simile 2
Calamus oxycarpus 2
Asplenium unilaterale 2
Schima argentea 2
Symplocos adenophylla 2
Common species of the beech forests in East Asia
Cornus controversa 1 2 1 I II II I II
Dryopteris crassirhizoma I
Euonymus fortunei var. radicans II
Disporum sessile II II I 2
3
Leptorumohra miqueliana II
Viola selkirkii I V 4
Tiarella polyphylla I I II I
Matteuccia orientalis 1 1 1 I
Gynostemma pentaphyllum I I I 2 I I
Cymbidium goeringii I 2 II
Synthetic Remarks
3.1
Tsuga sieboldii
Ampelopsis brevipedunculata 1
Viburnum furcatum V III
Rhus ambigua I III
Ardisia japonica 3 II V
Parathelypteris japonica III
Hydrangea paniculata III 2 II 1 2
Sorbus alnifolia II 4 1
Oxalis griffithii 2 II I I 2 II
Acer mono 1 1 I I 1 1
Athyrium wardii 1 1 I 1 II III
Clethra barbinervis 1 4
Ilex macropoda 1
The Flora of the East Asiatic Fagus Forests
Lindera umbellata I I II
Cornus kousa z 2
Vaccinium japonicum II 2 II
Carpinus cordata 3
Asarum sieboldii II 2
Helwingia japonica II
Akebia trifoliata 1
Actinidia kolomikta II 3
Carex siderosticta III 4
Styrax japonica II 1
Carpinus tschonoskii I
Oxalis acetosella II V
Schisandra chinensis 3 2
Ophiopogon japonicus 2 1 II
101
(continued)
Table 3.1b (continued)
102
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Viola grypoceras II
Pachysandra terminalis I
Lespedeza buergeri II 1
Athyrium otophorum I I
Eurya japonica 1
Rubus palmatus I 1
Pertya glabrescens 2
Dryopteris erythrosora I II
Viburnum plicatum var.tomentosum 4
Rubus pectinellus I
Celastrus orbiculatus II III 1
Cayratia japonica I I 3 II
Lindera erythrocarpa III
Acer palmatum 2
3
Pieris japonica 3
Lyonia ovalifolia var. elliptica 2 I 1 II 1
Lindera obtusiloba 2 1 I II 2 3 III
Meliosma myriantha I
Sapium japonicum 3 4
Ilex pedunculosa III 3 I I
Synthetic Remarks
3.1
Companions
Euonymus fortunei 1 I II I 1 II 2 1 II
Enkianthus chinensis 1 2 I I 1 I
Lonicera henryi 2 II I 1 II II
Lindera fragrans I II 2 II 1 I
Elaeagnus lanceolata II I 2 I 2 I
Dryopteris championii II I I 2 1 II
Symplocos adenopus I III III 2 IV 2
Litsea cubeba I 1 1 I 3 1
Symplocos lanrina 2 1 I 2 I
Sorbus xanthoneura 1 2 I I II
Oplismenus undulatifolius I II 1 I I
Smilax cocculoides I I I II 1
Smilax backii I 1 1 II I
103
(continued)
Table 3.1b (continued)
104
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Microtropis fokienensis I II II 1 III
Rhus sp. 1 1 2 I III
Rhododendron sp. 1 1 IV 2 III
Lithocarpus sp1. 1 I 3 II I
Arachniodes chinensis I I 1 I I
Tetrastigma hypoglaucum I I I I I
Helwingia chinensis 1 2 2 I
Eurya alata 1 1 1 II
Mahonia bealei 1 I I II
Ilex chinensis 2 2 II 1
Viola principis II II I 1
Actinidia chinensis II II 1 II
Aucuba chinensis II I I I
Magnolia sprengeri I v II 1
3
Toxicodendron succedaneum I I 1 II
Liriope spicata II II 2 2
Litsea coreana I I 2 3
Nyssa sinensis 1 II 1 2
Eurya nitida II I 1 2
Hydrangea umbellata I I 1 2
Synthetic Remarks
3.1
Kerria japonica II 1 I
Viburnum corymbiflorum II I I
Schisandra sphenanthera 1 II I
Viburnum setigerum II I II
Berberis jalianae I II I
Pellionia radicans I II 1
Rhamnus hemsleyana I II I
Schisandra sinensis II 1 1
Betula luminifera I I II
Hedera nepalensis var. sinensis 2 II I
Heterosmilax japonica I I II
Kalopanax septemlobus I I 2
Hydrangea davidii I II 2
Symplocos botryanta II 2 III
105
(continued)
Table 3.1b (continued)
106
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Athyrium sp. I I II
Pittosporum glabratum var. neriifolium I I I
Lindera communis I I I
Acer sp. 1 1 II
Ilex sp. II 1 I
Chrysosplenium lanuginosum I I I
Eurya sp2. I 1 I
Ilex ciliospinosa I I 1
Sorbus aronioides I I I
Celastrus sp. 2 1 I
Lepisorus contortus I 1 I
Smilax arisanensis I I I
Symplocos chinensis I 1 I
Carpinus fargesii 1 1 1
3
Dioscorea althaeoides I r I
Litsea ichangensis 1 II 1
Magnolia sp. I I 1
Rhododendron stamineum II 2 1
Rubus irenaeus II 1 2
Sorbus sp. 1 I II
Synthetic Remarks
3.1
Stauntonia sp. 1 I I
Trachelospermum cathayanum 1 II 2
Viburnum sp. 1 I I
Chloranthus henryi 1 I 2
Kadsura longepedunculata I I 2
Eurya obtusifolia 1 II
Ilex yunnanensis II I
Machilus rehderi II I
Eupatorium japonicum I II
Galium asperuloides I I
Indocalamus longiauritus II II
Dryopteris sp1. 2 I
Quercus sp. 2 I
The Flora of the East Asiatic Fagus Forests
Rubus sp1. 1 IV
Calamus sp. 2 III
Ophiopogon spp. 3 V
Carex sp.2 2 III
Selaginella sp. 1 II
Calanthe sp. 3 II
Euonymus sp1. 2 IV
Chirita sp. 3 II
Piper sp. 2 II
Lepisorus sp. I II
Camellia sp. IV 2
Carex sp.1 2 IV
Eurya sp1. 1 II
Photinia sp. I III
107
(continued)
Table 3.1b (continued)
108
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Lindera sp. III I
Asarum spp. II III
Chimaphila japonica 1 1
Goodyera schlechtendalina 3 II
Allantodia wichurae I I
Euonymus cornutus I I
Cephalanthera erecta 1 I
Goodyera velutina I I
Carex grandiligulata I I
Cinnamomum wilsonii I I
Lyonia ovalifolia var. ovalifolia 1 1
Lindera cercidifolia 1 II
Lithocarpus confinis I II
Neolitsea aurata I 2
3
Smilax glanco-china 1 1
Tripterospermum cordatum I I
Acanthopanax sp. 1 I
Acer grisema I 1
Actinidia sp. 1 I
Aralia chinensis I I
Synthetic Remarks
3.1
Ardisia sp. I 1
Aruncus sylvester 1 1
Asarum caudigerum I 2
Berchemia kulingensis II 2
Camellia caudata I II
Castanopsis chunii II 1
Celastrus gemmatus 2 I
Clematis finetiana 1 I
Clematis mantana 1 1
Corylopsis sinensis I 2
Quercus augustinii I I
Cyrtomium macrophyllum 1 I
Dipelta floribunda 1 1
The Flora of the East Asiatic Fagus Forests
Elatostema obtusum I I
Enkianthus deflexus 1 1
Ficus sp. 1 I
Ilex fargesii I 1
Ilex szechwanensis 2 II
Koelreuteria paniculata II I
Lasianthus henryi 1 1
Ligustrum sinense 1 I
Parathenocissus himalayana I I
Paris bashanensis 1 1
Castanopsis hystrix 3 1
Prunus brachypoda I II
Polystichum deltodon I 1
Prunus dielsiana 1 I
109
(continued)
Table 3.1b (continued)
110
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Prunus vaniotii I 1
Rhododendron fargesii 1 1
Rubia leiocaulis I I
Schizophragma sp. I I
Stauntonia chinensis I 1
Styrax sp. 1 1
Tilia oliveri I 1
Tricyrtis maculata II 1
Tripterospermum affine I I
Acer amplum II 2
Bretschneidera sinensis 1 II
Cymbidium faberi 1 2
Cyrtomium balansae I 1
Diospyros lotus I 1
3
Elatostema stewardii II 1
Epimedium davidii II 3
Euonymus hederaceus I 1
Euonymus sp2. I 1
Helenia elliptica I I
Helwingia himalaica I II
Synthetic Remarks
3.1
Lonicera similis I 1
Machilus sp2. I 1
Ophiopogon intermedius 1 II
Osmanthus yunanenesis I 1
Padus wilsonii II 2
Peracarpa sp. I I
Pilea martinii I II
Pilea sp. I 1
Rhus chinensis 1 2
Sargentodoxa cuneata I 2
Saxifraga sibirica I 1
Arctous alpinas 1 1
Woodsia polystichoides I 1
The Flora of the East Asiatic Fagus Forests
Alangium chinense 1 2
Toxicodendron vernicifluum 1 1
Quercus stenophylloides I I
Arisaema ssp. I I
Sarcopyramis napalensis I I
Viburnum integrifolium I I
Vaccinium japonicum var. lasiostemon I I
Ligustrum japonicum I I
Tripterospermum taiwanense I I
Viburnum taiwanianum I I
Lonicera acuminata II I
Callicarpa cathayana II
Schizophragma molle II
Tupistra wattii II
111
(continued)
Table 3.1b (continued)
112
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Lithocarpus sp2. IV
Schima sp. 3
Smilax sp. III
Rubus sp2. II
Schisandra sp. II
Dioscorea sp. II
Distylium sp. II
Polystichum sp. III
Embelia sp. III
Osmanthus sp. II
Viola sp1. II
Ribes sp. 2
Machilus sp1. 1
Begonia sp. 1
3
Dryopteris sp2. 1
Viola acuminata II
Dioscorea nipponica I
Oreorchis patens I
Aristolochia kaempferi 1
Ostrya japonica 1
Synthetic Remarks
3.1
Rubus flosculosus 1
Goodyera biflora I
Carpinus polyneura 2
Phymatopsis sp. I
Lonicera ligustrina I
Asteropyrum peltatum I
Hedera nepalansis var. sinensis I
Epipactis sp. I
Toona sinensis I
Ilex fragilis I
Pseudocystopteris atkinsonii I
Rhus delavayi I
Schizophragma integrifolia 1
The Flora of the East Asiatic Fagus Forests
Abelia parvifolia I
Abelia sp. 1
Acanthopanax henryi 1
Acanthopanax simonii I
Acer erianthum I
Acer shensiense 1
Aconitum sinomontanuns 1
Actinidia coriacea I
Adina pilulifera 1
Ainsliaea grossedentata 1
Ainsliaea rubrinervis 1
Ainsliaea yunnanensis I
Ajuga nipponensis 1
Akebia trifoliata var. australis 1
113
(continued)
Table 3.1b (continued)
114
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Allantodia chinensis 1
Alpinia sp. 1
Ampelopsis delavayana I
Arachniodes festina I
Arisaema erubescens 1
Asparagus filicinus 1
Asplenium crinicaule I
Asplenium wrightii I
Asteropyrum cavaleriei I
Aristolochia tubiflora 1
Berberis dasytachya 1
Betula chinensis I
Betula sp. 1
Broussonetia karinoki 1
3
Buxus henryi 1
Cacalia ainsliaeflora I
Cacalia profundorum 1
Cacalia sp. I
Callicarpa japonica var. angustata I
Carex baccans I
Synthetic Remarks
3.1
Carex brunnea I
Carex sutchanensis I
Castanopsis lamontii 1
Castanopsis umensis I
Cayratia oligocarpa 1
Centella asiatica I
Clematis otophora I
Clematis sp. I
Clematoclethra scandens 1
Cleyera incornuta I
Conandron sp. 1
Cornus paucinervis 1
Cornus sp. I
The Flora of the East Asiatic Fagus Forests
(continued)
Table 3.1b (continued)
116
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Desmodium sp. 1
Desmodium szechuenense 1
Deyeuxia arundinacea 1
Dictyocline griffithii 1
Diospyros morrisiana 1
Diplopterygium glaucum 1
Diplopterygium laevissimum 1
Disporum sp. I
Dryopteris subtriangularis 1
Elatostema sp. I
Embelia ribes var.pachyphylla 1
Eragrostis ferruginea I
Eulalia quadrinervis I
Euphorbia hylonoma I
3
Euptelea pleiospermum 1
Eurya distichophylla I
Euscaphis japonica I
Ficus harlandii I
Fragaria orientalis I
Grangea maderaspatana I
Synthetic Remarks
3.1
Hanceola sinensis I
Hedera sp. 1
Hemselya szechuenensis I
Homalium bhamoense 1
Hopea sp. I
Humata tyermanni 1
Idesia sp. I
Illicium sp. I
Impatiens sp. I
Indocalamus fanjinshanensis 1
Indocalamus sp. 1
Liquidamber formosana 1
Litsea ssp. 2
The Flora of the East Asiatic Fagus Forests
Lithocarpus thomsonii I
Lithocarpus variolosus I
Litsea grandiflora I
Litsea populifolia I
Litsea thomsinii I
Lonicera gynochlamydea I
Lonicera sp. I
Lyonia ovalifolia 1
Lysimachia clethroides I
Maackia hwashanensis 2
Machilus faberi 1
Mahonia gracilipes I
Meliosma flexuosa 1
Meliosma paupera I
117
(continued)
Table 3.1b (continued)
118
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Meliosma sp. I
Melothria maysorensis I
Michelia sp. 1
Microsorium hymenodes I
Myrsine sp. I
Millettia championii 1
Monochosorum henryi 1
Monotropa uniflora 1
Morinda umbellata 1
Neolitsea sp. I
Neolitsea zeylanica I
Nothopanax davidii I
Oplismenus compostius I
Panax transitorius I
3
Phoebe faberi I
Phymatopsis teneupes 1
Pieris tomentosa 1
Plagiogyria distinctissima 1
Pleuropus euchloron 1
Pollia sp. 1
Synthetic Remarks
3.1
Polystichum discretum I
Polystichum squarrosum 1
Polystichum tsus-sinense 1
Primula sp. I
Prismatomeris labordei 1
Pterocarya dalavayi I
Pterostyrax rosea I
Pyrola rugosa 1
Quercus asine 1
Rhododendron anthopogonoides 1
Rhododendron rivulare I
Rhododendron seniavinii 1
Schefflera glomerulata 1
The Flora of the East Asiatic Fagus Forests
Scutellaria baicalensis I
Scutellaria franchetiana I
Smilacina glabra I
Smilacina paniculata I
Smilacina sp. I
Smilax ferox 1
Smilax lanceiofolia I
Sorbus caloneura 1
Sorbus hemslaya 1
Sorbus rufopilosa I
Stachyurus himalaicus 1
Stauntonia duclouxii I
Stellaria alsine 1
Stewartia sp. 1
119
(continued)
Table 3.1b (continued)
120
Running number 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51
Class II III
Order C D E
Alliance C-a C-b C-c C-d D-a E-a
Association 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
No. of releves 3 3 3 10 10 7 3 6 4 23 4 3 3 7 5 2 5 3 4 17 8 3 8 4
Styrax rosea I
Symplocos glauca 1
Symplocos lucida I
Symplocos theaefolia I
Symplocos wikstroemiifolia 1
Synurus deltoides I
Ternstroemia luteoflora 1
Tetrastigma obtectum var. glabrum 1
Thladiantha glabra 1
Thalictrum uncatum 1
Tripterospermum sp. I
Tupistra tui I
Vaccinium sp. 1
Veratrum schindleri I
3
Viburnum cordifolium I
Viola cordifolia I
Viola schiensis I
Viola sp2. 1
Vitis sp. I
Zanthoxylum scandens 1
Synthetic Remarks
3.1
Zanthoxylum schinifolium I
Viola arcuta 1
Viola biflora 1
Vitis flexuosa 1
Elaeagnus pungens I
Asarum chinensis 1
Polystichum makinoi 1
Saxifraga flabellifolia 1
Schisandra incarnata 1
Prunus transarlsanensis I
Lysimachia ardisioides I
Arisaema formosanum I
Polystichum parvipinnulum I
The Flora of the East Asiatic Fagus Forests
Quercus morii I
Rubia lanceolata I
Rhododendron rubropilosum I
Athyrium arisaense I
Berberis aristato-serrulata I
Abelia serrata
121
122 3 Synthetic Remarks
It is difficult to manage this material, because this flora is scattered over large
territories (partly islands, partly on the mainland), from Hokkaido (45 N) in the
north of the Japanese Archipelago, to Yunnan Province in South China (23 N), and
a floristic survey, treating the flora of the whole region with unitary criteria does not
exist. As the support for our analysis we used the well known classical Floras of
Japan by Makino T. and by Ohwi J., and for China and Taiwan the recently
produced e-Floras (in some parts still unpublished).
Families with the most elevate number of species are:
Polypodiaceae 126
Liliaceaae 84
Rosaceae 82
Ericaceae 65
Asteraceae 62
Lauraceae 56
Caprifoliaceae 48
Saxifragaceae 46
Aceraceae 45
Theaceae 45
Fagaceae 44
Poaceae 42
Cyperaceae 37
Araliaceae 32
Magnoliaceae 32
Rubiaceae 30
Celastraceae 30
Betulaceae 29
The highest values are reached in the Polypodiaceae and Liliaceae, indeed, both
families were recently split in many smaller units; consequently, the only family
reaching more than 5 % are the Rosaceae, followed by Ericaceae, Asteraceae and
Lauraceae with 4 %. Among families with surprisingly low presence there are
Fabaceae (15 species), Brassicaceae and Boraginaceae (each 1 species).
The elevate presence of Ericaceae is the consequence of the high diversity in
Rhododendron: 40 species or infraspecific taxa; among the genera of woody plants
in the flora of the beech forests, only Acer is reaching a higher total (44 species).
Genera in the synoptic table (Tables 3.1a and 3.1b) are in total 442, which is
barely 14 % of the total registered for the entire flora of China (3,143 genera, see
Wang et al. 2006). The genera including more than 20 species or subspecies are:
Acer 44
Rhododendron 40
Carex 37
Symplocos 28
Viola 25
Viburnum 24
(continued)
3.2 Investigation of Plant Communities at the Level of Genus Composition 123
Quercus 23
Rubus 23
Evonymus 22
Ilex 22
Dryopteris 21
The majority of these genera belongs to the woody components of the flora of the
E-Asiatic beech forests (mainly Japan, China and Taiwan).
From these data it is clear, that the flora of the East-Asiatic beech forests is
highly diversified regarding species number, as well as genera and families. In the
phytosociological literature, there are only few examples of syntaxonomical
elaborations dealing with a similar complex floristic information. The complexity
depends from the large geographical range, but also from the previous selection,
based on the presence of forests, composed by the different species of Fagus
(in total seven species, plus some local varieties) endemic in East Asia.
In this section, similarities at the genus level between plant communities are
discussed. All the species in Tables 3.1a and 3.1b, including Pteridophyta and
Gymnosperms, are distributed among a total of 455 genera (Table 3.1d [Online]).
Considering their appearance or not in units classified by species composition, we
grouped genera displaying similar appearance patterns, and extracted 20 genus
groups. Each individual plant community unit is, therefore, recognizable from the
combination of constituent genus groups. Genera with low frequency were treated
as companions. Therefore, three classes are distinguished in accordance with the
vegetation units shown in Tables 3.1a, 3.1b, and 3.1d [Online]. Within the three
classes in Table 3.1d [Online], differences between class III and the rest of the two
classes are large. At the order level, characteristic genus groups are found at each
order, quite independent from each other. Among them the relatively small Island
Ulleungdo, belonging to Korea (A) is the poorest in distinctive species, followed by
Taiwan (D); in both cases we have to deal with insular floras (in the case of
Ulleungdo a flora with a relatively low species number, because of the isolation
of this very small island, far away from the coasts of the continent and of the
Japanese island of Honshu). Japan and China have many genera in common. Within
each of the lower vegetation units such as alliance, association and subassociation,
much lesser changes in the composition at the genus level exist, and therefore, in
this case it was difficult to point out significant differences among these units.
According to Table 3.1d [Online], the genera constituting groups 1 and 2 have a
quite uniform distribution across the entire range of beech forests in East Asia.
Among these, genera of the evergreen broad-leaved forest class are included in
genus group 1. On the contrary, genus group 2 contains genera, in prevalence
124 3 Synthetic Remarks
Table 3.2 Sum of presences of each family in the three classes of beech forests
Fagetea Litseo-
Code Element Family name crenatae Fagetea ass. 21-22
1 SE-Asiatic Bretschneideraceae 0 2 0
1 SE-Asiatic Cercidiphyllaceae 1 0 0
1 SE-Asiatic Dipterocarpaceae 0 1 0
1 SE-Asiatic Lithocarpaceae 0 30 10
1 SE-Asiatic Nyssaceae 0 6 2
1 SE-Asiatic Stachyuraceae 2 1 0
1 SE-Asiatic Trochodendraceae 1 3 0
4 43 12
2 E-Asiatic—N-temperate Ulmaceae 17 9 3
2 E-Asiatic—N-temperate Urticaceae 11 7 0
2 SE-Asiatic Arctic-alpine Ericaceae 145 94 2
2 SE-Asiatic Circumboreal Caprifoliaceae 128 86 0
2 SE-Asiatic Circumboreal Elaegnaceae 5 7 0
306 203 5
3 E-Asiatic—subcosmop. Oleaceae 61 18 2
3 E-Asiatic-Pantropical Buxaceae 1 2 0
3 E-Asiatic-Pantropical Hamamelidaceae 18 7 0
3 Pantrop—SE-Asiatic Icacinaceae 2 0 2
3 SE-Asiatic—pantropical Zingiberaceae 0 1 4
3 SE-Asiatic—pantropical Arecaceae 0 0 3
3 SE-Asiatic—pantropical Taccaceae 0 2 0
3 SE-Asiatic—pantropical Theaceae 30 86 11
3 Trop-Subtropical (SE-Asiat.) Celastraceae 100 40 10
3 W-Pacific-subcosmop. Balsaminaceae 1 5 0
213 161 32
4 E-Asiatic anfipacific Gesneriaceae 1 2 4
4 SE-Asiatic anfipacific Staphyleaceae 3 1 0
4 SE-Asiatic anfipacific Styracaceae 33 5 1
4 SE-Asiatic anfipacific Symplocaceae 26 64 9
4 SE-Asiatic anfipacific Sabiaceae 13 6 2
4 SE-Asiatic anfipacific Junglandaceae 6 4 0
4 SE-Asiatic anfipacific Lardizabalaceae 10 17 1
4 SE-Asiatic anfipacific Magnoliaceae 62 38 13
4 SE-Asiatic-Mesoamerican Chloranthaceae 6 8 2
4 SE-Asiatic-Mesoamerican Clethraceae 19 5 0
4 SE-Asiatic and S-American Elaeocarpaceae 0 1 5
4 Tropical amphi-pacific Actinidiaceae 17 10 0
4 Tropical amphi-pacific Melastomataceae 0 10 0
4 Tropical amphi-pacific Ochnaceae 0 1 0
196 172 37
5 Oceanic-subcosmopol. Myricaceae 0 1 0
5 Oceanic-trop. and subtrop. Aquifoliaceae 64 42 0
(continued)
3.2 Investigation of Plant Communities at the Level of Genus Composition 125
deciduous, common to the beech forest classes, and this group appears in opposition
to genus group 6, which is comprised in the class III, with genera characteristic of
evergreen broad-leaved forests. Genera such as Fagus, Acer, Prunus and Viola,
which are common to beech forests in Europe and North America (Braun 1974;
Ellenberg 1988) appear in genus group 1 and 2. Among them, members of the genus
Acer are abundant in East Asia, characteristically appearing with high frequency in
every vegetation unit. Other genera frequently appear in the beech forests of East
Asia. Genus group 3 includes a group of scrub bamboos that are characteristic of the
beech forests of East Asia; however, the eight genera in this group differ in terms of
vegetation unit and distribution.
Group 10 contains the genera characteristic of the class Fagetea crenatae which
are in common to the beech forests of Korea and Japan. However, while Trillium and
Lilium are characteristic genera of beech forests in Korea and Japan, the other genera
in the group are not centered in the beech forests and consequently they can not be
considered as particularly indicative for this class. However, in genus group 7,
formed of genera common to beech forests in China, Korea and Japan, eight genera
(Kalopanax, Tilia, Astilbe, Panax, Leptorumohra, Maianthemum, Leptogramma and
Mitchella) clearly appear with high frequency in the class Fagetea crenatae.
Genus group 11 is characteristic of the order Fagetalia multinervis, into which the
beech forests on Ulleungdo Island, Korea, are placed. This genus group is composed
of herbaceous species. In addition, the other distinctive character of this order
Fagetalia multinervis is that it lacks any of the genera found in genus groups 4, 5
and 8. Furthermore, among the genera which characterize the beech forests in
Taiwan, China and Japan, the following evergreen genera are missing from the
beech forests in Korea: Ilex, Daphniphyllum, Pieris, Skimmia, Eurya, Illicium,
Quercus, Symplocos, Neolitsea, Euonymus and Elaeagnus. As a general view, the
flora of these beech forests appears relatively poor, probably as a consequence of the
isolated position in the Sea of Japan and of the small surface of the Ulleungdo Island.
Genus group 12 characterizes the Japanese beech forests classified as order Saso-
Fagetalia crenatae. Although this genus group is characterized by the presence of
many deciduous and herbaceous genera, it also includes coniferous genera such as
Cephalotaxus, Abies, Torreya and Cryptomeria. This order also includes 15 genera
of high frequency from genus group 8, Magnolia, Ilex (D : deciduous), Paris,
Corylus, Cacalia, Smilacina, Hamamelis, Vaccinium, Tricyrtis, Akebia, Diplazium,
Sapium, Aesculus, Pertya, Pterocarya, which are common to beech forests in Japan
and China. This order is divided into two alliances. The two alliances are
characterised by the genus groups 13 and 14, but the presence of these does not
seem very significant. Fagion crenatae species on the Sea of Japan side of Japan are
characterized by the presence of Streptopus and Heloniopsis. In addition,
Arachniodes, Daphniphyllum of the genus group 4, Plagiogyria in the group 5, and
Shortia and Aconitum in the group 12 showed obviously higher frequency in the
alliance Fagion crenatae than in the opponent alliance Sasamorpho-Fagion crenatae.
On the other hand, in the alliance Sasamorpho-Fagion crenatae, distributed along the
Pacific Ocean side of the county and characterized by genus group 14, the evergreen
conifer Chamaecyparis is a very significant presence, although this genus can be
observed only with low frequency. However, scrub bamboo Sasamorpha in the
128 3 Synthetic Remarks
genus group 3, Celastrus and Sapium in the genus group 8, and evergreen of Illicium
(E), Quercus (E) and Symplocos (E) in the genus group 12 appear in this alliance with
high frequencies. Class Litseo elongatae-Fagetea sp. div., observed in China and
Taiwan, can be distinguished from class Fagetea crenatae in Japan and Korea by the
combination of various genera. Especially, genus groups in China showed low
frequency of these genera. As a result, in the wide-ranging and often isolated
beech forests of China, composition is various and a compositional unity is limited
in an area with a lack of any overlapping unity. The characteristic genera of the class
Litseo elongatae-Fagetea sp. div., the unifying class of beech forests of China and
Taiwan are gathered in genus group 15 and 16, which contains evergreen genera such
as Smilax, Lithocarpus, Litsea, Machilus, Stauntonia, Cleyera and Maesa. From this
it is clear that class Litseo elongatae-Fagetea sp. div., which is characteristic of
Chinese and Taiwanese beech forests, can be distinguished by the presence of
evergreen genera. The genera characteristic of the Chinese order Sinarundinario
nitidae-Fagetalia sp. div. are included in genus group 17. This genus group is also
distinguished by the presence of evergreen genera such as Lonicera, Pilea and
Pittosporum. Genus group 18 is characterized by the alliance Abelio englerianae-
Fagion, which is distributed in areas of Sichuan Province. Furthermore, this alliance
does not have the genera found in evergreen-rich genus group 19 and 20.
Two genus groups (19 and 20) contain the genera characterizing other three
alliances present in Chinese beech forests; Aceri davidii-Fagion lucidae all.,
Qiongzheo tumidinodae—Fagion all. and Indocalamo latifolii-Fagion hayatae var.
zhejiangensis all. These three alliances show variations in species composition, but
they appear quite uniform in terms of genus composition, and only few differences
can be observed. Further on, these alliances include 13 associations and 9
subassociations, and regarding the alliance level, there are few characteristic features
regarding the presence of genera in order to distinguish these associations and
subassociations. Genus group 9 characterizes the order Fagetalia hayatae, found in
Taiwan, and includes evergreen genera such as Damnacanthus and Trochodendron.
In addition, it clearly differs from the Chinese orders in that it lacks genera from
genus groups 7, 8 and 17. This order contains the largest number of evergreen genera
than any other orders in East Asian beech forests. Class III, an evergreen broad-
leaved class, is characterized by genus group 6, and one alliance and two associations
are classified. However, because of the lack of sufficient data, a final study still has
not been reached. Nevertheless, this class can be divided into two associations with
variable distributions on the basis of the genera from genus group 6, such as
Gomphandra, Parakmeria, Chirita, Vittaria, Adinandra, Eriobotrya, Fissistigma
and Meliosma, which characterize this class. There are many evergreen genera in
these associations, too.
Observing the genus composition of all genera which occur in the beech forests
of East Asia, clear similarities and differences can be seen between the three classes
and order levels divided on the basis of species composition. At the alliance level,
although the number of genera which characterize each alliance is small, some
genera show obviously higher frequency at a particular alliance. No genus level
characteristics can be observed at the association or subassociation level. This
3.3 Phytogegraphical Analysis at the Family Level 129
FE 1. Cosmopolitan
FE 2. Pantropical
FE 3. Tropical Asian–trop. American
FE 4. Palaeotropical
FE 5. Tropical Asian–Trop. Australian
FE 6. Tropical Asian–Tropical African
FE 7. Tropical Asian
FE 8. Holarctic
FE 9. Eastern Asian–North American
FE 10. Temperate Eurasian
FE 11. Temperate Asian
FE 12. Mediterranean, western to central Asian
FE 13. Central Asian
FE 14. Eastern Asian
While treating the data dealing with the beech forests in order to obtain a clearer
overview of the families, we condensed the elements into eight groups with the
following definitions (in brackets, the corresponding FE in the Chinese literature):
a) Endemic—the species of these families are distributed only in the SE-Asiatic
zone, from China to Vietnam, Malaya and eventually in Indonesia (FE 14).
b) SE-Asiatic—Northern—families having the centre of diversity in the SE-Asiatic
zone, with expansion in the Boreal zone and eventually to the Arctic (FE 9).
c) SE-Asiatic—Southern—families having the centre of diversity in the SE-Asiatic
zone, with expansion in the Subtropical and Tropical zones (FE 7).
d) Anfipacific—families with a bi-zonal distribution range: in Asia (on islands
and coastal areas of the continent) and North America on the western coastal
130 3 Synthetic Remarks
side (among them the family Chloranthaceae, one of the earliest forms of
Angiosperms!); absent at temperate and cold latitudes: in general corresponding
to FE 3.
e) Oceanic—worldwide distribution, but mostly on coastal areas and with large
gaps in areas with continental climate (some of them are probably considered
within FE 3, but with more or less worldwide distribution).
f) Boreal—the biomes of the lowland coniferous forest and tundra (FE 8).
g) Tropical—the cool mountain climate is not favourable for tropical plants,
therefore they are in the beech forests relatively rare and can be concentrated
here, not regarding their continental distribution (FE 2, FE 4, FE 5, FE 6).
h) Cosmopolitan—Families with world-wide distribution, in some cases limited to
the temperate and tropical zones, or with particular presence in warmer climates
(FE 1).
In any case, it has to be pointed out, that our proposal for the definition of
phytogeographical elements is an “ad hoc system”, useful for the particular argu-
ment of beech forests in East Asia, but which cannot be extended, without a careful
revision, to other arguments.
Methods—A complex procedure was necessary to obtain significant results. We
started from Tables 3.1a and 3.1b, with 1,535 rows (species ordered as ecological and
sociological indicators) and 51 columns (associations and subassociations ordered by
their position in the phytosociological system); in Tables 3.1a and 3.1b, for each
species values of frequency are given (from I to V). First, the rows were ordered by
families, in alphabetical order and Pteridophyta and Gymnospermae were excluded,
reducing the table to 1,370 rows. Then, for each family the number of species
occurring in each column (vertical sum) was calculated: this means, as many times
species of every family were present in each column (independently from the fact
whether they were present as I, II, III, IV or V). In this way, the table became reduced
to 114 rows (the families) by 51 columns. Successively, for every row (horizontal
sum) the total of species present in columns 1–27 (class Fagetea crenatae) was
calculated, in columns 28–48 (class Litseo-Fagetea) and in columns 49–51 (the last
two associations with number 21–22); with this transformation Table 3.2 was
obtained, with 114 rows and 3 columns. Finally, families were ordered by phytogeo-
graphical elements and for each element the sum of frequences was calculated: in this
way, Table 3.3 was obtained.
Table 3.3 reaches the maximum level of synthesis, giving a quantitative evalua-
tion for the presence of the different phytogeographical elements in the three
vegetation classes, i.e. reducing the dimensions from 1,370 by 51 to 8 by 3. In
Table 3.4 the raw data (totals) are given on the left side, but a comparison among
the columns remains difficult, because of the big numerical difference among the
three columns. The better outlook on the different incidence of each element is
given on the right side of Table 3.4, where data are normalized as %.
A short commentary to the data in Table 3.3. From a comparison of the normalized
data it is possible to distinguish that the class Fagetea crenatae (ass. 1–6) has a
prevalence in families belonging to the Boreal and Northern elements, but the largest
3.3 Phytogegraphical Analysis at the Family Level 131
Table 3.3 Frequency of the species (totals and percentages) distributed among the phytogeo-
graphical element of their families
Raw data (total) Normalized (%)
Associations (reference no.) 1–6 7–20 21–22 1–6 7–20 21–22
Total of columns in Tables 3.1a and 3.1b 27 21 3 27 21 3
Endemic 4 43 12 0.14 2.4 6.3
SE-Asiatic—Northern 306 203 5 10.9 11.5 2.6
SE-Asiatic—Southern 213 161 32 7.6 9.1 16.8
Anfipacific 196 172 37 7.0 9.8 19.4
Oceanic 76 53 1 2.7 3.0 0.5
Boreal 370 161 9 13.2 9.1 4.7
Tropical 341 348 43 12.2 19.8 22.6
Cosmopolitan 1,271 616 49 45.4 35.0 25.8
Gross total 2,777 1,757 188 99.14 99.7 98.7
component (ca. 50 %) are Cosmopolitan families, i.e. those not giving any phytogeo-
graphical information. The two associations from South Yunnan on the contrary have
elevate frequences of Endemic families, of Southern, Amphipacific and Tropical
families, whereas Cosmopolitan have reduced frequency (ca. 25 %). The class Litseo-
Fagetea occupy an intermediate position between the two extremes.
From the phytogeographical point of view, the most significant component is
represented by the endemic element. In East Asia 6 families can be considered as
endemic (Table 3.5), some of them are monotypic, other with hundreds of species,
mainly belonging to the arboreal flora (Dipterocarps, Lithocarpus). Six families are
a very conspicuous component, even if some of them possibly are to be connected
with other families with broader geographical distribution (e.g. Lithocarpus in
Fagaceae). For a comparison, we can remember that in the whole flora of Europe
(over 10,000 species) not even a single endemic family does exist.
A comparison with the investigation of Angiosperms in the flora of China is
difficult because we are analyzing endemic families, instead of genera. In addition,
Wang et al. (2006) consider as endemics only the genera restricted to China, whereas
we consider as endemic all families with a range extended to the whole SEAsiatic
zone. Indeed, in both cases some similar trends can be observed: the elevate
endemism in the SW provinces (Sichuan and Yunnan), elevate values of boreal
genera in the northern Provinces (Shaanxi, Henan and in particular Heilongjiang),
and a flora rich of tropical elements in the southern provinces (Guangxi, Guizhou
and Yunnan).
The present phytogeographical analysis shows an unexpected condition in the
distribution of Angiosperm taxa in SE-Asia. Previous research (Wang et al. 2006)
interprets the presence of genera of Angiosperms in China as strongly correlated
with the latitude, which for the authors of this study means climate i.e. from the
tropical climate to a cold-temperate one. From the present analysis it appears clear,
that families are not distributed along a geographical (or climatic) gradient, but with
an evident discontinuity, and the concentration of the most peculiar elements can be
Table 3.4 Climatic conditions for each vegetation unit of beech forests in East Asia
132
Mean Snow
Elevation temperature 1,000m 1,500m Precipitation depth
Class Order Alliance Association Location Latitude Longitude (m) ( C) ( C) ( C) (mm) (cm)
I A A-a 1. Hepatico- 1. Ulleungdo in 37 130 510 E 357 11.5 8.0 5.2 1,371
Fagetum Korea 290 N
multinervis
B B-a 2. Saso kurilensis- 2. Mt. Kariba in 42 139 590 E 500 5.8 3.1 0.3 1,755 277
Fagetum crenatae Hokkaido Dist. 350 N
3. Mt. Chokai in 39 140 010 E 727 8.0 6.5 3.7 3,039 253
Tohoku Dist. 080 N
4. Mt. Hakusan in 36 136 400 E 750 9.4 8.0 5.3 3,087 207
Hokuriku Dist. 080 N
3. Lindero 5. Mt. Daisen in 35 133 320 E 1,000 8.1 8.1 5.4 2,705 182
umbellatae- Chugou Dist. 230 N
Fagetum crenatae 6. Mt. Garyu in 34 132 110 E 800 9.3 8.2 5.5 2,301 116
Chugoku Dist. 410 N
B-b 4. Sasamorpho- 7. Nikko in Kanto 36 139 290 E 1,417 5.8 8.1 5.3 2,020 18
Fagetum crenatae Dist. 440 N
8. Chichibu in 36 138 500 E 1,230 7.1 8.4 5.6 1,347 18
Kanto Dist. 000 N
5. Corno-Fagetum 9. Mt. Kintoki in 35 139 000 E 982 9.1 9.0 6.3 2,932 15
crenatae Kanto Dist. 170 N
3
10. Mt. Fuji in 35 138 430 E 1,277 7.8 9.3 6.6 2,821 71
Kanto Dist. 170 N
6. Sapio japonici- 11. Hinodegatake 34 136 060 E 1,318 7.4 9.1 6.4 3,655 6
Fagetum crenatae in Kinki Dist. 110 N
12. Mt. Ishizuchi 33 133 070 E 1,676 5.4 9.1 6.4 2,840 14
in Shikoku Dist. 450 N
13. Mt. Aso in 32 131 060 E 1,200 9.0 10.1 7.4 3,060 16
Synthetic Remarks
II C C-a 7. Euonymo 14. Nanjan in 32 107 100 E 1,371 11.1 13.1 10.4 993 No
porphyrei-Fagetum Sichuan Prov. 450 N data
englerianae
8. Vaccinio
henryi—Fagetum
hayatae subsp.
pashanicae
C-b 9. Sinarundinario 15. Fanjingshan 27 108 420 E 1,758 10.2 14.4 11.6 1,421 No
chungii- Fagetum Natural Reserve 530 N data
lucidae in Guizhow Prov.
16. Kuankuoshui 28 107 080 E 1,464 13.3 15.9 13.1 1,203 No
Nature Reserve 130 N data
in Guizhou Prov.
10. Polypodio 17. Fanjingshan 27 108 440 E 1,185 13.2 14.2 11.5 1,293 No
argutum—Fagetum Natural Reserve 530 N data
longipetiolatae ass. in Guizhow Prov.
Phytogegraphical Analysis at the Family Level
nov.
11. Sinarundinario 18. Nanshan 26 110 060 E 1,648 10.9 14.5 11.7 1,630 No
bashersuto-Fagetum Nature Reserve 080 N data
lucidae in Hunan Prov.
12. Fagetum 19. Longmenhe 31 110 290 E 1,646 9.6 13.2 10.4 1,266 No
engleriano-lucidae Nature Reserve 180 N data
in Hubei Prov.
20. Dalaoling 31 110 560 E 1,534 9.9 12.8 10.1 1,289 No
Nature Reserve 030 N data
in Hubei Prov.
21. Houhe Nature 30 110 310 E 1,696 9.6 13.4 10.7 1,527 No
Reserve in Hubei 040 N data
Prov.
(continued)
133
134
C-d 18. Carici 26. 30º 118º 580 E 1,008 11.8 11.8 9.1 1,714 No
lanceolatae— Qingliangfeng 110 N data
Fagetum hayatae Nature Reserve
var. zhejiangensis in Zhejiang Prov.
19. Indocalamo 27. Sihaishan 28º 120º 160 E 927 13.6 13.2 10.4 1,864 No
latifolii-Fagetum Nature Reserve 260 N data
hayatae var. in Zhejiang Prov.
zhejiangensis
D D-a 20. Yushanio- 28. Mt. Lalashan, 24 121 260 E 1,529 14.2 17.1 14.4 2,663 No
Fagetum hayatae North-east 430 N data
Taiwan
III E E-a 21. Prismatomerio 29. Mt. 23 103 580 E 1,572 16.4 19.5 16.8 1,375 No
henryi— Caoguoshan in 070 N data
Lithocarpetum Yunnan Prov.
naiadari
22. Athyrio nardii— 30. Xiaoqiao 23º 104º 580 E 1,657 15.5 19.1 16.4 1,240 No
Phytogegraphical Analysis at the Family Level
Table 3.5 Endemic families in the flora of East Asia (mostly trees and shrubs)
Bretschneideraceae—one species: B. sinensis in SW-China, N-Vietnam and Taiwan, recently
connected with Akania (1 sp. in East-Australia)
Cercidiphyllaceae—one genus with two species in China mainly south of the Chang Jiang
(Yangtze) River and Japan (Katsura)
Dipterocarpaceae—over 700 species: main components of the tropical forests from East-Himalaya
to Yunnan, Hainan, Vietnam, Philippines, Indonesia, New Guinea
Lithocarpaceae—over 300 species, mainly in China, also in Japan and one species in North
America, now included in Fagaceae
Nyssaceae—six genera with ca. 30 species, mainly East-Asian and Indo-Malesian, recently
included in Cornaceae
Stachyuraceae—one genus with ten species: from East Himalaya to China mainly south of the
Chang Jiang (Yangtze) River and Japan
Trochodendraceae—Monotypic: T. aralioides in Japan and Taiwan
The life form of plant species provides significant information on the structure and
growth conditions of plant communities. In the phytogeographical literature, the
concept of life form was first introduced by A. von Humboldt at the beginning of the
nineteen century. The first classification was proposed by Raunkiaer, 100 years later
and is still used in Europe and in the Mediterranean area. Indeed, for the analysis of
the flora in other continents many difficulties arose, because of the underestimation
(in the European flora) of the ecophysiological difference between deciduous and
evergreen species and because other extraeuropean plant forms (e.g. succulents,
palms, bamboo) were not considered. Enlarged and improved classifications were
successively proposed by Ellenberg and Mueller-Dombois, by Box and others. In the
Japanese geobotanical tradition the classification proposed by Suzuki and Arakane
(1968) is used. This classification (cf. Table 3.6) differs in many aspects: by all
categories of woody plants evergreen species are separated from the deciduous ones;
in addition, some categories not considered by Raunkiaer are introduced: needle
leaved trees and shrubs and various types of bamboos.
In fact, one of the most peculiar characteristics of the beech forests in East Asia
is that evergreen and deciduous species mostly grow within mixed communities. In
consequence, also alliances, orders and classes of forest vegetation are composed of
3.4 Life Form Composition of the Plant Communities 137
Table 3.6 Number of species and percentage (%) within the contingency table of each life form
and phytosociological order
Orders
Life form abbreviations A B C D E
PD 15 (15.6) 101 (21.4) 135 (20.1) 5 (4.9) 9 (7.6)
PE 1 (1) 8 (1.7) 131 (19.5) 26 (25.5) 54 (45.8)
PEN 2 (2.1) 16 (3.4) 2 (0.3) 0 (0) 0 (0)
NPD 8 (8.3) 104 (22) 106 (15.8) 14 (13.7) 9 (7.6)
NPE 3 (3.1) 15 (3.2) 57 (8.5) 16 (15.7) 12 (10.2)
NPEN 0 (0) 2 (0.4) 0 (0) 0 (0) 0 (0)
GS 1 (1) 13 (2.7) 10 (1.5) 1 (1) 1 (0.8)
PL 12 (12.5) 28 (5.9) 84 (12.5) 9 (8.8) 8 (6.8)
Ch frut 1 (1) 3 (0.6) 3 (0.4) 2 (2) 0 (0)
Ch suffr 0 (0) 5 (1.1) 5 (0.7) 0 (0) 0 (0)
H scap 18 (18.8) 115 (24.3) 80 (11.9) 4 (3.9) 4 (3.4)
H scand 3 (3.1) 9 (1.9) 11 (1.6) 3 (2.9) 0 (0)
H caesp 2 (2.1) 22 (4.7) 26 (3.9) 1 (1) 3 (2.5)
H rosul 9 (9.4) 31 (6.6) 26 (3.9) 3 (2.9) 1 (0.8)
G rhiz 29 (30.2) 70 (14.8) 112 (16.7) 19 (18.6) 24 (20.3)
G rad 1 (1) 13 (2.7) 6 (0.9) 1 (1) 2 (1.7)
G bulb 4 (4.2) 15 (3.2) 9 (1.3) 2 (2) 0 (0)
G paras 0 (0) 2 (0.4) 1 (0.1) 0 (0) 0 (0)
T 2 (2.1) 2 (0.4) 2 (0.3) 1 (1) 0 (0)
E 2 (2.1) 5 (1.1) 4 (0.6) 2 (2) 0 (0)
Total 96 (100) 473 (100) 671 (100) 102 (100) 118 (100)
See Table 3.2 for details of the order
both elements: deciduous and evergreen; the proportion of both components can be
considered as an important basis for the definition of the superior vegetation units.
For this reason, after a detailed analysis of the genera present in the different
vegetation classes, we were able to reach a syntaxonomical scheme, based on
the differences in the ratio between evergreen and deciduous flora. However,
among those genera, 13 (Euonymus, Rhododendron, Smilax, Ligustrum, Meliosma,
Elaeagnus, Lonicera, Quercus, Ilex, Symplochos, Lindera, Magnolia, and Berchemia)
contain both evergreen and deciduous species. As it has already been clearly shown in
the description of plant communities, this particular difference is important in beech
forests of East Asia. Therefore, here, we treat in separated groups evergreen and
deciduous species even belonging to the same genus (Table 3.2).
We compared the life form composition of the species in each plant community
in order to analyze similarities between the communities. The life forms shown in
Tables 3.6, 3.7, and 3.8 were used for this comparison. However, there are signifi-
cant differences in the number of relevés for each plant community in the main
synoptic table of the vegetation (Tables 3.1a and 3.1b). In addition, there are data
that do not include complete flora lists, such as the data regarding Yunnan Province
138 3 Synthetic Remarks
Table 3.7 Number of species and percentage (%) within the contingency table of each life form
and phytosociological classes
Table 3.8 Number of species and percentage (%) within the contingency table of each life form and phytosociological association
Associations
Life form abbreviations A-a B-a B-b C-a C-b C-c C-d D-a E-a E-b
PD 15 (15.6) 77 (25.8) 86 (23.8) 39 (30.5) 81 (19.9) 37 (16.3) 26 (31.7) 5 (4.9) 7 (9.5) 4 (6.9)
PE 1 (1) 3 (1) 8 (2.2) 19 (14.8) 86 (21.1) 57 (25.1) 13 (15.9) 26 (25.5) 35 (47.3) 26 (44.8)
PEN 2 (2.1) 10 (3.3) 12 (3.3) 1 (0.8) 2 (0.5) 0 (0) 0 (0) 0 (0) 0 (0) 0 (0)
NPD 8 (8.3) 70 (23.4) 88 (24.3) 26 (20.3) 66 (16.2) 25 (11) 19 (23.2) 14 (13.7) 6 (8.1) 5 (8.6)
NPE 3 (3.1) 11 (3.7) 9 (2.5) 10 (7.8) 34 (8.3) 15 (6.6) 14 (17.1) 16 (15.7) 8 (10.8) 7 (12.1)
NPEN 0 (0) 2 (0.7) 0 (0) 0 (0) 0 (0) 0 (0) 0 (0) 0 (0) 0 (0) 0 (0)
GS 1 (1) 7 (2.3) 8 (2.2) 2 (1.6) 6 (1.5) 2 (0.9) 1 (1.2) 1 (1) 0 (0) 1 (1.7)
PL 12 (12.5) 20 (6.7) 25 (6.9) 19 (14.8) 48 (11.8) 34 (15) 13 (15.9) 9 (8.8) 6 (8.1) 3 (5.2)
Ch frut 1 (1) 1 (0.3) 3 (0.8) 2 (1.6) 1 (0.2) 0 (0) 0 (0) 2 (2) 0 (0) 0 (0)
Ch suffr 0 (0) 3 (1) 4 (1.1) 1 (0.8) 4 (1) 1 (0.4) 0 (0) 0 (0) 0 (0) 0 (0)
H scap 18 (18.8) 60 (20.1) 87 (24) 20 (15.6) 47 (11.5) 18 (7.9) 4 (4.9) 4 (3.9) 2 (2.7) 2 (3.4)
Life Form Composition of the Plant Communities
H scand 3 (3.1) 6 (2) 6 (1.7) 0 (0) 7 (1.7) 3 (1.3) 2 (2.4) 3 (2.9) 0 (0) 0 (0)
H caesp 2 (2.1) 15 (5) 15 (4.1) 2 (1.6) 17 (4.2) 6 (2.6) 2 (2.4) 1 (1) 2 (2.7) 1 (1.7)
H rosul 9 (9.4) 20 (6.7) 22 (6.1) 6 (4.7) 14 (3.4) 11 (4.8) 1 (1.2) 3 (2.9) 1 (1.4) 0 (0)
G rhiz 29 (30.2) 52 (17.4) 51 (14.1) 18 (14.1) 63 (15.4) 52 (22.9) 10 (12.2) 19 (18.6) 14 (18.9) 11 (19)
G rad 1 (1) 7 (2.3) 12 (3.3) 1 (0.8) 4 (1) 1 (0.4) 0 (0) 1 (1) 0 (0) 2 (3.4)
G bulb 4 (4.2) 9 (3) 10 (2.8) 1 (0.8) 7 (1.7) 1 (0.4) 2 (2.4) 2 (2) 0 (0) 0 (0)
G paras 0 (0) 2 (0.7) 1 (0.3) 0 (0) 1 (0.2) 0 (0) 0 (0) 0 (0) 0 (0) 0 (0)
T 2 (2.1) 1 (0.3) 1 (0.3) 0 (0) 1 (0.2) 1 (0.4) 1 (1.2) 1 (1) 0 (0) 0 (0)
E 2 (2.1) 1 (0.3) 5 (1.4) 0 (0) 3 (0.7) 1 (0.4) 0 (0) 2 (2) 0 (0) 0 (0)
Total 96 (100) 299 (100) 362 (100) 128 (100) 408 (100) 227 (100) 82 (100) 102 (100) 74 (100) 58 (100)
See Table 3.2 for details of the associtations
139
140 3 Synthetic Remarks
the proportions of evergreen trees (PE), evergreen shrubs (NPE) and herbs with
underground storage rhizome (G rhiz) is highest in Class III and lowest in Class I.
The total of herbaceous life forms is by far the highest in Class I, being 60.9 %,
compared to 39.9 % in Class II and 28.9 % in Class III.
Moreover the following eight life forms do not appear in Class III: evergreen
needleleaved trees (PEN), creeping low-shrubs (Ch frut), creeping sub-shrubs
(Ch-suffr), perennial herbs (climbing: H scand), herbs with underground storage
bulb (G bulb), herbs with underground parasitism (G paras), annual herbs (T) and
epiphytes (E). Among them, the absence of epiphytes (E) in Class III probably due
to lack of complete vegetation information, being Class III is the closest to tropical
areas. Nevertheless, there are important differences in life form compositions
between Class I, II and III. It is not reasonable to include Class III among the
communities of the beech forest vegetation, because beech forests are mainly
composed of deciduous flora, with Arcto-tertiary elements.
From these figures it can be said that the deciduous flora in Class I which is
distributed in Japan and Korea, is dominant and, therefore is the most similar to the
beech forests in other areas of the northern hemisphere in which the development of
the herb level has been observed.
The beech forests in China and Taiwan fit somewhere between Class I and
Class III in terms of overall life form composition, and they are very different from
those in Japan and Korea. This difference can already be observed in the species
composing the classes, and it provides an important argument in favour of
maintaining these two classes separately. As we have seen in the description of
the plant communities, while Class I is distributed in areas with low winter
temperatures and large amounts of snowfall, the beech forests in China and Taiwan
develop at high altitudes in warm mountainous regions where evergreen broad-
leaved forests grow, and its flora does not develop into deciduous broad-leaved
forests. Therefore, deciduous broad-leaved flora is mixed with evergreen broad-
leaved flora.
Further on, evergreen conifer life forms appear in Class I and II. This shows that
the relictic nature is preserved in the beech forests in both Classes I and II, as
evergreen conifers are to be considered surviving elements from the Neogene
period. This group is present with higher frequency in Class I, which, on the
other side, has also high proportion of deciduous trees. We consider that the elevate
presence of these groups is somehow related to the geological history of the island
system where Class I can be observed today, in which the evergreen coniferous
species had a chance to survive and the forest communities have reached a higher
degree of differentiation.
Table 3.6 shows the life form composition at an order level, classified by the
appearance of the same species in Tables 3.1a and 3.1b. From Table 3.6 we can see
the variations in each class. The numbers of relevés sampled in Order A and B in
Class I were not equal. However, in spite of the difference, more deciduous shrubs
(NPD) are seen in Order B in Japan, and the number of bamboo species is greater
too. Although the overall proportion of evergreen conifers is low, there exists a
wider variety of such species in Order B. On the other hand, the proportion of herbs
3.4 Life Form Composition of the Plant Communities 141
Although beech forests distributions in East Asia are disjunct with each other at a
macro scale, they form a distinct vegetation belt in each region. However, as the
beech forests are usually distant from human activity, it is not easy to obtain a
climatic data set of the beech forests. Moreover, even though a meteorological
observation point exists within a beech zone, it is not possible to represent the
climatic data as an entire representative climatic condition within the beech zone,
since one obtains from the observation point “punctual” data, which are not always
suitable to be extended to the entire area of a forest community. Nevertheless, we
believe that it is still worth while comparing climatic conditions at a large geo-
graphical scale, because in this way we are able to obtain a summarized outlook
on the climatic conditions within the entire beech zones of East Asia at least at a
macro scale.
We obtained mesh climatic data released by Japan Meteorological Agency
(1996) for Japan. This climatic data are given on a grid with spatial resolution of
c. 1 km. For the rest of the country, we obtained data set from World Climate
(Hijmans et al. 2005) at a 5 km spatial resolution. For each association, we selected
one or a few representative points on which relevés were obtained, and each of the
climatic information was obtained from those representative points. Occasionally,
elevation was different between the relevé and the mesh, because mesh resolution is
1 and 5 km and a variety of topography may exist within each mesh. In this case, we
tried to look up the climatic data from the neighbouring mesh cell with an elevation
value close to that of the relevé. Although elevation range of relevés ranges
between 357 and 1,954 m, most of the associations are distributed at an average
elevation of c. 1,000 m. Thus, we further estimated mean annual temperature at an
elevation of 1,000 and 1,500 m, by using rapse rate of 0.55 C/100 m. In this way,
it was easy to compare the temperature conditions between the associations.
Some of these temperatures were used also for the description of the different
vegetation units.
Mean annual temperature, annual precipitation, and maximum snow depth are
shown in Table 3.4. The locations of climatic data extracted for Table 3.4 are shown
in Fig. 3.1. Climatic conditions at 1,000 m elevation show clear differences between
the Class levels. That is, mean annual temperature for Fagetea crenatae Miyawaki,
Ohba et Murate 1964 (Class I) shows 10 C at most of the points, whereas Litseo
elongatae-Fagetea sp. div. cl. nov. (Class II) and an unnamed evergreen broad
leaved class (Class III) shows much higher temperature values (Table 3.4). The
lowest value is 3.1 C in Hokkaido, Japan, and the highest value is 19.2 C in
Yunnan, China. At elevation 1,500 m, which is close to the highest elevation of
beech forests in Japan, most of the temperature records in Japan show less than
7 C, whereas that in China and Taiwan show more than 10 C. Furthermore, at the
evergreen class (Class III), temperature shows over 16 C, which is the highest
among the three classes. This differences in temperature indicate that beech forests
in China and Taiwan grow under warmer climatic conditions than in Japan and
3.5 Relationships Between the Distributions of Vegetation and Climatic Conditions 143
Fig. 3.1 Locations of climatic data extracted. The numbers in the figure are the same as the
numbers assigned for the sites in Table 3.4
Korea, and moreover, that the evergreen class (Class III) is obviously under
different temperature conditions than the beech forest classes. At order and alliance
level, it is hard to grasp general trends in climatic conditions due to larger variations
in climate.
Beech forests in Korea and China are living under conditions of lower precipi-
tation, compared to those in Taiwan and Japan. Annual precipitation shows small
differences between the three classes; however, regional differences at the order
level become clear. We examine a first case: the class Fagetea crenatae Miyawaki,
Ohba et Murate 1964 (Class I); in the Japanese Saso-Fagetalia Suz.-Tok. 1966
(Order B:) a large range of precipitation is recorded, with values varying from 1,347
to 3,655 mm; in comparison, the Korean communities of Fagetalia multinervis
Kim, Kimura et Yim 1986 (Order A: 1,371 mm precipitation in the year), have a
low annual precipitation with the total value very close to the lowest observed in the
Saso-Fagetalia. A similar relationship can be measured within Litseo elongatae-
Fagetea sp. div. cl. nov. (Class II): Fagetalia hayatae Hukusima et al. 2005
(Order D) in Taiwan (2,663 mm), reaches a precipitation total, which is clearly
higher than in the mainland Chinese Sinarundinario nitidae—Fagetalia sp. div.
(Order C) where values between 917 and 1,864 mm are measured, and most of
the area is totalizing a yearly rainfall of less than 1,500 mm. Judging from the above
examples, it becomes clear that beech forests in Korea and China have been
developed in regions with relatively low rainfall, whereas beech forests in Taiwan
and Japan have been developed under conditions of elevate humidity.
At the alliance level, Abelio englerianae—Fagion all.nov. (Alliance C-a) in
Sichuan in China and Qiongzheo tumidinodae—Fagion all. nov. (Alliance C-c) in
144 3 Synthetic Remarks
Yunnan show precipitation values of less than 1,000 mm. The total rainfall in Japan
is mainly much higher and only for one locality a value under 1,500 mm is indicated
(see Table 3.4).
Information on snow depth was available only in Japan, but in Ulleungdo,
Korea, 1 m of snow depth is known (Hukusima pers. comm.), which is similar to
the snow conditions observed on the Sea of Japan side of Japan. Nanjyan in Sichuan
Province in China 30–40 cm of snow depth is known (Hukusima pers. comm.),
where Abelio englerianae—Fagion all.nov. (Alliance C-a) distributed. Sabhiankou
Nature Reseve in Yunnan province, c. 30 cm of snow depth is known (Hukusima
pers. comm.). It is speculated that snow depth of other areas in China is much less
than those reported above. Nevertheless duration period of snow accumulation
should be short, as it can be deduced from the high mean temperature in the region
(Table 3.4). In Japan, the areas of Fagion crenatae Suz.-Tok. 1952 (Alliance B-a),
an alliance on the Sea of Japan side, show much higher snow depth (116–277 cm)
than the areas of Sasamorpho-Fagion crenatae Miyawaki, Ohba et Murase 1964
(Alliance B-b), an alliance on the Pacific Ocean side (6–71 cm). In the areas of
Fagion crenatae Suz.-Tok. 1952 (Alliance B-a), Hokuriku, Tohoku, and Hokkaido
regions, that are the distribution areas of Saso kurilensis-Fagetum crenatae
Suz.-Tok. (Association 2), are particularly known for high snow fall, and over
2 m snow depth is recorded.
By considering the above-mentioned environmental conditions, it may be
concluded that vegetation units at the class level are more strongly associated
with temperature conditions. Indeed this does not match with the differences in
the proportions between evergreen and deciduous components of the flora. On the
other hand, the vegetation units at the order level are more associated with precipi-
tation conditions. This relation between temperature and precipitation shows that
beech forests in Japan are under the conditions of relatively cool climate with high
precipitation. Although beech forests in Korea (Ulleungdo) are under similar
conditions, the amount of precipitation is relatively lower (indeed this comparison
has to be judged with caution, because of the particular climatic conditions, e.g.
elevate atmospheric humidity in this rather small island). Although Chinese beech
forests are under warmer conditions compared to those in Korea or Japan, precipi-
tation is low, and thus forests in China are developing under dry conditions. Among
those Chinese beech forests, those living in Sichuan and Yunnan Provinces are
growing under particularly drier conditions. In contrast, beech forests in Taiwan
grow under relatively warm and wet conditions. As discussed above, differences in
climatic conditions for beech forests at each region are the most reflected within
each vegetation order which corresponds well to the different geographical
locations.
3.6 Some Reflections on Origin and Evolution of the Beech Forest in the. . . 145
The careful analysis of the diversity and regional specialization of beech forests in
East Asia allows to connect the different aspects (structure, ecology, geographical
distribution etc.) in one evolutionary hypothesis. This means to shift from the
synchronic view (all elements considered in the same time: the present) to the
diachronic view (how these elements interacted in the past).
The genus Fagus has a dozen of species occurring in different areas of the
northern hemisphere; in China 4 species occur (one of them with a variety endemic
in Taiwan), some other have been described (e.g. F. chienii) but remain still under
discussion. Anyway, China is the area where the genus Fagus is reaching its highest
diversity.
All Fagus species are deciduous trees. The genus Fagus belongs to the family
Fagaceae, mainly including genera of species with arboreal growth (Castanea,
Castanopsis, Fagus, Lithocarpus, Quercus) and closely related are other genera
of trees, e.g. Carpinus and Ostrya); some of these genera consist of evergreen
species, others are deciduous, whereas from the genus Quercus evergreen as well as
deciduous and “semi-evergreen” species are known. Most of these Fagaceae are
growing in mixed stands, often (e.g. in Quercus) several species can grow together
in the same community (often producing hybrids). All Fagaceae are anemophilous
(wind-pollinated), but as a secondary adaptation, deriving from a group of ento-
mophilous plants; the fruits have no adaptations for dissemination at distance, are
relatively heavy and fall at the basis of the stem (barochorous), but are edible and
often dispersed by animals. A peculiarity of the Fagus species is that their distribu-
tion is seldomly sympatric and in consequence, as a rule, in a single forest complex
only one species of Fagus occurs, even if in the vicinity other Fagus species grow
and in this case the one Fagus species has the tendency to invade the whole tree
canopy (an exception is the complex F. crenata–F. japonica in Japan). Seedlings of
Fagus sylvatica (Europe) can grow in the understory of a dense forest under
reduced solar irradiation, and this is an important factor to maintain a complete
dominance of the beech in the tree layer.
Another important indicator are the evergreen PEN and NPEN conifers, which can
also be interpreted as relict elements. Indeed, they are not widely diffused in the beech
forests of China. Conifers occur (with low frequency, up to 3 %) mostly in the beech
forests of Japan, under a cooler climate, as species of Abies and Picea, the latter
strictly connected with the boreal coniferous forest with circumboreal distribution.
In China conditions are quite peculiar: although four species of Fagus are
present, in general they grow in different, separate areas. The tree layer can consist
of mixed individuals of different species, several of them with evergreen leaves.
The presence of evergreen species is an important link with the evergreen woody
flora adapted to the subtropical climate of South China.
Under these conditions, China appears to be the centre of diversity (hot spot) for
this genus: several species occur in this area, and in general they remain with
regional distribution; with the progressive increasing of the distance from this hot
146 3 Synthetic Remarks
spot, the number of species decreases but their distribution range has the tendency
to expand: F. crenata on the whole Japanese Archipelago, F. orientalis from Iran to
the southernmost extremity of the Balcan Peninsula, F. sylvatica and F. grandifolia
with continental distribution in Europe and North America respectively.
A particular significance can be given to the South-western mountainous area in
the Yunnan Province, where the associations 21 and 22 are described. Following
Zhu (2008), the flora of this area is rich in endemic components and with the most
elevate frequency of subtropical evergreen elements in the arboreal vegetation; in
the Flora of South Yunnan the similarity with the tropical flora of Asia is 80 % at the
level of families and 64 % of genera. Indeed this is interpreted as a marginal type of
the Indo-Malaysian flora: “The tropical flora of southern Yunnan is supposed to be
derived from tropical Asian flora with the formation of the eastern monsoon climate
after the Tertiary”.
In fact, in this area it is possible to observe many endemic families and genera,
some of them appearing as relicts, which are considered as basal in the process of
the evolution of the eurasiatic flora. In this sense, the flira od South Yunnan
maintains a “Cathaysian” character.
In this frame it seems possible that such evergreen forest stands with Fagus
longepetiolata in the tree layer may offer a model for the earliest example of a
forest community, occurring in the subtropical-mountain environment, where a
deciduous tree species (namely Fagus) had the possibility to grow and to expand
in midst an evergreen broadlived community.
This deciduous tree species was successful and during the geological time slowly
expanded over the mountain areas of China, south of the Chang Jiang (Yangtze)
River. On the isolated mountain ranges followed a slow process of radiation, with
the evolving of several different Fagus species, as local endemics, characterizing the
communities of the class Litseo-Fagetea. In these beech forests there is still an
important presence of evergreen species remaining, but in the same time other
deciduous elements expand, as Acer, Carpinus, Fraxinus, Juglans, Prunus, Sorbus,
and the endemic and subtropical elements tend progressively to vanish.
The following event, of more recent geological age, is the expansion to the North
on the Japanese Archipelago (and Ulleungdo Island): here Fagus and other decidu-
ous trees are completely dominant in the forest canopy, and evergreen species
seem to have lost importance (or are substituted by the evergreen dwarf bamboo
Sasa, Arundinaria etc. which probably are also of Chinese origin). The area, in
present time occupied by the Fagetea crenatae communities was covered by the
pleistocenic glaciation, and later the expansion of these compact deciduous beech
forests in Honshu and Hokkaido is probably recent (postglacial). A similar recent
(holocene) expansion took place also in Europe and in the Eastern North America,
there probably from a still controversial species (F. mexicana, often reduced at
subspecies or variety level), rarely occurring in the mountains of Mexico, in contact
with the American subtropical evergreen forest.
Such relationships can be expressed also in a more abstract and quantitative
form, as part of two different evolutionary syndromes: on the one side (1), a
tendency to a condition of stability, with in situ survival of ancestral forms (relictual
3.6 Some Reflections on Origin and Evolution of the Beech Forest in the. . . 147
groups at the family, genus and species level), and on the other side (2) the natural
tendency to evolve, with the insurgence of new groups, with new ecomorphological
types, invading the marginal zones:
1. As indicators for stability, the following phytogeographical elements can be
considered: a—endemics; c—SE-Asiatic-tropical; d—Amphipacific; g—Tropical
2. As indicators for speciation and invasion of new areas we consider the following
elements: b—SE-Asiatic-Boreal; f—Boreal and Arctic-alpine; h—Cosmopolitan
The ratio between the sums of the frequences of the two groups of indicators for
the opposite tendencies can be expressed as a fraction where, in this case, in the
numerator is the sum of indicators of stability and in the denominator are those for
speciation and invasion. In this form we obtain an index of biological heritage.
aþcþdþg
Index of biological heritage ¼
bþfþh
From the sum of the values given in Table 3.3 for the single elements chosen as
indicators, we obtain the following figures:
Conclusions
The aim of the present study, is to propose a syntaxonomical synthesis for the
beech forests in East Asia. More than 20 plant communities with Fagus species
in the tree layer were compared and basing on their species composition, were
distributed among three groups (Table 3.1c, Fig. 2.1), corresponding to three
distinct vegetation classes.
In a first phase of our elaboration, it was possible to order the beech forests
into two groups: the first one with northern and insular distribution (Japan and
the Korean Island Ulleungdo), the other mostly in continental areas at lower
latitude, in China (a restricted presence also in Taiwan). The description of two
different classes is the result of a broad comparison among such beech forests.
The first group is organized by the class Fagetea crenatae, with insular distribu-
tion, and includes many deciduous tree and shrub species (and none or few
evergreen elements). The second group corresponds to the class Litseo
elongatae-Fagetea and consists of beech forests scattered among the continental
districts of China, which bear a progressively increasing number of evergreen
woody species. The two classes are very polymorphic and can be further
subdivided into four orders, eight alliances, and 20 associations, reaching a
gross total of 48 elementary units (subassociations) (Table 3.9). Tables 3.1a
and 3.1b are a large synoptic table (see also Table 3.1c, Online) giving particu-
lar evidence to the groups of character species which can be proposed for each
level (orders, alliances, associations). Among them beech forest associations of
Japan/Korea appear relatively homogeneous, whereas beech forests in China
show higher distances in species composition among the different associations.
This can be interpreted probably because the beech forests in China are
distributed over a large geographical area and are disjunct with each other,
occurring on isolated high mountain systems (Fig. 2.1).
Indeed, the last two associations (21 and 22, columns 49–51) do not fit with
these two classes: they belong to the type of the evergreen broad-leaved forest,
150 3 Synthetic Remarks
but with the penetration of beech species in the tree layer, together with some
other elements from the deciduous woodlands. In consequence, they are ascribed
to a third, still not defined class with subtropical mountain character.
To reach this result, a large amount of information was elaborated and
critically discussed. The synoptic association table is based on 657 relevés
obtained from previous literature and original data of the authors, selected
among several thousands of relevés. These relevés are considered the raw data
for the description of the Fagus vegetation types in East Asia. Further informa-
tion was added, through the definition for every association (and for every
species) of the life forms, life cycle (deciduous—evergreen) and of the compo-
sition at the level of genera: these are meta-data, very significant for the
interpretation of the relationships among the different associations.
The life form composition is a very distinctive characteristic of each unit, at
the class and order level, but as we move down the classification at the alliance
level or below, differences in the life forms became lesser significant. However
the global species composition as well as the presence of character species are
highly informative at the alliance, association and subassociation level.
According to Table 3.1d (Online), the diversity in genus combinations shows
more clearly the differences among the three classes and five orders as the analysis
of species composition. However the number of characteristic genera decreases at
the level of alliance and none are observed at association or subassociation level.
Again this indicates that the investigation at the association and subassociation
level should be focussed at the species, rather than at the genus level.
The current study of species composition and life forms confirms that the
beech forests of China and Taiwan and those of Japan and Korea are clearly
diverging from one another, so that the separation of the beech forests of East
Asia into two main classes appears largely justified. It seems that in the future it
will be necessary to extend forward this result in order to cover the beech forests
of the entire Northern Hemisphere through a comparison with the examples
existing in Europe and North America (see also, Pignatti 1977).
An important feature to understand the significance of the differences
between these classes are the relationships between the presence of evergreen
and deciduous species within the different types of beech forests. The number of
deciduous species regularly decreases from class I to II and III; at the same time
the presence of the evergreen flora increases.
At the order level and alliance level, regional flora including geographically
endemic species characterise each order and alliance. Especially in the islands
of Ulleungdo and Taiwan, where the distribution range is limited to small
areas, endemism is high: in this case, beech forests are considered relictual
communities. In contrast, in Japan beech forests are widely distributed. Espe-
cially in the middle part of Japan northwards, beech forest vigorously develop
and show a high uniformity in species composition. However as beech forests in
western Japan are sporadic on isolated mountains ranges, the presence of the
regional flora is often prevailing. A similar tendency can be observed in the
vegetation of China, where beech forests are distributed in a wide area, but occur
3.6 Some Reflections on Origin and Evolution of the Beech Forest in the. . . 151
Associations Tables
T. Hukusima et al., Phytosociology of the Beech (Fagus) Forests in East Asia, 153
Geobotany Studies, DOI 10.1007/978-3-642-35620-9,
# Springer-Verlag Berlin Heidelberg 2013
154 Appendix
Association number 7 8
Char.sp. of Vaccinio henryi-Fagetum hayatae subsp. pashanicae ass. nov.
Fagus pashanica ∙ ∙ ∙ ∙ ∙ ∙ 5.5 5.5 5.5
Rubus bambusarus ∙ ∙ ∙ ∙ ∙ ∙ + + +
Hugeria vaccinioides ∙ ∙ ∙ ∙ ∙ ∙ ∙ + +
Vaccinium henryi ∙ ∙ ∙ ∙ ∙ ∙ ∙ + +
Daphniphyllum ∙ ∙ ∙ ∙ ∙ ∙ + + ∙
angustifolium
Char.sp. of Abelio englerianae-Fagion all. nov.
Rhododendron + ∙ ∙ 1.2 ∙ ∙ + 2.2 ∙
micranthum
Prunus pilosiuscula + ∙ ∙ ∙ ∙ + + ∙ +
Euonymus giraldii ∙ ∙ + + + + + ∙ ∙
Calanthe fimbriata ∙ +0.2 + ∙ ∙ + + ∙ ∙
Abelia engleriana ∙ ∙ 3.3 ∙ ∙ ∙ + ∙ ∙
Quercus glandulifera 2.2 2.1 2.2 2.2 1.1 ∙ ∙ + ∙
Acer laxiflorum ∙ ∙ + ∙ + ∙ ∙ + ∙
Holboellia fargesii ∙ ∙ ∙ + + + ∙ + +
Epimedium sagittatum ∙ ∙ ∙ + + ∙ + ∙ ∙
Lonicera ∙ ∙ ∙ 1.1 + ∙ + ∙ ∙
pseudoproterantha
Ainsleaea triflora ∙ ∙ ∙ ∙ ∙ + ∙ + +
Berberis dielsiana ∙ ∙ ∙ ∙ ∙ + + ∙ +
Char.sp. of Sinarundinario nitidae-Fagetalia sp. divar.
Fraxinus chinensis ∙ ∙ ∙ ∙ ∙ ∙ ∙ + +
Pylora decorata ∙ ∙ ∙ ∙ + ∙ ∙ ∙ ∙
Stewartia sinensis 2.2 1.1 ∙ + ∙ ∙ + + +
Viburnum sympodiale 1.1 + 2.2 + 1.1 ∙ 1.1 2.2 2.2
Paederia scandens ∙ ∙ ∙ ∙ ∙ + ∙ ∙ ∙
Lindera glauca ∙ ∙ + ∙ + ∙ ∙ ∙ ∙
Acanthopanax ∙ ∙ ∙ ∙ ∙ ∙ ∙ + +
evodiaefolius
Ilex henryi ∙ ∙ 1.1 ∙ + + ∙ + +
Sinarundinaria nitida 4.4 4.4 ∙ 2.2 4.4 + ∙ ∙ 2.2
Sorbus folgneri ∙ + + ∙ 1.1 + ∙ + +
Acer sinense ∙ ∙ ∙ + + + ∙ ∙ +
Polystichum + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
neolobatum
Fagus lucida 4.4 5.5 ∙ ∙ ∙ ∙ ∙ ∙ ∙
Smilax stans + ∙ + ∙ ∙ + + ∙ ∙
Ophiopogon bodinieri ∙ ∙ + + ∙ + ∙ + ∙
Quercus oxyodon ∙ ∙ ∙ ∙ ∙ ∙ 2.2 + +
Pinus armandii ∙ ∙ + + + +0.2 ∙ + +
Dryopteris labordei ∙ ∙ ∙ + + ∙ ∙ ∙ ∙
(continued)
Associations Tables 155
Association number 7 8
Acer oliverianum ∙ + ∙ + + ∙ ∙ + ∙
Fagus engleriana 3.3 2.2 5.5 5.5 5.5 3.3 + ∙ ∙
Eurya brevistyla ∙ ∙ ∙ ∙ ∙ ∙ 2.2 ∙ 2.2
Char.sp. of Litseo elongatae-Fagetea sp.givar.cl.nov.
Smilax discotis ∙ + ∙ ∙ ∙ ∙ ∙ ∙ +
Parathelypteris ∙ ∙ ∙ ∙ ∙ + ∙ ∙ ∙
glanduligera
Common species of the beech forests in East Asia
Cornus controversa + + ∙ ∙ + ∙ ∙ ∙ ∙
Matteuccia orientalis ∙ ∙ ∙ ∙ + + ∙ ∙ ∙
Oxalis griffithii 1.1 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Acer mono ∙ + + ∙ ∙ ∙ ∙ ∙ ∙
Athyrium wardii ∙ ∙ ∙ +0.2 + ∙ ∙ ∙ ∙
Schisandra chinensis ∙ ∙ ∙ + + + ∙ + 1.1
Lyonia ovalifolia var. ∙ + ∙ ∙ ∙ ∙ 1.1 ∙ ∙
elliptica
Lindera obutisiloba + ∙ + ∙ ∙ + ∙ ∙ ∙
Companions
Quercus myrsinaefolia 1.1 1.2 1.1 1.1 ∙ 1.1 + + ∙
Carex sp. + + 3.3 + + 3.3 + ∙ ∙
Taxus chinensis + + +0.2 ∙ + 1.1 ∙ ∙ ∙
Symplocos paniculata ∙ 1.1 ∙ + 1.1 1.1 ∙ ∙ ∙
Helwingia chinensis ∙ + ∙ + + ∙ ∙ + +
Ilex chinensis ∙ ∙ ∙ ∙ + + 1.1 1.1 ∙
Celastrus sp. ∙ ∙ + + 1.1 ∙ + ∙ ∙
Abelia dielsii ∙ ∙ ∙ 1.1 1.1 2.2 ∙ ∙ ∙
Symplocos laurina 1.1 3.3 ∙ ∙ ∙ ∙ 1.1 ∙ ∙
Cornus kousa 1.1 ∙ ∙ 1.1 ∙ 1.1 ∙ ∙ ∙
Carpinus 1.1 + ∙ ∙ ∙ ∙ + ∙ ∙
turczanninowii
Eurya alata ∙ + ∙ + ∙ ∙ ∙ 3.3 ∙
Enkianthus chinensis ∙ ∙ ∙ ∙ + ∙ 2.2 ∙ +
Lonicera henryi ∙ ∙ + + ∙ ∙ ∙ + ∙
Sorbus sp. + ∙ ∙ ∙ ∙ ∙ +0.2 ∙ ∙
Lithocarpus sp. ∙ ∙ ∙ ∙ ∙ ∙ 1.1 ∙ ∙
Ophiopogon japonicus + ∙ +0.2 ∙ ∙ ∙ +0.2 ∙ ∙
Sorbus xanthoneura ∙ ∙ ∙ ∙ ∙ 1.1 1.1 ∙ +
Carpinus fargesii ∙ ∙ + + ∙ ∙ ∙ ∙ ∙
Carpinus polyneura 1.1 1.1 ∙ ∙ ∙ ∙ ∙ ∙ ∙
Rhododendron 2.2 ∙ ∙ + ∙ ∙ ∙ ∙ ∙
sutchuenense
Rhododendron sp. ∙ ∙ ∙ ∙ + ∙ ∙ 3.3 ∙
Rhus sp. + ∙ ∙ ∙ ∙ + + + ∙
(continued)
156 Appendix
Association number 7 8
Dipelta floribunda + ∙ ∙ ∙ ∙ ∙ ∙ ∙ +
Clematis mantana + ∙ ∙ ∙ + ∙ ∙ ∙ ∙
Rhododendron fargesii ∙ 2.2 ∙ ∙ ∙ ∙ ∙ ∙ 4.4
Enkianthus deflexus ∙ 1.1 ∙ 1.1 ∙ ∙ ∙ ∙ ∙
Dryopteris sp. + ∙ ∙ + ∙ ∙ ∙ ∙ ∙
Chimaphila japonica ∙ ∙ + ∙ ∙ ∙ + ∙ ∙
Arctous alpinas ∙ ∙ ∙ ∙ ∙ + + ∙ ∙
Paris bashanensis ∙ ∙ ∙ ∙ + ∙ + ∙ ∙
Ilex szechwanensis ∙ ∙ ∙ + + ∙ ∙ ∙ ∙
Acer palmatum ∙ ∙ ∙ ∙ + + ∙ ∙ ∙
Rhododendron ∙ ∙ ∙ ∙ ∙ ∙ 3.3 2.2 ∙
hypoglancum
Aruncus sylvester ∙ ∙ + ∙ ∙ + ∙ ∙ ∙
Smilax glanco-china + ∙ ∙ ∙ ∙ + ∙ ∙ ∙
Cornus macrophylla ∙ ∙ ∙ ∙ ∙ ∙ + 1.1 ∙
Viburnum sp. ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ 1.1
Michelia sp. + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Eurya obutsifolia + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Acanthopanax henryi + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Tupistra chinensis + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Lindera cercifolia + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Dactylicapnos sp. + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Viola biflora + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Asparagus filicinus + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Euptelea pleiospermum ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Cacalia profundorum ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Acer sp. ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Mahonia bealei ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Ainsleaea grossedentata ∙ ∙ 1.1 ∙ ∙ ∙ ∙ ∙ ∙
Prunus sp. ∙ ∙ 1.1 ∙ ∙ ∙ ∙ ∙ ∙
Rhododendron ∙ ∙ +0.2 ∙ ∙ ∙ ∙ ∙ ∙
anthopogonoides
Thaladiantha glabra ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙
Meliosma flexuosa ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙
Corylos heterophylla v. ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙
sutchuenensis
Cornus paucinervis ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙
Disporum cantoniense ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙
Cynanchum chinense ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙
Gllium aparine v. ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙
tenerum
Ajuga nipponensis ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙
Stellaria alsine ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙
(continued)
Associations Tables 157
Association number 7 8
Desmodium ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙
szechuenense
Litsea ichangensis ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙
Polystichum ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙
squarrosum
Pollia sp. ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙
Pylora rugosa ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙
Cyrtomium ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙
macrophyllum
Stauntonia sp. ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙
Actinidia sp. ∙ ∙ ∙ + ∙ ∙ ∙ ∙ ∙
Carpinus cordata v. ∙ ∙ ∙ + ∙ ∙ ∙ ∙ ∙
chinensis
Arisaema erubescens ∙ ∙ ∙ + ∙ ∙ ∙ ∙ ∙
Acanthopanax giraldii ∙ ∙ ∙ + ∙ ∙ ∙ ∙ ∙
Deyeuxia arundinacea ∙ ∙ ∙ + ∙ ∙ ∙ ∙ ∙
Sorbus hemslaya ∙ ∙ ∙ ∙ + ∙ ∙ ∙ ∙
Stewartia sp. ∙ ∙ ∙ ∙ + ∙ ∙ ∙ ∙
Berberis dasytachya ∙ ∙ ∙ ∙ + ∙ ∙ ∙ ∙
Rubus flosculosus ∙ ∙ ∙ ∙ + ∙ ∙ ∙ ∙
Acanthopanax sp. ∙ ∙ ∙ ∙ + ∙ ∙ ∙ ∙
Viola sp. ∙ ∙ ∙ ∙ + ∙ ∙ ∙ ∙
Clematocleththra ∙ ∙ ∙ ∙ + ∙ ∙ ∙ ∙
scandens
Chloranthus henryi ∙ ∙ ∙ ∙ + ∙ ∙ ∙ ∙
Betula sp. ∙ ∙ ∙ ∙ ∙ 1.1 ∙ ∙ ∙
Sorbus caloneura ∙ ∙ ∙ ∙ ∙ 3.3 ∙ ∙ ∙
Acer shensiense ∙ ∙ ∙ ∙ ∙ + ∙ ∙ ∙
Smilax ferox ∙ ∙ ∙ ∙ ∙ + ∙ ∙ ∙
Indocalamus sp. ∙ ∙ ∙ ∙ ∙ 1.2 ∙ ∙ ∙
Ligustrum sinense ∙ ∙ ∙ ∙ ∙ + ∙ ∙ ∙
Cynanchum julianae ∙ ∙ ∙ ∙ ∙ + ∙ ∙ ∙
Abelia engleriniana ∙ ∙ ∙ ∙ ∙ + ∙ ∙ ∙
Thalictrum uncatum ∙ ∙ ∙ ∙ ∙ + ∙ ∙ ∙
Desmodium sp. ∙ ∙ ∙ ∙ ∙ + ∙ ∙ ∙
Vitis flexuosa ∙ ∙ ∙ ∙ ∙ + ∙ ∙ ∙
Ostrya japonica ∙ ∙ ∙ ∙ ∙ + ∙ ∙ ∙
Photinia parviflora ∙ ∙ ∙ ∙ ∙ + ∙ ∙ ∙
Quercus asine ∙ ∙ ∙ ∙ ∙ ∙ 2.1 ∙ ∙
Vaccinium sp. ∙ ∙ ∙ ∙ ∙ ∙ 1.1 ∙ ∙
Buxus henryi ∙ ∙ ∙ ∙ ∙ ∙ 2.2 ∙ ∙
Pieris tomentosa ∙ ∙ ∙ ∙ ∙ ∙ 1.1 ∙ ∙
Polystichum tsus- ∙ ∙ ∙ ∙ ∙ ∙ +0.2 ∙ ∙
sinense
(continued)
158 Appendix
Association number 7 8
Machilus faberi ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙
Hedera sp. ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙
Ainsleaea rubrinervis ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙
Ficus sp. ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙
Phoebe sheareri ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙
Cymbidium faberi ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙
Aristolochia tubiflora ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙
Akebia trifoliata v. ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙
australis
Pleuropus euchloron ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙
Aconitum sinomontanus ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙
Euonymus fortunei ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙
Goodyera biflora ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙
Viola arcuta ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙
Pieris formosana ∙ ∙ ∙ ∙ ∙ ∙ ∙ 1.1 ∙
Daphniphillum sp. ∙ ∙ ∙ ∙ ∙ ∙ ∙ + ∙
Plagiogyria ∙ ∙ ∙ ∙ ∙ ∙ ∙ + ∙
distinctissima
Humata tyermanni ∙ ∙ ∙ ∙ ∙ ∙ ∙ + ∙
Rhododendron ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ 1.1
augustinii
Phymatopsis teneupes ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ +
Association table of 15, 16 and 17 in alliance, Qiongzheo tumidinodae—Fagion all. nov.; 15. Sinocalamo giganteus—Fagetum lucidae ass. nov.; 16. Viburno
flavescentis—Fagetum englerianae ass. nov.; 17. Tripterospermo cordifolium—Fagetum englerianae ass. nov.
Association number 15 16 17
Running number 1 2 3 4 5 1 2 1 2 3 4 5
Stand number YNN5 YNN2 YNN3 YNN4 YNN6
Associations Tables
Altitude (m) 2,010 1,780 1,800 1,900 1,930 1,700 1,600 1,800 1,750 1,800 1,850 1,640
Exposure N20E N80E S80E S20W S70W NE NW W NE
Inclination ( ) 31 35 25 42 32 45 40 4 38 25 15
Area 225 150 200 200 225 400 400 400 400 400 400 400
Tree layer Hight (m) 17 17 17 13 20 19 19 22 19 21 24 21
Cover (%) 80 80 75 95 60 90 80 95 85 85 90 85
Sub tree layer Hight(m) 8 7 8 6 7
Cover (%) 65 50 55 50 40
Shrub layer (m) 3.0 2.0 1.5 2.0 2.5
Cover (%) 45 20 65 20 70
Herb layer (m) 0.5 0.5 0.5 0.5 0.5
Cover (%) 20 10 15 50 50
Number of species 64 63 39 49 62
Char.sp. of Sinocalamo giganteus-Fagetum lucidae ass. nov.
Sinanocalamus giganteus 23 ∙ 44 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Galium asperuloides var. hoffmeisteris + + ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Eurya graffi ∙ 22 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Carpinus fangiana ∙ ∙ 11 ∙ 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙
Eurya semisenullata 11 ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Symplocos ramosissima ∙ ∙ + + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Allantodia squamigera ∙ + ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Microlepia marginata + ∙ ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Ligustrum delavayanum ∙ + ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
159
(continued)
160
Association number 15 16 17
Spatholirion longifolium ∙ ∙ + + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Acer prolificum ∙ ∙ + + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Hydrangea xanthoneura ∙ ∙ ∙ + + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Arachnioides pseudo-aristata ∙ ∙ ∙ + + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Char.sp. of Viburno flavescentis-Fagetum englerianae ass. nov.
Neolitsea chuii ∙ ∙ ∙ ∙ ∙ 22 33 ∙ ∙ ∙ ∙ ∙
Toxicodendron radicans var. hispidus ∙ ∙ ∙ ∙ ∙ 12 22 ∙ ∙ ∙ ∙ ∙
Athyrium delavayi ∙ ∙ ∙ ∙ ∙ 11 22 ∙ ∙ ∙ ∙ ∙
Dichroa febrifuga ∙ ∙ ∙ ∙ ∙ 11 11 ∙ ∙ ∙ ∙ ∙
Lepisorus bicolor ∙ ∙ ∙ ∙ ∙ +2 +2 ∙ ∙ ∙ ∙ ∙
Polypodiastrum dielsianum ∙ ∙ ∙ ∙ ∙ +2 +2 ∙ ∙ ∙ ∙ ∙
Sorbus sargentiana ∙ ∙ ∙ ∙ ∙ + + ∙ ∙ ∙ ∙ ∙
Dendrobenthamia melanotricha ∙ ∙ ∙ ∙ ∙ 33 ∙ ∙ ∙ ∙ ∙ ∙
Viburnum flavescens ∙ ∙ ∙ ∙ ∙ 22 ∙ ∙ ∙ ∙ ∙ ∙
Davidia involucrata var. vilmoriniana ∙ ∙ ∙ ∙ ∙ ∙ 11 ∙ ∙ ∙ ∙ ∙
Phyllagathis longipes ∙ ∙ ∙ ∙ ∙ ∙ 11 ∙ ∙ ∙ ∙ ∙
Smilacina yunnanensis ∙ ∙ ∙ ∙ ∙ ∙ 11 ∙ ∙ ∙ ∙ ∙
Schizophragma hypoleuca ∙ ∙ ∙ ∙ ∙ ∙ 11 ∙ ∙ ∙ ∙ ∙
Smilax opaca ∙ ∙ ∙ ∙ ∙ ∙ 11 ∙ ∙ ∙ ∙ ∙
Char.sp. of Tripterospermo cordifolium–Fagetum englerianae ass. nov.
Schima crenata ∙ ∙ ∙ ∙ ∙ ∙ ∙ 11 11 11 + 33
Neolitsea chinensis ∙ ∙ ∙ ∙ ∙ ∙ ∙ 22 + 33 + +
Lindera subcaudata var. hemsleyana ∙ ∙ ∙ ∙ ∙ ∙ ∙ + + + + 11
Panicum brevifolium ∙ ∙ ∙ ∙ ∙ ∙ ∙ + + ∙ + +
Manglietia duclouxii ∙ ∙ ∙ ∙ ∙ ∙ ∙ 11 ∙ + + +
Appendix
Tripterospermum cordifolium ∙ ∙ ∙ ∙ ∙ ∙ ∙ + 11 + ∙ +
Sorbus coronata ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ + + +
Microsorium buergerianum ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ + ∙ +2 11
Ophiopogon bockianus ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ 11 + ∙ +
Elatostema lineolatum ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ + 22 +
Associations Tables
Association number 15 16 17
Char.sp. of Sinarundinario nitidae-Fagetalia sp. div.
Fagus lucida 33 44 22 22 33 ∙ ∙ ∙ ∙ ∙ ∙ ∙
Lithocarpus cleistcarpus + 11 22 + 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙
Symplocos botryantha + 11 33 11 + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Acanthopanax evodiaefolius 22 + 22 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ +
Viola schneideri ∙ + + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Quercus oxyodon + 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Viburnum sympodiale 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Dryopteris labordei ∙ +2 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Smilax stans ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Ophiopogon bodinieri ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Eurya brevistyla ∙ ∙ ∙ ∙ ∙ 11 + + 44 11 33 33
Lithocarpus hancei ∙ ∙ ∙ ∙ ∙ 33 ∙ 11 ∙ 11 + 22
Machilus ichangensis ∙ ∙ ∙ ∙ ∙ 11 11 11 33 33 + 11
Symplocos anomala ∙ ∙ ∙ ∙ ∙ ∙ ∙ + ∙ 11 11 +
Fagus engleriana ∙ 11 ∙ ∙ ∙ 44 33 11 22 + + 33
Plagiogyria stenoptera + + ∙ + ∙ 22 + ∙ ∙ 22 +2 +
Acer oliverianum ∙ +2 ∙ ∙ ∙ 11 33 11 + + 22 +
Rubus malifolius ∙ ∙ ∙ ∙ ∙ 11 11 + ∙ ∙ + +
Hydrangea anomala ∙ ∙ ∙ ∙ 22 ∙ ∙ ∙ ∙ + 11 +
Ilex wilsonii ∙ ∙ ∙ ∙ ∙ ∙ ∙ + 11 + ∙ 11
Symplocos caudata ∙ ∙ ∙ ∙ ∙ + 11 ∙ 33 + 22 +
Reineckia carnea ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ 11 ∙ + ∙
Char.sp. of Litseo elongatae-Fagetea sp. div. cl. nov.
Litsea elongata ∙ ∙ ∙ ∙ ∙ ∙ ∙ + + + 11 +
Appendix
Parathelypteris glanduligara +2 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Dennstaedtia scabra ∙ + ∙ + ∙ ∙ 22 ∙ ∙ ∙ ∙ ∙
Common species of the beech forests in East Asia
Cornus controversa ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ + ∙ + ∙
Euonymus fortunei var. radicans ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ + ∙ +
Associations Tables
Tiarella polyphylla ∙ + ∙ ∙ + ∙ ∙ ∙ ∙ ∙ 11 ∙
Ardisia crenata ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ + + ∙ +
Plagiogyria euphlebia ∙ ∙ ∙ ∙ ∙ ∙ ∙ +2 11 + + +
Matteuccia orientalis ∙ 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Athyrium wardii + + ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Oxalis griffithii ∙ + ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Companions
Carex sp. ∙ 11 + ∙ ∙ + 11 ∙ 11 + + 11
Camellia sp. ∙ 11 + + 11 ∙ ∙ ∙ ∙ ∙ ∙
Lonicera henryi + ∙ + ∙ ∙ ∙ ∙ ∙ ∙ + ∙ +
Sarcopyramis bodinieti ∙ ∙ ∙ ∙ ∙ 11 ∙ ∙ 11 + ∙ +
Photinia sp. ∙ + 11 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Smilax sp. + ∙ + + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Symplocos sp. 33 + ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Rhododendron sp. 22 ∙ ∙ 22 + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Camellia pitardii + ∙ + 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙
Polystichum makinoi ∙ + ∙ + 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙
Polystichum sp. ∙ ∙ ∙ ∙ ∙ ∙ ∙ 22 ∙ ∙ 11 +
Embelia sp. ∙ ∙ ∙ ∙ ∙ ∙ ∙ + ∙ + ∙ +
Eurya sp. ∙ ∙ ∙ ∙ ∙ ∙ 11 44 11 ∙ ∙ ∙
Symplocos discolor ∙ ∙ ∙ ∙ ∙ 22 ∙ ∙ 22 11 ∙ ∙
Viola sp. ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ 11 ∙ + 11
163
(continued)
164
Association number 15 16 17
Quercus myrsinaefolia ∙ ∙ ∙ ∙ ∙ ∙ ∙ 11 ∙ 11 ∙ 22
Actinidia chinensis ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ + +
Rhus sp. + 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Quercus engleriana ∙ ∙ ∙ 11 + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Prunus sp. ∙ 11 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Arisaema sp. ∙ +2 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Acer sp. 11 ∙ ∙ ∙ 22 ∙ ∙ ∙ ∙ ∙ ∙ ∙
Sorbus sp. + ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Rubus swinhoei + ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Chinamomum sp. 11 ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Rubus sp. + ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Castanopsis sp. + ∙ ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Dystyllum sp. + ∙ ∙ ∙ 22 ∙ ∙ ∙ ∙ ∙ ∙ ∙
Hydrangea davidii + + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Viola brunneostipulosa ∙ 11 ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Ophiopogon intermedius ∙ ∙ ∙ + + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Illicium simonsii ∙ ∙ ∙ 11 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙
Dioscoea sp. ∙ + ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Schisandra sp. ∙ ∙ + ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Ilex franchetiana + ∙ ∙ ∙ ∙ 22 ∙ ∙ ∙ ∙ ∙ ∙
Hedera nepalensis var. sinensis ∙ ∙ ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ +
Symplocos paniculata ∙ ∙ ∙ ∙ ∙ 11 33 ∙ ∙ ∙ ∙ ∙
Camellia sp. ∙ ∙ ∙ ∙ ∙ 11 + ∙ ∙ ∙ ∙ ∙
Helwingia himalaica ∙ ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙ 22 ∙
Lonicera ligustrina ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙ 11
Appendix
Pilea martinii ∙ ∙ ∙ ∙ ∙ ∙ ∙ 11 ∙ ∙ + ∙
Meliosma kirkii ∙ ∙ ∙ ∙ ∙ ∙ ∙ 11 ∙ ∙ + ∙
Osmanthus sp. ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ + ∙ +
Beilschmiedia robusta ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ 11 +
Hopea sp. 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Associations Tables
Viburnum sp. 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Cremanthodium decaisnei 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Aucuba chinensis 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Lithocarpus variolosus 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Pterostyrax rosea + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Quercus angustinii + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Acanthopanax simonii + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Ophiopogon maiei + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Ilex yunnanensis + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Sorbus rufoilosa + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Cystopteris moupinensis + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Polystichum discretum + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Smilacina glabra + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Viburnum cordifolium + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Ainsleaea yunnanensis + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Panax transitorius + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Lindera communis + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Eurya sp. + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Diospyros sp. + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Lonicera sp. + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Neolitsea sp. + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Ophiopogon sp. + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
165
(continued)
166
Association number 15 16 17
Schizophragma spl + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Lindera sp. + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Primula sp. + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Acer laxiflorum + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Lithocarpus sp. + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Ficus sp. + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Rhus delavayi ∙ 11 ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Elatostema obtusum ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Eurya loquaina ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Acanthopanax sp. ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Ilex sp. ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Phymatopsis sp. ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Paederia scandens ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Oplismenus undulatifolius ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Schisandra sphananthera ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Smilax boskii ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Cyrtomium mecrophyllum ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Allantodia metteniana ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Ctenitis mariformis ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Arachniodes chinensis ∙ ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Pseudocystopteris atkinsonii ∙ ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Magnolia sp. ∙ ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Calastrus sp. ∙ ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙
Helwingia chinensis ∙ ∙ ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Stauntonia sp. ∙ ∙ ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Appendix
Actinidia sp. ∙ ∙ ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Peracarpa sp. ∙ ∙ ∙ ∙ + ∙ ∙ ∙ ∙ ∙ ∙ ∙
Machilus sp. ∙ ∙ ∙ ∙ ∙ 33 ∙ ∙ ∙ ∙ ∙ ∙
Begonia sp. ∙ ∙ ∙ ∙ ∙ 11 ∙ ∙ ∙ ∙ ∙ ∙
Ilex ciliospinosa ∙ ∙ ∙ ∙ ∙ 11 ∙ ∙ ∙ ∙ ∙ ∙
Associations Tables
Pellionia radicans ∙ ∙ ∙ ∙ ∙ ∙ 22 ∙ ∙ ∙ ∙ ∙
Dryopteris sp. ∙ ∙ ∙ ∙ ∙ ∙ 11 ∙ ∙ ∙ ∙ ∙
Impatiens siculifer ∙ ∙ ∙ ∙ ∙ ∙ 11 ∙ ∙ ∙ ∙ ∙
Ilex fragilis ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ 22 ∙ ∙ ∙
Prunus pilosiuscula ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ 11 ∙ ∙ ∙
Microtropis fokienensis ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙ ∙
Tetrastigma hypoglaucum ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ + ∙ ∙ ∙
Toona sinensis ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ 11
Asteropyrum peltatum ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ 11
Epipactis sp. ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ ∙ +
167
Appendix 1. Alphabetic List—Species of the
Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Abelia dielsii Caprifoliaceae NPD Char. sp. of the Carici lanceolatae-
Fagetum hayatae
Abelia engleriana Caprifoliaceae NPD Char.-sp. of Abelio englerianae-
Fagion sp. div.
Abelia macrotera Caprifoliaceae NPD Char. sp. of the Fagetum engleriano-
lucidae
Abelia parvifolia Caprifoliaceae NPD Companion species with sporadical
presence (mainly China)
Abelia serrata Caprifoliaceae NPD Companion species with sporadical
presence (mainly Japan)
Abelia sp. Caprifoliaceae NPD Companion species with sporadical
presence (mainly China)
Abelia spathulata Caprifoliaceae NPD Char.-sp. of Sasamorpho_Fagion
crenatae
Abelia spathulata var. Caprifoliaceae NPD Companion species with sporadical
sanguinea presence (mainly Japan)
Abelia spathulata var. Caprifoliaceae NPD Companion species with sporadical
stenophylla presence (mainly Japan)
Abies firma Pinaceae PEN Species of the Quercetalia serratae-
grosseserratae
Abies homolepis Pinaceae PEN Char.-sp. of Sasamorpho_Fagion
crenatae
Abies mariesii Pinaceae PEN Diff.-sp. of Saso kurilensis-Fagetum
crenatae
Acanthopanax evodiaefolius Araliaceae PD Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Acanthopanax evodiaefolius Araliaceae PD Char. sp. of the Quingzheo
var. gracilis tumidinodae-Fagion
Acanthopanax evodiaefolius Araliaceae PD Char. sp. of the Prismatomerio
var. pseudoevodiaefolius henryi—Lithocarpetum naiadar
Acanthopanax fargersii Araliaceae NPD Diff. sp. of the Sinarundinario nitidae-
Fagetum lucidae (subass.)
Acanthopanax giraldii Araliaceae NPD Char. sp. of the Athyrio nardii-
Michelietum balansae
(continued)
T. Hukusima et al., Phytosociology of the Beech (Fagus) Forests in East Asia, 169
Geobotany Studies, DOI 10.1007/978-3-642-35620-9,
# Springer-Verlag Berlin Heidelberg 2013
170 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Acanthopanax henryi Araliaceae NPD Companion species with sporadical
presence (mainly China)
Acanthopanax hypoleucus Araliaceae NPD Diff.-sp. of Sapio japonici-Fagetum
crenatae subass.)
Acanthopanax Araliaceae PD Char.-sp. of Saso-Fagetalia
sciadophylloides
Acanthopanax simonii Araliaceae NPD Companion species with sporadical
presence (mainly China)
Acanthopanax sp. Araliaceae NPD Companion species with sporadical
presence (mainly China)
Acanthopanax spinosus Araliaceae NPD Companion species with sporadical
presence (mainly Japan)
Acanthopanax trichodon Araliaceae NPD Companion species with sporadical
presence (mainly Japan)
Acer amplum Aceraceae PD Companion species with sporadical
presence (mainly China)
Acer argutum Aceraceae PD Companion species with sporadical
presence (mainly Japan)
Acer capillipes Aceraceae PD Diff.-sp. of Sasamorpho_Fagion
crenatae (subass.)
Acer carpinifolium Aceraceae PD Species of the Fraxino-Ulmetalia
Acer crataegifolium Aceraceae PD Species of the Quercetalia serratae-
grosseserratae
Acer davidii Aceraceae PD Char. sp. of the Aceri davidii-Fagion
lucidae
Acer diabolicum Aceraceae PD Diff.-sp. of Crnoo-Fagetum crenatae
(subass.)
Acer distylum Aceraceae PD Species of the Quercetalia serratae-
grosseserratae
Acer elegantulum Aceraceae PD Char. sp. of the Indocalamo latifolii—
Fagion hayatae var. zhejiangensis
Acer erianthum Aceraceae PD Companion species with sporadical
presence (mainly China)
Acer flabellatum Aceraceae PD Char. sp. of the Aceri davidii-Fagion
lucidae
Acer franchetii Aceraceae PD Diff. sp. of the Fagetum engleriano-
lucidae
Acer ginnola Aceraceae PD Char.-sp. of Euonymo porphyrei-
Fagetum englerianae
Acer grisema Aceraceae PD Companion species with sporadical
presence (mainly China)
Acer japonicum Aceraceae PD Char.-sp. of Saso-Fagetalia
Acer kawakamii Aceraceae PD Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Acer laxiflorum Aceraceae PD Char.-sp. of Abelio englerianae-
Fagion sp. div.
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 171
Life
Species name Family name form Ecological indication
Acer micranthum Aceraceae PD Char.-sp. of Saso-Fagetalia
Acer mono Aceraceae PD Widespead in the Fagus-forests of
East Asia
Acer mono f. dissectum Aceraceae PD Companion-species
Acer mono var. ambiguum Aceraceae PD Companion species with sporadical
presence (mainly Japan)
Acer mono var. connivens Aceraceae PD Companion species with sporadical
presence (mainly Japan)
Acer mono var. glabrum Aceraceae PD Companion species with sporadical
presence (mainly Japan)
Acer mono var. mayrii Aceraceae PD Char.-sp. of Saso kurilensis_Fagetum
crenatae
Acer mono var. savatieri Aceraceae PD Companion species with sporadical
presence (mainly Japan)
Acer nikoense Aceraceae PD Diff.-sp. of Sasamorpho_Fagion
crenatae (subass.)
Acer nipponicum Aceraceae PD Companion species with sporadical
presence (mainly Japan)
Acer okamotoana Aceraceae PD Char.-sp. of Fagion multinrvis and
Fagetalia multinervis
Acer oliverianum Aceraceae PD Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Acer palmatum Aceraceae PD Widespead in the Fagus-forests of
East Asia
Acer palmatum var. Aceraceae PD Char.-sp. of Sasamorpho_Fagion
amoenum crenatae
Acer palmatum var. Aceraceae PD Char.-sp. of Fagion crenatae
matsumurae
Acer prolificum Aceraceae PD Char. sp. of the Sinocalamus
giganteus-Fagetum lucidae
Acer pseudosieboldianum Aceraceae PD Char.-sp. of Hepatico-Fagetum
multinervis
Acer rufinerve Aceraceae PD Char.-sp. of Saso-Fagetalia
Acer shensiense Aceraceae PD Companion species with sporadical
presence (mainly China)
Acer shirasawanum Aceraceae PD Char.-sp. of Saso-Fagetalia
Acer sieboldianum Aceraceae PD Char.-sp. of Saso-Fagetalia
Acer sinense Aceraceae PD Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Acer sp. Aceraceae PD Companion species with sporadical
presence (mainly China)
Acer takesimense Aceraceae PD Char.-sp. of Fagion multinrvis and
Fagetalia multinervis
Acer tenuifolium Aceraceae PD Char.-sp. of Sasamorpho_Fagion
crenatae
(continued)
172 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Acer tschonoskii Aceraceae PD Char.-sp. of Saso kurilensis_Fagetum
crenatae
Acer ukurunduense Aceraceae PD Companion species with sporadical
presence (mainly Japan)
Acer wilsonii Aceraceae PD Char. sp. of the Ardisio hypargeriae—
Castanopsion fabrii
Achyranthes japonica Amaranthaceae H Companion species with sporadical
scap presence (mainly Japan)
Aconitum japonicum Ranunculaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Aconitum japonicum var. Ranunculaceae G rhiz Diff.-sp. of Crnoo-Fagetum crenatae
montanum (subass.)
Aconitum senanense Ranunculaceae NPD Companion-species
Aconitum sinomontanuns Ranunculaceae G rad Companion species with sporadical
presence (mainly China)
Acrophorus stipellatus Polypodiaceae G rhiz Char. sp. of the Yushania-Fagetum
hayatae
Actaea asiatica Ranunculaceae G rad Diff.-sp. of Sapio japonici-Fagetum
crenatae subass.)
Actinidia arguta Actinidiaceae PL Companion-species
Actinidia chinensis Actinidiaceae PL Companion-species
Actinidia coriacea Actinidiaceae PL Companion species with sporadical
presence (mainly China)
Actinidia kolomikta Actinidiaceae PL Widespead in the Fagus-forests of
East Asia
Actinidia polygama Actinidiaceae PL Companion species with sporadical
presence (mainly Japan)
Actinidia sp. Actinidiaceae PL Companion species with sporadical
presence (mainly China)
Actinidia vitifolia Actinidiaceae PL Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Actinodaphne reticulata Lauraceae PE Diff. sp. of the Sinarundinario
chungii-Fagetum lucidae (subass.)
Adenophora cordifolia Campanulaceae H Diff. sp. of the Fagetum engleriano-
scap lucidae
Adenophora hunanensis Campanulaceae H Char. sp. of the Fagetum engleriano-
scap lucidae
Adenophora polymorpha Campanulaceae H Diff. sp. of the Fagetum engleriano-
scap lucidae
Adenophora remotiflora Campanulaceae H Diff.-sp. of Saso kurilensis-Fagetum
scap crenatae
Adenophora trachelioides Campanulaceae H Diff. sp. of the Fagetum engleriano-
scap lucidae
Adiantum pedatum Polypodiaceae G rhiz Diff.-sp. of Hepatico-Fagetum
multinervis (subass.)
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 173
Life
Species name Family name form Ecological indication
Adina pilulifera Rubiaceae H Companion species with sporadical
scap presence (mainly China)
Adinandra wangii Theaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Aegopodium alpestre Apiaceae H Diff.-sp. of Saso kurilensis-Fagetum
scap crenatae
Aesculus turbinata Hippocastanaceae PD Species of the Fraxino-Ulmetalia
Aesculus wilsonii Hippocastanaceae PD Char. sp. of the Elatostemo
sessile_Fagetum lucidae
Agrostis scouleri Poaceae T Companion species with sporadical
presence (mainly Japan)
Ainsliaea acerifolia Asteraceae H Companion species with sporadical
scap presence (mainly Japan)
Ainsliaea acerifolia var. Asteraceae H Species of the Quercetalia serratae-
subapoda scap grosseserratae
Ainsliaea apiculata Asteraceae H Diff.-sp. of Sasamorpho_Fagion
scap crenatae (subass.)
Ainsliaea gracilis Asteraceae H Diff. sp. of the Sinarundinario
scap chungii-Fagetum lucidae (subass.)
Ainsliaea grossedentata Asteraceae H Companion species with sporadical
rosul presence (mainly China)
Ainsliaea henryi Asteraceae H Char. sp. of the Aceri davidii-Fagion
scap lucidae
Ainsliaea macroclinidioides Asteraceae H Char. sp. of the Indocalamo latifolii—
scap Fagion hayatae var. zhejiangensis
Ainsliaea rubrinervis Asteraceae H Companion species with sporadical
rosul presence (mainly China)
Ainsliaea triflora Asteraceae H Char.-sp. of Abelio englerianae-
rosul Fagion sp. div.
Ainsliaea yunnanensis Asteraceae H Companion species with sporadical
rosul presence (mainly China)
Ajuga nipponensis Lamiaceae H Companion species with sporadical
scap presence (mainly China)
Akebia trifoliata Lardizabalaceae PL Widespead in the Fagus-forests of
East Asia
Akebia trifoliata var. Lardizabalaceae PL Companion species with sporadical
australis presence (mainly China)
Alangium chinense Alangiaceae NPD Companion species with sporadical
presence (mainly China)
Alangium platanifolium var. Alangiaceae NPD Diff.-sp. of Sasamorpho_Fagion
trilobum crenatae (subass.)
Albizia kalkora Fabaceae PD Char. sp. of the Indocalamo latifolii—
Fagion hayatae var. zhejiangensis
Albizia turgida Fabaceae PD Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
(continued)
174 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Allantodia chinensis Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Allantodia hirtipes Polypodiaceae G rhiz Char. sp. of the Quingzheo
tumidinodae-Fagion
Allantodia metteniana Polypodiaceae G rhiz Diff. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
(subass.)
Allantodia petri Polypodiaceae G rhiz Char. sp. of the Athyrio nardii-
Michelietum balansae
Allantodia squamigera Polypodiaceae G rhiz Char. sp. of the Sinocalamus
giganteus-Fagetum lucidae
Allantodia wichurae Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Allium victorialis var. Liliaceae G Char.-sp. of Hepatico-Fagetum
platyphyllum bulb multinervis
Alnus firma Betulaceae PD Companion-species
Alnus hirsuta Betulaceae PD Companion species with sporadical
presence (mainly Japan)
Alnus hirsuta var.sibirica Betulaceae PD Companion species with sporadical
presence (mainly Japan)
Alnus maximowiczii Betulaceae PD Char.-sp. of Hepatico-Fagetum
multinervis
Alpinia chinensis Zingiberaceae G rhiz Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Alpinia sp. Zingiberaceae G rhiz Companion species with sporadical
presence (mainly China)
Ampelopsis Vitaceae PL Widespead in the Fagus-forests of
brevipedunculata East Asia
Ampelopsis delavayana Vitaceae PL Companion species with sporadical
presence (mainly China)
Anaphalis margaritacea Asteraceae H Diff. sp. of the Fagetum engleriano-
scap lucidae
Anemone debilis Ranunculaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Anemone nikoensis Ranunculaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Anemone pesudo-altaica Ranunculaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Anemonopsis macrophylla Ranunculaceae G rad Diff.-sp. of Crnoo-Fagetum crenatae
(subass.)
Angelica edulis Apiaceae H Companion species with sporadical
scap presence (mainly Japan)
Angelica polymorha Apiaceae H Companion species with sporadical
scap presence (mainly Japan)
Angelica pubescens Apiaceae H Diff.-sp. of Sapio japonici-Fagetum
scap crenatae subass.)
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 175
Life
Species name Family name form Ecological indication
Arachniodes chinensis Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Arachniodes festina Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Arachniodes mutica Polypodiaceae H Char.-sp. of Fagion crenatae
rosul
Arachniodes pseudo- Polypodiaceae G rhiz Char. sp. of the Sinocalamus
aristata giganteus-Fagetum lucidae
Arachniodes rhomboides Polypodiaceae G rhiz Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Arachniodes standishii Polypodiaceae H Species of the Fraxino-Ulmetalia
rosul
Araiostegia parripinnata Polypodiaceae G rhiz Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Aralia chinensis Araliaceae NPD Companion species with sporadical
presence (mainly China)
Arctous alpinas Ericaceae Ch Companion species with sporadical
frut presence (mainly China)
Ardisia affinis Myrsinaceae NPE Char. sp. of the Sinarundinario
bashersuto-Fagetum lucidae
Ardisia brevicaulis Myrsinaceae NPE Char. sp. of the Yushania-Fagetum
hayatae
Ardisia crenata Myrsinaceae NPE Widespead in the Fagus-forests of
East Asia
Ardisia crispa Myrsinaceae NPE Char.-sp. of Litseo elongatae-Fagetea
Ardisia hypargyera Myrsinaceae NPE Char. sp. of the Ardisio hypargeriae—
Castanopsion fabrii
Ardisia japonica Myrsinaceae NPE Widespead in the Fagus-forests of
East Asia
Ardisia sp. Myrsinaceae NPE Companion species with sporadical
presence (mainly China)
Arisaema amurense Araceae G Char.-sp. of Fagion multinrvis and
bulb Fagetalia multinervis
Arisaema angustatum Araceae G Diff.-sp. of Crnoo-Fagetum crenatae
bulb (subass.)
Arisaema consanguineum Araceae G Diff. sp. of the Sinarundinario
bulb chungii-Fagetum lucidae (subass.)
Arisaema erubescens Araceae G Companion species with sporadical
bulb presence (mainly China)
Arisaema formosanum Araceae G Companion species with sporadical
bulb presence (mainly China)
Arisaema japonicum Araceae G Companion species with sporadical
bulb presence (mainly Japan)
Arisaema limbatum Araceae G Companion species with sporadical
bulb presence (mainly Japan)
(continued)
176 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Arisaema peninsulae Araceae G Companion species with sporadical
bulb presence (mainly Japan)
Arisaema ssp. Araceae G Companion species with sporadical
bulb presence (mainly China)
Arisaema ternatipartitum Araceae G Diff.-sp. of Sapio japonici-Fagetum
bulb crenatae subass.)
Aristolochia kaempferi Aristolochiaceae PL Companion species with sporadical
presence (mainly Japan)
Aristolochia mollissima Aristolochiaceae PL Diff. sp. of the Sinarundinario nitidae-
Fagetum lucidae (subass.)
Aristolochia tubiflora Aristolochiaceae PL Companion species with sporadical
presence (mainly China)
Arthraxon hispidus Poaceae T Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Arthromeris lehmannii Polypodiaceae H Diff. sp. of the Sinarundinario
caesp bashersuto-Fagetum lucidae (subass.)
Aruncus dioicus Rosaceae H Char.-sp. of Hepatico-Fagetum
scap multinervis
Aruncus dioicus var. Rosaceae H Diff.-sp. of Saso kurilensis-Fagetum
tenuifolius scap crenatae
Aruncus sylvester Rosaceae H Companion species with sporadical
scap presence (mainly China)
Asarum caudigerum Aristolochiaceae G rad Companion species with sporadical
presence (mainly China)
Asarum chinensis Aristolochiaceae G rad Companion species with sporadical
presence (mainly China)
Asarum heterotropoides Aristolochiaceae G rad Companion species with sporadical
presence (mainly Japan)
Asarum ichangense Aristolochiaceae G rad Diff. sp. of the Sinarundinario nitidae-
Fagetum lucidae (subass.)
Asarum savatieri Aristolochiaceae G rad Companion species with sporadical
presence (mainly Japan)
Asarum sieboldii Aristolochiaceae G rad Widespead in the Fagus-forests of
East Asia
Asarum spp. Aristolochiaceae G rad Companion species with sporadical
presence (mainly China)
Asparagus filicinus Liliaceae G rhiz Companion species with sporadical
presence (mainly China)
Asperula odorata Rubiaceae H Char.-sp. of Fagion multinrvis and
scap Fagetalia multinervis
Asplenium crinicaule Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Asplenium incisum Polypodiaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Asplenium normale Polypodiaceae G rhiz Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 177
Life
Species name Family name form Ecological indication
Asplenium unilaterale Polypodiaceae G rhiz Char. sp. of the Athyrio nardii-
Michelietum balansae
Asplenium wrightii Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Aster ageratoides Asteraceae H Char. sp. of the Fagetum engleriano-
scap lucidae
Aster ageratoides ssp. Asteraceae H Diff.-sp. of Crnoo-Fagetum crenatae
amplexifolius scap (subass.)
Aster ageratoides var. harae Asteraceae H Char.-sp. of Corno-Fagetum crenatae
f. leucanthus scap
Aster dimorphophyllus Asteraceae H Char.-sp. of Corno-Fagetum crenatae
scap
Aster glehnii Asteraceae H Char.-sp. of Fagion multinrvis and
scap Fagetalia multinervis
Aster procerus Asteraceae H Char. sp. of the Carici lanceolatae-
scap Fagetum hayatae var. zhejiangensis
Aster scaber Asteraceae H Companion species with sporadical
scap presence (mainly Japan)
Asteropyrum cavaleriei Ranunculaceae G rhiz Companion species with sporadical
presence (mainly China)
Asteropyrum peltatum Ranunculaceae G rhiz Companion species with sporadical
presence (mainly China)
Astilbe koreana Saxifragaceae G rad Diff.-sp. of Hepatico-Fagetum
multinervis (subass.)
Astilbe rubra Saxifragaceae G rad Diff. sp. of the Fagetum engleriano-
lucidae
Astilbe thunbergii Saxifragaceae G rad Companion-species
Astilbe thunbergii var. Saxifragaceae G rad Companion-species
congesta
Astilbe thunbergii var. Saxifragaceae G rad Companion species with sporadical
fujisanensis presence (mainly Japan)
Astilbe thunbergii var. Saxifragaceae G rad Companion species with sporadical
shikokiana presence (mainly Japan)
Athyrium arisaense Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Athyrium brevifrons Polypodiaceae G rhiz Char.-sp. of Fagion multinrvis and
Fagetalia multinervis
Athyrium clivicola Polypodiaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Athyrium delavayi Polypodiaceae G rhiz Char. sp. of the Viburno flavescentis-
Fagetum englerianae
Athyrium epirachis Polypodiaceae G rhiz Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Athyrium henryi Polypodiaceae G rhiz Companion species with sporadical
presence (mainly Japan)
(continued)
178 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Athyrium malipoense Polypodiaceae G rhiz Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Athyrium nardii Polypodiaceae G rhiz Char. sp. of the Athyrio nardii-
Michelietum balansae
Athyrium niponicum Polypodiaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Athyrium otophorum Polypodiaceae G rhiz Widespead in the Fagus-forests of
East Asia
Athyrium sp. Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Athyrium strigillosum Polypodiaceae G rhiz Char. sp. of the Sinarundinario
chungii-Fagetum lucidae
Athyrium wardii Polypodiaceae G rhiz Widespead in the Fagus-forests of
East Asia
Athyrium yokoscense Polypodiaceae G rhiz Companion-species
Aucuba chinensis Cornaceae NPE Companion-species
Aucuba japonica Cornaceae NPE Diff.-sp. of Sasamorpho_Fagion
crenatae (subass.)
Aucuba japonica var. Cornaceae NPE Char.-sp. of Fagion crenatae
borealis
Aucuba obcordata Cornaceae NPE Char. sp. of the Sinarundinario
chungii-Fagetum lucidae
Barthea formosana Melastomataceae NPD Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Beccarinda tonkinensis Gesneriaceae H Diff. sp. of the Prismatomerio
scap henryi—Lithocarpetum naiadari
(subass.)
Begonia sp. Begoniaceae H Companion species with sporadical
scap presence (mainly China)
Beilschmiedia robusta Lauraceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Berberis aristato-serrulata Berberidaceae NPD Companion species with sporadical
presence (mainly China)
Berberis dasytachya Berberidaceae NPD Companion species with sporadical
presence (mainly China)
Berberis dielsiana Berberidaceae NPD Char.-sp. of Abelio englerianae-
Fagion sp. div.
Berberis jalianae Berberidaceae NPD Companion-species
Berberis mingetsuensys Berberidaceae NPD Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Berberis thunbergii Berberidaceae PL Companion species with sporadical
presence (mainly Japan)
Berchemia kulingensis Rhamnaceae PL Companion species with sporadical
presence (mainly China)
Berchemia racemosa Rhamnaceae PL Companion species with sporadical
presence (mainly Japan)
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 179
Life
Species name Family name form Ecological indication
Betula albo-sinensis Betulaceae PD Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Betula chinensis Betulaceae PD Companion species with sporadical
presence (mainly China)
Betula corylifolia Betulaceae PD Companion species with sporadical
presence (mainly Japan)
Betula ermanii Betulaceae PD Diff.-sp. of Saso kurilensis-Fagetum
crenatae
Betula grossa Betulaceae PD Char.-sp. of Saso-Fagetalia
Betula insignis Betulaceae PD Diff. sp. of the Sinarundinario nitidae-
Fagetum lucidae (subass.)
Betula luminifera Betulaceae PD Companion-species
Betula maximowicziana Betulaceae PD Companion species with sporadical
presence (mainly Japan)
Betula schmidtii Betulaceae PD Companion species with sporadical
presence (mainly Japan)
Betula sp. Betulaceae PD Companion species with sporadical
presence (mainly China)
Blastus pauciflorus Melastomataceae NPD Diff. sp. of the Sinarundinario
bashersuto-Fagetum lucidae (subass.)
Bletilla striata Orchidaceae G rhiz Char. sp. of the Elatostemo
sessile_Fagetum lucidae
Boehmeria spicata Urticaceae Ch Companion species with sporadical
frut presence (mainly Japan)
Boehmeria tricuspis Urticaceae Ch Companion species with sporadical
frut presence (mainly Japan)
Botrychium multifidum var. Ophioglossaceae H Char.-sp. of Hepatico-Fagetum
robustum caesp multinervis
Brachypodium sylvaticum Poaceae H Companion species with sporadical
caesp presence (mainly Japan)
Bredia amoena Melastomataceae NPE Char. sp. of the Carici lanceolatae-
Fagetum hayatae
Bretschneidera sinensis Bretschneideraceae PE Companion species with sporadical
presence (mainly China)
Broussonetia karinoki Moraceae PL Companion species with sporadical
presence (mainly China)
Brylkinia caudata Poaceae H Diff.-sp. of Sapio japonici-Fagetum
caesp crenatae subass.)
Buckleya henryi Santalaceae NPD Diff. sp. of the Fagetum engleriano-
lucidae
Buckleya lanceolata Santalaceae NPD Species of the Quercetalia serratae-
grosseserratae
Buxus henryi Buxaceae PE Companion species with sporadical
presence (mainly China)
Cacalia adenostyloides Asteraceae H Companion-species
scap
(continued)
180 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Cacalia ainsliaeflora Asteraceae H Companion species with sporadical
scap presence (mainly China)
Cacalia delphiniifolia Asteraceae H Companion-species
scap
Cacalia farfaraefolia var. Asteraceae H Companion species with sporadical
bulbifera scap presence (mainly Japan)
Cacalia hastata var. Asteraceae H Diff.-sp. of Crnoo-Fagetum crenatae
farfaraefolia scap (subass.)
Cacalia hastata var. Asteraceae H Diff.-sp. of Saso kurilensis-Fagetum
tanakae scap crenatae
Cacalia nikomontana Asteraceae H Companion species with sporadical
scap presence (mainly Japan)
Cacalia profundorum Asteraceae H Companion species with sporadical
scap presence (mainly China)
Cacalia roborowskii Asteraceae H Diff.-sp. of Euonymo porphyrei-
scap Fagetum englerianae (subass.)
Cacalia sp. Asteraceae H Companion species with sporadical
scap presence (mainly China)
Cacalia tebakoensis Asteraceae H Diff.-sp. of Sapio japonici-Fagetum
scap crenatae subass.)
Cacalia yatabei Asteraceae H Diff.-sp. of Crnoo-Fagetum crenatae
scap (subass.)
Cacalia yatabei var. Asteraceae H Diff.-sp. of Sapio japonici-Fagetum
occidentalis scap crenatae subass.)
Calamagrostis arundinacea Poaceae G rhiz Char. sp. of the Carici lanceolatae-
var. ciliata Fagetum hayatae
Calamagrostis hakonensis Poaceae G rhiz Diff.-sp. of Crnoo-Fagetum crenatae
(subass.)
Calamagrostis sylvatica Poaceae G rhiz Diff. sp. of the Fagetum engleriano-
lucidae
Calamus oxycarpus Arecaceae PL Char. sp. of the Athyrio nardii-
Michelietum balansae
Calamus sp. Arecaceae PL Companion species with sporadical
presence (mainly China)
Calanthe discolor Orchidaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Calanthe fimbriata Orchidaceae G rhiz Char.-sp. of Abelio englerianae-
Fagion sp. div.
Calanthe sp. Orchidaceae G rhiz Companion species with sporadical
presence (mainly China)
Calanthe tricarinata Orchidaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Callicarpa brevipes Verbenaceae NPD Char. sp. of the Sinarundinario
bashersuto-Fagetum lucidae
Callicarpa cathayana Verbenaceae NPD Companion-species
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 181
Life
Species name Family name form Ecological indication
Callicarpa giraldii Verbenaceae NPD Char. sp. of the Carici lanceolatae-
Fagetum hayatae var. zhejiangensis
Callicarpa japonica Verbenaceae NPD Species of the Quercetalia serratae-
grosseserratae
Callicarpa japonica var. Verbenaceae NPD Companion species with sporadical
angustata presence (mainly China)
Callicarpa mollis Verbenaceae NPD Companion species with sporadical
presence (mainly Japan)
Camellia brevistyla Theaceae PE Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Camellia caudata Theaceae PE Companion species with sporadical
presence (mainly China)
Camellia cuspidata Theaceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Camellia forrestii Theaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Camellia grijsii Theaceae PE Char. sp. of the Quingzheo
tumidinodae-Fagion
Camellia japonica Theaceae PE Diff.-sp. of Hepatico-Fagetum
multinervis (subass.)
Camellia japonica var. Theaceae NPE Diff.-sp. of Lindero umbellatae-
decumbens Fagetum crenatae
Camellia pitardii Theaceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Camellia rosthorniana Theaceae NPE Char. sp. of the Sinarundinario
chungii-Fagetum lucidae
Camellia sp. Theaceae PE Companion species with sporadical
presence (mainly China)
Camellia tenuifolia Theaceae PE Char. sp. of the Yushania-Fagetum
hayatae
Canthium simile Rubiaceae NPE Char. sp. of the Athyrio nardii-
Michelietum balansae
Cardiandra alternifolia Saxifragaceae H Companion species with sporadical
scap presence (mainly Japan)
Carex baccans Cyperaceae H Companion species with sporadical
caesp presence (mainly China)
Carex blepharicarpa Cyperaceae H Companion species with sporadical
caesp presence (mainly Japan)
Carex breviculmis Cyperaceae H Companion species with sporadical
caesp presence (mainly Japan)
Carex brunnea Cyperaceae H Companion species with sporadical
caesp presence (mainly China)
Carex capilliformis Cyperaceae H Char. sp. of the Fagetum engleriano-
caesp lucidae
Carex chinensis Cyperaceae H Char. sp. of the Carici lanceolatae-
caesp Fagetum hayatae
(continued)
182 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Carex conica Cyperaceae H Companion-species
caesp
Carex cruciata Cyperaceae H Companion-species
caesp
Carex dolichostachya var. Cyperaceae H Char.-sp. of Saso-Fagetalia
glaberrima caesp
Carex fernaldiana Cyperaceae H Char.-sp. of Sasamorpho_Fagion
caesp crenatae
Carex filicina Cyperaceae H Char. sp. of the Aceri davidii-Fagion
caesp lucidae
Carex filipes Cyperaceae H Diff.-sp. of Sapio japonici-Fagetum
caesp crenatae subass.)
Carex floribunda Cyperaceae H Companion species with sporadical
caesp presence (mainly Japan)
Carex foliosissima Cyperaceae H Char.-sp. of Fagion crenatae
caesp
Carex grallatoria Cyperaceae H Diff.-sp. of Sapio japonici-Fagetum
caesp crenatae subass.)
Carex grandiligulata Cyperaceae H Companion species with sporadical
caesp presence (mainly China)
Carex henryi Cyperaceae H Char. sp. of the Sinarundinario
caesp chungii-Fagetum lucidae
Carex insaniae Cyperaceae H Companion species with sporadical
caesp presence (mainly Japan)
Carex lanceolata Cyperaceae H Char. sp. of the Carici lanceolatae-
caesp Fagetum hayatae var. zhejiangensis
Carex lasiolepis Cyperaceae H Companion species with sporadical
caesp presence (mainly Japan)
Carex morrowii Cyperaceae H Companion-species
caesp
Carex morrowii var. Cyperaceae H Companion-species
temnolepis caesp
Carex omeiensis Cyperaceae H Char. sp. of the Sinarundinario
caesp chungii-Fagetum lucidae
Carex pachyrrhiza Cyperaceae H Diff. sp. of the Sinarundinario
caesp chungii-Fagetum lucidae (subass.)
Carex perakensis Cyperaceae H Char. sp. of the Athyrio nardii-
caesp Michelietum balansae
Carex pilosa Cyperaceae H Diff.-sp. of Saso kurilensis-Fagetum
caesp crenatae
Carex reinii Cyperaceae H Companion-species
caesp
Carex sachalinensis Cyperaceae H Companion-species
caesp
Carex sachalinensis var. Cyperaceae H Diff.-sp. of Saso kurilensis-Fagetum
alterniflora caesp crenatae
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 183
Life
Species name Family name form Ecological indication
Carex sachalinensis var. Cyperaceae H Diff.-sp. of Sapio japonici-Fagetum
sikokiana caesp crenatae subass.)
Carex sendaica Cyperaceae H Char. sp. of the Fagetum engleriano-
caesp lucidae
Carex siderosticta Cyperaceae G rhiz Widespead in the Fagus-forests of
East Asia
Carex sp.1 Cyperaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Carex sp.2 Cyperaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Carex stenostachys var. Cyperaceae H Companion species with sporadical
cuneata caesp presence (mainly Japan)
Carex subpediformis Cyperaceae H Diff. sp. of the Fagetum engleriano-
caesp lucidae
Carex sutchanensis Cyperaceae H Companion species with sporadical
caesp presence (mainly China)
Carpesium triste Asteraceae H Companion species with sporadical
scap presence (mainly Japan)
Carpinus cordata Betulaceae PD Widespead in the Fagus-forests of
East Asia
Carpinus cordata var. Betulaceae PD Diff. sp. of the Fagetum engleriano-
chinensis lucidae
Carpinus fangiana Betulaceae PD Char. sp. of the Sinocalamus
giganteus-Fagetum lucidae
Carpinus fargesii Betulaceae PD Companion species with sporadical
presence (mainly China)
Carpinus japonica Betulaceae PD Char.-sp. of Saso-Fagetalia
Carpinus laxiflora Betulaceae PD Species of the Quercetalia serratae-
grosseserratae
Carpinus polyneura Betulaceae PD Diff.-sp. of Euonymo porphyrei-
Fagetum englerianae (subass.)
Carpinus pubescens Betulaceae PD Char. sp. of the Sinarundinario
chungii-Fagetum lucidae
Carpinus tschonoskii Betulaceae PD Widespead in the Fagus-forests of
East Asia
Carpinus turczaninowii Betulaceae PD Diff. sp. of the Fagetum engleriano-
lucidae
Carpinus viminea Betulaceae PD Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Castanea crenata Fagaceae PD Species of the Quercetalia serratae-
grosseserratae
Castanea henryi Fagaceae PD Diff. sp. of the Fagetum engleriano-
lucidae
Castanopsis calathiformis Fagaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
(continued)
184 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Castanopsis carlesii Fagaceae PE Char. sp. of the Polypodio argutum-
Fagetum longepetiolatae
Castanopsis chunii Fagaceae PE Companion species with sporadical
presence (mainly China)
Castanopsis eryei Fagaceae PE Char. sp. of the Sinarundinario
bashersuto-Fagetum lucidae
Castanopsis fabrii Fagaceae PE Char. sp. of the Ardisio hypargeriae—
Castanopsion fabrii
Castanopsis hystrix Fagaceae PE Companion species with sporadical
presence (mainly China)
Castanopsis lamontii Fagaceae PE Companion species with sporadical
presence (mainly China)
Castanopsis pachyrachis Fagaceae PE Diff. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
(subass.)
Castanopsis rufotomentosa Fagaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Castanopsis umensis Fagaceae PE Companion species with sporadical
presence (mainly China)
Caulophyllum robustum Berberidaceae H Diff.-sp. of Sasamorpho_Fagion
scap crenatae (subass.)
Cayratia corniculata Vitaceae PL Diff. sp. of the Sinarundinario
bashersuto-Fagetum lucidae (subass.)
Cayratia japonica Vitaceae H Widespead in the Fagus-forests of
scand East Asia
Cayratia oligocarpa Vitaceae PL Companion species with sporadical
presence (mainly China)
Celastrus gemmatus Celastraceae PL Companion species with sporadical
presence (mainly China)
Celastrus orbiculatus Celastraceae PL Widespead in the Fagus-forests of
East Asia
Celastrus orbiculatus var. Celastraceae PL Companion species with sporadical
papillosus presence (mainly Japan)
Celastrus rosthornianus Celastraceae PL Char. sp. of the Sinarundinario
var. loeseneri chungii-Fagetum lucidae
Celastrus sp. Celastraceae PL Companion species with sporadical
presence (mainly China)
Celtis jessoensis Ulmaceae PD Diff.-sp. of Hepatico-Fagetum
multinervis (subass.)
Centella asiatica Apiaceae H Companion species with sporadical
scap presence (mainly China)
Cephalanthera erecta Orchidaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Cephalanthera falcata Orchidaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Cephalotaxus harringtonia Cephalotaxaceae PEN Companion-species
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 185
Life
Species name Family name form Ecological indication
Cephalotaxus harringtonia Cephalotaxaceae NPEN Char.-sp. of Fagion crenatae
var. nana
Cerasus conradinae Rosaceae PD Diff. sp. of the Fagetum engleriano-
lucidae
Cercidiphyllum japonicum Cercidiphyllaceae PD Companion species with sporadical
presence (mainly Japan)
Cercis chinensis Fabaceae NPD Char. sp. of the Elatostemo
sessile_Fagetum lucidae
Chamaecyparis obtusa Cupressaceae PEN Companion species with sporadical
presence (mainly Japan)
Chamaecyparis pisifera Cupressaceae PEN Companion species with sporadical
presence (mainly Japan)
Chamaele decumbens Apiaceae H Diff.-sp. of Crnoo-Fagetum crenatae
scap (subass.)
Chelonopsis moschata Lamiaceae H Companion species with sporadical
scap presence (mainly Japan)
Chimaphila japonica Pyrolaceae H Companion species with sporadical
scap presence (mainly Japan)
Chimonobambusa utilis Poaceae NPE Char. sp. of the Ardisio hypargeriae—
Castanopsion fabrii
Chirita sp. Gesneriaceae H Companion species with sporadical
scap presence (mainly China)
Chloranthus henryi Chloranthaceae H Companion species with sporadical
scap presence (mainly China)
Chloranthus serratus Chloranthaceae H Diff.-sp. of Sasamorpho_Fagion
scap crenatae (subass.)
Chrysosplenium grayanum Saxifragaceae H Companion species with sporadical
scap presence (mainly Japan)
Chrysosplenium henryi Saxifragaceae H Companion species with sporadical
scap presence (mainly China)
Chrysosplenium Saxifragaceae H Companion species with sporadical
lanuginosum scap presence (mainly China)
Cimicifuga acerina Ranunculaceae G rad Diff.-sp. of Crnoo-Fagetum crenatae
(subass.)
Cimicifuga simplex Ranunculaceae G rad Diff.-sp. of Saso kurilensis-Fagetum
crenatae
Cinnamomum bodinieri Lauraceae PE Char. sp. of the Sinarundinario
bashersuto-Fagetum lucidae
Cinnamomum burmannii Lauraceae PE Char. sp. of the Ardisio hypargeriae—
Castanopsion fabrii
Cinnamomum wilsonii Lauraceae PE Companion species with sporadical
presence (mainly China)
Circaea alpina Oenotheraceae H Companion species with sporadical
scap presence (mainly Japan)
Circaea erubescens Oenotheraceae H Diff.-sp. of Sasamorpho_Fagion
scap crenatae (subass.)
(continued)
186 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Circaea mollis Oenotheraceae H Companion species with sporadical
scap presence (mainly Japan)
Cirsium buergeri Asteraceae H Diff.-sp. of Sapio japonici-Fagetum
scap crenatae subass.)
Cirsium effusum Asteraceae H Diff.-sp. of Crnoo-Fagetum crenatae
scap (subass.)
Cirsium microspicatum Asteraceae H Diff.-sp. of Crnoo-Fagetum crenatae
scap (subass.)
Cirsium nipponicum var. Asteraceae H Diff.-sp. of Sapio japonici-Fagetum
shikokianum scap crenatae subass.)
Clematis finetiana Ranunculaceae PL Companion species with sporadical
presence (mainly China)
Clematis japonica Ranunculaceae PL Diff.-sp. of Sasamorpho_Fagion
crenatae (subass.)
Clematis mantana Ranunculaceae PL Companion species with sporadical
presence (mainly China)
Clematis otophora Ranunculaceae PL Companion species with sporadical
presence (mainly China)
Clematis sp. Ranunculaceae PL Companion species with sporadical
presence (mainly China)
Clematoclethra scandens Ochnaceae PL Companion species with sporadical
presence (mainly China)
Clerodendron Taccaceae G rhiz Diff. sp. of the Sinarundinario
colebrookianum bashersuto-Fagetum lucidae (subass.)
Clerodendron fortunatum Taccaceae G rhiz Diff. sp. of the Sinarundinario
bashersuto-Fagetum lucidae (subass.)
Clethra barbinervis Clethraceae PD Widespead in the Fagus-forests of
East Asia
Clethra fabri Clethraceae PD Char. sp. of the Sinarundinario
bashersuto-Fagetum lucidae
Clethra fargesii Clethraceae PD Companion-species
Cleyera incornuta Theaceae PE Companion species with sporadical
presence (mainly China)
Cleyera japonica Theaceae PE Widespead in the Fagus-forests of
East Asia
Clinopodium gracile var. Lamiaceae H Diff.-sp. of Crnoo-Fagetum crenatae
multicaule scap (subass.)
Clintonia udensis Liliaceae G rhiz Diff.-sp. of Saso kurilensis-Fagetum
crenatae
Conandron sp. Gesneriaceae H Companion species with sporadical
scap presence (mainly China)
Coptis quinquefolia Ranunculaceae G rhiz Char. sp. of the Yushania-Fagetum
hayatae
Coptis trifolia Ranunculaceae G rhiz Companion species with sporadical
presence (mainly Japan)
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 187
Life
Species name Family name form Ecological indication
Cornopteris decurrenti- Polypodiaceae G rhiz Diff.-sp. of Sapio japonici-Fagetum
alata crenatae subass.)
Cornus controversa Cornaceae PD Widespead in the Fagus-forests of
East Asia
Cornus kousa Cornaceae PD Widespead in the Fagus-forests of
East Asia
Cornus macrophylla Cornaceae PD Companion species with sporadical
presence (mainly Japan)
Cornus paucinervis Cornaceae PD Companion species with sporadical
presence (mainly China)
Cornus sp. Cornaceae PD Companion species with sporadical
presence (mainly China)
Corylopsis sinensis Hamamelidaceae PD Companion species with sporadical
presence (mainly China)
Corylus heterophylla Betulaceae PD Companion species with sporadical
presence (mainly China)
Corylus heterophylla var. Betulaceae PD Companion species with sporadical
sutchuenensis presence (mainly China)
Corylus sieboldiana Betulaceae NPD Char.-sp. of Saso-Fagetalia
Cremanthodium decaisnel Asteraceae H Companion species with sporadical
scap presence (mainly China)
Cremastra appendiculata Orchidaceae G rhiz Char. sp. of the Yushania-Fagetum
hayatae
Crypsinus echinosporus Polypodiaceae G rhiz Char. sp. of the Yushania-Fagetum
hayatae
Crypsinus quasioivaricata Polypodiaceae G rhiz Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Cryptomeria japonica Taxodiaceae PEN Char.-sp. of Lindero umbellatae-
Fagetum crenatae
Cylindrokelupha balansae Fabaceae H Char. sp. of the Athyrio nardii-
scap Michelietum balansae
Cymbidium faberi Orchidaceae G rhiz Companion species with sporadical
presence (mainly China)
Cymbidium goeringii Orchidaceae G rhiz Widespead in the Fagus-forests of
East Asia
Cymbidium sinense Orchidaceae G rhiz Char. sp. of the Polypodio argutum-
Fagetum longepetiolatae
Cynanchum chinense Apocynaceae H Companion species with sporadical
scap presence (mainly China)
Cynanchum julianae Apocynaceae H Companion species with sporadical
scap presence (mainly China)
Cynanchum sp. Apocynaceae H Companion species with sporadical
scap presence (mainly China)
Cynanchum sublanceolatum Apocynaceae H Companion species with sporadical
scand presence (mainly Japan)
(continued)
188 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Cypripedium japonica Orchidaceae G rhiz Companion species with sporadical
presence (mainly China)
Cyrtomium balansae Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Cyrtomium hookeranum Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Cyrtomium macrophyllum Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Cystopteris moupinensis Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Dactylicapnos sp. Papaveraceae H Companion species with sporadical
scap presence (mainly China)
Damnacanthus Rubiaceae NPE Char. sp. of the Yushania-Fagetum
angustifolius hayatae
Damnacanthus Rubiaceae NPE Diff. sp. of the Yushania-Fagetum
angustifolius var. hayatae (subass.)
stenophyllus
Damnacanthus indica Rubiaceae NPE Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Daphne miyabeana Thymelaeaceae NPE Companion species with sporadical
presence (mainly Japan)
Daphne papyracea Thymelaeaceae NPE Diff. sp. of the Sinarundinario
chungii-Fagetum lucidae (subass.)
Daphne sp. Thymelaeaceae NPE Companion species with sporadical
presence (mainly China)
Daphniphyllum Euphorbiaceae PE Char. sp. of Vaccinio henryi-Fagetum
angustifolium pashanicae
Daphniphyllum Euphorbiaceae PE Widespead in the Fagus-forests of
macropodum East Asia
Daphniphyllum Euphorbiaceae NPE Char.-sp. of Fagion crenatae
macropodum var. humile
Davallia mariesii Polypodiaceae E Companion species with sporadical
presence (mainly Japan)
Davidia involucrata Nyssaceae PE Companion species with sporadical
presence (mainly China)
Davidia involucrata var. Nyssaceae PE Char. sp. of the Viburno flavescentis-
vilmoriniana Fagetum englerianae
Deinanthe bifida Saxifragaceae H Diff.-sp. of Sapio japonici-Fagetum
scap crenatae subass.)
Dendrobenthamia japonica Cornaceae PD Companion-species
var. chinensis
Dendrobenthamia Cornaceae PD Char. sp. of the Viburno flavescentis-
melanotricha Fagetum englerianae
Dendropanax chevalieri Araliaceae PE Char.-sp. of Litseo elongatae-Fagetea
Dendropanax hainanensis Araliaceae PE Diff. sp. of the Sinarundinario
bashersuto-Fagetum lucidae (subass.)
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 189
Life
Species name Family name form Ecological indication
Dendropanax macrocarpus Araliaceae PE Char. sp. of the Athyrio nardii-
Michelietum balansae
Dennstaedtia scabra Polypodiaceae G rhiz Char.-sp. of Litseo elongatae-Fagetea
Deparia albosquamata Polypodiaceae G rhiz Diff.-sp. of Sapio japonici-Fagetum
crenatae subass.)
Deparia conilii Polypodiaceae G rhiz Species of the Fraxino-Ulmetalia
Deparia japonica Polypodiaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Deparia pycnosorum Polypodiaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Desmodium oxyphyllum Fabaceae H Companion-species
scap
Desmodium sp. Fabaceae H Companion species with sporadical
scap presence (mainly China)
Desmodium szechuenense Fabaceae H Companion species with sporadical
scap presence (mainly China)
Deutzia crenata Saxifragaceae NPD Companion-species
Deyeuxia arundinacea Arundinaceae H Companion species with sporadical
caesp presence (mainly China)
Deyeuxia sinelatior Arundinaceae H Diff. sp. of the Fagetum engleriano-
caesp lucidae
Diacalpe aspidioides Papaveraceae G rhiz Diff. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
(subass.)
Dichocarpum stoloniferum Ranunculaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Dichroa febrifuga Saxifragaceae NPD Char. sp. of the Viburno flavescentis-
Fagetum englerianae
Dictyocline griffithii Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Dioscorea bulbifera Dioscoreaceae G Char. sp. of the Indocalamo latifolii—
bulb Fagion hayatae var. zhejiangensis
Dioscorea althaeoides Dioscoreaceae G Companion species with sporadical
bulb presence (mainly China)
Dioscorea batatas Dioscoreaceae G Char. sp. of the Sinarundinario
bulb bashersuto-Fagetum lucidae
Dioscorea gracillima Dioscoreaceae G Companion species with sporadical
bulb presence (mainly Japan)
Dioscorea nipponica Dioscoreaceae G Companion species with sporadical
bulb presence (mainly Japan)
Dioscorea quinqueloba Dioscoreaceae G Diff.-sp. of Sasamorpho_Fagion
bulb crenatae (subass.)
Dioscorea septemloba Dioscoreaceae G Companion species with sporadical
bulb presence (mainly Japan)
Dioscorea sp. Dioscoreaceae G Companion species with sporadical
bulb presence (mainly China)
(continued)
190 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Dioscorea tokoro Dioscoreaceae G Companion species with sporadical
bulb presence (mainly Japan)
Diospyros lotus Ebenaceae G Companion species with sporadical
bulb presence (mainly China)
Diospyros morrisiana Ebenaceae NPD Companion species with sporadical
presence (mainly China)
Dipelta floribunda Caprifoliaceae NPD Companion species with sporadical
presence (mainly China)
Diphylleia grayi Berberidaceae H Companion species with sporadical
scap presence (mainly Japan)
Diplazium pinfaense Polypodiaceae G rhiz Char. sp. of the Carici lanceolatae-
Fagetum hayatae
Diplazium squamigerum Polypodiaceae G rhiz Species of the Fraxino-Ulmetalia
Diplopterygium glaucum Gleicheniaceae G rhiz Companion species with sporadical
presence (mainly China)
Diplopterygium Gleicheniaceae G rhiz Companion species with sporadical
laevissimum presence (mainly China)
Disporopsis pernyi Liliaceae G rhiz Char. sp. of the Elatostemo
sessile_Fagetum lucidae
Disporum cantoniense Liliaceae G rhiz Char. sp. of the Sinarundinario
bashersuto-Fagetum lucidae
Disporum sessile Liliaceae G rhiz Widespead in the Fagus-forests of
East Asia
Disporum smilacinum Liliaceae G rhiz Char.-sp. of Fagetea crenatae
Disporum sp. Liliaceae G rhiz Companion species with sporadical
presence (mainly China)
Distylium sp. Hamamelidaceae PE Companion species with sporadical
presence (mainly China)
Dryopteris bissetiana Polypodiaceae G rhiz Diff.-sp. of Sasamorpho_Fagion
crenatae (subass.)
Dryopteris championii Polypodiaceae G rhiz Companion-species
Dryopteris chinensis Polypodiaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Dryopteris crassirhizoma Polypodiaceae H Widespead in the Fagus-forests of
rosul East Asia
Dryopteris erythrosora Polypodiaceae G rhiz Widespead in the Fagus-forests of
East Asia
Dryopteris expansa Polypodiaceae G rhiz Companion-species
Dryopteris formosana Polypodiaceae G rhiz Char. sp. of the Yushania-Fagetum
hayatae
Dryopteris fuscipes Polypodiaceae G rhiz Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Dryopteris labordei Polypodiaceae G rhiz Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Dryopteris lacera Polypodiaceae H Companion species with sporadical
rosul presence (mainly Japan)
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 191
Life
Species name Family name form Ecological indication
Dryopteris livida Polypodiaceae G rhiz Char. sp. of the Athyrio nardii-
Michelietum balansae
Dryopteris mariformis Polypodiaceae G rhiz Diff. sp. of the Sinarundinario nitidae-
Fagetum lucidae (subass.)
Dryopteris maximowicziana Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Dryopteris monticola Polypodiaceae H Companion species with sporadical
rosul presence (mainly Japan)
Dryopteris polylepis Polypodiaceae H Diff.-sp. of Sasamorpho_Fagion
rosul crenatae (subass.)
Dryopteris sabaei Polypodiaceae H Char.-sp. of Saso-Fagetalia
rosul
Dryopteris saxifraga Polypodiaceae H Companion species with sporadical
rosul presence (mainly Japan)
Dryopteris sp1. Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Dryopteris sp2. Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Dryopteris subtriangularis Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Dumasia truncata Fabaceae H Companion species with sporadical
scand presence (mainly Japan)
Dystaenia takesimana Apiaceae H Char.-sp. of Fagion multinrvis and
scap Fagetalia multinervis
Elaeagnus lanceolata Elaegnaceae PL Companion-species
Elaeagnus macrophylla Elaegnaceae PL Companion species with sporadical
presence (mainly Japan)
Elaeagnus montana Elaegnaceae NPD Companion species with sporadical
presence (mainly Japan)
Elaeagnus montana var. Elaegnaceae NPD Companion species with sporadical
ovata presence (mainly Japan)
Elaeagnus pungens Elaegnaceae NPE Companion species with sporadical
presence (mainly China)
Elaeocarpus baceanus Elaeocarpaceae PE Diff. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
(subass.)
Elaeocarpus japonicus Elaeocarpaceae PE Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Elaeocarpus javanicus Elaeocarpaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Elatostema lineolatum Ulmaceae G rhiz Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Elatostema obtusum Ulmaceae G rhiz Companion species with sporadical
presence (mainly China)
Elatostema papillosum Ulmaceae G rhiz Char. sp. of the Ardisio hypargeriae—
Castanopsion fabrii
(continued)
192 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Elatostema sessile Ulmaceae G rhiz Char. sp. of the Elatostemo
sessile_Fagetum lucidae
Elatostema sp. Ulmaceae G rhiz Companion species with sporadical
presence (mainly China)
Elatostema stewardii Ulmaceae G rhiz Companion species with sporadical
presence (mainly China)
Elatostema trilobulatum Ulmaceae G rhiz Char. sp. of the Yushania-Fagetum
hayatae
Elatostema umbellatum Ulmaceae H Diff.-sp. of Crnoo-Fagetum crenatae
scap (subass.)
Elatostema umbellatum var. Ulmaceae H Diff.-sp. of Sapio japonici-Fagetum
majus scap crenatae subass.)
Embelia ribes var. Myrsinaceae NPE Companion species with sporadical
pachyphylla presence (mainly China)
Embelia sp. Myrsinaceae NPE Companion species with sporadical
presence (mainly China)
Enkianthus campanulatus Ericaceae NPD Char.-sp. of Corno-Fagetum crenatae
Enkianthus campanulatus Ericaceae NPD Companion species with sporadical
var. ikokianus presence (mainly Japan)
Enkianthus cernuus f. Ericaceae NPD Char.-sp. of Sapio japonici-Fagetum
rubens crenatae
Enkianthus chinensis Ericaceae NPD Companion-species
Enkianthus deflexus Ericaceae NPD Companion species with sporadical
presence (mainly China)
Enkianthus perlata Ericaceae NPD Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Enkianthus serrulatus Ericaceae NPD Char. sp. of the Aceri davidii-Fagion
lucidae
Enkianthus subsessilis Ericaceae NPD Diff.-sp. of Sasamorpho_Fagion
crenatae (subass.)
Ephippianthus schmidtii Orchidaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Epigaea asiatica Ericaceae H Companion species with sporadical
scand presence (mainly Japan)
Epimedium davidii Berberidaceae H Companion species with sporadical
scap presence (mainly China)
Epimedium grandiflorum Berberidaceae H Companion species with sporadical
var. thunberganum scap presence (mainly Japan)
Epimedium sagittatum Berberidaceae H Char.-sp. of Abelio englerianae-
scap Fagion sp. div.
Epimedium sempervirens Berberidaceae H Companion species with sporadical
scap presence (mainly Japan)
Epipactis papillosa Orchidaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Epipactis sp. Orchidaceae G rhiz Companion species with sporadical
presence (mainly China)
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 193
Life
Species name Family name form Ecological indication
Eragrostis ferruginea Poaceae H Companion species with sporadical
caesp presence (mainly China)
Eriobotrya bengalensis Rosaceae NPE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Erythronium japonicum Liliaceae G Companion species with sporadical
bulb presence (mainly Japan)
Erythroxylum kunthianum Erythroxylaceae PE Char. sp. of the Sinarundinario
bashersuto-Fagetum lucidae
Eulalia quadrinervis Poaceae H Companion species with sporadical
caesp presence (mainly China)
Euonymus alatus Celastraceae NPD Char. sp. of the Fagetum engleriano-
lucidae
Euonymus alatus f. ciliato- Celastraceae NPD Species of the Quercetalia serratae-
dentatus grosseserratae
Euonymus cornutus Celastraceae NPD Companion species with sporadical
presence (mainly China)
Euonymus forbesianus Celastraceae NPD Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Euonymus fortunei Celastraceae NPE Companion-species
Euonymus fortunei var. Celastraceae PL Widespead in the Fagus-forests of
radicans East Asia
Euonymus giraldii Celastraceae NPD Char.-sp. of Abelio englerianae-
Fagion sp. div.
Euonymus hederaceus Celastraceae NPD Companion species with sporadical
presence (mainly China)
Euonymus lanceolatus Celastraceae NPD Char.-sp. of Lindero umbellatae-
Fagetum crenatae
Euonymus leclerei Celastraceae NPD Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Euonymus macropterus Celastraceae NPD Companion-species
Euonymus melananthus Celastraceae NPD Species of the Fraxino-Ulmetalia
Euonymus mengtzeunus Celastraceae NPD Char. sp. of the Ardisio hypargeriae—
Castanopsion fabrii
Euonymus oxyphyllus Celastraceae NPD Species of the Quercetalia serratae-
grosseserratae
Euonymus planipes Celastraceae NPD Companion species with sporadical
presence (mainly Japan)
Euonymus pourphyreus Celastraceae NPD Char.-sp. of Euonymo porphyrei-
Fagetum englerianae
Euonymus praewarskii Celastraceae NPD Diff. sp. of the Fagetum engleriano-
lucidae
Euonymus sieboldianus Celastraceae NPD Species of the Quercetalia serratae-
grosseserratae
Euonymus sieboldianus var. Celastraceae NPD Companion species with sporadical
sanguineus presence (mainly Japan)
(continued)
194 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Euonymus sp1. Celastraceae NPD Companion species with sporadical
presence (mainly China)
Euonymus sp2. Celastraceae NPD Companion species with sporadical
presence (mainly China)
Euonymus spraguei Celastraceae NPD Char. sp. of the Yushania-Fagetum
hayatae
Eupatorium chinense var. Asteraceae H Companion species with sporadical
simplicifolium scap presence (mainly Japan)
Eupatorium japonicum Asteraceae H Companion-species
scap
Eupatorium shinense Asteraceae H Diff. sp. of the Sinarundinario
scap bashersuto-Fagetum lucidae (subass.)
Euphorbia hylonoma Euphorbiaceae H Companion species with sporadical
scap presence (mainly China)
Euphorbia sieboldiana Euphorbiaceae H Diff.-sp. of Sapio japonici-Fagetum
scap crenatae subass.)
Euptelea pleiospermum Trochodendraceae PD Companion species with sporadical
presence (mainly China)
Euptelea polyandra Trochodendraceae PD Companion species with sporadical
presence (mainly Japan)
Eurya alata Theaceae NPE Companion-species
Eurya brevistyla Theaceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Eurya crenatifolia Theaceae NPE Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Eurya distichophylla Theaceae NPE Companion species with sporadical
presence (mainly China)
Eurya graffi Theaceae NPE Char. sp. of the Sinocalamus
giganteus-Fagetum lucidae
Eurya handel-mazzettii Theaceae PE Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Eurya hebeclados Theaceae NPE Char. sp. of the Carici lanceolatae-
Fagetum hayatae var. zhejiangensis
Eurya japonica Theaceae PE Widespead in the Fagus-forests of
East Asia
Eurya leptophylla Theaceae NPE Char. sp. of the Yushania-Fagetum
hayatae
Eurya loquaiana Theaceae NPE Char. sp. of the Sinarundinario
bashersuto-Fagetum lucidae
Eurya muricata Theaceae NPE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Eurya nitida Theaceae NPE Companion-species
Eurya obtusifolia Theaceae NPE Companion-species
Eurya ribiginosa var. Theaceae NPE Char. sp. of the Indocalamo latifolii—
attenuata Fagion hayatae var. zhejiangensis
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 195
Life
Species name Family name form Ecological indication
Eurya semisenullata Theaceae NPE Char. sp. of the Sinocalamus
giganteus-Fagetum lucidae
Eurya sp1. Theaceae PE Companion species with sporadical
presence (mainly China)
Eurya sp2. Theaceae NPE Companion species with sporadical
presence (mainly China)
Eurya trichocarpa Theaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Euscaphis japonica Staphyleaceae PD Companion species with sporadical
presence (mainly China)
Eutrema japonica Brassicaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Evodia rutaecarpa var. Rutaceae G rhiz Char. sp. of the Tripterospermo
bodinieri cordifolium-Fagetum englerianae
Evodiopanax innovans Araliaceae PD Species of the Quercetalia serratae-
grosseserratae
Fagus crenata Fagaceae PD Char.-sp. of Saso-Fagetalia
Fagus engleriana Fagaceae PD Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Fagus hayatae Fagaceae PD Char. sp. of the Yushania-Fagetum
hayatae
Fagus hayatae subsp. Fagaceae PD Char. sp. of Vaccinio henryi-Fagetum
pashanica pashanicae
Fagus hayatae var. Fagaceae PD Char. sp. of the Indocalamo latifolii—
zhejiangensis Fagion hayatae var. zhejiangensis
Fagus japonica Fagaceae PD Species of the Quercetalia serratae-
grosseserratae
Fagus longipetiolata Fagaceae PD Char.-sp. of Litseo elongatae-Fagetea
Fagus lucida Fagaceae PD Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Fagus multinervis Fagaceae PD Char.-sp. of Hepatico-Fagetum
multinervis
Ficus harlandii Moraceae PD Companion species with sporadical
presence (mainly China)
Ficus sp. Moraceae PL Companion species with sporadical
presence (mainly China)
Filipendula multijuga Rosaceae H Diff.-sp. of Crnoo-Fagetum crenatae
scap (subass.)
Fissistigma Annonaceae PL Char. sp. of the Prismatomerio
acuminatissimum henryi—Lithocarpetum naiadari
Fordiophyton faberi Melastomataceae H Char. sp. of the Sinarundinario
scap bashersuto-Fagetum lucidae
Forsythia suspensa Oleaceae NPD Diff. sp. of the Fagetum engleriano-
lucidae
Fragaria orientalis Rosaceae H Companion species with sporadical
scap presence (mainly China)
(continued)
196 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Fraxinus apertisquamifera Oleaceae PD Companion species with sporadical
presence (mainly Japan)
Fraxinus chinensis Oleaceae PD Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Fraxinus lanuginosa Oleaceae PD Char.-sp. of Saso-Fagetalia
Fraxinus rhynchophylla Oleaceae PD Companion-species
Fraxinus sieboldiana Oleaceae PD Species of the Quercetalia serratae-
grosseserratae
Galium asperuloides Rubiaceae H Companion-species
scap
Galium asperuloides var. Rubiaceae H Char. sp. of the Sinocalamus
hoffmeisteris scap giganteus-Fagetum lucidae
Galium japonicum Rubiaceae H Diff.-sp. of Sasamorpho_Fagion
scap crenatae (subass.)
Galium kamtschaticum Rubiaceae H Companion species with sporadical
scap presence (mainly Japan)
Galium kamtschaticum var. Rubiaceae H Companion species with sporadical
acutifolium scap presence (mainly Japan)
Galium kikumugura Rubiaceae H Diff.-sp. of Crnoo-Fagetum crenatae
scap (subass.)
Galium kinuta Rubiaceae H Diff.-sp. of Sapio japonici-Fagetum
scap crenatae subass.)
Galium paradoxum Rubiaceae H Companion species with sporadical
scap presence (mainly Japan)
Galium pogonanthum Rubiaceae H Companion species with sporadical
scap presence (mainly Japan)
Galium trachyspermum Rubiaceae H Companion species with sporadical
scap presence (mainly Japan)
Galium trifloriforme Rubiaceae H Diff.-sp. of Saso kurilensis-Fagetum
scap crenatae
Gentiana zollingeri Gentianaceae T Companion species with sporadical
presence (mainly Japan)
Geum japonicum Rosaceae H Companion species with sporadical
scap presence (mainly Japan)
Glaucidium palmatum Ranunculaceae H Companion species with sporadical
scap presence (mainly Japan)
Globba barthei Zingiberaceae G rad Char. sp. of the Athyrio nardii-
Michelietum balansae
Gomphandra tetrandra Icacinaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Goodyera biflora Orchidaceae G rhiz Companion species with sporadical
presence (mainly China)
Goodyera cyrtoglossa Orchidaceae G rhiz Diff. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
(subass.)
Goodyera foliosa Orchidaceae G rhiz Companion-species
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 197
Life
Species name Family name form Ecological indication
Goodyera maximowicziana Orchidaceae G rhiz Companion-species
Goodyera repens Orchidaceae G rhiz Diff. sp. of the Fagetum engleriano-
lucidae
Goodyera schlechtendalina Orchidaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Goodyera velutina Orchidaceae G rhiz Companion species with sporadical
presence (mainly China)
Grangea maderaspatana Asteraceae H Companion species with sporadical
scap presence (mainly China)
Gynostemma pentaphyllum Cucurbitaceae H Widespead in the Fagus-forests of
scand East Asia
Hamamelis japonica Hamamelidaceae PD Species of the Quercetalia serratae-
grosseserratae
Hamamelis japonica var. Hamamelidaceae PD Char.-sp. of Fagion crenatae
obtusata
Hamamelis mollis Hamamelidaceae PD Diff. sp. of the Fagetum engleriano-
lucidae
Hanceola sinensis Lamiaceae H Companion species with sporadical
scap presence (mainly China)
Hedera nepalansis var. Araliaceae PL Companion species with sporadical
sinensis presence (mainly China)
Hedera nepalensis Araliaceae PL Char. sp. of the Sinarundinario
nitidae-Fagetum lucidae
Hedera nepalensis var. Araliaceae PL Companion-species
sinensis
Hedera rhombea Araliaceae PL Diff.-sp. of Hepatico-Fagetum
multinervis (subass.)
Hedera rhombera var. Araliaceae PL Diff. sp. of the Yushania-Fagetum
formosana hayatae (subass.)
Hedera sp. Araliaceae PL Companion species with sporadical
presence (mainly China)
Helenia elliptica Gentianaceae H Companion species with sporadical
scap presence (mainly China)
Helicia cochinchinensis Proteaceae PE Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Heloniopsis orientalis Liliaceae H Char.-sp. of Fagion crenatae
rosul
Helwingia chinensis Cornaceae NPD Companion-species
Helwingia himalaica Cornaceae NPD Companion species with sporadical
presence (mainly China)
Helwingia japonica Cornaceae NPD Widespead in the Fagus-forests of
East Asia
Hemselya szechuenensis Cucurbitaceae H Companion species with sporadical
scap presence (mainly China)
Hepatica maxima Ranunculaceae G rhiz Char.-sp. of Hepatico-Fagetum
multinervis
(continued)
198 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Heterosmilax japonica Liliaceae PL Companion-species
Holboellia angustifolia Lardizabalaceae PL Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Holboellia coriacea Lardizabalaceae PL Diff. sp. of the Sinarundinario
chungii-Fagetum lucidae (subass.)
Holboellia fargesii Lardizabalaceae PL Char.-sp. of Abelio englerianae-
Fagion sp. div.
Homalium bhamoense Flacourtiaceae PE Companion species with sporadical
presence (mainly China)
Hopea sp. Dipterocarpaceae PE Companion species with sporadical
presence (mainly China)
Hosiea japonica Icacinaceae PL Companion species with sporadical
presence (mainly Japan)
Hosta montana Liliaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Hugeria vaccinioides Ericaceae Ch Char. sp. of Vaccinio henryi-Fagetum
frut pashanicae
Humata tyermanni Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Hydrangea anomala Saxifragaceae PL Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Hydrangea davidii Saxifragaceae NPD Companion-species
Hydrangea hirta Saxifragaceae NPD Species of the Quercetalia serratae-
grosseserratae
Hydrangea involucrata Saxifragaceae NPD Companion species with sporadical
presence (mainly Japan)
Hydrangea luteo-venosa Saxifragaceae NPD Char.-sp. of Sapio japonici-Fagetum
crenatae
Hydrangea macrophylla Saxifragaceae NPD Species of the Fraxino-Ulmetalia
var. acuminata
Hydrangea macrophylla Saxifragaceae NPD Species of the Fraxino-Ulmetalia
var. megacarpa
Hydrangea paniculata Saxifragaceae PD Widespead in the Fagus-forests of
East Asia
Hydrangea petiolaris Saxifragaceae PL Char.-sp. of Fagetea crenatae
Hydrangea sikokiana Saxifragaceae NPD Diff.-sp. of Sapio japonici-Fagetum
crenatae subass.)
Hydrangea umbellata Saxifragaceae NPD Companion-species
Hydrangea xanthoneura Saxifragaceae NPD Char. sp. of the Sinocalamus
giganteus-Fagetum lucidae
Hydrangea yunnanensis Saxifragaceae NPD Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Idesia sp. Flacourtiaceae PD Companion species with sporadical
presence (mainly China)
Ilex chinensis Aquifoliaceae PE Companion-species
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 199
Life
Species name Family name form Ecological indication
Ilex ciliospinosa Aquifoliaceae PE Companion species with sporadical
presence (mainly China)
Ilex crenata Aquifoliaceae PE Species of the Quercetalia serratae-
grosseserratae
Ilex crenata var. paludosa Aquifoliaceae NPD Char.-sp. of Fagion crenatae
Ilex fargesii Aquifoliaceae PE Companion species with sporadical
presence (mainly China)
Ilex ficoidea Aquifoliaceae NPE Char. sp. of the Carici lanceolatae-
Fagetum hayatae
Ilex fragilis Aquifoliaceae PE Companion species with sporadical
presence (mainly China)
Ilex franchetiana Aquifoliaceae PE Companion species with sporadical
presence (mainly China)
Ilex geniculata Aquifoliaceae NPD Companion-species
Ilex intermedia var. fangli Aquifoliaceae PE Char. sp. of the Quingzheo
tumidinodae-Fagion
Ilex latifrons Aquifoliaceae PE Diff. sp. of the Sinarundinario nitidae-
Fagetum lucidae (subass.)
Ilex leucoclada Aquifoliaceae NPE Char.-sp. of Fagion crenatae
Ilex macropoda Aquifoliaceae PD Widespead in the Fagus-forests of
East Asia
Ilex pedunculosa Aquifoliaceae NPE Widespead in the Fagus-forests of
East Asia
Ilex penryi Aquifoliaceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Ilex rugosa Aquifoliaceae PL Diff.-sp. of Saso kurilensis-Fagetum
crenatae
Ilex sp. Aquifoliaceae PE Companion species with sporadical
presence (mainly China)
Ilex sugerokii var. Aquifoliaceae NPD Companion-species
brevipedunculata
Ilex szechwanensis Aquifoliaceae NPE Companion species with sporadical
presence (mainly China)
Ilex triflora Aquifoliaceae NPE Char. sp. of the Carici lanceolatae-
Fagetum hayatae
Ilex wilsonii Aquifoliaceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Ilex yunnanensis Aquifoliaceae PE Companion-species
Illicium religiosum Magnoliaceae PE Species of the Quercetalia serratae-
grosseserratae
Illicium simonsii Magnoliaceae PE Char. sp. of the Sinarundinario
chungii-Fagetum lucidae
Illicium sp. Magnoliaceae PE Companion species with sporadical
presence (mainly China)
Illicium tashiroi Magnoliaceae PE Char. sp. of the Yushania-Fagetum
hayatae
(continued)
200 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Illicium yunnanensis Magnoliaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Impatiens hypophylla Balsaminaceae H Diff.-sp. of Sapio japonici-Fagetum
scap crenatae subass.)
Impatiens siculifer Balsaminaceae H Companion species with sporadical
scap presence (mainly China)
Impatiens sp. Balsaminaceae H Companion species with sporadical
scap presence (mainly China)
Indigofera nigrescens Fabaceae NPD Char. sp. of the Carici lanceolatae-
Fagetum hayatae
Indocalamus Poaceae GS Companion species with sporadical
fanjinshanensis presence (mainly China)
Indocalamus latifolius Poaceae GS Char. sp. of the Indocalamo latifolii—
Fagion hayatae var. zhejiangensis
Indocalamus longiauritus Poaceae GS Companion-species
Indocalamus sp. Poaceae GS Companion species with sporadical
presence (mainly China)
Indosasa shibataeoides Poaceae GS Char. sp. of the Sinarundinario
bashersuto-Fagetum lucidae
Isodon excisus Lamiaceae H Companion species with sporadical
scap presence (mainly Japan)
Itea chinensis Saxifragaceae NPE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Itea chinensis var. oblonga Saxifragaceae NPE Char. sp. of the Carici lanceolatae-
Fagetum hayatae
Kadsura heteroclita Magnoliaceae PL Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Kadsura longepedunculata Magnoliaceae PL Companion species with sporadical
presence (mainly China)
Kalopanax pictus Araliaceae PD Char.-sp. of Fagetea crenatae
Kalopanax septemlobus Araliaceae PD Companion-species
Kerria japonica Rosaceae NPD Companion-species
Kilengeshoma palmata Saxifragaceae H Diff.-sp. of Sapio japonici-Fagetum
scap crenatae subass.)
Koelreuteria paniculata Sapindaceae PD Companion species with sporadical
presence (mainly China)
Lactuca graciliflora Asteraceae H Diff. sp. of the Sinarundinario nitidae-
scap Fagetum lucidae (subass.)
Lamium humile Lamiaceae H Diff.-sp. of Crnoo-Fagetum crenatae
scap (subass.)
Laportea bulbifera Urticaceae H Species of the Fraxino-Ulmetalia
scap
Lasianthus biermannii Rubiaceae NPE Char. sp. of the Ardisio hypargeriae—
Castanopsion fabrii
Lasianthus henryi Rubiaceae NPE Companion species with sporadical
presence (mainly China)
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 201
Life
Species name Family name form Ecological indication
Lasianthus japonicus Polypodiaceae NPE Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Lasianthus longicaudus Rubiaceae NPE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Lastrea quelpaertensis Polypodiaceae H Companion species with sporadical
rosul presence (mainly Japan)
Lepisorus bicolor Polypodiaceae E Char. sp. of the Viburno flavescentis-
Fagetum englerianae
Lepisorus contortus Polypodiaceae E Companion species with sporadical
presence (mainly China)
Lepisorus obscure- Polypodiaceae E Diff. sp. of the Yushania-Fagetum
venulosus hayatae (subass.)
Lepisorus onoei Polypodiaceae E Companion-species
Lepisorus sp. Polypodiaceae E Companion species with sporadical
presence (mainly China)
Lepisorus thunbergianus Polypodiaceae E Widespead in the Fagus-forests of
East Asia
Lepisorus ussuriensis var. Polypodiaceae E Companion-species
distans
Leptogramma mollissima Polypodiaceae G rhiz Species of the Fraxino-Ulmetalia
Leptogramma scallani Polypodiaceae G rhiz Char. sp. of the Polypodio argutum-
Fagetum longepetiolatae
Leptorumohra miqueliana Polypodiaceae G rhiz Widespead in the Fagus-forests of
East Asia
Lespedeza buergeri Fabaceae NPD Widespead in the Fagus-forests of
East Asia
Lespedeza formosa Fabaceae NPD Diff. sp. of the Fagetum engleriano-
lucidae
Leucosceptrum japonicum Lamiaceae H Diff.-sp. of Sapio japonici-Fagetum
scap crenatae subass.)
Leucosceptrum sp. Lamiaceae H Companion species with sporadical
scap presence (mainly Japan)
Leucosceptrum stellipilum Lamiaceae H Diff.-sp. of Sapio japonici-Fagetum
var. tosaense scap crenatae subass.)
Leucothoe grayana var. Ericaceae NPD Companion species with sporadical
oblongifolia presence (mainly Japan)
Ligularia dentata Asteraceae H Diff.-sp. of Saso kurilensis-Fagetum
scap crenatae
Ligularia fischeri Asteraceae H Companion species with sporadical
scap presence (mainly Japan)
Ligularia stenocephala Asteraceae H Diff.-sp. of Sapio japonici-Fagetum
scap crenatae subass.)
Ligularia veitchiana Asteraceae H Diff. sp. of the Fagetum engleriano-
scap lucidae
Ligustrum delavayanum Oleaceae NPE Char. sp. of the Sinocalamus
giganteus-Fagetum lucidae
(continued)
202 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Ligustrum foliosum Oleaceae NPD Char.-sp. of Fagion multinrvis and
Fagetalia multinervis
Ligustrum japonicum Oleaceae NPD Companion species with sporadical
presence (mainly China)
Ligustrum obtusifolium Oleaceae NPD Companion species with sporadical
presence (mainly Japan)
Ligustrum sinense Oleaceae NPD Companion species with sporadical
presence (mainly China)
Ligustrum tschonoskii Oleaceae NPD Species of the Fraxino-Ulmetalia
Ligustrum tschonoskii var. Oleaceae NPD Diff.-sp. of Saso kurilensis-Fagetum
glabrescens crenatae
Lilium auratum Liliaceae G Companion species with sporadical
bulb presence (mainly Japan)
Lilium cordatum Liliaceae G Companion-species
bulb
Lilium hansonii Liliaceae G Char.-sp. of Hepatico-Fagetum
bulb multinervis
Lilium medeoloides Liliaceae G Companion species with sporadical
bulb presence (mainly Japan)
Lindera cercidifolia Lauraceae PD Companion species with sporadical
presence (mainly China)
Lindera communis Lauraceae NPD Companion species with sporadical
presence (mainly China)
Lindera erythrocarpa Lauraceae PD Widespead in the Fagus-forests of
East Asia
Lindera floribunda Lauraceae PD Diff. sp. of the Fagetum engleriano-
lucidae
Lindera fragrans Lauraceae NPE Companion-species
Lindera fruticosa Lauraceae PD Companion species with sporadical
presence (mainly China)
Lindera glauca Lauraceae NPD Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Lindera metcalfiana Lauraceae PD Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Lindera obtusiloba Lauraceae NPD Widespead in the Fagus-forests of
East Asia
Lindera reflexa Lauraceae NPD Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Lindera sericea Lauraceae NPD Companion species with sporadical
presence (mainly Japan)
Lindera sericea var. Lauraceae NPD Char.-sp. of Sapio japonici-Fagetum
glabrata crenatae
Lindera sp. Lauraceae PD Companion species with sporadical
presence (mainly China)
Lindera subcaudata var. Lauraceae PD Char. sp. of the Tripterospermo
hemsleyana cordifolium-Fagetum englerianae
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 203
Life
Species name Family name form Ecological indication
Lindera umbellata Lauraceae NPD Widespead in the Fagus-forests of
East Asia
Lindera umbellata var. Lauraceae NPD Char.-sp. of Fagion crenatae
membranacea
Liparis japonica Orchidaceae G rhiz Char. sp. of the Athyrio nardii-
Michelietum balansae
Liparis krameri Orchidaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Liquidamber acalycina Hamamelidaceae PD Char. sp. of the Carici lanceolatae-
Fagetum hayatae var. zhejiangensis
Liquidamber formosana Hamamelidaceae PD Companion species with sporadical
presence (mainly China)
Liriope graminifolia Liliaceae G rhiz Char. sp. of the Indocalamo latifolii—
Fagion hayatae var. zhejiangensis
Liriope muscari Liliaceae G rhiz Diff. sp. of the Sinarundinario
chungii-Fagetum lucidae (subass.)
Liriope platyphylla Liliaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Liriope spicata Liliaceae G rhiz Companion-species
Lithocarpus carolinae Lithocarpaceae PE Char. sp. of the Athyrio nardii-
Michelietum balansae
Lithocarpus cleistocarpus Lithocarpaceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Lithocarpus confinis Lithocarpaceae PE Companion species with sporadical
presence (mainly China)
Lithocarpus dealbatus Lithocarpaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Lithocarpus fruncatus Lithocarpaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Lithocarpus glober Lithocarpaceae PE Diff. sp. of the Sinarundinario
bashersuto-Fagetum lucidae (subass.)
Lithocarpus hancei Lithocarpaceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Lithocarpus harlandii Lithocarpaceae PE Char. sp. of the Carici lanceolatae-
Fagetum hayatae
Lithocarpus henryi Lithocarpaceae PE Char. sp. of the Aceri davidii-Fagion
lucidae
Lithocarpus megalophyllus Lithocarpaceae PE Char. sp. of the Ardisio hypargeriae—
Castanopsion fabrii
Lithocarpus naiadarum Lithocarpaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Lithocarpus sp1. Lithocarpaceae PE Companion species with sporadical
presence (mainly China)
Lithocarpus sp2. Lithocarpaceae PE Companion species with sporadical
presence (mainly China)
(continued)
204 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Lithocarpus thomsonii Lithocarpaceae PE Companion species with sporadical
presence (mainly China)
Lithocarpus variolosus Lithocarpaceae PE Companion species with sporadical
presence (mainly China)
Litsea acuminata Lauraceae PE Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Litsea coreana Lauraceae PE Companion-species
Litsea cubeba Lauraceae PE Companion species with sporadical
presence (mainly China)
Litsea elongata Lauraceae PE Char.-sp. of Litseo elongatae-Fagetea
Litsea grandiflora Lauraceae PE Companion species with sporadical
presence (mainly China)
Litsea ichangensis Lauraceae PE Companion species with sporadical
presence (mainly China)
Litsea pedunculata Lauraceae PE Diff. sp. of the Sinarundinario
bashersuto-Fagetum lucidae (subass.)
Litsea populifolia Lauraceae PE Companion species with sporadical
presence (mainly China)
Litsea pungens Lauraceae PE Char. sp. of the Aceri davidii-Fagion
lucidae
Litsea ssp. Lauraceae PE Companion species with sporadical
presence (mainly China)
Litsea suberosa Lauraceae PE Diff. sp. of the Sinarundinario nitidae-
Fagetum lucidae (subass.)
Litsea thomsinii Lauraceae PE Companion species with sporadical
presence (mainly China)
Lonicera acuminata Caprifoliaceae NPD Companion species with sporadical
presence (mainly China)
Lonicera gracilipes Caprifoliaceae NPD Companion species with sporadical
presence (mainly Japan)
Lonicera gracilipes var. Caprifoliaceae NPD Companion species with sporadical
glandulosa presence (mainly Japan)
Lonicera gynochlamydea Caprifoliaceae NPD Companion species with sporadical
presence (mainly China)
Lonicera henryi Caprifoliaceae PL Companion-species
Lonicera japonica Caprifoliaceae PL Char. sp. of the Aceri davidii-Fagion
lucidae
Lonicera ligustrina Caprifoliaceae NPD Companion species with sporadical
presence (mainly China)
Lonicera Caprifoliaceae NPD Char.-sp. of Abelio englerianae-
pseudoproterantha Fagion sp. div.
Lonicera similis Caprifoliaceae PL Companion species with sporadical
presence (mainly China)
Lonicera sp. Caprifoliaceae NPD Companion species with sporadical
presence (mainly China)
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 205
Life
Species name Family name form Ecological indication
Lophatherum gracile Poaceae G rhiz Char. sp. of the Polypodio argutum-
Fagetum longepetiolatae
Loxogramme remote- Polypodiaceae G rhiz Char. sp. of the Yushania-Fagetum
frondigera hayatae
Luzula plumosa Poaceae G rhiz Diff. sp. of the Fagetum engleriano-
lucidae
Luzula plumosa var. Poaceae H Companion species with sporadical
macrocarpa caesp presence (mainly Japan)
Lycopodium clavatum Lycopodiaceae Ch Companion species with sporadical
suffr presence (mainly Japan)
Lycopodium obscurum Lycopodiaceae Ch Companion species with sporadical
suffr presence (mainly Japan)
Lycopodium serratum Lycopodiaceae Ch Companion-species
suffr
Lycopodium serratum var. Lycopodiaceae Ch Diff. sp. of the Yushania-Fagetum
longipetiolatum frut hayatae (subass.)
Lycopodium serratum var. Lycopodiaceae Ch Companion-species
serratum suffr
Lyonia ovalifolia Ericaceae PD Char. sp. of the Carici lanceolatae-
Fagetum hayatae
Lyonia ovalifolia Ericaceae NPD Companion species with sporadical
presence (mainly China)
Lyonia ovalifolia var. Ericaceae NPD Widespead in the Fagus-forests of
elliptica East Asia
Lyonia ovalifolia var. Ericaceae NPD Companion species with sporadical
ovalifolia presence (mainly China)
Lysimachia ardisioides Primulaceae H Companion species with sporadical
scap presence (mainly China)
Lysimachia clethroides Primulaceae H Companion species with sporadical
scap presence (mainly China)
Maackia hwashanensis Fabaceae PD Companion species with sporadical
presence (mainly China)
Machilus faberi Lauraceae PE Companion species with sporadical
presence (mainly China)
Machilus ichangensis Lauraceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Machilus kurzii Lauraceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Machilus rehderi Lauraceae PE Companion-species
Machilus salicina Lauraceae PE Char. sp. of the Athyrio nardii-
Michelietum balansae
Machilus sp1. Lauraceae PE Companion species with sporadical
presence (mainly China)
Machilus sp2. Lauraceae PE Companion species with sporadical
presence (mainly China)
(continued)
206 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Machilus thunbergii Lauraceae PE Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Maesa japonica Myrsinaceae NPE Widespead in the Fagus-forests of
East Asia
Magnolia biondii Magnoliaceae PE Diff. sp. of the Sinarundinario nitidae-
Fagetum lucidae (subass.)
Magnolia cylindrica Magnoliaceae PD Char. sp. of the Indocalamo latifolii—
Fagion hayatae var. zhejiangensis
Magnolia kobus Magnoliaceae PD Companion species with sporadical
presence (mainly Japan)
Magnolia kobus var. Magnoliaceae PD Companion species with sporadical
borealis presence (mainly Japan)
Magnolia obovata Magnoliaceae PD Char.-sp. of Saso-Fagetalia
Magnolia salicifolia Magnoliaceae NPD Char.-sp. of Fagion crenatae
Magnolia sieboldii Magnoliaceae NPD Companion species with sporadical
presence (mainly Japan)
Magnolia sp. Magnoliaceae PE Companion species with sporadical
presence (mainly China)
Magnolia sprengeri Magnoliaceae PE Companion-species
Mahonia bealei Berberidaceae NPE Companion-species
Mahonia gracilipes Berberidaceae NPE Companion species with sporadical
presence (mainly China)
Mahonia japonica Berberidaceae NPE Char. sp. of the Aceri davidii-Fagion
lucidae
Maianthemum dilatatum Liliaceae G rhiz Char.-sp. of Fagetea crenatae
Malus tschonoskii Rosaceae PD Companion species with sporadical
presence (mainly Japan)
Manglietia duclouxii Magnoliaceae PE Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Manglietia fordiana Magnoliaceae PE Char. sp. of the Sinarundinario
bashersuto-Fagetum lucidae
Manglietia rufibarbata Magnoliaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Matteuccia orientalis Polypodiaceae H Widespead in the Fagus-forests of
rosul East Asia
Melampyrum roseum Scrophulariaceae H Diff. sp. of the Fagetum engleriano-
scap lucidae
Meliosma flexuosa Sabiaceae PD Companion species with sporadical
presence (mainly China)
Meliosma myriantha Sabiaceae PD Widespead in the Fagus-forests of
East Asia
Meliosma myriantha var. Sabiaceae NPD Char. sp. of the Carici lanceolatae-
discolor Fagetum hayatae var. zhejiangensis
Meliosma paupera Sabiaceae PD Companion species with sporadical
presence (mainly China)
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 207
Life
Species name Family name form Ecological indication
Meliosma sichourensis Sabiaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Meliosma sp. Sabiaceae PD Companion species with sporadical
presence (mainly China)
Meliosma tenuis Sabiaceae NPD Species of the Quercetalia serratae-
grosseserratae
Meliosma veitchiorum Sabiaceae PD Diff.-sp. of Euonymo porphyrei-
Fagetum englerianae (subass.)
Melothria maysorensis Cucurbitaceae H Companion species with sporadical
scap presence (mainly China)
Menziesia ciliicalyx Ericaceae NPD Char.-sp. of Lindero umbellatae-
Fagetum crenatae
Menziesia multiflora Ericaceae NPD Companion species with sporadical
presence (mainly Japan)
Menziesia pentandra Ericaceae NPD Diff.-sp. of Saso kurilensis-Fagetum
crenatae
Michelia balansae Magnoliaceae PE Char. sp. of the Athyrio nardii-
Michelietum balansae
Michelia sp. Magnoliaceae PE Companion species with sporadical
presence (mainly China)
Michelia yunnanensis Magnoliaceae PE Char. sp. of the Athyrio nardii-
Michelietum balansae
Microlepia marginata Polypodiaceae G rhiz Char. sp. of the Sinocalamus
giganteus-Fagetum lucidae
Microsorium buergerianum Polypodiaceae G rhiz Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Microsorium hymenodes Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Microtropis fokienensis Celastraceae NPE Companion-species
Millettia championii Fabaceae PL Companion species with sporadical
presence (mainly China)
Miricacalia makineana Asteraceae H Diff.-sp. of Crnoo-Fagetum crenatae
scap (subass.)
Mitchella undulata Rubiaceae H Char.-sp. of Fagetea crenatae
scand
Monochosorum henryi Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Monotropa hypopitys Pyrolaceae G Companion species with sporadical
paras presence (mainly Japan)
Monotropa uniflora Pyrolaceae G Companion species with sporadical
paras presence (mainly China)
Monotropastrum globosum Pyrolaceae G Companion-species
paras
Morinda umbellata Rubiaceae NPD Companion species with sporadical
presence (mainly China)
(continued)
208 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Morus bombycis Moraceae PD Companion species with sporadical
presence (mainly Japan)
Morus mongolica Moraceae PD Diff. sp. of the Fagetum engleriano-
lucidae
Myrica rubra var. Myricaceae PE Diff. sp. of the Yushania-Fagetum
acuminata hayatae (subass.)
Myrsine sp. Celastraceae NPE Companion species with sporadical
presence (mainly China)
Neolitsea aciculata Lauraceae PE Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Neolitsea acuminatissima Lauraceae PE Char. sp. of the Yushania-Fagetum
hayatae
Neolitsea aurata Lauraceae PE Companion species with sporadical
presence (mainly China)
Neolitsea chinensis Lauraceae PE Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Neolitsea chuii Lauraceae PE Char. sp. of the Viburno flavescentis-
Fagetum englerianae
Neolitsea levinei Lauraceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Neolitsea sericea Lauraceae PE Companion species with sporadical
presence (mainly Japan)
Neolitsea sp. Lauraceae PE Companion species with sporadical
presence (mainly China)
Neolitsea zeylanica Lauraceae PE Companion species with sporadical
presence (mainly China)
Nothopanax davidii Araliaceae NPD Companion species with sporadical
presence (mainly China)
Nyssa javanica Nyssaceae PD Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Nyssa sinensis Nyssaceae PD Companion-species
Omphalodes japonica Boraginaceae H Companion-species
scap
Ophiopogon bockianus Liliaceae G rhiz Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Ophiopogon bodinieri Liliaceae G rhiz Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Ophiopogon clavatus Liliaceae G rhiz Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Ophiopogon intermedius Liliaceae G rhiz Companion species with sporadical
presence (mainly China)
Ophiopogon japonicus Liliaceae G rhiz Diff.-sp. of Hepatico-Fagetum
multinervis (subass.)
Ophiopogon mairei Liliaceae G rhiz Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 209
Life
Species name Family name form Ecological indication
Ophiopogon planiscapus Liliaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Ophiopogon spp. Liliaceae G rhiz Companion species with sporadical
presence (mainly China)
Ophiorrhiza japonica Rubiaceae H Char.-sp. of Sinarundinario nitidae-
scand Fagetalia sp. div.
Oplismenus compostius Poaceae H Companion species with sporadical
caesp presence (mainly China)
Oplismenus undulatifolius Poaceae H Companion-species
caesp
Oplismenus undulatifolius Poaceae Ch Companion species with sporadical
var. japonicus suffr presence (mainly Japan)
Oplopanax japonicus Araliaceae NPD Diff.-sp. of Saso kurilensis-Fagetum
crenatae
Oreocharis benthamii var. Gesneriaceae H Char. sp. of the Sinarundinario
reticulata rosul bashersuto-Fagetum lucidae
Oreorchis patens Orchidaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Orixa japonica Rutaceae NPD Companion species with sporadical
presence (mainly Japan)
Osmanthus corymbosus Oleaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Osmanthus heterophyllus Oleaceae PE Char. sp. of the Yushania-Fagetum
hayatae
Osmanthus sp. Oleaceae PE Companion species with sporadical
presence (mainly China)
Osmanthus yunanenesis Oleaceae PE Companion species with sporadical
presence (mainly China)
Osmorhiza aristata Apiaceae H Diff.-sp. of Saso kurilensis-Fagetum
scap crenatae
Osmunda cinnamomea var. Osmundaceae H Companion-species
fokiensis rosul
Osmunda japonica Osmundaceae H Companion-species
rosul
Ostrya japonica Betulaceae PD Companion species with sporadical
presence (mainly Japan)
Oxalis acetosella Oxalidaceae G rhiz Widespead in the Fagus-forests of
East Asia
Oxalis griffithii Oxalidaceae G rhiz Widespead in the Fagus-forests of
East Asia
Oxalis griffithii var. Oxalidaceae G rhiz Diff.-sp. of Sasamorpho_Fagion
kantoensis crenatae (subass.)
Pachysandra terminalis Buxaceae H Widespead in the Fagus-forests of
scap East Asia
Padus grayana Rosaceae PD Diff. sp. of the Sinarundinario
chungii-Fagetum lucidae (subass.)
(continued)
210 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Padus wilsonii Rosaceae PD Companion species with sporadical
presence (mainly China)
Paederia scandens Rubiaceae PL Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Paeonia japonica Ranunculaceae G rad Diff.-sp. of Sapio japonici-Fagetum
crenatae subass.)
Panax japonicus Araliaceae H Species of the Fraxino-Ulmetalia
scap
Panax transitorius Araliaceae H Companion species with sporadical
scap presence (mainly China)
Panicum brevifolium Poaceae T Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Parabenzoin praecox Lauraceae NPD Char.-sp. of Corno-Fagetum crenatae
Parabenzoin trilobum Lauraceae PD Char.-sp. of Sapio japonici-Fagetum
crenatae
Parakmeria yunnanensis Magnoliaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Parathelypteris Polypodiaceae G rhiz Char.-sp. of Litseo elongatae-Fagetea
glanduligera
Parathelypteris hirsutipes Polypodiaceae G rhiz Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Parathelypteris japonica Polypodiaceae G rhiz Widespead in the Fagus-forests of
East Asia
Parathelypteris nipponica Polypodiaceae G rhiz Diff. sp. of the Fagetum engleriano-
lucidae
Parathenocissus Vitaceae PL Companion species with sporadical
himalayana presence (mainly China)
Paris bashanensis Liliaceae H Companion species with sporadical
scap presence (mainly China)
Paris tetraphylla Liliaceae G rhiz Char.-sp. of Saso-Fagetalia
Parkmeria yunnanensis Magnoliaceae PE Char. sp. of the Athyrio nardii-
Michelietum balansae
Parthenocissus Vitaceae PL Char.-sp. of Sinarundinario nitidae-
heterophylla Fagetalia sp. div.
Patrinia scabiosaefolia Valerianaceae H Diff. sp. of the Fagetum engleriano-
scap lucidae
Patrinia triloba var. Valerianaceae H Companion species with sporadical
palmata scap presence (mainly Japan)
Pedicularis nasturtifolia Scrophulariaceae PD Diff.-sp. of Euonymo porphyrei-
Fagetum englerianae (subass.)
Pellionia arisanensis Urticaceae PL Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Pellionia radicans Urticaceae PL Companion-species
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 211
Life
Species name Family name form Ecological indication
Peracarpa carnosa Campanulaceae H Widespead in the Fagus-forests of
scap East Asia
Peracarpa carnosa var. Campanulaceae H Species of the Fraxino-Ulmetalia
circaeoides scap
Peracarpa sp. Campanulaceae H Companion species with sporadical
scap presence (mainly China)
Pertya cordifolia Asteraceae H Diff. sp. of the Fagetum engleriano-
scap lucidae
Pertya glabrescens Asteraceae NPD Widespead in the Fagus-forests of
East Asia
Pertya rigidula Asteraceae H Diff.-sp. of Lindero umbellatae-
scap Fagetum crenatae
Pertya robusta Asteraceae H Diff.-sp. of Sasamorpho_Fagion
scap crenatae (subass.)
Pertya scandens Asteraceae NPD Diff.-sp. of Sasamorpho_Fagion
crenatae (subass.)
Pertya triloba Asteraceae H Diff.-sp. of Sasamorpho_Fagion
scap crenatae (subass.)
Phanerophlebiopsis blinii Polypodiaceae G rhiz Diff. sp. of the Sinarundinario nitidae-
Fagetum lucidae (subass.)
Phegopteris connectilis Polypodiaceae H Companion-species
rosul
Phellodendron amurense Rutaceae PD Companion species with sporadical
presence (mainly Japan)
Phellodendron chinense Rutaceae PD Char. sp. of the Sinarundinario
bashersuto-Fagetum lucidae
Philadelphus incanus Saxifragaceae NPD Char. sp. of the Fagetum engleriano-
lucidae
Philadelphus satsumi Saxifragaceae NPD Diff.-sp. of Sapio japonici-Fagetum
crenatae subass.)
Phoebe faberi Lauraceae PE Companion species with sporadical
presence (mainly China)
Phoebe neurantha Lauraceae PE Char. sp. of the Elatostemo
sessile_Fagetum lucidae
Phoebe sheareri Lauraceae PE Char. sp. of the Elatostemo
sessile_Fagetum lucidae
Photinia paniculata Rosaceae PD Char. sp. of the Carici lanceolatae-
Fagetum hayatae
Photinia parvifolia Rosaceae PD Char. sp. of the Indocalamo latifolii—
Fagion hayatae var. zhejiangensis
Photinia sp. Rosaceae PD Companion species with sporadical
presence (mainly China)
Photinia villosa Rosaceae PD Diff. sp. of the Sinarundinario nitidae-
Fagetum lucidae (subass.)
Phryma leptostachya Phrymaceae H Companion-species
v .asiatica scap
(continued)
212 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Phyllagathis longipes Polypodiaceae H Char. sp. of the Viburno flavescentis-
caesp Fagetum englerianae
Phyllagathis ovalifolia Polypodiaceae H Diff. sp. of the Prismatomerio
caesp henryi—Lithocarpetum naiadari
(subass.)
Phymatopsis sp. Polypodiaceae H Companion species with sporadical
caesp presence (mainly China)
Phymatopsis teneupes Polypodiaceae H Companion species with sporadical
caesp presence (mainly China)
Picea polita Pinaceae PEN Companion species with sporadical
presence (mainly Japan)
Picrasma quassioides Simaroubaceae NPD Char. sp. of the Carici lanceolatae-
Fagetum hayatae var. zhejiangensis
Pieris formosa Ericaceae NPE Diff. sp. of the Fagetum engleriano-
lucidae
Pieris japonica Ericaceae NPE Widespead in the Fagus-forests of
East Asia
Pieris taiwanensis Ericaceae NPE Char. sp. of the Yushania-Fagetum
hayatae
Pieris tomentosa Ericaceae NPE Companion species with sporadical
presence (mainly China)
Pilea martinii Urticaceae NPE Companion species with sporadical
presence (mainly China)
Pilea sp. Urticaceae NPE Companion species with sporadical
presence (mainly China)
Pinus armandii Pinaceae PEN Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Pinus densiflora Pinaceae PEN Companion species with sporadical
presence (mainly Japan)
Pinus parviflora Pinaceae PEN Companion species with sporadical
presence (mainly Japan)
Piper cft. flaviiflorum Piperaceae PL Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Piper sp. Piperaceae PL Companion species with sporadical
presence (mainly China)
Pittosporum glabratum var. Pittosporaceae NPE Companion species with sporadical
neriifolium presence (mainly China)
Plagiogyria stenoptera Plagiogyriaceae H Diff. sp. of the Sinarundinario nitidae-
rosul Fagetum lucidae (subass.)
Plagiogyria distinctissima Plagiogyriaceae H Companion species with sporadical
rosul presence (mainly China)
Plagiogyria euphlebia Plagiogyriaceae H Widespead in the Fagus-forests of
rosul East Asia
Plagiogyria formosana Plagiogyriaceae H Char. sp. of the Yushania-Fagetum
rosul hayatae
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 213
Life
Species name Family name form Ecological indication
Plagiogyria matsumureana Plagiogyriaceae H Char.-sp. of Fagion crenatae
rosul
Plagiogyria maxima Plagiogyriaceae H Diff. sp. of the Prismatomerio
rosul henryi—Lithocarpetum naiadari
(subass.)
Plagiogyria stenoptera Plagiogyriaceae H Char.-sp. of Sinarundinario nitidae-
rosul Fagetalia sp. div.
Platanthera florentii Orchidaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Platanthera sachalinensis Orchidaceae G rhiz Diff.-sp. of Saso kurilensis-Fagetum
crenatae
Platycarya strobilacea Junglandaceae PD Char. sp. of the Fagetum engleriano-
lucidae
Plectranthus inflexus Lamiaceae H Companion species with sporadical
scap presence (mainly Japan)
Plectranthus kameba Lamiaceae H Companion species with sporadical
scap presence (mainly Japan)
Plectranthus longitubus Lamiaceae H Companion-species
scap
Plectranthus shikokianus Lamiaceae H Companion species with sporadical
var. intermedius scap presence (mainly Japan)
Plectranthus umbrosus Lamiaceae H Char.-sp. of Corno-Fagetum crenatae
scap
Pleioblastus chino Poaceae GS Companion species with sporadical
presence (mainly Japan)
Pleuropus euchloron Poaceae Ch Companion species with sporadical
suffr presence (mainly China)
Poa takeshimana Poaceae H Companion species with sporadical
caesp presence (mainly Japan)
Pollia sp. Commelinaceae H Companion species with sporadical
scap presence (mainly China)
Polygala tricornis Polygalaceae NPD Char. sp. of the Ardisio hypargeriae—
Castanopsion fabrii
Polygonatum cyrtonema Liliaceae H Char. sp. of the Aceri davidii-Fagion
scap lucidae
Polygonatum falcatum Liliaceae G Companion-species
bulb
Polygonatum lasianthum Liliaceae G rhiz Companion-species
Polygonatum macranthum Liliaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Polygonatum odoratum Liliaceae G Char.-sp. of Sinarundinario nitidae-
bulb Fagetalia sp. div.
Polygonatum odoratum var. Liliaceae G rhiz Companion species with sporadical
pluriflorum presence (mainly Japan)
Polygonatum sibiricum Liliaceae G rhiz Char. sp. of the Carici lanceolatae-
Fagetum hayatae
(continued)
214 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Polygonum cuspidatum Polygonaceae H Companion species with sporadical
scap presence (mainly Japan)
Polygonum debile Polygonaceae H Diff.-sp. of Sapio japonici-Fagetum
scap crenatae subass.)
Polygonum filiforme Polygonaceae H Companion species with sporadical
scap presence (mainly Japan)
Polygonum suffultum Polygonaceae H Diff. sp. of the Fagetum engleriano-
scap lucidae
Polygonum tenuicaule Polygonaceae H Diff.-sp. of Sapio japonici-Fagetum
scap crenatae subass.)
Polygonum thunbergii Polygonaceae T Companion species with sporadical
presence (mainly Japan)
Polypodiastrum dielsianum Polypodiaceae G rhiz Char. sp. of the Viburno flavescentis-
Fagetum englerianae
Polypodium amoenum Polypodiaceae G rhiz Char. sp. of the Yushania-Fagetum
hayatae
Polypodium argutum Polypodiaceae G rhiz Char. sp. of the Polypodio argutum-
Fagetum longepetiolatae
Polypodium vulgare Polypodiaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Polystichum braunii Polypodiaceae G rhiz Diff.-sp. of Saso kurilensis-Fagetum
crenatae
Polystichum deltodon Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Polystichum discretum Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Polystichum makinoi Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Polystichum neolobatum Polypodiaceae G rhiz Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Polystichum ovato- Polypodiaceae G rhiz Species of the Fraxino-Ulmetalia
paleaceum
Polystichum parvipinnulum Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Polystichum polyblepharum Polypodiaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Polystichum retroso- Polypodiaceae G rhiz Char.-sp. of Fagion multinrvis and
paleaceum Fagetalia multinervis
Polystichum sp. Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Polystichum squarrosum Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
Polystichum tripteron Polypodiaceae G rhiz Species of the Fraxino-Ulmetalia
Polystichum tsus-sinense Polypodiaceae G rhiz Companion species with sporadical
presence (mainly China)
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 215
Life
Species name Family name form Ecological indication
Potentilla yokusaiana Rosaceae H Diff.-sp. of Crnoo-Fagetum crenatae
rosul (subass.)
Pourthiaea villosa Rosaceae PD Companion species with sporadical
presence (mainly Japan)
Pourthiaea villosa var. Rosaceae PD Char. sp. of the Yushania-Fagetum
parvifolia hayatae
Pourthiaea villosa var. Rosaceae PD Species of the Quercetalia serratae-
laevis grosseserratae
Prenanthes acerifolia Asteraceae H Companion species with sporadical
scap presence (mainly Japan)
Primula ovalifolia Primulaceae H Char. sp. of the Elatostemo
rosul sessile_Fagetum lucidae
Primula sp. Primulaceae H Companion species with sporadical
rosul presence (mainly China)
Prismatomeris henryi Rubiaceae NPD Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Prismatomeris labordei Rubiaceae NPD Companion species with sporadical
presence (mainly China)
Prunus apetala Rosaceae PD Companion species with sporadical
presence (mainly Japan)
Prunus brachypoda Rosaceae PD Companion species with sporadical
presence (mainly China)
Prunus buergeriana Rosaceae PD Companion species with sporadical
presence (mainly Japan)
Prunus dielsiana Rosaceae PD Companion species with sporadical
presence (mainly China)
Prunus grayana Rosaceae PD Char.-sp. of Saso-Fagetalia
Prunus incisa Rosaceae PD Char.-sp. of Corno-Fagetum crenatae
Prunus incisa var. Rosaceae PD Diff.-sp. of Lindero umbellatae-
kinkiensis Fagetum crenatae
Prunus jamasakura Rosaceae PD Species of the Fraxino-Ulmetalia
Prunus maximowiczii Rosaceae PD Diff.-sp. of Saso kurilensis-Fagetum
crenatae
Prunus phaeosticta Rosaceae PD Companion-species
Prunus pilosiuscula Rosaceae PD Char.-sp. of Abelio englerianae-
Fagion sp. div.
Prunus serrulata Rosaceae PD Char. sp. of the Carici lanceolatae-
Fagetum hayatae
Prunus sp. Rosaceae PD Companion species with sporadical
presence (mainly China)
Prunus ssiori Rosaceae PD Diff.-sp. of Saso kurilensis-Fagetum
crenatae
Prunus takesimensis Rosaceae PD Char.-sp. of Fagion multinrvis and
Fagetalia multinervis
Prunus transarlsanensis Rosaceae PD Companion species with sporadical
presence (mainly China)
(continued)
216 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Prunus vaniotii Rosaceae PD Companion species with sporadical
presence (mainly China)
Prunus verecunda Rosaceae PD Diff.-sp. of Sasamorpho_Fagion
crenatae (subass.)
Pseudocystopteris Athyriaceae G rhiz Companion species with sporadical
atkinsonii presence (mainly China)
Pteridophyllum racemosum Papaveraceae H Companion species with sporadical
scap presence (mainly Japan)
Pternopetalum tanakae Apiaceae H Diff.-sp. of Sasamorpho_Fagion
scap crenatae (subass.)
Pterocarya dalavayi Junglandaceae PD Companion species with sporadical
presence (mainly China)
Pterocarya rhoifolia Junglandaceae PD Species of the Fraxino-Ulmetalia
Pterostyrax corymbosa Styracaceae PD Companion species with sporadical
presence (mainly Japan)
Pterostyrax rosea Styracaceae PD Companion species with sporadical
presence (mainly China)
Pterygocalyx volubilis Gentianaceae H Char. sp. of the Aceri davidii-Fagion
caesp lucidae
Pyrola decorata Pyrolaceae H Char.-sp. of Sinarundinario nitidae-
scap Fagetalia sp. div.
Pyrola japonica Pyrolaceae G rhiz Companion-species
Pyrola renifolia Pyrolaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Pyrola rugosa Pyrolaceae G rhiz Companion species with sporadical
presence (mainly China)
Pyrrosia lingua Polypodiaceae G rhiz Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Pyrrosia sheareri Polypodiaceae G rhiz Companion-species
Qiongzhea tumidinoda Poaceae GS Char. sp. of the Quingzheo
tumidinodae-Fagion
Quercus acuta Fagaceae PE Diff.-sp. of Sasamorpho_Fagion
crenatae (subass.)
Quercus aliena var. Fagaceae PD Char. sp. of the Fagetum engleriano-
acutidentata lucidae
Quercus asine Fagaceae PE Companion species with sporadical
presence (mainly China)
Quercus augustinii Fagaceae PE Companion species with sporadical
presence (mainly China)
Quercus bambusaefolia Fagaceae PE Char. sp. of the Sinarundinario
bashersuto-Fagetum lucidae
Quercus chapaensis Fagaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Quercus engleriana Fagaceae PE Companion-species
Quercus fabri Fagaceae PD Char. sp. of the Carici lanceolatae-
Fagetum hayatae
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 217
Life
Species name Family name form Ecological indication
Quercus glandulifera Fagaceae PE Char.-sp. of Abelio englerianae-
Fagion sp. div.
Quercus gracilis Fagaceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Quercus mongolica var. Fagaceae PD Char.-sp. of Saso-Fagetalia
grosseserrata
Quercus morii Fagaceae PE Companion species with sporadical
presence (mainly China)
Quercus multinervis Fagaceae PE Char. sp. of the Aceri davidii-Fagion
lucidae
Quercus nubium Fagaceae PE Char. sp. of the Indocalamo latifolii—
Fagion hayatae var. zhejiangensis
Quercus oxyodon Fagaceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Quercus salicina Fagaceae PE Companion species with sporadical
presence (mainly Japan)
Quercus serrata Fagaceae PD Species of the Quercetalia serratae-
grosseserratae
Quercus serrata var. Fagaceae PD Char. sp. of the Fagetum engleriano-
brevipetiolata lucidae
Quercus sessilifolia Fagaceae PE Char. sp. of the Yushania-Fagetum
hayatae
Quercus sp. Fagaceae PE Companion species with sporadical
presence (mainly China)
Quercus spinosa Fagaceae PE Char.-sp. of Euonymo porphyrei-
Fagetum englerianae
Quercus stenophylloides Fagaceae PE Companion species with sporadical
presence (mainly China)
Quercus stewardiana Fagaceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Reineckia carnea Liliaceae G rhiz Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Rhamnus costata Rhamnaceae NPD Diff.-sp. of Sasamorpho_Fagion
crenatae (subass.)
Rhamnus hemsleyana Rhamnaceae NPD Companion-species
Rhamnus japonica var. Rhamnaceae NPD Companion species with sporadical
decipiens presence (mainly Japan)
Rhaphiolepis indica Rosaceae NPD Char. sp. of the Sinarundinario
bashersuto-Fagetum lucidae
Rhododendron albrechtii Ericaceae NPD Char.-sp. of Saso kurilensis_Fagetum
crenatae
Rhododendron Ericaceae NPE Companion species with sporadical
anthopogonoides presence (mainly China)
Rhododendron augustinii Ericaceae NPE Diff. sp. of the Fagetum engleriano-
lucidae
(continued)
218 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Rhododendron Ericaceae NPE Diff.-sp. of Saso kurilensis-Fagetum
brachycarpum crenatae
Rhododendron Ericaceae NPE Diff.-sp. of Hepatico-Fagetum
brachycarpum var. roseum multinervis (subass.)
Rhododendron bricranthum Ericaceae NPE Char.-sp. of Abelio englerianae-
Fagion sp. div.
Rhododendron decandrum Ericaceae NPD Companion species with sporadical
presence (mainly Japan)
Rhododendron dilatatum Ericaceae NPD Companion species with sporadical
presence (mainly Japan)
Rhododendron fargesii Ericaceae PE Companion species with sporadical
presence (mainly China)
Rhododendron formosanum Ericaceae PE Char. sp. of the Yushania-Fagetum
hayatae
Rhododendron haofui Ericaceae PE Char. sp. of the Sinarundinario
bashersuto-Fagetum lucidae
Rhododendron Ericaceae PE Diff. sp. of the Fagetum engleriano-
hypoglancum lucidae
Rhododendron Ericaceae NPD Char. sp. of the Quingzheo
hypoglaucum tumidinodae-Fagion
Rhododendron kaempferi Ericaceae NPD Species of the Quercetalia serratae-
grosseserratae
Rhododendron lagopus Ericaceae NPD Diff.-sp. of Lindero umbellatae-
Fagetum crenatae
Rhododendron latoucheae Ericaceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Rhododendron leishanicum Ericaceae PE Diff. sp. of the Sinarundinario
chungii-Fagetum lucidae (subass.)
Rhododendron Ericaceae PE Char. sp. of the Prismatomerio
macrocarpum henryi—Lithocarpetum naiadari
Rhododendron mariesii Ericaceae NPD Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Rhododendron metternichii Ericaceae NPE Companion species with sporadical
presence (mainly Japan)
Rhododendron metternichii Ericaceae NPE Companion species with sporadical
var. hondoense presence (mainly Japan)
Rhododendron micranthum Ericaceae NPE Diff. sp. of the Fagetum engleriano-
lucidae
Rhododendron nudipes ssp. Ericaceae NPD Diff.-sp. of Lindero umbellatae-
niphophilum Fagetum crenatae
Rhododendron ovatum Ericaceae NPE Char. sp. of the Indocalamo latifolii—
Fagion hayatae var. zhejiangensis
Rhododendron Ericaceae NPD Companion species with sporadical
pentaphyllum presence (mainly Japan)
Rhododendron pseudo- Ericaceae PE Diff. sp. of the Yushania-Fagetum
chrysenthnum hayatae (subass.)
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 219
Life
Species name Family name form Ecological indication
Rhododendron Ericaceae NPD Char.-sp. of Sasamorpho_Fagion
quinquefolium crenatae
Rhododendron reticulatum Ericaceae NPD Companion species with sporadical
presence (mainly Japan)
Rhododendron reticulatum Ericaceae NPD Diff.-sp. of Lindero umbellatae-
var. ciliatum Fagetum crenatae
Rhododendron ririei Ericaceae PE Diff. sp. of the Sinarundinario
chungii-Fagetum lucidae (subass.)
Rhododendron rivulare Ericaceae NPE Companion species with sporadical
presence (mainly China)
Rhododendron Ericaceae PE Companion species with sporadical
rubropilosum presence (mainly China)
Rhododendron Ericaceae NPD Species of the Quercetalia serratae-
semibarbatum grosseserratae
Rhododendron seniavinii Ericaceae NPE1 Companion species with sporadical
presence (mainly China)
Rhododendron simsii Ericaceae NPD Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Rhododendron sp. Ericaceae PE Companion species with sporadical
presence (mainly China)
Rhododendron stamineum Ericaceae NPE Companion species with sporadical
presence (mainly China)
Rhododendron Ericaceae PE Companion-species
sutchuenense
Rhododendron wadanum Ericaceae NPD Companion-species
Rhododendron weyrichii Ericaceae NPD Companion species with sporadical
presence (mainly Japan)
Rhus ambigua Anacardiaceae PL Widespead in the Fagus-forests of
East Asia
Rhus chinensis Anacardiaceae PD Companion species with sporadical
presence (mainly China)
Rhus delavayi Anacardiaceae PD Companion species with sporadical
presence (mainly China)
Rhus punfabensis Anacardiaceae PD Diff. sp. of the Fagetum engleriano-
lucidae
Rhus sp. Anacardiaceae PD Companion species with sporadical
presence (mainly China)
Rhus trichocarpa Anacardiaceae PD Species of the Quercetalia serratae-
grosseserratae
Ribes longirucemosum var. Saxifragaceae NPD Diff. sp. of the Sinarundinario
davidii bashersuto-Fagetum lucidae (subass.)
Ribes sp. Saxifragaceae NPD Companion species with sporadical
presence (mainly China)
Rodgersia podophylla Saxifragaceae H Diff.-sp. of Saso kurilensis-Fagetum
scap crenatae
(continued)
220 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Rosa setipoda Rosaceae NPD Diff. sp. of the Fagetum engleriano-
lucidae
Rubia chinensis var. Rubiaceae G rad Diff.-sp. of Sapio japonici-Fagetum
glabrescens crenatae subass.)
Rubia cordifolia Rubiaceae H Diff. sp. of the Sinarundinario
scap chungii-Fagetum lucidae (subass.)
Rubia lanceolata Rubiaceae H Companion species with sporadical
scap presence (mainly China)
Rubia leiocaulis Rubiaceae H Companion species with sporadical
scap presence (mainly China)
Rubus alceaefolius Rosaceae Ch Char. sp. of the Sinarundinario
suffr bashersuto-Fagetum lucidae
Rubus amphidasya Rosaceae Ch Char. sp. of the Sinarundinario
suffr chungii-Fagetum lucidae
Rubus bambusarum Rosaceae NPD Char. sp. of Vaccinio henryi-Fagetum
pashanicae
Rubus buergeri Rosaceae H Char.-sp. of Litseo elongatae-Fagetea
scand
Rubus chroosepalus Rosaceae NPD Char. sp. of the Quingzheo
tumidinodae-Fagion
Rubus corchorifolius Rosaceae NPD Char. sp. of the Carici lanceolatae-
Fagetum hayatae
Rubus flosculosus Rosaceae NPD Companion species with sporadical
presence (mainly China)
Rubus illecebrosus Rosaceae Ch Companion species with sporadical
frut presence (mainly Japan)
Rubus irenaeus Rosaceae Ch Companion species with sporadical
suffr presence (mainly China)
Rubus malifolius Rosaceae Ch Char.-sp. of Sinarundinario nitidae-
suffr Fagetalia sp. div.
Rubus microphyllus Rosaceae NPD Companion species with sporadical
presence (mainly Japan)
Rubus pacificus Rosaceae NPD Char. sp. of the Sinarundinario
chungii-Fagetum lucidae
Rubus palmatus Rosaceae NPD Widespead in the Fagus-forests of
East Asia
Rubus palmatus var. Rosaceae NPD Companion-species
coptophyllus
Rubus pectinellus Rosaceae H Widespead in the Fagus-forests of
scand East Asia
Rubus peltatus Rosaceae NPD Diff.-sp. of Sapio japonici-Fagetum
crenatae subass.)
Rubus peseudo-japonicus Rosaceae NPD Diff.-sp. of Saso kurilensis-Fagetum
crenatae
Rubus pungens Rosaceae PL Char.-sp. of Euonymo porphyrei-
Fagetum englerianae
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 221
Life
Species name Family name form Ecological indication
Rubus shinkoensis Rosaceae NPD Char. sp. of the Yushania-Fagetum
hayatae
Rubus sp1. Rosaceae NPD Companion species with sporadical
presence (mainly China)
Rubus sp2. Rosaceae NPD Companion species with sporadical
presence (mainly China)
Rubus swinhoei Rosaceae NPD Companion-species
Rubus trianthus Rosaceae NPD Char. sp. of the Aceri davidii-Fagion
lucidae
Rumohra standishii Polypodiaceae H Diff.-sp. of Hepatico-Fagetum
rosul multinervis (subass.)
Salvia nipponica Lamiaceae H Companion-species
scap
Sambucus sieboldiana Caprifoliaceae NPD Companion-species
Sambucus sieboldiana var. Caprifoliaceae NPD Diff.-sp. of Saso kurilensis-Fagetum
miquelii crenatae
Sambucus williamsii var. Caprifoliaceae NPD Diff.-sp. of Hepatico-Fagetum
coreana multinervis (subass.)
Sanicula chinensis Apiaceae H Companion species with sporadical
scap presence (mainly Japan)
Sapium japonicum Euphorbiaceae PD Widespead in the Fagus-forests of
East Asia
Sarcandra glabra Chloranthaceae NPE Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Sarcandra hainanensis Chloranthaceae NPE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Sarcopyramis bodinieri Melastomataceae H Diff. sp. of the Sinarundinario nitidae-
scap Fagetum lucidae (subass.)
Sarcopyramis napalensis Melastomataceae H Companion species with sporadical
scap presence (mainly China)
Sargentodoxa cuneata Lardizabalaceae PL Companion species with sporadical
presence (mainly China)
Sasa hayatae Poaceae GS Diff.-sp. of Crnoo-Fagetum crenatae
(subass.)
Sasa ishizuchiana Poaceae GS Diff.-sp. of Sapio japonici-Fagetum
crenatae subass.)
Sasa kurilensis Poaceae GS Char.-sp. of Fagion crenatae
Sasa megalophylla Poaceae GS Companion species with sporadical
presence (mainly Japan)
Sasa nipponica Poaceae GS Companion-species
Sasa palmata Poaceae GS Char.-sp. of Fagion crenatae
Sasa ramosa Poaceae GS Companion species with sporadical
presence (mainly Japan)
Sasa senanensis Poaceae GS Char.-sp. of Saso-Fagetalia
Sasa tsuboiana Poaceae GS Companion species with sporadical
presence (mainly Japan)
(continued)
222 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Sasa tsukubensis Poaceae GS Diff.-sp. of Saso kurilensis-Fagetum
crenatae
Sasa veitchii var. Poaceae GS Diff.-sp. of Lindero umbellatae-
tyugokuensis Fagetum crenatae
Sasamorpha borealis Poaceae GS Char.-sp. of Sasamorpho_Fagion
crenatae
Saussurea cordifolia Asteraceae H Diff. sp. of the Fagetum engleriano-
scap lucidae
Saussurea grandifolia Asteraceae H Diff.-sp. of Hepatico-Fagetum
scap multinervis (subass.)
Saxifraga flabellifolia Saxifragaceae H Companion species with sporadical
rosul presence (mainly China)
Saxifraga fortunei var. Saxifragaceae H Companion species with sporadical
incisolobata rosul presence (mainly Japan)
Saxifraga fusca var. Saxifragaceae H Diff.-sp. of Sapio japonici-Fagetum
kikubuki rosul crenatae subass.)
Saxifraga sibirica Saxifragaceae H Companion species with sporadical
rosul presence (mainly China)
Saxifraga stolonifera Saxifragaceae H Char. sp. of the Elatostemo
rosul sessile_Fagetum lucidae
Schefflera bodinieri Araliaceae PE Char. sp. of the Athyrio nardii-
Michelietum balansae
Schefflera glomerulata Araliaceae PE Companion species with sporadical
presence (mainly China)
Schefflera producta Araliaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Schima argentea Theaceae PE Char. sp. of the Athyrio nardii-
Michelietum balansae
Schima crenata Theaceae PE Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Schima sinensis Theaceae PE Diff. sp. of the Sinarundinario
chungii-Fagetum lucidae (subass.)
Schima sp. Theaceae PE Companion species with sporadical
presence (mainly China)
Schima superba Theaceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Schima villosa Theaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Schisandra chinensis Magnoliaceae PL Widespead in the Fagus-forests of
East Asia
Schisandra henryi Magnoliaceae PL Char. sp. of the Carici lanceolatae-
Fagetum hayatae var. zhejiangensis
Schisandra incarnata Magnoliaceae PL Companion species with sporadical
presence (mainly China)
Schisandra repanda Magnoliaceae PL Species of the Quercetalia serratae-
grosseserratae
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 223
Life
Species name Family name form Ecological indication
Schisandra sinensis Magnoliaceae PL Companion-species
Schisandra sp. Magnoliaceae PL Companion species with sporadical
presence (mainly China)
Schisandra sphenanthera Magnoliaceae PL Companion-species
Schizophragma Saxifragaceae PL Char.-sp. of Fagetea crenatae
hydrangeoides
Schizophragma hypoleuca Saxifragaceae PL Char. sp. of the Viburno flavescentis-
Fagetum englerianae
Schizophragma integrifolia Saxifragaceae PL Companion species with sporadical
presence (mainly China)
Schizophragma Saxifragaceae PL Char. sp. of the Yushania-Fagetum
integrifolium var. hayatae
formosana
Schizophragma molle Saxifragaceae PL Companion species with sporadical
presence (mainly China)
Schizophragma sp. Saxifragaceae PL Companion species with sporadical
presence (mainly China)
Scrophularia duplicato- Scrophulariaceae H Companion species with sporadical
serrata scap presence (mainly Japan)
Scutellaria baicalensis Lamiaceae H Companion species with sporadical
scap presence (mainly China)
Scutellaria franchetiana Lamiaceae H Companion species with sporadical
scap presence (mainly China)
Scutellaria laeteviolacea Lamiaceae H Companion species with sporadical
scap presence (mainly Japan)
Selaginella labordei Selaginellaceae H Char. sp. of the Sinarundinario
caesp chungii-Fagetum lucidae
Selaginella remotifolia Selaginellaceae H Char. sp. of the Yushania-Fagetum
caesp hayatae
Selaginella sp. Selaginellaceae H Companion species with sporadical
caesp presence (mainly China)
Shortia soldanelloides Diapensiaceae H Companion species with sporadical
rosul presence (mainly Japan)
Shortia soldanelloides var. Diapensiaceae H Companion species with sporadical
magnus rosul presence (mainly Japan)
Shortia uniflora Diapensiaceae H Companion-species
rosul
Sinarundinaria bashersuta Poaceae GS Char. sp. of the Sinarundinario
bashersuto-Fagetum lucidae
Sinarundinaria chungii Poaceae GS Char. sp. of the Sinarundinario
chungii-Fagetum lucidae
Sinarundinaria nitida Poaceae GS Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Sinocalamus giganteus Poaceae GS Char. sp. of the Sinocalamus
giganteus-Fagetum lucidae
(continued)
224 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Skimmia arborescens Rutaceae NPE Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Skimmia japonica Rutaceae NPE Diff.-sp. of Sasamorpho_Fagion
crenatae (subass.)
Skimmia japonica f. rugosa Rutaceae NPE Diff.-sp. of Sasamorpho_Fagion
crenatae (subass.)
Skimmia japonica var. Rutaceae NPE Char.-sp. of Saso-Fagetalia
intermedia f. repens
Skimmia reevesiana Rutaceae NPE Widespead in the Fagus-forests of
East Asia
Sloanea elegans Elaeocarpaceae PE Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Smilacina glabra Liliaceae G rhiz Companion species with sporadical
presence (mainly China)
Smilacina hondoensis Liliaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Smilacina japonica Liliaceae G rhiz Char.-sp. of Saso-Fagetalia
Smilacina paniculata Liliaceae G rhiz Companion species with sporadical
presence (mainly China)
Smilacina sp. Liliaceae G rhiz Companion species with sporadical
presence (mainly China)
Smilacina yesoensis Liliaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Smilacina yunnanensis Liliaceae G rhiz Char. sp. of the Viburno flavescentis-
Fagetum englerianae
Smilax arisanensis Liliaceae PL Companion species with sporadical
presence (mainly China)
Smilax austro-zhejiangensis Liliaceae PL Char. sp. of the Carici lanceolatae-
Fagetum hayatae
Smilax backii Liliaceae PL Companion-species
Smilax china Liliaceae PL Companion-species
Smilax cocculoides Liliaceae PL Companion-species
Smilax discotis Liliaceae PL Char.-sp. of Litseo elongatae-Fagetea
Smilax elongato-reticullata Liliaceae PL Char. sp. of the Yushania-Fagetum
hayatae
Smilax elongato-umbelata Liliaceae PL Char. sp. of the Yushania-Fagetum
hayatae
Smilax ferox Liliaceae PL Companion species with sporadical
presence (mainly China)
Smilax glabra Liliaceae PL Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Smilax glanco-china Liliaceae PL Companion species with sporadical
presence (mainly China)
Smilax granulicaulis Liliaceae PL Char. sp. of the Ardisio hypargeriae—
Castanopsion fabrii
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 225
Life
Species name Family name form Ecological indication
Smilax lanceaefolia var. Liliaceae PL Widespead in the Fagus-forests of
opaca East Asia
Smilax lanceiofolia Liliaceae PL Companion species with sporadical
presence (mainly China)
Smilax lebrunii Liliaceae PL Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Smilax nervo-marginata Liliaceae PL Char. sp. of the Indocalamo latifolii—
Fagion hayatae var. zhejiangensis
Smilax nipponica Liliaceae PL Char.-sp. of Fagetea crenatae
Smilax opaca Liliaceae PL Char. sp. of the Viburno flavescentis-
Fagetum englerianae
Smilax polycorea Liliaceae PL Diff. sp. of the Fagetum engleriano-
lucidae
Smilax riparia var. Liliaceae PL Companion species with sporadical
ussuriensis presence (mainly Japan)
Smilax sieboldii Liliaceae PL Diff.-sp. of Sasamorpho_Fagion
crenatae (subass.)
Smilax sp. Liliaceae PL Companion species with sporadical
presence (mainly China)
Smilax stans Liliaceae PL Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Smilax trachypoda Liliaceae PL Diff. sp. of the Fagetum engleriano-
lucidae
Solidago virga-aurea var. Asteraceae H Companion-species
asiatica scap
Solidago virga-aurea var. Asteraceae H Char.-sp. of Fagion multinrvis and
gigantea scap Fagetalia multinervis
Solidago virga-aurea var. Asteraceae H Companion species with sporadical
leiocarpa scap presence (mainly Japan)
Sorbus alnifolia Rosaceae PD Widespead in the Fagus-forests of
East Asia
Sorbus aronioides Rosaceae PD Companion species with sporadical
presence (mainly China)
Sorbus caloneura Rosaceae PD Companion species with sporadical
presence (mainly China)
Sorbus commixta Rosaceae PD Char.-sp. of Fagetea crenatae
Sorbus coronata Rosaceae PD Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Sorbus folgneri Rosaceae PD Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Sorbus gracilis Rosaceae PD Companion species with sporadical
presence (mainly Japan)
Sorbus hemslaya Rosaceae PD Companion species with sporadical
presence (mainly China)
Sorbus japonica Rosaceae PD Species of the Quercetalia serratae-
grosseserratae
(continued)
226 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Sorbus rufopilosa Rosaceae PD Companion species with sporadical
presence (mainly China)
Sorbus sargentiana Rosaceae PD Char. sp. of the Viburno flavescentis-
Fagetum englerianae
Sorbus sp. Rosaceae PD Companion species with sporadical
presence (mainly China)
Sorbus xanthoneura Rosaceae PD Companion-species
Spatholirion longifolium Commelinaceae G rhiz Char. sp. of the Sinocalamus
giganteus-Fagetum lucidae
Spiraea chinensis Rosaceae NPD Char. sp. of the Carici lanceolatae-
Fagetum hayatae
Spiraea dasyantha Rosaceae NPD Diff. sp. of the Fagetum engleriano-
lucidae
Spiraea prunifolia Rosaceae NPD Diff. sp. of the Fagetum engleriano-
lucidae
Spuriopimpinella calycina Apiaceae H Diff.-sp. of Saso kurilensis-Fagetum
scap crenatae
Spuriopimpinella nikoensis Apiaceae H Companion species with sporadical
scap presence (mainly Japan)
Stachyurus himalaicus Stachyuraceae NPD Companion species with sporadical
presence (mainly China)
Stachyurus praecox Stachyuraceae NPD Companion species with sporadical
presence (mainly Japan)
Staphylea bumalda Staphyleaceae NPD Companion species with sporadical
presence (mainly Japan)
Stauntonia chinensis Lardizabalaceae PL Companion species with sporadical
presence (mainly China)
Stauntonia duclouxii Lardizabalaceae PL Companion species with sporadical
presence (mainly China)
Stauntonia purpurea Lardizabalaceae PL Char. sp. of the Yushania-Fagetum
hayatae
Stauntonia sp. Lardizabalaceae PL Companion species with sporadical
presence (mainly China)
Stellaria alsine Caryophyllaceae H Companion species with sporadical
scap presence (mainly China)
Stellaria diversiflora Caryophyllaceae H Companion species with sporadical
scap presence (mainly Japan)
Stellaria palustris Caryophyllaceae H Diff. sp. of the Fagetum engleriano-
scap lucidae
Stellaria sessiliflora Caryophyllaceae H Diff.-sp. of Crnoo-Fagetum crenatae
scap (subass.)
Stellaria wushanensis Caryophyllaceae H Diff. sp. of the Sinarundinario
scap chungii-Fagetum lucidae (subass.)
Stephanandra incisa Rosaceae NPD Companion-species
Stewartia monadelpha Theaceae PD Char.-sp. of Sasamorpho_Fagion
crenatae
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 227
Life
Species name Family name form Ecological indication
Stewartia pseudo-camellia Theaceae PD Char.-sp. of Saso-Fagetalia
Stewartia serrata Theaceae PD Diff.-sp. of Crnoo-Fagetum crenatae
(subass.)
Stewartia sinensis Theaceae PD Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Stewartia sp. Theaceae PD Companion species with sporadical
presence (mainly China)
Stranvaesia amphidoxa Rosaceae PE Char. sp. of the Quingzheo
tumidinodae-Fagion
Streptopus amplexifolius Liliaceae G rhiz Companion species with sporadical
var. papillatus presence (mainly Japan)
Streptopus streptoides Liliaceae G rhiz Companion species with sporadical
presence (mainly Japan)
Streptopus streptopoides Liliaceae G rhiz Char.-sp. of Saso kurilensis_Fagetum
var. japonicus crenatae
Struthiopteris castanea Polypodiaceae H Companion species with sporadical
rosul presence (mainly Japan)
Struthiopteris nipponica Polypodiaceae H Species of the Quercetalia serratae-
rosul grosseserratae
Styrax hemsleyana Styracaceae PD Diff. sp. of the Fagetum engleriano-
lucidae
Styrax japonica Styracaceae PD Widespead in the Fagus-forests of
East Asia
Styrax obassia Styracaceae PD Char.-sp. of Fagetea crenatae
Styrax rosea Styracaceae PD Companion species with sporadical
presence (mainly China)
Styrax shiraiana Styracaceae PD Char.-sp. of Sapio japonici-Fagetum
crenatae
Styrax sp. Styracaceae PD Companion species with sporadical
presence (mainly China)
Symplocos adenophylla Symplocaceae PE Char. sp. of the Athyrio nardii-
Michelietum balansae
Symplocos adenopus Symplocaceae PE Companion-species
Symplocos anomala Symplocaceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Symplocos botryantha Symplocaceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Symplocos caudata Symplocaceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Symplocos chinensis Symplocaceae NPD Companion species with sporadical
presence (mainly China)
Symplocos chinensis var. Symplocaceae PD Companion species with sporadical
leucocarpa f. pilosa presence (mainly Japan)
Symplocos cochinchinensis Symplocaceae PE Diff. sp. of the Yushania-Fagetum
subsp. laurina hayatae (subass.)
Symplocos coreana Symplocaceae PD Char.-sp. of Saso-Fagetalia
(continued)
228 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Symplocos crassifolia Symplocaceae PE Diff. sp. of the Sinarundinario nitidae-
Fagetum lucidae (subass.)
Symplocos discolor Symplocaceae PE Char. sp. of the Tripterospermo
cordifolium-Fagetum englerianae
Symplocos glandulifera Symplocaceae PE Char. sp. of the Ardisio hypargeriae—
Castanopsion fabrii
Symplocos glauca Symplocaceae PE Companion species with sporadical
presence (mainly China)
Symplocos groffii Symplocaceae PE Diff. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
(subass.)
Symplocos lancifolia Symplocaceae PE Char.-sp. of Litseo elongatae-Fagetea
Symplocos lancilimba Symplocaceae PE Diff. sp. of the Sinarundinario
chungii-Fagetum lucidae (subass.)
Symplocos lanrina Symplocaceae PE Companion-species
Symplocos lucida Symplocaceae PE Companion species with sporadical
presence (mainly China)
Symplocos macrostroma Symplocaceae PE Diff. sp. of the Yushania-Fagetum
hayatae (subass.)
Symplocos myrtacea Symplocaceae PE Companion species with sporadical
presence (mainly Japan)
Symplocos paniculata Symplocaceae NPD Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Symplocos ramosissima Symplocaceae PE Char. sp. of the Sinocalamus
giganteus-Fagetum lucidae
Symplocos sp. Symplocaceae PE Companion species with sporadical
presence (mainly China)
Symplocos stapfiana Symplocaceae PE Diff. sp. of the Sinarundinario nitidae-
Fagetum lucidae (subass.)
Symplocos stellaris Symplocaceae PE Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Symplocos sumuntia Symplocaceae PE Widespead in the Fagus-forests of
East Asia
Symplocos theaefolia Symplocaceae PE Companion species with sporadical
presence (mainly China)
Symplocos wikstroemiifolia Symplocaceae PE Companion species with sporadical
presence (mainly China)
Syneilesis palmata Asteraceae H Companion species with sporadical
scap presence (mainly Japan)
Synurus deltoides Asteraceae H Companion species with sporadical
scap presence (mainly China)
Synurus pungens Asteraceae H Diff.-sp. of Crnoo-Fagetum crenatae
scap (subass.)
Taxus cuspidata Taxaceae PEN Char.-sp. of Hepatico-Fagetum
multinervis
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 229
Life
Species name Family name form Ecological indication
Ternstroemia Theaceae PE Char. sp. of the Sinarundinario
kwangtungensis bashersuto-Fagetum lucidae
Ternstroemia luteoflora Theaceae PE Companion species with sporadical
presence (mainly China)
Tetracentron sinense Magnoliaceae PD Companion species with sporadical
presence (mainly China)
Tetrastigma hypoglaucum Vitaceae PL Companion species with sporadical
presence (mainly China)
Tetrastigma obtectum var. Vitaceae PL Companion species with sporadical
glabrum presence (mainly China)
Thalictrum minus var. Ranunculaceae H Diff.-sp. of Saso kurilensis-Fagetum
hypoleucum scap crenatae
Thalictrum tuberiferum Ranunculaceae H Diff.-sp. of Saso kurilensis-Fagetum
scap crenatae
Thalictrum uncatum Ranunculaceae H Companion species with sporadical
scap presence (mainly China)
Thladiantha glabra Cucurbitaceae H Companion species with sporadical
scap presence (mainly China)
Thujopsis dolabrata Cupressaceae PEN Companion species with sporadical
presence (mainly Japan)
Thujopsis dolabrata var. Cupressaceae PEN Companion species with sporadical
hondae presence (mainly Japan)
Tiarella polyphylla Saxifragaceae H Widespead in the Fagus-forests of
scap East Asia
Tilia insularis Tiliaceae PD Char.-sp. of Hepatico-Fagetum
multinervis
Tilia japonica Tiliaceae PD Char.-sp. of Saso-Fagetalia
Tilia maximowicziana Tiliaceae PD Companion species with sporadical
presence (mainly Japan)
Tilia oliveri Tiliaceae PD Companion species with sporadical
presence (mainly China)
Tilingia holopetala Apiaceae H Companion species with sporadical
scap presence (mainly Japan)
Toona ciliata Meliaceae PD Char. sp. of the Elatostemo
sessile_Fagetum lucidae
Toona sinensis Meliaceae PD Companion species with sporadical
presence (mainly China)
Torreya nucifera Taxaceae PEN Diff.-sp. of Sasamorpho_Fagion
crenatae (subass.)
Torreya nucifera var. Taxaceae NPEN Char.-sp. of Lindero umbellatae-
radicans Fagetum crenatae
Torricellia tillifolia Cornaceae PD Diff. sp. of the Sinarundinario
bashersuto-Fagetum lucidae (subass.)
Toxicodendron radicans Cornaceae PD Char. sp. of the Viburno flavescentis-
var. hispidus Fagetum englerianae
(continued)
230 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Toxicodendron Cornaceae PD Companion-species
succedaneum
Toxicodendron trichocarpa Cornaceae PD Char. sp. of the Indocalamo latifolii—
Fagion hayatae var. zhejiangensis
Toxicodendron vernicifluum Cornaceae PD Companion species with sporadical
presence (mainly China)
Trachelospermum Apocynaceae PL Companion species with sporadical
asiaticum var. intermedium presence (mainly Japan)
Trachelospermum Apocynaceae PL Companion species with sporadical
cathayanum presence (mainly China)
Trachelospermum Apocynaceae H Char. sp. of the Elatostemo
jasminoides scand sessile_Fagetum lucidae
Tricyrtis affinis Liliaceae H Companion-species
scap
Tricyrtis latifolia Liliaceae H Companion species with sporadical
scap presence (mainly Japan)
Tricyrtis macropoda Liliaceae H Companion-species
scap
Tricyrtis maculata Liliaceae H Companion species with sporadical
scap presence (mainly China)
Trillium smallii Liliaceae G rhiz Char.-sp. of Fagetea crenatae
Trillium tschonoskii Liliaceae G rhiz Companion-species
Tripterospermum affine Gentianaceae H Companion species with sporadical
scand presence (mainly China)
Tripterospermum chinense Gentianaceae H Char. sp. of the Indocalamo latifolii—
scand Fagion hayatae var. zhejiangensis
Tripterospermum cordatum Gentianaceae H Companion species with sporadical
scand presence (mainly China)
Tripterospermum Gentianaceae H Char. sp. of the Tripterospermo
cordifolium scand cordifolium-Fagetum englerianae
Tripterospermum Gentianaceae H Char.-sp. of Fagetea crenatae
japonicum scand
Tripterospermum Gentianaceae H Char. sp. of the Yushania-Fagetum
lanceolata scand hayatae
Tripterospermum sp. Gentianaceae H Companion species with sporadical
scand presence (mainly China)
Tripterospermum Gentianaceae H Companion species with sporadical
taiwanense scand presence (mainly China)
Tripterygium regelii Celastraceae PL Companion species with sporadical
presence (mainly Japan)
Trochodendron aralioides Trochodendraceae PE Char. sp. of the Yushania-Fagetum
hayatae
Tsuga chinensis Pinaceae PEN Diff. sp. of the Fagetum engleriano-
lucidae
Tsuga diversifolia Pinaceae PEN Companion species with sporadical
presence (mainly Japan)
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 231
Life
Species name Family name form Ecological indication
Tsuga sieboldii Pinaceae PEN Widespead in the Fagus-forests of
East Asia
Tupistra chinensis Liliaceae G rhiz Diff. sp. of the Fagetum engleriano-
lucidae
Tupistra tui Liliaceae G rhiz Companion species with sporadical
presence (mainly China)
Tupistra wattii Liliaceae G rhiz Companion species with sporadical
presence (mainly China)
Tylophora aristolochioides Asclepiadaceae H Companion species with sporadical
scand presence (mainly Japan)
Ulmus japonica Ulmaceae PD Companion species with sporadical
presence (mainly Japan)
Ulmus laciniata Ulmaceae PD Char.-sp. of Hepatico-Fagetum
multinervis
Vaccinium henryi Ericaceae NPD Char. sp. of Vaccinio henryi-Fagetum
pashanicae
Vaccinium hirtum Ericaceae NPD Companion-species
Vaccinium japonicum Ericaceae Ch Widespead in the Fagus-forests of
frut East Asia
Vaccinium japonicum var. Ericaceae Ch Companion species with sporadical
lasiostemon frut presence (mainly China)
Vaccinium oldhamii Ericaceae NPD Companion-species
Vaccinium smallii Ericaceae NPD Companion-species
Vaccinium smallii var. Ericaceae NPD Companion species with sporadical
glabrum presence (mainly Japan)
Vaccinium sp. Ericaceae NPD Companion species with sporadical
presence (mainly China)
Valeriana degeletiana Valerianaceae H Companion species with sporadical
scap presence (mainly Japan)
Veratrum grandiflorum Liliaceae G rhiz Diff.-sp. of Sapio japonici-Fagetum
crenatae subass.)
Veratrum schindleri Liliaceae G rhiz Companion species with sporadical
presence (mainly China)
Veronica cana var. Scrophulariaceae H Companion species with sporadical
miqueliana scap presence (mainly Japan)
Viburnum betulifolium Caprifoliaceae NPD Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Viburnum cordifolium Caprifoliaceae PD Companion species with sporadical
presence (mainly China)
Viburnum corymbiflorum Caprifoliaceae NPD Companion-species
Viburnum dilatatum Caprifoliaceae NPD Species of the Quercetalia serratae-
grosseserratae
Viburnum erosum Caprifoliaceae NPD Widespead in the Fagus-forests of
East Asia
Viburnum flavescens Caprifoliaceae NPD Char. sp. of the Viburno flavescentis-
Fagetum englerianae
(continued)
232 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Viburnum foetidum var. Caprifoliaceae NPE Char. sp. of the Sinarundinario
rectangulatum bashersuto-Fagetum lucidae
Viburnum furcatum Caprifoliaceae NPD Widespead in the Fagus-forests of
East Asia
Viburnum hengshenicum Caprifoliaceae NPD Char. sp. of the Carici lanceolatae-
Fagetum hayatae var. zhejiangensis
Viburnum ichangense Caprifoliaceae NPD Char. sp. of the Aceri davidii-Fagion
lucidae
Viburnum integrifolium Caprifoliaceae NPD Companion species with sporadical
presence (mainly China)
Viburnum luzonicum Caprifoliaceae NPD Char. sp. of the Yushania-Fagetum
hayatae
Viburnum phlebotrichum Caprifoliaceae NPD Species of the Quercetalia serratae-
grosseserratae
Viburnum plicatum var. Caprifoliaceae NPD Widespead in the Fagus-forests of
tomentosum East Asia
Viburnum rhytidophyllum Caprifoliaceae NPE Diff. sp. of the Fagetum engleriano-
lucidae
Viburnum setigerum Caprifoliaceae NPD Companion-species
Viburnum sieboldii Caprifoliaceae NPD Diff.-sp. of Crnoo-Fagetum crenatae
(subass.)
Viburnum sp. Caprifoliaceae NPD Companion species with sporadical
presence (mainly China)
Viburnum sympodiale Caprifoliaceae NPD Char.-sp. of Sinarundinario nitidae-
Fagetalia sp. div.
Viburnum taiwanianum Caprifoliaceae NPD Companion species with sporadical
presence (mainly China)
Viburnum urceolatum Caprifoliaceae NPD Widespead in the Fagus-forests of
East Asia
Viburnum urceolatum var. Caprifoliaceae NPD Companion-species
procumbens
Viburnum willeanum Caprifoliaceae NPD Char. sp. of the Quingzheo
tumidinodae-Fagion
Viburnum wrightii Caprifoliaceae NPD Char.-sp. of Fagetea crenatae
Viola acuminata Violaceae H Companion species with sporadical
scap presence (mainly Japan)
Viola arcuta Violaceae H Companion species with sporadical
scap presence (mainly China)
Viola biflora Violaceae H Companion species with sporadical
scap presence (mainly China)
Viola bissetii Violaceae H Companion-species
rosul
Viola boissieuana Violaceae H Companion-species
rosul
Viola brunneostipulosa Violaceae H Diff. sp. of the Sinarundinario
rosul chungii-Fagetum lucidae (subass.)
(continued)
Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia 233
Life
Species name Family name form Ecological indication
Viola cordifolia Violaceae H Companion species with sporadical
rosul presence (mainly China)
Viola davidii Violaceae H Char. sp. of the Aceri davidii-Fagion
rosul lucidae
Viola dissecta Violaceae H Companion species with sporadical
rosul presence (mainly Japan)
Viola eizanensis Violaceae H Diff.-sp. of Crnoo-Fagetum crenatae
rosul (subass.)
Viola formosana var. Violaceae H Char. sp. of the Yushania-Fagetum
formosana rosul hayatae
Viola grypoceras Violaceae H Widespead in the Fagus-forests of
scap East Asia
Viola hondoensis Violaceae H Char.-sp. of Hepatico-Fagetum
rosul multinervis
Viola kusanoana Violaceae H Char.-sp. of Fagion multinrvis and
scap Fagetalia multinervis
Viola principis Violaceae H Companion-species
rosul
Viola rossii Violaceae H Char. sp. of the Carici lanceolatae-
rosul Fagetum hayatae var. zhejiangensis
Viola schiensis Violaceae H Companion species with sporadical
rosul presence (mainly China)
Viola schneideri Violaceae H Char.-sp. of Sinarundinario nitidae-
rosul Fagetalia sp. div.
Viola selkirkii Violaceae H Widespead in the Fagus-forests of
rosul East Asia
Viola shikokiana Violaceae H Diff.-sp. of Sapio japonici-Fagetum
rosul crenatae subass.)
Viola sp1. Violaceae H Companion species with sporadical
rosul presence (mainly China)
Viola sp2. Violaceae H Companion species with sporadical
rosul presence (mainly China)
Viola takeshimana Violaceae H Diff.-sp. of Hepatico-Fagetum
rosul multinervis (subass.)
Viola vaginata Violaceae H Char.-sp. of Fagion crenatae
rosul
Viola violacea Violaceae H Companion species with sporadical
rosul presence (mainly Japan)
Viscum aibum var. Loranthaceae E Companion species with sporadical
coloratum presence (mainly Japan)
Vitis amurensis f. Vitaceae PL Companion species with sporadical
glabrescens presence (mainly Japan)
Vitis coignetiae Vitaceae PL Companion species with sporadical
presence (mainly Japan)
Vitis flexuosa Vitaceae PL Companion species with sporadical
presence (mainly China)
(continued)
234 Appendix 1. Alphabetic List—Species of the Beech Forests in East Asia
Life
Species name Family name form Ecological indication
Vitis sp. Vitaceae PL Companion species with sporadical
presence (mainly China)
Vittaria yunnanensis Vittariaceae G rhiz Char. sp. of the Prismatomerio
henryi—Lithocarpetum naiadari
Waldsteinia ternata Rosaceae H Diff.-sp. of Saso kurilensis-Fagetum
rosul crenatae
Weigela hortensis Caprifoliaceae NPD Companion species with sporadical
presence (mainly Japan)
Wisteria floribunda Fabaceae PL Companion species with sporadical
presence (mainly Japan)
Woodsia manchuriensis Polypodiaceae H Companion species with sporadical
rosul presence (mainly Japan)
Woodsia polystichoides Polypodiaceae E Companion species with sporadical
presence (mainly China)
Yushania nitakayamensis Lauraceae GS Char. sp. of the Yushania-Fagetum
hayatae
Zanthoxylum piperitum Rutaceae NPD Companion-species
Zanthoxylum scandens Rutaceae PD Companion species with sporadical
presence (mainly China)
Zanthoxylum schinifolium Rutaceae NPD Companion species with sporadical
presence (mainly China)
Zelkova serrata Ulmaceae PD Companion species with sporadical
presence (mainly Japan)
Zingiber mioga Zingiberaceae G rhiz Char. sp. of the Elatostemo
sessile_Fagetum lucidae
Appendix 2. Taxonomical Outlook on the Flora
of the East Asiatic Beech Forests
The present Appendix 2 is the complete list of all species (and infraspecific taxa)
mentioned in Table 3.1a, 3.1b, distributed among the 114 families of vascular
plants, which form the flora of the Esat Asiatic beech forests. The sequence of
families in general follows the recent concepts of the phylogenetical taxonomy, as
exposed in the Angiosperm Phylogeny Group Website, with minor differences in
order to maintain some units of frequent use in the special literature on the
vegetation of China and of Japan.
Pteridophyta
Lycopodiaceae Lycopodium clavatum Lycopodium serratum var.
longipetiolatum
Lycopodium obscurum Lycopodium serratum var. serratum
Lycopodium serratum
Selaginellaceae Selaginella labordei Selaginella sp. 1
Selaginella remotifolia Selaginella sp. 2
Ophioglossaceae Botrychium multifidum var. robustum
Osmundaceae Osmunda cinnamomea var. fokiensis
Osmunda japonica
Gleicheniaceae Diplopterygium glaucum Diplopterygium laevissimum
Polypodiaceae s. l. Acrophorus stipellatus Dryopteris maximowicziana
Adiantum pedatum Dryopteris monticola
Allantodia chinensis Dryopteris polylepis
Allantodia hirtipes Dryopteris sabaei
Allantodia metteniana Dryopteris saxifraga
Allantodia petri Dryopteris sp1.
Allantodia squamigera Dryopteris sp2.
Allantodia wichurae Dryopteris subtriangularis
Arachniodes chinensis Humata tyermanni
Arachniodes festina Lasianthus japonicus
Arachniodes mutica Lastrea quelpaertensis
Arachniodes pseudo-aristata Lepisorus bicolor
Arachniodes rhomboides Lepisorus contortus
Arachniodes standishii Lepisorus obscure-venulosus
Araiostegia parripinnata Lepisorus onoei
(continued)
T. Hukusima et al., Phytosociology of the Beech (Fagus) Forests in East Asia, 235
Geobotany Studies, DOI 10.1007/978-3-642-35620-9,
# Springer-Verlag Berlin Heidelberg 2013
236 Appendix 2. Taxonomical Outlook on the Flora of the East Asiatic Beech Forests
T. Hukusima et al., Phytosociology of the Beech (Fagus) Forests in East Asia, 255
Geobotany Studies, DOI 10.1007/978-3-642-35620-9,
# Springer-Verlag Berlin Heidelberg 2013
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Relevant Websites
Angiosperm Phylogeny Group Website: http://www.mobot.org/mobot/research/APweb/
Flora of China: http://www.efloras.org/flora_page.aspx?flora_id¼2
Flora of Taiwan: http://www.efloras.org/flora_page.aspx?flora_id¼101