Journal of Biogeography (J. Biogeogr.) (2006) 33, 1804–1819
ORIGINAL
ARTICLE
Climatic limits for the present distribution
of beech (Fagus L.) species in the world
Jingyun Fang1* and Martin J. Lechowicz2
1
Department of Ecology, College of
Environmental Sciences, and Centre for
Ecological Research & Education, Peking
University, Beijing 100871, China and 2Biology
Department, McGill University, 1205 Dr
Penfield Avenue, Montreal, Quebec, Canada
H3A 1B1
ABSTRACT
Aim Beech (Fagus L., Fagaceae) species are representative trees of temperate
deciduous broadleaf forests in the Northern Hemisphere. We focus on the
distributional limits of beech species, in particular on identifying climatic factors
associated with their present range limits.
Location Beech species occur in East Asia, Europe and West Asia, and North
America. We collated information on both the southern and northern range
limits and the lower and upper elevational limits for beech species in each region.
Methods In total, 292 lower/southern limit and 310 upper/northern limit sites
with available climatic data for all 11 extant beech species were collected by
reviewing the literature, and 13 climatic variables were estimated for each site
from climate normals at nearby stations. We used principal components analysis
(PCA) to detect climatic variables most strongly associated with the distribution
of beech species and to compare the climatic spaces for the different beech
species.
Results Statistics for thermal and moisture climatic conditions at the lower/
southern and upper/northern limits of all world beech species are presented. The
first two PCA components accounted for 70% and 68% of the overall variance in
lower/southern and upper/northern range limits, respectively. The first PCA axis
represented a thermal gradient, and the second a moisture gradient associated
with the world-wide distribution pattern of beech species. Among thermal
variables, growing season warmth was most important for beech distribution, but
winter low temperature (coldness and mean temperature for the coldest month)
and climatic continentality were also coupled with beech occurrence. The
moisture gradient, indicated by precipitation and moisture indices, showed
regional differences. American beech had the widest thermal range, Japanese
beeches the most narrow; European beeches occurred in the driest climate,
Japanese beeches the most humid. Climatic spaces for Chinese beech species were
between those of American and European species.
*Correspondence: Jingyun Fang, Department of
Ecology, College of Environmental Sciences,
Peking University, Beijing 100871, China.
E-mail: [email protected]
1804
Main conclusions The distributional limits of beech species were primarily
associated with thermal factors, but moisture regime also played a role. There
were some regional differences in the climatic correlates of distribution. The
growing season temperature regime was most important in explaining
distribution of Chinese beeches, whilst their northward distribution was mainly
limited by shortage of precipitation. In Japan, distribution limits of beech species
were correlated with summer temperature, but the local dominance of beech was
likely to be dependent on snowfall and winter low temperature. High summer
temperature was probably a limiting factor for southward extension of American
beech, while growing season warmth seemed critical for its northward
distribution. Although the present distribution of beech species corresponded
www.blackwellpublishing.com/jbi
doi:10.1111/j.1365-2699.2006.01533.x
ª 2006 The Authors
Journal compilation ª 2006 Blackwell Publishing Ltd
Climatic limits for world beech distribution
well to the contemporary climate in most areas, climatic factors could not
account for some distributions, e. g., that of F. mexicana compared to its close
relative F. grandifolia. It is likely that historical factors play a secondary role in
determining the present distribution of beech species. The lack of F. grandifolia
on the island of Newfoundland, Canada, may be due to inadequate growing
season warmth. Similarly, the northerly distribution of beech in Britain has not
reached its potential limit, perhaps due to insufficient time since deglaciation to
expand its range.
Keywords
Climatic index, climatic space, continentality, Fagus, growing season warmth,
precipitation, principal components analysis, range limit, temperate forest.
INTRODUCTION
Beech (Fagus L., Fagaceae) are among the most representative
trees in the temperate deciduous broadleaf forests of the
Northern Hemisphere (Shen, 1992; Denk, 2003). The genus
includes ten primary species and two minor segregates (Willis,
1966) broadly distributed in three isolated regions: East Asia,
Europe and West Asia, and North America. Six species occur
in East Asia (F. engleriana, F. longipetiolata, F. lucida and
F. hayatae in China and F. crenata and F. japonica in Japan)
(Horikawa, 1972; Editorial Committee for Flora of China,
1999), two in Europe and West Asia (F. sylvatica and
F. orientalis) (Jalas & Suominen, 1972–91), and one in North
America (F. grandifolia) (Little, 1965, 1979). Some studies have
recognized two additional beech species, one on a small island
off Korea: F. multinervis (Kim et al., 1986; Kim, 1988) and
another one in the north-eastern mountains of Mexico:
F. mexicana (e.g. Miranda & Sharp, 1950; Rzedowski, 1983;
Maycock, 1994; Peters, 1995). Fagus multinervis is a segregate
of F. engleriana in China (Okubo et al., 1988; Peters, 1992;
Shen, 1992; Denk, 2003); F. mexicana is a segregate of the
North American F. grandifolia. For the purposes of the present
study we accept the validity of both segregate species.
In East Asia, beech occurs primarily in mountain areas. All
Chinese beeches are restricted to remote subtropical/warmtemperate mountain areas, ranging from south China to the
Yangtze River (c. 33! N), and from the south-east coast of the
East China Sea to the eastern edge of the Tibetan Plateau
(Tsien et al., 1975; Wu, 1980; Hou, 1988; Hong & An, 1993;
Cao et al., 1995). Compared with the continuous distribution
of other Chinese beeches, F. hayatae is isolated in three very
limited mountain areas: north-east Taiwan, eastern Zhejiang
and north-west Sichuan. Unlike beech in other regions,
Chinese beeches rarely form pure stands but typically occur
mixed with various deciduous broad-leaved trees (Betula, Acer,
Liriodendron, Davidia, Tilia, Carpinus and Nyssa), evergreen
broad-leaved trees (Lithocarpus, Cyclobalanopsis, Manglietia
and Castanopsis) and evergreen needle-leaved (Tsuga) trees
(Wu, 1980; Hou, 1983, 1988; Hsieh, 1989; Cao, 1995; Fang
et al., 1996).
Two beech species, F. crenata and F. japonica, are native to
Japan. The former is distributed from Kyushu (c. 30.5! N) to
southern Hokkaido (c. 42.8! N), whereas the latter is limited
to the south of Iwate-ken (Horikawa, 1972; Miyawaki, 1980–
89). Beech forest in Japan falls into two broad forest types: the
Pacific-Ocean-side type that grows intermixed with many
other temperate tree species, and the Japan-Sea-side type
where beech is frequently dominant (Yamazaki, 1983; Maeda,
1991). Forests with F. multinervis are restricted to a small
island, Ulreung-do in South Korea, and they are more or less
similar in community composition and structure to Japanese
beech forests of the Japan-Sea-side type (Kim et al., 1986; Kim,
1988).
In North America, F. grandifolia is one of the most
widespread species among temperate trees, covering almost
all the temperate zone along the Appalachian Mountains from
the northern edge of the subtropical zone almost to the
southern edge of the boreal zone (USDA Forest Service, 1975).
A number of studies have shown large differences in community composition and structure in different climatic regions
(Braun, 1967; Barnes, 1991; Maycock, 1994). Based on
differences in geographical distribution and morphological
characters, three races of F. grandifolia (grey beech, white
beech, and red beech) have been identified (Camp, 1951;
Braun, 1967; Maycock, 1994) but these have never been given
species status.
Another beech species in North America, F. mexicana, is
found in only four montane localities in north-eastern Mexico
(Little, 1965; Rzedowski, 1983). Its community characters are
more or less similar to those of Chinese beech forests, usually
mixing with many species of Quercus, Magnolia, Acer and
Carya (Miranda & Sharp, 1950; Rzedowski, 1983; Peters, 1995;
Williams-Linera et al., 2000).
In Europe, F. sylvatica spreads from Sicily in southern Italy
(c. 37.7! N) to Bergen in south Norway (c. 60.7! N) (Jalas &
Suominen, 1972–91; Feoli & Lagonegro, 1982; Jahn, 1991); this
is the most widely distributed of all beech species. Fagus
orientalis replaces F. sylvatica in a small region of southeast
Europe and spreads into West Asia: northern Turkey, the
Caucasus and the Elburz Mountains of northern Iran, where
Journal of Biogeography 33, 1804–1819
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1805
J. Fang and M. J. Lechowicz
the climate is more or less continental (Jalas & Suominen,
1972–91; Davis, 1982).
The three regions where beech species occur are controlled
by different air masses and have various topographic settings. A
monsoon climate prevails in East Asia with cold and dry air
masses from Siberia predominant in winter and hot, humid
subtropical highs from the Pacific Ocean predominant in
summer. This causes abundant summer rainfall and high
temperature after a dry and cold winter (Arakawa, 1969; Shen,
1986). The Himalayan Range, higher to the west and lower
eastwards in mainland China, intensifies this climatic difference
between winter and summer (Chang, 1983; Fang et al., 1996)
on the continent in comparison to the Japanese archipelago.
In eastern North America, the climate is governed by two
major circulations: cold and dry air masses from the Arctic and
moist warm air masses from the southern tropical seas, and
therefore the climatic patterns resemble more or less those in
East Asia. However, there is no topographic barrier akin to the
Himalayas to block meridional air mass exchange (Lydolph,
1985).
The European and West Asian region has a narrower
seasonal cycle of temperatures and rainfall than the two other
regions. In Europe, the air masses resemble those of western
Northern America, but are much influenced by topography.
The east–west trend of European mountain ranges reduces
northward invasion of large subtropical bodies of warm air,
and the Mediterranean Sea also plays a role in the generation
and routes of cyclonic storms (Lydolph, 1985).
The floristic, climatic and topographic patterns of these
regions where beech occurs have attracted the attention of
previous investigators. For example, using Fagus pollen data
and monthly mean temperature, Huntley et al. (1989) studied
climatic control of beech distribution and abundance in
Europe and North America. Peters (1992) and Peters &
Poulson (1994) compared tree growth, and community
structure and dynamics of the world beech species. Maycock
(1994) documented detailed information on differences in
community composition between North American and
Japanese beeches. Iverson & Prasad (1998) used forest
inventory data to estimate the climate envelope for
F. grandifolia and predict its range extension under climate
change. Piovesan & Adams (2001) compared masting behaviours of beech from Europe, eastern North America and
Japan, and discussed their links to climatic variations. In
addition, some case studies on relationships between beech
distribution and climates have been conducted, but most are
restricted to single species in a region (e.g. Birks, 1989; Cao
et al., 1995; Sykes et al., 1996).
Taken together, the studies mentioned above provide useful
ecological comparisons among beech species in the three
regions where they occur, but many questions remain about
relationships between beech distribution and climate. For
example, how different are the geographical patterns shown in
the three separate regions where beech species occur? What
climatic factors control such patterns? Can contemporary
climate explain the present distribution of beech species?
1806
Answering these questions is not only important in biogeography, but also will provide a firmer basis for predicting the
effects of climate change on the future distribution of beech
species. Focusing on distribution limits (lower or southern and
upper or northern limits) of the world beech species, we
therefore compare their climatic spaces to detect possible
factors limiting their distributions.
DATA AND METHODS
Data sets for beech distribution
Data sets for beech distribution were assembled by reviewing
ecological, botanical and geographical journals, books and
reports to record the geographical location, lower and upper
elevational limits, and associated information (e.g. topography,
climate and community characteristics) for each site where a
beech species was reported to occur. Vertical distribution ranges
were reported especially frequently in East Asia, but less often in
Europe and North America where topographic relief at the range
limits of beech species is less extreme. We assembled data for
F. grandifolia from a silvicultural atlas (USDA Forest Service,
1975) and ancillary literature reports. We found relatively little
data for F. sylvatica and F. orientalis; the map of Jalas &
Suominen (1972–91) yielded data on northern range limits for
these species, but high relief and uncertain elevational distributions left the southern limits undefined.
When latitude and longitude were not reported for study
locations, we used gazetteers to georeference the sites: (1)
China Places Name (Anon., 1986) and Atlas of Land Use in
China (Editorial Committee of 1/1,000,000 Land-use Map of
China, 1990) for Chinese beeches; (2) Gazetteer to AMS
1 : 250,000 Maps of Japan (Corps of Engineers, US Army,
1956) and The National Atlas of Japan (Geographical Survey
Institute, 1977) for Japanese beeches; (3) American Places
Dictionary (Abate, 1994) for American beech; and (4) Atlas of
the World (Times, 1992) for others. In total we identified 350
sites with reliable data for the lower/southern limits and 353
sites for the upper/northern limits of beech species (Table 1;
Fang, 2003). For detailed information on the location and
elevation of distribution limits for each beech species, see
Appendix S1 in Supplementary Material.
Climatic data
We used monthly mean temperatures and monthly precipitation records from the following data sources: (1) China
Meteorological Agency (1984); (2) Japan Meteorological
Agency (1972); (3) Central Meteorological Office of Korea
(1972); (4) National Climatic Center, NOAA (1983) for USA;
(5) Atmospheric Environmental Service, Environment Canada
(1982) for Canada; and (6) Wernstedt (1972) for Mexico,
Europe and West Asia. The period of record for most stations
was 1951–80 or 1941–71.
Temperatures at altitudes of the upper and lower
elevation limits of beech were estimated for each locality
Journal of Biogeography 33, 1804–1819
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Climatic limits for world beech distribution
Table 1 Sample size for distribution limits of all world beech
species. Data set for Fagus grandifolia was mainly extracted from
its southern and northern distributional edges, and the northern
limit of F. sylvatica actually lies to the north-east according to the
map of Jalas & Suominen (1972–91)
Species
Distribution
F.
F.
F.
F.
Pacific Ocean side, Japan
Japanese Sea side, Japan
South and southwest China
Taiwan, Zhejiang,
and Sichuan, China
South, east, and
southwest China
East, and Central China
Ulreung-do, South Korea
Eastern North America
Northeastern Mexico
Europe
Southeast Europe
and West Asia
crenata
japonica
engleriana
hayatae
F. longipetiolata
F.
F.
F.
F.
F.
F.
lucida
multinervis
grandifolia*
mexicana
sylvatica!
orientalis
Total
Lower
limit
Upper
limit
42
28
34
8
46
22
36
10
55
55
60
1
80
4
32
6
54
1
80
4
40
5
350
353
*Most data for southern or northern extremes.
!Most data for northern edge.
Climatic parameters
The distribution limits of many tree species are closely related
to growing season temperature (e.g. Kira, 1945, 1991; Holdridge, 1947; Tuhkanen, 1980; Woodward, 1987; Prentice et al.,
1992; Sykes et al., 1996). We used Kira’s Warmth Index (WI)
(Kira, 1945, 1991) and Holdridge’s annual biotemperature
(ABT) (Holdridge, 1947) as proxies for growing season
warmth, given respectively by:
X
WI ¼
ðT # 5Þ ðfor months in which T > 5 % CÞ
ð1Þ
T
ðfor months in which 0 < T < 30 % CÞ
12
Annual mean temperature (AMT) and mean temperature for
the warmest month (MTWM) are often used to explain
northward and upward distribution of northern tree species
(Walter, 1979; Tuhkanen, 1980; Ohsawa, 1990, 1991).
Climatic continentality
Wolfe (1979), Tuhkanen (1980), Ohsawa (1990) and Fang
et al. (1996) have addressed the effect of climatic continentality
on distribution of some tree species. We used the annual range
of monthly mean temperatures (ART) and Gorcynski’s (1922)
continentality index (K):
!
"
R
K ¼ 1:7 &
# 20:4
ð4Þ
sin L
where R is the annual range of monthly mean temperature in
!C and L is the latitude in degrees.
Growing season warmth
P
A number of studies have shown the importance of minimum
winter temperatures in controlling distributional limits of
plant species (e.g. Sakai & Weiser, 1973; Woodward, 1987;
Sykes et al., 1996; Pederson et al., 2004). Mean temperature for
the coldest month (MTCM) is often used as a surrogate for the
minimum winter temperature (Solomon, 1986; Ohsawa, 1990;
Prentice et al., 1992; Sykes et al., 1996). In this study, the
coefficient of determination (R2) between these two variables
was 0.94 (P < 0.0001, n ¼ 602), so we also used the more
readily available MTCM as a measure of coldness.
Previous studies also suggest that cumulative winter temperature is important for the northward/upward distributions
of warm-temperate and tropical tree species (Kira, 1948, 1991;
Hattori & Nakanishi, 1985; Fang & Yoda, 1991), and for spring
budburst of many northern tree species (Prentice et al., 1992;
Lechowicz, 2001). We used Kira’s Coldness Index (CI) (Kira,
1948, 1991) to express the cumulative winter temperature:
X
CI ¼ #
ð5 # TÞ ðfor months in which T < 5 % CÞ
ð3Þ
Annual mean temperature and mean temperature for the
warmest month
using a mean lapse rate of 0.6 !C per 100 m (Barry, 1992)
together with data from the nearest climatic station. The
rainfall data for each beech site were estimated from
relationships between precipitation and elevation regressed
by using rainfall and altitude data at more than five climatic
stations close to the beech site. Similar to Huntley et al.
(1989), distances between beech sites and climatic stations
were less than 0.5! of latitude and 1.0! of longitude (c. 50–
100 km radius). In total, we secured reliable climate data for
292 sites at the lower/southern limit and 310 sites at the
upper/northern limit of beech species.
ABT ¼
Coldness and winter temperature summation
ð2Þ
Moisture variables
To assess moisture regime, we considered annual precipitation (AP, mm), potential evapotranspiration (PET, mm),
annual actual evapotranspiration (AAE, mm), moisture
index (Im), and the Ellenberg quotient (EQ, !C mm)1).
With the exception of AP and EQ, all moisture parameters
were estimated by the Thornthwaite (1948) method, which
uses two climatic variables commonly recorded at climatic
stations around the world: monthly mean temperature and
monthly precipitation. The Thornthwaite index has proven a
good correlate of vegetation and plant distribution at both
Journal of Biogeography 33, 1804–1819
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1807
J. Fang and M. J. Lechowicz
regional and global scales (e.g. Mather & Yoshioka, 1968;
Fang & Yoda, 1990; O’Brien, 1993; Frank & Inouye, 1994).
For calculation of PET, AA and Im, see Fang (1989) and
Fang & Yoda (1990).
The EQ is the ratio of the MTWM to annual precipitation
and is frequently used to show the climatic limit of beech in
Europe (Ellenberg, 1986; Jahn, 1991). According to Jahn
(1991), values below 20 show a pure ‘beech climate’; the
competitive vigour of beech slowly decreases with an increase
from 20 to 30; and in Europe beech disappears in regions with
an EQ over 30:
EQ ¼
Warmest month’s mean temperature in % C
& 1000
Annual precipitation ðmmÞ
ð5Þ
Principal components analysis
We used principal components analysis (PCA) (Wilks, 1995)
to: (1) examine which climatic variables are most associated
with the distribution of beech species, and (2) compare the
climatic space at distributional limits for beech species in the
three regions.
To assess the influence of the diverse climatic variables, we
compared the eigenvalues of different PCA axes for each
species and the axis loadings of climatic variables for the
species that have a large sample size. In general, variables with
bigger loadings are considered more important in placement
along PCA axes when variables are standardized to allow for
differences in units and magnitude (Wilks, 1995). We analysed
13 climatic variables (AMT, WI, CI, ABT, MTWM, MTCM,
ART, K, AP, PET, AAE, Im and EQ) using PC-ORD software
(McCune & Mefford, 1999). To minimize the differences
associated with differing units and magnitude, all the variables
were standardized:
xi0 ¼
xi # !x
r
ð6Þ
where xi and xi0 are the original and standardized value of a
climatic variable for the ith site, !x is average of the climatic
variable for all the sites, and r is the standard deviation of the
climatic variable. To compare climate spaces among the beech
species, we used the eigenvalues of the first two components
for each beech site. To illustrate the climate space for species
with sufficiently large sample sizes, we calculated a 50%
Gaussian bivariate confidence ellipse (ELL) using sysgraph
(SYSTAT Inc., 1996).
RESULTS
Climatic statistics at the distribution limits
Table 2 lists the average for thermal and moisture climatic
parameters at lower/southern and upper/northern limits for
all beech species; the details for the statistics (average, SD and
range) of these variables are displayed in Table S1 in the
1808
Supplementary Material. The descriptions that follow are
supported by both Table 2 and Table S1.
In East Asia, in spite of large fluctuations within and among
species, average values of growing season warmth at the lower
limits of beech distribution were between 74.8 !C (F. crenata)
and 115.9 !CÆmonth (F. longipetiolata) for Warmth Index
(WI) and between 9.8 and 14.3 !C for annual biotemperature
(ABT) (Table 2). Interestingly, most of the seven Asian beech
species had lower limits associated with very similar average
thermal parameters (AMT of 11.7–12.8 !C, WI of 91.8–
100.6 !CÆmonth, ABT of 11.7–12.8 !C, MTWM of 22.4–
23.9 !C and MTCM of 0.1–2.1 !C) even though the
species have widely disparate ranges (F. japonica in Japan,
F. multinervis in Korea and others in China). Only two species,
F. crenata from colder climates and F. longipetiolata in warmer
regions, are exceptions. The thermal variables at the upper
limit of East Asian beeches showed smaller fluctuation for
most species with an AMT value of 7.3–9.2 !C, WI of 56.0–
70.3 !CÆmonth, and ABT of 8.1–9.5 !C (Table 2), but
F. crenata is an exception. Fagus crenata has a WI value
of 45.1 !CÆmonth, approximately the climatic threshold
(45 !CÆmonth) of the cool-temperate zone defined by Kira
(1991).
With regard to moisture regime, all beech species locations
in East Asia fall within an average Im range of 61.4–189.9 for
their lower limits and 70–241.3 for their upper limits, denoting
a humid or perhumid climate according to the Thornthwaite
(1948) system (Table 2).
In North America, F. grandifolia showed the widest climatic
space, with average WI ranging from 50.7 !CÆmonth (northern
limit) to 173.4 !CÆmonth (southern limit), and ABT from 7 to
19.5 !C. This spans two bioclimatic zones: cool-temperate
(45–85 !CÆmonth for WI and 6–12 !C for ABT) and warmtemperate zone (85–180 !CÆmonth for WI and over 12 !C for
ABT). Although F. mexicana is distributed much farther south
than F. grandifolia, it has a smaller climatic range (117.1–
127.3 !CÆmonth in WI and 14.8–15.6 !C in ABT) and much
smaller thermal variables at its lower limit than those of
F. grandifolia (Table 2). Fagus grandifolia has more moist
conditions at its northern limit with an average Im value of
91.1 compared with 40.3 at its southern limit. Mexican beech
has a much larger Im average value (Im ¼ 103) at its
elevational and southern limit.
In Europe, the climatic spaces of beech species span almost
the entire temperate zone. Growing season warmth in the
range of beech ranged from 47.7 to 104.3 !CÆmonth for
WI and 7.2–13.5 !C for ABT for F. sylvatica, and 46.3–
78.3 !CÆmonth for WI and 7.1–10.4 !C for ABT for F. orientalis
(Table 2). It is noteworthy that F. sylvatica has a rather narrow
climatic range despite having the widest latitudinal range
among the world beech species (spanning c. 23!) (Jalas &
Suominen, 1972–91). In comparison with beech species in
other regions, the moisture regime in the area of European
beech species is relatively dry, with average precipitation of
905.9 mm, mean Im value of 38, and mean EQ of 29 !C mm)1
for the lower limit of F. sylvatica, and 1272.3 mm, 119.3 and
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Journal of Biogeography 33, 1804–1819
ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd
F. orientalis
F. sylvatica
F. mexicana
F. grandifolia
F. multinervis
F. japonica
F. crenata
F. lucida
F. longipetiolata
F. hayatae
F. engleriana
Species
Lower
Upper
Lower
Upper
Lower
Upper
Lower
Upper
Lower
Upper
Lower
Upper
Lower
Upper
Lower
Upper
Lower
Upper
Lower
Upper
Lower
Upper
Lower/upper
limits
12.4
7.3
12.8
9.2
14.3
8.9
12.7
8.8
9.5
5.0
11.7
9.0
11.5
7.6
19.5
4.2
15.6
14.8
13.5
6.6
10.2
6.5
AMT
(!C)
99.0
56.0
99.4
67.0
115.9
66.2
100.6
66.4
74.8
45.1
91.8
70.3
90.2
59.0
173.4
50.7
127.3
117.1
104.3
47.7
78.3
46.3
WI
(!CÆmonth)
ABT
(!C)
12.5
8.1
12.8
9.5
14.3
9.2
12.8
9.2
9.8
6.7
11.7
9.4
11.5
8.3
19.5
7.0
15.6
14.8
13.5
7.2
10.4
7.1
CI
(!CÆmonth)
)10.1
)28.1
)6.4
)16.6
)4.1
)19.8
)8.2
)21.3
)21.3
)45.2
)12.0
)21.9
)11.9
)27.5
0
)60.5
0
0
)2.7
)28.3
)16.2
)28.5
23.0
17.8
22.4
18.6
24.2
18.9
23.0
19.1
21.8
17.6
23.9
21.2
23.4
19.5
27.5
18.4
19.6
18.7
23.0
16.9
20.5
16.1
MTWM
(!C)
1.1
)3.9
2.1
)0.7
3.5
)2.0
1.5
)2.4
)2.1
)6.8
0.3
)2.2
0.1
)3.8
10.4
)11.4
10.5
9.7
4.7
)2.7
)1.5
)3.2
MTCM
(!C)
21.9
21.7
20.3
19.3
20.8
20.9
21.5
21.5
23.9
24.4
23.7
23.4
23.3
23.3
17.1
29.8
9.1
9.1
18.2
19.6
22.0
19.3
ART
(!C)
53.3
52.8
50.7
46.3
54.1
54.2
57.8
57.8
48.5
48.8
48.8
48.6
45.0
45.0
36.8
49.5
22.2
22.2
27.7
25.8
38.1
31.5
K
1230
1366
1670
2233
1421
1738
1419
1757
2047
2660
1805
2329
1485
–
1426
1021
1741
–
906
1272
745
912
AP
(mm)
805.4
795.6
914.6
887.0
845.4
621.2
874.0
674.4
706.2
606.8
746.6
716.4
702.6
–
1024.0
537.8
949.0
–
749.8
577.6
717.2
668.9
PET
(mm)
803.3
793.5
914.6
887.0
831.8
619.2
862.7
671.7
706.2
606.8
746.6
716.4
702.6
–
996.9
530.5
906.0
–
497.1
496.7
486.5
460.3
AAE
(mm)
61.4
70.0
85.5
109.8
77.5
132.6
70.0
168.5
189.9
241.3
142.0
206.5
111.3
–
40.3
91.1
103.1
–
38.1
119.3
15.4
23.8
Im
21.7
19.4
17.0
13.2
19.0
14.7
19.5
13.8
12.5
9.5
15.3
11.5
16.1
–
19.5
18.5
14.5
–
29.0
16.8
32.7
27.9
EQ
(!C mm)1)
Table 2 Average for climatic variables at lower (southern) and upper (northern) limits of distribution for world beech (Fagus) species. AMT: annual mean temperature; WI: warmth index; CI:
coldness index; ABT: annual biotemperature; MTWM: mean temperature for the warmest month; MTCM: mean temperature for the coldest month; ART: annual range of mean temperature; K:
continentality index; AP: annual precipitation; PET: annual potential evapotranspiration; AAE: actual annual evapotranspiration; Im: moisture index; and EQ: Ellenberg quotient. – indicates no
estimation
Climatic limits for world beech distribution
1809
J. Fang and M. J. Lechowicz
16.8 !C mm)1 for AP, Im and EQ for its upper/northern limit.
For F. orientalis, AP, Im and EQ were, respectively, 744.8 mm
and 32.7 !C mm)1 for its lower limits, and 911.8 mm, 23.8
and 27.9 !C mm)1 for its upper limit. It is apparent that
climate at the lower limits of both beeches is rather dry, close
to the EQ threshold of 30 !C mm)1 suggested by Jahn (1991).
The EQ value of European beeches is much smaller than for
beech species in other regions: ranging from 15 to 20 !C mm)1
for the lower/southern limit and from 11.5 to 19.4 !C mm)1
for the upper/northern limit for other regions, with the
smallest value (12.5 and 9.5 !C mm)1 for its lower and upper
limits, respectively) for F. crenata (Table 2).
Climatic factors controlling beech species
distributions
We used the 13 climatic variables in a PCA to identify limiting
factors for beech distribution. To assess regional variations in
the relationships between geographic distribution and climatic
space, we combined four Chinese beeches and two European
beeches into the same group because they have similar climate
ranges (cf. Table S1). The two Japanese beeches occur
primarily in two different climatic regions (Japan Sea side
and Pacific Ocean side), and thus we did not deal with them as
a single group. The first two principal components account for
70% and 68% of overall variance for southern/lower and
northern/upper limits of all species, respectively (Table 3).
Accordingly we considered the loadings on these axes to be
most important in delimiting beech distribution. For each
beech region, the first two PCA axes explained more than
70% of the variance; for example, 80% and 87% for the
southern and northern limits of Amerian beech, respectively,
and 73–80% for F. japonica and the European beeches.
The first PCA axis represents a thermal gradient, and the
second a moisture gradient in the overall distribution of world
beech species (Table S2 in Supplementary Material). In general
the thermal climate played a leading role and precipitation a
secondary role in controlling the large-scale distribution of
Table 3 Proportion (%) of cumulative variance on the first four
principal components in a principal components analysis of the
distribution limits for world beech (Fagus) species
Species
Chinese
beeches
F. crenata
F. japonica
F. grandifolia
European
beeches
All beech
species
1810
Lower (southern) limit
Upper (northern) limit
PCA
1
PCA
1
PCA
2
PCA
3
PCA
4
PCA
2
PCA
3
PCA
4
46.34 69.89 83.71 96.69 41.19 73.56 89.04 96.12
37.60
47.98
59.25
43.65
71.86
72.96
79.76
76.86
90.25
88.70
91.69
87.73
97.27
98.34
98.77
94.73
50.48
49.27
54.46
40.45
74.57
80.20
86.65
71.79
88.08
90.85
96.99
88.01
96.49
97.71
99.12
97.26
50.71 69.89 84.98 94.28 44.89 67.57 83.72 93.79
beech species. Among thermal variables, AMT, WI and ABT
always showed larger loadings at both lower/southern and
upper/northern limits, but CI and MTCM exhibited a large
value for world beech species (Table S2). This indicates that
growing season warmth is most important for beech distribution, and CI and MTCM are also closely coupled with the
potential for their range expansion. The second PCA axis
exhibited a different trend: for the lower/southern limit the
loadings of AP, Im and EQ were 0.83 mm, 0.84 and
)0.89 !C mm)1, while for the upper limit/northern they were
)0.83 mm, )0.92 and 0.89 !C mm)1, respectively (Table S2).
The negative loadings of AP and Im indicate that altitude
limits for beech species are negatively correlated with moisture
climate, implying that the altitude of the upper elevational
limit decreases with an increase of precipitation.
In spite of these overall trends, some component loadings
varied across regions, suggesting that limiting climatic factors
shifted somewhat among beech in different regions. In general,
both the lower/southern and upper/northern limits in all three
beech regions depended on seasonal thermal regimes (loadings
of growing season warmth (WI and ABT) on the first PCA axis
were largest, and winter temperatures (CI and MTCM) also
showed a large influence), but the two Japanese beech species
were an exception. For the second PCA axis, the loadings of
moisture climate variables were largest for the lower/southern
limit, indicating the general influence of a moisture gradient.
However, for the upper/northern limit, the climatic variables
with the largest loading showed a large regional difference. The
second axis for Chinese beech suggests a moisture regime
gradient, while the loading of winter coldness (CI) was largest
for European beeches, and MTCM had the largest loading for
American beech.
For the lower limit of the two Japanese beeches, winter
temperature (CI and MTCM) had a large loading on the first
axis, summer temperature (MTWT) on the second. This
suggests that moisture regime is not a limiting factor for the
downward distribution of these two species due to abundant
precipitation in Japan, which agrees with other studies (e.g.
Maeda, 1991). For the upper limit of these two species, on the
second axis, the loadings of PET and AAE for F. crenata and of
MTWT and moisture indices (Im and EQ) for F. japonica were
largest.
It is noteworthy that PET, an indicator of total solar energy,
showed the largest loading (0.98) on the first axis for the
northern limit of American beech; this is consistent with the
correlation between PET and vegetation distribution and
overall tree species richness in North America (Stephenson,
1990; Francis & Currie, 2003).
DISCUSSION
Zonal distribution of world beech species
Although most beech species are considered typical trees of the
temperate zone, they in fact showed different climatic ranges in
the three regions where they occur (Table 2; Table S1). In East
Journal of Biogeography 33, 1804–1819
ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd
Climatic limits for world beech distribution
Asia, most species, excluding F. crenata, concentrated within a
climatic range of 90–116 !CÆmonth in WI and 11.7–14.3 !C in
ABT in their lower limits, and 56–70 !CÆmonth in WI and
c. 8.1–9.5 !C in ABT in their upper limits. This is to say, the
boundaries for most Asian beech species are located in warmer
places than the cool-temperate zone defined by Kira (1945,
1991) and Holdridge (1947); the climatic parameters at the
southern/lower edge of the cool-temperate zone was set at
85 !CÆmonth in WI by Kira, and 12 !C in ABT by Holdridge,
and 45 !CÆmonth in WI and 8 !C in ABT at the northern or
upper limit. This suggests that beech species occupy an ecotone
between cool-temperate deciduous broadleaf forest (a WI
range of 45–85 !CÆmonth) and warm-temperate evergreen
broadleaf forest (WI > 85 !CÆmonth) (Kira, 1991). However,
the WI value at the upper limit of F. crenata in Japan
(45.1 !CÆmonth) coincided well with Kira’s criterion of 45 !C
month; this supports the idea that F. crenata is an indicator of
the Japanese temperate zone (e.g. Miyawaki, 1980–89).
American beech occupies a large climatic space, with a range
of 50.7–173.4 !CÆmonth in WI and 7.0–19.5 !C in ABT
(Table S1). This spans two bioclimatic zones: the cooltemperate zone [45–85 !CÆmonth for WI (Kira, 1945, 1991)
and 6–12 !C for ABT (Holdridge, 1947)], and the warmtemperate zone (85–180 !C month for WI and > 12 !C for
ABT).
In Europe, climatic parameters at the northern/upper limit
of beech distribution indicated good agreement with criteria
defining the cool-temperate zone: a WI value of 47.7–
104.3 !CÆmonth, and an ABT value of 7.2–13.5 !C for
F. sylvatica, and 46.3–78.3 !CÆmonth and 7.1–10.4 !C for
F. orientalis (Table S1). On the other hand, an average Im
greater than 15.4 for all beech sites suggest a perhumid or
humid climate, defined by Thornthwaite (1948) as Im values of
20–100 and > 100, respectively.
Climates and present distribution of East Asian
beeches
As shown in Table 3, the first two PCA components account
for more than 70% of the variance in parameters associated
with the distribution of Asian beech species. No one variable
alone can explain beech distributional patterns; growing season
warmth (WI and ABT), winter low temperature (CI) and
annual mean temperature (AMT) all showed almost equal
loadings (Table S2). Thermal regime is clearly paramount in
the relationships between beech distribution and climatic
factors in East Asia, but the strong correlation among different
climatic elements (see Table S3 in Supplementary Material)
precludes identification of a single, dominant aspect of thermal
regime that affects the distribution of East Asian beech species.
There are a few viewpoints on the relationships between the
present distribution of Chinese beech species and limiting
factors. Hong & An (1993) pointed out that the climatic
factors affecting beech distribution varied from place by place;
for example, in northern regions, coldness and short growing
season were major limiting factors, whereas water deficit was
more important for southward migration. Cao et al. (1995)
demonstrated the importance of the moisture deficit in the
northern range of beech species and the importance of high
temperature and insufficient water supply in the south.
Focusing on the relationship between Fagus- and Tsugadominated forests, Fang et al. (1996) suggested possible effects
of the annual temperature range (ART) and high winter
temperature on beech distribution. They found that the beechdominated forests did not appear in places where hemlock
dominates, and that their boundary was consistent with an
ART isotherm of 23 !C; beech-dominated forests lay north of
this isotherm and hemlock-dominated forests lay south. This
implies the importance of high winter temperatures in
determining the distribution of Chinese beech because the
ART is closely correlated with winter temperature in southern
mountain areas in China.
Two effects of high winter temperatures that can influence
the distribution of temperate tree species may be playing a role
in beech distribution in China. First, temperate trees require a
sufficient period of winter cold (a period of chilling) before
warming will induce budburst in spring (Cannell & Smith,
1986; Lechowicz, 2001). Second, higher winter temperatures
can reduce the competitive ability of deciduous trees with
more warmth tolerant evergreen broadleaf trees (Woodward,
1987). These effects of winter temperature may explain an
unresolved question in Asian biogeography: why beech species
do not spread westward into the Himalayas and south-eastern
Tibet where growing season warmth and precipitation appear
satisfactory. Although Asian beech and hemlock have similar
heat requirements during the growing season (Liu & Qiu,
1980; Hou, 1983; Fang et al., 1996), hemlock is favoured by a
warm-winter climate (Sakai, 1975). While Chinese beeches
co-exist with evergreen broad-leaved tree species in genera
such as Lithocarpus, Cyclobalanopsis and Castanopsis (Wu,
1980; Hou, 1983; Fang, 1999), there may be a point where a
warm-winter climate tips the competitive balance to evergreen
tree species. Observations from Woodward (1987), Cao
(1995), Williams-Linera et al. (2000) and Miyazawa &
Kikuzawa (2005) support this hypothesis that warm winters
can favour evergreens and limit range expansion in beech
species.
In Japan, growing season warmth rather than winter
temperatures is generally seen as the control on the distributions of beech species (Kira, 1945; Miyawaki, 1980–89; and
many others), and Japanese cool-temperate vegetation is
sometimes termed the beech forest zone (e.g. Miyawaki,
1980–89). Precipitation is also used to explain differences in
community composition and structure of Japanese beech
forests (Kure & Yoda, 1984; Hattori & Nakanishi, 1985; Fang &
Yoda, 1990; Matsui et al., 2004). In particular, distribution of
F. crenata along the Pacific Ocean side and the Japan Sea side
of Japan usually has been explained by accumulated snowfall
(Yamazaki, 1983; Maeda, 1991; Matsui et al., 2004), with pure
beech forests found only on the Japan Sea side where snowfall
is extremely abundant. Given the situation in China, however,
we should not discount out of hand the possible influence of
Journal of Biogeography 33, 1804–1819
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1811
J. Fang and M. J. Lechowicz
low winter temperatures in the development of pure beech
forests on the Japan Sea side. Winter temperature there is
much lower than on the Pacific Ocean side; this could
strengthen the competitive ability of beech against other tree
species (Tsuga and Quercus) and understorey bamboo (Sasa),
and enable its dominance.
A question remains with regard to Japanese beech distribution: why is there no beech on Yakushima Island (30!27¢ N,
130!30¢ E, 1935 m a.s.l.) in southern Japan where the climate
is suitable for beech growth and there are many species that
co-occur with beech in Kyushu, Shikoku and Honshu
(Miyawaki, 1980–89). This may be related to some topographic barriers or climatic limitations at the time during
glaciation when Yakushima was part of the major Japanese
islands, or simply to dispersal limitation since sea levels rose
and isolated the island. Another possibility worth investigating
is that a combination of warm winters and a photoperiodic
influence on the timing of budburst in beech (Falusi &
Calamassi, 1996) lead to poor synchrony between spring
budburst and the early part of the growing season that puts
beech at a competitive disadvantage.
Climates and the present distribution of American
beech
The PCA showed that MTCM and PET have almost equivalent
loadings to growing-season warmth (WI and ABT) for the
southern limit of F. grandifolia, whereas heat (WI and ABT)
and energy (PET) are most important for the northern limit
(Table S2). Although Huntley et al. (1989) demonstrated the
role of January and July mean temperatures in the present
distribution and abundance of beech species in North America
and Europe using pollen percentages of surface samples, they
used only monthly mean temperatures as thermal parameters.
Our findings based on survey of many more climatic
parameters are generally consistent with their conclusions.
Although our study suggested that the growing season
warmth was associated with the northerly distribution limit,
some physiological observations emphasize the adverse effects
of excessively low winter temperatures for many temperate
North American trees (Sakai & Weiser, 1973; Hicks & Chabot,
1985; Denton & Barnes, 1987; Maycock, 1994). Many studies
stress the influence of low winter temperature, but perhaps
only because it is easier to do experimental manipulations of
chilling effects than warmth during the growing season. We
can, however, consider biogeographic evidence supporting the
importance of growing season temperature. The lack of
American beech in Newfoundland, Canada, where winter
temperatures are much higher than at the northern edge of
beech distribution (Table 4) suggests growing season warmth
is a greater limitation than winter cold. Climatic statistics of
thermal variables at 18 stations located between 47! and 48! N
in Newfoundland where the latitudes were coincident with the
northern limit in east Quebec show far higher winter
temperatures (CI and MTCM) in Newfoundland than at the
beech northern limit (in the former, )34.8 !CÆmonth for CI
1812
Table 4 Thermal variables for Newfoundland, Canada, based on
18 climatic stations (Atmospheric Environmental Service, Environment Canada, 1982). For comparison, the estimated mean
value of climate variables at the northern limit of Fagus grandifolia
in North America (‘Mean at northern limit’ column) is also
tabulated. See Table 2 for abbreviations for climatic variables
Variable
Mean at
Mean SD Minimum Maximum northern limit
AMT (!C)
5.2 0.5
4.1
WI (!C month) 36.7 3.0 30.2
CI (!C month) )34.8 4.2 )42.2
ABT (!C)
6.1 0.3
5.3
MTWM (!C)
15.6 0.5 14.7
MTCM (!C)
)4.2 1.1 )6.1
ART (!C)
19.8 1.1 17.2
K
25.3 2.6 19.5
5.8
41.6
)26.3
6.4
16.4
)1.9
22.5
31.4
4.2
50.7
)60.5
7.0
18.4
)11.4
29.8
49.5
and )4.2 !C for MTCM, and the latter )60.5 !CÆmonth and
)11.4 !C). In contrast, the growing season temperatures were
much lower in Newfoundland than at the beech northern
limit; for the former, WI, ABT and MTWM were 36.7 !C,
6.1 !C and 15.6 !CÆmonth, and for the latter those are 50.7 !C,
7.0 !C and 18.4 !CÆmonth, respectively (Table 4). This suggests that insufficient warmth during the growing season may
be a factor limiting the expansion of beech into Newfoundland. This hypothesis is supported by eco-physiological studies
of flowering and seed production showing that a certain
minimum degree of heat is required for floral initiation, and
flower and seed production in many temperate tree species
(Matyes, 1969; Owens & Blake, 1985). The comparative study
of masting behaviours of beech species also shows the
importance of summer heat in controlling beech seed
production (Piovesan & Adams, 2001).
Although temperature can account for the distribution
limits of American beech, Fig. 1 suggests that continentality
(K) also is important for limiting the southern and northern
distribution limits. The fact that WI and MTCM at the
southern limit decrease markedly with increasing K value
shows that beech requires more heat and higher winter
temperature in an oceanic climate than in a continental one.
However, growing season warmth tends to increase as K
increases at the northern limit (Fig. 1a), suggesting higher
summer temperatures in the continental than in the oceanic
climate. The relationship between winter temperature and K
values at the northern limit shows the same pattern as at the
southern limit (Fig. 1b). Similar results were found for the
distributions of some tree species and vegetation zones in East
Asia (Ohsawa, 1990; Fang & Yoda, 1991; Fang et al., 1996).
Climates and present distribution of beech species in
Europe
The relationships between beech distribution and environments in Europe have been discussed from the viewpoint of
soil, topography and climate (Ellenberg, 1986; Jahn, 1991).
Journal of Biogeography 33, 1804–1819
ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd
Climatic limits for world beech distribution
Figure 1 Relationships between (a) warmth index (WI) and (b)
mean temperature for the coldest month (MTCM) and continentality index (K) at the southern (filled circles) and northern (open
circles) limits of Fagus grandifolia in North America. The relationships for the northern limits are fit by a nonlinear regression.
These earlier studies supported the importance of growing
season temperatures, but did not focus in any detail on the
array of thermal parameters that might be involved. Both
thermal climate and continentality (K) contributed to European beech distributions, much as in North America. Figure 2
expresses the relationships between the thermal variables (WI
and MTCM) and K value at the northern limit of F. sylvatica.
With an increase of the K values, WI increased (Fig. 2a), and
MTCM decreased (Fig. 2b).
Although beech has been present in south-eastern England
since at least 3000 yr bp (Birks, 1989), its present distribution
in the British Isles does not appear to be in equilibrium with
present climate. Climatic analysis shows that beech still has not
reached its potential northern range limit (Table 5). Compared
with averages of climatic variables at the upper/northern
limits on the European mainland (AMT of 6.6 !C, WI of
47.7 !CÆmonth, and ABT of 7.2 !C), those at the present
northern limit in Britain were much larger (more southerly),
by c. 3.4 !C for AMT, 15 !CÆmonth for WI and 2.8 !C for ABT
(Table 5). Assuming a decrease in mean temperatures of
0.5 !C per degree latitude in higher latitudes (Fang, 1996),
beech should arrive at its range limit c. 6! to the north of its
present range boundary. Spreading north at a rate of 100–
200 m yr)1 (Birks, 1989), beech should reach its potential
natural distribution in Britain 3500–7000 years in the future if
one assumes an unchanging climate scenario. This implies that
most of Britain eventually should be covered by beech.
Figure 2 Relationships between (a) warmth index (WI) and (b)
mean temperature for the coldest month (MTCM) and continentality index (K) at the northern limit of Fagus sylvatica in
Europe.
Table 5 Climatic parameters at the northern limit for Fagus
sylvatica in England based on 11 climatic stations. For comparison,
the estimated mean value of climate variables at the northern limit
of F. sylvatica on the European mainland (‘Mean at northern limit’
column) is also tabulated. See Table 2 for abbreviations for
climatic variables
Mean at
Minimum Maximum northern limit
Variables
Mean SD
AMT (!C)
WI (!C month)
CI (!C month)
ABT (!C)
MTWM (!C)
MTCM (!C)
ART (!C)
K
AP (mm)
PET (mm)
AAE (mm)
Im
EQ (!C mm)1)
10.0
0.4
9.1
62.6
3.4 53.8
)2.3
1.2 )4.2
10.0
0.4
9.1
16.8
0.4 15.8
3.9
0.6
3.1
12.9
0.7 10.6
7.7
1.6
2.8
707.2 148.4 540.0
646.8
8.6 625.0
560.6 44.3 497.0
14.7 20.4 )8.8
24.7
4.8 15.4
10.7
69.2
0.0
10.7
17.6
5.5
13.9
9.8
1068.0
661.0
652.0
64.2
32.4
6.6
47.7
)28.3
7.2
16.9
)2.7
19.6
25.8
1272.3
577.6
496.7
119.3
16.8
Comparison of climatic space for world beech species
The present distributions of most tree species are strongly related
to climate (Woodward, 1987; Huntley et al., 1989; Francis &
Currie, 2003). Simple isotherm methods have long been used to
assess distributional limits in relation to climate (Hutchinson,
1918; Koppen, 1936; Kira, 1945, 1991; Thornthwaite, 1948;
Journal of Biogeography 33, 1804–1819
ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd
1813
J. Fang and M. J. Lechowicz
Bryson, 1966; Tuhkanen, 1980; Grace, 1987; Arris & Eagleson,
1989; and many others), but the nuances of climatic controls on
range limits are not assessed fully by isotherms. In this study, we
used a more comprehensive PCA approach to detect the single
and joint importance of diverse climatic parameters in explaining the distribution of beech species. We are thus able to
compare the climatic space of beech distribution among species
in the three separate geographic regions where beech are major
components of forest ecosystems.
Because the first two PCA axes were most important in
explaining the distribution of beech species around the world,
sample scores of these axes were used to compile scatter
diagrams comparing the climatic spaces (climatic niches) of
the respective beech species (Figs 3 and 4). Because of a large
sample size and rather scattered score values, 50% Gaussian
bivariate confidence ellipses (ELL) were drawn for the species
with a large sample size to more easily compare climatic
conditions among beech species. Also, because four Chinese
beeches and two European beeches have similar moisture and
warmth requirements (see Table 2; Table S1), sample data
were combined.
Figures 3 and 4 show climatic gradients associated with
the lower (southern) limit and the upper (northern) limit of
beech. At its lower or southern limits (Fig. 3), F. grandifolia
extends to warmer regions, while F. crenata requires less
warmth, but both Japanese beeches occur in more moist
climates than F. grandifolia. Chinese and European beeches
occupy similar temperature ranges, but the latter is in a
drier climate. At the upper or northern edges (Fig. 4), both
F. grandifolia and F. crenata occupy colder areas, while
Chinese beeches have similar warmth demand to F. japonica.
Along the moisture axis, F. crenata occurs in the most
humid conditions, and European beech in the driest
habitats. Although the actual influence of climatic factors
on species distributions may be nonlinear and only partly
reflected in the present analyses (Austin, 2002), it is clear
that there is a degree of climatic niche differentiation among
the extant beech species.
CONCLUSIONS
Focusing on distribution limits of the world beech species, we
compare their climatic spaces in three different regions
globally, and explore the climatic correlates of these distribution patterns. The results suggest that thermal climate is most
important overall in determining the distribution of beech
Figure 3 Climatic spaces for world beech (Fagus) species at their lower/southern distribution limit. The first two principal components are
plotted. The first axis indicates a gradient in thermal climate and the second a moisture gradient in the overall distribution of world beech
species. The 50% ELL is drawn for major beech species to show their primary ranges. Inset graph shows climatic scores of four Chinese beech
species that have similar climatic ranges.
1814
Journal of Biogeography 33, 1804–1819
ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd
Climatic limits for world beech distribution
Figure 4 Climatic spaces for world beech (Fagus) species at their upper/northern distribution limit. The second axis, indicated by
precipitation and moisture index for most beech species, shows a negative correlation with beech distribution (Table S2 in Supplementary
Material); we have inverted the scale of this axis to more easily compare with Fig. 3. The occurrence of F. grandifolia to the lower-left does
not indicate its northern limit is set by dry conditions; the second axis indicates low winter temperature, but not moisture regime, with the
loadings of )0.98 and )0.91 for MTCM (mean temperature for the coldest month) and CI (coldness index) (Table S2), respectively. For
additional explanations see Fig. 3.
species, and that moisture effects are secondary. The degree
and duration of low winter temperature (MTCM and CI), and
annually available solar energy (PET) sometimes also played a
role. At the lower or southern limits, F. grandifolia occurred in
much warmer regions, and F. crenata in colder regions;
Chinese and European beeches have similar, intermediate heat
requirements. Along moisture gradients, Japanese beeches
appeared in more moist conditions and Chinese and European
beeches in drier situations. At the upper or northern limits,
F. crenata and American beeches had similar, relatively low
warmth demands, while F. japonica and Chinese beeches were
found only in warmer regions. Along a moisture gradient, the
Japanese beech species again occupied the most moist regions,
with European beech in contrast occupying the most dry
(Fig. 4).
Growing season temperature was most important in
explaining overall distribution of Chinese beeches, but their
northern limits were mainly set by low precipitation. The
climatic factor controlling their westward expansion (southeast Tibet and Himalaya) may be higher winter temperatures
that influence their budburst in spring and weaken their
competitive ability with evergreen hemlock and broad-leaved
evergreen trees. Although the distribution limits of beech
species in Japan were controlled by summer temperature, their
dominance may depend on regional climatic factors such as
snowfall and winter low temperature. Winter low temperature
may enhance the competitive ability of F. crenata with other
co-existing species, allowing it to form pure beech forests in
western Japan.
High summer temperature was considered to be the limiting
factor for southward extension of American beech, while
adequate growing season warmth was critical for its northward
distribution. Continentality (K) played an important part in
delimiting its range expansion, but lack of growing season
warmth was the most important climatic factor precluding its
migration to the Atlantic Islands (such as Newfoundland,
Canada). Summer temperature is a limiting factor for the
distribution of beeches in Europe, but continentality was also
associated with limits to their north-western distribution. The
northerly distribution of beech in Britain has apparently not
reached its potential limit due to lack of time since deglaciation.
Although the present-day distribution patterns of beech
species showed good correspondence to contemporary climate,
Journal of Biogeography 33, 1804–1819
ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd
1815
J. Fang and M. J. Lechowicz
isolated exceptions exist. For example, despite favourable
climatic conditions there are no records of beech ever having
grown on Yakushima Island in Japan. Similarly, F. mexicana,
isolated in a small mountainous area of north-eastern Mexico,
shows no clear association with contemporary climate. Therefore, a historical view on beech distribution is an essential
complement to the climatic analyses emphasized in this paper.
ACKNOWLEDGEMENTS
Assistance from many colleagues enabled this study. JYF is
greatly indebted to Y.H. Tang for assistance in collecting beech
data in Japan and Korea, C.F. Hsieh for F. hayatae in Taiwan,
and F. Reygadas for Mexican beech. Thanks are extended to
Z.H. Wang and X.P. Wang for their assistance in data analysis,
and to S.P. Wang for her help in compiling climatic data
sets and checking place locations. We also thank Robert
Whittaker and two anonymous referees for their helpful
comments and suggestions on the earlier version of this paper.
This work was mostly done in MJL’s laboratory when JFY
worked as a postdoctoral researcher in 1996–97, and supported
by a Natural Sciences and Engineering Research Council of
Canada grant to MJL and by the National Natural Science
Foundation of China to JYF.
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SUPPLEMENTARY MATERIAL
The following supplementary material is available online from
http://www.Blackwell-Synergy.com
Appendix S1 Location and elevation of distribution limits of
beech (Fagus L.) species in the present study.
Table S1 Statistics for climatic variables at lower (southern)
and upper (northern) limits of distribution for world beech
species.
Table S2 Loadings of climatic variables derived from Principal Components Analysis for the first three principal compo-
nents associated with the distribution limits of world beech
species.
Table S3 Coefficient of correlation between annual mean
temperature (AMT) and other thermal variables across the
global range of beech species.
BIOSKETCHES
Jingyun Fang is a professor and chair of the Department of
Ecology, Peking University. His research interests cover
biogeography of plants, terrestrial ecosystem productivity
and remote sensing of vegetation.
Martin J. Lechowicz is a professor in the Department of
Biology, McGill University, and Director of the University’s
Gault Nature Reserve. His research interests centre on the
comparative ecology of trees and on the ecology and
conservation of forest communities.
Editor: Robert J. Whittaker
Journal of Biogeography 33, 1804–1819
ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd
1819
Supplementary Material for Paper of Fang and Lechowicz
Climatic limits for the present distribution of beech (Fagus L.) species in
the world
Jingyun Fang 1 and Martin J. Lechowicz 2
1
Department of Ecology, College of Environmental Sciences, and Center for Ecological Research &
Education, Peking University, Beijing 100871, China
2
Biology Department, McGill University, 1205 Dr. Penfield Avenue, Montreal, Quebec, Canada H3A 1B1.
Journal of Biogeography, 2006, 33, 1804–1819
Supplementary Materials
Appendix S1
Table S1
Table S2
Table S3
Appendix S1. Locations and elevation of distribution limits of beech (Fagus) species used in the present study. “-“: no
available records.
Longitude Lower limit (m) Upper limit
No.
Location
Latitude
(o.’)
(m)
(o.’)
F. engleriana
1
Daheshan, Yiliang, Yunnan
27.34
104.20
1200
2500
2
Taiyanggong, Zhenxiong, Yunnan
27.48
104.54
1200
2000
3
Daguang, Yunnan
27.52
103.54
1200
2100
4
Fanjianshan, Guizhou
27.55
108.41
1600
2200
5
Yongshan, Yunnan
28.01
103.36
1200
2500
6
Baimashan, Suichang, Zhejiang
28.12
119.12
800
1000
7
Sanjiangkou, Yunnan
28.15
103.58
!
2550
8
Daliangshan, Meigu, Sichuan
28.18
103.06
1000
2400
9
Jinfushan, Nanchuan, Sichuan
29.02
107.06
!
2100
10
Datungyan, Ebian, Sichuan
29.06
103.25
1000
11
Gutianshan, Kaihua, Zhejiang
29.06
118.24
800
2500
!
12
Emeishan, Sichuan
29.31
103.21
1900
2100
13
Qiyueshan, Lichuan, Hubei
30.02
108.30
!
1800
14
Guniujiang, Shitai, Anhui
30.03
117.28
1100
1600
15
mt-qingliang, jixi, anhui
30.07
118.55
1000
1500
16
Huangshan, Anhui
30.08
118.09
1200
1550
17
Tianquan, Sichuan
30.10
102.34
1000
2400
18
jiuhuashan, Qingyang, Anhui
30.36
117.48
1200
1360
19
Tianzhushan, Qianshan, Anhui
30.40
116.30
1000
1400
20
Daozhijian, Yuexi, Anhui
30.48
116.04
1000
1400
21
Xiningxia, Hubei
31.00
110.35
1300
1800
22
Guanxian, Sichuan
31.00
103.36
1000
2500
23
Tiantangzhai, Dabieshan, Anhui
31.11
115.44
1200
1700
24
Dabao, Penxian, Sichuan
31.14
103.44
1000
2400
25
Leigutai, Xingshan, Hubei
31.22
110.35
1000
2000
s1
26
Wuoxi, Sichuan
31.24
109.36
1000
1800
27
Dafushan, Jinzhai, Anhui
31.36
115.38
1000
1500
28
Shenlongja, Hubei
31.42
110.35
1400
2200
29
Mt-Baimajian
31.44
116.22
900
1600
30
Jiulongshan, Zhenping, Shanxi
31.48
109.30
1100
2100
31
Chengkou, Sichuan
31.50
108.40
1000
2400
32
Tongjiang, Sichuan
31.56
107.14
1500
2200
33
Micangshan, Nanzhen, Sichuan
32.02
107.04
1400
2000
34
Houhekou, Fangxian, Hubei
32.13
110.32
1000
1500
35
North Dabashan, Ziyang, Shanxi
32.15
108.30
1100
2100
36
Cailinbao, Pingwu, Sichuan
32.17
104.20
1000
2400
37
Moutianling, Qingchuan, Sichuan
32.30
104.52
1000
2400
F. longipetiolata
1
Mongci, Yunnan
23.14
103.15
800
!
2
Wenshan, Yunnan
23.20
104.13
800
2000
3
Yuyuan, Guangdong
23.58
113.56
1200
1800
4
Xinchang, Xinyi, Guizhou
24.08
104.48
1000
!
5
Yangshan, Guangdong
24.18
112.29
1200
1800
6
Laoshan, Tianlin, Guanxi
24.18
106.00
1500
1900
7
Yuanbaoshan, Guangxi
25.27
109.12
1300
2200
8
Dupuangling, Daoxian, Hunan
25.30
111.30
1000
1700
9
Miaoershan, Guangxi
25.52
110.28
1000
2200
10
YangMingshan, Shuangpai, Hunan
25.54
111.36
1000
1700
11
Bamienshan, Guidong, Hunan
26.00
113.44
1200
1700
12
Zhengbaoding, Hunan
26.07
110.27
800
1500
13
Leigongshan, Guizhou
26.24
108.12
900
2100
14
Louxiaoshan, Hunan
26.26
113.54
800
1500
15
Jinggangshan, Jiangxi
26.38
113.11
900
1500
16
Baishifong, Wuyi Mts, Fujian
26.47
116.51
1100
1400
17
Chaling, Hunan
27.05
113.38
1200
1400
18
Xuefengshan, Hunan
27.12
110.04
800
1500
19
Zhengxong, Yunnan
27.29
104.59
!
1900
20
South Yandangshan, Taixun, Zhejiang
27.32
120.06
400
1200
21
Weixing, Yunnan
27.49
105.01
!
2000
22
MT-fangjinshan
27.55
108.41
1100
1900
23
Guixi, Jiangxi
28.01
117.30
800
!
24
Yunhe, Zhejiang
28.01
119.25
400
1400
25
Zhulong, Longquan, Zhejiang
28.01
118.50
400
1400
26
Suzishan, Songdao, Guizhou
28.06
109.06
1000
1800
27
Daliangshan, Miegu, Sichuan
28.18
103.16
1000
2500
28
Dabaoding, Leibou, Sichuan
28.26
103.37
1000
2400
29
Xushan, Sichuan
28.32
108.50
1000
1600
30
Dalingshan, Shangrao, Jiangxi
28.34
117.54
800
!
31
Baimashan, Suichang, Zhejiang
28.48
119.21
400
1400
s2
32
Mabian, Sichuan
28.55
103.15
1000
2600
33
Sanqingshan, Jiangxi
28.56
118.04
800
1400
34
Jinfushan, Nanchuan, Sichuan
29.02
107.06
1000
2500
35
Datuanya, Ebian, Sichuan
29.06
103.25
1000
2200
36
Huadingshan, Tiantai, Zhejiang
29.10
121.04
400
1000
37
Tiantaishan, Fonghua, Zhejiang
29.24
121.14
400
1000
38
Qixi, Kaihua, Zhejiang
29.26
118.24
400
1100
39
Hefeng, Hubei
29.38
109.48
800
1900
40
Qiongan, Zhejiang
29.51
118.59
400
1200
41
Hongya, Sichuan
29.54
103.18
!
2000
42
Huoshaobao, Xuanen, Hubei
30.01
109.44
!
1900
43
Qiyueshan, Lichuan, Hubei
30.02
108.30
800
1800
44
Tianquan, Sichuan
30.06
102.42
1000
2600
45
mt-qingliang,jixi,anhui
30.07
118.55
1000
1500
46
Huangshan, Anhui
30.08
118.09
600
1400
47
Longwangshan, Linan, Zhejiang
30.27
119.26
400
1200
48
Jianshi, Hubei
30.43
109.40
800
2000
49
Wushan, Sichuan
31.00
109.48
1000
1800
50
Xilingxia, Hubei
31.00
110.30
1200
1700
51
Xiningxia, Hubei
31.00
110.35
1200
1700
52
Guangxian, Sichuan
31.00
103.36
1000
!
53
Leigutai, Xingshan, Hubei
31.22
110.35
800
2000
54
Wenchuan, Sichuan
31.24
103.36
1000
2400
55
Wuxi, Sichuan
31.28
109.26
1000
1800
56
Shenlongjia, Hubei
31.42
110.35
!
2100
57
Changkou, Sichuan
31.50
108.48
1000
2300
58
Micanshan, Nanzhen, Sichuan
32.02
107.04
1300
1900
59
Sijiemeishan, Pingwu, Sichuan
32.34
104.24
1000
1600
60
Sungpan, Sichuan
32.36
103.36
1300
2500
61
Dabashan, Sichuan
32.38
107.30
1300
1900
1
Dayiaoshan, Guangxi
24.00
110.00
1000
1600
2
Shueshanzhang, Guangdong
24.16
113.30
600
!
3
Laoshan, Tianlin, Guangxi
24.18
106.00
1000
!
4
Shanmatangding, Jianghua, Hunan
24.40
111.36
800
1700
5
Mangshan, Yizhang, Hunan
24.57
113.00
800
1900
6
Linwu, Hunan
25.12
112.30
800
1700
7
Wuzhifeng, Yucheng, Hunan
25.23
113.30
800
1700
8
Baojieling, Guanyang, Guangxi
25.24
110.58
1000
1600
9
Jiucailing, Daoxian, Hunan
25.30
111.25
800
1900
10
Bamainshan, Zixing, Hunan
26.00
113.38
800
1900
11
Jigongdong, Jiangxi
26.02
116.21
900
1300
12
Yangmingshan, Shuangpai, Hunan
26.04
111.54
1000
1500
13
Zhenbaoding, Ziyuan, Guangxi
26.08
110.50
1000
1600
F. lucida
s3
14
Mingzhulaoshan, Chengbu, Hunan
26.16
110.50
1200
1950
15
Mingjushan, Chengpu, Hunan
26.18
110.18
1400
1600
16
Nanshan, Chengbu, Hunan
26.22
110.23
1400
1850
17
Louxiaoshan, Hunan
26.24
113.52
1350
1500
18
Leigongshan, Guizhou
26.24
108.12
1000
2100
19
Leigongshan, Guizhou
26.25
108.08
1300
1600
20
Dongan, Hunan
26.38
111.08
800
1600
21
Salaxi, Bijie, Guizhou
27.08
105.04
1300
1600
22
Wugongshan, Jiangxi
South Yandangshan, Taishun, Zhejiang
27.31
27.32
114.10
120.06
800
1000
1400
1200
24
Congan, Fujian
27.38
118.18
1100
!
25
Baishanzuo, Qingyuan, Zhejiang
27.46
119.10
1300
1700
26
Huanggang, Zhejiang
27.52
117.49
1300
1700
27
Guankoushui, Suiyang, Guizhou
27.54
107.06
1000
1750
28
Fanjianshan, Guizhou
27.55
108.41
1300
2100
29
Huangmoujian, Longquan, Zhejiang
27.56
119.10
1300
1700
30
Xinjieping, Gulan, Sichuan
28.00
105.42
1300
1800
31
Kuankuoshui, Guizhou
28.11
107.11
1400
1750
32
Daliangshan, Meigu, Sichuan
28.18
103.06
1300
2300
33
Mt-jiulongshan,Zhangjiang
28.21
118.52
1400
1700
34
Dabaoding, Leibuo, Sichuan
28.22
103.36
1300
2300
35
Jiulongshan, Suichang, Zhejiang
28.23
118.56
1300
1700
36
Mabian, Sichuan
28.32
103.15
1300
2300
37
Baimashan, Suichang, Zhejiang
28.36
119.10
1300
1600
38
Daozhen, Guizhou
28.38
107.36
1000
1700
39
Jianshanke, Yongxun, Hunan
29.01
109.38
700
2200
40
Datangyan, Ebian, Sichuan
29.04
103.22
1300
2300
41
Erbian, Sichuan
29.10
103.20
1550
2400
42
Shimen, Hunan
29.36
111.18
700
!
43
Hefeng, Hubei
29.38
109.48
1000
1800
44
Badagongshan, Hunan
29.40
119.49
1400
1850
45
Zhangcui, Hongya, Sichuan
29.40
103.04
1300
1700
46
Tongzi, Shizhu, Sichuan
29.44
107.43
1300
1900
47
Huoshaobao, Xuanen, Hubei
30.01
109.44
1000
2000
48
Longtangba, Hubei
30.02
109.06
1000
1600
49
Qiyueshan, Lichuan, Hubei
30.02
108.30
1000
1800
50
Guniujiang, Shitai, Anhui
30.03
117.28
1300
!
51
Tianmushan, Zhejiang
30.25
119.30
1000
1400
23
52
Tianzhoushan, Qianshan, Anhui
30.40
116.30
1000
1300
53
Yuexi, Anhui
30.48
116.04
1000
1400
54
Xilingxia, Hubei
31.00
110.35
1300
1800
55
Huoshan, Anhui
31.20
116.05
1000
1600
56
Leigutai, Xingshan, Hubei
31.22
110.35
1000
2000
57
Shenlongja, Hubei
31.42
110.35
1400
2200
s4
58
mt-baimajian, heshan, anhui
31.44
116.22
700
!
59
Guangwushan, Nanjiang, Sichuan
32.38
106.43
1300
2300
60
Pingwu, Sichuan
32.24
104.30
1300
1300
F. hayatae
1
Nanjiang, Sichuan
32.38
106.43
1300
1900
2
Tongjiang, Sichuan
31.50
108.20
1100
1900
1850
3
Dabashan, Sichuan
32.38
107.30
!
4
Longtangshan, Linan, Zhenjiang
30.28
119.42
900
1300
5
Taiixun, Zhejiang
27.03
120.06
900
1200
6
Shimen, Hunan
29.36
111.18
900
1300
7
Dabashan, Nanzheng, Shanxi
33.00
106.54
!
1600
8
Lalashan, Taipei
24.45
121.26
1300
2000
9
Sanhsingsan-Tungshan
24.31
121.33
1800
1800
10
Sihaishan, Yongjia, Zhejiang
28.08
120.33
850
1000
F. crenata
1
Tsubamenosawa, Hokkaido
42.47
140.2
!
620
2
Taiheizan, Hokkaido
42.45
140.18
100
900
3
Oshamanbedake, Hokkaido
42.30
140.18
320
800
4
Shimokita
41.26
141.10
10
850
5
Hakkoda-san
40.40
140.53
200
1200
6
Iwaki-san
40.38
140.18
!
1150
7
Ashiro-cho, Iwate
40.05
140.00
460
1000
8
Hachimantai
39.58
140.51
500
1100
9
Iwaizumicho, Shimoheigun, Iwate
39.51
141.47
!
1200
10
Hayachine
39.29
141.31
400
1150
11
Chokai-san
39.06
140.04
450
1120
12
Yakeishi-dake
39.05
140.50
300
1100
13
Kurikoma-yama
38.58
140.49
400
1200
14
Gassa
38.33
140.02
100
1350
15
Funagata-yama
38.25
140.35
!
1300
16
Asahi-dake
38.15
139.56
300
1200
17
Zao-san
38.09
140.27
250
1375
18
Ihde-san
37.51
139.43
350
1500
19
Azumayama
37.44
140.09
300
1450
20
Asakusa-dake
37.21
139.15
400
1450
21
Aizuasahi-dake
37.13
139.20
600
1550
22
Aizukomagadake
37.02
139.25
!
1600
23
Hiuchigadake
36.55
139.20
!
1550
24
Tanigawa-dake, Gumma
36.55
138.55
680
1550
25
jap2
36.53
138.53
!
1600
26
Naebasan,
36.50
138.41
!
1640
27
Nikko, Tochigi
36.45
139.35
850
1700
28
jap3
36.34
137.37
400
1600
29
Tsukubasan, Ibaraki
36.10
140.05
820
!
s5
30
Chichibu, Saitama
35.55
138.50
1000
1650
31
Hyonosen, Okayama
35.35
134.30
1150
!
32
Okutama,
35.35
139.20
850
1650
1700
33
jap5
35.30
138.49
!
34
Ooginosen, Tottori
35.25
134.25
850
!
35
Mt Fuji
35.22
138.44
1000
1580
36
Daisen, Tottori
35.2
133.30
800
!
37
Jap6
35.17
138.01
!
1550
38
Sanjogatake, Nara
34.15
135.55
!
1590
39
Hakkenzan
34.15
135.55
900
1760
40
Hakkenzan, Nara
34.10
135.50
!
1660
41
Ohdaigahara
34.10
136.05
850
1690
42
Hatenashi, Wakayama
33.55
135.4
940
!
43
Miune, Kochi
33.53
133.58
1300
1550
44
Kumosoyama, Tokushima
33.50
134.15
1250
!
45
Tsurugisan, Tokushima
33.50
134.05
1240
1650
46
Ishizuchiyama, Kochi
33.46
133.07
1350
1670
47
Sasagamine, Eihimei
33.45
133.15
1250
1500
48
Kanpuzan, Kochi
33.45
133.25
1380
1660
49
Hikosan
33.35
130.10
650
!
50
Kujusan
33.05
131.10
1000
!
51
Sobosan
32.49
131.22
1200
1700
52
Kunimidake
32.30
131.00
1350
1650
53
Ichifusayama
32.15
131.05
!
1630
54
Kirishima
31.50
130.50
1150
1400
55
Takakumayama
31.29
130.49
1200
!
F. japonica
1
Yamadamachi, Miyakoshi, Iwate
39.30
141.55
100
600
2
Hanamaki City, Iwate
39.23
141.07
100
!
3
Sendai, Miyagi
38.15
140.53
200
!
4
North Abukuma Santi, Hukushima
5
Central Abukuma Santi, Hukushima
37.46
37.20
140.42
140.43
150
200
700
750
6
Naganuma-mati, Hukushima
37.18
140.13
200
750
7
Mt.Yamizo, Ibaraki
36.54
140.10
450
!
8
Nikko, Tochigi
36.45
139.35
500
1100
9
Okukinu, Tochigi
36.36
139.56
400
1100
10
Numata, Gumma
36.35
139.00
500
1000
11
Chichibu, Saitama
35.55
139.00
500
1100
12
Hyonosen
35.35
134.30
200
800
13
Okutama,
35.35
139.20
600
850
14
Onzui-Mt.Mimuro
35.15
134.25
560
930
15
Hagacho, Hyogo
35.10
134.35
200
810
16
Oozorayama,
35.10
133.50
400
910
17
Hieizan, Shiga
35.07
135.49
500
!
s6
18
Gozaishosan,
35.00
136.25
520
1020
19
Mengamesan,
34.55
132.40
650
!
20
Hikimi-Sandankyo, Hiroshima
34.35
132.10
450
880
21
Osorakanzan,
34.35
132.05
750
!
22
Anzojiyama
34.30
132.00
800
!
23
Jipposan, Hinoshima
34.30
132.05
750
1050
24
Naganoyama
34.15
131.55
800
!
25
Gomadan, Wakayama
34.03
135.29
!
1050
26
Yazurayama, Tokushima
34.00
134.05
850
1250
27
Miune, Kochi
33.53
133.58
!
1400
28
Ohtasan, Wakayama
33.44
135.45
500
1000
29
Nomuracho, Ehime
Kunimidake, Shiibamura, Miyazaki
33.25
32.30
132.35
131.00
900
1000
1000
1300
37.29
130.54
300
960
30
F. multinervis
1
Ulreung-do, South Korea
F. sylvatica
1
Pindhos Mts, Greece
39.20
21.35
!
2000
2
Olympus, Greece
40.05
22.21
!
2000
3
Harz-Mts, Germany
51.47
10.39
!
800
4
Campania, Apennines, Italy
40.40
15.02
500
1300
5
Lazio, Apennines, Italy
41.55
13.20
350
1500
6
Abruzzi, Apennines, Italy
42.10
13.30
!
1500
7
Sicilia, Apennines, Italy
37.50
14.00
950
1750
8
Romagna, Apennines, Italy
44.25
10.00
!
1600
9
Saphane Da., Kutahya, Turkey
39.02
29.14
!
1500
10
Simav, Kutahya, Turkey
39.05
28.59
!
1700
11
Kaz-Dagi, Turkey
39.41
26.52
!
1300
12
Bergen, Norway
60.24
5.140
43
43
13
Oslo, Norway
59.56
10.44
94
94
14
Jomfruland, Norway
58.52
9.36
45
45
15
Kyrkerud, Sweden
59.23
12.07
110
110
16
Linkoping, Sweden
58.25
15.38
64
64
17
Skara,Sweden
58.24
13.27
115
115
18
Ketrzyn, Poland
54.06
21.23
!
!
19
Blalystok, Poland
53.09
23.09
!
!
20
Lublin, Poland
51.15
22.35
!
!
21
Suwalki, Poland
54.07
22.56
!
!
22
Zamosc, Poland
50.44
23.15
!
!
23
Gorodenka, Ukraine
48.40
25.30
265
265
24
Kamenetz-Podolsk, Ukraine
48.40
26.36
258
258
25
Nemirov, Ukraine
48.58
28.50
285
285
26
Nizhniy, Olchedaev, Ukraine
48.38
27.40
187
187
27
Simferopol, Ukraine
44.57
34.06
205
205
28
Staro, Konstantinov
49.45
27.13
279
279
s7
29
Tarnopol, Ukraine
49.33
25.36
320
320
30
Zdolbunovo, Ukraine
50.30
26.10
20
20
31
Cuenca, Spain
40.05
-2.08
987
987
32
Gerona, Spain
41.59
2.50
95
95
33
Marseille, Observ, France
43.18
5.23
75
75
34
Albertacce, Corsica
42.17
8.55
1074
1074
35
Sartene, Corsica
41.36
8.59
50
50
36
Catanzaro, Italy
38.55
16.37
343
343
37
Enna, Italy
37.34
14.18
950
950
38
Gambarie, Utaly
38.10
15.51
1300
1300
39
Linguaglossa, Italy
37.50
15.10
560
560
40
Petrlia, Sottana, Italy
37.48
14.06
930
930
41
Tindari, Italy
38.08
15.04
280
280
42
Komotini, Greece
41.07
25.24
30
30
43
Trikala, Greece
39.33
21.46
150
150
44
Bourgas, Bulgaria
42.30
27.28
17
17
45
Kazanlak, Bulgaria
42.37
25.24
372
372
46
Kolarovgrad ,Bulgaria
43.16
26.55
!
!
F. orientalis
1
Ulu-dagi, turkey
40.12
29.04
!
1600
2
Murat Da., Kutahya, Turkey
38.56
29.43
1700
2000
3
4
Turkmen Da., Turkey
Duldul Da., Gokcayir to Atlik Y., Adana, Turkey
39.50
37.04
30.10
36.15
1400
1600
1600
1700
5
Buyukduz, Turkey
41.20
32.30
!
1450
6
Zonguldak, south Anatolia, Turkey
41.26
31.47
200
!
7
Bolu, south Anatolia, Turkey
40.35
31.50
900
!
8
Yalnizcam Mts, northeast Turkey
41.03
42.28
50
!
F. grandifolia
1
Grand Anse, Cape Breton island, NS
46.49
60.48
11
11
2
Northeast Margaree, Cape Breton Island
46.20
61.00
61
61
3
Barney’s River region, Pictou Co, NS
45.36
62.16
76
76
4
Tignish, Prince Edward Island, NB
46.55
64.02
3
3
5
Richibucto, NB
46.41
64.52
38
38
6
Bonaventure, NB
48.03
65.29
8
8
7
Newcastle, NB
47
65.34
34
34
8
Causapscal, NB
48.22
67.14
26
26
9
NW NEW Brunswick
48
67.55
152
152
10
Rimouski, New Brunswick (NB)
48.26
68.33
411
411
11
Trois-Pistoles, Riviere-du-Loup, PQ
48.05
69.1
61
61
12
Cantons-de-l’ Est, Que
47.21
69.56
15
15
13
Henri, Levis Co
46.42
70.04
351
351
14
Baie-St-Paul, PQ d’orleans, Quebec City
47.25
70.32
15
15
15
d’ orleans, Quebec City
47.2
70.45
15
15
16
Cap-Rouge, Ile d’ orleans, Que
47
70.52
320
320
17
Saint-Joseph-de-la-Pointe, Levis Co
46.48
71.05
145
145
s8
18
Mont Megantic
45.27
71.1
354
354
19
Levis City, Levis Co
46.48
71.11
184
184
20
Saint-Lambert, Levis Co
46.35
71.13
152
152
21
Saint-Nicolas, Levis Co
46.42
71.27
184
184
22
Ste-Agathe, PQ
46.14
73.38
59
59
23
Lac-Cayamant, Que
46.08
76.15
170
170
24
Fort Coulonge, PQ
45.51
76.44
168
168
25
South of Temiskaming, ON
46.45
78.1
172
172
26
Mattawa, ON
46.19
78.42
172
172
27
Temiscaming, Que
46.43
79.06
181
181
28
North Bay City, ON
46.19
79.28
201
201
29
Sturgeon Falls, ON
46.22
79.55
358
358
30
Cache Lake, Algonquin Park, ON
46.22
79.59
198
198
31
Capreol, ON
46.43
80.56
237
237
32
Sudbury, ON
46.3
81
259
259
33
Espanola, ON
46.15
81.46
206
206
34
SE-Shore of Lake Superior
46.55
84.2
212
212
35
Sault Ste. Marie, ON
46.31
84.2
192
192
36
Univ of Michigan Biol Station, MI
45.34
84.42
216
216
37
Bay-Mills TP, Chipperwa Co, MI
46.27
84.46
220
220
38
New-Berry-Luce-co-MI
46.21
85.3
270
270
39
Washington Island, Door Co, Wisconsin
45.23
86.55
189
189
40
Chatham, Alger Co, MI
46.21
86.56
267
267
41
Ishpeming, Marquette Co, Mi
46.29
87.4
431
431
42
Iron Mountain City, Dickinson Co, MI
45.49
88.04
352
352
43
Northern Great Lakes region, Menominee Co, WIS
44.53
88.38
246
246
44
Iron River City, Iron Co, MI
46.06
88.38
452
452
45
Grand-marais, MI
46.4
85.59
230
230
46
Crivitz, high-fall, WIS
45.17
88.12
252
252
47
Fairhope, AL
30.33
87.53
7
7
48
Mobile, AL
30.41
88.15
64.3
64.3
49
Robertsdale, AL
30.32
87.4
47
47
50
Apalachicola
29.44
82.02
4
4
51
De Funiak, FL
30.44
86.07
70
70
52
Gainesville, FL
29.38
82.21
26.2
26.2
53
Lake City, FL
30.11
82.36
59
59
54
Madison, FL
30.28
83.25
58
58
55
Milton, FL
30.47
87.08
66
66
56
monticello, FL
30.32
83.55
45
45
57
Nicelle, FL
30.31
86.3
18
18
58
Pensacola
30.28
87.12
34
34
59
Saint Marks, FL
30.05
84.1
5
5
60
Tallahassee, FL
30.23
84.22
17
17
61
Albany, GA
31.32
84.08
55
55
62
Brooklet. GA
32.23
81.41
58
58
63
Camilla,GA
31.14
84.13
53
53
s9
64
Dublin, GA
32.3
82.54
66
66
65
Eastman, GA
32.12
83.12
122
122
66
Hawkinsville, GA
32.17
83.28
74
74
67
Savannah, GA
32.08
81.12
14
14
68
Swainsboro, GA
32.35
82.22
99
99
69
Warrenton, GA
30.48
83.54
64
64
70
Baton-rouge, LA
30.32
91.08
20
20
71
Carville, LA
30.12
91.07
8
8
72
Jennings
30.15
92.4
9
9
73
Lake-charles, LA
30.07
93.13
3
3
74
Melville, LA
30.41
91.45
9
9
75
Reserve, LA
30.04
90.34
4
4
76
Biloxi-City, MISS
30.24
88.54
5
5
77
Gulfport-naval
30.23
89.08
11
11
78
Picayune, Miss
30.31
89.41
15
15
79
Wiggins, Miss
30.48
89.06
61
61
80
Liberty, Tex
30.03
94.49
11
11
81
Port-Arthur
29.57
94.01
5
5
F. mexicana
1
Zacatlamaya Mts, Hidolgo
20.4
98.4
1800
1920
2
Cerro de Tutotepec, Hidolgo
20.2
98.2
1800
1920
3
Ojo-de-Agua, Temaulipas
24.15
98.25
1200
1520
4
Teziutlan, Puebla
19.5
97.2
2000
2000
s10
Table S1. Statistics for climatic variables at lower (southern) and upper (northern) limits of distribution for world beech
species. AMT: annual mean temperature; WI: warmth index; CI: coldness index; ABT: annual biotemperature; MTWM:
mean temperature for the warmest month; MTCM: mean temperature for the coldest month; ART: annual range of mean
temperature; K: continental index; AP: annual precipitation; PET: annual potential evapotranspiration; AAE: actual annual
evapotransporation; Im: moisture index; and EQ: Ellenberg quotient. Symbol “-“ means no estimation.
Climatic index
Distribution limits
Lower or southern limit
Upper or northern limit
Mean
SD
Min
Max
Mean
SD
Min
Max
AMT (oC)
12.4
2.51
8.3
17.7
7.3
1.88
3.7
12.8
o
99.0
22.40
57.6
152.4
56.0
14.19
32.2
99.9
F. engleriana
WI ( C·month)
o
-10.1
8.61
-27.3
0.0
-28.1
10.40
-52.9
-6.1
ABT (oC)
12.5
2.37
8.5
17.7
8.1
1.50
5.6
12.8
MTWM (oC)
CI ( C·month)
23.0
1.72
17.1
26.0
17.8
2.23
13.6
23.1
o
1.1
3.55
-4.1
8.2
-3.9
2.36
-8.9
1.7
ART ( C)
21.9
2.70
17.8
26.1
21.7
2.65
17.8
26.1
K
53.3
8.12
41.3
66.1
52.8
7.75
41.3
66.1
AP (mm)
1229.6
359.34
738.1
2394.5
1366.4
241.65
1180.4
2394.5
PET (mm)
805.4
139.40
421.3
940.8
795.6
137.16
421.3
911.5
AAE (mm)
803.3
137.8
421.3
940.8
793.5
136.0
421.3
908.5
61.4
82.32
0.0
356.4
70.0
76.98
36.3
356.4
21.7
5.80
6.1
32.0
19.4
3.98
6.1
22.5
AMT (oC)
12.8
2.06
9.1
15.4
9.2
2.07
6.3
11.9
o
99.4
22.36
67.6
132.1
67.0
13.26
52.4
89.2
CI ( C·month)
-6.4
6.08
-17.9
0.0
-16.6
13.73
-36.4
0.0
o
12.8
2.00
9.3
15.4
9.5
1.66
7.5
11.9
MTCM ( C)
o
Im
o
EQ ( C/mm)
F. hayatae
WI ( C·month)
o
ABT ( C)
o
MTWM ( C)
22.4
4.31
16.9
29.2
18.6
2.14
16.3
21.9
o
2.1
3.26
-2.3
7.3
-0.7
4.15
-5.9
5.4
ART ( C)
20.3
6.10
11.5
28.3
19.3
4.85
11.5
23.5
MTCM ( C)
o
K
50.7
16.74
27.1
75.0
46.3
11.21
27.1
56.1
AP (mm)
1669.9
515.72
1141.0
2558.0
2232.5
340.65
1694.5
2558.0
PET (mm)
914.6
188.11
587.0
1133.5
887.0
193.10
587.0
1133.5
AAE (mm)
914.6
188.11
587.0
1133.5
887.0
193.10
587.0
1133.5
85.5
54.20
24.4
182.1
109.8
34.81
84.4
182.1
17.0
5.20
10.8
24.1
13.2
1.74
10.8
16.5
14.3
2.15
9.6
21.6
8.9
1.76
4.9
13.3
115.9
22.64
75.5
198.9
66.2
13.75
36.5
99.3
CI ( C·month)
-4.1
4.20
-19.9
0.0
-19.8
8.42
-39.8
0.0
o
14.3
2.13
9.9
21.6
9.2
1.48
6.1
13.3
Im
o
EQ ( C/mm)
F. longipetiolata
AMT (oC)
o
WI ( C·month)
o
ABT ( C)
o
MTWM ( C)
24.2
1.69
20.7
27.4
18.9
2.08
14.5
22.4
o
3.5
3.16
-2.6
15.1
-2.0
2.19
-6.3
6.0
ART ( C)
20.8
2.97
10.7
25.5
20.9
2.47
12.0
25.5
K
54.1
9.81
25.9
72.2
54.2
8.44
31.4
72.2
MTCM ( C)
o
s11
AP (mm)
1421.0
386.11
729.5
2394.5
1738.2
165.37
1582.8
2394.5
PET(mm)
845.4
163.31
421.3
1190.7
621.2
93.91
421.3
811.6
AAE (mm)
831.8
154.9
421.3
1114.9
619.2
88.5
430.9
798.6
77.5
77.07
-4.4
356.4
132.6
63.35
97.0
356.4
19.0
5.08
6.1
29.4
14.7
2.60
6.1
17.2
12.7
2.07
8.5
17.6
8.8
1.81
3.6
12.5
100.6
18.45
68.8
150.8
66.4
12.49
35.1
95.3
CI ( C·month)
-8.2
7.26
-27.3
0.0
-21.3
10.20
-52.9
-5.3
o
12.8
1.97
9.2
17.6
9.2
1.40
5.7
12.5
Im
o
EQ ( C/mm)
F. lucida
AMT (oC)
o
WI ( C·month)
o
ABT ( C)
o
MTWM ( C)
23.0
1.40
20.5
25.5
19.1
1.57
14.9
21.7
o
1.5
2.89
-4.2
8.4
-2.4
2.39
-8.9
1.9
ART ( C)
21.5
2.40
15.3
26.1
21.5
2.12
17.7
25.8
MTCM ( C)
o
K
57.8
7.91
41.8
72.9
57.8
7.93
41.8
72.9
AP (mm)
1418.6
311.55
822.0
2205.7
1756.7
114.30
1574.9
2205.7
PET(mm)
874.0
145.23
421.3
1190.7
674.4
94.88
421.3
854.4
AAE (mm)
862.7
138.0
421.3
1114.9
671.7
90.1
431.4
842.6
70.0
67.93
-3.8
356.5
168.5
51.14
92.2
356.5
19.5
4.82
6.1
32.0
13.8
2.32
6.1
17.0
9.5
1.26
5.7
12.0
5.0
1.25
2.3
7.6
74.8
10.21
52.0
95.2
45.1
5.73
31.9
56.1
-21.3
6.68
-43.7
-6.9
-45.2
10.21
-66.3
-25.3
9.8
1.02
7.3
12.0
6.7
0.68
5.3
8.1
Im
o
EQ ( C/mm)
F. crenata
AMT (oC)
o
WI ( C·month)
o
CI ( C·month)
o
ABT ( C)
o
MTWM ( C)
21.8
1.62
19.1
24.8
17.6
0.69
15.6
18.8
o
MTCM ( C)
-2.1
1.57
-6.8
1.8
-6.8
1.81
-10.4
-3.3
o
ART ( C)
23.9
1.94
19.8
26.9
24.4
1.75
20.9
26.9
K
48.5
3.02
41.2
54.5
48.8
3.34
41.2
55.3
AP (mm)
2047.2
525.76
1135.0
3323.0
2660.3
280.00
1948.6
3323.0
PET(mm)
706.2
85.46
548.3
900.7
606.8
33.30
548.3
688.3
AAE (mm)
706.2
85.5
548.3
900.7
606.8
33.30
548.3
688.3
Im
189.9
64.52
62.9
357.3
241.3
30.11
182.8
357.3
12.5
3.10
7.3
20.6
9.5
1.20
7.3
12.8
AMT (oC)
11.7
1.03
10.3
14.8
9.0
0.99
7.3
11.8
o
91.8
8.71
81.1
117.5
70.3
7.09
55.3
87.2
-19.1
-0.3
-21.9
5.18
-29.1
-6.0
o
EQ ( C/mm)
F. jopanica
WI ( C·month)
o
CI ( C·month)
-12.0
4.09
o
ABT ( C)
11.7
1.00
10.5
14.8
9.4
0.82
7.9
11.8
MTWM (oC)
23.9
1.11
21.9
26.0
21.2
0.84
19.5
22.4
MTCM (oC)
0.3
1.31
-1.8
4.7
-2.2
1.25
-4.1
1.7
ART ( C)
23.7
1.25
20.0
26.8
23.4
1.22
20.0
25.0
K
48.8
3.08
41.3
53.1
48.6
3.03
41.3
53.1
AP (mm)
1805.2
617.07
1148.0
4006.0
2328.7
452.47
1807.0
4006.0
PET(mm)
746.6
62.51
543.5
854.7
716.4
44.50
543.5
752.1
o
s12
AAE (mm)
746.6
62.51
543.5
854.7
716.4
44.50
543.5
752.1
Im
142.0
78.51
52.1
373.9
206.5
51.49
153.0
373.9
15.3
4.21
6.5
22.7
11.5
1.99
6.5
14.7
AMT (oC)
11.5
-
-
-
7.6
-
-
-
o
90.2
-
-
-
59.0
-
-
-
CI ( C·month)
-11.9
-
-
-
-27.5
-
-
-
o
11.5
-
-
-
8.3
-
-
-
o
EQ ( C/mm)
F. multinervis
WI ( C·month)
o
ABT ( C)
o
MTWM ( C)
23.4
-
-
-
19.5
-
-
-
MTCM (oC)
0.1
-
-
-
-3.8
-
-
-
ART (oC)
23.3
-
-
-
23.3
-
-
-
K
45.0
-
-
-
45.0
-
-
-
AP (mm)
1485.0
-
-
-
-
-
-
-
PET(mm)
702.6
-
-
-
-
-
-
-
AAE (mm)
702.6
-
-
-
-
-
-
-
Im
111.3
-
-
-
-
-
-
-
16.1
-
-
-
-
-
-
-
19.5
0.76
17.2
21.0
4.2
0.88
1.6
6.0
173.4
9.14
146.5
192.0
50.7
4.74
36.1
61.2
0.0
0.00
0.0
0.0
-60.5
8.17
-76.9
-38.1
19.5
0.76
17.2
21.0
7.0
0.48
5.5
8.1
o
EQ ( C/mm)
F. grandifolia
AMT (oC)
o
WI ( C·month)
o
CI ( C·month)
o
ABT ( C)
o
MTWM ( C)
27.5
0.41
26.7
28.4
18.4
0.84
16.0
20.1
o
MTCM ( C)
10.4
1.20
7.2
13.6
-11.4
1.95
-13.7
-6.6
o
ART ( C)
17.1
1.10
13.8
19.5
29.8
2.13
24.5
32.2
K
36.8
3.04
27.4
45.0
49.5
5.03
37.4
55.3
AP (mm)
1425.7
166.81
1148.0
1668.0
1020.9
168.18
790.3
1426.5
PET (mm)
1024.2
40.43
907.8
1100.3
537.8
19.86
475.9
583.8
AAE (mm)
996.9
69.0
840.5
1100.3
530.5
18.6
475.9
575.5
40.3
14.64
20.6
65.6
91.1
34.34
43.3
168.2
19.5
2.27
16.3
23.6
18.5
3.29
12.5
24.9
Im
o
EQ ( C/mm)
F. mexicana
AMT (oC)
15.6
1.51
14.3
17.4
14.8
1.79
13.6
17.4
WI (oC·month)
127.3
1.52
111.8
149.0
117.1
1.80
103.4
149.0
CI (oC·month)
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
15.6
1.51
14.3
17.4
14.8
1.8
13.6
17.4
o
ABT ( C)
o
MTWM ( C)
19.6
1.59
18.1
21.1
18.7
1.6
17.4
20.8
o
10.5
2.59
7.9
14.1
9.7
3.1
7.2
14.1
ART ( C)
9.1
2.10
6.7
11.2
9.1
2.1
6.7
11.2
22.2
7.46
13.7
13.7
22.2
7.5
13.7
13.7
AP (mm)
1741.0
470.00
1109.0
2240.0
-
-
-
-
PET(mm)
949.0
906.0
264.80
203.1
715.0
715.0
1263.0
1088.0
-
-
-
-
-
-
-
-
103.1
89.80
-6.7
200.4
-
-
-
-
MTCM ( C)
o
K
AAE (mm)
Im
s13
EQ (oC/mm)
14.5
7.50
8.6
25.3
-
-
-
-
13.5
2.23
9.1
17.0
6.6
1.40
4.5
8.9
104.3
23.39
59.3
144.2
47.7
9.75
28.0
65.5
CI ( C·month)
-2.7
4.35
-13.9
0.0
-28.3
9.36
-42.7
-12.4
o
13.5
2.22
9.1
17.0
7.2
1.07
5.2
8.9
F. sylvatica
AMT (oC)
o
WI ( C·month)
o
ABT ( C)
o
MTWM ( C)
23.0
2.58
17.4
28.8
16.9
1.75
12.5
20.2
o
4.7
2.75
-1.0
9.8
-2.7
2.13
-6.3
1.6
ART ( C)
18.2
2.60
14.2
23.0
19.6
2.72
12.8
22.9
MTCM ( C)
o
K
27.7
6.61
16.1
41.0
25.8
9.74
4.7
41.2
AP (mm)
905.9
387.48
573.0
1864.0
1272.3
492.57
524.0
1958.0
PET(mm)
749.8
82.24
608.4
946.5
577.6
52.29
445.9
652.1
AAE (mm)
497.1
55.8
414.4
625.1
496.7
19.2
466.3
549.0
38.1
65.17
-8.2
211.4
119.3
82.62
-2.7
240.3
29.0
9.54
9.3
39.4
16.8
9.55
5.4
34.9
AMT (oC)
10.2
2.74
7.5
14.3
6.5
1.98
4.9
9.9
o
78.3
26.52
56.3
126.1
46.3
14.50
29.9
69.1
CI ( C·month)
-16.2
10.20
-27.7
-0.8
-28.5
10.45
-37.6
-10.7
o
10.4
2.57
8.0
14.4
7.1
1.68
5.4
9.9
Im
o
EQ ( C/mm)
F. orientalis
WI ( C·month)
o
ABT ( C)
o
MTWM ( C)
20.5
3.73
17.6
27.8
16.1
1.95
13.3
18.4
o
MTCM ( C)
-1.5
3.64
-6.0
4.2
-3.2
2.26
-5.0
0.7
o
ART ( C)
22.0
4.88
16.6
29.9
19.3
2.25
16.6
22.2
K
38.1
12.11
22.4
57.0
31.5
6.58
22.4
40.1
AP (mm)
744.8
335.68
526.0
1261.0
911.8
246.55
709.6
1261.0
PET(mm)
717.2
175.77
524.1
1044.1
668.9
9.14
653.8
676.8
AAE (mm)
486.5
144.5
344.7
673.6
460.3
4.9
452.1
464.6
15.4
31.20
-5.1
76.7
23.8
29.93
7.3
76.7
32.7
9.77
17.2
44.7
27.9
6.53
17.2
32.4
Im
o
EQ ( C/mm)
s14
Table S2. Loadings of climatic variables derived from Principal Component Analysis for the first three principal components
for distribution limits of world beech species. For abbreviations of climatic variables see Table S1.
Variable
Lower (southern) limit
Upper (northern) limit
PCA 1
All beech species
AMT (oC)
WI (oC·month)
CI (oC·month)
ABT (oC)
MTWM (oC)
MTCM (oC)
ART (oC)
K
AP (mm)
PET (mm)
AAE (mm)
Im
EQ (oC/mm)
0.97
0.95
0.85
0.97
0.73
0.96
-0.73
-0.42
-0.34
0.54
0.42
-0.50
0.28
PCA 2
0.15
0.14
0.16
0.15
0.06
0.23
-0.26
-0.22
0.83
-0.22
-0.05
0.84
-0.89
PCA 3
0.00
0.03
-0.10
0.00
0.16
-0.14
0.29
0.68
0.35
0.68
0.84
-0.08
-0.25
PCA 1
PCA 2
PCA 3
0.98
0.89
0.91
0.93
0.59
0.88
-0.61
0.22
0.23
0.63
0.62
0.03
0.02
0.24
-0.17
0.15
0.34
-0.25
0.41
0.16
-0.83
0.31
0.23
-0.92
-0.12
0.13
-0.31
0.02
0.51
-0.38
0.62
0.80
0.38
0.22
0.37
0.29
-0.13
0.89
-0.35
Chinese beeches
AMT (oC)
WI (oC·month)
CI (oC·month)
ABT (oC)
MTWM (oC)
MTCM (oC)
ART (oC)
K
AP (mm)
PET (mm)
AAE (mm)
Im
0.96
0.93
0.90
0.96
0.58
0.97
-0.73
-0.61
0.00
-0.32
-0.33
0.21
0.20
0.24
0.03
0.21
0.22
0.12
0.01
0.07
-0.69
0.62
0.62
-0.95
0.07
0.03
0.20
0.06
0.01
0.09
-0.09
0.28
0.71
0.70
0.69
0.10
0.99
0.92
0.95
0.97
0.67
0.93
-0.39
-0.09
0.33
0.09
0.08
0.14
0.12
0.30
-0.13
0.21
0.50
-0.17
0.56
0.53
-0.63
0.83
0.83
-0.82
0.03
0.20
-0.20
0.08
0.52
-0.30
0.70
0.77
0.25
-0.24
-0.25
0.42
EQ (oC/mm)
-0.27
0.85
-0.41
-0.23
0.86
-0.35
F. crenata
AMT (oC)
WI (oC·month)
CI (oC·month)
ABT (oC)
MTWM (oC)
MTCM (oC)
ART (oC)
K
AP (mm)
PET (mm)
AAE (mm)
Im
0.91
0.71
0.96
0.82
0.38
0.95
-0.45
0.05
0.38
0.43
0.43
0.25
0.42
0.68
-0.10
0.55
0.89
-0.09
0.82
0.45
-0.70
-0.70
-0.70
-0.40
-0.03
0.07
-0.17
0.00
0.09
-0.21
0.25
0.50
0.59
-0.39
-0.39
0.87
0.95
0.80
0.95
0.88
0.43
0.92
-0.83
-0.06
0.69
-0.36
-0.36
0.63
0.31
0.44
-0.21
0.39
0.58
0.23
-0.02
-0.42
-0.62
0.73
0.73
-0.51
0.01
0.34
-0.20
0.21
0.62
-0.27
0.54
0.86
0.09
0.08
0.08
0.21
EQ (oC/mm)
-0.38
0.47
-0.77
-0.69
0.60
-0.10
F. japonica
AMT (oC)
WI (oC·month)
CI (oC·month)
ABT (oC)
MTWM (oC)
0.92
0.86
0.96
0.92
0.46
0.38
0.46
0.16
0.38
0.84
-0.01
-0.06
0.08
-0.02
-0.09
0.93
0.84
0.98
0.92
0.40
0.31
0.47
0.07
0.33
0.80
-0.18
-0.24
0.07
-0.19
-0.38
s15
MTCM (oC)
ART (oC)
K
AP (mm)
PET (mm)
AAE (mm)
Im
0.97
-0.61
-0.52
0.64
0.30
0.30
0.55
0.09
0.65
0.08
-0.68
-0.31
-0.31
-0.63
0.05
-0.14
0.11
-0.19
0.88
0.88
-0.49
0.97
-0.71
-0.40
0.71
0.33
0.33
0.57
-0.02
0.57
0.53
-0.56
0.67
0.67
-0.75
-0.06
-0.20
0.23
0.32
0.66
0.66
-0.02
EQ (oC/mm)
F. grandifolia
AMT (oC)
WI (oC·month)
CI (oC·month)
ABT (oC)
MTWM (oC)
MTCM (oC)
ART (oC)
K
AP (mm)
PET (mm)
AAE (mm)
Im
-0.43
0.73
0.39
-0.46
0.77
-0.25
0.97
0.97
-0.08
0.97
0.54
0.97
-0.86
-0.68
0.57
0.91
0.94
0.23
-0.19
-0.19
-0.36
-0.19
-0.18
-0.14
0.08
0.19
0.81
-0.30
0.09
0.97
0.11
0.11
0.24
0.11
0.79
-0.18
0.49
0.68
0.12
0.27
0.00
0.06
0.75
0.98
0.39
0.97
0.90
0.15
0.26
0.38
-0.67
0.98
0.82
-0.78
-0.66
-0.02
-0.91
-0.14
0.09
-0.98
0.93
0.88
-0.35
-0.05
-0.16
-0.29
0.08
0.19
0.01
0.18
0.33
-0.11
0.25
0.27
0.65
0.17
0.26
0.55
EQ (oC/mm)
European beeches
AMT (oC)
WI (oC·month)
CI (oC·month)
ABT (oC)
MTWM (oC)
MTCM (oC)
ART (oC)
K
AP (mm)
PET (mm)
AAE (mm)
Im
-0.57
-0.81
-0.04
0.86
0.25
-0.43
0.95
0.89
0.92
0.94
0.72
0.94
-0.44
-0.43
0.04
0.58
0.31
-0.13
0.24
0.38
-0.26
0.27
0.56
-0.23
0.73
0.72
-0.91
0.02
-0.49
-0.86
0.17
0.20
-0.04
0.17
0.38
-0.12
0.45
0.46
0.35
-0.48
-0.07
0.44
0.63
0.89
0.04
0.79
0.90
-0.20
0.73
0.56
-0.63
0.37
0.34
-0.73
0.76
0.40
0.95
0.59
0.20
0.88
-0.52
-0.08
0.55
0.46
-0.32
0.44
-0.11
0.06
-0.29
-0.02
0.16
-0.39
0.39
0.82
0.46
0.32
-0.69
0.34
EQ (oC/mm)
-0.09
0.87
-0.43
0.76
-0.42
-0.34
s16
Table S3. Coefficient of correlation between annual mean temperature (AMT) and other thermal variables in the distribution
range of beech species. Sample size is 310 for the upper (northern) limit, and 292 for lower (southern) limit.
Parameter
Lower limit
Upper limit
Warmth index (oC.month)
0.99
0.93
Coldness index (oC.month)
0.88
0.95
Annual biotemperature (oC)
1.00
0.96
Mean temperature for the warmest month (oC)
0.85
0.61
o
Mean temperature for the coldest month ( C)
0.98
0.93
Minimum mean temperature (oC)
0.95
0.90
Annual range of temperature (oC)
-0.79
-0.69
s17