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Journal of Biogeography (J. Biogeogr.) (2006) 33, 1804–1819
ORIGINAL Climatic limits for the present distribution ARTICLE of beech (Fagus L.) species in the world Jingyun Fang1* and Martin J. Lechowicz2
1Department of Ecology, College of ABSTRACT Environmental Sciences, and Centre for Aim Beech (Fagus L., Fagaceae) species are representative trees of temperate Ecological Research & Education, Peking University, Beijing 100871, China and 2Biology deciduous broadleaf forests in the Northern Hemisphere. We focus on the Department, McGill University, 1205 Dr distributional limits of beech species, in particular on identifying climatic factors Penfield Avenue, Montreal, Quebec, Canada associated with their present range limits. H3A 1B1 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. 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 *Correspondence: Jingyun Fang, Department of temperature was probably a limiting factor for southward extension of American Ecology, College of Environmental Sciences, Peking University, Beijing 100871, China. beech, while growing season warmth seemed critical for its northward E-mail: [email protected] distribution. Although the present distribution of beech species corresponded
1804 www.blackwellpublishing.com/jbi ª 2006 The Authors doi:10.1111/j.1365-2699.2006.01533.x Journal compilation ª 2006 Blackwell Publishing Ltd
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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.
Two beech species, F. crenata and F. japonica, are native to INTRODUCTION Japan. The former is distributed from Kyushu (c. 30.5° N) to Beech (Fagus L., Fagaceae) are among the most representative southern Hokkaido (c. 42.8° N), whereas the latter is limited trees in the temperate deciduous broadleaf forests of the to the south of Iwate-ken (Horikawa, 1972; Miyawaki, 1980– Northern Hemisphere (Shen, 1992; Denk, 2003). The genus 89). Beech forest in Japan falls into two broad forest types: the includes ten primary species and two minor segregates (Willis, Pacific-Ocean-side type that grows intermixed with many 1966) broadly distributed in three isolated regions: East Asia, other temperate tree species, and the Japan-Sea-side type Europe and West Asia, and North America. Six species occur where beech is frequently dominant (Yamazaki, 1983; Maeda, in East Asia (F. engleriana, F. longipetiolata, F. lucida and 1991). Forests with F. multinervis are restricted to a small F. hayatae in China and F. crenata and F. japonica in Japan) island, Ulreung-do in South Korea, and they are more or less (Horikawa, 1972; Editorial Committee for Flora of China, similar in community composition and structure to Japanese 1999), two in Europe and West Asia (F. sylvatica and beech forests of the Japan-Sea-side type (Kim et al., 1986; Kim, F. orientalis) (Jalas & Suominen, 1972–91), and one in North 1988). America (F. grandifolia) (Little, 1965, 1979). Some studies have In North America, F. grandifolia is one of the most recognized two additional beech species, one on a small island widespread species among temperate trees, covering almost off Korea: F. multinervis (Kim et al., 1986; Kim, 1988) and all the temperate zone along the Appalachian Mountains from another one in the north-eastern mountains of Mexico: the northern edge of the subtropical zone almost to the F. mexicana (e.g. Miranda & Sharp, 1950; Rzedowski, 1983; southern edge of the boreal zone (USDA Forest Service, 1975). Maycock, 1994; Peters, 1995). Fagus multinervis is a segregate A number of studies have shown large differences in commu- of F. engleriana in China (Okubo et al., 1988; Peters, 1992; nity composition and structure in different climatic regions Shen, 1992; Denk, 2003); F. mexicana is a segregate of the (Braun, 1967; Barnes, 1991; Maycock, 1994). Based on North American F. grandifolia. For the purposes of the present differences in geographical distribution and morphological study we accept the validity of both segregate species. characters, three races of F. grandifolia (grey beech, white In East Asia, beech occurs primarily in mountain areas. All beech, and red beech) have been identified (Camp, 1951; Chinese beeches are restricted to remote subtropical/warm- Braun, 1967; Maycock, 1994) but these have never been given temperate mountain areas, ranging from south China to the species status. Yangtze River (c. 33° N), and from the south-east coast of the Another beech species in North America, F. mexicana, is East China Sea to the eastern edge of the Tibetan Plateau found in only four montane localities in north-eastern Mexico (Tsien et al., 1975; Wu, 1980; Hou, 1988; Hong & An, 1993; (Little, 1965; Rzedowski, 1983). Its community characters are Cao et al., 1995). Compared with the continuous distribution more or less similar to those of Chinese beech forests, usually of other Chinese beeches, F. hayatae is isolated in three very mixing with many species of Quercus, Magnolia, Acer and limited mountain areas: north-east Taiwan, eastern Zhejiang Carya (Miranda & Sharp, 1950; Rzedowski, 1983; Peters, 1995; and north-west Sichuan. Unlike beech in other regions, Williams-Linera et al., 2000). Chinese beeches rarely form pure stands but typically occur In Europe, F. sylvatica spreads from Sicily in southern Italy mixed with various deciduous broad-leaved trees (Betula, Acer, (c. 37.7° N) to Bergen in south Norway (c. 60.7° N) (Jalas & Liriodendron, Davidia, Tilia, Carpinus and Nyssa), evergreen Suominen, 1972–91; Feoli & Lagonegro, 1982; Jahn, 1991); this broad-leaved trees (Lithocarpus, Cyclobalanopsis, Manglietia is the most widely distributed of all beech species. Fagus and Castanopsis) and evergreen needle-leaved (Tsuga) trees orientalis replaces F. sylvatica in a small region of southeast (Wu, 1980; Hou, 1983, 1988; Hsieh, 1989; Cao, 1995; Fang Europe and spreads into West Asia: northern Turkey, the et al., 1996). Caucasus and the Elburz Mountains of northern Iran, where
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the climate is more or less continental (Jalas & Suominen, Answering these questions is not only important in biogeog- 1972–91; Davis, 1982). raphy, but also will provide a firmer basis for predicting the The three regions where beech species occur are controlled effects of climate change on the future distribution of beech by different air masses and have various topographic settings. A species. Focusing on distribution limits (lower or southern and monsoon climate prevails in East Asia with cold and dry air upper or northern limits) of the world beech species, we masses from Siberia predominant in winter and hot, humid therefore compare their climatic spaces to detect possible subtropical highs from the Pacific Ocean predominant in factors limiting their distributions. summer. This causes abundant summer rainfall and high temperature after a dry and cold winter (Arakawa, 1969; Shen, DATA AND METHODS 1986). The Himalayan Range, higher to the west and lower eastwards in mainland China, intensifies this climatic difference Data sets for beech distribution between winter and summer (Chang, 1983; Fang et al., 1996) on the continent in comparison to the Japanese archipelago. Data sets for beech distribution were assembled by reviewing In eastern North America, the climate is governed by two ecological, botanical and geographical journals, books and major circulations: cold and dry air masses from the Arctic and reports to record the geographical location, lower and upper moist warm air masses from the southern tropical seas, and elevational limits, and associated information (e.g. topography, therefore the climatic patterns resemble more or less those in climate and community characteristics) for each site where a East Asia. However, there is no topographic barrier akin to the beech species was reported to occur. Vertical distribution ranges Himalayas to block meridional air mass exchange (Lydolph, were reported especially frequently in East Asia, but less often in 1985). Europe and North America where topographic relief at the range The European and West Asian region has a narrower limits of beech species is less extreme. We assembled data for seasonal cycle of temperatures and rainfall than the two other F. grandifolia from a silvicultural atlas (USDA Forest Service, regions. In Europe, the air masses resemble those of western 1975) and ancillary literature reports. We found relatively little Northern America, but are much influenced by topography. data for F. sylvatica and F. orientalis; the map of Jalas & The east–west trend of European mountain ranges reduces Suominen (1972–91) yielded data on northern range limits for northward invasion of large subtropical bodies of warm air, these species, but high relief and uncertain elevational distribu- and the Mediterranean Sea also plays a role in the generation tions left the southern limits undefined. and routes of cyclonic storms (Lydolph, 1985). When latitude and longitude were not reported for study The floristic, climatic and topographic patterns of these locations, we used gazetteers to georeference the sites: (1) regions where beech occurs have attracted the attention of China Places Name (Anon., 1986) and Atlas of Land Use in previous investigators. For example, using Fagus pollen data China (Editorial Committee of 1/1,000,000 Land-use Map of and monthly mean temperature, Huntley et al. (1989) studied China, 1990) for Chinese beeches; (2) Gazetteer to AMS climatic control of beech distribution and abundance in 1 : 250,000 Maps of Japan (Corps of Engineers, US Army, Europe and North America. Peters (1992) and Peters & 1956) and The National Atlas of Japan (Geographical Survey Poulson (1994) compared tree growth, and community Institute, 1977) for Japanese beeches; (3) American Places structure and dynamics of the world beech species. Maycock Dictionary (Abate, 1994) for American beech; and (4) Atlas of (1994) documented detailed information on differences in the World (Times, 1992) for others. In total we identified 350 community composition between North American and sites with reliable data for the lower/southern limits and 353 Japanese beeches. Iverson & Prasad (1998) used forest sites for the upper/northern limits of beech species (Table 1; inventory data to estimate the climate envelope for Fang, 2003). For detailed information on the location and F. grandifolia and predict its range extension under climate elevation of distribution limits for each beech species, see change. Piovesan & Adams (2001) compared masting beha- Appendix S1 in Supplementary Material. viours of beech from Europe, eastern North America and Japan, and discussed their links to climatic variations. In Climatic data addition, some case studies on relationships between beech distribution and climates have been conducted, but most are We used monthly mean temperatures and monthly precipita- restricted to single species in a region (e.g. Birks, 1989; Cao tion records from the following data sources: (1) China et al., 1995; Sykes et al., 1996). Meteorological Agency (1984); (2) Japan Meteorological Taken together, the studies mentioned above provide useful Agency (1972); (3) Central Meteorological Office of Korea ecological comparisons among beech species in the three (1972); (4) National Climatic Center, NOAA (1983) for USA; regions where they occur, but many questions remain about (5) Atmospheric Environmental Service, Environment Canada relationships between beech distribution and climate. For (1982) for Canada; and (6) Wernstedt (1972) for Mexico, example, how different are the geographical patterns shown in Europe and West Asia. The period of record for most stations the three separate regions where beech species occur? What was 1951–80 or 1941–71. climatic factors control such patterns? Can contemporary Temperatures at altitudes of the upper and lower climate explain the present distribution of beech species? elevation limits of beech were estimated for each locality
1806 Journal of Biogeography 33, 1804–1819 ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd 中国科技论文在线 http://www.paper.edu.cn
Climatic limits for world beech distribution
Table 1 Sample size for distribution limits of all world beech Coldness and winter temperature summation species. Data set for Fagus grandifolia was mainly extracted from its southern and northern distributional edges, and the northern A number of studies have shown the importance of minimum limit of F. sylvatica actually lies to the north-east according to the winter temperatures in controlling distributional limits of map of Jalas & Suominen (1972–91) plant species (e.g. Sakai & Weiser, 1973; Woodward, 1987; Sykes et al., 1996; Pederson et al., 2004). Mean temperature for Lower Upper the coldest month (MTCM) is often used as a surrogate for the Species Distribution limit limit minimum winter temperature (Solomon, 1986; Ohsawa, 1990; F. crenata Pacific Ocean side, Japan 42 46 Prentice et al., 1992; Sykes et al., 1996). In this study, the 2 F. japonica Japanese Sea side, Japan 28 22 coefficient of determination (R ) between these two variables F. engleriana South and southwest China 34 36 was 0.94 (P < 0.0001, n 602), so we also used the more ¼ F. hayatae Taiwan, Zhejiang, 8 10 readily available MTCM as a measure of coldness. and Sichuan, China Previous studies also suggest that cumulative winter tem- F. longipetiolata South, east, and 55 55 perature is important for the northward/upward distributions southwest China of warm-temperate and tropical tree species (Kira, 1948, 1991; F. lucida East, and Central China 60 54 Hattori & Nakanishi, 1985; Fang & Yoda, 1991), and for spring F. multinervis Ulreung-do, South Korea 1 1 budburst of many northern tree species (Prentice et al., 1992; F. grandifolia* Eastern North America 80 80 Lechowicz, 2001). We used Kira’s Coldness Index (CI) (Kira, F. mexicana Northeastern Mexico 4 4 F. sylvatica Europe 32 40 1948, 1991) to express the cumulative winter temperature: F. orientalis Southeast Europe 6 5 CI 5 T for months in which T < 5 C 3 and West Asia ¼ À ð À Þ ð Þ ð Þ Total 350 353 X
*Most data for southern or northern extremes. Annual mean temperature and mean temperature for the Most data for northern edge. warmest month Annual mean temperature (AMT) and mean temperature for the warmest month (MTWM) are often used to explain using a mean lapse rate of 0.6 °C per 100 m (Barry, 1992) northward and upward distribution of northern tree species together with data from the nearest climatic station. The (Walter, 1979; Tuhkanen, 1980; Ohsawa, 1990, 1991). rainfall data for each beech site were estimated from relationships between precipitation and elevation regressed Climatic continentality by using rainfall and altitude data at more than five climatic stations close to the beech site. Similar to Huntley et al. Wolfe (1979), Tuhkanen (1980), Ohsawa (1990) and Fang (1989), distances between beech sites and climatic stations et al. (1996) have addressed the effect of climatic continentality were less than 0.5° of latitude and 1.0° of longitude (c. 50– on distribution of some tree species. We used the annual range 100 km radius). In total, we secured reliable climate data for of monthly mean temperatures (ART) and Gorcynski’s (1922) 292 sites at the lower/southern limit and 310 sites at the continentality index (K): upper/northern limit of beech species. R K 1:7 20:4 4 ¼ Â sin L À ð Þ Climatic parameters where R is the annual range of monthly mean temperature in °C and L is the latitude in degrees. Growing season warmth
The distribution limits of many tree species are closely related Moisture variables to growing season temperature (e.g. Kira, 1945, 1991; Hold- ridge, 1947; Tuhkanen, 1980; Woodward, 1987; Prentice et al., To assess moisture regime, we considered annual precipita- 1992; Sykes et al., 1996). We used Kira’s Warmth Index (WI) tion (AP, mm), potential evapotranspiration (PET, mm), (Kira, 1945, 1991) and Holdridge’s annual biotemperature annual actual evapotranspiration (AAE, mm), moisture )1 (ABT) (Holdridge, 1947) as proxies for growing season index (Im), and the Ellenberg quotient (EQ, °C mm ). warmth, given respectively by: With the exception of AP and EQ, all moisture parameters were estimated by the Thornthwaite (1948) method, which WI T 5 for months in which T > 5 C 1 uses two climatic variables commonly recorded at climatic ¼ ð À Þ ð Þ ð Þ X stations around the world: monthly mean temperature and T monthly precipitation. The Thornthwaite index has proven a ABT for months in which 0 < T < 30 C 2 ¼ 12 ð Þ ð Þ P good correlate of vegetation and plant distribution at both
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regional and global scales (e.g. Mather & Yoshioka, 1968; Supplementary Material. The descriptions that follow are Fang & Yoda, 1990; O’Brien, 1993; Frank & Inouye, 1994). supported by both Table 2 and Table S1. For calculation of PET, AA and Im, see Fang (1989) and In East Asia, in spite of large fluctuations within and among Fang & Yoda (1990). species, average values of growing season warmth at the lower The EQ is the ratio of the MTWM to annual precipitation limits of beech distribution were between 74.8 °C (F. crenata) and is frequently used to show the climatic limit of beech in and 115.9 °CÆmonth (F. longipetiolata) for Warmth Index Europe (Ellenberg, 1986; Jahn, 1991). According to Jahn (WI) and between 9.8 and 14.3 °C for annual biotemperature (1991), values below 20 show a pure ‘beech climate’; the (ABT) (Table 2). Interestingly, most of the seven Asian beech competitive vigour of beech slowly decreases with an increase species had lower limits associated with very similar average from 20 to 30; and in Europe beech disappears in regions with thermal parameters (AMT of 11.7–12.8 °C, WI of 91.8– an EQ over 30: 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 Warmest month’s mean temperature in C EQ 1000 species have widely disparate ranges (F. japonica in Japan, ¼ Annual precipitation mm  ð Þ F. multinervis in Korea and others in China). Only two species, 5 ð Þ 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 Principal components analysis most species with an AMT value of 7.3–9.2 °C, WI of 56.0– We used principal components analysis (PCA) (Wilks, 1995) 70.3 °CÆmonth, and ABT of 8.1–9.5 °C (Table 2), but to: (1) examine which climatic variables are most associated F. crenata is an exception. Fagus crenata has a WI value with the distribution of beech species, and (2) compare the of 45.1 °CÆmonth, approximately the climatic threshold climatic space at distributional limits for beech species in the (45 °CÆmonth) of the cool-temperate zone defined by Kira three regions. (1991). To assess the influence of the diverse climatic variables, we With regard to moisture regime, all beech species locations compared the eigenvalues of different PCA axes for each in East Asia fall within an average Im range of 61.4–189.9 for species and the axis loadings of climatic variables for the their lower limits and 70–241.3 for their upper limits, denoting species that have a large sample size. In general, variables with a humid or perhumid climate according to the Thornthwaite bigger loadings are considered more important in placement (1948) system (Table 2). along PCA axes when variables are standardized to allow for In North America, F. grandifolia showed the widest climatic differences in units and magnitude (Wilks, 1995). We analysed space, with average WI ranging from 50.7 °CÆmonth (northern 13 climatic variables (AMT, WI, CI, ABT, MTWM, MTCM, limit) to 173.4 °CÆmonth (southern limit), and ABT from 7 to ART, K, AP, PET, AAE, Im and EQ) using PC-ORD software 19.5 °C. This spans two bioclimatic zones: cool-temperate (McCune & Mefford, 1999). To minimize the differences (45–85 °CÆmonth for WI and 6–12 °C for ABT) and warm- associated with differing units and magnitude, all the variables temperate zone (85–180 °CÆmonth for WI and over 12 °C for were standardized: ABT). Although F. mexicana is distributed much farther south than F. grandifolia, it has a smaller climatic range (117.1– xi x x0 À 6 127.3 °CÆmonth in WI and 14.8–15.6 °C in ABT) and much i ¼ r ð Þ smaller thermal variables at its lower limit than those of where xi and xi0 are the original and standardized value of a F. grandifolia (Table 2). Fagus grandifolia has more moist climatic variable for the ith site, x is average of the climatic conditions at its northern limit with an average Im value of variable for all the sites, and r is the standard deviation of the 91.1 compared with 40.3 at its southern limit. Mexican beech climatic variable. To compare climate spaces among the beech has a much larger Im average value (Im 103) at its ¼ species, we used the eigenvalues of the first two components elevational and southern limit. for each beech site. To illustrate the climate space for species In Europe, the climatic spaces of beech species span almost with sufficiently large sample sizes, we calculated a 50% the entire temperate zone. Growing season warmth in the Gaussian bivariate confidence ellipse (ELL) using sysgraph range of beech ranged from 47.7 to 104.3 °CÆmonth for (SYSTAT Inc., 1996). 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 RESULTS climatic range despite having the widest latitudinal range among the world beech species (spanning c. 23°) (Jalas & Climatic statistics at the distribution limits Suominen, 1972–91). In comparison with beech species in Table 2 lists the average for thermal and moisture climatic other regions, the moisture regime in the area of European parameters at lower/southern and upper/northern limits for beech species is relatively dry, with average precipitation of all beech species; the details for the statistics (average, SD and 905.9 mm, mean Im value of 38, and mean EQ of 29 °C mm)1 range) of these variables are displayed in Table S1 in the for the lower limit of F. sylvatica, and 1272.3 mm, 119.3 and
1808 Journal of Biogeography 33, 1804–1819 ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd 中国科技论文在线 ª Journal 2006 of The Biogeograph Authors.
Journal 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: y
33 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: ,
1804–1819 continentality index; AP: annual precipitation; PET: annual potential evapotranspiration; AAE: actual annual evapotranspiration; Im: moisture index; and EQ: Ellenberg quotient. – indicates no compilation estimation
Lower/upper AMT WI CI ABT MTWM MTCM ART AP PET AAE EQ Species limits (°C) (°CÆmonth) (°CÆmonth) (°C) (°C) (°C) (°C) K (mm) (mm) (mm) Im (°C mm)1) ª
2006 F. engleriana Lower 12.4 99.0 )10.1 12.5 23.0 1.1 21.9 53.3 1230 805.4 803.3 61.4 21.7 Upper 7.3 56.0 )28.1 8.1 17.8 )3.9 21.7 52.8 1366 795.6 793.5 70.0 19.4 Blackwell F. hayatae Lower 12.8 99.4 )6.4 12.8 22.4 2.1 20.3 50.7 1670 914.6 914.6 85.5 17.0 Upper 9.2 67.0 )16.6 9.5 18.6 )0.7 19.3 46.3 2233 887.0 887.0 109.8 13.2 F. longipetiolata Lower 14.3 115.9 )4.1 14.3 24.2 3.5 20.8 54.1 1421 845.4 831.8 77.5 19.0 Publishing Upper 8.9 66.2 )19.8 9.2 18.9 )2.0 20.9 54.2 1738 621.2 619.2 132.6 14.7 F. lucida Lower 12.7 100.6 )8.2 12.8 23.0 1.5 21.5 57.8 1419 874.0 862.7 70.0 19.5 Upper 8.8 66.4 )21.3 9.2 19.1 )2.4 21.5 57.8 1757 674.4 671.7 168.5 13.8
Ltd F. crenata Lower 9.5 74.8 )21.3 9.8 21.8 )2.1 23.9 48.5 2047 706.2 706.2 189.9 12.5 Upper 5.0 45.1 )45.2 6.7 17.6 )6.8 24.4 48.8 2660 606.8 606.8 241.3 9.5 F. japonica Lower 11.7 91.8 )12.0 11.7 23.9 0.3 23.7 48.8 1805 746.6 746.6 142.0 15.3 Upper 9.0 70.3 )21.9 9.4 21.2 )2.2 23.4 48.6 2329 716.4 716.4 206.5 11.5 F. multinervis Lower 11.5 90.2 )11.9 11.5 23.4 0.1 23.3 45.0 1485 702.6 702.6 111.3 16.1 Upper 7.6 59.0 )27.5 8.3 19.5 )3.8 23.3 45.0 – – – – –
F. grandifolia Lower 19.5 173.4 0 19.5 27.5 10.4 17.1 36.8 1426 1024.0 996.9 40.3 19.5 Climatic Upper 4.2 50.7 )60.5 7.0 18.4 )11.4 29.8 49.5 1021 537.8 530.5 91.1 18.5 F. mexicana Lower 15.6 127.3 0 15.6 19.6 10.5 9.1 22.2 1741 949.0 906.0 103.1 14.5 Upper 14.8 117.1 0 14.8 18.7 9.7 9.1 22.2 – – – – – limits F. sylvatica Lower 13.5 104.3 )2.7 13.5 23.0 4.7 18.2 27.7 906 749.8 497.1 38.1 29.0
Upper 6.6 47.7 )28.3 7.2 16.9 )2.7 19.6 25.8 1272 577.6 496.7 119.3 16.8 http://www.paper.edu.cn for F. orientalis Lower 10.2 78.3 )16.2 10.4 20.5 )1.5 22.0 38.1 745 717.2 486.5 15.4 32.7 Upper 6.5 46.3 )28.5 7.1 16.1 )3.2 19.3 31.5 912 668.9 460.3 23.8 27.9 world beech distribution 1809 中国科技论文在线 http://www.paper.edu.cn
J. Fang and M. J. Lechowicz
16.8 °C mm)1 for AP, Im and EQ for its upper/northern limit. beech species. Among thermal variables, AMT, WI and ABT For F. orientalis, AP, Im and EQ were, respectively, 744.8 mm always showed larger loadings at both lower/southern and and 32.7 °C mm)1 for its lower limits, and 911.8 mm, 23.8 upper/northern limits, but CI and MTCM exhibited a large and 27.9 °C mm)1 for its upper limit. It is apparent that value for world beech species (Table S2). This indicates that climate at the lower limits of both beeches is rather dry, close growing season warmth is most important for beech distribu- to the EQ threshold of 30 °C mm)1 suggested by Jahn (1991). tion, and CI and MTCM are also closely coupled with the The EQ value of European beeches is much smaller than for potential for their range expansion. The second PCA axis beech species in other regions: ranging from 15 to 20 °C mm)1 exhibited a different trend: for the lower/southern limit the for the lower/southern limit and from 11.5 to 19.4 °C mm)1 loadings of AP, Im and EQ were 0.83 mm, 0.84 and for the upper/northern limit for other regions, with the )0.89 °C mm)1, while for the upper limit/northern they were smallest value (12.5 and 9.5 °C mm)1 for its lower and upper )0.83 mm, )0.92 and 0.89 °C mm)1, respectively (Table S2). limits, respectively) for F. crenata (Table 2). 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 Climatic factors controlling beech species limit decreases with an increase of precipitation. distributions In spite of these overall trends, some component loadings We used the 13 climatic variables in a PCA to identify limiting varied across regions, suggesting that limiting climatic factors factors for beech distribution. To assess regional variations in shifted somewhat among beech in different regions. In general, the relationships between geographic distribution and climatic both the lower/southern and upper/northern limits in all three space, we combined four Chinese beeches and two European beech regions depended on seasonal thermal regimes (loadings beeches into the same group because they have similar climate of growing season warmth (WI and ABT) on the first PCA axis ranges (cf. Table S1). The two Japanese beeches occur were largest, and winter temperatures (CI and MTCM) also primarily in two different climatic regions (Japan Sea side showed a large influence), but the two Japanese beech species and Pacific Ocean side), and thus we did not deal with them as were an exception. For the second PCA axis, the loadings of a single group. The first two principal components account for moisture climate variables were largest for the lower/southern 70% and 68% of overall variance for southern/lower and limit, indicating the general influence of a moisture gradient. northern/upper limits of all species, respectively (Table 3). However, for the upper/northern limit, the climatic variables Accordingly we considered the loadings on these axes to be with the largest loading showed a large regional difference. The most important in delimiting beech distribution. For each second axis for Chinese beech suggests a moisture regime beech region, the first two PCA axes explained more than gradient, while the loading of winter coldness (CI) was largest 70% of the variance; for example, 80% and 87% for the for European beeches, and MTCM had the largest loading for southern and northern limits of Amerian beech, respectively, American beech. and 73–80% for F. japonica and the European beeches. For the lower limit of the two Japanese beeches, winter The first PCA axis represents a thermal gradient, and the temperature (CI and MTCM) had a large loading on the first second a moisture gradient in the overall distribution of world axis, summer temperature (MTWT) on the second. This beech species (Table S2 in Supplementary Material). In general suggests that moisture regime is not a limiting factor for the the thermal climate played a leading role and precipitation a downward distribution of these two species due to abundant secondary role in controlling the large-scale distribution of 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 Table 3 Proportion (%) of cumulative variance on the first four MTWT and moisture indices (Im and EQ) for F. japonica were principal components in a principal components analysis of the distribution limits for world beech (Fagus) species largest. It is noteworthy that PET, an indicator of total solar energy, Lower (southern) limit Upper (northern) limit showed the largest loading (0.98) on the first axis for the northern limit of American beech; this is consistent with the PCA PCA PCA PCA PCA PCA PCA PCA correlation between PET and vegetation distribution and Species 1 2 3 4 1 2 3 4 overall tree species richness in North America (Stephenson, Chinese 46.34 69.89 83.71 96.69 41.19 73.56 89.04 96.12 1990; Francis & Currie, 2003). beeches F. crenata 37.60 71.86 90.25 97.27 50.48 74.57 88.08 96.49 F. japonica 47.98 72.96 88.70 98.34 49.27 80.20 90.85 97.71 DISCUSSION F. grandifolia 59.25 79.76 91.69 98.77 54.46 86.65 96.99 99.12 European 43.65 76.86 87.73 94.73 40.45 71.79 88.01 97.26 Zonal distribution of world beech species beeches Although most beech species are considered typical trees of the All beech 50.71 69.89 84.98 94.28 44.89 67.57 83.72 93.79 temperate zone, they in fact showed different climatic ranges in species the three regions where they occur (Table 2; Table S1). In East
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Asia, most species, excluding F. crenata, concentrated within a more important for southward migration. Cao et al. (1995) climatic range of 90–116 °CÆmonth in WI and 11.7–14.3 °C in demonstrated the importance of the moisture deficit in the ABT in their lower limits, and 56–70 °CÆmonth in WI and northern range of beech species and the importance of high c. 8.1–9.5 °C in ABT in their upper limits. This is to say, the temperature and insufficient water supply in the south. boundaries for most Asian beech species are located in warmer Focusing on the relationship between Fagus- and Tsuga- places than the cool-temperate zone defined by Kira (1945, dominated forests, Fang et al. (1996) suggested possible effects 1991) and Holdridge (1947); the climatic parameters at the of the annual temperature range (ART) and high winter southern/lower edge of the cool-temperate zone was set at temperature on beech distribution. They found that the beech- 85 °CÆmonth in WI by Kira, and 12 °C in ABT by Holdridge, dominated forests did not appear in places where hemlock and 45 °CÆmonth in WI and 8 °C in ABT at the northern or dominates, and that their boundary was consistent with an upper limit. This suggests that beech species occupy an ecotone ART isotherm of 23 °C; beech-dominated forests lay north of between cool-temperate deciduous broadleaf forest (a WI this isotherm and hemlock-dominated forests lay south. This range of 45–85 °CÆmonth) and warm-temperate evergreen implies the importance of high winter temperatures in broadleaf forest (WI > 85 °CÆmonth) (Kira, 1991). However, determining the distribution of Chinese beech because the the WI value at the upper limit of F. crenata in Japan ART is closely correlated with winter temperature in southern (45.1 °CÆmonth) coincided well with Kira’s criterion of 45 °C mountain areas in China. month; this supports the idea that F. crenata is an indicator of Two effects of high winter temperatures that can influence the Japanese temperate zone (e.g. Miyawaki, 1980–89). the distribution of temperate tree species may be playing a role American beech occupies a large climatic space, with a range in beech distribution in China. First, temperate trees require a of 50.7–173.4 °CÆmonth in WI and 7.0–19.5 °C in ABT sufficient period of winter cold (a period of chilling) before (Table S1). This spans two bioclimatic zones: the cool- warming will induce budburst in spring (Cannell & Smith, temperate zone [45–85 °CÆmonth for WI (Kira, 1945, 1991) 1986; Lechowicz, 2001). Second, higher winter temperatures and 6–12 °C for ABT (Holdridge, 1947)], and the warm- can reduce the competitive ability of deciduous trees with temperate zone (85–180 °C month for WI and > 12 °C for more warmth tolerant evergreen broadleaf trees (Woodward, ABT). 1987). These effects of winter temperature may explain an In Europe, climatic parameters at the northern/upper limit unresolved question in Asian biogeography: why beech species of beech distribution indicated good agreement with criteria do not spread westward into the Himalayas and south-eastern defining the cool-temperate zone: a WI value of 47.7– Tibet where growing season warmth and precipitation appear 104.3 °CÆmonth, and an ABT value of 7.2–13.5 °C for satisfactory. Although Asian beech and hemlock have similar F. sylvatica, and 46.3–78.3 °CÆmonth and 7.1–10.4 °C for heat requirements during the growing season (Liu & Qiu, F. orientalis (Table S1). On the other hand, an average Im 1980; Hou, 1983; Fang et al., 1996), hemlock is favoured by a greater than 15.4 for all beech sites suggest a perhumid or warm-winter climate (Sakai, 1975). While Chinese beeches humid climate, defined by Thornthwaite (1948) as Im values of co-exist with evergreen broad-leaved tree species in genera 20–100 and > 100, respectively. 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 Climates and present distribution of East Asian tree species. Observations from Woodward (1987), Cao beeches (1995), Williams-Linera et al. (2000) and Miyazawa & As shown in Table 3, the first two PCA components account Kikuzawa (2005) support this hypothesis that warm winters for more than 70% of the variance in parameters associated can favour evergreens and limit range expansion in beech with the distribution of Asian beech species. No one variable species. alone can explain beech distributional patterns; growing season In Japan, growing season warmth rather than winter warmth (WI and ABT), winter low temperature (CI) and temperatures is generally seen as the control on the distribu- annual mean temperature (AMT) all showed almost equal tions of beech species (Kira, 1945; Miyawaki, 1980–89; and loadings (Table S2). Thermal regime is clearly paramount in many others), and Japanese cool-temperate vegetation is the relationships between beech distribution and climatic sometimes termed the beech forest zone (e.g. Miyawaki, factors in East Asia, but the strong correlation among different 1980–89). Precipitation is also used to explain differences in climatic elements (see Table S3 in Supplementary Material) community composition and structure of Japanese beech precludes identification of a single, dominant aspect of thermal forests (Kure & Yoda, 1984; Hattori & Nakanishi, 1985; Fang & regime that affects the distribution of East Asian beech species. Yoda, 1990; Matsui et al., 2004). In particular, distribution of There are a few viewpoints on the relationships between the F. crenata along the Pacific Ocean side and the Japan Sea side present distribution of Chinese beech species and limiting of Japan usually has been explained by accumulated snowfall factors. Hong & An (1993) pointed out that the climatic (Yamazaki, 1983; Maeda, 1991; Matsui et al., 2004), with pure factors affecting beech distribution varied from place by place; beech forests found only on the Japan Sea side where snowfall for example, in northern regions, coldness and short growing is extremely abundant. Given the situation in China, however, season were major limiting factors, whereas water deficit was we should not discount out of hand the possible influence of
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low winter temperatures in the development of pure beech Table 4 Thermal variables for Newfoundland, Canada, based on forests on the Japan Sea side. Winter temperature there is 18 climatic stations (Atmospheric Environmental Service, Envi- much lower than on the Pacific Ocean side; this could ronment Canada, 1982). For comparison, the estimated mean strengthen the competitive ability of beech against other tree value of climate variables at the northern limit of Fagus grandifolia species (Tsuga and Quercus) and understorey bamboo (Sasa), in North America (‘Mean at northern limit’ column) is also tabulated. See Table 2 for abbreviations for climatic variables and enable its dominance. A question remains with regard to Japanese beech distribu- Mean at tion: why is there no beech on Yakushima Island (30°27¢ N, Variable Mean SD Minimum Maximum northern limit 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 AMT (°C) 5.2 0.5 4.1 5.8 4.2 co-occur with beech in Kyushu, Shikoku and Honshu WI (°C month) 36.7 3.0 30.2 41.6 50.7 CI (°C month) )34.8 4.2 )42.2 )26.3 )60.5 (Miyawaki, 1980–89). This may be related to some topo- ABT (°C) 6.1 0.3 5.3 6.4 7.0 graphic barriers or climatic limitations at the time during MTWM (°C) 15.6 0.5 14.7 16.4 18.4 glaciation when Yakushima was part of the major Japanese MTCM (°C) )4.2 1.1 )6.1 )1.9 )11.4 islands, or simply to dispersal limitation since sea levels rose ART (°C) 19.8 1.1 17.2 22.5 29.8 and isolated the island. Another possibility worth investigating K 25.3 2.6 19.5 31.4 49.5 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 and )4.2 °C for MTCM, and the latter )60.5 °CÆmonth and budburst and the early part of the growing season that puts )11.4 °C). In contrast, the growing season temperatures were beech at a competitive disadvantage. 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, Climates and the present distribution of American 7.0 °C and 18.4 °CÆmonth, respectively (Table 4). This sug- beech gests that insufficient warmth during the growing season may The PCA showed that MTCM and PET have almost equivalent be a factor limiting the expansion of beech into Newfound- loadings to growing-season warmth (WI and ABT) for the land. This hypothesis is supported by eco-physiological studies southern limit of F. grandifolia, whereas heat (WI and ABT) of flowering and seed production showing that a certain and energy (PET) are most important for the northern limit minimum degree of heat is required for floral initiation, and (Table S2). Although Huntley et al. (1989) demonstrated the flower and seed production in many temperate tree species role of January and July mean temperatures in the present (Matyes, 1969; Owens & Blake, 1985). The comparative study distribution and abundance of beech species in North America of masting behaviours of beech species also shows the and Europe using pollen percentages of surface samples, they importance of summer heat in controlling beech seed used only monthly mean temperatures as thermal parameters. production (Piovesan & Adams, 2001). Our findings based on survey of many more climatic Although temperature can account for the distribution parameters are generally consistent with their conclusions. limits of American beech, Fig. 1 suggests that continentality Although our study suggested that the growing season (K) also is important for limiting the southern and northern warmth was associated with the northerly distribution limit, distribution limits. The fact that WI and MTCM at the some physiological observations emphasize the adverse effects southern limit decrease markedly with increasing K value of excessively low winter temperatures for many temperate shows that beech requires more heat and higher winter North American trees (Sakai & Weiser, 1973; Hicks & Chabot, temperature in an oceanic climate than in a continental one. 1985; Denton & Barnes, 1987; Maycock, 1994). Many studies However, growing season warmth tends to increase as K stress the influence of low winter temperature, but perhaps increases at the northern limit (Fig. 1a), suggesting higher only because it is easier to do experimental manipulations of summer temperatures in the continental than in the oceanic chilling effects than warmth during the growing season. We climate. The relationship between winter temperature and K can, however, consider biogeographic evidence supporting the values at the northern limit shows the same pattern as at the importance of growing season temperature. The lack of southern limit (Fig. 1b). Similar results were found for the American beech in Newfoundland, Canada, where winter distributions of some tree species and vegetation zones in East temperatures are much higher than at the northern edge of Asia (Ohsawa, 1990; Fang & Yoda, 1991; Fang et al., 1996). beech distribution (Table 4) suggests growing season warmth is a greater limitation than winter cold. Climatic statistics of Climates and present distribution of beech species in thermal variables at 18 stations located between 47° and 48° N Europe in Newfoundland where the latitudes were coincident with the northern limit in east Quebec show far higher winter The relationships between beech distribution and environ- temperatures (CI and MTCM) in Newfoundland than at the ments in Europe have been discussed from the viewpoint of beech northern limit (in the former, )34.8 °CÆmonth for CI soil, topography and climate (Ellenberg, 1986; Jahn, 1991).
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Figure 2 Relationships between (a) warmth index (WI) and (b) Figure 1 Relationships between (a) warmth index (WI) and (b) mean temperature for the coldest month (MTCM) and conti- mean temperature for the coldest month (MTCM) and continen- nentality index (K) at the northern limit of Fagus sylvatica in tality index (K) at the southern (filled circles) and northern (open Europe. circles) limits of Fagus grandifolia in North America. The rela- tionships for the northern limits are fit by a nonlinear regression. Table 5 Climatic parameters at the northern limit for Fagus sylvatica in England based on 11 climatic stations. For comparison, These earlier studies supported the importance of growing the estimated mean value of climate variables at the northern limit season temperatures, but did not focus in any detail on the of F. sylvatica on the European mainland (‘Mean at northern limit’ array of thermal parameters that might be involved. Both column) is also tabulated. See Table 2 for abbreviations for thermal climate and continentality (K) contributed to Euro- climatic variables pean beech distributions, much as in North America. Figure 2 Mean at expresses the relationships between the thermal variables (WI Variables Mean SD Minimum Maximum northern limit and MTCM) and K value at the northern limit of F. sylvatica. With an increase of the K values, WI increased (Fig. 2a), and AMT (°C) 10.0 0.4 9.1 10.7 6.6 MTCM decreased (Fig. 2b). WI (°C month) 62.6 3.4 53.8 69.2 47.7 Although beech has been present in south-eastern England CI (°C month) )2.3 1.2 )4.2 0.0 )28.3 since at least 3000 yr bp (Birks, 1989), its present distribution ABT (°C) 10.0 0.4 9.1 10.7 7.2 in the British Isles does not appear to be in equilibrium with MTWM (°C) 16.8 0.4 15.8 17.6 16.9 MTCM ( C) 3.9 0.6 3.1 5.5 2.7 present climate. Climatic analysis shows that beech still has not ° ) ART (°C) 12.9 0.7 10.6 13.9 19.6 reached its potential northern range limit (Table 5). Compared K 7.7 1.6 2.8 9.8 25.8 with averages of climatic variables at the upper/northern AP (mm) 707.2 148.4 540.0 1068.0 1272.3 limits on the European mainland (AMT of 6.6 °C, WI of PET (mm) 646.8 8.6 625.0 661.0 577.6 47.7 °CÆmonth, and ABT of 7.2 °C), those at the present AAE (mm) 560.6 44.3 497.0 652.0 496.7 northern limit in Britain were much larger (more southerly), Im 14.7 20.4 )8.8 64.2 119.3 by c. 3.4 °C for AMT, 15 °CÆmonth for WI and 2.8 °C for ABT EQ (°C mm)1) 24.7 4.8 15.4 32.4 16.8 (Table 5). Assuming a decrease in mean temperatures of 0.5 °C per degree latitude in higher latitudes (Fang, 1996), Comparison of climatic space for world beech species beech should arrive at its range limit c. 6° to the north of its present range boundary. Spreading north at a rate of 100– The present distributions of most tree species are strongly related 200 m yr)1 (Birks, 1989), beech should reach its potential to climate (Woodward, 1987; Huntley et al., 1989; Francis & natural distribution in Britain 3500–7000 years in the future if Currie, 2003). Simple isotherm methods have long been used to one assumes an unchanging climate scenario. This implies that assess distributional limits in relation to climate (Hutchinson, most of Britain eventually should be covered by beech. 1918; Koppen, 1936; Kira, 1945, 1991; Thornthwaite, 1948;
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Bryson, 1966; Tuhkanen, 1980; Grace, 1987; Arris & Eagleson, beech. At its lower or southern limits (Fig. 3), F. grandifolia 1989; and many others), but the nuances of climatic controls on extends to warmer regions, while F. crenata requires less range limits are not assessed fully by isotherms. In this study, we warmth, but both Japanese beeches occur in more moist used a more comprehensive PCA approach to detect the single climates than F. grandifolia. Chinese and European beeches and joint importance of diverse climatic parameters in explain- occupy similar temperature ranges, but the latter is in a ing the distribution of beech species. We are thus able to drier climate. At the upper or northern edges (Fig. 4), both compare the climatic space of beech distribution among species F. grandifolia and F. crenata occupy colder areas, while in the three separate geographic regions where beech are major Chinese beeches have similar warmth demand to F. japonica. components of forest ecosystems. Along the moisture axis, F. crenata occurs in the most Because the first two PCA axes were most important in humid conditions, and European beech in the driest explaining the distribution of beech species around the world, habitats. Although the actual influence of climatic factors sample scores of these axes were used to compile scatter on species distributions may be nonlinear and only partly diagrams comparing the climatic spaces (climatic niches) of reflected in the present analyses (Austin, 2002), it is clear the respective beech species (Figs 3 and 4). Because of a large that there is a degree of climatic niche differentiation among sample size and rather scattered score values, 50% Gaussian the extant beech species. bivariate confidence ellipses (ELL) were drawn for the species with a large sample size to more easily compare climatic CONCLUSIONS conditions among beech species. Also, because four Chinese beeches and two European beeches have similar moisture and Focusing on distribution limits of the world beech species, we warmth requirements (see Table 2; Table S1), sample data compare their climatic spaces in three different regions were combined. globally, and explore the climatic correlates of these distribu- Figures 3 and 4 show climatic gradients associated with tion patterns. The results suggest that thermal climate is most the lower (southern) limit and the upper (northern) limit of 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.
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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 evergreen trees. Although the distribution limits of beech and duration of low winter temperature (MTCM and CI), and species in Japan were controlled by summer temperature, their annually available solar energy (PET) sometimes also played a dominance may depend on regional climatic factors such as role. At the lower or southern limits, F. grandifolia occurred in snowfall and winter low temperature. Winter low temperature much warmer regions, and F. crenata in colder regions; may enhance the competitive ability of F. crenata with other Chinese and European beeches have similar, intermediate heat co-existing species, allowing it to form pure beech forests in requirements. Along moisture gradients, Japanese beeches western Japan. appeared in more moist conditions and Chinese and European High summer temperature was considered to be the limiting beeches in drier situations. At the upper or northern limits, factor for southward extension of American beech, while F. crenata and American beeches had similar, relatively low adequate growing season warmth was critical for its northward warmth demands, while F. japonica and Chinese beeches were distribution. Continentality (K) played an important part in found only in warmer regions. Along a moisture gradient, the delimiting its range expansion, but lack of growing season Japanese beech species again occupied the most moist regions, warmth was the most important climatic factor precluding its with European beech in contrast occupying the most dry migration to the Atlantic Islands (such as Newfoundland, (Fig. 4). Canada). Summer temperature is a limiting factor for the Growing season temperature was most important in distribution of beeches in Europe, but continentality was also explaining overall distribution of Chinese beeches, but their associated with limits to their north-western distribution. The northern limits were mainly set by low precipitation. The northerly distribution of beech in Britain has apparently not climatic factor controlling their westward expansion (south- reached its potential limit due to lack of time since deglaci- east Tibet and Himalaya) may be higher winter temperatures ation. that influence their budburst in spring and weaken their Although the present-day distribution patterns of beech competitive ability with evergreen hemlock and broad-leaved species showed good correspondence to contemporary climate,
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isolated exceptions exist. For example, despite favourable Camp, W.H. (1951) A biogeographical and paragenetic ana- climatic conditions there are no records of beech ever having lysis of the American beech (Fagus). American Philosophy grown on Yakushima Island in Japan. Similarly, F. mexicana, Society Yearbook, 1950, 166–199. isolated in a small mountainous area of north-eastern Mexico, Cannell, M.G.R. & Smith, R.I. (1986) Climatic warming, shows no clear association with contemporary climate. There- spring budburst and frost damage of trees. Journal of Applied fore, a historical view on beech distribution is an essential Ecology, 23, 177–191. complement to the climatic analyses emphasized in this paper. Cao, K.F. (1995) Fagus dominance in Chinese montane forests: natural regeneration of Fagus lucida and Fagus hayatae var. pashanica. PhD Thesis, Wageningen Agricultural University, ACKNOWLEDGEMENTS Wageningen. Assistance from many colleagues enabled this study. JYF is Cao, K.F., Peters, R. & Oldeman, R.A.A. (1995) Climatic greatly indebted to Y.H. Tang for assistance in collecting beech ranges and distribution of Chinese Fagus species. Journal of data in Japan and Korea, C.F. Hsieh for F. hayatae in Taiwan, Vegetation Science, 6, 317–324. and F. Reygadas for Mexican beech. Thanks are extended to Central Meteorological Office of Korea (1972) Climatic table of Z.H. Wang and X.P. Wang for their assistance in data analysis, Korea. Central Meteorological Office of Korea, Seoul. and to S.P. Wang for her help in compiling climatic data Chang, D.H.S. (1983) The Tibetan Plateau in relation to the sets and checking place locations. We also thank Robert vegetation of China. Annals of Missouri Botanical Garden, Whittaker and two anonymous referees for their helpful 70, 564–570. comments and suggestions on the earlier version of this paper. 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Climatic limits for world beech distribution
Wolfe, J.A. (1979) Temperature parameters of humid to mesic nents associated with the distribution limits of world beech forests of Eastern Asia and relation to forest of other regions of species. the northern Hemisphere and Australasia. Geological Survey Table S3 Coefficient of correlation between annual mean Professional Paper 1106, Washington, DC. temperature (AMT) and other thermal variables across the Woodward, F.I. (1987) Climate and plant distribution. Cam- global range of beech species. bridge University Press, Cambridge. Wu, Z.Y. (ed.) (1980) Vegetation of China. Scientific Press, Beijing. Yamazaki, K. (1983) Plant distribution in Japan. Modern biology series, 7a, Higher plants (ed. by M. Honda and K. B I O S K E T CH E S Yamazaki), pp. 119–155. Nakayama-syoten, Tokyo. Jingyun Fang is a professor and chair of the Department of SUPPLEMENTARY MATERIAL Ecology, Peking University. His research interests cover biogeography of plants, terrestrial ecosystem productivity The following supplementary material is available online from and remote sensing of vegetation. http://www.Blackwell-Synergy.com Martin J. Lechowicz is a professor in the Department of Appendix S1 Location and elevation of distribution limits of Biology, McGill University, and Director of the University’s beech (Fagus L.) species in the present study. Gault Nature Reserve. His research interests centre on the Table S1 Statistics for climatic variables at lower (southern) comparative ecology of trees and on the ecology and and upper (northern) limits of distribution for world beech conservation of forest communities. species. Table S2 Loadings of climatic variables derived from Princi- pal Components Analysis for the first three principal compo- Editor: Robert J. Whittaker
Journal of Biogeography 33, 1804–1819 1819 ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd 中国科技论文在线 http://www.paper.edu.cn
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. No. Location Latitude Longitude Lower limit (m) Upper limit (o.’) (o.’) (m) 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 2500 ! 11 Gutianshan, Kaihua, Zhejiang 29.06 118.24 800 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
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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
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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 F. lucida 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
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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 27.31 114.10 800 1400 South Yandangshan, Taishun, Zhejiang 27.32 120.06 1000 1200 23 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 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
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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 3 Dabashan, Sichuan 32.38 107.30 ! 1850 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 !
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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 33 jap5 35.30 138.49 ! 1700 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 37.46 140.42 150 700 37.20 140.43 200 750 5 Central Abukuma Santi, Hukushima 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 !
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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 33.25 132.35 900 1000 Kunimidake, Shiibamura, Miyazaki 32.30 131.00 1000 1300 30 F. multinervis 1 Ulreung-do, South Korea 37.29 130.54 300 960 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
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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 Turkmen Da., Turkey 39.50 30.10 1400 1600 4 Duldul Da., Gokcayir to Atlik Y., Adana, Turkey 37.04 36.15 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
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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
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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
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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
F. engleriana AMT (oC) 12.4 2.51 8.3 17.7 7.3 1.88 3.7 12.8 WI (oC·month) 99.0 22.40 57.6 152.4 56.0 14.19 32.2 99.9 CI (oC·month) -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) 23.0 1.72 17.1 26.0 17.8 2.23 13.6 23.1 MTCM (oC) 1.1 3.55 -4.1 8.2 -3.9 2.36 -8.9 1.7 ART (oC) 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 Im 61.4 82.32 0.0 356.4 70.0 76.98 36.3 356.4 EQ (oC/mm) 21.7 5.80 6.1 32.0 19.4 3.98 6.1 22.5 F. hayatae AMT (oC) 12.8 2.06 9.1 15.4 9.2 2.07 6.3 11.9 WI (oC·month) 99.4 22.36 67.6 132.1 67.0 13.26 52.4 89.2 CI (oC·month) -6.4 6.08 -17.9 0.0 -16.6 13.73 -36.4 0.0 ABT (oC) 12.8 2.00 9.3 15.4 9.5 1.66 7.5 11.9 MTWM (oC) 22.4 4.31 16.9 29.2 18.6 2.14 16.3 21.9 MTCM (oC) 2.1 3.26 -2.3 7.3 -0.7 4.15 -5.9 5.4 ART (oC) 20.3 6.10 11.5 28.3 19.3 4.85 11.5 23.5 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 Im 85.5 54.20 24.4 182.1 109.8 34.81 84.4 182.1 EQ (oC/mm) 17.0 5.20 10.8 24.1 13.2 1.74 10.8 16.5 F. longipetiolata AMT (oC) 14.3 2.15 9.6 21.6 8.9 1.76 4.9 13.3 WI (oC·month) 115.9 22.64 75.5 198.9 66.2 13.75 36.5 99.3 CI (oC·month) -4.1 4.20 -19.9 0.0 -19.8 8.42 -39.8 0.0 ABT (oC) 14.3 2.13 9.9 21.6 9.2 1.48 6.1 13.3 MTWM (oC) 24.2 1.69 20.7 27.4 18.9 2.08 14.5 22.4 MTCM (oC) 3.5 3.16 -2.6 15.1 -2.0 2.19 -6.3 6.0 ART (oC) 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
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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 Im 77.5 77.07 -4.4 356.4 132.6 63.35 97.0 356.4 EQ (oC/mm) 19.0 5.08 6.1 29.4 14.7 2.60 6.1 17.2 F. lucida AMT (oC) 12.7 2.07 8.5 17.6 8.8 1.81 3.6 12.5 WI (oC·month) 100.6 18.45 68.8 150.8 66.4 12.49 35.1 95.3 CI (oC·month) -8.2 7.26 -27.3 0.0 -21.3 10.20 -52.9 -5.3 ABT (oC) 12.8 1.97 9.2 17.6 9.2 1.40 5.7 12.5 MTWM (oC) 23.0 1.40 20.5 25.5 19.1 1.57 14.9 21.7 MTCM (oC) 1.5 2.89 -4.2 8.4 -2.4 2.39 -8.9 1.9 ART (oC) 21.5 2.40 15.3 26.1 21.5 2.12 17.7 25.8 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 Im 70.0 67.93 -3.8 356.5 168.5 51.14 92.2 356.5 EQ (oC/mm) 19.5 4.82 6.1 32.0 13.8 2.32 6.1 17.0 F. crenata AMT (oC) 9.5 1.26 5.7 12.0 5.0 1.25 2.3 7.6 WI (oC·month) 74.8 10.21 52.0 95.2 45.1 5.73 31.9 56.1 CI (oC·month) -21.3 6.68 -43.7 -6.9 -45.2 10.21 -66.3 -25.3 ABT (oC) 9.8 1.02 7.3 12.0 6.7 0.68 5.3 8.1 MTWM (oC) 21.8 1.62 19.1 24.8 17.6 0.69 15.6 18.8 MTCM (oC) -2.1 1.57 -6.8 1.8 -6.8 1.81 -10.4 -3.3 ART (oC) 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 EQ (oC/mm) 12.5 3.10 7.3 20.6 9.5 1.20 7.3 12.8 F. jopanica AMT (oC) 11.7 1.03 10.3 14.8 9.0 0.99 7.3 11.8 WI (oC·month) 91.8 8.71 81.1 117.5 70.3 7.09 55.3 87.2 CI (oC·month) -12.0 4.09 -19.1 -0.3 -21.9 5.18 -29.1 -6.0 ABT (oC) 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 (oC) 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
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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 EQ (oC/mm) 15.3 4.21 6.5 22.7 11.5 1.99 6.5 14.7 F. multinervis AMT (oC) 11.5 - - - 7.6 - - - WI (oC·month) 90.2 - - - 59.0 - - - CI (oC·month) -11.9 - - - -27.5 - - - ABT (oC) 11.5 - - - 8.3 - - - MTWM (oC) 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 ------EQ (oC/mm) 16.1 ------F. grandifolia AMT (oC) 19.5 0.76 17.2 21.0 4.2 0.88 1.6 6.0 WI (oC·month) 173.4 9.14 146.5 192.0 50.7 4.74 36.1 61.2 CI (oC·month) 0.0 0.00 0.0 0.0 -60.5 8.17 -76.9 -38.1 ABT (oC) 19.5 0.76 17.2 21.0 7.0 0.48 5.5 8.1 MTWM (oC) 27.5 0.41 26.7 28.4 18.4 0.84 16.0 20.1 MTCM (oC) 10.4 1.20 7.2 13.6 -11.4 1.95 -13.7 -6.6 ART (oC) 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 Im 40.3 14.64 20.6 65.6 91.1 34.34 43.3 168.2 EQ (oC/mm) 19.5 2.27 16.3 23.6 18.5 3.29 12.5 24.9 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 ABT (oC) 15.6 1.51 14.3 17.4 14.8 1.8 13.6 17.4 MTWM (oC) 19.6 1.59 18.1 21.1 18.7 1.6 17.4 20.8 MTCM (oC) 10.5 2.59 7.9 14.1 9.7 3.1 7.2 14.1 ART (oC) 9.1 2.10 6.7 11.2 9.1 2.1 6.7 11.2 K 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) 949.0 264.80 715.0 1263.0 - - - - 906.0 203.1 715.0 1088.0 AAE (mm) - - - - Im 103.1 89.80 -6.7 200.4 - - - -
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EQ (oC/mm) 14.5 7.50 8.6 25.3 - - - - F. sylvatica AMT (oC) 13.5 2.23 9.1 17.0 6.6 1.40 4.5 8.9 WI (oC·month) 104.3 23.39 59.3 144.2 47.7 9.75 28.0 65.5 CI (oC·month) -2.7 4.35 -13.9 0.0 -28.3 9.36 -42.7 -12.4 ABT (oC) 13.5 2.22 9.1 17.0 7.2 1.07 5.2 8.9 MTWM (oC) 23.0 2.58 17.4 28.8 16.9 1.75 12.5 20.2 MTCM (oC) 4.7 2.75 -1.0 9.8 -2.7 2.13 -6.3 1.6 ART (oC) 18.2 2.60 14.2 23.0 19.6 2.72 12.8 22.9 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 Im 38.1 65.17 -8.2 211.4 119.3 82.62 -2.7 240.3 EQ (oC/mm) 29.0 9.54 9.3 39.4 16.8 9.55 5.4 34.9 F. orientalis AMT (oC) 10.2 2.74 7.5 14.3 6.5 1.98 4.9 9.9 WI (oC·month) 78.3 26.52 56.3 126.1 46.3 14.50 29.9 69.1 CI (oC·month) -16.2 10.20 -27.7 -0.8 -28.5 10.45 -37.6 -10.7 ABT (oC) 10.4 2.57 8.0 14.4 7.1 1.68 5.4 9.9 MTWM (oC) 20.5 3.73 17.6 27.8 16.1 1.95 13.3 18.4 MTCM (oC) -1.5 3.64 -6.0 4.2 -3.2 2.26 -5.0 0.7 ART (oC) 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 Im 15.4 31.20 -5.1 76.7 23.8 29.93 7.3 76.7 EQ (oC/mm) 32.7 9.77 17.2 44.7 27.9 6.53 17.2 32.4
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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 PCA 2 PCA 3 PCA 1 PCA 2 PCA 3 All beech species AMT (oC) 0.97 0.15 0.00 0.98 0.02 -0.12 WI (oC·month) 0.95 0.14 0.03 0.89 0.24 0.13 CI (oC·month) 0.85 0.16 -0.10 0.91 -0.17 -0.31 ABT (oC) 0.97 0.15 0.00 0.93 0.15 0.02 MTWM (oC) 0.73 0.06 0.16 0.59 0.34 0.51 MTCM (oC) 0.96 0.23 -0.14 0.88 -0.25 -0.38 ART (oC) -0.73 -0.26 0.29 -0.61 0.41 0.62 K -0.42 -0.22 0.68 0.22 0.16 0.80 AP (mm) -0.34 0.83 0.35 0.23 -0.83 0.38 PET (mm) 0.54 -0.22 0.68 0.63 0.31 0.22 AAE (mm) 0.42 -0.05 0.84 0.62 0.23 0.37 Im -0.50 0.84 -0.08 0.03 -0.92 0.29 EQ (oC/mm) 0.28 -0.89 -0.25 -0.13 0.89 -0.35 Chinese beeches AMT (oC) 0.96 0.20 0.07 0.99 0.12 0.03 WI (oC·month) 0.93 0.24 0.03 0.92 0.30 0.20 CI (oC·month) 0.90 0.03 0.20 0.95 -0.13 -0.20 ABT (oC) 0.96 0.21 0.06 0.97 0.21 0.08 MTWM (oC) 0.58 0.22 0.01 0.67 0.50 0.52 MTCM (oC) 0.97 0.12 0.09 0.93 -0.17 -0.30 ART (oC) -0.73 0.01 -0.09 -0.39 0.56 0.70 K -0.61 0.07 0.28 -0.09 0.53 0.77 AP (mm) 0.00 -0.69 0.71 0.33 -0.63 0.25 PET (mm) -0.32 0.62 0.70 0.09 0.83 -0.24 AAE (mm) -0.33 0.62 0.69 0.08 0.83 -0.25 Im 0.21 -0.95 0.10 0.14 -0.82 0.42 EQ (oC/mm) -0.27 0.85 -0.41 -0.23 0.86 -0.35 F. crenata AMT (oC) 0.91 0.42 -0.03 0.95 0.31 0.01 WI (oC·month) 0.71 0.68 0.07 0.80 0.44 0.34 CI (oC·month) 0.96 -0.10 -0.17 0.95 -0.21 -0.20 ABT (oC) 0.82 0.55 0.00 0.88 0.39 0.21 MTWM (oC) 0.38 0.89 0.09 0.43 0.58 0.62 MTCM (oC) 0.95 -0.09 -0.21 0.92 0.23 -0.27 ART (oC) -0.45 0.82 0.25 -0.83 -0.02 0.54 K 0.05 0.45 0.50 -0.06 -0.42 0.86 AP (mm) 0.38 -0.70 0.59 0.69 -0.62 0.09 PET (mm) 0.43 -0.70 -0.39 -0.36 0.73 0.08 AAE (mm) 0.43 -0.70 -0.39 -0.36 0.73 0.08 Im 0.25 -0.40 0.87 0.63 -0.51 0.21 EQ (oC/mm) -0.38 0.47 -0.77 -0.69 0.60 -0.10 F. japonica AMT (oC) 0.92 0.38 -0.01 0.93 0.31 -0.18 WI (oC·month) 0.86 0.46 -0.06 0.84 0.47 -0.24 CI (oC·month) 0.96 0.16 0.08 0.98 0.07 0.07 ABT (oC) 0.92 0.38 -0.02 0.92 0.33 -0.19 MTWM (oC) 0.46 0.84 -0.09 0.40 0.80 -0.38
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MTCM (oC) 0.97 0.09 0.05 0.97 -0.02 -0.06 ART (oC) -0.61 0.65 -0.14 -0.71 0.57 -0.20 K -0.52 0.08 0.11 -0.40 0.53 0.23 AP (mm) 0.64 -0.68 -0.19 0.71 -0.56 0.32 PET (mm) 0.30 -0.31 0.88 0.33 0.67 0.66 AAE (mm) 0.30 -0.31 0.88 0.33 0.67 0.66 Im 0.55 -0.63 -0.49 0.57 -0.75 -0.02 EQ (oC/mm) -0.43 0.73 0.39 -0.46 0.77 -0.25 F. grandifolia AMT (oC) 0.97 -0.19 0.11 0.75 -0.66 0.08 WI (oC·month) 0.97 -0.19 0.11 0.98 -0.02 0.19 CI (oC·month) -0.08 -0.36 0.24 0.39 -0.91 0.01 ABT (oC) 0.97 -0.19 0.11 0.97 -0.14 0.18 MTWM (oC) 0.54 -0.18 0.79 0.90 0.09 0.33 MTCM (oC) 0.97 -0.14 -0.18 0.15 -0.98 -0.11 ART (oC) -0.86 0.08 0.49 0.26 0.93 0.25 K -0.68 0.19 0.68 0.38 0.88 0.27 AP (mm) 0.57 0.81 0.12 -0.67 -0.35 0.65 PET (mm) 0.91 -0.30 0.27 0.98 -0.05 0.17 AAE (mm) 0.94 0.09 0.00 0.82 -0.16 0.26 Im 0.23 0.97 0.06 -0.78 -0.29 0.55 EQ (oC/mm) -0.57 -0.81 -0.04 0.86 0.25 -0.43 European beeches AMT (oC) 0.95 0.24 0.17 0.63 0.76 -0.11 WI (oC·month) 0.89 0.38 0.20 0.89 0.40 0.06 CI (oC·month) 0.92 -0.26 -0.04 0.04 0.95 -0.29 ABT (oC) 0.94 0.27 0.17 0.79 0.59 -0.02 MTWM (oC) 0.72 0.56 0.38 0.90 0.20 0.16 MTCM (oC) 0.94 -0.23 -0.12 -0.20 0.88 -0.39 ART (oC) -0.44 0.73 0.45 0.73 -0.52 0.39 K -0.43 0.72 0.46 0.56 -0.08 0.82 AP (mm) 0.04 -0.91 0.35 -0.63 0.55 0.46 PET (mm) 0.58 0.02 -0.48 0.37 0.46 0.32 AAE (mm) 0.31 -0.49 -0.07 0.34 -0.32 -0.69 Im -0.13 -0.86 0.44 -0.73 0.44 0.34 EQ (oC/mm) -0.09 0.87 -0.43 0.76 -0.42 -0.34
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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 Mean temperature for the coldest month (oC) 0.98 0.93 Minimum mean temperature (oC) 0.95 0.90 Annual range of temperature (oC) -0.79 -0.69
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