Appendix B. Additional details on the protocol followed and the original sources used to build the tolerance database and to standardize the rankings of tolerance obtained from different sources and for species from different continents.
Shade tolerance rankings for North America
Shade-tolerance rankings, each containing a different set of species, were employed to develop a common shade tolerance ranking for North-American species (Table 1). We used the five-level (very intolerant, 1; intolerant, 2; moderately tolerant, 3; tolerant, 4; very tolerant, 5) shade tolerance scale of Baker (1949) as the starting point. This shade tolerance ranking is based on actual measurements of minimum light availability at species location in the field (Wiesner 1907, Zon and Graves 1911) , that were revised and expanded to include more species based on questionnaires sent out to foresters. Very intolerant species require full light for growth and grow best in the complete absence of competition, while very tolerant species can persist in forest understories for many years at light availabilities as low as 0.5–2% of incident light (Küppers 1989, Walters and Reich 1999) . Because it is based on actual light measurements and includes a large number of important species, the scale of Baker (1949) is commonly used in classifying tree light requirements in comparative studies of life history traits in North-American tree species (Kobe et al. 1995, Coomes and Grubb 2000) .
However, Baker (1949) provides a lumped estimate of shade tolerance for several diverse genera such as Aesculus, Alnus, Carya, Fraxinus, and many species from important forest genera such as Acer, Pinus, Populus, Quercus, Ulmus are not scored. In addition, this scale includes few understory species and broadleaf evergreen temperate species. In addition to these limitations, every score includes a certain degree of subjective error. Averaging the opinions of foresters from different North-American forest types that may not always encompass the extremes of shade tolerance can have led to systematic over- or underscoring of shade-tolerance for the less frequent species in the scale of Baker (1949) . To increase the species coverage and reduce the subjective error of species shade tolerance scoring, we included 9 additional published shade tolerance scales. In addition, four up-to-date online databases were employed to derive the shade-tolerance estimates of less frequent species (Table 1).
Because the number of shade-tolerance levels as well as the criteria of scoring differed among the data sets, we converted all sets of data to the 5-scale shade-tolerance scale of Baker (1949) . For every data set, this was done using a regression analysis between the estimates of shade tolerance for species present in both data sets, and then using this regression to calculate a Baker-equivalent shade tolerance rank for all species in a specific data set. Simple linear regression analysis was employed for most data sets. For the studies reporting actual minimum light intensities, either an exponential function (Baker vs. Wiesner, Table 1) or a power function (Baker vs. Graham; Table 1) was used. A final value of species shade tolerance was obtained as an average of all available data sets for a species. Because the shade tolerance scale of Baker (1949) is derived from that of Zon and Graves (1911) , these two data sets were averaged before calculating the overall mean.
Scoring shade tolerance in European species
For European species, the species ranking of Ellenberg (1991) is commonly employed (Niinemets and Kull 1994, Coomes and Grubb 2000, Valladares et al. 2002, Cornwell and Grubb 2003) . The Ellenberg’s ecological indicator values for light characterize species natural dispersal along the habitats of varying light availability, and vary for woody species from 3 to 9, giving a seven-level scale (Ellenberg 1991) . These values are derived from actual measurements of light availability in species habitat and correspond to the approximate light requirements (% of full light) of : 3: 2–5%, 4; 5–10%, 5; 10–20%, 6; 20–30%, 7; 30–40%, 8; 40–50%, 9; >50% (Ellenberg 1991, Niinemets and Kull 1994) . Recently, the ecological indicator values of Ellenberg (1991) were revised for the British Isles (Hill et al. 1999, Hill et al. 2000) , and we calculated an average value of light requirement for these two estimations. The ecological indicator values are determined for seedling and sapling stages of plant development, and may potentially change during the course of plant development (Yevstigneyev 1990, Ellenberg 1991) .
To improve the shade tolerance estimates of important forest trees and increase the scope of the data set, eleven additional shade tolerance scorings (Table 1) based on direct light measurements in forest understory (Wiesner 1907, Yevstigneyev 1990) , foliage physiological characteristics (Ivanov and Kossovich 1932) , and foresters’ and ecologists’ knowledge of species biology (Gayer 1898, Morozov 1903, Warming 1909, Walter 1968, Jahn 1991, Brzeziecki and Kienast 1994, Otto 1994, Ellenberg 1996) were used. As for the North-American data set, regression analysis was employed to convert the various species rankings to the Ellenberg scale using species that were common to Ellenberg's data set and the specific data set. Linear regressions were used in all cases, except for Wiesner (1907) vs. Ellenberg, which was fitted by an exponential function. From these data, a common initial mean value of species light requirement was calculated.For some rankings, the values for certain species significantly differed from the calculated mean value. For instance, intolerant to very intolerant species Pinus cembra and P. mugo (Wiesner 1907, Warming 1909) and Populus tremula (Ellenberg 1991) were classified as intermediately shade tolerant, and tolerant to very tolerant species Acer platanoides as intolerant or intermediate (Gayer 1898, Otto 1994) in some assessments. To control for such clearly erroneous data, estimates of species light requirement in any single data set that differed by more than two levels from the general species mean were removed, and the corrected species mean value was calculated.
Shade tolerance of East-Asian species
For East-Asian species, we used the study of Kikuzawa (1984) as the starting point. This study provided a five-level scale of species dispersal across the understory-open continuum. This set of data was augmented using the assessments of species successional position in Koike (1988) and Maruyama (1978) . Again, a common shade tolerance scale was obtained using linear regressions developed for overlapping species in specific data sets. Species shade tolerance estimates for a more limited set of species were also obtained from a large set of studies reporting data of forest succession and species ecophysiological characteristics (e.g. Kohyama 1984, Ohsawa et al. 1986, Kikuzawa 1988, Peters 1992, Kamijo and Okutomi 1995b, 1995a, Ozaki and Ohsawa 1995, Peters et al. 1995, Sumida 1995, Tanouchi and Yamamoto 1995, Nakashizuka and Iida 1996, Tanouchi 1996, Ohsawa and Nitta 1997, Peters 1997, Suzuki 1997, Hiroki and Ichino 1998, Lei et al. 1998, Ke and Werger 1999, Masaki 2002, Hiroki 2003, Ishii et al. 2003) . Studies that reported broad differentiation of species groups during succession (Ohsawa et al. 1986, Kamijo and Okutomi 1995b, 1995a, Ozaki and Ohsawa 1995, Nakashizuka and Iida 1996, Ohsawa and Nitta 1997, Masaki 2002) were used first to develop the initial shade tolerance ranking for specific species. Detailed studies reporting survival and growth of species of similar successional sequence (Peters et al. 1995, Tanouchi and Yamamoto 1995, Tanouchi 1996, Peters 1997, Hiroki and Ichino 1998, Lei et al. 1998, Ke and Werger 1999, Hiroki 2003, Ishii et al. 2003) were further employed to fine-tune the species rankings. An average was calculated from all shade tolerance estimates per species. The final tolerance ranking was critically revised by Professors Kihachiro Kikuzawa (Kyoto University, Kyoto, Japan), Tohru Nakashizuka (Research Institute for Humanity & Nature, Kyoto, Japan), Masahiko Ohsawa (The University of Tokyo, Tokyo, Japan) and Tsutom Hiura (University of Hokkaido, Sapporo, Japan), and in the response to these expert assessments the tolerance rankings were changed by ± 0.25–1.0 tolerance units for a total of 26% of species. These modifications altered the overall ranking somewhat, but for these 26% of East-Asian species, the modified rankings were strongly correlated with the rankings before modification (r = 0.88, P < 0.001; r = 0.95 for the entire data set). Thus, while these revisions significantly enhanced the reliability of the East-Asian species ranking, they did not qualitatively alter the literature-based ranking.
Waterlogging tolerance
Waterlogging tolerance rankings for the North-American species were obtained from nine primary data sources: Barnes (1991) (73 North-American native species), Bell and Johnson (1974, Bratkovich et al. 1993) (23 North-American native species), Kuhns and Rupp (2000) (210 species of which 115 were native, 37 introduced from Europe and 57 from East Asia), Minore (1979) (14 North-American native species), online USDA Plants database (USDA NRCS 2005) (altogether 369 species, of which 314 were native to North America, 23 to Europe and 29 to East Asia), Tesche (1992) (21 North-American native species), U.S. Department of Agriculture Natural Resources Conservation Service (1996) (56 native species) and Whitlow and Harris (1979, Bratkovich et al. 1993) (57 North-American native species).
Although the USDA Plants database was the most extensive, the data for different genera have been collected and revised by different authors such that the overall scoring scales for various families differed. This database also scored the species anaerobic resistance on a four-level scale. Therefore, this database was used to get the initial estimates of species waterlogging tolerance, and the waterlogging estimates of this database were revised using the data from White (1973) and Iles and Gleason (1994) , and our own knowledge of species biology. As with the shade tolerance, different sets of data were cross-calibrated using linear regression analyses with waterlogging tolerance estimates in common species among the data sets, and an average waterlogging tolerance value was calculated. Despite the definitions of waterlogging tolerance differed from study to study, the correlation among the data sets was generally good (r > 0.80, average ± SE r = 0.86 ± 0.01, P < 0.001 for all comparisons), suggesting that we obtained reliable average estimates of species waterlogging tolerance. As several extensive data sets used in our study provide insufficient separation for several species, the final rankings were further refined using a series of studies reporting information of dispersal of species along wetland-upland continua as well as using reports of ecophysiological common garden investigations (Hosner 1958, Harms et al. 1980, Jones and Sharitz 1989, Jones et al. 1994, Ranney 1994, Ranney and Bir 1994, Yin et al. 1994, Hoagland et al. 1996, Naiman et al. 1998, Bendix and Hupp 2000, Dale and Ware 2004) .For the European species, waterlogging tolerance estimates were obtained from Glenz (2005) (65 native species), Merritt (1994) (24 native species), Prentice and Helmisaari (1991) (20 native species) and Schaffrath (2000) (48 species of which 44 were native to Europe, 6 to North America and one to East Asia) and Tesche (1992) (8 native species). For 42 European native species, data of the ecological requirements, including the waterlogging tolerance, were available in the Biological Flora of British Isles review series published regularly by The Journal of Ecology (1941–2005) . Specific studies of comparative flooding tolerance (Frye and Grosse 1992, Tapper 1993, Ranney 1994, Ranney and Bir 1994, van Splunder et al. 1995, Anonymous 1996, Tapper 1996, Siebel and Blom 1998, Siebel et al. 1998, van Splunder 1998, Burkart 2001, Karrenberg et al. 2002, Kreuzwieser et al. 2002) were also examined to extend the database and identify potentially erroneous species scorings. For instance, the moderately waterlogging tolerant species Fraxinus excelsior is assigned a low waterlogging tolerance estimate of 1.5–2 in some multispecies rankings (Merritt 1994, Anonymous 1996) , and the relatively tolerant species Alnus glutinosa a value of 2–3 (Brzeziecki and Kienast 1994, Schaffrath 2000) ; other rankings (Prentice and Helmisaari 1991, Glenz 2005) along with comparative ecophysiological studies (Tapper 1993, 1996, Siebel and Blom 1998, Siebel et al. 1998) resulted in corrected estimates of 2.7 ± 0.3 for F. excelsior and 3.9 ± 0.2 for A. glutinosa.
For species-rich families such as Betulaceae, Ericaceae, Rosaceae, and Salicaceae, waterlogging tolerance of less frequent species was estimated on the basis of species dispersal patterns across wet to dry habitats using country-specific floras (e.g. Vaga et al. 1960, Oberdorfer et al. 1994) , and our own knowledge of species biology. All these estimates were cross-calibrated using linear regressions (r > 0.77 for data sets including more than 20 species, average ± SE r = 0.80 ± 0.03, P < 0.001 for all comparisons), and average waterlogging tolerance estimates were calculated. Finally, the resulting overall rankings were converted to the 5-level scale derived for North-American species using the waterlogging estimates of the species common in both North-American and European waterlogging tolerance assessments (Fig. 2A).
Drought tolerance
For the North-American species, the drought tolerance rankings (very intolerant, 1; intolerant, 2; moderately tolerant, 3; tolerant, 4; very tolerant, 5) were derived from four main data sources: Kuhns and Rupp (2000) and (Cerny et al. 2002) (altogether 214 species, of which 119 were native to North America, 37 to Europe and 57 to East Asia), Meerow and Norcini (1997) (73 native species), Minore (1979) (23 native species) and online USDA Plants database (USDA NRCS 2005) (altogether 366 species, of which 309 were native to North America, 22 to Europe and 32 to East Asia). As with waterlogging tolerance, online sources were used to get initial estimates of drought tolerance. All drought-tolerance estimates were strongly correlated with each other (r > 0.80, average ± SE r = 0.86 ± 0.02, P <0.001 for all comparisons). 6}. Comparative studies on species drought tolerance (e.g., Abrams 1990, Ni and Pallardy 1991, e.g., Ranney et al. 1991, Tyree and Alexander 1993, Abrams et al. 1994, Kubiske and Abrams 1994, Sperry et al. 1994, Kubiske et al. 1996, Linton et al. 1998, Loewenstein and Pallardy 1998) were further employed to refine the drought tolerance rankings. As with shade and waterlogging tolerance, different drought tolerance scales were cross-calibrated using species common in specific data sets, and an average drought tolerance score was determined for each species.
For the European species, data on species water requirements were available from a series of studies that also provided the data for shade tolerance (Table 1 for the number of species): Brzeziecki and Kienast (1994) , Ellenberg (1996) , Ellenberg (1991) , Hill et al. (1999) , Jahn (1991) and Otto (1994) . In addition, Brzeziecki’s (1995) estimates of drought tolerance for 41 native European species were also included. While most of these studies have scored species drought tolerance, the ecological indicator values for soil moisture by Ellenberg (Ellenberg 1991, Hill et al. 1999) characterize species occurrence along the gradient of water availability. Thus, the ecological indicator value for soil moisture actually combines both species drought and waterlogging tolerances rather than measures the drought tolerance per se. The indicator values for soil moisture varied from 3 (very dry) to 9 (very wet) for the European woody species, and the species with high waterlogging tolerance generally had a large indicator value despite of potentially high drought tolerance. For instance, most species of Ericaceae family and several species of Pinaceae were characterized by high values of soil moisture indicator value, but are actually both waterlogging and drought tolerant and can growth in mires as well as dry heaths. Such polytolerant species were identified on the basis of species waterlogging tolerance and drought tolerance estimates obtained from other studies and our own knowledge on species biology. The polytolerant species were removed from Ellenberg (1991) and Hill et al. (1999) data sets, and the drought tolerance of these species was assessed using other data sets.Ellenberg’s indicator values for soil moisture of Central Europe (Ellenberg 1991) and Great Britain species (Hill et al. 1999) were averaged, and all species drought tolerance rankings were cross-calibrated by linear regressions using species common in specific data sets. All different scales were strongly correlated, but the scatter was somewhat larger than among North-American data sets (r > 0.62, average ± SE r = 0.74 ± 0.02, P <0.001 for all comparisons). To enhance the reliability of species scorings with the largest discrepancy among the studies, assessments from several comparative ecophysiological studies (e.g. Ranney et al. 1991, Acherar and Rambal 1992, Epron et al. 1993, Epron 1997, Aasamaa and Sõber 2001, Aasamaa et al. 2004, Cochard et al. 2004) were employed. The final drought tolerance scoring was taken as an average of all available estimates.
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