Ecological Archives M076-020-A2

Ülo Niinemets and Fernando Valladares. 2006. Tolerance to shade, drought, and waterlogging of temperate Northern Hemisphere trees and shrubs. Ecological Monographs 76:521–547.

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.


Aasamaa, K., and A. Sõber. 2001. Hydraulic conductance and stomatal sensitivity to changes of leaf water status in six deciduous tree species. Biologia Plantarum 44:65–73.

Aasamaa, K., A. Sõber, W. Hartung, and Ü. Niinemets. 2004. Drought acclimation of two deciduous tree species of different layers in a temperate forest canopy. Trees: Structure and Function 18:93–101.

Abrams, M. D. 1990. Adaptations and responses to drought in Quercus species of North America. Tree Physiology 7:227–238.

Abrams, M. D., M. E. Kubiske, and S. A. Mostoller. 1994. Relating wet and dry year ecophysiology to leaf structure in contrasting temperate tree species. Ecology 75:123–133.

Acherar, M., and S. Rambal. 1992. Comparative water relations of four Mediterranean oak species. Vegetatio 99-100:177–184.

Anonymous. 1996. Auwälder in Südbayern. Standörtliche Grundlagen und Bestockungsverhältnisse im Staatswald. LWF Bericht, 9. Bayerische Landesanstalt für Wald und Forstwirtschaft, Freising.

Baker, F. S. 1949. A revised tolerance table. Journal of Forestry 47:179–181.

Barnes, B. V. 1991. Deciduous forests of North America. Pages 219–344 in E. Röhrig and B. Ulrich, editors. Temperate deciduous forests. Ecosystems of the world, 7. Elsevier, Amsterdam - London - New York - Tokyo.

Bell, D. T., and E. L. Johnson. 1974. Flood-caused tree mortality around Illinois reservoirs. Transactions of the Illinois State Academy of Science 67:28–37.

Bendix, J., and C. R. Hupp. 2000. Hydrological and geomorphological impacts on riparian plant communities. Hydrological Processes 14:2977–2990.

Bratkovich, S., L. Burban, S. Katovich, C. Locey, J. Pokorny, and R. Wiest. 1993. Flooding and its effect on trees: Information packet. Forest Resources Management and Forest Health Protection, USDA Forest Service, Northeastern Area State and Private Forestry, St. Paul, Minnesota, USA.

Brzeziecki, B. 1995. Skale nominalne wymagan klimatycznych gatunków drzew lesnych. (Nominal scales for climatic requirements of forest tree species). Sylwan 139:53–65.

Brzeziecki, B., and F. Kienast. 1994. Classifying the life-history strategies of trees on the basis of the Grimian model. Forest Ecology and Management 69:167–187.

Burkart, M. 2001. River corridor plants (Stromtalpflanzen) in Central European lowland: a review of a poorly understood plant distribution pattern. Global Ecology and Biogeography 10:449–468.

Cerny, T. A., M. Kuhns, K. L. Kopp, and M. Johnson. 2002. Efficient irrigation of trees and shrubs. Utah State University Extension, Logan, UT. Electronic Publishing, HG-523.

Cochard, H., F. Froux, S. Mayr, and C. Coutand. 2004. Xylem wall collapse in water-stressed pine needles. Plant Physiology 134:401–408.

Coomes, D. A., and P. J. Grubb. 2000. Impacts of root competition in forests and woodlands: a theoretical framework and review of experiments. Ecological Monographs 70:171–207.

Cornwell, W. K., and P. J. Grubb. 2003. Regional and local patterns in plant species richness with respect to resource availability. Oikos 100:417–428.

Dale, E. E., Jr., and S. Ware. 2004. Distribution of wetland tree species in relation to a flooding gradient and backwater versus streamside location in Arkansas, U.S.A. The Journal of the Torrey Botanical Society 131:177–186.

Ellenberg, H. 1991. Zeigerwerte der Gefäßpflanzen (ohne Rubus). Pages 9-166 in H. Ellenberg, R. Düll, V. Wirth, W. Werner, and D. Paulißen, editors. Zeigerwerte von Pflanzen in Mitteleuropa. Scripta Geobotanica, 18. Erich Goltze KG, Göttingen.

Ellenberg, H. 1996. Vegetation Mitteleuropas mit den Alpen in ökologischer, dynamischer und historischer Sicht. 5th edition. Verlag Eugen Ulmer, Stuttgart. 1096.

Epron, D. 1997. Effects of drought on photosynthesis and on the thermotolerance of photosystem II in seedlings of cedar (Cedrus atlantica and C. libani). Journal of Experimental Botany 48:1835–1841.

Epron, D., E. Dreyer, and G. Aussenac. 1993. A comparison of photosynthetic responses to water stress in seedlings from 3 oak species: Quercus petraea (Matt.) Liebl., Q. rubra L. and Q. cerris L. Annales des Sciences Forestieres 50:48–60.

Frye, J., and W. Grosse. 1992. Growth responses to flooding and recovery of deciduous trees. Zeitschrift für Naturforschung, Section C Journal of Biosciences 47c:683–689.

Gayer, K. 1898. Der Waldbau. vierte, verbesserte Auflage edition. Verlagsbuchhandlung Paul Parey, Berlin, Germany.

Glenz, C. 2005. Process-based, spatially-explicit modelling of riparian forest dynamics in Central Europe – tool for decisionmaking in river restoration. École Polytechnique Fédérale de Lausanne, Lausanne.

Harms, W. R., H. T. Schreuder, D. D. Hook, and C. L. Brown. 1980. The effects of flooding on the swamp forest in Lake Ocklawaha, Florida. Ecology 61:1412–1421.

Hill, M. O., J. O. Mountford, D. B. Roy, and R. G. H. Bunce. 1999. Ellenberg's indicator values for British plants. Centre for Ecology & Hydrology, Huntingdon, Cambs. 46.

Hill, M. O., D. B. Roy, J. O. Mountford, and R. G. H. Bunce. 2000. Extending Ellenberg's indicator values to a new area: an algorithmic approach. Journal of Applied Ecology 37:3–15.

Hiroki, S. 2003. Early life history stage and segregative distribution of Fagaceae in Japan. Pages 14-14 in Integration of silviculture and genetics in creating and sustaining oak forests. Joint meeting of IUFRO working groups. Genetics of Quercus & improvement and silviculture of oaks. OAK 2003, Japan, Tsukuba, Japan, 29 Sep.~ 3 Oct. 2003.

Hiroki, S., and K. Ichino. 1998. Comparison of growth habits under various light conditions between two climax species, Castanopsis sieboldii and Castanopsis cuspidata, with special reference to their shade tolerance. Ecological Research 13:65–72.

Hoagland, B. W., L. R. Sorrels, and S. M. Glenn. 1996. Woody species composition of floodplain forests of the Little River, McCurtain and LeFlore Counties, Oklahoma. Proc. Okla. Acad. Sci. 76:23–29.

Hosner, J. F. 1958. The effects of complete inundation upon seedlings of six bottomland tree species. Ecology 39:371–373.

Iles, J., and M. Gleason. 1994. Understanding the effects of flooding on trees. Sustainable urban landscapes, 1. Iowa State University Extension, Urbandale.

Ishii, H., M. Ooishi, Y. Maruyama, and T. Koike. 2003. Acclimation of shoot and needle morphology and photosynthesis of two Picea species to differences in soil nutrient availability. Tree Physiology 23:453–461.

Ivanov, L. A., and N. L. Kossovich. 1932. O rabote assimilyatsionnovo apparata drevesnyh porod. II. Botanicheskii Zhurnal 17:3–71.

Jahn, G. 1991. Temperate deciduous forests of Europe. Pages 377-502 in E. Röhrig and B. Ulrich, editors. Temperate deciduous forests. Ecosystems of the world, 7. Elsevier, Amsterdam - London - New York - Tokyo.

Jones, R. H., and R. R. Sharitz. 1989. Potential advantages and disadvantages of germinating early for trees in floodplain forests. Oecologia 81:443–449.

Jones, R. H., R. R. Sharitz, P. M. Dixon, D. S. Segal, and R. L. Schneider. 1994. Woody plant regeneration in four floodplain forests. Ecological Monographs 64:345–367.

Journal of Ecology. 1941–2005. The Biological Flora. British Ecological Society, London.

Kamijo, T., and K. Okutomi. 1995a. Distribution of Castanopsis forest and Persea forest and its causal factors on the southern part of Izu Islands. J. Phytogeogr. & Taxon. 43:67–73.

Kamijo, T., and K. Okutomi. 1995b. Seedling establishment of Castanopsis cuspidata var. sieboldii and Persea thunbergii on lava and scoria of the 1962 eruption on Miyake-jima Island, the Izu Islands. Ecological Research 10:235–242.

Karrenberg, S., P. J. Edwards, and J. Kollmann. 2002. The life history of Salicaceace living in the active zone of floodplains. Freshwater Biology 47:733–748.

Ke, G., and M. J. A. Werger. 1999. Different responses to shade of evergreen and deciduous oak seedlings and the effect of acorn size. Acta Oecologica 20:579–586.

Kikuzawa, K. 1984. Leaf survival of woody plants in deciduous broad-leaved forests. 2. Small trees and shrubs. Canadian Journal of Botany 62:2551–2556.

Kikuzawa, K. 1988. Leaf survivals of tree species in deciduous broad-leaved forests. Plant Species Biology 3:67–76.

Kobe, R. K., S. W. Pacala, J. A. Silander, Jr., and C. D. Canham. 1995. Juvenile tree survivorship as a component of shade tolerance. Ecological Applications 5:517–532.

Kohyama, T. 1984. Regeneration and coexistence of two Abies species dominating subalpine forests in central Japan. Oecologia 62:156–161.

Koike, T. 1988. Leaf structure and photosynthetic performance as related to the forest succession of deciduous broad-leaved trees. Plant Species Biology 3:77–87.

Kreuzwieser, J., S. Fürniss, and H. Rennenberg. 2002. Impact of waterlogging on the N-metabolism of flood tolerant and non-tolerant tree species. Plant, Cell and Environment 25:1039–1049.

Kubiske, M. E., and M. D. Abrams. 1994. Ecophysiological analysis of woody species in contrasting temperate communities during wet and dry years. Oecologia 98:303–312.

Kubiske, M. E., M. D. Abrams, and S. A. Mostoller. 1996. Stomatal and nonstomatal limitations of photosynthesis in relation to the drought and shade tolerance of tree species in open and understory environments. Trees: Structure and Function 11:76–82.

Kuhns, M., and L. Rupp. 2000. Selecting and planting landscape trees. Electronic Publishing, NR-460. Utah State University Extension, Logan, UTah, USA.

Küppers, M. 1989. Ecological significance of aboveground architectural patterns in woody plants: a question of cost-benefit relationships. Trends in Ecology & Evolution 4:375–379.

Lei, T. T., R. Tabuchi, M. Kitao, K. Takahashi, and T. Koike. 1998. Effects of season, weather and vertical position on the variation in light quantity and quality in a Japanese decidous broadleaf forest. Journal of Sustainable Forestry 6(1/2):35–55.

Linton, M. J., J. S. Sperry, and D. G. Williams. 1998. Limits to water transport in Juniperus osteosperma and Pinus edulis: implications for drought tolerance and regulation of transpiration. Functional Ecology 12:906–911.

Loewenstein, N. J., and S. G. Pallardy. 1998. Drought tolerance, xylem sap abscisic acid and stomatal conductance during soil drying: a comparison of young plants of four temperate deciduous angiosperms. Tree Physiology 18:421–430.

Maruyama, K. 1978. Ecological studies on natural beech forest. 32. Shoot elongation characteristics and phenological behavior of forest trees in natural beech forest. Bulletin of the Niigata University Forests 11:1–30.

Masaki, T. 2002. Structure and dynamics. Pages 53–65 in T. Nakashizuka and Y. Matsumoto, editors. Diversity and interaction in a temperate forest community - Ogawa Forest Reserve of Japan.   Ecological Studies, 158. Springer Verlag, Berlin, Germany.

Meerow, A. W., and J. G. Norcini. 1997. Native trees for North Florida. Pages Date first printed: September 1989. Reviewed: June 1997. in. Department of Environmental Horticulture, Florida Cooperative Extension Service, Institute of Food and Agricultural Sciences, University of Florida, Gainesville, Florida, USA.

Merritt, A. 1994. Wetlands, industry & wildlife. A manual of principles and practices. The Wildfowl & Wetlands Trust, Slimbridge, Gloucester, UK.

Minore, D. 1979. Comparative autecological characteristics of northwestern tree species. A literature review. United States Department of Agriculture Pacific Northwest Forest and Range Experiment Station Technical Report, 87. Pacific Northwest Forest and Range Experiment Station, U. S. Department of Agriculture, Forest Service, Portland, Oregon. 72.

Morozov, G. F. 1903. Forest Dendrology. Attitude of tree species to light. On the types of of tree stands. A conspectus of lectures. S.-Peterb. Lesnoi Inst., St. Peterburg.

Naiman, R. J., K. L. Fetherston, S. McKay, and J. Chen. 1998. Riparian forests. Pages 289–323 in R. J. Naiman and E. Bilby, editors. River ecology and management: lessons from the Pacific coastal ecoregion. Springer Verlag, New York, New York, USA.

Nakashizuka, T., and S. Iida. 1996. Composition, dynamics and disturbance regime of temperate deciduous forests in monsoon Asia. Pages 23–30 in T. Hirose, B. H. Walker, H. A. Mooney, and A. Kratochwil, editors. Global change and terrestrial ecosystems in monsoon Asia.   Tasks for vegetation science, 33. Kluwer Academic Publishers, Dordrecht, Boston, London.

Ni, B.-R., and S. G. Pallardy. 1991. Response of gas exchange to water stress in seedlings of woody angiosperms. Tree Physiology 8:1–9.

Niinemets, Ü., and K. Kull. 1994. Leaf weight per area and leaf size of 85 Estonian woody species in relation to shade tolerance and light availability. Forest Ecology and Management 70:1–10.

Oberdorfer, E., T. Müller, D. Korneck, W. Lippert, E. Patzke, and H. E. Weber. 1994. Pflanzensoziologische Exkursionsflora. UTB für Wissenschaft: Uni-Taschenbücher, 1828, 7th edition. Verlag Eugen Ulmer, Stuttgart. 1050.

Ohsawa, M., and I. Nitta. 1997. Patterning of subtropical/warm-temperate evergreen broad-leaved forests in East Asian mountains with special reference to shoot phenology. Tropics 6:317–334.

Ohsawa, M., P. R. Shakya, and M. Numata. 1986. Distribution and succession of west Himalaya forest types in the eastern part of the Nepal Himalaya. Mountain Research and Development 6:143–157.

Otto, H.-J. 1994. Waldökologie. Verlag Eugen Ulmer, Stuttgart. 391.

Ozaki, K., and M. Ohsawa. 1995. Successional change of forest pattern along topographical gradients in warm-temperate mixed forests in Mt Kiyosumi, central Japan. Ecological Research 10:223–234.

Peters, R. 1992. Ecology of beech forests in the northern hemisphere. Ph.D. Thesis. Landbouwuniversiteit Wageningen, Wageningen.

Peters, R. 1997. Beech forests. Geobotany, 24. Kluwer Academic Publishers, Dordrecht - Boston - London. 169.

Peters, R., H. Tanaka, M. Shibata, and T. Nakashizuka. 1995. Light climate and growth in shade-tolerant Fagus crenata, Acer mono and Carpinus cordata. Ecoscience 2:67–74.

Prentice, I. C., and H. Helmisaari. 1991. Silvics of North-European trees: compilation, comparisons and implications for forest succession modelling. Forest Ecology and Management 42:79–93.

Ranney, T. G. 1994. Differential tolerance of eleven Prunus taxa to root zone flooding. Journal of Environmental Horticulture 12:138–141.

Ranney, T. G., and R. E. Bir. 1994. Comparative flood tolerance of birch rootstocks. Journal of the American Society for Horticultural Science 119:43–48.

Ranney, T. G., R. E. Bir, and W. A. Skroch. 1991. Comparative drought resistance among six species of birch (Betula): influence of mild water stress on water relations and leaf gas exchange. Tree Physiology 8:351–360.

Schaffrath, J. 2000. Auswirkungen des extremen Sommerhochwassers des Jahres 1997 auf die Gehölzwegetation in der Oderaue bei Frakfurt (O.). Naturschutz und Landespflege in Brandenburg 9:4–13.

Siebel, H. N., and C. W. P. M. Blom. 1998. Effects of irregular flooding on the establishment of tree species. Acta Botanica Neerlandica 47:231–240.

Siebel, H. N., M. v. Wijk, and C. W. P. M. Blom. 1998. Can tree seedlings survive increased flood levels of rivers? Acta Botanica Neerlandica 46:219–230.

Sperry, J. S., K. L. Nichols, J. E. M. Sullivan, and S. E. Eastlack. 1994. Xylem embolism in ring-porous, diffuse-porous, and coniferous trees of northern Utah and interior Alaska. Ecology 75:1736–1752.

Sumida, A. 1995. Three-dimensional structure of a mixed broad-leaved forest in Japan. Vegetatio 119:67–80.

Suzuki, E. 1997. The dynamics of old Cryptomeria japonica forest on Yakushima Island. Tropics 6:421–428.

Tanouchi, H. 1996. Survival and growth of two coexisting evergreen oak species after germination under different light conditions. International Journal of Plant Sciences 157:516–522.

Tanouchi, H., and S. Yamamoto. 1995. Structure and regeneration of canopy species in an old-growth evergreen broad-leaved forest of Aya district, southwestern Japan. Vegetatio 117:51–60.

Tapper, P.-G. 1993. The replacement of Alnus glutinosa by Fraxinus excelsior during succession related to regenerative differences. Ecography 16:212–218.

Tapper, P.-G. 1996. Tree dynamics in a successional Alnus-Fraxinus woodland. Ecography 19:237–244.

Tesche, M. 1992. Klimaresistenz. Pages 279–306 in H. Lyr, H. J. Fiedler, and W. Tranquillini, editors. Physiologie und Ökologie der Gehölze. Gustav Fischer Verlag, Jena.

Tyree, M. T., and J. D. Alexander. 1993. Hydraulic conductivity of branch junctions in three temperate tree species. Trees: Structure and Function 7:156–159.

U. S. Department of Agriculture Natural Resources Conservation Service. 1996. Chapter 16. streambank and shoreline protection. in National engineering handbook, Part 650 - engineering field handbook. USDA NRCS, Washington, D.C., USA.

USDA NRCS. 2005. The PLANTS Database, Version 3.5. Data compiled from various sources by Mark W. Skinner. National Plant Data Center, Information Technology Center, Baton Rouge, LA, USA.

Vaga, A., K. Eichwald, and et al., editors. 1960. Eesti NSV floora. Eesti Riiklik Kirjastus/Valgus, Tallinn.

Valladares, F., J. M. Chico, I. Aranda, L. Balaguer, P. Dizengremel, E. Manrique, and E. Dreyer. 2002. The greater seedling high-light tolerance of Quercus robur over Fagus sylvatica is linked to a greater physiological plasticity. Trees: Structure and Function 16:395–403.

van Splunder, I. 1998. Floodplain forest recovery. Softwood development in relation to hydrology, riverbank morphology and management. PhD Dissertation. Katholieke Universiteit Nijmgen, Nijmgen.

van Splunder, I., H. Coops, L. A. C. J. Voesenek, and C. W. P. M. Blom. 1995. Establishment of alluvial forest species in floodplains: the role of dispersal timing, germination characteristics and water-level fluctuations. Acta Botanica Neerlandica 44:269–278.

Walter, H. 1968. Die Vegetation der Erde in öko-physiologischer Betrachtung. G. Fischer, Stuttgart.

Walters, M. B., and P. B. Reich. 1999. Research review. Low-light carbon balance and shade tolerance in the seedlings of woody plants: do winter deciduous and broad-leaved evergreen species differ? The New Phytologist 143:143–154.

Warming, E. 1909. Oecology of plants. An introduction to the study of plant communities. Oxford University Press, Oxford. 422.

White, R. M. 1973. Plant tolerance for standing water: an assessment. Cornell Plantations 28:50–52.

Whitlow, T. H., and R. W. Harris. 1979. Flood tolerance in plants: a state-of-the-art review. National Technical Information Service, U.S. Department of Commerce, Washington, D.C. 161.

Wiesner, J. 1907. Der Lichtgenuss der Pflanzen. Photometrische und physiologische Untersuchungen mit besonderer Rücksichtnahme auf Lebensweise, geographische Verbreitung und Kultur der Pflanzen. Verlag von Wilhelm Engelmann, Leipzig.

Yevstigneyev, O. I. 1990. Fitotsenotipy i otnosheniye listvennyh derevyev k svetu. (Phytocoenotypes and the behaviour of deciduous trees with respect to light). Cand. Biol. Dissertation. Moskovskii Gosudarstvennyi Pedagogicheskii Institut imeni V. I. Lenina, Moscow, Russia.

Yin, Y., J. C. Nelson, G. V. Swenson, H. A. Langrehr, and T. A. Blackburn. 1994. Tree mortality in the upper Mississippi river and floodplain following an extreme flood in 1993. Pages 41–60 in Long term resource monitoring program 1993 flood observations. National Biological Service, Environmental Management Technical Center, Onalaska, Wisconsin, USA.

Zon, R., and H. S. Graves. 1911. Light in relation to tree growth. U.S. Department of Agriculture, Forest Service - Bulletin, 92. Government Printing Office, Washington. 59.

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