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Hackett-Pain, A. J., Ascoli, D., Vacchiano, G., Biondi, F., Cavin, L., Conedera, M., Drobyshev, I., Liñán, I. D., Friend, A. D., Grabner, M., Hartl, C., Kreyling, J., Lebourgeois, F., Levanič, T., Menzel, A., Maaten, E., Maaten-Theunissen, M., Muffler, L., Motta, R., ... Zang, C. S. (2018). Climatically controlled reproduction drives interannual growth variability in a temperate tree species. *Ecology Letters*, 21(12), 1833-1844. https://doi.org/10.1111/ele.13158

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Han, Q., Kabeya, D., & Hoch, G. (2011). Leaf traits, shoot growth and seed production in mature *Fagus sylvatica* trees after 8 years of CO2 enrichment. *Annals of Botany*, 107(8), 1405-1411. https://doi.org/10.1093/aob/mcr082

Hanner, M., Aitken, S. N., Ericsson, T., & Ying, C. C. (2001). Inheritance of strobili production at genetic correlation with growth in lodgepole pine. *Forest Genetics*, 8(4), 323-329.

Hansen, M. M., Olivieri, I., Waller, D. M., & Nielsen, E. E. (2012). Monitoring adaptive genetic responses to environmental change. *Molecular Ecology*, 21, 1311-1329.

Hansen, T. F., Pélabon, C., & Houle, D. (2011). Heritability is not evolvability. *Evolutionary Biology*, 38(3), 258-277. https://doi.org/10.1007/s11692-011-9127-6

Hermida, L., Sánchez, J. L., López, L., Berthet, C., Dessens, J., García-Ortega, E., & Merino, A. (2013). Climatic trends in hail precipitation in France: Spatial, altitudinal, and temporal variability. *The Scientific World Journal*, 2013, 1-10. https://doi.org/10.1155/2013/494971

Hoch, G., Siegwolf, R. T. W., Keel, S. G., Körner, C., & Han, Q. (2013). Fruit production in three masting tree species does not rely on stored carbon reserves. *Oecologia*, 171(3), 653-662. https://doi.org/10.1007/s00442-012-2579-2

Hoffmann, A. A., & Sgrò, C. M. (2011). Climate change and evolutionary adaptation. *Nature*, 470(7335), 479-485. https://doi.org/10.1038/nature09670

Houston Durrant, T., de Rigo, D., & Caudullo, G. (2016). *Fagus sylvatica* in Europe: Distribution, habitat, usage and threats. In J. San-Miguel-Ayanz, D. de Rigo, G. Caudullo, T. Houston Durrant, & A. Mauri (Eds.), *European atlas of forest tree species* (p. e012b90+). Publications Office of the European Union.

Howe, G. T., Aitken, S. N., Neale, D. B., Jermstad, K. D., Wheeler, N. C., & Chen, T. H. (2003). From genotype to phenotype: Unraveling the complexities of cold adaptation in forest trees. *Canadian Journal of Botany*, 81(12), 1247-1266. https://doi.org/10.1139/b03-141

Hufford, K. M., & Hamrick, J. L. (2003). Viability selection at three early life stages of the tropical tree, *Platypodium elegans* (Fabaceae, Papilionoideae). *Evolution*, 57(3), 518-526. https://doi.org/10.1111/j.0014-3820.2003.tb01543.x

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Inouye, D. W. (2000). The ecological and evolutionary significance of frost in the context of climate change. *Ecology Letters*, 3(5), 457-463. https://doi.org/10.1046/j.1461-0248.2000.00165.x

Keenan, T. F., Gray, J., Friedl, M. A., Toomey, M., Bohrer, G., Hollinger, D. Y., Munger, J. W., O'Keefe, J., Schmid, H. P., Wing, I. S., Yang, B., & Richardson, A. D. (2014). Net carbon uptake has increased through warming-induced changes in temperate forest phenology. *Nature Climate Change*, 4(7), 598-604. https://doi.org/10.1038/nclimate2253

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Kruuk, L. E., Clutton-Brock, T. H., Slate, J., Pemberton, J. M., Brotherstone, S., & Guinness, F. E. (2000). Heritability of fitness in a wild mammal population. *Proceedings of the National Academy of Sciences of the United States of America*, 97(2), 698-703. https://doi.org/10.1073/pnas.97.2.698

Kruuk, L. E. B., Slate, J., & Wilson, A. J. (2008). New answers for old questions: The evolutionary quantitative genetics of wild animal populations. *Annual Review of Ecology, Evolution, and Systematics*, 39(1), 525-548. https://doi.org/10.1146/annurev.ecolsys.39.110707.173542

Kulbaba, M. W., Sheth, S. N., Pain, R. E., Eckhart, V. M., & Shaw, R. G. (2019). Additive genetic variance for lifetime fitness and the capacity for adaptation in an annual plant. *Evolution*, 73(9), 1746-1758. https://doi.org/10.1111/evo.13830

Lande, R., & Arnold, S. J. (1983). The measurement of selection on correlated characters. *Evolution*, 37(6), 1210-1226.

Lee, E. A., Deen, W., Hooyer, M. E., Chambers, A., Parkin, G., Gordon, R., & Singh, A. K. (2016). Involvement of year-to-year variation in thermal time, solar radiation and soil available moisture in genotype-by-environment effects in maize. *Crop Science*, 56, 2180-2192. https://doi.org/10.2135/cropsci2015.04.0231

Lefevre, S., Wagner, S., Petit, R. J., & De Lafontaine, G. (2012). Multiplexed microsatellite markers for genetic studies of beech. *Molecular Ecology Resources*, 12, 484-491. https://doi.org/10.1111/j.1755-0998.2011.03094.x

Li, Y., Suontama, M., Burdon, R. D., & Dungey, H. S. (2017). Genotype by environment interactions in forest tree breeding: Review of methodology and perspectives on research and application. *Tree Genetics & Genomes*, 13, 60. https://doi.org/10.1007/s11295-017-1144-x
Gienapp, P., Fior, S., Guillaume, F., Lasky, J. R., Sork, V. L., & Csilléry, K. (2017). Genomic quantitative genetics to study evolution in the wild. *Trends in Ecology & Evolution, 32*(12), 897-908. https://doi.org/10.1016/j.tree.2017.09.004

González-Martínez, S. C., Burczyk, J., Nathan, R., Gil, L., & Alía, R. (2006). Effective gene dispersal and female reproductive success in Mediterranean maritime pine (Pinus pinaster Aiton). *Molecular Ecology, 15*, 4577-4588.

Gömöry, D., & Paule, L. (2011). Trade-off between height growth and spring flushing in common beech (*Fagus sylvatica* L.). *Annals of Forest Science, 68*(5), 975-984. https://doi.org/10.1007/s13595-011-0103-1

Gömöry, D., Paule, L., & Vyšný, J. (2007). Patterns of allozyme variation in western Eurasian *Fagus*. *Botanical Journal of the Linnean Society, 154*(2), 165-174. https://doi.org/10.1111/j.1095-8339.2007.00666.x

Grant, P. R., & Grant, B. R. (2014). Synergism of natural selection and introgression in the origin of a new species. *The American Naturalist, 183*(5), 671-681. https://doi.org/10.1086/675496

Gyllenstrand, N., Clapham, D., Källman, T., & Lagercrantz, U. (2007). A Norway spruce flowering locus T homolog is implicated in control of growth rhythm in conifers. *Plant Physiology, 144*(1), 248-257. https://doi.org/10.1104/pp.107.095802

Hackett-Pain, A., & Bogdziewicz, M. (2021). Climate change and plant reproduction: Trends and drivers of mast seeding change. *Philosophical Transactions of the Royal Society, B: Biological Sciences, 376*(1839), 20200379. https://doi.org/10.1098/rstb.2020.0379

Hackett-Pain, A. J., Ascoli, D., Vacchiano, G., Biondi, F., Cavin, L., Conedera, M., Drobyshev, I., Liñán, I. D., Friend, A. D., Grabner, M., Hartl, C., Kreyling, J., Lebourgeois, F., Levanič, T., Menzel, A., Maaten, E., Maaten-Theunissen, M., Muffler, L., Motta, R., ... Zang, C. S. (2018). Climatically controlled reproduction drives interannual growth variability in a temperate tree species. *Ecology Letters, 21*(12), 1833-1844. https://doi.org/10.1111/ele.13158

Hadfield, J. D. (2010). MCMC methods for multi-response generalized linear mixed models: The MCMCglmm R Package. *Journal of Statistical Software, 33*(2), i02. https://doi.org/10.18637/jss.v033.i02

Han, Q., Kabeya, D., & Hoch, G. (2011). Leaf traits, shoot growth and seed production in mature *Fagus sylvatica* trees after 8 years of CO2 enrichment. *Annals of Botany, 107*(8), 1405-1411. https://doi.org/10.1093/aob/mcr082

Hanner, M., Aitken, S. N., Ericsson, T., & Ying, C. C. (2001). Inheritance of strobili production at genetic correlation with growth in lodgepole pine. *Forest Genetics, 8*(4), 323-329.

Hansen, M. M., Olivieri, I., Waller, D. M., & Nielsen, E. E. (2012). Monitoring adaptive genetic responses to environmental change. *Molecular Ecology, 21*, 1311-1329.

Hansen, T. F., Pélabon, C., & Houle, D. (2011). Heritability is not evolvability. *Evolutionary Biology, 38*(3), 258-277. https://doi.org/10.1007/s11692-011-9127-6

Hermida, L., Sánchez, J. L., López, L., Berthet, C., Dessens, J., García-Ortega, E., & Merino, A. (2013). Climatic trends in hail precipitation in France: Spatial, altitudinal, and temporal variability. *The Scientific World Journal, 2013*, 1-10. https://doi.org/10.1155/2013/494971

Hoch, G., Siegwolf, R. T. W., Keel, S. G., Körner, C., & Han, Q. (2013). Fruit production in three masting tree species does not rely on stored carbon reserves. *Oecologia, 171*(3), 653-662. https://doi.org/10.1007/s00442-012-2579-2

Hoffmann, A. A., & Sgrò, C. M. (2011). Climate change and evolutionary adaptation. *Nature, 470*(7335), 479-485. https://doi.org/10.1038/nature09670

Houston Durrant, T., de Rigo, D., & Caudullo, G. (2016). *Fagus sylvatica* in Europe: Distribution, habitat, usage and threats. In J. San-Miguel-Ayanz, D. de Rigo, G. Caudullo, T. Houston Durrant, & A. Mauri (Eds.), *European atlas of forest tree species* (p. e012b90+). Publications Office of the European Union.

Howe, G. T., Aitken, S. N., Neale, D. B., Jermstad, K. D., Wheeler, N. C., & Chen, T. H. (2003). From genotype to phenotype: Unraveling the complexities of cold adaptation in forest trees. *Canadian Journal of Botany, 81*(12), 1247-1266. https://doi.org/10.1139/b03-141

Hufford, K. M., & Hamrick, J. L. (2003). Viability selection at three early life stages of the tropical tree, *Platypodium elegans* (Fabaceae, Papilionoideae). *Evolution, 57*(3), 518-526. https://doi.org/10.1111/j.0014-3820.2003.tb01543.x

Innes, J. L. (1994). The occurrence of flowering and fruiting on individual trees over 3 years and their effects on subsequent crown condition. *Trees, 8*(3), 139-150. https://doi.org/10.1007/BF00196638

Inouye, D. W. (2000). The ecological and evolutionary significance of frost in the context of climate change. *Ecology Letters, 3*(5), 457-463. https://doi.org/10.1046/j.1461-0248.2000.00165.x

Keenan, T. F., Gray, J., Friedl, M. A., Toomey, M., Bohrer, G., Hollinger, D. Y., Munger, J. W., O'Keefe, J., Schmid, H. P., Wing, I. S., Yang, B., & Richardson, A. D. (2014). Net carbon uptake has increased through warming-induced changes in temperate forest phenology. *Nature Climate Change, 4*(7), 598-604. https://doi.org/10.1038/nclimate2253

Klinkhamer, P. G. L., de Jong, T. J., & Metz, H. (1997). Sex and size in co-sexual plants. *Trends in Ecology & Evolution, 12*(7), 260-265. https://doi.org/10.1016/S0169-5347(97)01078-1

Koenig, W. D., Funk, K. A., Kraft, T. S., Carmen, W. J., Barringer, B. C., & Knops, J. M. H. (2012). Stabilizing selection for within-season flowering phenology confirms pollen limitation in a wind-pollinated tree: Phenology and acorn production in *Quercus lobata*. *Journal of Ecology, 100*(3), 758-763. https://doi.org/10.1111/j.1365-2745.2011.01941.x

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Kruuk, L. E., Clutton-Brock, T. H., Slate, J., Pemberton, J. M., Brotherstone, S., & Guinness, F. E. (2000). Heritability of fitness in a wild mammal population. *Proceedings of the National Academy of Sciences of the United States of America, 97*(2), 698-703. https://doi.org/10.1073/pnas.97.2.698

Kruuk, L. E. B., Slate, J., & Wilson, A. J. (2008). New answers for old questions: The evolutionary quantitative genetics of wild animal populations. *Annual Review of Ecology, Evolution, and Systematics, 39*(1), 525-548. https://doi.org/10.1146/annurev.ecolsys.39.110707.173542

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Vitasse, Y., Delzon, S., Bresson, C. C., Michalet, R., & Kremer, A. (2009). Altitudinal differentiation in growth and phenology among populations of temperate-zone tree species growing in a common garden. *Canadian Journal of Forest Research*, 39(7), 1259-1269. https://doi.org/10.1139/X09-054

Walsh, B., & Lynch, M. (2018). *Evolution and selection of quantitative traits*. Oxford University Press. https://doi.org/10.1093/oso/9780198830870.001.0001

Warwell, M. V., & Shaw, R. G. (2018). Phenotypic selection on growth rhythm in whitebark pine under climatic conditions warmer than seed origins. *Journal of Evolutionary Biology*, 31(9), 1284-1299. https://doi.org/10.1111/jeb.13301

Warwell, M. V., & Shaw, R. G. (2019). Phenotypic selection on ponderosa pine seed and seedling traits in the field under three experimentally manipulated drought treatments. *Evolutionary Applications*, 12, 159-174.

Westergren, M., Archambeau, J., Bajc, M., Damjanić, R., Theraroz, A., Kraigher, H., Oddou-Muratorio, S., & González-Martínez, S. C. (2023). Low but significant evolutionary potential for growth, phenology and reproduction traits in European beech. *Data Dryad*. [dataset]. https://doi.org/10.5061/dryad.x0k6djhrn

Westergren, M., Kavaliauskas, D., Alizoti, P., Bajc, M., Aravanopoulos, F. A., Bozic, G., Damjanic, R., Dovc, N., Finzgar, D., Fussi, B., Kiourtsis, F., & Kraigher, H. (2020). Guidelines for genetic monitoring of European beech (Fagus sylvatica L.). In F. Aravanopoulos, M. Westergren, B. Fussi, D. Kavaliauskas, P. Alizoti, F. Kiourtsis, & H. Kraigher (Eds.), *Manual for forest genetic monitoring* (pp. 179-194). Slovenian Forestry Institute. https://doi.org/10.20315/SFS.167

Wheelwright, N. T., Keller, L. F., & Postma, E. (2014). The effect of trait type and strength of selection on heritability and evolvability in an Island bird population. *Evolution*, 68, 3325-3336. https://doi.org/10.1111/evo.12499

Wickert, K. L., O'Neal, E. S., Davis, D. D., & Kasson, M. T. (2017). Seed production, viability, and reproductive limits of the invasive *Ailanthus altissima* (Tree-of-Heaven) within invaded environments. *Forests*, 8(7), 226. https://doi.org/10.3390/f8070226

Zenni, R. D., & Hoban, S. M. (2015). Loci under selection during multiple range expansions of an invasive plant are mostly population specific, but patterns are associated with climate. *Molecular Ecology*, 24(13), 3360-3371. https://doi.org/10.1111/mec.13234

Zenni, R. D., Lacerda da Cunha, W., & Sena, G. (2016). Rapid increase in growth and productivity can aid invasions by a non-native tree. *AoB Plants*, 8, plw048. https://doi.org/10.1093/aobpla/plw048

## SUPPORTING INFORMATION

Additional supporting information can be found online in the Supporting Information section at the end of this article.

**How to cite this article:** Westergren, M., Archambeau, J., Bajc, M., Damjanić, R., Theraroz, A., Kraigher, H., Oddou-Muratorio, S., & González-Martínez, S. C. (2023). Low but significant evolutionary potential for growth, phenology and reproduction traits in European beech. *Molecular Ecology*, 00, 1-20. https://doi.org/10.1111/mec.17196
Richardson, A. D., Bailey, A. S., Denny, E. G., Martin, C. W., & O'Keefe, J. (2006). Phenology of a northern hardwood forest canopy. *Global Change Biology*, 12(7), 1174-1188. https://doi.org/10.1111/j.1365-2486.2006.01164.x

Rousset, F. (2008). *genepop'007: A complete re-implementation of the genepop software for Windows and Linux*. *Molecular Ecology Resources*, 8, 103-106.

Schlichting, C. D. (1989). Phenotypic integration and environmental change. What are the consequences of differential phenotypic plasticity of traits? *Bioscience*, 39(7), 460-464. https://doi.org/10.2307/1311138

Schulter, D., Price, T. D., & Rowe, L. (1991). Conflicting selection pressures and life history trade-offs. *Proceedings of the Royal Society of London. Series B: Biological Sciences*, 246(1315), 11-17. https://doi.org/10.1098/rspb.1991.0118

Scotti-Saintagne, C., Bodénès, C., Barreneche, T., Bertocchi, E., Plomion, C., & Kremer, A. (2004). Detection of quantitative trait loci controlling bud burst and height growth in *Quercus robur* L. *Theoretical and Applied Genetics*, 109(8), 1648-1659. https://doi.org/10.1007/s00122-004-1789-3

Sedlacek, J., Cortés, A. J., Wheeler, J., Bossdorf, O., Hoch, G., Klápště, J., Lerx, C., Rixen, C., Wipf, S., Karrenberg, S., & Kleunen, M. (2016). Evolutionary potential in the Alpine: Trait heritabilities and performance variation of the dwarf willow *Salix herbacea* from different elevations and microhabitats. *Ecology and Evolution*, 6(12), 3940-3952. https://doi.org/10.1002/ece3.2171

Šercelj, A. (1996). Začetki in razvoj gozdv v Sloveniji (p. 142). Slovenska Akademija Znanosti in Umetnosti.

Sgrò, C. M., & Hoffmann, A. A. (1998). Heritable variation for fecundity in field-collected Drosophila melanogaster and their offspring reared under different environmental temperatures. *Evolution*, 52(1), 134-143. https://doi.org/10.1111/j.1558-5646.1998.tb05146.x

Siepielski, A. M., DiBattista, J. D., & Carlson, S. M. (2009). It's about time: The temporal dynamics of phenotypic selection in the wild. *Ecology Letters*, 12(11), 1261-1276. https://doi.org/10.1111/j.1461-0248.2009.01381.x

Staaf, H., & Stjernquist, I. (1986). Seasonal dynamics, especially autumnal retranslocation, of nitrogen and phosphorus in foliage of dominant and suppressed trees of beech, *Fagus sylvatica*. *Scandinavian Journal of Forest Research*, 1(1-4), 333-342. https://doi.org/10.1080/02827588609382425

Thomas, S. C. (2011). Age-related changes in tree growth and functional biology: The role of reproduction. In F. C. Meinzer, B. Lachenbruch, & T. E. Dawson (Eds.), *Size- and age-related changes in tree structure and function* (Vol. 4, pp. 33-64). Springer. https://doi.org/10.1007/978-94-007-1242-3_2

Tonnabel, J., David, P., Klein, E. K., & Pannell, J. R. (2019). Sex-specific selection on plant architecture through "budget" and "direct" effects in experimental populations of the wind-pollinated herb, *Mercurialis annua*. *Evolution*, 73(5), 897-912. https://doi.org/10.1111/evo.13714

Toomey, M., Friedl, M. A., Frolking, S., Hufkens, K., Klosterman, S., Sonnentag, O., Baldocchi, D. D., Bernacchi, C. J., Biraud, S. C., Bohrer, G., Brzostek, E., Burns, S. P., Coursolle, C., Hollinger, D. Y., Margolis, H. A., McCaughey, H., Monson, R. K., Munger, J. W., Pallardy, S., ... Richardson, A. D. (2015). Greenness indices from digital cameras predict the timing and seasonal dynamics of canopy-scale photosynthesis. *Ecological Applications*, 25(1), 99-115. https://doi.org/10.1890/14-0005.1

Visscher, P. M., Hill, W. G., & Wray, N. R. (2008). Heritability in the genomics era—Concepts and misconceptions. *Nature Reviews Genetics*, 9(4), 255-266. https://doi.org/10.1038/nrg2322

Vitasse, Y. (2013). Ontogenic changes rather than difference in temperature cause understory trees to leaf out earlier. *New Phytologist*, 198(1), 149-155. https://doi.org/10.1111/nph.12130

Vitasse, Y., Delzon, S., Bresson, C. C., Michalet, R., & Kremer, A. (2009). Altitudinal differentiation in growth and phenology among populations of temperate-zone tree species growing in a common garden. *Canadian Journal of Forest Research*, 39(7), 1259-1269. https://doi.org/10.1139/X09-054

Walsh, B., & Lynch, M. (2018). *Evolution and selection of quantitative traits*. Oxford University Press. https://doi.org/10.1093/oso/9780198830870.001.0001

Warwell, M. V., & Shaw, R. G. (2018). Phenotypic selection on growth rhythm in whitebark pine under climatic conditions warmer than seed origins. *Journal of Evolutionary Biology*, 31(9), 1284-1299. https://doi.org/10.1111/jeb.13301

Warwell, M. V., & Shaw, R. G. (2019). Phenotypic selection on ponderosa pine seed and seedling traits in the field under three experimentally manipulated drought treatments. *Evolutionary Applications*, 12, 159-174.

Westergren, M., Archambeau, J., Bajc, M., Damjanić, R., Theraroz, A., Kraigher, H., Oddou-Muratorio, S., & González-Martínez, S. C. (2023). Low but significant evolutionary potential for growth, phenology and reproduction traits in European beech. *Data Dryad*. [dataset]. https://doi.org/10.5061/dryad.x0k6djhrn

Westergren, M., Kavaliauskas, D., Alizoti, P., Bajc, M., Aravanopoulos, F. A., Bozic, G., Damjanic, R., Dovc, N., Finzgar, D., Fussi, B., Kiourtsis, F., & Kraigher, H. (2020). Guidelines for genetic monitoring of European beech (Fagus sylvatica L.). In F. Aravanopoulos, M. Westergren, B. Fussi, D. Kavaliauskas, P. Alizoti, F. Kiourtsis, & H. Kraigher (Eds.), *Manual for forest genetic monitoring* (pp. 179-194). Slovenian Forestry Institute. https://doi.org/10.20315/SFS.167

Wheelwright, N. T., Keller, L. F., & Postma, E. (2014). The effect of trait type and strength of selection on heritability and evolvability in an Island bird population. *Evolution*, 68, 3325-3336. https://doi.org/10.1111/evo.12499

Wickert, K. L., O'Neal, E. S., Davis, D. D., & Kasson, M. T. (2017). Seed production, viability, and reproductive limits of the invasive *Ailanthus altissima* (Tree-of-Heaven) within invaded environments. *Forests*, 8(7), 226. https://doi.org/10.3390/f8070226

Zenni, R. D., & Hoban, S. M. (2015). Loci under selection during multiple range expansions of an invasive plant are mostly population specific, but patterns are associated with climate. *Molecular Ecology*, 24(13), 3360-3371. https://doi.org/10.1111/mec.13234

Zenni, R. D., Lacerda da Cunha, W., & Sena, G. (2016). Rapid increase in growth and productivity can aid invasions by a non-native tree. *AoB Plants*, 8, plw048. https://doi.org/10.1093/aobpla/plw048

## SUPPORTING INFORMATION

Additional supporting information can be found online in the Supporting Information section at the end of this article.

**How to cite this article:** Westergren, M., Archambeau, J., Bajc, M., Damjanić, R., Theraroz, A., Kraigher, H., Oddou-Muratorio, S., & González-Martínez, S. C. (2023). Low but significant evolutionary potential for growth, phenology and reproduction traits in European beech. *Molecular Ecology*, 00, 1-20. https://doi.org/10.1111/mec.17196