Functional traits are organismal attributes that can respond to environmental cues, thereby providing important ecological functions. In addition, an organism's potential for adaptation is defined by the patterns of covariation among groups of functionally related traits. Whether an organism is evolutionarily constrained or has the potential for adaptation is based on the phenotypic integration or modularity of these traits. Here, we revisited leaf morphology in two European sympatric white oaks (Quercus petraea (Matt.) Liebl. and Quercus robur L.), sampling 2098 individuals, across much of their geographical distribution ranges. At the phenotypic level, leaf morphology traditionally encompasses discriminant attributes among different oak species. Here, we estimated in situ heritability, genetic correlation, and integration across such attributes. Also, we performed Selection Response Decomposition to test these traits for potential differences in oak species' evolutionary responses. Based on the uncovered functional units of traits (modules) in our study, the morphological module "leaf size gradient" was highlighted among functionally integrated traits. Equally, this module was defined in both oaks as being under "global regulation" in vegetative bud establishment and development. Lamina basal shape and intercalary veins' number were not, or, less integrated within the initially defined leaf functional unit, suggesting more than one module within the leaf traits' ensemble. Since these traits generally show the greatest species discriminatory power, they potentially underwent effective differential response to selection among oaks. Indeed, the selection of these traits could have driven the ecological preferences between the two sympatric oaks growing under different microclimates.

Département des sciences du bois et de la forêt Université Laval 1030 Avenue de la Médecine Québec QC G1V 0A6 Canada

Department of Forest and Conservation Sciences University of British Columbia 2424 Main Mall Vancouver BC V6T 1Z4 Canada

Department of Genetics and Physiology of Forest Trees Faculty of Forestry and Wood Sciences Czech University of Life Sciences Prague Kamýcká 129 165 21 Prague 6 Czechia

Department of Health and Environment AIT Austrian Institute of Technology Konrad Lorenz Straβe 24 3430 Tulln Austria

INRA UMR Biodiversité Gènes et Communautés 69 route d'Arcachon 33612 Cestas Cedex France

Pharmacognosy Department Faculty of Pharmacy Alexandria University Alexandria Egypt

Scion 49 Sala Street Whakarewarewa Rotorua 3010 New Zealand

University of Bordeaux UMR 1202 Biodiversité Gènes et Communautés F 33400 Talence France

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