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2. Adaptive responses of animals to climate change are most likely insufficient

Author

Radchuk, Viktoriia, Reed, Thomas, Teplitsky, Céline, van de Pol, Martijn, Charmantier, Anne, Hassall, Christopher, Adamík, Peter, Adriaensen, Frank, Ahola, Markus P, Arcese, Peter, Miguel Avilés, Jesús, Balbontin, Javier, Berg, Karl S, Borras, Antoni, Burthe, Sarah, Clobert, Jean, Dehnhard, Nina, de Lope, Florentino, Dhondt, André A, Dingemanse, Niels J, Doi, Hideyuki, Eeva, Tapio, Fickel, Joerns, Filella, Iolanda, Fossøy, Frode, Goodenough, Anne E, Hall, Stephen JG, Hansson, Bengt, Harris, Michael, Hasselquist, Dennis, Hickler, Thomas, Joshi, Jasmin, Kharouba, Heather, Martínez, Juan Gabriel, Mihoub, Jean-Baptiste, Mills, James A, Molina-Morales, Mercedes, Moksnes, Arne, Ozgul, Arpat, Parejo, Deseada, Pilard, Philippe, Poisbleau, Maud, Rousset, Francois, Rödel, Mark-Oliver, Scott, David, Senar, Juan Carlos, Stefanescu, Constanti, Stokke, Bård G, Kusano, Tamotsu, Tarka, Maja, Tarwater, Corey E, Thonicke, Kirsten, Thorley, Jack, Wilting, Andreas, Tryjanowski, Piotr, Merilä, Juha, Sheldon, Ben C, Pape Møller, Anders, Matthysen, Erik, Janzen, Fredric, Dobson, F Stephen, Visser, Marcel E, Beissinger, Steven R, Courtiol, Alexandre, and Kramer-Schadt, Stephanie

Subjects
Climate Change Impacts and Adaptation, Biological Sciences, Environmental Sciences, Evolutionary Biology, Climate Action, Acclimatization, Animals, Birds, Climate Change, Phenotype, Selection, Genetic, Time Factors
Abstract

Biological responses to climate change have been widely documented across taxa and regions, but it remains unclear whether species are maintaining a good match between phenotype and environment, i.e. whether observed trait changes are adaptive. Here we reviewed 10,090 abstracts and extracted data from 71 studies reported in 58 relevant publications, to assess quantitatively whether phenotypic trait changes associated with climate change are adaptive in animals. A meta-analysis focussing on birds, the taxon best represented in our dataset, suggests that global warming has not systematically affected morphological traits, but has advanced phenological traits. We demonstrate that these advances are adaptive for some species, but imperfect as evidenced by the observed consistent selection for earlier timing. Application of a theoretical model indicates that the evolutionary load imposed by incomplete adaptive responses to ongoing climate change may already be threatening the persistence of species.

Published
2019

4. Effects of interspecific coexistence on laying date and clutch size in two closely related species of hole-nesting birds

Author

Møller, Anders Pape, Adriaensen, Frank, Balbontín, Javier, Dhondt, André A., Remeš, Vladimir, Biard, Clotilde, Camprodon, Jordi, Cichoń, Mariusz, Gustafsson, Lars, Doligez, Blandine, Dubiec, Anna, Eens, Marcel, Eeva, Tapio, Goodenough, Anne E., Gosler, Andrew G., Heeb, Philipp, Hinsley, Shelley A., Jacob, Staffan, Juškaitis, Rimvydas, Laaksonen, Toni, Leclercq, Bernard, Massa, Bruno, Mazgajski, Tomasz D., Nager, Ruedi G., Nilsson, Jan-Åke, Nilsson, Sven G., Norte, Ana C., Lambrechts, Marcel M., Pinxten, Rianne, Robles, Hugo, Solonen, Tapio, Sorace, Alberto, and van Noordwijk, Arie J

Published
2018

18. Interaction of climate change with effects of conspecific and heterospecific density on reproduction

Author

Møller, Anders Pape, Balbontín, Javier, Dhondt, André A., Adriaensen, Frank, Artemyev, Alexandr, Bańbura, Jerzy, Barba, Emilio, Biard, Clotilde, Blondel, Jacques, Bouvier, Jean-Charles, Camprodon, Jordi, Cecere, Francesco, Charter, Motti, Cichoń, Mariusz, Cusimano, Camillo, Dubiec, Anna, Eens, Marcel, Eeva, Tapio, Ferns, Peter N., Forsman, Jukka T., Goldshtein, Aya, Goodenough, Anne E., Gosler, Andrew G., Gustafsson, Lars, Harnist, Iga, Hartley, Ian R., Heeb, Philipp, Hinsley, Shelley A., Jacob, Staffan, Järvinen, Antero, Juškaitis, Rimvydas, Korpimäki, Erkki, Krams, Indrikis, Laaksonen, Toni, Leclercq, Bernard, Lehikoinen, Esa, Loukola, Olli, Mainwaring, Mark C., Mänd, Raivo, Massa, Bruno, Matthysen, Erik, Mazgajski, Tomasz D., Merino, Santiago, Mitrus, Cezary, Mönkkönen, Mikko, Nager, Ruedi G., Nilsson, Jan-Åke, Nilsson, Sven G., Norte, Ana C., von Numers, Mikael, Orell, Markku, Pimentel, Carla S., Pinxten, Rianne, Priedniece, Ilze, Remeš, Vladimir, Richner, Heinz, Robles Díez, Hugo, Rytkönen, Seppo, Senar, Juan Carlos, Seppänen, Janne T., da Silva, Luís P., Slagsvold, Tore, Solonen, Tapio, Sorace, Alberto, Stenning, Martyn J., Török, János, Tryjanowski, Piotr, van Noordwijk, Arie J., Walankiewicz, Wiesław, Lambrechts, Marcel M., Moller, Anders Pape, Balbontin, Javier, Dhondt, Andre A., Banbura, Jerzy, Cichon, Mariusz, Jarvinen, Antero, Juskaitis, Rimvydas, Korpimaki, Erkki, Mand, Raivo, Monkkonen, Mikko, Nilsson, Jan-ake, Remes, Vladimir, Rytkonen, Seppo, Seppanen, Janne T., da Silva, Luis P., Torok, Janos, Walankiewicz, Wieslaw, Helsinki Institute of Sustainability Science (HELSUS), Organismal and Evolutionary Biology Research Programme, Kilpisjärvi Biological Station, Ecologie Systématique et Evolution (ESE), AgroParisTech-Université Paris-Saclay-Centre National de la Recherche Scientifique (CNRS), Universidad de Sevilla / University of Sevilla, Cornell University [New York], University of Antwerp (UA), Russian Academy of Sciences [Moscow] (RAS), University of Lódź, Cavanilles Institute of Biodiversity and Evolutionary Biology, University of Valencia, Institut d'écologie et des sciences de l'environnement de Paris (iEES Paris ), Institut de Recherche pour le Développement (IRD)-Sorbonne Université (SU)-Université Paris-Est Créteil Val-de-Marne - Paris 12 (UPEC UP12)-Centre National de la Recherche Scientifique (CNRS)-Institut National de Recherche pour l’Agriculture, l’Alimentation et l’Environnement (INRAE), Centre d’Ecologie Fonctionnelle et Evolutive (CEFE), Université Paul-Valéry - Montpellier 3 (UPVM)-École Pratique des Hautes Études (EPHE), Université Paris sciences et lettres (PSL)-Université Paris sciences et lettres (PSL)-Université de Montpellier (UM)-Centre National de la Recherche Scientifique (CNRS)-Institut de Recherche pour le Développement (IRD [France-Sud])-Institut National de Recherche pour l’Agriculture, l’Alimentation et l’Environnement (INRAE)-Institut Agro - Montpellier SupAgro, Institut national d'enseignement supérieur pour l'agriculture, l'alimentation et l'environnement (Institut Agro)-Institut national d'enseignement supérieur pour l'agriculture, l'alimentation et l'environnement (Institut Agro), Unité de recherche Plantes et Systèmes de Culture Horticoles (PSH), Institut National de Recherche pour l’Agriculture, l’Alimentation et l’Environnement (INRAE), Centre de Ciència i Tecnologia Forestal de Catalunya (CTFC), University of Haifa [Haifa], Uniwersytet Jagielloński w Krakowie = Jagiellonian University (UJ), Stazione Ornitologica di Palermo, Polish Academy of Sciences (PAN), Université de Lyon, University of Turku, School of Biosciences [Cardiff], Cardiff University, University of Oulu, Tel Aviv University (TAU), University of Gloucestershire (Cheltenham, GB), Edward Grey Institute, Department of Zoology, University of Oxford, Uppsala University, Lancaster University, Evolution et Diversité Biologique (EDB), Institut de Recherche pour le Développement (IRD)-Université Toulouse III - Paul Sabatier (UT3), Université de Toulouse (UT)-Université de Toulouse (UT)-Centre National de la Recherche Scientifique (CNRS), Centre for Ecology and Hydrology [Wallingford] (CEH), Natural Environment Research Council (NERC), Station d'écologie théorique et expérimentale (SETE), Université Toulouse III - Paul Sabatier (UT3), Université de Toulouse (UT)-Université de Toulouse (UT)-Observatoire Midi-Pyrénées (OMP), Université de Toulouse (UT)-Université de Toulouse (UT)-Institut national des sciences de l'Univers (INSU - CNRS)-Centre National d'Études Spatiales [Toulouse] (CNES)-Centre National de la Recherche Scientifique (CNRS)-Météo-France -Institut de Recherche pour le Développement (IRD)-Institut national des sciences de l'Univers (INSU - CNRS)-Centre National d'Études Spatiales [Toulouse] (CNES)-Centre National de la Recherche Scientifique (CNRS)-Météo-France -Centre National de la Recherche Scientifique (CNRS), Helsingin yliopisto = Helsingfors universitet = University of Helsinki, Nature Research Centre, Institute of Ecology, Akademijos str. 2, LT-08412, Vilnius, Lithuania., University of Tartu, University of Montana, Consejo Superior de Investigaciones Científicas [Madrid] (CSIC), Uniwersytet Wroclawski, University of Jyväskylä (JYU), University of Glasgow, Lund University [Lund], Marine and Environmental Sciences Centre (MARE UC), Universidade de Coimbra [Coimbra], Åbo Akademi University [Turku], Université de Lisbonne, Latvian Fund for Nature, Palacky University Olomouc, University of Bern, University of A Coruña (UDC), Museu de Ciències Naturals de Barcelona, Universidade do Porto = University of Porto, University of Oslo (UiO), Luontotutkimus Solonen Oy, SROPU, University of Sussex, Eötvös Loránd University (ELTE), Poznan University of Life Sciences (Uniwersytet Przyrodniczy w Poznaniu) (PULS), Netherlands Institute of Ecology - NIOO-KNAW (NETHERLANDS), IB KRC RAS no. 0218-2019-0080, Academy of Finland (project 265859), Ministry of Economy and Competitivity, Spanish Research Council: research project CGL-2016-79568-C3-3-P, ANR-10-LABX-0041,TULIP,Towards a Unified theory of biotic Interactions: the roLe of environmental(2010), Animal Ecology (AnE), Ministry of Education Key Laboratory for Biodiversity Science and Ecological Engineering, College of Life Sciences, Beijing Normal University, Beijing 100875, Universidad de Sevilla, Cavanilles Institute of Biodiversity and Evolutionary Biology, Terrestrial Vertebrates Group, Institut de Recherche pour le Développement (IRD)-Sorbonne Université (SU)-Université Paris-Est Créteil Val-de-Marne - Paris 12 (UPEC UP12)-Centre National de la Recherche Scientifique (CNRS)-Université de Paris (UP)-Institut National de Recherche pour l’Agriculture, l’Alimentation et l’Environnement (INRAE), Université Paul-Valéry - Montpellier 3 (UPVM)-Centre international d'études supérieures en sciences agronomiques (Montpellier SupAgro)-École pratique des hautes études (EPHE), Université Paris sciences et lettres (PSL)-Université Paris sciences et lettres (PSL)-Université de Montpellier (UM)-Centre National de la Recherche Scientifique (CNRS)-Institut de Recherche pour le Développement (IRD [France-Sud])-Institut national d’études supérieures agronomiques de Montpellier (Montpellier SupAgro), Institut national d'enseignement supérieur pour l'agriculture, l'alimentation et l'environnement (Institut Agro)-Institut national d'enseignement supérieur pour l'agriculture, l'alimentation et l'environnement (Institut Agro)-Institut National de Recherche pour l’Agriculture, l’Alimentation et l’Environnement (INRAE), Tel Aviv University [Tel Aviv], University of Oxford [Oxford], Centre National de la Recherche Scientifique (CNRS)-Institut de Recherche pour le Développement (IRD)-Université Toulouse III - Paul Sabatier (UT3), Université Fédérale Toulouse Midi-Pyrénées-Université Fédérale Toulouse Midi-Pyrénées, Université Fédérale Toulouse Midi-Pyrénées-Université Fédérale Toulouse Midi-Pyrénées-Centre National de la Recherche Scientifique (CNRS)-Observatoire Midi-Pyrénées (OMP), Météo France-Centre National d'Études Spatiales [Toulouse] (CNES)-Université Fédérale Toulouse Midi-Pyrénées-Centre National de la Recherche Scientifique (CNRS)-Institut de Recherche pour le Développement (IRD)-Météo France-Centre National d'Études Spatiales [Toulouse] (CNES)-Centre National de la Recherche Scientifique (CNRS)-Institut de Recherche pour le Développement (IRD), University of Helsinki, Universidade do Porto, Russian Academy of Sciences, Academy of Finland, Ministerio de Economía y Competitividad (España), Consejo Superior de Investigaciones Científicas (España), Université Paul-Valéry - Montpellier 3 (UPVM)-École pratique des hautes études (EPHE), Université Fédérale Toulouse Midi-Pyrénées-Université Fédérale Toulouse Midi-Pyrénées-Centre National de la Recherche Scientifique (CNRS), Université Fédérale Toulouse Midi-Pyrénées-Université Fédérale Toulouse Midi-Pyrénées-Observatoire Midi-Pyrénées (OMP), and Université Fédérale Toulouse Midi-Pyrénées-Université Fédérale Toulouse Midi-Pyrénées-Institut national des sciences de l'Univers (INSU - CNRS)-Centre National d'Études Spatiales [Toulouse] (CNES)-Centre National de la Recherche Scientifique (CNRS)-Météo-France -Institut de Recherche pour le Développement (IRD)-Institut national des sciences de l'Univers (INSU - CNRS)-Centre National d'Études Spatiales [Toulouse] (CNES)-Centre National de la Recherche Scientifique (CNRS)-Météo-France -Centre National de la Recherche Scientifique (CNRS)

Subjects
BREEDING SUCCESS, 0106 biological sciences, Avian clutch size, clutch size, Q1, 01 natural sciences, DEPENDENCE, Parus major, sinitiainen, POPULATION, QL_671, [SDV.EE]Life Sciences [q-bio]/Ecology, environment, education.field_of_study, GE, biology, Ecology, Blue tit, tiaiset, Cyanistes, blue tit, Plan_S-Compliant_NO, talitiainen, Spatial heterogeneity, Chemistry, great tit, international, 1181 Ecology, evolutionary biology, lämpötila, laying date, CLUTCH-SIZE, intraspecific competition, Population, HABITAT HETEROGENEITY, PARUS-MAJOR, 010603 evolutionary biology, Ecology and Environment, Intraspecific competition, temperature anomaly, muninta, QH301, BLUE, Cyanistes caeruleus, education, Biology, Ecology, Evolution, Behavior and Systematics, Interspecioc competition, FICEDULA, Parus, QL, pesintä, lisääntymiskäyttäytyminen, BIRDS, 010604 marine biology & hydrobiology, interspecific competition, Ficedula, Interspecific competition, ilmastonmuutokset, biology.organism_classification, 13. Climate action, GREAT TITS
Abstract

We studied the relationship between temperature and the coexistence of great tit Parus major and blue tit Cyanistes caeruleus, breeding in 75 study plots across Europe and North Africa. We expected an advance in laying date and a reduction in clutch size during warmer springs as a general response to climate warming and a delay in laying date and a reduction in clutch size during warmer winters due to density-dependent effects. As expected, as spring temperature increases laying date advances and as winter temperature increases clutch size is reduced in both species. Density of great tit affected the relationship between winter temperature and laying date in great and blue tit. Specifically, as density of great tit increased and temperature in winter increased both species started to reproduce later. Density of blue tit affected the relationship between spring temperature and blue and great tit laying date. Thus, both species start to reproduce earlier with increasing spring temperature as density of blue tit increases, which was not an expected outcome, since we expected that increasing spring temperature should advance laying date, while increasing density should delay it cancelling each other out. Climate warming and its interaction with density affects clutch size of great tits but not of blue tits. As predicted, great tit clutch size is reduced more with density of blue tits as temperature in winter increases. The relationship between spring temperature and density on clutch size of great tits depends on whether the increase is in density of great tit or blue tit. Therefore, an increase in temperature negatively affected the coexistence of blue and great tits differently in both species. Thus, blue tit clutch size was unaffected by the interaction effect of density with temperature, while great tit clutch size was affected in multiple ways by these interactions terms., A. Artemyev acknowledges funding by IB KRC RAS no. 0218-2019-0080 and T. Eeva acknowledges funding by the Academy of Finland (project 265859). This study was funded by research project CGL-2016-79568-C3-3-P (to J. C. Senar), from the Ministry of Economy and Competitivity, Spanish Research Council.

Published
2020
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21. Assessing Multivariate Constraints to Evolution across Ten Long-Term Avian Studies

Author

Teplitsky, Celine, Tarka, Maja, Moller, Anders P., Nakagawa, Shinichi, Balbontin, Javier, Burke, Terry A., Doutrelant, Claire, Gregoire, Arnaud, Hansson, Bengt, Hasselquist, Dennis, Gustafsson, Lars, de Lope, Florentino, Marzal, Alfonso, Mills, James A., Wheelwright, Nathaniel T., Yarrall, John W., Charmantier, Anne, Teplitsky, Celine, Tarka, Maja, Moller, Anders P., Nakagawa, Shinichi, Balbontin, Javier, Burke, Terry A., Doutrelant, Claire, Gregoire, Arnaud, Hansson, Bengt, Hasselquist, Dennis, Gustafsson, Lars, de Lope, Florentino, Marzal, Alfonso, Mills, James A., Wheelwright, Nathaniel T., Yarrall, John W., and Charmantier, Anne

Abstract

Background: In a rapidly changing world, it is of fundamental importance to understand processes constraining or facilitating adaptation through microevolution. As different traits of an organism covary, genetic correlations are expected to affect evolutionary trajectories. However, only limited empirical data are available. Methodology/Principal Findings: We investigate the extent to which multivariate constraints affect the rate of adaptation, focusing on four morphological traits often shown to harbour large amounts of genetic variance and considered to be subject to limited evolutionary constraints. Our data set includes unique long-term data for seven bird species and a total of 10 populations. We estimate population-specific matrices of genetic correlations and multivariate selection coefficients to predict evolutionary responses to selection. Using Bayesian methods that facilitate the propagation of errors in estimates, we compare (1) the rate of adaptation based on predicted response to selection when including genetic correlations with predictions from models where these genetic correlations were set to zero and (2) the multivariate evolvability in the direction of current selection to the average evolvability in random directions of the phenotypic space. We show that genetic correlations on average decrease the predicted rate of adaptation by 28%. Multivariate evolvability in the direction of current selection was systematically lower than average evolvability in random directions of space. These significant reductions in the rate of adaptation and reduced evolvability were due to a general nonalignment of selection and genetic variance, notably orthogonality of directional selection with the size axis along which most (60%) of the genetic variance is found. Conclusions: These results suggest that genetic correlations can impose significant constraints on the evolution of avian morphology in wild populations. This could have important impacts on evolutiona

Published
2014
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25. Adaptive responses of animals to climate change are most likely insufficient

Author

Radchuk, Viktoriia, Reed, Thomas, Teplitsky, Céline, Van De Pol, Martijn, Charmantier, Anne, Hassall, Christopher, Adamík, Peter, Adriaensen, Frank, Ahola, Markus P., Arcese, Peter, Miguel Avilés, Jesús, Balbontin, Javier, Berg, Karl S., Borras, Antoni, Burthe, Sarah, Clobert, Jean, Dehnhard, Nina, De Lope, Florentino, Dhondt, André A., Dingemanse, Niels J., Doi, Hideyuki, Eeva, Tapio, Fickel, Joerns, Filella, Iolanda, Fossøy, Frode, Goodenough, Anne E., Hall, Stephen J. G., Hansson, Bengt, Harris, Michael, Hasselquist, Dennis, Hickler, Thomas, Joshi, Jasmin, Kharouba, Heather, Martínez, Juan Gabriel, Mihoub, Jean-Baptiste, Mills, James A., Molina-Morales, Mercedes, Moksnes, Arne, Ozgul, Arpat, Parejo, Deseada, Pilard, Philippe, Poisbleau, Maud, Rousset, Francois, Rödel, Mark-Oliver, Scott, David, Senar, Juan Carlos, Stefanescu, Constanti, Stokke, Bård G., Kusano, Tamotsu, Tarka, Maja, Tarwater, Corey E., Thonicke, Kirsten, Thorley, Jack, Wilting, Andreas, Tryjanowski, Piotr, Merilä, Juha, Sheldon, Ben C., Pape Møller, Anders, Matthysen, Erik, Janzen, Fredric, Dobson, F. Stephen, Visser, Marcel E., Beissinger, Steven R., Courtiol, Alexandre, and Kramer-Schadt, Stephanie

Subjects
141, 631/158/857, 13. Climate action, 704/158/2165, article, sense organs, 15. Life on land, skin and connective tissue diseases, 704/158/672
Abstract

Biological responses to climate change have been widely documented across taxa and regions, but it remains unclear whether species are maintaining a good match between phenotype and environment, i.e. whether observed trait changes are adaptive. Here we reviewed 10,090 abstracts and extracted data from 71 studies reported in 58 relevant publications, to assess quantitatively whether phenotypic trait changes associated with climate change are adaptive in animals. A meta-analysis focussing on birds, the taxon best represented in our dataset, suggests that global warming has not systematically affected morphological traits, but has advanced phenological traits. We demonstrate that these advances are adaptive for some species, but imperfect as evidenced by the observed consistent selection for earlier timing. Application of a theoretical model indicates that the evolutionary load imposed by incomplete adaptive responses to ongoing climate change may already be threatening the persistence of species.

26. Adaptive responses of animals to climate change are most likely insufficient

Author

Radchuk, Viktoriia, Reed, Thomas, Teplitsky, Céline, Van De Pol, Martijn, Charmantier, Anne, Hassall, Christopher, Adamík, Peter, Adriaensen, Frank, Ahola, Markus P, Arcese, Peter, Miguel Avilés, Jesús, Balbontin, Javier, Berg, Karl S, Borras, Antoni, Burthe, Sarah, Clobert, Jean, Dehnhard, Nina, De Lope, Florentino, Dhondt, André A, Dingemanse, Niels J, Doi, Hideyuki, Eeva, Tapio, Fickel, Joerns, Filella, Iolanda, Fossøy, Frode, Goodenough, Anne E, Hall, Stephen JG, Hansson, Bengt, Harris, Michael, Hasselquist, Dennis, Hickler, Thomas, Joshi, Jasmin, Kharouba, Heather, Martínez, Juan Gabriel, Mihoub, Jean-Baptiste, Mills, James A, Molina-Morales, Mercedes, Moksnes, Arne, Ozgul, Arpat, Parejo, Deseada, Pilard, Philippe, Poisbleau, Maud, Rousset, Francois, Rödel, Mark-Oliver, Scott, David, Senar, Juan Carlos, Stefanescu, Constanti, Stokke, Bård G, Kusano, Tamotsu, Tarka, Maja, Tarwater, Corey E, Thonicke, Kirsten, Thorley, Jack, Wilting, Andreas, Tryjanowski, Piotr, Merilä, Juha, Sheldon, Ben C, Pape Møller, Anders, Matthysen, Erik, Janzen, Fredric, Dobson, F Stephen, Visser, Marcel E, Beissinger, Steven R, Courtiol, Alexandre, and Kramer-Schadt, Stephanie

Subjects
Birds, Phenotype, Time Factors, 13. Climate action, Acclimatization, Climate Change, Animals, sense organs, Selection, Genetic, skin and connective tissue diseases
Abstract

Biological responses to climate change have been widely documented across taxa and regions, but it remains unclear whether species are maintaining a good match between phenotype and environment, i.e. whether observed trait changes are adaptive. Here we reviewed 10,090 abstracts and extracted data from 71 studies reported in 58 relevant publications, to assess quantitatively whether phenotypic trait changes associated with climate change are adaptive in animals. A meta-analysis focussing on birds, the taxon best represented in our dataset, suggests that global warming has not systematically affected morphological traits, but has advanced phenological traits. We demonstrate that these advances are adaptive for some species, but imperfect as evidenced by the observed consistent selection for earlier timing. Application of a theoretical model indicates that the evolutionary load imposed by incomplete adaptive responses to ongoing climate change may already be threatening the persistence of species.

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