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      Differences in spatial versus temporal reaction norms for spring and autumn phenological events

      research-article
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      Proceedings of the National Academy of Sciences of the United States of America
      National Academy of Sciences
      chilling, climate change, heating, phenology, plasticity

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          Significance

          To do the right thing at the right time, organisms need to glean cues from their environment. How they respond can then be described by reaction norms, i.e., by the relationship between the phenotype expressed (the phenology of an event) and the environment (the date when a given number of degree-days are achieved). We use information on 178 phenological events across the former Soviet Union. We found the timing of events to differ more between sites in spring and less in autumn. These patterns of local adaptation translate to a massive imprint on nature’s calendar: geographic variation in phenology is more pronounced in spring and less pronounced in autumn than if organisms were to respond equally everywhere.

          Abstract

          For species to stay temporally tuned to their environment, they use cues such as the accumulation of degree-days. The relationships between the timing of a phenological event in a population and its environmental cue can be described by a population-level reaction norm. Variation in reaction norms along environmental gradients may either intensify the environmental effects on timing (cogradient variation) or attenuate the effects (countergradient variation). To resolve spatial and seasonal variation in species’ response, we use a unique dataset of 91 taxa and 178 phenological events observed across a network of 472 monitoring sites, spread across the nations of the former Soviet Union. We show that compared to local rates of advancement of phenological events with the advancement of temperature-related cues (i.e., variation within site over years), spatial variation in reaction norms tend to accentuate responses in spring (cogradient variation) and attenuate them in autumn (countergradient variation). As a result, among-population variation in the timing of events is greater in spring and less in autumn than if all populations followed the same reaction norm regardless of location. Despite such signs of local adaptation, overall phenotypic plasticity was not sufficient for phenological events to keep exact pace with their cues—the earlier the year, the more did the timing of the phenological event lag behind the timing of the cue. Overall, these patterns suggest that differences in the spatial versus temporal reaction norms will affect species’ response to climate change in opposite ways in spring and autumn.

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          Most cited references49

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          Regularization Paths for Generalized Linear Models via Coordinate Descent

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            Regularization and variable selection via the elastic net

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              Ecological responses to recent climate change.

              There is now ample evidence of the ecological impacts of recent climate change, from polar terrestrial to tropical marine environments. The responses of both flora and fauna span an array of ecosystems and organizational hierarchies, from the species to the community levels. Despite continued uncertainty as to community and ecosystem trajectories under global change, our review exposes a coherent pattern of ecological change across systems. Although we are only at an early stage in the projected trends of global warming, ecological responses to recent climate change are already clearly visible.
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                Author and article information

                Journal
                Proc Natl Acad Sci U S A
                Proc Natl Acad Sci U S A
                pnas
                pnas
                PNAS
                Proceedings of the National Academy of Sciences of the United States of America
                National Academy of Sciences
                0027-8424
                1091-6490
                8 December 2020
                23 November 2020
                23 November 2020
                : 117
                : 49
                : 31249-31258
                Affiliations
                [1] aResearch Unit of Biodiversity, Oviedo University , 33600 Mieres, Spain;
                [2] bDepartment of Ecology, Swedish University of Agricultural Sciences , SE-75007 Uppsala, Sweden;
                [3] cUniversity of Helsinki , 00014 Helsinki, Finland;
                [4] dEarthCape , 00790 Helsinki, Finland;
                [5] eUniversity of Maryland , College Park, MD 20742;
                [6] fNational Park Orlovskoe Polesie , 303943, Orel Region, Hotynetskiy District, Zhuderskiy Village, Russian Federation;
                [7] gInstitute of Botany, Academy of Sciences of the Republic of Uzbekistan , 100053, Tashkent, 232 V, Uzbekistan;
                [8] hKostomuksha Nature Reserve , 186930, Karelia Republic, Kostomuksha, Russian Federation;
                [9] iAltai State Nature Biosphere Reserve , 649000, Altai Republic, Gorno-Altaysk, Russian Federation;
                [10] jKabardino-Balkarski Nature Reserve , 360000, Kabardino-Balkaria, Cherek District, Russian Federation;
                [11] kFederal State Enterprise Zapovednoe Podlemorye , 671623, Republic of Buryatia, Ust-Bargizin, Russian Federation;
                [12] lInstitute of General and Experimental Biology, Siberian Department, Russian Academy of Sciences , 670047, Republic of Buryatia, Ulan-Ude, Russian Federation;
                [13] mState Nature Reserve Stolby , 660006, Krasnoyarsk Region, Krasnoyarsk, Russian Federation;
                [14] nCarpathian Biosphere Reserve , 90600, Zakarpatska Oblast, Rakhiv, Ukraine;
                [15] oNizhne-Svirsky State Nature Reserve , 18700, Leningrad Region, Lodeinoe Pole, Russian Federation;
                [16] pState Nature Reserve Prisursky , 428034, Cheboksary, Russian Federation;
                [17] qZapovednoe Pribajkalje (Bajkalo-Lensky State Nature Reserve, Pribajkalsky National Park) , 664050, Irkutsk, Russian Federation;
                [18] rDarwin Nature Biosphere Reserve , 162723, Cherepovets District, Vologda Region, Borok, Russian Federation;
                [19] sVolzhsko-Kamsky National Nature Biosphere Rezerve , 422537, Tatarstan Republic, Zelenodolsk District, Russian Federation;
                [20] tFederal State Budget Institution National Park Shushenskiy Bor , 662710, Krasnoyarsk Region, Shushenskoe, Russian Federation;
                [21] uVoronezhsky Nature Biosphere Reserve , 394080, Centralnaja Usadba, Goszapovednik, Voronezh, Russian Federation;
                [22] vBaikalsky State Nature Biosphere Reserve , 671220, Buryatia Republic, Kabansky District, Tankhoy, Russian Federation;
                [23] wVisimsky Nature Biosphere Reserve , 624140, Kirovgrad, Russian Federation;
                [24] xKondinskie Lakes National Park, Named after L. F. Stashkevich , 628240, Hanty-Mansijsk District, City Sovietsky, Russian Federation;
                [25] yFederal State Budget Institution United Administration of the Kedrovaya Pad’ State Biosphere Nature Reserve and Leopard’s Land National Park , 690068, Primorskiy Kray, Vladivostok, Russian Federation;
                [26] zPechoro-Ilych State Nature Reserve , 169436, Komi Republic, Trinity-Pechora Region, Yaksha, Russian Federation;
                [27] aaA. N. Severtsov Institute of Ecology and Evolution , 119071, Moscow, Russian Federation;
                [28] bbKomsomolskiy Department, FGBU Zapovednoye Priamurye , 681000, Khabarovskyi Krai, Komsomolsk-on-Amur, Russian Federation;
                [29] ccTigirek State Nature Reserve , 656043, Barnaul, Russian Federation;
                [30] ddInstitute of Systematics and Ecology of Animals, Siberian Branch, Russian Academy of Sciences , 930091, Novosibirsk, Russian Federation;
                [31] eeState Nature Reserve Bolshaya Kokshaga , 424038, Mary El Republic, Yoshkar-Ola, Russian Federation;
                [32] ffInstitute of Plant and Animal Ecology, Ural Branch, Russian Academy of Sciences , 620100, Ekaterinburg, Russian Federation;
                [33] ggSikhote-Alin State Nature Biosphere Reserve, Named after K. G. Abramov , 692150, Primorsky Krai, Terney, Russian Federation;
                [34] hhFSBI Prioksko-Terrasniy State Reserve , 142200, Moscow Region, Serpukhov District, Danky, Russian Federation;
                [35] iiLomonosov Moscow State University , 119991, Moscow, Leninskie Gory, Russian Federation;
                [36] jjNational Park Meshchera , 601501, Vladimir Region, Gus-Hrustalnyi, Russian Federation;
                [37] kkSouth Urals Federal Research Center of Mineralogy and Geoecology, Ural Branch, Russian Academy of Sciences , Ilmeny State Reserve, 456317, Chelyabinskaya Oblast, Miass, Russian Federation;
                [38] llFGBU National Park Kenozersky , 163000, Arkhangelsk, Embankment of the Northern Dvina, Russian Federation;
                [39] mmFGBU GPZ Kologrivskij les im. M. G. Sinicina , 157440, Kostromskaja Oblast, Kologriv, Russian Federation;
                [40] nnAltai State University , 656049, Barnaul, Russian Federation;
                [41] ooPryazovskyi National Nature Park , 72312, Zaporiz’ka Oblast, Melitopol’, Ukraine;
                [42] ppState Nature Reserve Privolzhskaya Lesostep , 440031, Penza, Russian Federation;
                [43] qqKomarov Botanical Institute, Russian Academy of Sciences , 197376, Saint Petersburg, Russian Federation;
                [44] rrSary-Chelek State Nature Reserve , 715705, Dzalal-Abad Region, Aksu District, Arkyt Village, Kyrgyzstan;
                [45] ssInstitute for Evolutionary Ecology, National Academy of Sciences of Ukraine , 03143, Kiev, Ukraine;
                [46] ttFGBU State Nature Reserve Kuznetsk Alatau , 652888, Kemerovo Region, Mezhdurechensk, Russian Federation;
                [47] uuKerzhenskiy State Nature Biosphere Reserve , 603001, Nizhny Novgorod, Russian Federation;
                [48] vvBryansk Forest Nature Reserve , 242180, Bryansk Region, Suzemka District, Russian Federation;
                [49] wwPinezhsky State Nature Reserve , 164610, Arhangel Region, Pinezkiy District, Pinega, Russian Federation;
                [50] xxThe Central Chernozem State Biosphere Nature Reserve, Named after Professor V. V. Alyokhin , 305528 Kurskiy Region, Kurskiy District, Russian Federation;
                [51] yyTyumen State University , 625043, Tyumen, Russian Federation;
                [52] zzReserves of Taimyr , 666300, Norilsk, Russian Federation;
                [53] aaaChatkalski National Park , 100059, Toshkent, Uzbekistan;
                [54] bbbNational Park Ugra , 248007, Kaluga, Russian Federation;
                [55] cccKaniv Nature Reserve , 19000, Kaniv, Ukraine;
                [56] dddSmolenskoe Poozerje National Park , 216270, Smolensk Region, Demidovskiy District, Przhevalskoe, Russian Federation;
                [57] eeeFSBI Zeya State Nature Reserve , 676246, Zeya, Amurskaya Oblast, Russian Federation;
                [58] fffPolistovsky State Nature Reserve , 182840, Pskov Region, Bezhanitsy District, Russian Federation;
                [59] gggUral State Pedagogical University , 620017, Yekaterinburg, Russian Federation;
                [60] hhhInstitute of Mathematical Problems of Biology, Branch of the Keldysh Institute of Applied Mathematics, Russian Academy of Sciences , 142290, Moscow Region, Pushchino, Russian Federation;
                [61] iiiKronotsky Federal Nature Biosphere Reserve , 684000, Kamchatka Region, Yelizovo, Russian Federation;
                [62] jjjZhiguli Nature Reserve , 445362, Samara Region, P. Bakhilova Polyana, Russian Federation;
                [63] kkkInstitute for Ecology and Geography, Siberian Federal University , 660041, Krasnoyarsk, Russian Federation;
                [64] lllCentral Forest State Nature Biosphere Reserve , 172521, Tver Region, Nelidovo District, Zapovedniy Village, Russian Federation;
                [65] mmmNational Park Bashkirija , 453870, Bashkortostan Republic, Meleuzovskiy District, Nurgush, Russian Federation;
                [66] nnnState Nature Reserve Kurilsky , 694500, Sakhalin, Juzhno-Kurilsk, Russian Federation;
                [67] oooVodlozersky National Park , 185002, Karelia, Petrozavodsk, Russian Federation;
                [68] pppState Nature Reserve Kivach , 186220, Kondopoga District, Republic of Karelia, Russian Federation;
                [69] qqqSouth-Ural Federal University , 4563304, Chelyabinskaya Oblast, Miass, Russian Federation;
                [70] rrrSaint-Petersburg State Forest Technical University , 194021, St. Petersburg, Russian Federation;
                [71] sssAstrakhan Biosphere Reserve , 414021, Astrakhan, Russian Federation;
                [72] tttFSBI United Administration of the Lazovsky State Reserve and National Park Zov Tigra , 692980, Primorskiy Krai, Lazovskiy District, Lazo, Russian Federation;
                [73] uuuState Nature Reserve Tungusskiy , 660028, Krasnoyarsk Region, Russian Federation;
                [74] vvvKrasnoyarsk State Pedagogical University, Named after V. P. Astafyev , 660049, Krasnoyarsk, Russian Federation;
                [75] wwwInstitute of Geography, Russian Academy of Sciences , 119017, Moscow, Russian Federation;
                [76] xxxKoltzov Institute of Developmental Biology, Russian Academy of Sciences , 119334, Moscow, Russian Federation;
                [77] yyyCarpathian National Nature Park , 78500, Ivano-Frankivsk Region, Ukraine;
                [78] zzzState Environmental Institution National Park Braslav Lakes , 211970, Vitebsk Region, Braslav, Belarus;
                [79] aaaaNational Park Synevyr , 90041, Zakarpattia Region, Mizhhirs’kyi District, Synevyr-Ostriki, Ukraine;
                [80] bbbbPasvik State Nature Reserve , 184421, Murmansk Region, Nikel, Russian Federation;
                [81] ccccMari Chodra National Park , 425090, Mari El Republic, Zvenigovsky District, Krasnogorsky Settlement, Russian Federation;
                [82] ddddState Nature Reserve Vishersky , 618590, Perm Region, Krasnovishersk, Russian Federation;
                [83] eeeeState Nature Reserve Olekminsky , 678100, Republic Sakha, Olekminsk, Russian Federation;
                [84] ffffCrimea Nature Reserve , 298514, Alushta, Republic of Crimea;
                [85] ggggForest Research Institute, Karelian Research Centre, Russian Academy of Sciences , 185910, Karelia, Petrozavodsk, Russian Federation;
                [86] hhhhBlack Sea Biosphere Reserve , 75600, Khersons’ka Oblast, Hola Prystan’, Ukraine;
                [87] iiiiInstitute of Physicochemical and Biological Problems in Soil Sciences, Russian Academy of Science , 142290, Moscow Region, Pushchino, Russian Federation;
                [88] jjjjState Nature Reserve Nurgush , 610002, Kirov, Russian Federation;
                [89] kkkkCaucasian State Biosphere Reserve of the Ministry of Natural Resources , 385000, Adygea Republik, Maykop, Russian Federation;
                [90] llllNational Nature Park Vyzhnytskiy , 59200, Chernivtsi Region, Vyzhnytsya District, Berehomet, Ukraine;
                [91] mmmmNational Park Khvalynsky , 412780, Region Saratov, Khvalynsk Sity, Russian Federation;
                [92] nnnnArctic and Antarctic Research Institute, State Research Center , 199397, Saint Petersburg, Russian Federation;
                [93] ooooInformation-Analytical Centre for Protected Areas , 123242, Moscow;
                [94] ppppFSBI United Administration of the Mordovia State Nature Reserve and National Park Smolny , 430005, Republic of Mordovia, Saransk, Russian Federation;
                [95] qqqqOgarev Mordovia State University , 430005, Republic of Mordovia, Saransk, Russian Federation;
                [96] rrrrState Nature Reserve Malaya Sosva , 628242, Tjumen Region, Sovetskiy, Russian Federation;
                [97] ssssSurhanskiy State Nature Reserve , 191404, Surhandarja Region, Sherabad, Uzbekistan;
                [98] ttttMordovia State Nature Reserve , 431230, Mordovia Republic, Temnikov Region, Village Pushta, Russian Federation;
                [99] uuuuCentre for Biodiversity Dynamics, Department of Biology, Norwegian University of Science and Technology , N-7491 Trondheim, Norway
                Author notes
                1To whom correspondence may be addressed. Email: delgadomar@ 123456uniovi.es .

                Edited by Susan P. Harrison, University of California, Davis, CA, and approved October 6, 2020 (received for review February 27, 2020)

                Author contributions: M.M.D., T.R., E.M., E. Gurarie, and O.O. designed research; M.M.D., T.R., E.M., C.L., E. Gurarie, M. Abadonova, O. Abduraimov, O. Adrianova, T.A., M. Akkiev, A.A., E.A., N.A., M. Antipin, K.A., S. Babina, M.B., O.B., A. Barabancova, I.B., N. Belova, N. Belyaeva, T.B., E. Bisikalova, A. Bobretsov, V. Bobrov, V. Bobrovskyi, E. Bochkareva, G.B., V. Bolshakov, S. Bondarchuk, E. Bukharova, A. Butunina, Y. Buyvolov, A. Buyvolova, Y. Bykov, E.C., O.C., N.C., S. Chistjakov, S. Chuhontseva, E.A.D., V.D., E.D., A. Dobrolyubov, L.D., S.D., Z.D., A. Dubanaev, Y.D., S.E., L.E., O.S.E., O. Ermakova, A.E., O. Evstigneev, I.F., V.F., T.F., S.G., A.G., I.G., D.G., N.G., E. Gorbunova, T.G., V.G., L.G., V.H., A.H., E.I., S.I., U.I., N.I., Y. Kalinkin, E. Kaygorodova, F.K., D.K., A. Knorre, L.K., E. Korobov, H.K., N. Korotkikh, G.K., S. Kossenko, E. Kotlugalyamova, E. Kozlovsky, V.K., A. Kozurak, I.K., A. Krasnopevtseva, S. Kruglikov, O.K., A. Kudryavtsev, E. Kulebyakina, Y. Kulsha, M. Kupriyanova, M. Kurbanbagamaev, A. Kutenkov, N. Kutenkova, N. Kuyantseva, A. Kuznetsov, E.L., P.L., K.L., N.L., A. Mahmudov, L.M., V.M., S.M., I.M., A. Meydus, A. Minin, O.M., M.M., A. Myslenkov, N. Nasonova, N. Nemtseva, I.N., T. Nezdoliy, T. Niroda, T. Novikova, D.P., A.P., K.P., P.V., S.P., N.P., T.P., I. Pospelov, E.P., I. Prokhorov, I. Prokosheva, L.P., I. Putrashyk, J.R., Y.R., O.R., M.R., I.R., S.R., M. Sahnevich, A. Samoylov, V. Sanko, I.S., S. Sazonov, Z.S., K.S., M. Shashkov, A. Shcherbakov, V. Shevchyk, S. Shubin, E. Shujskaja, R.S., N. Sikkila, E. Sitnikova, A. Sivkov, N. Skok, S. Skorokhodova, E. Smirnova, G. Sokolova, V. Sopin, Y.S., S. Stepanov, V. Stratiy, V. Strekalovskaya, A. Sukhov, G. Suleymanova, L.S., V. Teleganova, V. Teplov, V. Teplova, T.T., V. Timoshkin, D.T., A. Tolmachev, A. Tomilin, L.T., M.T., Y.T., V. Van, E.E., A. Vasin, A. Vasina, A. Vekliuk, L.V., V. Vinogradov, N.V., I.V., T.X., E.Y.-G., V.Y., M.Y., O.Y., Y.Y., A. Zahvatov, V.Z., N.Z., A. Zheltukhin, T.Z., J.K., and O.O. performed research; G.T. and O.O. contributed new reagents/analytic tools; M.M.D., T.R., G.T., and O.O. analyzed data; M.M.D., T.R., E. Gurarie, and O.O. wrote the paper; E.M., C.L., and J.K. performed database organization; and M. Abadonova, O. Abduraimov, O. Adrianova, T.A., M. Akkiev, A.A., E.A., N.A., M. Antipin, K.A., S. Babina, M.B., O.B., A. Barabancova, I.B., N. Belova, N. Belyaeva, T.B., E. Bisikalova, A. Bobretsov, V. Bobrov, V. Bobrovskyi, E. Bochkareva, G.B., V. Bolshakov, S. Bondarchuk, E. Bukharova, A. Butunina, Y. Buyvolov, A. Buyvolova, Y. Bykov, E.C., O.C., N.C., S. Chistjakov, S. Chuhontseva, E.A.D., V.D., E.D., A. Dobrolyubov, L.D., S.D., Z.D., A. Dubanaev, Y.D., S.E., L.E., O.S.E., O. Ermakova, A.E., O. Evstigneev, I.F., V.F., T.F., S.G., A.G., I.G., D.G., N.G., E. Gorbunova, T.G., V.G., L.G., V.H., A.H., E.I., S.I., U.I., N.I., Y. Kalinkin, E. Kaygorodova, F.K., D.K., A. Knorre, L.K., E. Korobov, H.K., N. Korotkikh, G.K., S. Kossenko, E. Kotlugalyamova, E. Kozlovsky, V.K., A. Kozurak, I.K., A. Krasnopevtseva, S. Kruglikov, O.K., A. Kudryavtsev, E. Kulebyakina, Y. Kulsha, M. Kupriyanova, M. Kurbanbagamaev, A. Kutenkov, N. Kutenkova, N. Kuyantseva, A. Kuznetsov, E.L., P.L., K.L., N.L., A. Mahmudov, L.M., V.M., S.M., I.M., A. Meydus, A. Minin, O.M., M.M., A. Myslenkov, N. Nasonova, N. Nemtseva, I.N., T. Nezdoliy, T. Niroda, T. Novikova, D.P., A.P., K.P., P.V., S.P., N.P., T.P., I. Pospelov, E.P., I. Prokhorov, I. Prokosheva, L.P., I. Putrashyk, J.R., Y.R., O.R., M.R., I.R., S.R., M. Sahnevich, A. Samoylov, V. Sanko, I.S., S. Sazonov, Z.S., K.S., M. Shashkov, A. Shcherbakov, V. Shevchyk, S. Shubin, E. Shujskaja, R.S., N. Sikkila, E. Sitnikova, A. Sivkov, N. Skok, S. Skorokhodova, E. Smirnova, G. Sokolova, V. Sopin, Y.S., S. Stepanov, V. Stratiy, V. Strekalovskaya, A. Sukhov, G. Suleymanova, L.S., V. Teleganova, V. Teplov, V. Teplova, T.T., V. Timoshkin, D.T., A. Tolmachev, A. Tomilin, L.T., M.T., Y.T., V. Van, E.E., A. Vasin, A. Vasina, A. Vekliuk, L.V., V. Vinogradov, N.V., I.V., T.X., E.Y.-G., V.Y., M.Y., O.Y., Y.Y., A. Zahvatov, V.Z., N.Z., A. Zheltukhin, T.Z., and J.K. performed data collection.

                2M.M.D. and T.R. contributed equally to this work.

                3Deceased 30 August 2020.

                4Deceased 16 October 2016.

                5Deceased 26 June 2015.

                6Deceased 2 July 2018.

                7Deceased 10 August 2011.

                8Deceased 12 January 2020.

                Author information
                http://orcid.org/0000-0002-3009-738X
                http://orcid.org/0000-0002-2957-4791
                http://orcid.org/0000-0003-1986-2631
                http://orcid.org/0000-0002-8666-9674
                http://orcid.org/0000-0001-9087-8949
                http://orcid.org/0000-0003-4751-7587
                http://orcid.org/0000-0002-6611-353X
                http://orcid.org/0000-0002-9251-0563
                http://orcid.org/0000-0003-1893-4728
                http://orcid.org/0000-0001-6756-516X
                http://orcid.org/0000-0003-3428-944X
                http://orcid.org/0000-0002-3641-5203
                http://orcid.org/0000-0002-6726-9368
                http://orcid.org/0000-0003-1350-2088
                http://orcid.org/0000-0002-7097-5378
                http://orcid.org/0000-0003-0962-7697
                http://orcid.org/0000-0002-7843-4189
                http://orcid.org/0000-0002-2316-8506
                http://orcid.org/0000-0003-0225-3207
                http://orcid.org/0000-0002-2620-0858
                http://orcid.org/0000-0002-3724-9761
                http://orcid.org/0000-0002-5196-4438
                http://orcid.org/0000-0002-6052-3263
                http://orcid.org/0000-0002-5774-0427
                http://orcid.org/0000-0001-6211-8213
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                http://orcid.org/0000-0001-9033-2292
                http://orcid.org/0000-0003-4782-2979
                http://orcid.org/0000-0003-4785-898X
                http://orcid.org/0000-0003-1757-1734
                http://orcid.org/0000-0002-1848-7588
                http://orcid.org/0000-0003-4974-7909
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                http://orcid.org/0000-0001-9564-5589
                http://orcid.org/0000-0001-6109-4977
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                Article
                202002713
                10.1073/pnas.2002713117
                7733824
                33229550
                8b14f411-6233-406f-99bf-d87c2d5f3c91
                Copyright © 2020 the Author(s). Published by PNAS.

                This open access article is distributed under Creative Commons Attribution-NonCommercial-NoDerivatives License 4.0 (CC BY-NC-ND).

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                Page count
                Pages: 10
                Categories
                Biological Sciences
                Ecology

                chilling,climate change,heating,phenology,plasticity
                chilling, climate change, heating, phenology, plasticity

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