{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,3]],"date-time":"2026-10-03T05:53:06Z","timestamp":1791006786110,"version":"4.1.0"},"reference-count":112,"publisher":"Wiley","issue":"8","license":[{"start":{"date-parts":[[2026,8,10]],"date-time":"2026-08-10T00:00:00Z","timestamp":1786320000000},"content-version":"vor","delay-in-days":9,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"},{"start":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T00:00:00Z","timestamp":1785542400000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/501100013265","name":"Norsk Polarinstitutt","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100013265","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100001399","name":"World Wildlife Fund","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100001399","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100005416","name":"Norges Forskningsr\u00e5d","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100005416","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Global Change Biology"],"published-print":{"date-parts":[[2026,8]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>\n                    Climate change is disrupting myriad biological processes, including the ability of species to synchronize life\u2010history events with optimal environmental conditions, with potential fitness impacts. Changes in migratory and breeding phenology and their consequences on fitness have been well studied in birds and some mammals. By contrast, changes in the phenology of overwinter sheltering and their influence on fitness have rarely been investigated. Pregnant female polar bears (\n                    <jats:styled-content style=\"fixed-case\">\n                      <jats:italic>Ursus maritimus<\/jats:italic>\n                    <\/jats:styled-content>\n                    ) spend the winter fasting in a maternity den where they give birth and nurse their cubs until emergence in early spring. To sustain their fast, females rely on energy stores built primarily through consumption of ice\u2010associated seals. Here, we assess the impact of sea\u2010ice loss driven by climate change on early reproductive success acting through disrupted denning phenology and other unspecified factors. To do so, we develop a Bayesian hierarchical model combining capture\u2013recapture and path modeling. We apply it to 38\u2009years of capture\u2013recapture data along with biologging data on denning status and phenology, while accounting for imperfect detection and uncertainty in individual denning schedules. We found that low sea\u2010ice availability led to earlier den emergence, which in turn reduced early litter survival and possibly litter size. By contrast, there was little evidence that sea\u2010ice availability affected early reproductive success through pathways independent of denning phenology (e.g., litter mass). Considering both direct and indirect effects, low sea\u2010ice availability reduced early litter survival but not the size of surviving litters. Given the strong link between sea\u2010ice conditions and body condition in female polar bears, we conclude that sea\u2010ice loss exerts an energetic constraint on denning females, forcing them to depart from dens early, which jeopardizes their cubs, ultimately reducing maternal fitness. Similar impacts of climate change may occur in other species where phenology is determined by body condition.\n                  <\/jats:p>","DOI":"10.1111\/gcb.71042","type":"journal-article","created":{"date-parts":[[2026,8,11]],"date-time":"2026-08-11T04:53:16Z","timestamp":1786423996000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Denning Phenology Mediates Sea\u2010Ice Loss Impacts on Early Reproductive Success in Polar Bears"],"prefix":"10.1111","volume":"32","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1489-9363","authenticated-orcid":false,"given":"Marwan","family":"Naciri","sequence":"first","affiliation":[{"name":"CEFE University of Montpellier, CNRS, EPHE, IRD  Montpellier France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0574-6712","authenticated-orcid":false,"given":"Jon","family":"Aars","sequence":"additional","affiliation":[{"name":"Norwegian Polar Institute, FRAM Centre  Troms\u00f8 Norway"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9433-2369","authenticated-orcid":false,"given":"Marl\u00e8ne","family":"Gamelon","sequence":"additional","affiliation":[{"name":"Universit\u00e9 de Lyon, Universit\u00e9 Lyon 1, CNRS, VetAgro Sup, Laboratoire de Biom\u00e9trie et Biologie Evolutive, UMR 5558  Villeurbanne France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Magnus","family":"Andersen","sequence":"additional","affiliation":[{"name":"Norwegian Polar Institute, FRAM Centre  Troms\u00f8 Norway"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1104-7774","authenticated-orcid":false,"given":"Andrew E.","family":"Derocher","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences University of Alberta  Edmonton Alberta Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0395-5251","authenticated-orcid":false,"given":"\u00d8ystein","family":"Wiig","sequence":"additional","affiliation":[{"name":"Natural History Museum  Oslo Norway"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sarah","family":"Cubaynes","sequence":"additional","affiliation":[{"name":"CEFE Univ Montpellier, CNRS, EPHE\u2010PSL University, IRD  Montpellier France"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,8,10]]},"reference":[{"key":"e_1_2_11_2_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41598\u2010025\u201033227\u20109"},{"key":"e_1_2_11_3_1","doi-asserted-by":"publisher","DOI":"10.1080\/17518369.2017.1374125"},{"key":"e_1_2_11_4_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469\u20107998.2008.00415.x"},{"key":"e_1_2_11_5_1","doi-asserted-by":"publisher","DOI":"10.1111\/gcb.13435"},{"key":"e_1_2_11_6_1","first-page":"587","volume-title":"Wild Mammals of North America: Biology, Management, and Conservation","author":"Amstrup S. C.","year":"2003"},{"key":"e_1_2_11_7_1","doi-asserted-by":"publisher","DOI":"10.1093\/jmammal\/gyae010"},{"key":"e_1_2_11_8_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00300\u2010011\u20101094\u2010y"},{"key":"e_1_2_11_9_1","doi-asserted-by":"publisher","DOI":"10.3354\/meps14382"},{"key":"e_1_2_11_10_1","doi-asserted-by":"publisher","DOI":"10.1016\/0306\u20104565(91)90029\u20102"},{"key":"e_1_2_11_11_1","doi-asserted-by":"publisher","DOI":"10.1086\/physzool.69.2.30164186"},{"key":"e_1_2_11_12_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.gecco.2021.e01925"},{"key":"e_1_2_11_13_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41598\u2010024\u201053575\u20102"},{"key":"e_1_2_11_14_1","doi-asserted-by":"publisher","DOI":"10.1086\/375680"},{"key":"e_1_2_11_15_1","doi-asserted-by":"publisher","DOI":"10.1152\/ajpregu.1979.236.1.R67"},{"key":"e_1_2_11_16_1","doi-asserted-by":"publisher","DOI":"10.1111\/eva.12121"},{"key":"e_1_2_11_17_1","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2009.0140"},{"key":"e_1_2_11_18_1","first-page":"449","article-title":"Evaluation of Age Determination of Polar Bears by Counts of Cementum Growth Layer Groups","volume":"10","author":"Calvert W.","year":"1998","journal-title":"Ursus"},{"key":"e_1_2_11_19_1","volume-title":"Sea Ice Concentrations From Nimbus\u20107 SMMR and DMSP SSM\/I\u2010SSMIS Passive Microwave Data, Version 1","author":"Cavalieri D. J.","year":"1996"},{"key":"e_1_2_11_20_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41558\u2010018\u20100067\u20103"},{"key":"e_1_2_11_21_1","doi-asserted-by":"publisher","DOI":"10.1890\/14\u20101365.1"},{"key":"e_1_2_11_22_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0008677"},{"key":"e_1_2_11_23_1","doi-asserted-by":"publisher","DOI":"10.1002\/ece3.72914"},{"key":"e_1_2_11_24_1","doi-asserted-by":"publisher","DOI":"10.1890\/11\u20100258.1"},{"key":"e_1_2_11_25_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2009003117"},{"key":"e_1_2_11_26_1","doi-asserted-by":"publisher","DOI":"10.3354\/meps09406"},{"key":"e_1_2_11_27_1","doi-asserted-by":"publisher","DOI":"10.1139\/z95\u2010197"},{"key":"e_1_2_11_28_1","doi-asserted-by":"publisher","DOI":"10.1111\/gcb.13057"},{"key":"e_1_2_11_29_1","doi-asserted-by":"publisher","DOI":"10.3354\/cr033271"},{"key":"e_1_2_11_30_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00300\u2010007\u20100300\u20104"},{"key":"e_1_2_11_31_1","doi-asserted-by":"publisher","DOI":"10.1098\/rsbl.2019.0070"},{"key":"e_1_2_11_32_1","doi-asserted-by":"publisher","DOI":"10.3354\/meps09516"},{"key":"e_1_2_11_33_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365\u20102656.2009.01615.x"},{"key":"e_1_2_11_34_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0101410"},{"key":"e_1_2_11_35_1","doi-asserted-by":"publisher","DOI":"10.1111\/gcb.14817"},{"key":"e_1_2_11_36_1","doi-asserted-by":"publisher","DOI":"10.1006\/anbe.2001.1803"},{"key":"e_1_2_11_37_1","doi-asserted-by":"publisher","DOI":"10.1086\/physzool.68.6.30163788"},{"key":"e_1_2_11_38_1","doi-asserted-by":"publisher","DOI":"10.1214\/ss\/1177011136"},{"key":"e_1_2_11_39_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.2041\u2010210X.2011.00150.x"},{"key":"e_1_2_11_40_1","doi-asserted-by":"publisher","DOI":"10.1086\/589520"},{"key":"e_1_2_11_41_1","doi-asserted-by":"publisher","DOI":"10.1093\/icb\/28.3.863"},{"key":"e_1_2_11_42_1","doi-asserted-by":"publisher","DOI":"10.1017\/CBO9780511617799"},{"key":"e_1_2_11_43_1","doi-asserted-by":"publisher","DOI":"10.1093\/beheco\/arac116"},{"key":"e_1_2_11_44_1","doi-asserted-by":"publisher","DOI":"10.1111\/1365\u20102664.13021"},{"key":"e_1_2_11_45_1","volume-title":"Global Daily Gridded 5\u2010Day Median\u2010Filtered, Gap\u2010Filled ASI Algorithm SSMI\u2010SSMIS Sea Ice Concentration Data","author":"Kern S.","year":"2020"},{"key":"e_1_2_11_46_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1541\u20100420.2008.00991.x"},{"key":"e_1_2_11_47_1","doi-asserted-by":"publisher","DOI":"10.1086\/319193"},{"key":"e_1_2_11_48_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ecolmodel.2024.110790"},{"key":"e_1_2_11_49_1","doi-asserted-by":"publisher","DOI":"10.3389\/fcosc.2024.1439386"},{"key":"e_1_2_11_50_1","doi-asserted-by":"publisher","DOI":"10.1002\/eap.2071"},{"key":"e_1_2_11_51_1","doi-asserted-by":"publisher","DOI":"10.1111\/gcb.15286"},{"key":"e_1_2_11_52_1","doi-asserted-by":"publisher","DOI":"10.3389\/fmars.2022.816666"},{"key":"e_1_2_11_53_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature11335"},{"key":"e_1_2_11_54_1","doi-asserted-by":"publisher","DOI":"10.1111\/ele.12437"},{"key":"e_1_2_11_55_1","doi-asserted-by":"publisher","DOI":"10.1111\/ele.13634"},{"key":"e_1_2_11_56_1","doi-asserted-by":"publisher","DOI":"10.2307\/2937171"},{"key":"e_1_2_11_57_1","doi-asserted-by":"publisher","DOI":"10.1890\/09\u20100766.1"},{"key":"e_1_2_11_58_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41598\u2010018\u201027947\u20104"},{"key":"e_1_2_11_59_1","unstructured":"L\u00f8n\u00f8 O.1970.\u201cPolar Bear Fetuses Found in Svalbard.\u201dhttps:\/\/eurekamag.com\/research\/023\/377\/023377480.php."},{"key":"e_1_2_11_60_1","doi-asserted-by":"publisher","DOI":"10.1890\/15\u20101256"},{"key":"e_1_2_11_61_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.jmarsys.2013.09.006"},{"key":"e_1_2_11_62_1","doi-asserted-by":"publisher","DOI":"10.1139\/z01\u2010126"},{"key":"e_1_2_11_63_1","doi-asserted-by":"publisher","DOI":"10.1046\/j.1365\u20102664.2002.00690.x"},{"key":"e_1_2_11_64_1","doi-asserted-by":"publisher","DOI":"10.1093\/conphys\/coaf058"},{"key":"e_1_2_11_65_1","doi-asserted-by":"publisher","DOI":"10.1111\/ele.12599"},{"key":"e_1_2_11_66_1","doi-asserted-by":"publisher","DOI":"10.1186\/s40317\u2010023\u201000323\u20104"},{"key":"e_1_2_11_67_1","doi-asserted-by":"publisher","DOI":"10.33265\/polar.v39.3447"},{"key":"e_1_2_11_68_1","doi-asserted-by":"publisher","DOI":"10.2307\/1382563"},{"key":"e_1_2_11_69_1","doi-asserted-by":"publisher","DOI":"10.1002\/ecs2.70370"},{"key":"e_1_2_11_70_1","doi-asserted-by":"publisher","DOI":"10.1111\/2041\u2010210x.70229"},{"key":"e_1_2_11_71_1","unstructured":"NIMBLE Development Team.2026.\u201cNIMBLE: MCMC Particle Filtering and Programmable Hierarchical Modeling (Version 1.4.2) [Computer Software].\u201dZenodo.https:\/\/doi.org\/10.5281\/ZENODO.1211190."},{"key":"e_1_2_11_72_1","doi-asserted-by":"publisher","DOI":"10.1139\/as\u20102015\u20100027"},{"key":"e_1_2_11_73_1","doi-asserted-by":"publisher","DOI":"10.3354\/meps12000"},{"key":"e_1_2_11_74_1","doi-asserted-by":"crossref","unstructured":"Pedersen T. L.2020.\u201cpatchwork: The Composer of Plots (R Package Version 1.1.1)\u201d[Computer Software].https:\/\/CRAN.R\u2010project.org\/package=patchwork.","DOI":"10.32614\/CRAN.package.patchwork"},{"key":"e_1_2_11_75_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.95.1.213"},{"key":"e_1_2_11_76_1","doi-asserted-by":"publisher","DOI":"10.1111\/1365\u20102656.13593"},{"key":"e_1_2_11_77_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00265\u2010016\u20102180\u20105"},{"key":"e_1_2_11_78_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.1001828"},{"key":"e_1_2_11_79_1","doi-asserted-by":"publisher","DOI":"10.3389\/fevo.2015.00033"},{"key":"e_1_2_11_80_1","volume-title":"R: A Language and Environment for Statistical Computing","author":"R Core Team","year":"2025"},{"key":"e_1_2_11_81_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469\u20107998.1988.tb03762.x"},{"key":"e_1_2_11_82_1","doi-asserted-by":"publisher","DOI":"10.1038\/s43247\u2010022\u201000498\u20103"},{"key":"e_1_2_11_83_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365\u20102656.2009.01603.x"},{"key":"e_1_2_11_84_1","doi-asserted-by":"publisher","DOI":"10.1644\/11\u2010MAMM\u2010A\u2010155.1"},{"key":"e_1_2_11_85_1","doi-asserted-by":"publisher","DOI":"10.1644\/11\u2010MAMM\u2010A\u2010406.1"},{"key":"e_1_2_11_86_1","doi-asserted-by":"publisher","DOI":"10.1093\/jmammal\/gyx181"},{"key":"e_1_2_11_87_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10144\u2010011\u20100299\u20109"},{"key":"e_1_2_11_88_1","doi-asserted-by":"publisher","DOI":"10.1111\/gcb.15572"},{"key":"e_1_2_11_89_1","doi-asserted-by":"publisher","DOI":"10.1111\/gcb.12339"},{"key":"e_1_2_11_90_1","doi-asserted-by":"publisher","DOI":"10.1111\/gcb.13933"},{"key":"e_1_2_11_91_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365\u20102656.2009.01537.x"},{"key":"e_1_2_11_92_1","doi-asserted-by":"publisher","DOI":"10.1086\/285627"},{"key":"e_1_2_11_93_1","doi-asserted-by":"publisher","DOI":"10.2307\/1382569"},{"key":"e_1_2_11_94_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature13696"},{"key":"e_1_2_11_95_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365\u20102907.2007.00096.x"},{"key":"e_1_2_11_96_1","volume-title":"The Evolution of Life Histories","author":"Stearns S. C.","year":"1992"},{"key":"e_1_2_11_97_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.212519399"},{"key":"e_1_2_11_98_1","doi-asserted-by":"publisher","DOI":"10.5194\/tc\u201010\u20102027\u20102016"},{"key":"e_1_2_11_99_1","doi-asserted-by":"publisher","DOI":"10.1002\/ecs2.4826"},{"key":"e_1_2_11_100_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365\u20102486.2012.02753.x"},{"key":"e_1_2_11_101_1","doi-asserted-by":"publisher","DOI":"10.7589\/0090\u20103558\u201025.2.159"},{"key":"e_1_2_11_102_1","doi-asserted-by":"publisher","DOI":"10.1890\/12\u20100833.1"},{"key":"e_1_2_11_103_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature18608"},{"key":"e_1_2_11_104_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1227548"},{"key":"e_1_2_11_105_1","doi-asserted-by":"publisher","DOI":"10.14430\/arctic615"},{"key":"e_1_2_11_106_1","doi-asserted-by":"publisher","DOI":"10.1139\/cjz\u20102014\u20100084"},{"key":"e_1_2_11_107_1","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.1998.0514"},{"key":"e_1_2_11_108_1","doi-asserted-by":"publisher","DOI":"10.1111\/ecog.06056"},{"key":"e_1_2_11_109_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-319-24277-4"},{"key":"e_1_2_11_110_1","first-page":"25","article-title":"Survival and Reproductive Rates for Polar Bears at Svalbard","volume":"10","author":"Wiig \u00d8.","year":"1998","journal-title":"Ursus"},{"key":"e_1_2_11_111_1","doi-asserted-by":"publisher","DOI":"10.1002\/eap.2112"},{"key":"e_1_2_11_112_1","doi-asserted-by":"publisher","DOI":"10.1111\/1365\u20102656.12790"},{"key":"e_1_2_11_113_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2221961"}],"container-title":["Global Change Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/gcb.71042","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1111\/gcb.71042","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/gcb.71042","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,3]],"date-time":"2026-10-03T05:05:37Z","timestamp":1791003937000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1111\/gcb.71042"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8]]},"references-count":112,"journal-issue":{"issue":"8","published-print":{"date-parts":[[2026,8]]}},"alternative-id":["10.1111\/gcb.71042"],"URL":"https:\/\/doi.org\/10.1111\/gcb.71042","archive":["Portico"],"relation":{},"ISSN":["1354-1013","1365-2486"],"issn-type":[{"value":"1354-1013","type":"print"},{"value":"1365-2486","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8]]},"assertion":[{"value":"2025-11-17","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-07-27","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-08-10","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e71042"}}