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Migratory animals may respond by advancing their phenology or colonising colder areas where spring starts later. The role of climate change in range expansion can be both driving (making traditional areas suboptimal) and facilitating (making new areas suitable).<\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>\n                          Recently, Pink\u2010footed Geese (\n                          <jats:italic>Anser brachyrhynchus<\/jats:italic>\n                          ) from Svalbard showed extreme range expansion by colonising the colder Novaya Zemlya as breeding ground, involving a new migration route. We examine potential costs and benefits associated with breeding in this new area.\n                        <\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>We use GPS\u2010tracking, long\u2010term population monitoring and remote sensing, to compare spring onset, migration timing and breeding performance between both flyways, and to evaluate how spring onset affects different reproductive stages in both breeding areas in these capital breeders.<\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>The traditional route showed challenges for migration timing, as spring had advanced in Svalbard (while arrival date had not kept up) but not on stopovers, and recently, early spring on Svalbard correlated with late spring on stopovers. On new stopovers, spring did advance. Spring was later in Novaya Zemlya than Svalbard, yet arrival dates were similar. In Novaya Zemlya, egg laying occurred later than in Svalbard, but still earlier relative to local spring onset (e.g. snowmelt and green\u2010up), suggesting a smaller mismatch. The period between arrival and egg laying was longer in Novaya Zemlya than Svalbard, but breeding performance was similar. Finally, on the new route, geese were larger and (relatively) heavier than on the traditional route, thus possibly carrying larger capital body stores to cover harsher pre\u2010laying periods.<\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>Our results suggest that colonising new breeding areas enables populations to regain phenological match with the environment, especially when advancement of migration timing was limited. Still, breeding in a colder area may require more parental investment, such as body stores. Thus, a benefit for offspring comes at a cost for parents.<\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>This mechanism can cause climate change to drive and facilitate colonisation especially in individuals capable of large investments. Consequently, variation in individual quality leads to heterogeneous effects of climate change within a population. These processes may play at the \u2018cold\u2019 edge of any range shift.<\/jats:p>\n                      <\/jats:list-item>\n                    <\/jats:list>\n                  <\/jats:p>","DOI":"10.1111\/1365-2656.70172","type":"journal-article","created":{"date-parts":[[2025,11,12]],"date-time":"2025-11-12T05:17:16Z","timestamp":1762924636000},"page":"97-114","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Arctic geese in newly colonised, colder breeding areas have higher spring body mass and breed earlier relative to the onset of spring"],"prefix":"10.1111","volume":"95","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-0550-7447","authenticated-orcid":false,"given":"Kees H. T.","family":"Schreven","sequence":"first","affiliation":[{"name":"Department of Animal Ecology Netherlands Institute of Ecology (NIOO\u2010KNAW)  Wageningen The Netherlands"},{"name":"Department of Theoretical and Computational Ecology Institute of Biodiversity and Ecosystem Dynamics (IBED), University of Amsterdam  Amsterdam The Netherlands"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6938-663X","authenticated-orcid":false,"given":"Tom S. L.","family":"Versluijs","sequence":"additional","affiliation":[{"name":"Department of Coastal Systems Royal Netherlands Institute for Sea, Research (NIOZ), \u2018t Horntje  Texel The Netherlands"},{"name":"Groningen Institute for Evolutionary Life Sciences (GELIFES), University of Groningen  Groningen The Netherlands"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6448-2446","authenticated-orcid":false,"given":"Michiel P.","family":"Boom","sequence":"additional","affiliation":[{"name":"Department of Animal Ecology Netherlands Institute of Ecology (NIOO\u2010KNAW)  Wageningen The Netherlands"},{"name":"Department of Theoretical and Computational Ecology Institute of Biodiversity and Ecosystem Dynamics (IBED), University of Amsterdam  Amsterdam The Netherlands"},{"name":"Sovon Dutch Center for Field Ornithology  Nijmegen The Netherlands"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Fred","family":"Cottaar","sequence":"additional","affiliation":[{"name":"Independent Researcher  Haarlem The Netherlands"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Eckhart","family":"Kuijken","sequence":"additional","affiliation":[{"name":"Terrestrial Ecology Unit, Department of Biology Ghent University  Ghent Belgium"},{"name":"Independent Researcher  Beernem Belgium"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Jorma","family":"Pessa","sequence":"additional","affiliation":[{"name":"ELY\u2010Keskus\u2010Centre for Economic Development, Transport and the Environment  Oulu Finland"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ingunn M.","family":"Tombre","sequence":"additional","affiliation":[{"name":"Department for Arctic Ecology The Norwegian Institute for Nature Research (NINA)  Troms\u00f8 Norway"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Christine","family":"Verscheure","sequence":"additional","affiliation":[{"name":"Independent Researcher  Beernem Belgium"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3246-0215","authenticated-orcid":false,"given":"Jesper","family":"Madsen","sequence":"additional","affiliation":[{"name":"Department of Ecoscience Aarhus University  Aarhus Denmark"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7437-4879","authenticated-orcid":false,"given":"Bart A.","family":"Nolet","sequence":"additional","affiliation":[{"name":"Department of Animal Ecology Netherlands Institute of Ecology (NIOO\u2010KNAW)  Wageningen The Netherlands"},{"name":"Department of Theoretical and Computational Ecology Institute of Biodiversity and Ecosystem Dynamics (IBED), University of Amsterdam  Amsterdam The Netherlands"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"311","published-online":{"date-parts":[[2025,11,11]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2016.0252"},{"key":"e_1_2_9_3_1","doi-asserted-by":"publisher","DOI":"10.21105\/joss.02272"},{"key":"e_1_2_9_4_1","doi-asserted-by":"publisher","DOI":"10.18637\/jss.v067.i01"},{"key":"e_1_2_9_5_1","doi-asserted-by":"publisher","DOI":"10.1139\/cjz\u20102013\u20100036"},{"key":"e_1_2_9_6_1","doi-asserted-by":"publisher","DOI":"10.1111\/1365\u20102656.14020"},{"key":"e_1_2_9_7_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365\u20102656.2008.01458.x"},{"key":"e_1_2_9_8_1","doi-asserted-by":"publisher","DOI":"10.1038\/35077063"},{"key":"e_1_2_9_9_1","doi-asserted-by":"publisher","DOI":"10.5751\/ACE\u201000708\u2010100101"},{"key":"e_1_2_9_10_1","doi-asserted-by":"publisher","DOI":"10.1002\/ece3.563"},{"key":"e_1_2_9_11_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1206432"},{"key":"e_1_2_9_12_1","doi-asserted-by":"publisher","DOI":"10.1111\/jav.00899"},{"key":"e_1_2_9_13_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00442\u2010013\u20102681\u20100"},{"key":"e_1_2_9_14_1","doi-asserted-by":"publisher","DOI":"10.1111\/gcb.14061"},{"key":"e_1_2_9_15_1","first-page":"13","article-title":"Effects of capture and marking on the behaviour of moulting pink\u2010footed geese Anser brachyrhynchus on Svalbard","volume":"70","author":"Clausen K. 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