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Understanding tree functional constraints under drought and their impacts on resilience and mortality among species is crucial to assess the impacts of global change on forests. We analyzed the long\u2010term drought and atmospheric CO<jats:sub>2<\/jats:sub> responses in three Mediterranean co\u2010occurring species with differing drought tolerances (<jats:styled-content style=\"fixed-case\"><jats:italic>Pinus pinaster<\/jats:italic><\/jats:styled-content>\u2009&lt;\u2009<jats:styled-content style=\"fixed-case\"><jats:italic>Pinus pinea<\/jats:italic><\/jats:styled-content>\u2009&lt;\u2009<jats:styled-content style=\"fixed-case\"><jats:italic>Juniperus oxycedrus<\/jats:italic><\/jats:styled-content>). In this mixed forest, <jats:styled-content style=\"fixed-case\"><jats:italic>P. pinaster<\/jats:italic><\/jats:styled-content> exhibited widespread mortality and mistletoe infection, <jats:styled-content style=\"fixed-case\"><jats:italic>P. pinea<\/jats:italic><\/jats:styled-content> showed scattered mortality, and <jats:styled-content style=\"fixed-case\"><jats:italic>J. oxycedrus<\/jats:italic><\/jats:styled-content> showed no decline. Using tree\u2010ring data (1978\u20132016), we compared intrinsic water\u2010use efficiency (iWUE) and xylem hydraulic traits in healthy and non\u2010healthy individuals of both pine species and healthy junipers. Healthy <jats:styled-content style=\"fixed-case\"><jats:italic>P. pinaster<\/jats:italic><\/jats:styled-content> trees produced a more hydraulically efficient xylem, with wider lumen tracheids, than non\u2010healthy trees, whereas <jats:styled-content style=\"fixed-case\"><jats:italic>P. pinea<\/jats:italic><\/jats:styled-content> showed no anatomical differences between health statuses. Healthy <jats:styled-content style=\"fixed-case\"><jats:italic>P. pinaster<\/jats:italic><\/jats:styled-content> displayed greater anatomical plasticity, adjusting hydraulic conductivity and cell\u2010wall thickness to water availability. Despite small differences in average iWUE, the response of iWUE to rising CO<jats:sub>2<\/jats:sub> and drought differed between species and health statuses. <jats:styled-content style=\"fixed-case\"><jats:italic>J. oxycedrus<\/jats:italic><\/jats:styled-content> and <jats:styled-content style=\"fixed-case\"><jats:italic>P. pinea<\/jats:italic><\/jats:styled-content> showed steady iWUE increases, but <jats:styled-content style=\"fixed-case\"><jats:italic>P. pinea<\/jats:italic><\/jats:styled-content> experienced periods of stagnation following an extreme drought, later recovering regardless of health status. In contrast, iWUE in <jats:styled-content style=\"fixed-case\"><jats:italic>P. pinaster<\/jats:italic><\/jats:styled-content> plateaued for over 20\u2009years after a decline\u2010inducing drought, particularly in non\u2010healthy, mistletoe\u2010infected trees. Differences in iWUE response to CO<jats:sub>2<\/jats:sub> and anatomical plasticity to drought may explain the contrasting mortality patterns among these coniferous species. Our results suggest a long\u2010term decline spiral in <jats:styled-content style=\"fixed-case\"><jats:italic>P. pinaster<\/jats:italic><\/jats:styled-content> induced by low hydraulic efficiency in drought\u2010induced defoliated trees and limited physiological responses to rising CO<jats:sub>2<\/jats:sub> and drought. Increasing drought stress makes pine recovery increasingly unlikely.<\/jats:p>","DOI":"10.1111\/gcb.70221","type":"journal-article","created":{"date-parts":[[2025,5,20]],"date-time":"2025-05-20T07:16:35Z","timestamp":1747725395000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":10,"title":["Loss of Stomatal Regulation Sensitivity to <scp>CO<sub>2<\/sub><\/scp> and Reduced Xylem Hydraulic Conductivity Contribute to Long\u2010Term Tree Decline and Mortality"],"prefix":"10.1111","volume":"31","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6738-3312","authenticated-orcid":false,"given":"Dario","family":"Martin\u2010Benito","sequence":"first","affiliation":[{"name":"Institute of Forest Sciences ICIFOR Inia\u2010CSIC  Madrid Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2512-5187","authenticated-orcid":false,"given":"Macarena","family":"F\u00e9rriz","sequence":"additional","affiliation":[{"name":"Institute of Forest Sciences ICIFOR Inia\u2010CSIC  Madrid Spain"},{"name":"Department of Geography Indiana University  Bloomington Indiana USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0194-2617","authenticated-orcid":false,"given":"Mar\u00eda","family":"Conde","sequence":"additional","affiliation":[{"name":"Institute of Forest Sciences ICIFOR Inia\u2010CSIC  Madrid Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8566-4599","authenticated-orcid":false,"given":"Georg","family":"von Arx","sequence":"additional","affiliation":[{"name":"Swiss Federal Institute for Forest, Snow and Landscape Research WSL  Birmensdorf Switzerland"},{"name":"Oeschger Centre for Climate Change Research University of Bern  Bern Switzerland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7070-3292","authenticated-orcid":false,"given":"Patrick","family":"Fonti","sequence":"additional","affiliation":[{"name":"Swiss Federal Institute for Forest, Snow and Landscape Research WSL  Birmensdorf Switzerland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4526-5462","authenticated-orcid":false,"given":"Jos\u00e9 Miguel","family":"Olano","sequence":"additional","affiliation":[{"name":"iuFOR, EiFAB Universidad de Valladolid  Soria Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0148-3721","authenticated-orcid":false,"given":"Guillermo","family":"Gea\u2010Izquierdo","sequence":"additional","affiliation":[{"name":"Institute of Forest Sciences ICIFOR Inia\u2010CSIC  Madrid Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2025,5,20]]},"reference":[{"key":"e_1_2_10_2_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41559\u2010017\u20100248\u2010x"},{"key":"e_1_2_10_3_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41558\u2010020\u20100747\u20107"},{"key":"e_1_2_10_4_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.95.25.14839"},{"key":"e_1_2_10_5_1","doi-asserted-by":"publisher","DOI":"10.1890\/es15\u201000203.1"},{"key":"e_1_2_10_6_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1525678113"},{"key":"e_1_2_10_7_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-8137.2005.01587.x"},{"key":"e_1_2_10_8_1","doi-asserted-by":"publisher","DOI":"10.1111\/ppl.14562"},{"key":"e_1_2_10_9_1","doi-asserted-by":"publisher","DOI":"10.18637\/jss.v111.i09"},{"key":"e_1_2_10_10_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365\u20103040.2010.02269.x"},{"key":"e_1_2_10_11_1","doi-asserted-by":"publisher","DOI":"10.18637\/jss.v067.i01"},{"key":"e_1_2_10_12_1","unstructured":"Beguer\u00eda S. andS. 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