{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,17]],"date-time":"2026-09-17T23:44:08Z","timestamp":1789688648163,"version":"4.0.1"},"reference-count":112,"publisher":"The Royal Society","issue":"2078","license":[{"start":{"date-parts":[[2026,9,2]],"date-time":"2026-09-02T00:00:00Z","timestamp":1788307200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,9,2]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Extinction debt and colonization credit describe the delayed impacts of environmental change on biodiversity, yet their significance largely remains unrecognized by those shaping conservation efforts. To address this gap, we conducted a systematic review of 127 empirical studies quantifying these biological lags across taxa, habitats and anthropogenic drivers. Our synthesis revealed geographic biases, with research concentrated in Northern Hemisphere terrestrial ecosystems, alongside taxonomic and habitat biases that continue to limit our understanding of these dynamics. Evaluation of anthropogenic drivers revealed a similarly skewed research effort, with habitat fragmentation dominating extinction debt studies and land-use change leading colonization credit research. By synthesizing proposed conservation strategies, we identified pathways for integrating time-lagged ecological responses into conservation frameworks. Despite the conceptual importance of these lags, most empirical efforts remain focused on isolated taxonomic groups, with limited consideration of how debts and credits propagate through communities or influence ecosystem services. This narrow focus restricts the utility of current findings for broad-scale conservation. We recommend that future research prioritizes the mechanisms driving lags across understudied ecosystems and environmental contexts. Crucially, conservation interventions must move beyond static biodiversity snapshots and explicitly account for extinction debt and colonization credit to ensure long-term ecosystem resilience.<\/jats:p>","DOI":"10.1098\/rspb.2026.0403","type":"journal-article","created":{"date-parts":[[2026,9,2]],"date-time":"2026-09-02T05:22:26Z","timestamp":1788326546000},"source":"Crossref","is-referenced-by-count":1,"title":["Extinction debt and colonization credit in conservation science"],"prefix":"10.1098","volume":"293","author":[{"ORCID":"https:\/\/orcid.org\/0009-0008-8298-2429","authenticated-orcid":false,"given":"Katy L.","family":"Rylance","sequence":"first","affiliation":[{"name":"School of Biology, Faculty of Biological Science, University of Leeds 1 , Leeds, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Steven","family":"Sait","sequence":"additional","affiliation":[{"name":"School of Biology, Faculty of Biological Science, University of Leeds 1 , Leeds, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1363-3571","authenticated-orcid":false,"given":"Maria","family":"Beger","sequence":"additional","affiliation":[{"name":"School of Biology, Faculty of Biological Science, University of Leeds 1 , Leeds, UK"},{"name":"Centre for Biodiversity and Conservation Science, University of Queensland 2 , Brisbane, QLD 4072, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"175","published-online":{"date-parts":[[2026,9,2]]},"reference":[{"key":"2026090201222111500_RSPB20260403C1","first-page":"727","article-title":"What is conservation biology? A new synthetic discipline addresses the dynamics and problems of perturbed species, communities, and ecosystems","volume":"35","author":"Soul\u00e9","year":"1985","journal-title":"Bioscience"},{"key":"2026090201222111500_RSPB20260403C2","doi-asserted-by":"crossref","first-page":"10497","DOI":"10.1073\/pnas.1702111114","article-title":"Waiting can be an optimal conservation strategy, even in a crisis discipline","volume":"114","author":"Iacona","year":"2017","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2026090201222111500_RSPB20260403C4","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1016\/j.tree.2024.12.002","article-title":"Advances in systematic conservation planning to meet global biodiversity goals","volume":"40","author":"Giakoumi","year":"2025","journal-title":"Trends Ecol. Evol."},{"key":"2026090201222111500_RSPB20260403C6","doi-asserted-by":"crossref","first-page":"1877","DOI":"10.1111\/ele.14303","article-title":"Accelerating local extinction associated with very recent climate change","volume":"26","author":"Holzmann","year":"2023","journal-title":"Ecol. Lett."},{"key":"2026090201222111500_RSPB20260403C8","volume-title":"The theory of island biogeography","author":"MacArthur","year":"1967"},{"key":"2026090201222111500_RSPB20260403C9","doi-asserted-by":"crossref","first-page":"563","DOI":"10.1111\/gcb.13976","article-title":"Lags in the response of mountain plant communities to climate change","volume":"24","author":"Alexander","year":"2018","journal-title":"Glob. Change Biol."},{"key":"2026090201222111500_RSPB20260403C10","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1038\/371065a0","article-title":"Habitat destruction and the extinction debt","volume":"371","author":"Tilman","year":"1994","journal-title":"Nature"},{"key":"2026090201222111500_RSPB20260403C11","doi-asserted-by":"crossref","first-page":"3199","DOI":"10.1073\/pnas.69.11.3199","article-title":"Biogeographic kinetics: estimation of relaxation times for avifaunas of southwest Pacific islands","volume":"69","author":"Diamond","year":"1972","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2026090201222111500_RSPB20260403C12","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1111\/j.1752-4598.2009.00075.x","article-title":"Colonisation credit in recent wet heathland butterfly communities","volume":"3","author":"Cristofoli","year":"2010","journal-title":"Insect Conserv. Divers."},{"key":"2026090201222111500_RSPB20260403C13","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1016\/j.tree.2009.10.001","article-title":"Balancing biodiversity in a changing environment: extinction debt, immigration credit and species turnover","volume":"25","author":"Jackson","year":"2010","journal-title":"Trends Ecol. Evol."},{"key":"2026090201222111500_RSPB20260403C14","doi-asserted-by":"crossref","first-page":"564","DOI":"10.1016\/j.tree.2009.04.011","article-title":"Extinction debt: a challenge for biodiversity conservation","volume":"24","author":"Kuussaari","year":"2009","journal-title":"Trends Ecol. Evol."},{"key":"2026090201222111500_RSPB20260403C15","doi-asserted-by":"crossref","first-page":"1973","DOI":"10.1111\/ecog.04740","article-title":"Understanding extinction debts: spatio-temporal scales, mechanisms and a roadmap for future research","volume":"42","author":"Ludmilla","year":"2019","journal-title":"Ecography"},{"key":"2026090201222111500_RSPB20260403C16","doi-asserted-by":"crossref","first-page":"383","DOI":"10.1002\/1438-390X.12009","article-title":"Relative importance of landscape and species characteristics on extinction debt, immigration credit and relaxation time after habitat turnover","volume":"61","author":"Lalech\u00e8re","year":"2019","journal-title":"Popul. Ecol."},{"key":"2026090201222111500_RSPB20260403C17","doi-asserted-by":"crossref","first-page":"823","DOI":"10.1007\/s00442-015-3377-4","article-title":"Assessing coexisting plant extinction debt and colonization credit in a grassland-forest change gradient","volume":"179","author":"Bagaria","year":"2015","journal-title":"Oecologia"},{"key":"2026090201222111500_RSPB20260403C18","doi-asserted-by":"crossref","first-page":"1460","DOI":"10.1890\/06-1032.1","article-title":"Minimum viable metapopulation size, extinction debt, and the conservation of a declining species","volume":"17","author":"Bulman","year":"2007","journal-title":"Ecol. Appl."},{"key":"2026090201222111500_RSPB20260403C20","doi-asserted-by":"crossref","first-page":"1619","DOI":"10.1016\/j.biocon.2011.02.013","article-title":"Plant species extinction debt in a temperate biodiversity hotspot: community, species and functional traits approaches","volume":"144","author":"Piqueray","year":"2011","journal-title":"Biol. Conserv."},{"key":"2026090201222111500_RSPB20260403C21","doi-asserted-by":"crossref","first-page":"228","DOI":"10.1126\/science.1219013","article-title":"Extinction debt and windows of conservation opportunity in the Brazilian Amazon","volume":"337","author":"Wearn","year":"2012","journal-title":"Science"},{"key":"2026090201222111500_RSPB20260403C22","doi-asserted-by":"crossref","first-page":"178","DOI":"10.1080\/10383441.2017.1355873","article-title":"Halting and restoring species loss: incorporating the concepts of extinction debt, ecological trap and dark diversity into conservation and restoration law","volume":"26","author":"Cliquet","year":"2017","journal-title":"Griffith Law Rev."},{"key":"2026090201222111500_RSPB20260403C23","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1016\/j.biocon.2014.12.002","article-title":"Colonization credit of post-agricultural forest patches in NE Germany remains 130-230 years after reforestation","volume":"182","author":"Naaf","year":"2015","journal-title":"Biol. Conserv."},{"key":"2026090201222111500_RSPB20260403C24","doi-asserted-by":"crossref","first-page":"4293","DOI":"10.1038\/s41467-019-12343-x","article-title":"Extinction debts and colonization credits of non-forest plants in the European Alps","volume":"10","author":"Rumpf","year":"2019","journal-title":"Nat. Commun."},{"key":"2026090201222111500_RSPB20260403C25","doi-asserted-by":"crossref","first-page":"1259","DOI":"10.1111\/ecog.05926","article-title":"Sampling biases shape our view of the natural world","volume":"44","author":"Hughes","year":"2021","journal-title":"Ecography"},{"key":"2026090201222111500_RSPB20260403C27","doi-asserted-by":"crossref","first-page":"172","DOI":"10.1016\/j.biocon.2012.08.034","article-title":"Using assisted colonisation to conserve biodiversity and restore ecosystem function under climate change","volume":"157","author":"Lunt","year":"2013","journal-title":"Biol. Conserv."},{"key":"2026090201222111500_RSPB20260403C29","doi-asserted-by":"crossref","first-page":"640","DOI":"10.1111\/brv.12816","article-title":"The Sixth Mass Extinction: fact, fiction or speculation?","volume":"97","author":"Cowie","year":"2022","journal-title":"Biol. Rev."},{"key":"2026090201222111500_RSPB20260403C32","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1007\/s10531-009-9761-9","article-title":"Re-assessing current extinction rates","volume":"19","author":"Stork","year":"2010","journal-title":"Biodivers. Conserv."},{"key":"2026090201222111500_RSPB20260403C33","doi-asserted-by":"crossref","first-page":"e13","DOI":"10.1017\/ext.2023.11","article-title":"A review of recent and future marine extinctions","volume":"1","author":"del Monte-Luna","year":"2023","journal-title":"Camb. Prism Extinct."},{"key":"2026090201222111500_RSPB20260403C34","doi-asserted-by":"crossref","first-page":"485","DOI":"10.1007\/s00227-010-1596-0","article-title":"Marine extinctions and conservation","volume":"158","author":"Briggs","year":"2011","journal-title":"Mar. Biol."},{"key":"2026090201222111500_RSPB20260403C36","doi-asserted-by":"crossref","first-page":"286","DOI":"10.1016\/j.tree.2024.11.007","article-title":"The ecology of plant extinctions","volume":"40","author":"Corlett","year":"2025","journal-title":"Trends Ecol. Evol."},{"key":"2026090201222111500_RSPB20260403C37","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1016\/j.tree.2015.05.002","article-title":"Delayed biodiversity change: no time to waste","volume":"30","author":"Essl","year":"2015","journal-title":"Trends Ecol. Evol."},{"key":"2026090201222111500_RSPB20260403C38","doi-asserted-by":"crossref","first-page":"70","DOI":"10.1007\/s40823-019-00040-w","article-title":"Temporal lag in ecological responses to landscape change: where are we now?","volume":"4","author":"Lira","year":"2019","journal-title":"Curr. Landsc. Ecol. Rep."},{"key":"2026090201222111500_RSPB20260403C39","doi-asserted-by":"crossref","first-page":"1647","DOI":"10.1111\/j.1523-1739.2006.00485.x","article-title":"Guidelines for systematic review in conservation and environmental management","volume":"20","author":"Pullin","year":"2006","journal-title":"Conserv. Biol."},{"key":"2026090201222111500_RSPB20260403C40","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1186\/s13750-017-0099-6","article-title":"Systematic searching for environmental evidence using multiple tools and sources","volume":"6","author":"Livoreil","year":"2017","journal-title":"Environ. Evid."},{"key":"2026090201222111500_RSPB20260403C41","doi-asserted-by":"crossref","first-page":"515","DOI":"10.1111\/jvs.12501","article-title":"Interactive effects of land-use change and distance-to-edge on the distribution of species in plant communities at the forest-grassland interface","volume":"28","author":"Burst","year":"2017","journal-title":"J. Veg. Sci."},{"key":"2026090201222111500_RSPB20260403C43","first-page":"44","article-title":"Evidence and mapping of extinction debts for global forest-dwelling reptiles, amphibians and mammals","volume":"7","author":"Chen","year":"2017","journal-title":"Sci. Rep."},{"key":"2026090201222111500_RSPB20260403C44","doi-asserted-by":"crossref","first-page":"1183","DOI":"10.1046\/j.1523-1739.1999.98433.x","article-title":"Predicting the pattern of decline of African primate diversity: an extinction debt from historical deforestation","volume":"13","author":"Cowlishaw","year":"1999","journal-title":"Conserv. Biol."},{"key":"2026090201222111500_RSPB20260403C45","doi-asserted-by":"crossref","first-page":"e0166409","DOI":"10.1371\/journal.pone.0166409","article-title":"Extinction debt and colonizer credit on a habitat perturbed fishing bank","volume":"11","author":"Duplisea","year":"2016","journal-title":"PLoS ONE"},{"key":"2026090201222111500_RSPB20260403C46","doi-asserted-by":"crossref","first-page":"324","DOI":"10.1038\/s41559-021-01653-3","article-title":"Widespread extinction debts and colonization credits in United States breeding bird communities","volume":"6","author":"Haddou","year":"2022","journal-title":"Nat. Ecol. Evol."},{"key":"2026090201222111500_RSPB20260403C47","doi-asserted-by":"crossref","first-page":"e13134","DOI":"10.1111\/jvs.13134","article-title":"Recruitment credit cannot compensate for extinction debt in a degraded dry afromontane forest","volume":"33","author":"Hishe","year":"2022","journal-title":"J. Veg. Sci."},{"key":"2026090201222111500_RSPB20260403C48","doi-asserted-by":"crossref","first-page":"735","DOI":"10.1007\/s10531-016-1271-y","article-title":"Paying the colonization credit: converging plant species richness in ancient and post-agricultural forests in NE Germany over five decades","volume":"26","author":"Kolk","year":"2017","journal-title":"Biodivers. Conserv."},{"key":"2026090201222111500_RSPB20260403C49","doi-asserted-by":"crossref","first-page":"1218","DOI":"10.1111\/1365-2664.13074","article-title":"Mapping extinction debt highlights conservation opportunities for birds and mammals in the South American Chaco","volume":"55","author":"Semper-Pascual","year":"2018","journal-title":"J. Appl. Ecol."},{"key":"2026090201222111500_RSPB20260403C51","doi-asserted-by":"crossref","first-page":"1705","DOI":"10.1111\/2041-210X.13654","article-title":"A practical guide to question formation, systematic searching and study screening for literature reviews in ecology and evolution","volume":"12","author":"Foo","year":"2021","journal-title":"Methods Ecol. Evol."},{"key":"2026090201222111500_RSPB20260403C52","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1002\/jrsm.1378","article-title":"Which academic search systems are suitable for systematic reviews or meta-analyses? Evaluating retrieval qualities of Google Scholar, PubMed, and 26 other resources","volume":"11","author":"Gusenbauer","year":"2020","journal-title":"Res. Synth. Methods"},{"key":"2026090201222111500_RSPB20260403C54","volume-title":"Publish or perish","author":"Harzing","year":"2007"},{"key":"2026090201222111500_RSPB20260403C55","first-page":"230","article-title":"Year Google Scholar's ranking algorithm: an introductory overview","volume-title":"Proc. 12th Int. Conf. Scientometrics Informetrics (ISSI\u201909)","author":"Beel","year":"2009"},{"key":"2026090201222111500_RSPB20260403C56","volume-title":"R: a language and environment for statistical computing","author":"Team","year":"2024"},{"key":"2026090201222111500_RSPB20260403C57","doi-asserted-by":"crossref","first-page":"1686","DOI":"10.21105\/joss.01686","article-title":"Welcome to the Tidyverse","volume":"4","author":"Wickham","year":"2019","journal-title":"J. Open Source Softw."},{"key":"2026090201222111500_RSPB20260403C58","doi-asserted-by":"crossref","first-page":"704","DOI":"10.1086\/508296","article-title":"On testing the competition-colonization trade-off in a multispecies assemblage","volume":"168","author":"Cadotte","year":"2006","journal-title":"Am. Nat."},{"key":"2026090201222111500_RSPB20260403C59","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1046\/j.1523-1739.1997.95381.x","article-title":"Extinction debts and risks faced by abundant species","volume":"11","author":"McCarthy","year":"1997","journal-title":"Conserv. Biol."},{"key":"2026090201222111500_RSPB20260403C60","doi-asserted-by":"crossref","first-page":"299","DOI":"10.3354\/meps134299","article-title":"Modelling coral reef biodiversity and habitat destruction","volume":"134","author":"Stone","year":"1996","journal-title":"Mar. Ecol. Prog. Ser."},{"key":"2026090201222111500_RSPB20260403C61","first-page":"297","article-title":"Biodiversity hotspots: a shortcut for a more complicated concept","volume":"3","author":"Marchese","year":"2015","journal-title":"Glob. Ecol. Conserv."},{"key":"2026090201222111500_RSPB20260403C62","author":"IUCNb","year":"2024","journal-title":"The IUCN red list of threatened species"},{"key":"2026090201222111500_RSPB20260403C64","doi-asserted-by":"crossref","first-page":"32","DOI":"10.5751\/ES-01609-110232","article-title":"Special feature on scenarios for ecosystem services","volume":"11","author":"Carpenter","year":"2006","journal-title":"Ecol Soc"},{"key":"2026090201222111500_RSPB20260403C66","doi-asserted-by":"crossref","first-page":"eabm9982","DOI":"10.1126\/sciadv.abm9982","article-title":"The direct drivers of recent global anthropogenic biodiversity loss","volume":"8","author":"Jaureguiberry","year":"2022","journal-title":"Sci. Adv."},{"key":"2026090201222111500_RSPB20260403C67","doi-asserted-by":"crossref","DOI":"10.5751\/ES-01826-110229","article-title":"Anthropogenic drivers of ecosystem change: an overview","volume":"11","author":"Nelson","year":"2006","journal-title":"Ecol. Soc."},{"key":"2026090201222111500_RSPB20260403C68","doi-asserted-by":"crossref","first-page":"105219","DOI":"10.1016\/j.landurbplan.2024.105219","article-title":"Evidence of taxonomic but not functional diversity extinction debt in bird assemblages in an urban area in the Cerrado hotspot","volume":"253","author":"Almeida-Gomes","year":"2025","journal-title":"Landsc. Urban Plan"},{"key":"2026090201222111500_RSPB20260403C70","doi-asserted-by":"crossref","first-page":"1211","DOI":"10.1007\/s11756-023-01394-7","article-title":"Biological invasions in the twenty-first century: a global risk","volume":"78","author":"Vantarov\u00e1","year":"2023","journal-title":"Biologia"},{"key":"2026090201222111500_RSPB20260403C72","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1007\/s10980-015-0312-3","article-title":"Habitat fragmentation and biodiversity conservation: key findings and future challenges","volume":"31","author":"Wilson","year":"2016","journal-title":"Landsc. Ecol."},{"key":"2026090201222111500_RSPB20260403C76","doi-asserted-by":"crossref","first-page":"784","DOI":"10.1111\/geb.13246","article-title":"Climate-driven range shifts of montane species vary with elevation","volume":"30","author":"Mamantov","year":"2021","journal-title":"Glob. Ecol. Biogeogr."},{"key":"2026090201222111500_RSPB20260403C77","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1146\/annurev-marine-010419-010916","article-title":"Climate-driven shifts in marine species ranges: scaling from organisms to communities","volume":"12","author":"Pinsky","year":"2020","journal-title":"Ann. Rev. Mar. Sci."},{"key":"2026090201222111500_RSPB20260403C78","doi-asserted-by":"crossref","first-page":"e17408","DOI":"10.1111\/gcb.17408","article-title":"Insufficient and biased representation of species geographic responses to climate change","volume":"30","author":"Parker","year":"2024","journal-title":"Glob. Change Biol."},{"key":"2026090201222111500_RSPB20260403C79","doi-asserted-by":"crossref","first-page":"e3001336","DOI":"10.1371\/journal.pbio.3001336","article-title":"Global and national trends, gaps, and opportunities in documenting and monitoring species distributions","volume":"19","author":"Oliver","year":"2021","journal-title":"PLoS Biol."},{"key":"2026090201222111500_RSPB20260403C80","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1016\/j.biocon.2023.110118","article-title":"The biased distribution of existing information on biodiversity hinders its use in conservation, and we need an integrative approach to act urgently","volume":"283","author":"Garc\u00eda-Rosell\u00f3","year":"2023","journal-title":"Biol. Conserv."},{"key":"2026090201222111500_RSPB20260403C81","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1111\/cobi.13577","article-title":"The challenge of biased evidence in conservation","volume":"35","author":"Christie","year":"2021","journal-title":"Conserv. Biol."},{"key":"2026090201222111500_RSPB20260403C82","doi-asserted-by":"crossref","first-page":"1311","DOI":"10.1038\/s42003-022-04277-w","article-title":"Half-millennium evidence suggests that extinction debts of global vertebrates started in the Second Industrial Revolution","volume":"5","author":"Liao","year":"2022","journal-title":"Commun. Biol."},{"key":"2026090201222111500_RSPB20260403C83","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1016\/j.biocon.2016.04.036","article-title":"Extinction debt on reservoir land-bridge islands","volume":"199","author":"Jones","year":"2016","journal-title":"Biol. Conserv."},{"key":"2026090201222111500_RSPB20260403C84","doi-asserted-by":"crossref","first-page":"741","DOI":"10.1126\/science.adq9512","article-title":"Limited evidence for range shift\u2013driven extinction in mountain biota","volume":"388","author":"Chen","year":"2025","journal-title":"Science"},{"key":"2026090201222111500_RSPB20260403C85","doi-asserted-by":"crossref","first-page":"e13078","DOI":"10.1111\/conl.13078","article-title":"A 150-year Avian extinction debt forewarns a global species crisis and highlights conservation opportunities","volume":"18","author":"Wang","year":"2025","journal-title":"Conserv. Lett."},{"key":"2026090201222111500_RSPB20260403C86","doi-asserted-by":"crossref","first-page":"5484","DOI":"10.3390\/su16135484","article-title":"Predicting the global extinction risk for 6569 species by applying the life cycle impact assessment method to the impact of future land use changes","volume":"16","author":"Liu","year":"2024","journal-title":"Sustainability"},{"key":"2026090201222111500_RSPB20260403C87","doi-asserted-by":"crossref","first-page":"651","DOI":"10.1007\/s10531-009-9725-0","article-title":"The prominence of and biases in biodiversity and ecosystem functioning research","volume":"19","author":"Caliman","year":"2010","journal-title":"Biodivers. Conserv."},{"key":"2026090201222111500_RSPB20260403C89","doi-asserted-by":"crossref","first-page":"666","DOI":"10.1046\/j.1523-1739.2002.00342.x","article-title":"Extinction debt at extinction threshold","volume":"16","author":"Hanski","year":"2002","journal-title":"Conserv Biol."},{"key":"2026090201222111500_RSPB20260403C90","first-page":"14","article-title":"Extinction debt \u2013 time-delayed extinction of species: a phenomenon that has hardly been considered in nature conservation practice","volume":"53","author":"Fartmann","year":"2021","journal-title":"Natschutz Landschplan"},{"key":"2026090201222111500_RSPB20260403C91","doi-asserted-by":"crossref","first-page":"591","DOI":"10.1111\/ddi.12187","article-title":"Extinction debt for plants and flower-visiting insects in landscapes with contrasting land use history","volume":"20","author":"Bommarco","year":"2014","journal-title":"Divers. Distrib."},{"key":"2026090201222111500_RSPB20260403C92","doi-asserted-by":"crossref","first-page":"1031","DOI":"10.1111\/ddi.12590","article-title":"Unpaid extinction debts for endemic plants and invertebrates as a legacy of habitat loss on oceanic islands","volume":"23","author":"Otto","year":"2017","journal-title":"Divers. Distrib."},{"key":"2026090201222111500_RSPB20260403C93","doi-asserted-by":"crossref","first-page":"e12767","DOI":"10.1111\/maec.12767","article-title":"La Ni\u00f1a-related coral death triggers biodiversity loss of associated communities in the Galapagos","volume":"44","author":"Rhoades","year":"2023","journal-title":"Mar. Ecol. Evol. Persp."},{"key":"2026090201222111500_RSPB20260403C94","doi-asserted-by":"crossref","first-page":"21","DOI":"10.2980\/19-1-3460","article-title":"An orchid colonization credit in restored calcareous grasslands","volume":"19","author":"Gijbels","year":"2012","journal-title":"Ecoscience"},{"key":"2026090201222111500_RSPB20260403C95","doi-asserted-by":"crossref","first-page":"1455","DOI":"10.1111\/ddi.12380","article-title":"Conservation in Brazil needs to include non-forest ecosystems","volume":"21","author":"Overbeck","year":"2015","journal-title":"Diver. Distrib"},{"key":"2026090201222111500_RSPB20260403C96","doi-asserted-by":"crossref","first-page":"e13858.","DOI":"10.1111\/rec.13858","article-title":"Grassland conservation and restoration in India: a governance crisis","volume":"31","author":"Lahiri","year":"2023","journal-title":"Restor. Ecol."},{"key":"2026090201222111500_RSPB20260403C97","doi-asserted-by":"crossref","first-page":"1491","DOI":"10.1007\/s13280-025-02155-3","article-title":"Seven ways to prevent biomism","volume":"54","author":"Silveira","year":"2025","journal-title":"Ambio"},{"key":"2026090201222111500_RSPB20260403C98","first-page":"328","article-title":"Forest-biased terminology does not help to include open ecosystems in conservation policies","volume":"22","author":"Overbeck","year":"2024","journal-title":"Persp. Ecol. Conserv."},{"key":"2026090201222111500_RSPB20260403C100","doi-asserted-by":"crossref","first-page":"1636","DOI":"10.1111\/nph.19151","article-title":"Global hotspots of plant phylogenetic diversity","volume":"240","author":"Tietje","year":"2023","journal-title":"New Phytol."},{"key":"2026090201222111500_RSPB20260403C101","doi-asserted-by":"crossref","first-page":"1280","DOI":"10.1126\/science.1067728","article-title":"Marine biodiversity hotspots and conservation priorities for tropical reefs","volume":"295","author":"Roberts","year":"2002","journal-title":"Science"},{"key":"2026090201222111500_RSPB20260403C102","doi-asserted-by":"crossref","first-page":"1205","DOI":"10.1111\/geb.13681","article-title":"Regional species gains outpace losses across North American continental shelf regions","volume":"32","author":"Kitchel","year":"2023","journal-title":"Glob. Ecol. Biogeogr."},{"key":"2026090201222111500_RSPB20260403C103","first-page":"100082","article-title":"Global trends and biases in biodiversity conservation research","volume":"1","author":"Caldwell","year":"2024","journal-title":"Cell Rep. Sustain."},{"key":"2026090201222111500_RSPB20260403C105","doi-asserted-by":"crossref","first-page":"152","DOI":"10.1016\/j.gloenvcha.2014.04.002","article-title":"Changes in the global value of ecosystem services","volume":"26","author":"Costanza","year":"2014","journal-title":"Glob. Environ. Chang."},{"key":"2026090201222111500_RSPB20260403C107","doi-asserted-by":"crossref","first-page":"330","DOI":"10.1007\/978-1-4613-0059-5_18","volume-title":"Applying landscape ecology in biological conservation","author":"Nagelkerke","year":"2002"},{"key":"2026090201222111500_RSPB20260403C108","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1016\/j.tree.2013.01.010","article-title":"The mechanisms causing extinction debts","volume":"28","author":"Hylander","year":"2013","journal-title":"Trends Ecol. Evol."},{"key":"2026090201222111500_RSPB20260403C109","doi-asserted-by":"crossref","first-page":"e3547","DOI":"10.7717\/peerj.3547","article-title":"A demographic history of a population of howler monkeys (Alouatta palliata) living in a fragmented landscape in Mexico","volume":"5","author":"Jurgi","year":"2017","journal-title":"PeerJ"},{"key":"2026090201222111500_RSPB20260403C110","doi-asserted-by":"crossref","first-page":"432","DOI":"10.1111\/acv.12648","article-title":"Evaluating extinction debt in fragmented forests: the rapid recovery of a critically endangered primate","volume":"24","author":"Alcocer-Rodr\u00edguez","year":"2021","journal-title":"Anim. Conserv."},{"key":"2026090201222111500_RSPB20260403C111","doi-asserted-by":"crossref","first-page":"277","DOI":"10.1111\/acv.12979","article-title":"Delayed responses and extinction debt: an opportunity for the conservation of Chaco Serrano forest birds","volume":"28","author":"Silvetti","year":"2025","journal-title":"Anim. Conserv."},{"key":"2026090201222111500_RSPB20260403C112","doi-asserted-by":"crossref","first-page":"156","DOI":"10.1034\/j.1600-0706.2002.980116.x","article-title":"Functional redundancy in ecology and conservation","volume":"98","author":"Rosenfeld","year":"2002","journal-title":"Oikos"},{"key":"2026090201222111500_RSPB20260403C113","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1086\/327796","article-title":"The ecological relations of the vegetation on the Sand Dunes of Lake Michigan. Part I.-geographical relations of the dune floras","volume":"27","author":"Cowles","year":"1899","journal-title":"Bot. Gaz."},{"key":"2026090201222111500_RSPB20260403C115","doi-asserted-by":"crossref","first-page":"2195","DOI":"10.1111\/1365-2664.13975","article-title":"Immigration credit of temperate forest herbs in fragmented landscapes-implications for restoration of habitat connectivity","volume":"58","author":"Brunet","year":"2021","journal-title":"J. Appl. Ecol."},{"key":"2026090201222111500_RSPB20260403C116","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1038\/s41559-019-1058-0","article-title":"Ocean warming compresses the three-dimensional habitat of marine life","volume":"4","author":"Jorda","year":"2020","journal-title":"Nat. Ecol. Evol."},{"key":"2026090201222111500_RSPB20260403C118","doi-asserted-by":"crossref","first-page":"456","DOI":"10.1666\/07034.1","article-title":"Press-pulse: a general theory of mass extinction?","volume":"34","author":"Arens","year":"2008","journal-title":"Paleobiology"},{"key":"2026090201222111500_RSPB20260403C121","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1016\/j.tree.2020.10.002","article-title":"Using climatic credits to pay the climatic debt","volume":"36","author":"Vaughan","year":"2021","journal-title":"Trends Ecol. Evol."},{"key":"2026090201222111500_RSPB20260403C122","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1177\/0309133315598713","article-title":"Habitat connectivity in biodiversity conservation: a review of recent studies and applications","volume":"40","author":"Correa Ayram","year":"2016","journal-title":"Prog. Phys. Geogr."},{"key":"2026090201222111500_RSPB20260403C123","first-page":"1","article-title":"The role of landscape connectivity in planning and implementing conservation and restoration priorities","volume":"16","author":"Rudnick","year":"2012","journal-title":"Issues Ecol."},{"key":"2026090201222111500_RSPB20260403C124","first-page":"271","article-title":"Year extinction debt and species credit in boreal forests: modelling the consequences of different approaches to biodiversity conservation","volume":"2000","author":"Hanski","year":"2000","journal-title":"Annales Zoologici Fennici"},{"key":"2026090201222111500_RSPB20260403C125","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1016\/j.jeem.2008.10.002","article-title":"Optimal conservation, extinction debt, and the augmented quasi-option value","volume":"58","author":"Leroux","year":"2009","journal-title":"J Environ. Econ. Manage."},{"key":"2026090201222111500_RSPB20260403C126","doi-asserted-by":"crossref","first-page":"2149","DOI":"10.1111\/1365-2664.13717","article-title":"Interactive persistent effects of past land-cover and its trajectory on tropical freshwater biodiversity","volume":"57","author":"Santos","year":"2020","journal-title":"J. Appl. Ecol."},{"key":"2026090201222111500_RSPB20260403C129","doi-asserted-by":"crossref","first-page":"266","DOI":"10.1071\/PC040266","article-title":"The role of connectivity in Australian conservation","volume":"10","author":"Soul\u00e9","year":"2004","journal-title":"Pac. Conserv. Biol."},{"key":"2026090201222111500_RSPB20260403C130","doi-asserted-by":"crossref","first-page":"1478","DOI":"10.1126\/science.aax8992","article-title":"Ongoing accumulation of plant diversity through habitat connectivity in an 18-year experiment","volume":"365","author":"Damschen","year":"2019","journal-title":"Science"},{"key":"2026090201222111500_RSPB20260403C131","doi-asserted-by":"crossref","first-page":"e03865","DOI":"10.1002\/ecs2.3865","article-title":"Connectivity affects species turnover in soil microarthropod communities during Mediterranean forest establishment","volume":"12","author":"Doblas-Miranda","year":"2021","journal-title":"Ecosphere"},{"key":"2026090201222111500_RSPB20260403C132","doi-asserted-by":"crossref","first-page":"e0206738","DOI":"10.1371\/journal.pone.0206738","article-title":"Unprecedented plant species loss after a decade in fragmented subtropical Chaco Serrano forests","volume":"13","author":"Aguilar","year":"2018","journal-title":"PLoS ONE"},{"key":"2026090201222111500_RSPB20260403C134","doi-asserted-by":"crossref","first-page":"1110","DOI":"10.1111\/j.1523-1739.2004.00083.x","article-title":"Extinction debt of protected areas in developing landscapes","volume":"18","author":"Carroll","year":"2004","journal-title":"Conserv. Biol."},{"key":"2026090201222111500_RSPB20260403C135","unstructured":"IUCNa\n          \n          2024\n          Guidelines for using the IUCN Red List categories and criteria. Version 16.\n          https:\/\/www.iucnredlist.org\/resources\/redlistguidelines (accessed 7 February 2026)."},{"key":"2026090201222111500_RSPB20260403C136","doi-asserted-by":"crossref","first-page":"20150843","DOI":"10.1098\/rsbl.2015.0843","article-title":"Clarifying misconceptions of extinction risk assessment with the IUCN Red List","volume":"12","author":"Collen","year":"2016","journal-title":"Biol. Lett."},{"key":"2026090201222111500_RSPB20260403C137","doi-asserted-by":"crossref","first-page":"110880","DOI":"10.1016\/j.biocon.2024.110880","article-title":"2025 IUCN Red List criteria fail to recognise most threatened and extinct species","volume":"301","author":"Edgar","journal-title":"Biol. Conserv."},{"key":"2026090201222111500_RSPB20260403C138","doi-asserted-by":"crossref","first-page":"e13942","DOI":"10.1111\/cobi.13942","article-title":"Time-lagged effects of habitat fragmentation on terrestrial mammals in Madagascar","volume":"36","author":"Broekman","year":"2022","journal-title":"Conserv. Biol."},{"key":"2026090201222111500_RSPB20260403C139","doi-asserted-by":"crossref","first-page":"e0147909","DOI":"10.1371\/journal.pone.0147909","article-title":"Time-lag in responses of birds to Atlantic forest fragmentation: restoration opportunity and urgency","volume":"11","author":"Uezu","year":"2016","journal-title":"PLoS ONE"},{"key":"2026090201222111500_RSPB20260403C142","doi-asserted-by":"crossref","first-page":"109932","DOI":"10.1016\/j.biocon.2023.109932","article-title":"The application of assisted migration as a climate change adaptation tactic: an evidence map and synthesis","volume":"280","author":"Twardek","year":"2023","journal-title":"Biol. Conserv."},{"key":"2026090201222111500_RSPB20260403C144","doi-asserted-by":"crossref","first-page":"63","DOI":"10.3897\/natureconservation.45.72554","article-title":"Risks and opportunities of assisted colonization: the perspectives of experts","volume":"45","author":"Kracke","year":"2021","journal-title":"Nat. Conserv."},{"key":"2026090201222111500_RSPB20260403C145","article-title":"Extinction debt and colonization credit in conservation science","author":"Rylance","year":"2026"}],"container-title":["Proceedings B"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/royalsocietypublishing.org\/rspb\/article-pdf\/doi\/10.1098\/rspb.2026.0403\/6163752\/rspb.2026.0403.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/royalsocietypublishing.org\/rspb\/article-pdf\/doi\/10.1098\/rspb.2026.0403\/6163752\/rspb.2026.0403.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,2]],"date-time":"2026-09-02T05:22:43Z","timestamp":1788326563000},"score":1,"resource":{"primary":{"URL":"https:\/\/royalsocietypublishing.org\/rspb\/article\/293\/2078\/20260403\/483173\/Extinction-debt-and-colonization-credit-in"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,2]]},"references-count":112,"journal-issue":{"issue":"2078","published-print":{"date-parts":[[2026,9,2]]}},"URL":"https:\/\/doi.org\/10.1098\/rspb.2026.0403","relation":{},"ISSN":["1471-2954"],"issn-type":[{"value":"1471-2954","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,2]]},"article-number":"20260403"}}