{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T03:15:33Z","timestamp":1790565333905,"version":"4.1.0"},"reference-count":111,"publisher":"The Royal Society","issue":"2075","license":[{"start":{"date-parts":[[2026,7,29]],"date-time":"2026-07-29T00:00:00Z","timestamp":1785283200000},"content-version":"vor","delay-in-days":28,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,7,29]],"date-time":"2026-07-29T00:00:00Z","timestamp":1785283200000},"content-version":"vor","delay-in-days":28,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100004189","name":"Max-Planck-Gesellschaft","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100004189","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000288","name":"Royal Society","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100000288","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000848","name":"University of Edinburgh","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100000848","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,7,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Although strong evidence links social connectedness to improved fitness, there is pronounced among-individual variation in social behaviour. This variation may be maintained if different types of social relationships (i.e. having many versus strongly connected partners) have environment-dependent fitness benefits, but this has rarely been tested. We applied network analysis to 37 years of social association data in female Soay sheep to test the relative importance of relationship quantity (mean group size, network degree and participation coefficient) and quality (mean strength and social differentiation) for overwinter survival. We find that both relationship quantity and quality predict survival the following winter. Crucially, the relative importance of relationship quantity and quality depends on weather conditions: harsh conditions favour quantity (specifically large mean group sizes), likely owing to thermoregulatory benefits, while benign conditions favour quality (specifically mean association strength), possibly because conspecific familiarity reduces conflict. Our second measure of relationship quality\u2014social differentiation\u2014had a positive association with survival that was not environment-dependent, illustrating the value of strong, stable relationships in fluid societies. These findings advance our understanding of social benefits in fission\u2013fusion systems and show that changing environmental conditions may be an important mechanism maintaining variation in social behaviour in natural populations.<\/jats:p>","DOI":"10.1098\/rspb.2025.2821","type":"journal-article","created":{"date-parts":[[2026,7,29]],"date-time":"2026-07-29T00:20:42Z","timestamp":1785284442000},"source":"Crossref","is-referenced-by-count":1,"title":["Environment-dependent benefits of sociality in Soay sheep"],"prefix":"10.1098","volume":"293","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4205-7278","authenticated-orcid":false,"given":"Erin R.","family":"Siracusa","sequence":"first","affiliation":[{"name":"University of Exeter, 1","place":["Exeter EX4 4QG,, UK"],"department":["Centre for Research in Animal Behaviour,"]},{"name":"The University of Edinburgh School of Biological Sciences, 2","place":["Edinburgh EH9 3FL,, UK"],"department":["Institute of Ecology and Evolution,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2206-2099","authenticated-orcid":false,"given":"Xavier","family":"Bal","sequence":"additional","affiliation":[{"name":"The University of Edinburgh School of Biological Sciences, 2","place":["Edinburgh EH9 3FL,, UK"],"department":["Institute of Ecology and Evolution,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2474-6125","authenticated-orcid":false,"given":"Delphine","family":"De Moor","sequence":"additional","affiliation":[{"name":"Max-Planck-Institute for Evolutionary Anthropology, 3","place":["04103 Leipzig,, Germany"],"department":["Department of Primate Behavior and Evolution,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6260-2662","authenticated-orcid":false,"given":"Gregory F.","family":"Albery","sequence":"additional","affiliation":[{"name":"Trinity College Dublin, 4","place":["Dublin D02 PN40,, Ireland"],"department":["School of Natural Sciences,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ross","family":"Beattie","sequence":"additional","affiliation":[{"name":"The University of Edinburgh School of Biological Sciences, 2","place":["Edinburgh EH9 3FL,, UK"],"department":["Institute of Ecology and Evolution,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0996-0262","authenticated-orcid":false,"given":"Sanjana","family":"Ravindran","sequence":"additional","affiliation":[{"name":"The University of Edinburgh School of Biological Sciences, 2","place":["Edinburgh EH9 3FL,, UK"],"department":["Institute of Ecology and Evolution,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ellis","family":"Wiersma","sequence":"additional","affiliation":[{"name":"The University of Edinburgh School of Biological Sciences, 2","place":["Edinburgh EH9 3FL,, UK"],"department":["Institute of Ecology and Evolution,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Amelie","family":"Renwick Wilson","sequence":"additional","affiliation":[{"name":"The University of Edinburgh School of Biological Sciences, 2","place":["Edinburgh EH9 3FL,, UK"],"department":["Institute of Ecology and Evolution,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Josephine M.","family":"Pemberton","sequence":"additional","affiliation":[{"name":"The University of Edinburgh School of Biological Sciences, 2","place":["Edinburgh EH9 3FL,, UK"],"department":["Institute of Ecology and Evolution,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9985-0317","authenticated-orcid":false,"given":"Daniel H.","family":"Nussey","sequence":"additional","affiliation":[{"name":"The University of Edinburgh School of Biological Sciences, 2","place":["Edinburgh EH9 3FL,, UK"],"department":["Institute of Ecology and Evolution,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Matthew J.","family":"Silk","sequence":"additional","affiliation":[{"name":"The University of Edinburgh School of Biological Sciences, 2","place":["Edinburgh EH9 3FL,, UK"],"department":["Institute of Ecology and Evolution,"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"175","published-online":{"date-parts":[[2026,7,29]]},"reference":[{"key":"2026072820203354300_R1","doi-asserted-by":"crossref","first-page":"326","DOI":"10.1016\/j.tree.2014.04.002","article-title":"The evolutionary and ecological consequences of animal social networks: emerging issues","volume":"29","author":"Kurvers","year":"2014","journal-title":"Trends Ecol. Evol."},{"key":"2026072820203354300_R2","doi-asserted-by":"crossref","DOI":"10.1007\/s00265-021-03055-8","article-title":"The role of social structure and dynamics in the maintenance of endemic disease","volume":"75","author":"Silk","year":"2021","journal-title":"Behav. Ecol. Sociobiol."},{"key":"2026072820203354300_R3","doi-asserted-by":"crossref","first-page":"879","DOI":"10.1098\/rsif.2007.1100","article-title":"When individual behaviour matters: homogeneous and network models in epidemiology","volume":"4","author":"Bansal","year":"2007","journal-title":"J. R. Soc. Interface"},{"key":"2026072820203354300_R4","doi-asserted-by":"crossref","DOI":"10.1098\/rsos.181493","article-title":"Evolution of non-kin cooperation: social assortment by cooperative phenotype in guppies","volume":"6","author":"Brask","year":"2019","journal-title":"R. Soc. Open Sci."},{"key":"2026072820203354300_R5","doi-asserted-by":"crossref","first-page":"546","DOI":"10.1111\/j.1461-0248.2011.01615.x","article-title":"Co-evolution of behaviour and social network structure promotes human cooperation: cooperation and self-organization in networks","volume":"14","author":"Fehl","year":"2011","journal-title":"Ecol. Lett."},{"key":"2026072820203354300_R6","doi-asserted-by":"crossref","first-page":"4199","DOI":"10.1098\/rspb.2012.1591","article-title":"Social networks predict patch discovery in a wild population of songbirds","volume":"279","author":"Aplin","year":"2012","journal-title":"Proc. R. Soc. B"},{"key":"2026072820203354300_R7","doi-asserted-by":"crossref","DOI":"10.1038\/s42003-020-01597-7","article-title":"Causal evidence for the adaptive benefits of social foraging in the wild","volume":"4","author":"Snijders","year":"2021","journal-title":"Commun. Biol."},{"key":"2026072820203354300_R8","doi-asserted-by":"crossref","first-page":"475","DOI":"10.1002\/ece3.1377","article-title":"Social effects on foraging behavior and success depend on local environmental conditions","volume":"5","author":"Marshall","year":"2015","journal-title":"Ecol. Evol."},{"key":"2026072820203354300_R9","doi-asserted-by":"crossref","DOI":"10.1098\/rspb.2017.1313","article-title":"Mortality risk and social network position in resident killer whales: sex differences and the importance of resource abundance","volume":"284","author":"Ellis","year":"2017","journal-title":"Proc. R. Soc. B"},{"key":"2026072820203354300_R10","doi-asserted-by":"crossref","first-page":"E80","DOI":"10.1086\/655216","article-title":"Structure of social networks in a passerine bird: consequences for sexual selection and the evolution of mating strategies","volume":"176","author":"Oh","year":"2010","journal-title":"Am. Nat."},{"key":"2026072820203354300_R11","doi-asserted-by":"crossref","first-page":"801","DOI":"10.1016\/j.anbehav.2007.06.017","article-title":"Not only mate choice matters: fitness consequences of social partner choice in female house mice","volume":"75","author":"Weidt","year":"2008","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R12","doi-asserted-by":"crossref","DOI":"10.1126\/science.aax9553","article-title":"Social determinants of health and survival in humans and other animals","volume":"368","author":"Snyder-Mackler","year":"2020","journal-title":"Science"},{"key":"2026072820203354300_R13","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1111\/j.1420-9101.2011.02411.x","article-title":"Fitness consequences of social network position in a wild population of forked fungus beetles (Bolitotherus cornutus)","volume":"25","author":"Formica","year":"2012","journal-title":"J. Evol. Biol."},{"key":"2026072820203354300_R14","doi-asserted-by":"crossref","DOI":"10.1098\/rspb.2018.0130","article-title":"Stable social relationships between unrelated females increase individual fitness in a cooperative bird","volume":"285","author":"Riehl","year":"2018","journal-title":"Proc. R. Soc. B"},{"key":"2026072820203354300_R15","doi-asserted-by":"crossref","first-page":"10910","DOI":"10.1073\/pnas.0701159104","article-title":"Predicting fate from early connectivity in a social network","volume":"104","author":"McDonald","year":"2007","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2026072820203354300_R16","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1016\/j.anbehav.2019.12.013","article-title":"More social female yellow-bellied marmots, Marmota flaviventer, have enhanced summer survival","volume":"160","author":"Montero","year":"2020","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R17","doi-asserted-by":"crossref","first-page":"521","DOI":"10.1007\/s11284-008-0563-4","article-title":"Complexity, dynamics and diversity of sociality in group-living mammals","volume":"24","author":"Kutsukake","year":"2009","journal-title":"Ecol. Res."},{"key":"2026072820203354300_R18","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1016\/j.anbehav.2015.02.018","article-title":"Measuring social complexity","volume":"103","author":"Bergman","year":"2015","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R19","doi-asserted-by":"crossref","first-page":"1129","DOI":"10.1111\/ele.13079","article-title":"Social complexity and kinship in animal societies","volume":"21","author":"Lukas","year":"2018","journal-title":"Ecol. Lett."},{"key":"2026072820203354300_R20","doi-asserted-by":"crossref","DOI":"10.1098\/rspb.2019.1991","article-title":"Deconstructing sociality: the types of social connections that predict longevity in a group-living primate","volume":"286","author":"Ellis","year":"2019","journal-title":"Proc. R. Soc. B"},{"key":"2026072820203354300_R21","doi-asserted-by":"crossref","first-page":"2993","DOI":"10.1073\/pnas.1913284117","article-title":"Reciprocity and behavioral heterogeneity govern the stability of social networks","volume":"117","author":"Dakin","year":"2020","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2026072820203354300_R22","doi-asserted-by":"crossref","DOI":"10.1007\/s00265-022-03133-5","article-title":"Quantifying within-group variation in sociality\u2014covariation among metrics and patterns across primate groups and species","volume":"76","author":"Sch\u00fclke","year":"2022","journal-title":"Behav. Ecol. Sociobiol."},{"key":"2026072820203354300_R23","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1098\/rstb.2009.0150","article-title":"Fluctuating selection: the perpetual renewal of adaptation in variable environments","volume":"365","author":"Bell","year":"2010","journal-title":"Phil. Trans. R. Soc. B"},{"key":"2026072820203354300_R24","doi-asserted-by":"crossref","first-page":"3381","DOI":"10.1111\/evo.12532","article-title":"The temporal distribution of directional gradients under selection for an optimum","volume":"68","author":"Chevin","year":"2014","journal-title":"Evolution"},{"key":"2026072820203354300_R25","doi-asserted-by":"crossref","first-page":"5113","DOI":"10.1073\/pnas.1600218113","article-title":"Fluctuating survival selection explains variation in avian group size","volume":"113","author":"Brown","year":"2016","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2026072820203354300_R26","doi-asserted-by":"crossref","DOI":"10.1038\/s41467-017-01299-5","article-title":"Direct benefits explain interspecific variation in helping behaviour among cooperatively breeding birds","volume":"8","author":"Kingma","year":"2017","journal-title":"Nat. Commun."},{"key":"2026072820203354300_R27","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1038\/s41559-016-0057","article-title":"Cooperation facilitates the colonization of harsh environments","volume":"1","author":"Cornwallis","year":"2017","journal-title":"Nat. Ecol. Evol."},{"key":"2026072820203354300_R28","doi-asserted-by":"crossref","DOI":"10.1098\/rsos.172406","article-title":"Multiple benefits of alloparental care in a fluctuating environment","volume":"5","author":"Guindre-Parker","year":"2018","journal-title":"R. Soc. Open Sci."},{"key":"2026072820203354300_R29","doi-asserted-by":"crossref","first-page":"1027","DOI":"10.1086\/708496","article-title":"Survival benefits of group living in a fluctuating environment","volume":"195","author":"Guindre-Parker","year":"2020","journal-title":"Am. Nat."},{"key":"2026072820203354300_R30","doi-asserted-by":"crossref","first-page":"168","DOI":"10.1111\/1365-2656.13322","article-title":"Social network position experiences more variable selection than weaponry in wild subpopulations of forked fungus beetles","volume":"90","author":"Formica","year":"2021","journal-title":"J. Anim. Ecol."},{"key":"2026072820203354300_R31","doi-asserted-by":"crossref","DOI":"10.1038\/s41467-021-23672-1","article-title":"Selection on heritable social network positions is context-dependent in Drosophila melanogaster","volume":"12","author":"Wice","year":"2021","journal-title":"Nat. Commun."},{"key":"2026072820203354300_R32","doi-asserted-by":"crossref","first-page":"1330","DOI":"10.1126\/science.adk0606","article-title":"Ecological disturbance alters the adaptive benefits of social ties","volume":"384","author":"Testard","year":"2024","journal-title":"Science"},{"key":"2026072820203354300_R33","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1016\/j.anbehav.2019.08.018","article-title":"Seasonal variation and stability across years in a social network of wild giraffe","volume":"157","author":"Prehn","year":"2019","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R34","doi-asserted-by":"crossref","first-page":"731","DOI":"10.1016\/j.anbehav.2011.12.021","article-title":"Social network correlates of food availability in an endangered population of killer whales, Orcinus orca","volume":"83","author":"Foster","year":"2012","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R35","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1111\/ele.12898","article-title":"Contact networks structured by sex underpin sex-specific epidemiology of infection","volume":"21","author":"Silk","year":"2018","journal-title":"Ecol. Lett."},{"key":"2026072820203354300_R36","doi-asserted-by":"crossref","first-page":"1015","DOI":"10.1007\/s00265-009-0720-y","article-title":"Cyclicity in the structure of female baboon social networks","volume":"63","author":"Henzi","year":"2009","journal-title":"Behav. Ecol. Sociobiol."},{"key":"2026072820203354300_R37","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1016\/j.anbehav.2017.10.012","article-title":"Seasonal variation in female Asian elephant social structure in Nagarahole-Bandipur, southern India","volume":"134","author":"Nandini","year":"2017","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R38","doi-asserted-by":"crossref","DOI":"10.1098\/rspb.2023.1187","article-title":"Dolphin social phenotypes vary in response to food availability but not the North Atlantic Oscillation index","volume":"290","author":"Fisher","year":"2023","journal-title":"Proc. R. Soc. B"},{"key":"2026072820203354300_R39","doi-asserted-by":"crossref","DOI":"10.1007\/s00265-023-03300-2","article-title":"Network reaction norms: taking account of network position and plasticity in response to environmental change","volume":"77","author":"Bonnell","year":"2023","journal-title":"Behav. Ecol. Sociobiol."},{"key":"2026072820203354300_R40","doi-asserted-by":"crossref","first-page":"2661","DOI":"10.1111\/brv.12772","article-title":"Anticipated effects of abiotic environmental change on intraspecific social interactions","volume":"96","author":"Fisher","year":"2021","journal-title":"Biol. Rev."},{"key":"2026072820203354300_R41","first-page":"1","article-title":"Stronger together? A framework for studying population resilience to climate change impacts via social shielding","author":"Warrington","year":"2024","journal-title":"EcoEvoRxiv"},{"key":"2026072820203354300_R42","doi-asserted-by":"crossref","DOI":"10.1098\/rsbl.2013.0428","article-title":"Coping with the cold: predictors of survival in wild Barbary macaques, Macaca sylvanus","volume":"9","author":"McFarland","year":"2013","journal-title":"Biol. Lett."},{"key":"2026072820203354300_R43","doi-asserted-by":"crossref","first-page":"1199","DOI":"10.1016\/j.tree.2025.09.004","article-title":"Quality, quantity, and the adaptive function of social relationships","volume":"40","author":"De Moor","year":"2025","journal-title":"Trends Ecol. Evol."},{"key":"2026072820203354300_R44","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1016\/bs.asb.2017.12.001","article-title":"Linking sociality to fitness in primates: a call for mechanisms","author":"Ostner","year":"2018","journal-title":"Adv. Study Behav."},{"key":"2026072820203354300_R45","doi-asserted-by":"crossref","first-page":"1664","DOI":"10.1016\/j.cub.2022.03.027","article-title":"Social integration influences fitness in allied male dolphins","volume":"32","author":"Gerber","year":"2022","journal-title":"Curr. Biol."},{"key":"2026072820203354300_R46","doi-asserted-by":"crossref","DOI":"10.1098\/rsbl.2017.0112","article-title":"Social bet-hedging in vampire bats","volume":"13","author":"Carter","year":"2017","journal-title":"Biol. Lett."},{"key":"2026072820203354300_R47","doi-asserted-by":"crossref","DOI":"10.1186\/1472-6785-11-17","article-title":"The dynamics of social networks among female Asian elephants","volume":"11","author":"de Silva","year":"2011","journal-title":"BMC Ecol."},{"key":"2026072820203354300_R48","article-title":"Social thermoregulation as a potential mechanism linking sociality and fitness: Barbary macaques with more social partners form larger huddles","volume":"8","author":"Campbell","year":"2018","journal-title":"Scient. Rep."},{"key":"2026072820203354300_R49","doi-asserted-by":"crossref","first-page":"749","DOI":"10.1111\/jav.00916","article-title":"Communal roosting, thermoregulatory benefits and breeding group size predictability in cooperatively breeding sociable weavers","volume":"47","author":"Paquet","year":"2016","journal-title":"J. Avian Biol."},{"key":"2026072820203354300_R50","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1016\/j.anbehav.2024.08.008","article-title":"Selective social tolerance drives differentiated relationships among wild female chimpanzees","volume":"217","author":"Fox","year":"2024","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R51","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1016\/j.anbehav.2014.11.001","article-title":"Predators, food and social context shape the types of vigilance exhibited by kangaroos","volume":"99","author":"Favreau","year":"2015","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R52","doi-asserted-by":"crossref","first-page":"695","DOI":"10.1098\/rspb.2003.2648","article-title":"Coping with divided attention: the advantage of familiarity","volume":"271","author":"Griffiths","year":"2004","journal-title":"Proc. R. Soc. Lond. B"},{"key":"2026072820203354300_R53","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1016\/j.anbehav.2019.02.014","article-title":"North American red squirrels mitigate costs of territory defence through social plasticity","volume":"151","author":"Siracusa","year":"2019","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R54","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1098\/rspb.2000.1357","article-title":"Complex social structure, alliance stability and mating access in a bottlenose dolphin \u2018super-alliance\u2019","volume":"268","author":"Connor","year":"2001","journal-title":"Proc. R. Soc. Lond. B"},{"key":"2026072820203354300_R55","doi-asserted-by":"crossref","first-page":"1829","DOI":"10.1007\/s00265-014-1792-x","article-title":"Predictors of food-related aggression in wild Assamese macaques and the role of conflict avoidance","volume":"68","author":"Heesen","year":"2014","journal-title":"Behav. Ecol. Sociobiol."},{"key":"2026072820203354300_R56","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1007\/s00265-021-03025-0","article-title":"Strongly bonded individuals prefer to forage together in cooperatively breeding dwarf mongoose groups","volume":"75","author":"Kern","year":"2021","journal-title":"Behav. Ecol. Sociobiol."},{"key":"2026072820203354300_R57","doi-asserted-by":"crossref","DOI":"10.1007\/s00265-017-2339-8","article-title":"The influence of social relationship on food tolerance in wolves and dogs","volume":"71","author":"Dale","year":"2017","journal-title":"Behav. Ecol. Sociobiol."},{"key":"2026072820203354300_R58","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1016\/j.anbehav.2017.01.012","article-title":"Understanding repeatability and plasticity in multiple dimensions of the sociability of wild female kangaroos","volume":"126","author":"Menz","year":"2017","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R59","doi-asserted-by":"crossref","first-page":"839","DOI":"10.1016\/j.anbehav.2008.12.007","article-title":"Structured association patterns and their energetic benefits in female eastern grey kangaroos, Macropus giganteus","volume":"77","author":"Carter","year":"2009","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R60","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1111\/jzo.12531","article-title":"Seasonal changes in social networks of giraffes","volume":"305","author":"Wolf","year":"2018","journal-title":"J. Zool."},{"key":"2026072820203354300_R61","doi-asserted-by":"crossref","first-page":"631","DOI":"10.1890\/14-1320.1","article-title":"Sex-based differences in the adaptive value of social behavior contrasted against morphology and environment","volume":"96","author":"Vander Wal","year":"2015","journal-title":"Ecology"},{"key":"2026072820203354300_R62","doi-asserted-by":"crossref","first-page":"701","DOI":"10.1111\/ibi.12191","article-title":"The importance of fission\u2013fusion social group dynamics in birds","volume":"156","author":"Silk","year":"2014","journal-title":"Ibis"},{"key":"2026072820203354300_R63","doi-asserted-by":"crossref","unstructured":"Clutton-Brock\n              TH\n            , PembertonJM. 2004Soay sheep: dynamics and selection in an island population. Cambridge, UK: Cambridge University Press. Seehttps:\/\/play.google.com\/store\/books\/details?id=3VyekpldE8EC.","DOI":"10.1017\/CBO9780511550669"},{"key":"2026072820203354300_R64","doi-asserted-by":"crossref","first-page":"1528","DOI":"10.1126\/science.292.5521.1528","article-title":"Age, sex, density, winter weather, and population crashes in Soay sheep","volume":"292","author":"Coulson","year":"2001","journal-title":"Science"},{"key":"2026072820203354300_R65","doi-asserted-by":"crossref","first-page":"453","DOI":"10.1111\/1467-9876.00205","article-title":"Factors influencing Soay sheep survival","volume":"49","author":"Catchpole","year":"2000","journal-title":"J. R. Stat. Soc. Ser. C. Appl. Stat."},{"key":"2026072820203354300_R66","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1016\/j.anbehav.2014.07.023","article-title":"Social networks created with different techniques are not comparable","volume":"96","author":"Castles","year":"2014","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R67","doi-asserted-by":"crossref","first-page":"1231","DOI":"10.1038\/s41559-022-01817-9","article-title":"Ageing red deer alter their spatial behaviour and become less social","volume":"6","author":"Albery","year":"2022","journal-title":"Nat. Ecol. Evol."},{"key":"2026072820203354300_R68","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.anbehav.2015.03.005","article-title":"The consequences of unidentifiable individuals for the analysis of an animal social network","volume":"104","author":"Silk","year":"2015","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R69","doi-asserted-by":"crossref","first-page":"1001","DOI":"10.1111\/ele.12965","article-title":"Declining home range area predicts reduced late-life survival in two wild ungulate populations","volume":"21","author":"Froy","year":"2018","journal-title":"Ecol. Lett."},{"key":"2026072820203354300_R70","doi-asserted-by":"crossref","first-page":"493","DOI":"10.1007\/s00265-009-0865-8","article-title":"Sampling animal association networks with the gambit of the group","volume":"64","author":"Franks","year":"2010","journal-title":"Behav. Ecol. Sociobiol."},{"key":"2026072820203354300_R71","doi-asserted-by":"crossref","DOI":"10.7208\/chicago\/9780226895246.001.0001","volume-title":"Analyzing animal societies","author":"Whitehead","year":"2008"},{"key":"2026072820203354300_R72","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1163\/156853974X00345","article-title":"The social organisation of antelope in relation to their ecology","volume":"48","author":"Jarman","year":"1974","journal-title":"Behaviour"},{"key":"2026072820203354300_R73","doi-asserted-by":"crossref","first-page":"676","DOI":"10.1111\/ele.13684","article-title":"Multiple spatial behaviours govern social network positions in a wild ungulate","volume":"24","author":"Albery","year":"2021","journal-title":"Ecol. Lett."},{"key":"2026072820203354300_R74","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1111\/ele.12550","article-title":"Sex differences in relationships between habitat use and reproductive performance in Soay sheep (Ovis aries)","volume":"19","author":"Regan","year":"2015","journal-title":"Ecol. Lett."},{"key":"2026072820203354300_R75","doi-asserted-by":"crossref","first-page":"1869","DOI":"10.1111\/1365-2656.13978","article-title":"Age-specific impacts of vegetation functional traits on gastrointestinal nematode parasite burdens in a large herbivore","volume":"92","author":"Wiersma","year":"2023","journal-title":"J. Anim. Ecol."},{"key":"2026072820203354300_R76","doi-asserted-by":"crossref","first-page":"516","DOI":"10.1016\/j.ecolmodel.2006.03.017","article-title":"The package \u2018adehabitat\u2019 for the R software: a tool for the analysis of space and habitat use by animals","volume":"197","author":"Calenge","year":"2006","journal-title":"Ecol. Modell."},{"key":"2026072820203354300_R77","doi-asserted-by":"crossref","first-page":"715","DOI":"10.1017\/S0031182025100383","article-title":"Local and global density have distinct and parasite-dependent effects on infection in wild sheep","volume":"152","author":"Albery","year":"2025","journal-title":"Parasitology"},{"key":"2026072820203354300_R78","doi-asserted-by":"crossref","first-page":"2002","DOI":"10.1038\/s41559-025-02843-z","article-title":"Density-dependent network structuring within and across wild animal systems","volume":"9","author":"Albery","year":"2025","journal-title":"Nat. Ecol. Evol."},{"key":"2026072820203354300_R79","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1007\/s101440200013","article-title":"Long-term responses in arctic ungulate dynamics to changes in climatic and trophic processes","volume":"44","author":"Forchhammer","year":"2002","journal-title":"Popul. Ecol."},{"key":"2026072820203354300_R80","doi-asserted-by":"crossref","first-page":"543","DOI":"10.1111\/oik.01727","article-title":"Analysis of phenotypic change in relation to climatic drivers in a population of Soay sheep Ovis aries","volume":"124","author":"Simmonds","year":"2015","journal-title":"Oikos"},{"key":"2026072820203354300_R81","doi-asserted-by":"crossref","DOI":"10.1007\/s00265-022-03254-x","article-title":"Common permutation methods in animal social network analysis do not control for non-independence","volume":"76","author":"Hart","year":"2022","journal-title":"Behav. Ecol. Sociobiol."},{"key":"2026072820203354300_R82","volume-title":"R: a language and environment for statistical computing","author":"R Core Team","year":"2025"},{"key":"2026072820203354300_R83","unstructured":"Stan Development Team\n          . 2025RStan: the R interface to Stan. R package version 2.26.24. Seehttps:\/\/mc-stan.org\/."},{"key":"2026072820203354300_R84","doi-asserted-by":"crossref","first-page":"1","DOI":"10.18637\/jss.v080.i01","article-title":"brms: An R package for Bayesian multilevel models using Stan","volume":"80","author":"B\u00fcrkner","year":"2017","journal-title":"J. Stat. Softw."},{"key":"2026072820203354300_R85","doi-asserted-by":"crossref","DOI":"10.1201\/9780429029608","volume-title":"Statistical rethinking: a Bayesian course with examples in R and Stan","author":"McElreath","year":"2020","edition":"2nd edn"},{"key":"2026072820203354300_R86","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1146\/annurev-statistics-031017-100307","article-title":"On p-values and Bayes factors","volume":"5","author":"Held","year":"2018","journal-title":"Annu. Rev. Stat. Appl."},{"key":"2026072820203354300_R87","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1016\/j.tree.2021.10.009","article-title":"Rewriting results sections in the language of evidence","volume":"37","author":"Muff","year":"2022","journal-title":"Trends Ecol. Evol."},{"key":"2026072820203354300_R88","doi-asserted-by":"crossref","DOI":"10.1098\/rsos.160255","article-title":"Network connections, dyadic bonds and fitness in wild female baboons","volume":"3","author":"Cheney","year":"2016","journal-title":"R. Soc. Open Sci."},{"key":"2026072820203354300_R89","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1016\/j.anbehav.2020.02.009","article-title":"How does individual variation in sociality influence fitness in prairie voles?","volume":"163","author":"Sabol","year":"2020","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R90","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1016\/j.anbehav.2017.02.002","article-title":"The \u2018strength of weak ties\u2019 among female baboons: fitness-related benefits of social bonds","volume":"126","author":"McFarland","year":"2017","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R91","doi-asserted-by":"crossref","first-page":"651","DOI":"10.1098\/rsbl.2007.0278","article-title":"Function of weaponry in females: the use of horns in intrasexual competition for resources in female Soay sheep","volume":"3","author":"Robinson","year":"2007","journal-title":"Biol. Lett."},{"key":"2026072820203354300_R92","article-title":"Structure and function in human and primate social networks: implications for diffusion, network stability and health","volume":"476","author":"Dunbar","year":"2020","journal-title":"Proc. Math. Phys. Eng. Sci."},{"key":"2026072820203354300_R93","doi-asserted-by":"crossref","first-page":"613","DOI":"10.1007\/s10682-006-9141-9","article-title":"Fission\u2013fusion social systems as a strategy for coping with ecological constraints: a primate case","volume":"21","author":"Lehmann","year":"2007","journal-title":"Evol. Ecol."},{"key":"2026072820203354300_R94","doi-asserted-by":"crossref","first-page":"683","DOI":"10.1007\/s00265-006-0212-2","article-title":"Time as a constraint on group size in spider monkeys","volume":"60","author":"Korstjens","year":"2006","journal-title":"Behav. Ecol. Sociobiol."},{"key":"2026072820203354300_R95","doi-asserted-by":"crossref","first-page":"553","DOI":"10.1016\/j.anbehav.2012.12.010","article-title":"Correcting for the impact of gregariousness in social network analyses","volume":"85","author":"Godde","year":"2013","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R96","doi-asserted-by":"crossref","DOI":"10.1098\/rspb.2025.0978","article-title":"Social and spatial drivers of the multitiered structure of zebra finch social networks","volume":"292","author":"Zhang","year":"2025","journal-title":"Proc. R. Soc. B"},{"key":"2026072820203354300_R97","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1007\/s002650050390","article-title":"The evolution of female social relationships in nonhuman primates","volume":"41","author":"Sterck","year":"1997","journal-title":"Behav. Ecol. Sociobiol."},{"key":"2026072820203354300_R98","doi-asserted-by":"crossref","first-page":"872","DOI":"10.1111\/eth.13143","article-title":"On the importance of defendable resources for social evolution: applying new techniques to a long-standing question","volume":"127","author":"Jungwirth","year":"2021","journal-title":"Ethology"},{"key":"2026072820203354300_R99","doi-asserted-by":"crossref","first-page":"531","DOI":"10.1016\/j.anbehav.2012.06.003","article-title":"Antipredator and social monitoring functions of vigilance behaviour in blue monkeys","volume":"84","author":"Gaynor","year":"2012","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R100","doi-asserted-by":"crossref","first-page":"988","DOI":"10.1093\/beheco\/arw003","article-title":"Working the crowd: sociable vervets benefit by reducing exposure to risk","volume":"27","author":"Josephs","year":"2016","journal-title":"Behav. Ecol."},{"key":"2026072820203354300_R101","article-title":"Social security: does social position influence flight initiation distance?","volume":"35","author":"Szulanski","year":"2023","journal-title":"Behav. Ecol."},{"key":"2026072820203354300_R102","doi-asserted-by":"crossref","DOI":"10.1098\/rspb.2021.0617","article-title":"Age and sex influence social interactions, but not associations, within a killer whale pod","volume":"288","author":"Weiss","year":"2021","journal-title":"Proc. R. Soc. B"},{"key":"2026072820203354300_R103","doi-asserted-by":"crossref","first-page":"868","DOI":"10.1111\/brv.12934","article-title":"Behavioural ecology at the spatial\u2013social interface","volume":"98","author":"Webber","year":"2023","journal-title":"Biol. Rev."},{"key":"2026072820203354300_R104","doi-asserted-by":"crossref","first-page":"274","DOI":"10.1017\/S0031182020002048","article-title":"Bidirectional interactions between host social behaviour and parasites arise through ecological and evolutionary processes","volume":"148","author":"Hawley","year":"2021","journal-title":"Parasitology"},{"key":"2026072820203354300_R105","first-page":"113","volume-title":"Soay sheep: dynamics and selection in an island population","author":"Wilson","year":"2004"},{"key":"2026072820203354300_R106","doi-asserted-by":"crossref","first-page":"1114","DOI":"10.2307\/1383163","article-title":"Optimal foraging and risk of predation: effects on behavior and social structure in ungulates","volume":"80","author":"Kie","year":"1999","journal-title":"J. Mammal."},{"key":"2026072820203354300_R107","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1016\/j.anbehav.2015.02.019","article-title":"Male dolphin alliances in Shark Bay: changing perspectives in a 30-year study","volume":"103","author":"Connor","year":"2015","journal-title":"Anim. Behav."},{"key":"2026072820203354300_R108","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0047508","article-title":"Early social networks predict survival in wild bottlenose dolphins","volume":"7","author":"Stanton","year":"2012","journal-title":"PLoS One"},{"key":"2026072820203354300_R109","doi-asserted-by":"crossref","DOI":"10.1016\/j.isci.2021.102864","article-title":"Social bonds provide multiple pathways to reproductive success in wild male chimpanzees","volume":"24","author":"Feldblum","year":"2021","journal-title":"iScience"},{"key":"2026072820203354300_R110","article-title":"Data for: Environment-dependent benefits of sociality in Soay sheep. Figshare.","author":"Siracusa","year":"2026"},{"key":"2026072820203354300_R111","article-title":"Supplementary material from: Environment-dependent benefits of sociality in Soay sheep. Figshare.","author":"Siracusa","year":"2026"}],"container-title":["Proceedings of the Royal Society B: Biological Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/royalsocietypublishing.org\/rspb\/article-pdf\/doi\/10.1098\/rspb.2025.2821\/6154181\/rspb.2025.2821.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/royalsocietypublishing.org\/rspb\/article-pdf\/doi\/10.1098\/rspb.2025.2821\/6154181\/rspb.2025.2821.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,29]],"date-time":"2026-07-29T00:20:57Z","timestamp":1785284457000},"score":1,"resource":{"primary":{"URL":"https:\/\/royalsocietypublishing.org\/rspb\/article\/293\/2075\/20252821\/482645\/Environment-dependent-benefits-of-sociality-in"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,1]]},"references-count":111,"journal-issue":{"issue":"2075","published-print":{"date-parts":[[2026,7,1]]}},"URL":"https:\/\/doi.org\/10.1098\/rspb.2025.2821","relation":{},"ISSN":["1471-2954"],"issn-type":[{"value":"1471-2954","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,1]]},"article-number":"20252821"}}