{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,2]],"date-time":"2026-06-02T05:14:16Z","timestamp":1780377256984,"version":"3.54.1"},"reference-count":81,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,1,11]],"date-time":"2025-01-11T00:00:00Z","timestamp":1736553600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2025,1,11]],"date-time":"2025-01-11T00:00:00Z","timestamp":1736553600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["IBN-9874523"],"award-info":[{"award-number":["IBN-9874523"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["CNS- 025214"],"award-info":[{"award-number":["CNS- 025214"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["IOB-9874523"],"award-info":[{"award-number":["IOB-9874523"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Commun Biol"],"DOI":"10.1038\/s42003-024-07169-3","type":"journal-article","created":{"date-parts":[[2025,1,11]],"date-time":"2025-01-11T13:28:42Z","timestamp":1736602122000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["\u201cAge of risk\u201d shapes simpler multimodal communication in the juvenile plains zebra (Equus quagga)"],"prefix":"10.1038","volume":"8","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9142-6917","authenticated-orcid":false,"given":"Severine B. S. W.","family":"Hex","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9049-5219","authenticated-orcid":false,"given":"Daniel I.","family":"Rubenstein","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,1,11]]},"reference":[{"key":"7169_CR1","doi-asserted-by":"crossref","unstructured":"Joffe, T. H. Social pressures have selected for an extended juvenile period in primates. J. Hum. Evol. 32, 593\u2013605 (1997).","DOI":"10.1006\/jhev.1997.0140"},{"key":"7169_CR2","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1016\/S0047-2484(85)80005-1","volume":"14","author":"M Tomasello","year":"1985","unstructured":"Tomasello, M., George, B. L., Kruger, A. C., Jeffrey, M. & Evans, A. The development of gestural communication in young chimpanzees. J. Hum. Evol. 14, 175\u2013186 (1985).","journal-title":"J. Hum. Evol."},{"key":"7169_CR3","doi-asserted-by":"publisher","first-page":"621","DOI":"10.1016\/j.anbehav.2009.12.008","volume":"79","author":"C Arnold","year":"2010","unstructured":"Arnold, C. & Taborsky, B. Social experience in early ontogeny has lasting effects on social skills in cooperatively breeding cichlids. Anim. Behav. 79, 621\u2013630 (2010).","journal-title":"Anim. Behav."},{"key":"7169_CR4","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s00265-018-2619-y","volume":"72","author":"M Fr\u00f6hlich","year":"2018","unstructured":"Fr\u00f6hlich, M. & Hobaiter, C. The development of gestural communication in great apes. Behav. Ecol. Sociobiol. 72, 1\u201314 (2018).","journal-title":"Behav. Ecol. Sociobiol."},{"key":"7169_CR5","doi-asserted-by":"publisher","first-page":"1869","DOI":"10.1098\/rstb.2011.0221","volume":"367","author":"KA Pollard","year":"2012","unstructured":"Pollard, K. A. & Blumstein, D. T. Evolving communicative complexity: insights from rodents and beyond. Philos. Trans. R. Soc. B Biol. Sci. 367, 1869\u20131878 (2012).","journal-title":"Philos. Trans. R. Soc. B Biol. Sci."},{"key":"7169_CR6","doi-asserted-by":"crossref","unstructured":"Taborsky, M. Cichlid fishes: a model for the integrative study of social behaviour. In Cooperative Breeding in Vertebrates 272\u2013293 (Cambridge University Press, Cambridge, 2016).","DOI":"10.1017\/CBO9781107338357.017"},{"key":"7169_CR7","doi-asserted-by":"publisher","first-page":"RP87008","DOI":"10.7554\/eLife.87008","volume":"12","author":"AV Rincon","year":"2023","unstructured":"Rincon, A. V. et al. Higher social tolerance is associated with more complex facial behaviour in macaques. Elife 12, RP87008 (2023).","journal-title":"Elife"},{"key":"7169_CR8","unstructured":"Bradbury, J. W. & Vehrencamp, S. L. Principles of Animal Communication Vol. 132 (Sinauer Associates, Sunderland, MA, 1998)."},{"key":"7169_CR9","doi-asserted-by":"publisher","first-page":"756","DOI":"10.2307\/1380802","volume":"66","author":"DA Conner","year":"1985","unstructured":"Conner, D. A. & Whitworth, M. R. The ontogeny of vocal communication in the pika. J. Mammal. 66, 756\u2013763 (1985).","journal-title":"J. Mammal."},{"key":"7169_CR10","doi-asserted-by":"publisher","first-page":"226","DOI":"10.1037\/0735-7036.111.3.226","volume":"111","author":"EF Brittan-Powell","year":"1997","unstructured":"Brittan-Powell, E. F., Dooling, R. J. & Farabaugh, S. M. Vocal development in budgerigars (Melopsittacus undulatus): contact calls. J. Comp. Psychol. 111, 226 (1997).","journal-title":"J. Comp. Psychol."},{"key":"7169_CR11","doi-asserted-by":"publisher","first-page":"223","DOI":"10.1075\/eoc.1.2.04tom","volume":"1","author":"M Tomasello","year":"1997","unstructured":"Tomasello, M. et al. The ontogeny of chimpanzee gestural signals: a comparison across groups and generations. Evol. Commun. 1, 223\u2013259 (1997).","journal-title":"Evol. Commun."},{"key":"7169_CR12","doi-asserted-by":"publisher","first-page":"827","DOI":"10.1007\/s10071-011-0416-3","volume":"14","author":"C Hobaiter","year":"2011","unstructured":"Hobaiter, C. & Byrne, R. W. Serial gesturing by wild chimpanzees: its nature and function for communication. Anim. Cognit. 14, 827\u2013838 (2011).","journal-title":"Anim. Cognit."},{"key":"7169_CR13","doi-asserted-by":"publisher","first-page":"923","DOI":"10.1126\/science.abf9279","volume":"373","author":"AA Fernandez","year":"2021","unstructured":"Fernandez, A. A., Burchardt, L. S., Nagy, M. & Kn\u00f6rnschild, M. Babbling in a vocal learning bat resembles human infant babbling. Science 373, 923\u2013926 (2021).","journal-title":"Science"},{"key":"7169_CR14","doi-asserted-by":"publisher","first-page":"20200239","DOI":"10.1098\/rstb.2020.0239","volume":"376","author":"SM Ter Haar","year":"2021","unstructured":"Ter Haar, S. M. et al. Cross-species parallels in babbling: animals and algorithms. Philos. Trans. R. Soc. B 376, 20200239 (2021).","journal-title":"Philos. Trans. R. Soc. B"},{"key":"7169_CR15","doi-asserted-by":"publisher","first-page":"679","DOI":"10.1016\/j.tree.2012.09.003","volume":"27","author":"B Taborsky","year":"2012","unstructured":"Taborsky, B. & Oliveira, R. F. Social competence: an evolutionary approach. Trends Ecol. Evol. 27, 679\u2013688 (2012).","journal-title":"Trends Ecol. Evol."},{"key":"7169_CR16","doi-asserted-by":"publisher","first-page":"225","DOI":"10.1146\/annurev-devpsych-051820-121446","volume":"2","author":"KM Faust","year":"2020","unstructured":"Faust, K. M., Carouso-Peck, S., Elson, M. R. & Goldstein, M. H. The origins of social knowledge in altricial species. Annu. Rev. Dev. Psychol. 2, 225\u2013246 (2020).","journal-title":"Annu. Rev. Dev. Psychol."},{"key":"7169_CR17","doi-asserted-by":"publisher","first-page":"230740","DOI":"10.1098\/rsos.230740","volume":"11","author":"O La Loggia","year":"2024","unstructured":"La Loggia, O., Wilson, A. J. & Taborsky, B. Early social complexity influences social behaviour but not social trajectories in a cooperatively breeding cichlid fish. R. Soc. Open Sci. 11, 230740 (2024).","journal-title":"R. Soc. Open Sci."},{"key":"7169_CR18","doi-asserted-by":"publisher","first-page":"12","DOI":"10.1007\/s00265-023-03426-3","volume":"78","author":"M Fr\u00f6hlich","year":"2024","unstructured":"Fr\u00f6hlich, M., van Noordwijk, M. A., Mitra Setia, T., van Schaik, C. P. & Knief, U. Wild and captive immature orangutans differ in their non-vocal communication with others, but not with their mothers. Behav. Ecol. Sociobiol. 78, 12 (2024).","journal-title":"Behav. Ecol. Sociobiol."},{"key":"7169_CR19","doi-asserted-by":"publisher","first-page":"e0231343","DOI":"10.1371\/journal.pone.0231343","volume":"15","author":"CM Nu\u00f1ez","year":"2020","unstructured":"Nu\u00f1ez, C. M. & Rubenstein, D. I. Communication is key: mother-offspring signaling can affect behavioural responses and offspring survival in feral horses (Equus caballus). PLoS ONE 15, e0231343 (2020).","journal-title":"PLoS ONE"},{"key":"7169_CR20","doi-asserted-by":"publisher","first-page":"319","DOI":"10.1016\/S0168-1591(99)00126-4","volume":"68","author":"JW Weeks","year":"2000","unstructured":"Weeks, J. W., Crowell-Davis, S. L., Caudle, A. B. & Heusner, G. L. Aggression and social spacing in light horse (Equus caballus) mares and foals. Appl. Anim. Behav. Sci. 68, 319\u2013337 (2000).","journal-title":"Appl. Anim. Behav. Sci."},{"key":"7169_CR21","unstructured":"Altmann, J. Baboon Mothers and Infants (University of Chicago Press, 2001)."},{"key":"7169_CR22","doi-asserted-by":"publisher","first-page":"478","DOI":"10.1093\/beheco\/arp020","volume":"20","author":"ML East","year":"2009","unstructured":"East, M. L. et al. Maternal effects on offspring social status in spotted hyenas. Behav. Ecol. 20, 478\u2013483 (2009).","journal-title":"Behav. Ecol."},{"key":"7169_CR23","doi-asserted-by":"publisher","first-page":"138","DOI":"10.1093\/beheco\/aru163","volume":"26","author":"CM Nu\u00f1ez","year":"2015","unstructured":"Nu\u00f1ez, C. M., Adelman, J. S. & Rubenstein, D. I. Sociality increases juvenile survival after a catastrophic event in the feral horse (Equus caballus). Behav. Ecol. 26, 138\u2013147 (2015).","journal-title":"Behav. Ecol."},{"key":"7169_CR24","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s00265-018-2547-x","volume":"72","author":"D Maestripieri","year":"2018","unstructured":"Maestripieri, D. Maternal influences on primate social development. Behav. Ecol. Sociobiol. 72, 1\u201312 (2018).","journal-title":"Behav. Ecol. Sociobiol."},{"key":"7169_CR25","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.anbehav.2018.07.002","volume":"143","author":"SZ Goldenberg","year":"2018","unstructured":"Goldenberg, S. Z. & Wittemyer, G. Orphaning and natal group dispersal are associated with social costs in female elephants. Anim. Behav. 143, 1\u20138 (2018).","journal-title":"Anim. Behav."},{"key":"7169_CR26","doi-asserted-by":"publisher","first-page":"325","DOI":"10.1006\/anbe.2002.3071","volume":"64","author":"F Aureli","year":"2002","unstructured":"Aureli, F., Cords, M. & Van Schaik, C. P. Conflict resolution following aggression in gregarious animals: a predictive framework. Anim. Behav. 64, 325\u2013343 (2002).","journal-title":"Anim. Behav."},{"key":"7169_CR27","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1644\/05-MAMM-A-138R1.1","volume":"87","author":"RJ Sarno","year":"2006","unstructured":"Sarno, R. J., Bank, M. S., Stern, H. S. & Franklin, W. L. Effects of age, sex, season, and social dynamics on juvenile guanaco subordinate behaviour. J. Mammal. 87, 41\u201347 (2006).","journal-title":"J. Mammal."},{"key":"7169_CR28","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1093\/icb\/14.1.249","volume":"14","author":"RL Trivers","year":"1974","unstructured":"Trivers, R. L. Parent-offspring conflict. Am. Zool. 14, 249\u2013264 (1974).","journal-title":"Am. Zool."},{"key":"7169_CR29","unstructured":"Bowker, M. H. Behaviour of Kirk\u2019s dikdik (Madoqua kirki hindei) in Kenya (Northern Arizona University, 1978)."},{"key":"7169_CR30","doi-asserted-by":"publisher","first-page":"665","DOI":"10.1016\/0003-3472(89)90045-6","volume":"37","author":"GE Hill","year":"1989","unstructured":"Hill, G. E. Late spring arrival and dull nuptial plumage: aggression avoidance by yearling males? Anim. Behav. 37, 665\u2013673 (1989).","journal-title":"Anim. Behav."},{"key":"7169_CR31","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1016\/j.anbehav.2015.01.004","volume":"102","author":"B Fresnillo","year":"2015","unstructured":"Fresnillo, B., Belliure, J. & Cuervo, J. J. Red coloration in juvenile spiny-footed lizards, Acanthodactylus erythrurus, reduces adult aggression. Anim. Behav. 102, 59\u201367 (2015).","journal-title":"Anim. Behav."},{"key":"7169_CR32","doi-asserted-by":"publisher","first-page":"235","DOI":"10.1111\/j.1439-0310.1943.tb00655.x","volume":"5","author":"K Lorenz","year":"1943","unstructured":"Lorenz, K. Die angeborenen formen m\u00f6glicher erfahrung. Z. Tierpsychol. 5, 235\u2013409 (1943).","journal-title":"Z. Tierpsychol."},{"key":"7169_CR33","doi-asserted-by":"publisher","first-page":"333","DOI":"10.1353\/ect.2008.0045","volume":"4","author":"SJ Gould","year":"2008","unstructured":"Gould, S. J. A biological homage to Mickey Mouse. Ecotone 4, 333\u2013340 (2008).","journal-title":"Ecotone"},{"key":"7169_CR34","doi-asserted-by":"publisher","first-page":"257","DOI":"10.1111\/j.1439-0310.2008.01603.x","volume":"115","author":"ML Glocker","year":"2009","unstructured":"Glocker, M. L. et al. Baby schema in infant faces induces cuteness perception and motivation for caretaking in adults. Ethology 115, 257\u2013263 (2009).","journal-title":"Ethology"},{"key":"7169_CR35","doi-asserted-by":"crossref","unstructured":"Frodi, A. When empathy fails: aversive infant crying and child abuse. in Infant Crying: Theoretical and Research Perspectives 263\u2013277 (Springer, Boston, MA, 1985).","DOI":"10.1007\/978-1-4613-2381-5_12"},{"key":"7169_CR36","doi-asserted-by":"publisher","DOI":"10.1038\/srep27585","volume":"6","author":"AS Stoeger","year":"2016","unstructured":"Stoeger, A. S. & Baotic, A. Information content and acoustic structure of male African elephant social rumbles. Sci. Rep. 6, 27585 (2016).","journal-title":"Sci. Rep."},{"key":"7169_CR37","doi-asserted-by":"crossref","unstructured":"Anthony, C. D. & Wicknick, J. A. Aggressive interactions and chemical communication between adult and juvenile salamanders. J. Herpetol. 27, 261\u2013264 (1993).","DOI":"10.2307\/1565145"},{"key":"7169_CR38","doi-asserted-by":"publisher","first-page":"916","DOI":"10.1093\/beheco\/ars056","volume":"23","author":"CN Templeton","year":"2012","unstructured":"Templeton, C. N., Campbell, S. E. & Beecher, M. D. Territorial song sparrows tolerate juveniles during the early song-learning phase. Behav. Ecol. 23, 916\u2013923 (2012).","journal-title":"Behav. Ecol."},{"key":"7169_CR39","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1016\/j.anbehav.2023.03.020","volume":"201","author":"E Doherty","year":"2023","unstructured":"Doherty, E., Davila-Ross, M. & Clay, Z. Multimodal communication development in semiwild chimpanzees. Anim. Behav. 201, 175\u2013190 (2023).","journal-title":"Anim. Behav."},{"key":"7169_CR40","doi-asserted-by":"publisher","first-page":"446","DOI":"10.1111\/j.1439-0310.2007.01348.x","volume":"113","author":"BT Hirsch","year":"2007","unstructured":"Hirsch, B. T. Spoiled brats: Is extreme juvenile agonism in ring\u2010tailed coatis (Nasua nasua) dominance or tolerated aggression? Ethology 113, 446\u2013456 (2007).","journal-title":"Ethology"},{"key":"7169_CR41","doi-asserted-by":"crossref","unstructured":"Ames, A. E. & Vergara, V. Trajectories of vocal repertoire development in beluga (Delphinapterus leucas) calves: insights from studies a decade apart. Aquat. Mamm. 46, 344\u2013366 (2020).","DOI":"10.1578\/AM.46.4.2020.344"},{"key":"7169_CR42","doi-asserted-by":"publisher","first-page":"619","DOI":"10.1007\/s10071-018-1197-8","volume":"21","author":"M Fr\u00f6hlich","year":"2018","unstructured":"Fr\u00f6hlich, M. & van Schaik, C. P. The function of primate multimodal communication. Anim. Cognit. 21, 619\u2013629 (2018).","journal-title":"Anim. Cognit."},{"key":"7169_CR43","doi-asserted-by":"publisher","first-page":"295","DOI":"10.1016\/j.anbehav.2023.11.002","volume":"207","author":"SB Hex","year":"2024","unstructured":"Hex, S. B. & Rubenstein, D. I. Using networks to visualize, analyse and interpret multimodal communication. Anim. Behav. 207, 295\u2013317 (2024a).","journal-title":"Anim. Behav."},{"key":"7169_CR44","doi-asserted-by":"publisher","DOI":"10.1007\/s00265-021-03046-9","volume":"75","author":"SB Hex","year":"2021","unstructured":"Hex, S. B., Tombak, K. & Rubenstein, D. I. A new classification of mammalian uni-male multi-female groups based on the fundamental principles governing inter-and intrasexual relationships. Behav. Ecol. Sociobiol. 75, 157 (2021).","journal-title":"Behav. Ecol. Sociobiol."},{"key":"7169_CR45","doi-asserted-by":"publisher","first-page":"2059","DOI":"10.1163\/1568539X-00003314","volume":"152","author":"W Tong","year":"2015","unstructured":"Tong, W., Shapiro, B. & Rubenstein, D. I. Genetic relatedness in two-tiered plains zebra societies suggests that females choose to associate with kin. Behaviour 152, 2059\u20132078 (2015).","journal-title":"Behaviour"},{"key":"7169_CR46","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1111\/j.1365-2028.1965.tb00742.x","volume":"3","author":"JS King","year":"1965","unstructured":"King, J. S. A field guide to the reproduction of Grant\u2019s zebra and Grevy\u2019s zebra. East Afr. Wildl. J. 3, 99\u2013117 (1965).","journal-title":"East Afr. Wildl. J."},{"key":"7169_CR47","doi-asserted-by":"publisher","first-page":"89","DOI":"10.4102\/koedoe.v19i1.1186","volume":"19","author":"GL Smuts","year":"1976","unstructured":"Smuts, G. L. Reproduction in the zebra mare Equus burchelli antiquorum from the Kruger National Park. Koedoe 19, 89\u2013132 (1976b).","journal-title":"Koedoe"},{"key":"7169_CR48","doi-asserted-by":"crossref","unstructured":"Klingel, H. The social organisation and population ecology of the plains zebra (Equus quagga). Afr. Zool. 4, 149\u2013163 (1969).","DOI":"10.1080\/00445096.1969.11447374"},{"key":"7169_CR49","unstructured":"Klingel, H. A comparison of the social behaviour of the Equidae. In Symposium on the Behaviour of Ungulates and its Relation to Management 124\u2013132 (IUCN Publications, 1974)."},{"key":"7169_CR50","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1007\/s13364-011-0061-x","volume":"57","author":"HI Simpson","year":"2012","unstructured":"Simpson, H. I., Rands, S. A. & Nicol, C. J. Social structure, vigilance and behaviour of plains zebra (Equus burchellii): a 5-year case study of individuals living on a managed wildlife reserve. Acta Theriol. 57, 111\u2013120 (2012).","journal-title":"Acta Theriol."},{"key":"7169_CR51","doi-asserted-by":"publisher","first-page":"1035","DOI":"10.1007\/s00265-009-0723-8","volume":"63","author":"IR Fischhoff","year":"2009","unstructured":"Fischhoff, I. R., Dushoff, J., Sundaresan, S. R., Cordingley, J. E. & Rubenstein, D. I. Reproductive status influences group size and persistence of bonds in male plains zebra (Equus burchelli). Behav. Ecol. Sociobiol. 63, 1035\u20131043 (2009).","journal-title":"Behav. Ecol. Sociobiol."},{"key":"7169_CR52","doi-asserted-by":"publisher","first-page":"42","DOI":"10.1111\/j.1439-0310.1985.tb00755.x","volume":"69","author":"SL Crowell\u2010Davis","year":"1985","unstructured":"Crowell\u2010Davis, S. L., Houpt, K. A. & Burnham, J. S. Snapping by foals of Equus caballus. Z. Tierpsychol. 69, 42\u201354 (1985).","journal-title":"Z. Tierpsychol."},{"key":"7169_CR53","unstructured":"McCusker, M. E. Investigation of the Effects of Social Experience on Snapping Intensity in Equus caballus Foals. Doctoral dissertation, Virginia Tech. (2003)."},{"key":"7169_CR54","doi-asserted-by":"crossref","unstructured":"Ransom, J. I. & Kaczensky, P. (eds) Wild Equids: Ecology, Management, and Conservation (JHU Press, 2016).","DOI":"10.1353\/book.47460"},{"key":"7169_CR55","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1111\/j.1439-0310.1984.tb01352.x","volume":"66","author":"MB Schilder","year":"1984","unstructured":"Schilder, M. B., van Hooff, J. A., van Geer\u2010Plesman, C. J. & Wensing, J. B. A quantitative analysis of facial expression in the plains zebra. Z. Tierpsychol. 66, 11\u201332 (1984).","journal-title":"Z. Tierpsychol."},{"key":"7169_CR56","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.anbehav.2019.06.006","volume":"154","author":"GM Kohn","year":"2019","unstructured":"Kohn, G. M. How social systems persist: learning to build a social network in an uncertain world. Anim. Behav. 154, 1\u20136 (2019).","journal-title":"Anim. Behav."},{"key":"7169_CR57","doi-asserted-by":"crossref","unstructured":"Maglieri, V., Scopa, C., Modica, G. & Palagi, E. All in good time: play development and tactics in foals (Equus calabus). Appl. Anim. Behav. Sci. 273, 106244 (2024).","DOI":"10.1016\/j.applanim.2024.106244"},{"key":"7169_CR58","doi-asserted-by":"publisher","first-page":"425","DOI":"10.1038\/35044191","volume":"408","author":"R Slotow","year":"2000","unstructured":"Slotow, R., Van Dyk, G., Poole, J., Page, B. & Klocke, A. Older bull elephants control young males. Nature 408, 425\u2013426 (2000).","journal-title":"Nature"},{"key":"7169_CR59","doi-asserted-by":"publisher","first-page":"e4888","DOI":"10.1371\/journal.pone.0004888","volume":"4","author":"M Bourjade","year":"2009","unstructured":"Bourjade, M. et al. Adult-young ratio, a major factor regulating social behaviour of young: a horse study. PLoS ONE 4, e4888 (2009).","journal-title":"PLoS ONE"},{"key":"7169_CR60","doi-asserted-by":"publisher","first-page":"1067","DOI":"10.1016\/j.anbehav.2012.01.037","volume":"83","author":"B Taborsky","year":"2012","unstructured":"Taborsky, B., Arnold, C., Junker, J. & Tschopp, A. The early social environment affects social competence in a cooperative breeder. Anim. Behav. 83, 1067\u20131074 (2012).","journal-title":"Anim. Behav."},{"key":"7169_CR61","doi-asserted-by":"publisher","first-page":"1847","DOI":"10.1098\/rstb.2011.0218","volume":"367","author":"ML Gustison","year":"2012","unstructured":"Gustison, M. L., le Roux, A. & Bergman, T. J. Derived vocalizations of geladas (Theropithecus gelada) and the evolution of vocal complexity in primates. Philos. Trans. R. Soc. B Biol. Sci. 367, 1847\u20131859 (2012).","journal-title":"Philos. Trans. R. Soc. B Biol. Sci."},{"key":"7169_CR62","doi-asserted-by":"publisher","first-page":"123","DOI":"10.1086\/681636","volume":"186","author":"S Fischer","year":"2015","unstructured":"Fischer, S., Bessert-Nettelbeck, M., Kotrschal, A. & Taborsky, B. Rearing-group size determines social competence and brain structure in a cooperatively breeding cichlid. Am. Nat. 186, 123\u2013140 (2015).","journal-title":"Am. Nat."},{"key":"7169_CR63","doi-asserted-by":"publisher","first-page":"119","DOI":"10.1080\/12265071.2003.9647692","volume":"7","author":"JR Rho","year":"2003","unstructured":"Rho, J. R. & Choe, J. C. Difference of early experience of foals according to dominance ranks of their mothers in the Jeju pony (Equus caballus). Korean J. Biol. Sci. 7, 119\u2013122 (2003).","journal-title":"Korean J. Biol. Sci."},{"key":"7169_CR64","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1016\/j.applanim.2007.11.005","volume":"113","author":"F Heitor","year":"2008","unstructured":"Heitor, F. & Vicente, L. Maternal care and foal social relationships in a herd of Sorraia horses: influence of maternal rank and experience. Appl. Anim. Behav. Sci. 113, 189\u2013205 (2008).","journal-title":"Appl. Anim. Behav. Sci."},{"key":"7169_CR65","doi-asserted-by":"publisher","first-page":"781","DOI":"10.1007\/s10071-020-01388-x","volume":"23","author":"K \u0160andlov\u00e1","year":"2020","unstructured":"\u0160andlov\u00e1, K., Kom\u00e1rkov\u00e1, M. & Ceacero, F. Daddy, daddy cool: stallion\u2013foal relationships in a socially-natural herd of Exmoor ponies. Anim. Cognit. 23, 781\u2013793 (2020).","journal-title":"Anim. Cognit."},{"key":"7169_CR66","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1016\/0168-1591(88)90100-1","volume":"21","author":"LE Boyd","year":"1988","unstructured":"Boyd, L. E. Ontogeny of behavior in Przewalski horses. Appl. Anim. Behav. Sci. 21, 41\u201369 (1988).","journal-title":"Appl. Anim. Behav. Sci."},{"key":"7169_CR67","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s00265-017-2426-x","volume":"72","author":"JW Turner","year":"2018","unstructured":"Turner, J. W., Bills, P. S. & Holekamp, K. E. Ontogenetic change in determinants of social network position in the spotted hyena. Behav. Ecol. Sociobiol. 72, 1\u201315 (2018).","journal-title":"Behav. Ecol. Sociobiol."},{"key":"7169_CR68","doi-asserted-by":"publisher","first-page":"353","DOI":"10.1007\/s10764-015-9826-4","volume":"36","author":"L Kulik","year":"2015","unstructured":"Kulik, L., Amici, F., Langos, D. & Widdig, A. Sex differences in the development of social relationships in rhesus macaques (Macaca mulatta). Int. J. Primatol. 36, 353\u2013376 (2015).","journal-title":"Int. J. Primatol."},{"key":"7169_CR69","doi-asserted-by":"publisher","first-page":"737","DOI":"10.1016\/S0003-3472(89)80106-X","volume":"38","author":"M Nakamichi","year":"1989","unstructured":"Nakamichi, M. Sex differences in social development during the first 4 years in a free-ranging group of Japanese monkeys, Macaca fuscata. Anim. Behav. 38, 737\u2013748 (1989).","journal-title":"Anim. Behav."},{"key":"7169_CR70","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s00265-016-2246-4","volume":"71","author":"MA Whiteside","year":"2017","unstructured":"Whiteside, M. A. et al. Differences in social preference between the sexes during ontogeny drive segregation in a precocial species. Behav. Ecol. Sociobiol. 71, 1\u201313 (2017).","journal-title":"Behav. Ecol. Sociobiol."},{"key":"7169_CR71","doi-asserted-by":"crossref","unstructured":"Rubenstein, D. I. & Nunez, C. M. in Reproductive Skew in Vertebrates: Proximate and Ultimate Causes 196\u2013226 (Cambridge University Press, 2009).","DOI":"10.1017\/CBO9780511641954.010"},{"key":"7169_CR72","unstructured":"Brugman, H., Russel, A. & Nijmegen, X. Annotating multi-media\/multi-modal resources with ELAN. In International Conference on Language Resources and Evaluation 2065\u20132068 (European Language Resources Association, 2004)."},{"key":"7169_CR73","first-page":"1","volume":"1695","author":"G Csardi","year":"2006","unstructured":"Csardi, G. & Nepusz, T. The igraph software. Complex Syst. 1695, 1\u20139 (2006).","journal-title":"Complex Syst."},{"key":"7169_CR74","doi-asserted-by":"crossref","unstructured":"Briatte, F. ggnetwork: geometries to plot networks with \u2018ggplot2\u2019. R Package Version 0.5 1 (2016).","DOI":"10.32614\/CRAN.package.ggnetwork"},{"key":"7169_CR75","doi-asserted-by":"publisher","first-page":"271","DOI":"10.1016\/j.anbehav.2016.12.018","volume":"134","author":"M Fr\u00f6hlich","year":"2017","unstructured":"Fr\u00f6hlich, M., M\u00fcller, G., Zeitr\u00e4g, C., Wittig, R. M. & Pika, S. Gestural development of chimpanzees in the wild: the impact of interactional experience. Anim. Behav. 134, 271\u2013282 (2017).","journal-title":"Anim. Behav."},{"key":"7169_CR76","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1016\/j.anbehav.2016.12.003","volume":"124","author":"M Araya-Salas","year":"2017","unstructured":"Araya-Salas, M., Wojczulanis-Jakubas, K., Phillips, E. M., Mennill, D. J. & Wright, T. F. To overlap or not to overlap: context-dependent coordinated singing in lekking long-billed hermits. Anim. Behav. 124, 57\u201364 (2017).","journal-title":"Anim. Behav."},{"key":"7169_CR77","doi-asserted-by":"crossref","unstructured":"Hex, S. B. & Rubenstein, D. I. A visual signal to achieve joint attention in wild plains zebras, Equus quagga. Anim. Behav. 211, 131\u2013145 (2024b).","DOI":"10.1016\/j.anbehav.2024.02.019"},{"key":"7169_CR78","doi-asserted-by":"publisher","first-page":"1144","DOI":"10.1111\/1365-2656.12418","volume":"84","author":"DR Farine","year":"2015","unstructured":"Farine, D. R. & Whitehead, H. Constructing, conducting and interpreting animal social network analysis. J. Anim. Ecol. 84, 1144\u20131163 (2015).","journal-title":"J. Anim. Ecol."},{"key":"7169_CR79","doi-asserted-by":"publisher","first-page":"066111","DOI":"10.1103\/PhysRevE.70.066111","volume":"70","author":"A Clauset","year":"2004","unstructured":"Clauset, A., Newman, M. E. & Moore, C. Finding community structure in very large networks. Phys. Rev. E 70, 066111 (2004).","journal-title":"Phys. Rev. E"},{"key":"7169_CR80","doi-asserted-by":"publisher","unstructured":"Hex, S. & Rubenstein, D. Multimodal communication of adult, subadult, and infant plains zebras (Equus quagga). Dryad https:\/\/doi.org\/10.5061\/dryad.c866t1ggk (2024).","DOI":"10.5061\/dryad.c866t1ggk"},{"key":"7169_CR81","doi-asserted-by":"publisher","unstructured":"Hex, S. & Rubenstein, D. Multimodal communication of adult, subadult, and infant plains zebras (Equus quagga). Zenodo https:\/\/doi.org\/10.5281\/zenodo.13714766 (2024).","DOI":"10.5281\/zenodo.13714766"}],"container-title":["Communications Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s42003-024-07169-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s42003-024-07169-3","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s42003-024-07169-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,1,11]],"date-time":"2025-01-11T15:03:46Z","timestamp":1736607826000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s42003-024-07169-3"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,1,11]]},"references-count":81,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["7169"],"URL":"https:\/\/doi.org\/10.1038\/s42003-024-07169-3","relation":{},"ISSN":["2399-3642"],"issn-type":[{"value":"2399-3642","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,1,11]]},"assertion":[{"value":"22 May 2024","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"29 October 2024","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"11 January 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"44"}}