{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T18:07:04Z","timestamp":1788977224424,"version":"build-2803163510"},"reference-count":93,"publisher":"Public Library of Science (PLoS)","issue":"9","license":[{"start":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T00:00:00Z","timestamp":1788912000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100011730","name":"Templeton World Charity Foundation","doi-asserted-by":"publisher","award":["TWCF-2022-30066"],"award-info":[{"award-number":["TWCF-2022-30066"]}],"id":[{"id":"10.13039\/501100011730","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100004870","name":"Research Foundation of The City University of New York","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100004870","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Animal Behavior and Conservation Program at Hunter College"}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS One"],"abstract":"<jats:p>\n                    Inhibitory control\u2014or the suppression of an impulsive, or prepotent, response in favor of a less obvious, but more effective behavior\u2014is an important aspect of executive function, decision-making, and cognitive flexibility. Although elephants have large brains and behave flexibly, little is known about their capacity for inhibitory control, which limits our understanding of its evolution across species. We tested 16 (8M, 8F, 4\u201360 yrs) Asian elephants (\n                    <jats:italic>Elephas maximus<\/jats:italic>\n                    ) at the National Elephant Institute in Thailand using a classic detour paradigm. Elephants first learned to reach their trunk around an opaque box to obtain food from a single opening. We then presented them with a transparent box with front-facing holes, where subjects needed to continue to reach around the box while inhibiting the impulse to reach straight toward the visible food and front-facing odor cues. We found that elephants performed similarly when detouring around both opaque and transparent boxes. This may indicate a strong ability to inhibit the impulse to reach straight forward, but may also indicate that the transparent box did not produce cues strong enough to elicit a prepotent response. In addition, we conducted trials in which we changed the box orientation so that elephants needed to inhibit using a previously learned response (i.e., a direction in which they had previously detoured) in favor of detouring in the newly changed direction. Here, we found evidence for a strong prepotent response: elephants\u2019 latency to detour around the box significantly increased when the box orientation changed. However, detour performance quickly improved across trials in which the orientation was constant, indicating effective inhibition of the previously learned response. Interestingly, performance in these trials worsened with age, revealing potential evidence for cognitive decline. This study provides some of the first evidence for inhibitory control in this species, and furthers our understanding of convergent cognitive evolution across distantly related taxa.\n                  <\/jats:p>","DOI":"10.1371\/journal.pone.0356549","type":"journal-article","created":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T17:23:26Z","timestamp":1788974606000},"page":"e0356549","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":0,"title":["Asian elephants (Elephas maximus) demonstrate inhibitory control in a detour task"],"prefix":"10.1371","volume":"21","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3711-8593","authenticated-orcid":true,"given":"Sydney F.","family":"Hope","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sarah L.","family":"Jacobson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sangpa","family":"Dittakul","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gillian","family":"Schmitt","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mananya","family":"Pla-ard","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marnoch","family":"Yindee","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Taweepoke","family":"Angkawanish","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7597-8818","authenticated-orcid":true,"given":"Joshua M.","family":"Plotnik","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2026,9,9]]},"reference":[{"key":"pone.0356549.ref001","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1146\/annurev-psych-113011-143750","article-title":"Executive functions","volume":"64","author":"A Diamond","year":"2013","journal-title":"Annu Rev Psychol"},{"issue":"3","key":"pone.0356549.ref002","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1016\/j.jecp.2011.04.015","article-title":"The role of inhibitory control in children\u2019s cooperative behaviors during a structured puzzle task","volume":"110","author":"F Giannotta","year":"2011","journal-title":"J Exp Child Psychol"},{"issue":"2","key":"pone.0356549.ref003","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1016\/j.tine.2013.12.001","article-title":"Skills underlying mathematics: The role of executive function in the development of mathematics proficiency","volume":"3","author":"L Cragg","year":"2014","journal-title":"Trends Neurosci Educ"},{"issue":"4","key":"pone.0356549.ref004","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1002\/rrq.54","article-title":"Roles of Attention Shifting and Inhibitory Control in Fourth-Grade Reading Comprehension","volume":"48","author":"MJ Kieffer","year":"2013","journal-title":"Read Res Q"},{"issue":"18","key":"pone.0356549.ref005","doi-asserted-by":"crossref","first-page":"1415","DOI":"10.1016\/j.cub.2008.08.020","article-title":"Fission-fusion dynamics, behavioral flexibility, and inhibitory control in primates","volume":"18","author":"F Amici","year":"2008","journal-title":"Curr Biol"},{"issue":"20","key":"pone.0356549.ref006","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.1323533111","article-title":"The evolution of self-control","volume":"111","author":"EL MacLean","year":"2014","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"4","key":"pone.0356549.ref007","doi-asserted-by":"crossref","first-page":"160104","DOI":"10.1098\/rsos.160104","article-title":"Ravens, New Caledonian crows and jackdaws parallel great apes in motor self-regulation despite smaller brains","volume":"3","author":"C Kabadayi","year":"2016","journal-title":"R Soc Open Sci"},{"issue":"6","key":"pone.0356549.ref008","doi-asserted-by":"crossref","first-page":"1209","DOI":"10.1016\/j.anbehav.2011.03.004","article-title":"Song repertoire size in male song sparrows correlates with detour reaching, but not with other cognitive measures","volume":"81","author":"NJ Boogert","year":"2011","journal-title":"Anim Behav"},{"issue":"7692","key":"pone.0356549.ref009","doi-asserted-by":"crossref","first-page":"364","DOI":"10.1038\/nature25503","article-title":"Cognitive performance is linked to group size and affects fitness in Australian magpies","volume":"554","author":"BJ Ashton","year":"2018","journal-title":"Nature"},{"issue":"1","key":"pone.0356549.ref010","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1007\/s10071-017-1152-0","article-title":"The detour paradigm in animal cognition","volume":"21","author":"C Kabadayi","year":"2018","journal-title":"Anim Cogn"},{"key":"pone.0356549.ref011","doi-asserted-by":"crossref","first-page":"849","DOI":"10.3389\/fpsyg.2017.00849","article-title":"Measures of Dogs\u2019 Inhibitory Control Abilities Do Not Correlate across Tasks","volume":"8","author":"D Brucks","year":"2017","journal-title":"Front Psychol"},{"key":"pone.0356549.ref012","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1016\/j.anbehav.2022.02.007","article-title":"Inhibitory control performance is repeatable over time and across contexts in a wild bird population","volume":"187","author":"GL Davidson","year":"2022","journal-title":"Anim Behav"},{"issue":"20","key":"pone.0356549.ref013","doi-asserted-by":"crossref","first-page":"7816","DOI":"10.1523\/JNEUROSCI.20-20-07816.2000","article-title":"Stress-level cortisol treatment impairs inhibitory control of behavior in monkeys","volume":"20","author":"DM Lyons","year":"2000","journal-title":"J Neurosci"},{"issue":"1","key":"pone.0356549.ref014","doi-asserted-by":"crossref","DOI":"10.1177\/0165025411406864","article-title":"Hypothalamic-pituitary-adrenal axis physiology and cognitive control of behavior in stress inoculated monkeys","volume":"36","author":"KJ Parker","year":"2012","journal-title":"Int J Behav Dev"},{"issue":"1875","key":"pone.0356549.ref015","first-page":"20180150","article-title":"Do detour tasks provide accurate assays of inhibitory control?","volume":"285","author":"JO van Horik","year":"2018","journal-title":"Proc Biol Sci"},{"issue":"3","key":"pone.0356549.ref016","article-title":"Proactive common waxbills make fewer mistakes in a cognitive assay, the detour-reaching task","volume":"74","author":"ACR Gomes","year":"2020","journal-title":"Behav Ecol Sociobiol"},{"issue":"2","key":"pone.0356549.ref017","doi-asserted-by":"crossref","first-page":"187","DOI":"10.1093\/cz\/zoz039","article-title":"Personality traits covary with individual differences in inhibitory abilities in 2 species of fish","volume":"66","author":"T Lucon-Xiccato","year":"2020","journal-title":"Curr Zool"},{"key":"pone.0356549.ref018","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1016\/j.anbehav.2021.08.013","article-title":"Ontogeny and personality affect inhibitory control in guppies, Poecilia reticulata","volume":"180","author":"BB Sava\u015f\u00e7\u0131","year":"2021","journal-title":"Anim Behav"},{"key":"pone.0356549.ref019","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1016\/j.beproc.2016.06.004","article-title":"Testing cognition in the wild: factors affecting performance and individual consistency in two measures of avian cognition","volume":"134","author":"RC Shaw","year":"2017","journal-title":"Behav Processes"},{"issue":"1","key":"pone.0356549.ref020","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1007\/s10071-019-01330-w","article-title":"Response learning confounds assays of inhibitory control on detour tasks","volume":"23","author":"JO van Horik","year":"2020","journal-title":"Anim Cogn"},{"issue":"1","key":"pone.0356549.ref021","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1007\/s10071-013-0633-z","article-title":"Context specificity of inhibitory control in dogs","volume":"17","author":"EE Bray","year":"2014","journal-title":"Anim Cogn"},{"issue":"4","key":"pone.0356549.ref022","doi-asserted-by":"crossref","first-page":"522","DOI":"10.3758\/s13420-018-0354-x","article-title":"Individual performance across motoric self-regulation tasks are not correlated for pet dogs","volume":"46","author":"AAA Vernouillet","year":"2018","journal-title":"Learn Behav"},{"issue":"6","key":"pone.0356549.ref023","doi-asserted-by":"crossref","first-page":"2482","DOI":"10.1111\/brv.70055","article-title":"Inhibitory control tests in non-human animals: validity, reliability, and perspectives","volume":"100","author":"L Loyant","year":"2025","journal-title":"Biol Rev Camb Philos Soc"},{"key":"pone.0356549.ref024","doi-asserted-by":"crossref","first-page":"1835","DOI":"10.3389\/fpsyg.2020.01835","article-title":"Validity of Cognitive Tests for Non-human Animals: Pitfalls and Prospects","volume":"11","author":"MN Schubiger","year":"2020","journal-title":"Front Psychol"},{"key":"pone.0356549.ref025","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1007\/s10071-014-0787-3","article-title":"Individual differences in learning speed, performance accuracy and exploratory behaviour in black-capped chickadees","volume":"18","author":"LM Guillette","year":"2015","journal-title":"Animal Cognition"},{"issue":"11","key":"pone.0356549.ref026","doi-asserted-by":"crossref","first-page":"231476","DOI":"10.1098\/rsos.231476","article-title":"Repeatability and heritability of inhibitory control performance in wild toutouwai (Petroica longipes)","volume":"10","author":"E McCallum","year":"2023","journal-title":"R Soc Open Sci"},{"key":"pone.0356549.ref027","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.anbehav.2019.03.003","article-title":"The effect of experience and olfactory cue in an inhibitory control task in guppies, Poecilia reticulata","volume":"151","author":"M Santac\u00e0","year":"2019","journal-title":"Anim Behav"},{"key":"pone.0356549.ref028","doi-asserted-by":"crossref","first-page":"1125","DOI":"10.1007\/s10764-011-9564-1","article-title":"A comparison of social organization in Asian elephants and African savannah elephants","volume":"33","author":"S de Silva","year":"2012","journal-title":"Int J Primatol"},{"key":"pone.0356549.ref029","doi-asserted-by":"crossref","first-page":"1215","DOI":"10.1163\/1568539X-00003465","article-title":"Collective behaviour of wild Asian elephants in risky situations: how do social groups cross roads?","volume":"154","author":"K Mizuno","year":"2017","journal-title":"Behaviour"},{"issue":"1","key":"pone.0356549.ref030","doi-asserted-by":"crossref","first-page":"8678","DOI":"10.1038\/s41598-019-45130-1","article-title":"All-Male Groups in Asian Elephants: A Novel, Adaptive Social Strategy in Increasingly Anthropogenic Landscapes of Southern India","volume":"9","author":"N Srinivasaiah","year":"2019","journal-title":"Sci Rep"},{"issue":"3","key":"pone.0356549.ref031","doi-asserted-by":"crossref","first-page":"406","DOI":"10.1037\/com0000281","article-title":"Multiple assessments of personality and problem-solving performance in captive Asian elephants (Elephas maximus) and African savanna elephants (Loxodonta africana)","volume":"135","author":"LP Barrett","year":"2021","journal-title":"J Comp Psychol"},{"key":"pone.0356549.ref032","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1016\/j.anbehav.2023.08.019","article-title":"Innovating to solve a novel puzzle: wild Asian elephants vary in their ability to problem solve","volume":"205","author":"SL Jacobson","year":"2023","journal-title":"Anim Behav"},{"issue":"3","key":"pone.0356549.ref033","doi-asserted-by":"crossref","first-page":"657","DOI":"10.1007\/s10071-021-01576-3","article-title":"Persistence is key: investigating innovative problem solving by Asian elephants using a novel multi-access box","volume":"25","author":"SL Jacobson","year":"2022","journal-title":"Anim Cogn"},{"issue":"9","key":"pone.0356549.ref034","article-title":"Cooperating elephants mitigate competition until the stakes get too high","volume":"19","author":"L-L Li","year":"2021","journal-title":"PLoS Biol"},{"issue":"12","key":"pone.0356549.ref035","doi-asserted-by":"crossref","first-page":"5116","DOI":"10.1073\/pnas.1101765108","article-title":"Elephants know when they need a helping trunk in a cooperative task","volume":"108","author":"JM Plotnik","year":"2011","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"5","key":"pone.0356549.ref036","doi-asserted-by":"crossref","first-page":"645","DOI":"10.1007\/s10071-019-01261-6","article-title":"Inhibitory control and memory in the search process for a modified problem in grey squirrels, Sciurus carolinensis","volume":"22","author":"PKY Chow","year":"2019","journal-title":"Anim Cogn"},{"issue":"2","key":"pone.0356549.ref037","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0118469","article-title":"The effect of domestication on inhibitory control: wolves and dogs compared","volume":"10","author":"S Marshall-Pescini","year":"2015","journal-title":"PLoS One"},{"issue":"2","key":"pone.0356549.ref038","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0147753","article-title":"Inhibitory Control, but Not Prolonged Object-Related Experience Appears to Affect Physical Problem-Solving Performance of Pet Dogs","volume":"11","author":"CA M\u00fcller","year":"2016","journal-title":"PLoS One"},{"key":"pone.0356549.ref039","doi-asserted-by":"crossref","first-page":"131","DOI":"10.3819\/CCBR.2020.150006","article-title":"The importance of sensory perception in an elephant\u2019s cognitive world","volume":"15","author":"SL Jacobson","year":"2020","journal-title":"CCBR"},{"issue":"6","key":"pone.0356549.ref040","doi-asserted-by":"crossref","first-page":"907","DOI":"10.1007\/s10071-019-01283-0","article-title":"The use of a human\u2019s location and social cues by Asian elephants in an object-choice task","volume":"22","author":"O Ketchaisri","year":"2019","journal-title":"Anim Cogn"},{"issue":"4","key":"pone.0356549.ref041","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0061174","article-title":"Visual cues given by humans are not sufficient for Asian elephants (Elephas maximus) to find hidden food","volume":"8","author":"JM Plotnik","year":"2013","journal-title":"PLoS One"},{"issue":"5516","key":"pone.0356549.ref042","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1126\/science.1057895","article-title":"Matriarchs as repositories of social knowledge in African elephants","volume":"292","author":"K McComb","year":"2001","journal-title":"Science"},{"key":"pone.0356549.ref043","doi-asserted-by":"crossref","DOI":"10.1093\/oso\/9780195107784.001.0001","volume-title":"The living elephants: evolutionary ecology, behavior, and conservation","author":"R Sukumar","year":"2003"},{"issue":"6","key":"pone.0356549.ref044","doi-asserted-by":"crossref","first-page":"1357","DOI":"10.1016\/j.anbehav.2004.08.018","article-title":"The socioecology of elephants: analysis of the processes creating multitiered social structures","volume":"69","author":"G Wittemyer","year":"2005","journal-title":"Anim Behav"},{"issue":"6","key":"pone.0356549.ref045","doi-asserted-by":"crossref","first-page":"1093","DOI":"10.1016\/j.anbehav.2011.02.013","article-title":"Association patterns of African elephants in all-male groups: the role of age and genetic relatedness","volume":"81","author":"PI Chiyo","year":"2011","journal-title":"Anim Behav"},{"issue":"1","key":"pone.0356549.ref046","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/j.1748-1090.2006.00001.x","article-title":"A brief review of the status, distribution and biology of wild Asian elephants Elephas maximus","volume":"40","author":"R Sukumar","year":"2006","journal-title":"Int Zoo Yearb"},{"key":"pone.0356549.ref047","doi-asserted-by":"crossref","DOI":"10.3389\/fragi.2021.726714","article-title":"Aging: what we can learn from elephants","volume":"2","author":"DE Chusyd","year":"2021","journal-title":"Front Aging"},{"issue":"4","key":"pone.0356549.ref048","doi-asserted-by":"crossref","first-page":"755","DOI":"10.1007\/s10071-020-01387-y","article-title":"The development of problem-solving abilities in a population of candidate detection dogs (Canis familiaris)","volume":"23","author":"L Lazarowski","year":"2020","journal-title":"Anim Cogn"},{"issue":"1","key":"pone.0356549.ref049","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1016\/0273-2297(92)90003-K","article-title":"The rise and fall of the inhibitory mechanism: Toward a unified theory of cognitive development and aging","volume":"12","author":"FN Dempster","year":"1992","journal-title":"Dev Rev"},{"key":"pone.0356549.ref050","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.neurobiolaging.2018.10.003","article-title":"Key periods of cognitive decline in a nonhuman primate model of cognitive aging, the common marmoset (Callithrix jacchus)","volume":"74","author":"A Sadoun","year":"2019","journal-title":"Neurobiol Aging"},{"issue":"1722","key":"pone.0356549.ref051","first-page":"3270","article-title":"Leadership in elephants: the adaptive value of age","volume":"278","author":"K McComb","year":"2011","journal-title":"Proc Biol Sci"},{"issue":"9","key":"pone.0356549.ref052","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0162018","article-title":"Detour Behavior of Mice Trained with Transparent, Semitransparent and Opaque Barriers","volume":"11","author":"GR Juszczak","year":"2016","journal-title":"PLoS One"},{"issue":"4","key":"pone.0356549.ref053","doi-asserted-by":"crossref","first-page":"463","DOI":"10.1111\/1467-7687.00188","article-title":"Going around transparent and grid\u2010like barriers: detour ability as a perception\u2013action skill","volume":"4","author":"JJ Lockman","year":"2001","journal-title":"Dev Sci"},{"issue":"2","key":"pone.0356549.ref054","doi-asserted-by":"crossref","first-page":"594","DOI":"10.1016\/S0003-3472(83)80083-9","article-title":"Route planning in cats, in relation to the visibility of the goal","volume":"31","author":"B Poucet","year":"1983","journal-title":"Anim Behav"},{"issue":"11","key":"pone.0356549.ref055","doi-asserted-by":"crossref","first-page":"211564","DOI":"10.1098\/rsos.211564","article-title":"Heterogeneity of performances in several inhibitory control tasks: male rhesus macaques are more easily distracted than females","volume":"8","author":"L Loyant","year":"2021","journal-title":"R Soc Open Sci"},{"key":"pone.0356549.ref056","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1186\/s13293-016-0065-y","article-title":"Sex dependency of inhibitory control functions","volume":"7","author":"FA Mansouri","year":"2016","journal-title":"Biol Sex Differ"},{"issue":"6","key":"pone.0356549.ref057","doi-asserted-by":"crossref","first-page":"986","DOI":"10.1111\/j.1469-8986.2008.00693.x","article-title":"Gender differences in behavioral inhibitory control: ERP evidence from a two-choice oddball task","volume":"45","author":"J Yuan","year":"2008","journal-title":"Psychophysiology"},{"key":"pone.0356549.ref058","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1016\/j.anbehav.2022.09.011","article-title":"The effect of sex, age and boldness on inhibitory control","volume":"193","author":"IM Vinogradov","year":"2022","journal-title":"Anim Behav"},{"issue":"8","key":"pone.0356549.ref059","doi-asserted-by":"crossref","first-page":"2448","DOI":"10.3390\/ani11082448","article-title":"Effect of Sex and Reproductive Status on Inhibitory Control and Social Cognition in the Domestic Dog (Canis familiaris)","volume":"11","author":"S Junttila","year":"2021","journal-title":"Animals (Basel)"},{"key":"pone.0356549.ref060","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1016\/j.bbr.2016.06.032","article-title":"Sex differences in inhibitory control in socially-housed baboons (Papio papio)","volume":"312","author":"A Lacreuse","year":"2016","journal-title":"Behav Brain Res"},{"key":"pone.0356549.ref061","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1007\/s10071-025-02009-1","article-title":"Do Asian elephants plan for mutually-exclusive outcomes?","volume":"28","author":"SF Hope","year":"2025","journal-title":"Anim Cogn"},{"key":"pone.0356549.ref062","author":"G Schmitt","year":"2025"},{"issue":"11","key":"pone.0356549.ref063","doi-asserted-by":"crossref","first-page":"1325","DOI":"10.1111\/2041-210X.12584","article-title":"BORIS: a free, versatile open\u2010source event\u2010logging software for video\/audio coding and live observations","volume":"7","author":"O Friard","year":"2016","journal-title":"Methods Ecol Evol"},{"key":"pone.0356549.ref064","volume-title":"R: A language and environment for statistical computing","author":"R Core Team","year":"2025"},{"issue":"1","key":"pone.0356549.ref065","doi-asserted-by":"crossref","DOI":"10.18637\/jss.v067.i01","article-title":"Fitting Linear Mixed-Effects Models Using lme4","volume":"67","author":"D Bates","year":"2015","journal-title":"J Stat Soft"},{"key":"pone.0356549.ref066","volume-title":"An R companion to applied regression","author":"J Fox","year":"2019","edition":"2"},{"key":"pone.0356549.ref067","author":"H Wickham","year":"2023"},{"issue":"1","key":"pone.0356549.ref068","doi-asserted-by":"crossref","DOI":"10.18637\/jss.v040.i01","article-title":"The Split-Apply-Combine Strategy for Data Analysis","volume":"40","author":"H Wickham","year":"2011","journal-title":"J Stat Soft"},{"key":"pone.0356549.ref069","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-319-24277-4","volume-title":"ggplot2: Elegant graphics for data analysis","author":"H Wickham","year":"2016"},{"key":"pone.0356549.ref070","author":"JB Arnold","year":"2025"},{"issue":"11","key":"pone.0356549.ref071","doi-asserted-by":"crossref","first-page":"1639","DOI":"10.1111\/2041-210X.12797","article-title":"rptR: repeatability estimation and variance decomposition by generalized linear mixed\u2010effects models","volume":"8","author":"MA Stoffel","year":"2017","journal-title":"Methods Ecol Evol"},{"key":"pone.0356549.ref072","author":"M Gamer","year":"2019"},{"issue":"2","key":"pone.0356549.ref073","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1016\/j.jcm.2016.02.012","article-title":"A Guideline of Selecting and Reporting Intraclass Correlation Coefficients for Reliability Research","volume":"15","author":"TK Koo","year":"2016","journal-title":"J Chiropr Med"},{"key":"pone.0356549.ref074","doi-asserted-by":"crossref","DOI":"10.7551\/mitpress\/9383.001.0001","article-title":"Convergent cognitive evolution across animal taxa: Comparisons of chimpanzees, corvids, and elephants.","volume-title":"The Conceptual Mind: New Directions in the Study of Concepts","author":"Margolis, Eric, Laurence, Stephen","year":"2015"},{"issue":"1","key":"pone.0356549.ref075","doi-asserted-by":"crossref","first-page":"32623","DOI":"10.1038\/s41598-025-16994-3","article-title":"Asian elephants (Elephas maximus) recognise human visual attention from body and face orientation","volume":"15","author":"H-L Jim","year":"2025","journal-title":"Sci Rep"},{"issue":"45","key":"pone.0356549.ref076","doi-asserted-by":"crossref","first-page":"17053","DOI":"10.1073\/pnas.0608062103","article-title":"Self-recognition in an Asian elephant","volume":"103","author":"JM Plotnik","year":"2006","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"3","key":"pone.0356549.ref077","doi-asserted-by":"crossref","first-page":"279","DOI":"10.26451\/abc.05.03.03.2018","article-title":"Asian Elephants (Elephas maximus) Discriminate Between Familiar and Unfamiliar Human Visual and Olfactory Cues","volume":"5","author":"EJ Polla","year":"2018","journal-title":"AB&C"},{"issue":"4","key":"pone.0356549.ref078","doi-asserted-by":"crossref","first-page":"329","DOI":"10.26451\/abc.10.04.03.2023","article-title":"The Role of Umwelt in Animal Curiosity: A Within and Between Species Comparison of Novelty Exploration in Mongooses","volume":"10","author":"K Birchmeier","year":"2023","journal-title":"AB&C"},{"issue":"4","key":"pone.0356549.ref079","doi-asserted-by":"crossref","first-page":"599","DOI":"10.1016\/S0003-3472(05)80243-X","article-title":"The seeking of novelty and its relation to play","volume":"42","author":"DGM Wood-Gush","year":"1991","journal-title":"Anim Behav"},{"issue":"1","key":"pone.0356549.ref080","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1037\/0278-7393.17.1.163","article-title":"Age and inhibition","volume":"17","author":"L Hasher","year":"1991","journal-title":"J Exp Psychol Learn Mem Cogn"},{"issue":"12","key":"pone.0356549.ref081","doi-asserted-by":"crossref","first-page":"5085","DOI":"10.1002\/hbm.24347","article-title":"Structural and functional cerebral bases of diminished inhibitory control during healthy aging","volume":"39","author":"S Hu","year":"2018","journal-title":"Hum Brain Mapp"},{"issue":"2","key":"pone.0356549.ref082","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1002\/ajp.20136","article-title":"Age differences in neophilia, exploration, and innovation in family groups of callitrichid monkeys","volume":"66","author":"RL Kendal","year":"2005","journal-title":"Am J Primatol"},{"issue":"5","key":"pone.0356549.ref083","doi-asserted-by":"crossref","first-page":"701","DOI":"10.1007\/s10071-010-0319-8","article-title":"Inter-individual and age differences in exploration, neophobia and problem-solving ability in a Neotropical raptor (Milvago chimango)","volume":"13","author":"LM Biondi","year":"2010","journal-title":"Anim Cogn"},{"key":"pone.0356549.ref084","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1093\/acprof:oso\/9780198526223.003.0008","article-title":"The Role of Neophobia and Neophilia in the Development of Innovative Behaviour of Birds.","volume-title":"Animal Innovation","author":"R. Greenberg","year":"2003"},{"issue":"1","key":"pone.0356549.ref085","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1163\/156853966X00065","article-title":"Responsiveness to objects in free-ranging Japanese monkeys","volume":"26","author":"EW Menzel","year":"1966","journal-title":"Behaviour"},{"key":"pone.0356549.ref086","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1016\/j.anbehav.2018.01.025","article-title":"The adaptive significance of age-dependent changes in the tendency of individuals to explore","volume":"138","author":"TN Sherratt","year":"2018","journal-title":"Anim Behav"},{"issue":"1","key":"pone.0356549.ref087","doi-asserted-by":"crossref","first-page":"15480","DOI":"10.1038\/s41598-021-95048-w","article-title":"Handler familiarity helps to improve working performance during novel situations in semi-captive Asian elephants","volume":"11","author":"O Liehrmann","year":"2021","journal-title":"Sci Rep"},{"issue":"7","key":"pone.0356549.ref088","doi-asserted-by":"crossref","first-page":"250896","DOI":"10.1098\/rsos.250896","article-title":"Wild elephants vary in their attraction to novelty across an anthropogenic landscape gradient","volume":"12","author":"SL Jacobson","year":"2025","journal-title":"R Soc Open Sci"},{"issue":"1963","key":"pone.0356549.ref089","first-page":"20211728","article-title":"Linking cognition to ecology in wild sympatric mouse lemur species","volume":"288","author":"J Henke-von der Malsburg","year":"2021","journal-title":"Proc Biol Sci"},{"issue":"25","key":"pone.0356549.ref090","doi-asserted-by":"crossref","first-page":"12566","DOI":"10.1073\/pnas.1818284116","article-title":"Elephants have a nose for quantity","volume":"116","author":"JM Plotnik","year":"2019","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pone.0356549.ref091","author":"C Williams","year":"2020"},{"key":"pone.0356549.ref092","volume-title":"Review of human-elephant conflict mitigation measures practiced in South Asia","author":"P Fernando","year":"2008"},{"key":"pone.0356549.ref093","doi-asserted-by":"crossref","first-page":"122","DOI":"10.3389\/fevo.2018.00122","article-title":"Taking the Elephants\u2019 Perspective: Remembering Elephant Behavior, Cognition and Ecology in Human-Elephant Conflict Mitigation","volume":"6","author":"HS Mumby","year":"2018","journal-title":"Front Ecol Evol"}],"container-title":["PLOS One"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0356549","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T17:23:37Z","timestamp":1788974617000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0356549"}},"subtitle":[],"editor":[{"given":"Gregg","family":"Roman","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2026,9,9]]},"references-count":93,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,9,9]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pone.0356549","relation":{},"ISSN":["1932-6203"],"issn-type":[{"value":"1932-6203","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,9]]}}}