{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T01:44:23Z","timestamp":1792028663836,"version":"4.4.0"},"reference-count":87,"publisher":"Springer Science and Business Media LLC","issue":"3","license":[{"start":{"date-parts":[[2026,2,21]],"date-time":"2026-02-21T00:00:00Z","timestamp":1771632000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,2,21]],"date-time":"2026-02-21T00:00:00Z","timestamp":1771632000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"name":"JSPS KAKENHI","award":["(#25K22873, #23K27254 and #19KK0191"],"award-info":[{"award-number":["(#25K22873, #23K27254 and #19KK0191"]}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Primates"],"published-print":{"date-parts":[[2026,5]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    Obesity is an often-stated issue in zoo animals, including primates, and has historically been particularly emphasized for lemurs. By contrast, similar reports for colobine species\u2014which are literally called \u2018slim monkeys\u2019 in at least one other language\u2014are absent. This has been hypothsized to be linked to the colobine foregut fermentation system; diets high in easily digestible carbohydrates might disturb the microbiome in the foregut, similar to acidosis in domestic ruminants, leading to reduced intake and disease rather than obesity, but empirical data is lacking. We compared body mass (BM) data for 14 lemur species (7187 individuals) and 10 colobine species (1443 individuals) with literature data on free-ranging specimens to test this concept. In contrast to expectations, a similar percentage of species had a majority of individuals above the \u2018healthy\u2019 BM range in at least one sex (36% of lemur and 40% of colobine species). However, colobines had a higher percentage of individuals considered \u2018underweight\u2019, especially in the two rarely kept species with the more complex \u2018quadripartite\u2019 stomachs (\n                    <jats:italic>Nasalis larvatus<\/jats:italic>\n                    ,\n                    <jats:italic>Pygathrix nemaeus<\/jats:italic>\n                    ). Neither sample size, sexual size dimorphism, the degree of folivory, nor indications for a seasonal physiology were clearly associated with species\u2019 propensity for zoo obesity. There were no clear historical trends in the development of obesity in these species. We conclude that no general rules about species\u2019 susceptibility to obesity appear to apply. Body mass and condition of zoo primates should be monitored continuously, and feeding regimes possibly be adapted to contain diets of higher fibre levels more similar to those reported in natural diets.\n                  <\/jats:p>","DOI":"10.1007\/s10329-026-01245-4","type":"journal-article","created":{"date-parts":[[2026,2,21]],"date-time":"2026-02-21T06:32:34Z","timestamp":1771655554000},"page":"313-329","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["No systematic difference in obesity susceptibility between two groups of zoo primates with distinct digestive physiologies: hindgut-fermenting lemurs and foregut-fermenting colobines"],"prefix":"10.1007","volume":"67","author":[{"ORCID":"https:\/\/orcid.org\/0009-0001-2691-8763","authenticated-orcid":false,"given":"Christiane","family":"Krauss","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-9142-9551","authenticated-orcid":false,"given":"Elisa","family":"Garand","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3995-419X","authenticated-orcid":false,"given":"Max","family":"Hahn-Klimroth","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9996-064X","authenticated-orcid":false,"given":"Dennis W. H.","family":"M\u00fcller","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6046-6406","authenticated-orcid":false,"given":"Paul W.","family":"Dierkes","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0861-7801","authenticated-orcid":false,"given":"Ikki","family":"Matsuda","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3841-6207","authenticated-orcid":false,"given":"Marcus","family":"Clauss","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-8920-6236","authenticated-orcid":false,"given":"Jo\u00e3o Pedro","family":"Meireles","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,2,21]]},"reference":[{"issue":"6","key":"1245_CR1","doi-asserted-by":"publisher","first-page":"533","DOI":"10.1002\/zoo.20018","volume":"23","author":"G Agoramoorthy","year":"2004","unstructured":"Agoramoorthy G, Alagappasamy C, Hsu MJ (2004) Can proboscis monkeys be successfully maintained in captivity? A case of swings and roundabouts. Zoo Biol 23(6):533\u2013544. https:\/\/doi.org\/10.1002\/zoo.20018","journal-title":"Zoo Biol"},{"key":"1245_CR2","doi-asserted-by":"publisher","first-page":"225","DOI":"10.1016\/j.gecco.2016.06.004","volume":"7","author":"KR Amato","year":"2016","unstructured":"Amato KR, Metcalf JL, Song SJ, Hale VL, Clayton J, Ackermann G, Humphrey G, Niu K, Cui D, Zhao H, Schrenzel MD, Tan CL, Knight R, Braun J (2016) Using the gut microbiota as a novel tool for examining colobine primate GI health. Glob Ecol Conserv 7:225\u2013237. https:\/\/doi.org\/10.1016\/j.gecco.2016.06.004","journal-title":"Glob Ecol Conserv"},{"key":"1245_CR3","doi-asserted-by":"publisher","first-page":"583","DOI":"10.1007\/s10764-020-00163-6","volume":"41","author":"KE Bailey","year":"2020","unstructured":"Bailey KE, Winking JW, Carlson DL, Van Bang T, Long HT (2020) Arm-swinging in the red-shanked douc (Pygathrix nemaeus): implications of body mass. Int J Primatol 41:583\u2013595","journal-title":"Int J Primatol"},{"key":"1245_CR4","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1638\/2021-0023","volume":"53","author":"KC Baker","year":"2022","unstructured":"Baker KC, Georoff TA, Ialeggio DM, Trupkiewicz J, Hopper J, Sutherland-Smith M (2022) Retrospective review of urolithiasis-related morbidity and mortality in Asian colobine monkeys. J Zoo Wildl Med 53:1\u201310","journal-title":"J Zoo Wildl Med"},{"issue":"4","key":"1245_CR5","doi-asserted-by":"publisher","first-page":"763","DOI":"10.1002\/ajpa.24230","volume":"174","author":"N Beeby","year":"2021","unstructured":"Beeby N, Baden AL (2021) Seasonal variability in the diet and feeding ecology of black-and-white ruffed lemurs (Varecia variegata) in Ranomafana National Park, southeastern Madagascar. Am J Phys Anthropol 174(4):763\u2013775. https:\/\/doi.org\/10.1002\/ajpa.24230","journal-title":"Am J Phys Anthropol"},{"key":"1245_CR6","doi-asserted-by":"publisher","first-page":"233","DOI":"10.1007\/BF02737402","volume":"9","author":"EL Bennett","year":"1988","unstructured":"Bennett EL, Sebastian AC (1988) Social organization and ecology of proboscis monkeys (Nasalis larvatus) in mixed coastal forest in Sarawak. Int J Primatol 9:233\u2013255","journal-title":"Int J Primatol"},{"key":"1245_CR7","doi-asserted-by":"crossref","first-page":"e1","DOI":"10.1111\/jpn.12694","volume":"102","author":"F Cabana","year":"2018","unstructured":"Cabana F, Dierenfeld E, Widateti W, Donati G, Nekaris KAI (2018a) Trialling nutrient recommendations for slow lorises (Nycticebus spp.) based on wild feeding ecology. J Anim Physiol Anim Nutr 102:e1\u2013e10","journal-title":"J Anim Physiol Anim Nutr"},{"key":"1245_CR8","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1111\/izy.12172","volume":"52","author":"F Cabana","year":"2018","unstructured":"Cabana F, Jasmi R, Maguire R (2018b) Great ape nutrition: low-sugar and high-fibre diets can lead to increased natural behaviours, decreased regurgitation and reingestion, and reversal of prediabetes. Int Zoo Yearb 52:48\u201361","journal-title":"Int Zoo Yearb"},{"issue":"3","key":"1245_CR9","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1002\/1098-2345(200011)52:3<133::AID-AJP2>3.0.CO;2-#","volume":"52","author":"JL Campbell","year":"2000","unstructured":"Campbell JL, Eisemann JH, Williams CV, Glenn KM (2000) Description of the gastrointestinal tract of five lemur species: Propithecus tattersalli, Propithecus verreauxi coquereli, Varecia variegata, Hapalemur griseus, and Lemur catta. Am J Primatol 52(3):133\u2013142","journal-title":"Am J Primatol"},{"key":"1245_CR10","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1002\/ajp.20080","volume":"64","author":"JL Campbell","year":"2004","unstructured":"Campbell JL, Williams CV, Eisemann JH (2004a) Characterizing gastrointestinal transit time in four lemur species using barium\u2010impregnated polyethylene spheres (BIPS). Am J Primatol 64:309\u2013321","journal-title":"Am J Primatol"},{"key":"1245_CR11","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1002\/ajp.20081","volume":"64","author":"JL Campbell","year":"2004","unstructured":"Campbell JL, Williams CV, Eisemann JH (2004b) Use of total dietary fiber across four lemur species (Propithecus verreauxi coquereli, Hapalemur griseus griseus, Varecia variegata, and Eulemur fulvus): does fiber type affect digestive efficiency? Am J Primatol 64:323\u2013335","journal-title":"Am J Primatol"},{"key":"1245_CR12","first-page":"41","volume":"6","author":"A Caravaggi","year":"2018","unstructured":"Caravaggi A, Plowman A, Wright DJ, Bishop CM (2018) The composition of captive ruffed lemur (Varecia spp.) diets in UK zoological collections, with reference to the problems of obesity and iron storage disease. J Zoo Aquar Res 6:41\u201349","journal-title":"J Zoo Aquar Res"},{"key":"1245_CR13","doi-asserted-by":"crossref","first-page":"444","DOI":"10.1016\/B978-141604047-7.50058-0","volume-title":"Zoo and wild animal medicine. Current therapy 6","author":"M Clauss","year":"2008","unstructured":"Clauss M, Dierenfeld ES (2008) The nutrition of browsers. In: Fowler ME, Miller RE (eds) Zoo and wild animal medicine. Current therapy 6. Saunders Elsevier, St. Louis, pp 444\u2013454"},{"key":"1245_CR14","doi-asserted-by":"crossref","first-page":"274","DOI":"10.1016\/j.cbpa.2008.03.012","volume":"150","author":"M Clauss","year":"2008","unstructured":"Clauss M, Streich WJ, Nunn CL, Ortmann S, Hohmann G, Schwarm A, Hummel J (2008) The influence of natural diet composition, food intake level, and body size on ingesta passage in primates. Comp Biochem Physiol A 150:274\u2013281","journal-title":"Comp Biochem Physiol A"},{"key":"1245_CR15","unstructured":"Clauss M, Janssens GPJ, Pritchard Moore O, Byrne S, Przyby\u0142o M (2025) Adjusting our thinking about feeding animals: humans and low-fiber diets. Zoo Biol (under review)"},{"key":"1245_CR16","first-page":"5","volume-title":"The ecology of arboreal folivores","author":"L Collins","year":"1978","unstructured":"Collins L, Roberts M (1978) Arboreal folivores in captivity - maintenance of a delicate minority. In: Montgomery GG (ed) The ecology of arboreal folivores. Smithsonian Institution, Washington DC, pp 5\u201311"},{"issue":"6","key":"1245_CR17","first-page":"261","volume":"41","author":"D Cousins","year":"1972","unstructured":"Cousins D (1972) Body measurements and weights of wild and captive gorillas, Gorilla gorilla. Zool Garten NF 41(6):261\u2013277","journal-title":"Zool Garten NF"},{"issue":"1","key":"1245_CR18","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1002\/zoo.21718","volume":"42","author":"BA Curry","year":"2023","unstructured":"Curry BA, Drane AL, Atencia R, Feltrer Y, Howatson G, Calvi T, Palmer C, Moittie S, Unwin S, Tremblay JC (2023) Body mass and growth rates in captive chimpanzees (Pan troglodytes) cared for in African wildlife sanctuaries, zoological institutions, and research facilities. Zoo Biol 42(1):98\u2013106","journal-title":"Zoo Biol"},{"key":"1245_CR19","doi-asserted-by":"publisher","DOI":"10.1186\/1472-6793-10-11","volume":"10","author":"A Dal-Pan","year":"2010","unstructured":"Dal-Pan A, Blanc S, Aujard F (2010) Resveratrol suppresses body mass gain in a seasonal non-human primate model of obesity. BMC Physiol 10:11","journal-title":"BMC Physiol"},{"key":"1245_CR20","doi-asserted-by":"publisher","DOI":"10.1638\/2015-0080.1","author":"BC Donadeo","year":"2016","unstructured":"Donadeo BC, Kerr KR, Morris CL, Swanson KS (2016) Estimated composition of diets fed to captive black-and-white ruffed lemurs (Varecia variegata) at 33 US zoological institutions. J Zoo Wildl Med. https:\/\/doi.org\/10.1638\/2015-0080.1","journal-title":"J Zoo Wildl Med"},{"key":"1245_CR21","doi-asserted-by":"publisher","first-page":"1237","DOI":"10.1007\/s00265-007-0354-x","volume":"61","author":"G Donati","year":"2007","unstructured":"Donati G, Bollen A, Borgognini-Tarli S, Ganzhorn JU (2007) Feeding over the 24-h cycle: dietary flexibility of cathemeral collared lemurs (Eulemur collaris). Behav Ecol Sociobiol 61:1237\u20131251","journal-title":"Behav Ecol Sociobiol"},{"key":"1245_CR22","volume":"17","author":"NT Dunham","year":"2022","unstructured":"Dunham NT, Koester DC, Dierenfeld ES, Rimbach R, Pontzer H (2022) Balancing the scales: preliminary investigation of total energy expenditure and daily metabolizable energy intake in Matschie\u2019s tree kangaroo (Dendrolagus matschiei). PLoS ONE 17:e0270570","journal-title":"PLoS ONE"},{"key":"1245_CR23","doi-asserted-by":"crossref","first-page":"537","DOI":"10.1002\/(SICI)1098-2361(1999)18:6<537::AID-ZOO8>3.0.CO;2-F","volume":"18","author":"MS Edwards","year":"1999","unstructured":"Edwards MS, Ullrey DE (1999) Effect of dietary fiber concentration on apparent digestibility and digesta passage in non-human primates. II. Hindgut- and foregut-fermenting folivores. Zoo Biol 18:537\u2013549","journal-title":"Zoo Biol"},{"key":"1245_CR24","first-page":"107","volume-title":"Zoo animal nutrition vol. I","author":"ATC Feistner","year":"1999","unstructured":"Feistner ATC, Mutschler TW (1999) Feeding fussy folivores: nutrition of gentle lemurs. In: Nijboer J, Hatt J-M, Kaumanns W, Beijnen A, Ganslosser U (eds) Zoo animal nutrition vol. I. Filander, F\u00fcrth, pp 107\u2013117"},{"key":"1245_CR25","doi-asserted-by":"publisher","first-page":"157","DOI":"10.1007\/s004420050917","volume":"121","author":"J Fietz","year":"1999","unstructured":"Fietz J, Ganzhorn JU (1999) Feeding ecology of the hibernating primate Cheirogaleus medius: how does it get so fat? Oecologia 121:157\u2013164","journal-title":"Oecologia"},{"key":"1245_CR26","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1111\/ahe.12494","volume":"49","author":"MDO Firmino","year":"2020","unstructured":"Firmino MDO, Pereira HCDS, Carvalho LRRA, Guerra RR (2020) External and digestive system morphology of the Tamandua tetradactyla. Anat Histol Embryol 49:97\u2013104","journal-title":"Anat Histol Embryol"},{"key":"1245_CR27","doi-asserted-by":"crossref","DOI":"10.1017\/CBO9780511975912","volume-title":"The changing body: health, nutrition, and human development in the western world since 1700","author":"R Floud","year":"2011","unstructured":"Floud R, Fogel RW, Harris B, Hong SC (2011) The changing body: health, nutrition, and human development in the western world since 1700. Cambridge University Press, Cambridge"},{"key":"1245_CR28","doi-asserted-by":"crossref","first-page":"537","DOI":"10.1002\/zoo.21865","volume":"43","author":"E Garand","year":"2024","unstructured":"Garand E, Krauss C, M\u00fcller DWH, Davis LR, Cordron D, Clauss M, Miranda F (2024) Larger than life? Body mass records of zoo-managed giant anteaters (Myrmecophaga tridactyla). Zoo Biol 43:537\u2013544","journal-title":"Zoo Biol"},{"issue":"6","key":"1245_CR29","first-page":"353","volume":"55","author":"S Goodchild","year":"2008","unstructured":"Goodchild S, Schwitzer C (2008) The problem of obesity in captive lemurs. Int Zoo News 55(6):353\u2013357","journal-title":"Int Zoo News"},{"key":"1245_CR30","doi-asserted-by":"crossref","first-page":"977","DOI":"10.1111\/joa.14016","volume":"244","author":"H Hanegraef","year":"2024","unstructured":"Hanegraef H, Spoor F (2024) Maxillary morphology of chimpanzees: captive versus wild environments. J Anat 244:977\u2013994","journal-title":"J Anat"},{"key":"1245_CR31","doi-asserted-by":"crossref","first-page":"66","DOI":"10.1111\/brv.12646","volume":"96","author":"SA Heldstab","year":"2021","unstructured":"Heldstab SA, van Schaik CP, M\u00fcller DWH, Rensch E, Bingaman Lackey L, Zerbe P, Hatt J-M, Clauss M, Matsuda I (2021) Reproductive seasonality in primates: patterns, concepts and unsolved questions. Biol Rev 96:66\u201388","journal-title":"Biol Rev"},{"key":"1245_CR32","doi-asserted-by":"crossref","unstructured":"Hladik CM, Charles-Dominique PC, Petter J-J (1980) Feeding strategies of five nocturnal prosimians in the dry forest of the west coast of Madagascar. In: Nocturnal malagasy primates: scology, physiology, and behavior, pp 41\u201373.","DOI":"10.1016\/B978-0-12-169350-3.50007-1"},{"key":"1245_CR33","volume-title":"Feeding ecology in apes and other primates: Ecological, physiological and behavioural aspects","author":"G Hohmann","year":"2006","unstructured":"Hohmann G, Robbins MM, Boesch C (2006) Feeding ecology in apes and other primates: Ecological, physiological and behavioural aspects. Cambridge University Press"},{"key":"1245_CR34","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1111\/j.1748-1090.1973.tb02146.x","volume":"13","author":"U Hollihn","year":"1973","unstructured":"Hollihn U (1973) Remarks on the breeding and maintenance of Colobus monkeys (Colobus guereza), Proboscis monkeys (Nasalis larvatus), and Douc langurs (Pygathrix nemaeus) in zoos. Int Zoo Yearb 13:185\u2013188","journal-title":"Int Zoo Yearb"},{"key":"1245_CR35","doi-asserted-by":"publisher","first-page":"123","DOI":"10.1007\/s10329-022-01031-y","volume":"64","author":"S Hoshino","year":"2023","unstructured":"Hoshino S, Seino S, Azumano A, Tuuga A, Nathan SK, Ramirez Saldivar DA, Goossens B, Salgado-Lynn M, Matsuda I, Yayota M (2023) Modifying the diets of captive proboscis monkeys in a temperate zoo to reduce weight loss and renal disease. Primates 64:123\u2013141","journal-title":"Primates"},{"key":"1245_CR36","doi-asserted-by":"crossref","DOI":"10.1002\/ece3.71069","volume":"15","author":"MT Irwin","year":"2025","unstructured":"Irwin MT, Rahalinarivo V, Ramorasata B, Raharison JL, Ranaivoarisoa JF, Gherardi CN, Rothman JM (2025) More choosy for minor foods: diet, seasonality and food selection in sympatric frugivorous and folivorous lemurs. Ecol Evol 15:e71069","journal-title":"Ecol Evol"},{"key":"1245_CR37","doi-asserted-by":"crossref","first-page":"244","DOI":"10.1080\/10888705.2021.1910032","volume":"25","author":"SN Kamaluddin","year":"2022","unstructured":"Kamaluddin SN, Matsuda I, Md-Zain BM (2022) Activity budget and postural behaviors in orangutans on Bukit Merah Orang Utan island for assessing captive great ape welfare. J Appl Anim Welf Sci 25:244\u2013255","journal-title":"J Appl Anim Welf Sci"},{"key":"1245_CR38","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1159\/000155017","volume":"2","author":"HJ Kuhn","year":"1964","unstructured":"Kuhn HJ (1964) Zur Kenntnis von Bau und Funktion des Magens der Schlankaffen (Colobinae). Folia Primatol 2:193\u2013221","journal-title":"Folia Primatol"},{"key":"1245_CR39","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1159\/000369581","volume":"86","author":"M LaFleur","year":"2015","unstructured":"LaFleur M, Sauther ML (2015) Seasonal feeding ecology of ring-tailed lemurs: a comparison of spiny and gallery forest habitats. Folia Primatol 86:25\u201334","journal-title":"Folia Primatol"},{"issue":"1","key":"1245_CR40","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1002\/(SICI)1520-6505(1998)7:1<8::AID-EVAN3>3.0.CO;2-C","volume":"7","author":"JE Lambert","year":"1998","unstructured":"Lambert JE (1998) Primate digestion: interactions among anatomy, physiology, and feeding ecology. Evol Anthropol Issues News Rev 7(1):8\u201320","journal-title":"Evol Anthropol Issues News Rev"},{"issue":"1","key":"1245_CR41","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1002\/zoo.1430130105","volume":"13","author":"SR Leigh","year":"1994","unstructured":"Leigh SR (1994) Relations between captive and noncaptive weights in anthropoid primates. Zoo Biol 13(1):21\u201343","journal-title":"Zoo Biol"},{"key":"1245_CR42","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1002\/ajp.20187","volume":"67","author":"RJ Lewis","year":"2005","unstructured":"Lewis RJ, Kappeler PM (2005) Seasonality, body condition, and timing of reproduction in Propithecus verreauxi verreauxi in the Kirindy Forest. Am J Primatol 67:347\u2013364. https:\/\/doi.org\/10.1002\/ajp.20187","journal-title":"Am J Primatol"},{"key":"1245_CR43","doi-asserted-by":"crossref","DOI":"10.7717\/peerj.3668","volume":"5","author":"KL Lewton","year":"2017","unstructured":"Lewton KL (2017) The effects of captive versus wild rearing environments on long bone articular surfaces in common chimpanzees (Pan troglodytes). PeerJ 5:e3668","journal-title":"PeerJ"},{"key":"1245_CR44","doi-asserted-by":"crossref","first-page":"873","DOI":"10.1086\/284385","volume":"125","author":"SL Lindstedt","year":"1985","unstructured":"Lindstedt SL, Boyce MS (1985) Seasonality, fasting endurance, and body size in mammals. Am Nat 125:873\u2013878","journal-title":"Am Nat"},{"key":"1245_CR45","doi-asserted-by":"crossref","first-page":"64","DOI":"10.1017\/9781108347150.007","volume-title":"The colobines. Natural history, behaviour and ecological diversity","author":"I Matsuda","year":"2022","unstructured":"Matsuda I, Clauss M (2022) Morphology and physiology of colobine digestive tracts. In: Matsuda I, Grueter CC, Teichroeb JA (eds) The colobines. Natural history, behaviour and ecological diversity. Cambridge University Press, Cambridge, pp 64\u201377"},{"key":"1245_CR46","doi-asserted-by":"publisher","DOI":"10.1038\/srep42774","volume":"7","author":"I Matsuda","year":"2017","unstructured":"Matsuda I, Clauss M, Tuuga A, Sugau J, Hanya G, Yumoto T, Bernard H, Hummel J (2017) Factors affecting leaf selection by foregut-fermenting proboscis monkeys: new insight from in vitro digestibility and toughness of leaves. Sci Rep 7:42774","journal-title":"Sci Rep"},{"key":"1245_CR47","doi-asserted-by":"crossref","DOI":"10.3389\/fvets.2017.00246","volume":"4","author":"I Matsuda","year":"2018","unstructured":"Matsuda I, Bernard H, Tuuga A, Nathan SKSS, Sha J, Osman I, Sipangkui R, Seino S, Asano S, Wong A, Kreuzer M, Ramirez Saldivar DA, Clauss M (2018) Fecal nutrients suggest diets of higher fiber levels in free-ranging than in captive proboscis monkeys (Nasalis larvatus). Front Vet Sci 4:246","journal-title":"Front Vet Sci"},{"key":"1245_CR48","doi-asserted-by":"crossref","first-page":"1608","DOI":"10.1002\/jmor.21052","volume":"280","author":"I Matsuda","year":"2019","unstructured":"Matsuda I, Chapman CA, Clauss M (2019) Colobine forestomach anatomy and diet. J Morphol 280:1608\u20131616","journal-title":"J Morphol"},{"issue":"3","key":"1245_CR49","doi-asserted-by":"publisher","first-page":"473","DOI":"10.1007\/s10329-020-00794-6","volume":"61","author":"I Matsuda","year":"2020","unstructured":"Matsuda I, Ihobe H, Tashiro Y, Yumoto T, Baranga D, Hashimoto C (2020) The diet and feeding behavior of the black-and-white colobus (Colobus guereza) in the Kalinzu Forest, Uganda. Primates 61(3):473\u2013484. https:\/\/doi.org\/10.1007\/s10329-020-00794-6","journal-title":"Primates"},{"key":"1245_CR50","doi-asserted-by":"crossref","DOI":"10.3389\/fevo.2022.795015","volume":"10","author":"I Matsuda","year":"2022","unstructured":"Matsuda I, Hashimoto C, Ihobe H, Yumoto T, Baranga D, Clauss M, Hummel J (2022a) Dietary choices of a foregut-fermenting primate, Colobus guereza: a comprehensive approach including leaf chemical and mechanical properties, digestibility and abundance. Front Ecol Evol 10:795015","journal-title":"Front Ecol Evol"},{"key":"1245_CR51","doi-asserted-by":"crossref","first-page":"735","DOI":"10.1002\/ajpa.24476","volume":"177","author":"I Matsuda","year":"2022","unstructured":"Matsuda I, Takano T, Shintaku Y, Clauss M (2022b) Gastrointestinal morphology and ontogeny of foregut-fermenting primates. Am J Biol Anthropol 177:735\u2013747","journal-title":"Am J Biol Anthropol"},{"issue":"2","key":"1245_CR52","first-page":"108","volume":"13","author":"JP Meireles","year":"2025","unstructured":"Meireles JP, Hahn-Klimroth M, M\u00fcller D, Dierkes P, Clauss M (2025) Body mass of adult zoo hippos Hippopotamidae and how they compare to data from populations in the wild. J Zoo Aquar Res 13(2):108\u2013116","journal-title":"J Zoo Aquar Res"},{"key":"1245_CR53","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1017\/S0962728600028165","volume":"11","author":"VA Melfi","year":"2002","unstructured":"Melfi VA, Feistner ATC (2002) A comparison of the activity budgets of wild and captive Sulawesi crested black macaques (Macaca nigra). Anim Welf 11:213\u2013222","journal-title":"Anim Welf"},{"key":"1245_CR54","volume":"10","author":"EL Mellor","year":"2020","unstructured":"Mellor EL, Cuthill IC, Schwitzer C, Mason GJ, Mendl M (2020) Large lemurs: ecological, demographic and environmental risk factors for weight gain in captivity. Animals 10:1443","journal-title":"Animals"},{"issue":"5","key":"1245_CR55","doi-asserted-by":"publisher","DOI":"10.1002\/ajp.23609","volume":"86","author":"TH Mihaminekena","year":"2024","unstructured":"Mihaminekena TH, Rakotonanahary AN, Frasier CL, Randriahaingo HNT, Sefczek TM, Tinsman J, Randrianarimanana HL, Ravaloharimanitra M, Rakotoarinivo TH, Ratsimbazafy J, King T, Louis EE (2024) Dietary flexibility of the greater bamboo lemur (Prolemur simus), a specialized feeder, in eastern Madagascar. Am J Primatol 86(5):e23609. https:\/\/doi.org\/10.1002\/ajp.23609","journal-title":"Am J Primatol"},{"key":"1245_CR56","doi-asserted-by":"crossref","first-page":"499","DOI":"10.1002\/(SICI)1098-2361(1996)15:5<499::AID-ZOO6>3.0.CO;2-6","volume":"15","author":"J Nijboer","year":"1996","unstructured":"Nijboer J, Dierenfeld ES (1996) Comparison of diets fed to Southeast Asian colobines in North American and European zoos, with emphasis on temperate browse composition. Zoo Biol 15:499\u2013507","journal-title":"Zoo Biol"},{"key":"1245_CR57","doi-asserted-by":"crossref","first-page":"366","DOI":"10.1638\/05-113.1","volume":"37","author":"J Nijboer","year":"2006","unstructured":"Nijboer J, Clauss M, Olsthoorn M, Noordermeer W, Huisman TR, Verheyen C, van der Kuilen J, Streich WJ, Beynen AC (2006b) Effect of diet on the feces quality in Javan langur (Trachypithecus auratus auratus). J Zoo Wildl Med 37:366\u2013372","journal-title":"J Zoo Wildl Med"},{"key":"1245_CR58","unstructured":"Nijboer J, Clauss M (2006) The digestive physiology of colobine primates. In: Nijboer J (ed) Fibre intake and faeces quality in leaf-eating monkeys. PhD thesis, University of Utrecht, The Netherlands, pp 9\u201328"},{"key":"1245_CR59","unstructured":"Nijboer J, Clauss M, Everts H, Beynen A, Fidgett A, Eulenberger K, Hatt J-M, Hume I, Janssens GPJ (2006a) Effect of dietary fibre on the faeces score in colobine monkeys at Dutch Zoos. Zool Libr, pp 145\u2013155"},{"key":"1245_CR60","doi-asserted-by":"publisher","first-page":"1001","DOI":"10.1007\/s10764-006-9056-x","volume":"27","author":"I Norscia","year":"2006","unstructured":"Norscia I, Carrai V, Borgognini-Tarli SM (2006) Influence of dry season and food quality and quantity on behavior and feeding strategy of Propithecus verreauxi in Kirindy, Madagascar. Int J Primatol 27:1001\u20131022","journal-title":"Int J Primatol"},{"key":"1245_CR61","volume-title":"Nutrient requirements nonhuman primates","author":"NRC","year":"2003","unstructured":"NRC (2003) Nutrient requirements nonhuman primates, Second revised edition. National Research Council, The National Academy Press, Washington D.C.","edition":"Second revised"},{"key":"1245_CR62","doi-asserted-by":"crossref","first-page":"222","DOI":"10.1159\/000155018","volume":"2","author":"WC Osman Hill","year":"1964","unstructured":"Osman Hill WC (1964) The maintenance of langurs (Colobidae) in captivity: experiences and some suggestions. Folia Primatol 2:222\u2013231","journal-title":"Folia Primatol"},{"key":"1245_CR63","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1007\/978-1-4899-2412-4_15","volume-title":"Lemur social systems and their ecological basis","author":"ME Pereira","year":"1993","unstructured":"Pereira ME (1993) Seasonal adjustment of growth rate and adult body weight in ringtailed lemurs. In: Kappeler PM, Ganzhorn JU (eds) Lemur social systems and their ecological basis. Plenum Press, New York, pp 205\u2013221"},{"key":"1245_CR64","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1002\/ajp.1350350102","volume":"35","author":"ME Pereira","year":"1995","unstructured":"Pereira ME, Pond CM (1995) Organization of white adipose tissue in lemuridae. Am J Primatol 35:1\u201313","journal-title":"Am J Primatol"},{"key":"1245_CR65","doi-asserted-by":"crossref","first-page":"345","DOI":"10.1002\/zoo.21901","volume":"44","author":"FMAM Pereira","year":"2025","unstructured":"Pereira FMAM, Bicudo ALDC, Mangueira DKA, Yogui DR, Costa MR, Desbiez ALJ, Testa CAEP, Guimar\u00e3es-Okamoto PTC, Dierenfeld ES, Melchert A (2025) Evaluation and ultrasonographic validation of the body condition score in giant anteaters (Myrmecophaga tridactyla). Zoo Biol 44:345\u2013352","journal-title":"Zoo Biol"},{"key":"1245_CR66","doi-asserted-by":"crossref","first-page":"1027","DOI":"10.1016\/S0140-6736(23)02750-2","volume":"403","author":"NH Phelps","year":"2024","unstructured":"Phelps NH, Singleton RK, Zhou B, NCD Risk Factor Collaboration (2024) Worldwide trends in underweight and obesity from 1990 to 2022: a pooled analysis of 3663 population-representative studies with 222 million children, adolescents, and adults. Lancet 403:1027\u20131050","journal-title":"Lancet"},{"key":"1245_CR67","first-page":"171","volume-title":"Zoo animal nutrition IV","author":"S Polowinsky","year":"2009","unstructured":"Polowinsky S, Schwitzer C (2009) Nutritional ecology of the blue-eyed black lemur (Eulemur flavifrons): Integrating in situ and ex situ research to assist the conservation of a critically endangered species. In: Clauss M, Fidgett AL, Hatt J-M, Huisman T, Hummel J, Janssen G, Nijboer J, Plowman A (eds) Zoo animal nutrition IV. Filander Verlag, F\u00fcrth, pp 171\u2013180"},{"key":"1245_CR68","doi-asserted-by":"crossref","DOI":"10.1098\/rstb.2022.0218","volume":"378","author":"H Pontzer","year":"2023","unstructured":"Pontzer H (2023) The provisioned primate: patterns of obesity across lemurs, monkeys, apes and humans. Philos Trans R Soc Lond B Biol Sci 378:20220218","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"issue":"4","key":"1245_CR69","doi-asserted-by":"crossref","first-page":"381","DOI":"10.1006\/jhev.2000.0427","volume":"39","author":"AF Richard","year":"2000","unstructured":"Richard AF, Dewar RE, Schwartz M, Ratsirarson J (2000) Mass change, environmental variability and female fertility in wild Propithecus verreauxi. J Hum Evol 39(4):381\u2013391","journal-title":"J Hum Evol"},{"key":"1245_CR70","volume":"86","author":"R Rimbach","year":"2023","unstructured":"Rimbach R, Pontzer H (2023) Increased physical activity is not related to markers of cardiometabolic health in two lemur species. Am J Primatol 86:e23564","journal-title":"Am J Primatol"},{"key":"1245_CR71","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1002\/zoo.1430020103","volume":"2","author":"CD Schaaf","year":"1983","unstructured":"Schaaf CD, Stuart MD (1983) Reproduction of the mongoose lemur (Lemur mongoz) in captivity. Zoo Biol 2:23\u201338","journal-title":"Zoo Biol"},{"issue":"6","key":"1245_CR72","doi-asserted-by":"crossref","first-page":"429","DOI":"10.1111\/j.1439-0264.2009.00964.x","volume":"38","author":"C Schwitzer","year":"2009","unstructured":"Schwitzer C (2009) Gastrointestinal morphology of the crowned lemur (Eulemur coronatus). Anat Histol Embryol 38(6):429\u2013431","journal-title":"Anat Histol Embryol"},{"key":"1245_CR73","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1002\/zoo.1026","volume":"20","author":"C Schwitzer","year":"2001","unstructured":"Schwitzer C, Kaumanns W (2001) Body weights of ruffed lemurs (Varecia variegata) in European zoos with reference to the problem of obesity. Zoo Biol 20:261\u2013269","journal-title":"Zoo Biol"},{"key":"1245_CR74","doi-asserted-by":"crossref","first-page":"201","DOI":"10.3354\/esr00210","volume":"8","author":"C Schwitzer","year":"2009","unstructured":"Schwitzer C, Kaumanns W (2009) Litter size, infant mortality and female body weight in captive black-and-white ruffed lemurs Varecia variegata. Endangered Species Res 8:201\u2013209","journal-title":"Endangered Species Res"},{"key":"1245_CR75","first-page":"109","volume-title":"Zoo animal nutrition III","author":"C Schwitzer","year":"2006","unstructured":"Schwitzer C, Klumpe K, Kaumanns W (2006a) Energy and nutrient intake, feeding behaviour and activity budget of captive Douc langurs (Pygrathrix n. nemaeus). In: Fidgett A, Clauss M, Eulenberger K, Hatt J, Hume M I, Janssens G, Nijboer J (eds) Zoo animal nutrition III. Filander Verlag, F\u00fcrth, pp 109\u2013124"},{"key":"1245_CR76","unstructured":"Schwitzer C, Schwitzer N, Randriatahina G, Rabarivola C, Kaumanns W (2006b) Programme Sahamalaza: New perspectives for the in situ and ex situ study and conservation of the blue-eyed black lemur (Eulemur macaco flavifrons) in a fragmented habitat. In: Proceedings of the German-Malagasy Research Cooperation in Life and Earth Sciences, 11(135\u2013149)"},{"key":"1245_CR77","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0009860","volume":"5","author":"B Simmen","year":"2010","unstructured":"Simmen B, Bayart F, Rasamimanana H, Zahariev A, Blanc S, Pasquet P (2010) Total energy expenditure and body composition in two free-living sympatric lemurs. PLoS ONE 5:e9860","journal-title":"PLoS ONE"},{"issue":"6","key":"1245_CR78","doi-asserted-by":"crossref","first-page":"523","DOI":"10.1006\/jhev.1996.0122","volume":"32","author":"RJ Smith","year":"1997","unstructured":"Smith RJ, Jungers WL (1997) Body mass in comparative primatology. J Hum Evol 32(6):523\u2013559","journal-title":"J Hum Evol"},{"key":"1245_CR79","doi-asserted-by":"crossref","DOI":"10.1002\/ajp.22996","volume":"81","author":"EH Sterck","year":"2019","unstructured":"Sterck EH, Zijlmans DG, de Vries H, van den Berg LM, van Schaik CP, Langermans JA (2019) Determining overweight and underweight with a new weight\u2010for\u2010height index in captive group\u2010housed macaques. Am J Primatol 81:e22996","journal-title":"Am J Primatol"},{"key":"1245_CR80","unstructured":"Taylor LA, Clauss M, Schwitzer C (2012) Fit or fat: obesity in captive lemurs. In: Proceedings of the European Zoo Nutrition Conference, 7:14"},{"issue":"1","key":"1245_CR81","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1002\/(SICI)1098-2361(1997)16:1<17::AID-ZOO4>3.0.CO;2-E","volume":"16","author":"CJ Terranova","year":"1997","unstructured":"Terranova CJ, Coffman BS (1997) Body weights of wild and captive lemurs. Zoo Biol 16(1):17\u201330","journal-title":"Zoo Biol"},{"key":"1245_CR82","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1007\/s10329-015-0509-8","volume":"57","author":"TR Turner","year":"2016","unstructured":"Turner TR, Cramer JD, Nisbett A, Patrick Gray J (2016) A comparison of adult body size between captive and wild vervet monkeys (Chlorocebus aethiops sabaeus) on the island of St. Kitts. Primates 57:211\u2013220","journal-title":"Primates"},{"key":"1245_CR83","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1017\/S0954422424000076","volume":"38","author":"L Van Mulders","year":"2024","unstructured":"Van Mulders L, Locquet L, Kaandorp C, Janssens GP (2024) An overview of nutritional factors in the aetiopathogenesis of myocardial fibrosis in great apes. Nutr Res Rev 38:37\u201352","journal-title":"Nutr Res Rev"},{"key":"1245_CR84","doi-asserted-by":"crossref","first-page":"14789","DOI":"10.1002\/ece3.8164","volume":"11","author":"L Veylit","year":"2021","unstructured":"Veylit L, S\u00e6ther BE, Gaillard J-M, Baubet E, Gamelon M (2021) Many lifetime growth trajectories for a single mammal. Ecol Evol 11:14789\u201314804","journal-title":"Ecol Evol"},{"key":"1245_CR85","unstructured":"WHO (2025) World Health Organisation. Obesity and overweight. https:\/\/www.who.int\/news-room\/fact-sheets\/detail\/obesity-and-overweight"},{"key":"1245_CR86","doi-asserted-by":"crossref","first-page":"1231","DOI":"10.1002\/ajp.20995","volume":"73","author":"Y Yamanashi","year":"2011","unstructured":"Yamanashi Y, Hayashi M (2011) Assessing the effects of cognitive experiments on the welfare of captive chimpanzees (Pan troglodytes) by direct comparison of activity budget between wild and captive chimpanzees. Am J Primatol 73:1231\u20131238","journal-title":"Am J Primatol"},{"issue":"4","key":"1245_CR87","doi-asserted-by":"crossref","first-page":"607","DOI":"10.2307\/1380844","volume":"65","author":"EM Zullinger","year":"1984","unstructured":"Zullinger EM, Ricklefs RE, Redford KH, Mace GM (1984) Fitting sigmoidal equations to mammalian growth curves. J Mammal 65(4):607\u2013636","journal-title":"J Mammal"}],"container-title":["Primates"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s10329-026-01245-4.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s10329-026-01245-4","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s10329-026-01245-4.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T00:49:45Z","timestamp":1792025385000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s10329-026-01245-4"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,21]]},"references-count":87,"journal-issue":{"issue":"3","published-print":{"date-parts":[[2026,5]]}},"alternative-id":["1245"],"URL":"https:\/\/doi.org\/10.1007\/s10329-026-01245-4","relation":{},"ISSN":["0032-8332","1610-7365"],"issn-type":[{"value":"0032-8332","type":"print"},{"value":"1610-7365","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,2,21]]},"assertion":[{"value":"24 November 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"8 February 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"21 February 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"3 August 2026","order":5,"name":"change_date","label":"Change Date","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Update","order":6,"name":"change_type","label":"Change Type","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The wrong Supplementary file was originally published with this article; it has now been removed.","order":7,"name":"change_details","label":"Change Details","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"14 October 2026","order":8,"name":"change_date","label":"Change Date","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Update","order":9,"name":"change_type","label":"Change Type","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"This article was originally published under the subscription model but it is now published under an Open Access license.","order":10,"name":"change_details","label":"Change Details","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no conflict of interest.","order":1,"name":"Ethics","label":"Conflict of interest","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Not applicable.","order":2,"name":"Ethics","label":"Ethical approval","group":{"name":"EthicsHeading","label":"Declarations"}}]}}