{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T17:59:10Z","timestamp":1791395950673,"version":"4.3.3"},"reference-count":148,"publisher":"Public Library of Science (PLoS)","issue":"10","license":[{"start":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T00:00:00Z","timestamp":1791331200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100010841","name":"Paleontological Society","doi-asserted-by":"publisher","award":["Allison R. \u201cPete\u201d Palmer Award"],"award-info":[{"award-number":["Allison R. \u201cPete\u201d Palmer Award"]}],"id":[{"id":"10.13039\/100010841","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100007859","name":"University of Kansas","doi-asserted-by":"publisher","award":["University of Kansas Biodiversity Institute and Natural History Museum Panorama Small Grant"],"award-info":[{"award-number":["University of Kansas Biodiversity Institute and Natural History Museum Panorama Small Grant"]}],"id":[{"id":"10.13039\/100007859","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100025772","name":"David B. Jones Foundation","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100025772","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS One"],"abstract":"<jats:p>\n                    Uintatheres, mammals belonging to the extinct order Dinocerata, are among the most recognizable of all Paleogene (~66\u201323 Ma) organisms. Unmistakable for their bizarre skulls with multiple pairs of horns and saber-like upper canines, uintatheres have captivated paleontologists since the late nineteenth century. Since their initial discovery, uintatheres have been regarded as a classic example of dramatic sexual dimorphism in the fossil record, with males purported to be larger and possess more prominent horns and canines than females. However, the hypothesis that uintatheres were highly sexually dimorphic has never been formally tested. Here, traditional, linear morphometrics are implemented with a collection including most known skulls of\n                    <jats:italic>Uintatherium anceps<\/jats:italic>\n                    to quantify patterns of cranial variation within this taxon. Use of univariate and multivariate statistical approaches provides some evidence of dimorphism in\n                    <jats:italic>Uintatherium<\/jats:italic>\n                    , particularly in the length of the canines, but the expected signals of strong sexual dimorphism are generally lacking. To verify the approaches used, the same suite of analyses was performed on a similarly sized dataset from\n                    <jats:italic>Bison bison<\/jats:italic>\n                    . Statistical signals of sexual dimorphism are clearer in this extant taxon. Comparison with other extant mammalian herbivores reveals that the estimated level of sexual dimorphism in skull length for\n                    <jats:italic>Uintatherium<\/jats:italic>\n                    is not especially high. In light of these findings, as well as the current understanding of uintathere systematics and paleoecology, it is argued that extreme sexual dimorphism should not be treated as the null hypothesis for this clade. These results may point to a broader pattern in mammalian evolution in which traits like horns and large canines emerge in both males and females of a species despite being primarily selected for in one sex.\n                  <\/jats:p>","DOI":"10.1371\/journal.pone.0345236","type":"journal-article","created":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T17:26:53Z","timestamp":1791394013000},"page":"e0345236","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":0,"title":["Linear morphometrics fail to support strong sexual dimorphism in Uintatherium anceps"],"prefix":"10.1371","volume":"21","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6739-7554","authenticated-orcid":true,"given":"Kevin D.","family":"Mulcahy","sequence":"first","affiliation":[{"name":"University of Kansas","place":["Lawrence, Kansas, United States of America"],"department":["Biodiversity Institute and Natural History Museum"]},{"name":"University of Kansas","place":["Lawrence, Kansas, United States of America"],"department":["Department of Ecology and Evolutionary Biology"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2026,10,7]]},"reference":[{"key":"pone.0345236.ref001","volume-title":"On the origin of species","author":"C Darwin","year":"1859"},{"key":"pone.0345236.ref002","volume-title":"The descent of man, and selection in relation to sex","author":"C Darwin","year":"1871"},{"key":"pone.0345236.ref003","doi-asserted-by":"crossref","first-page":"429","DOI":"10.1146\/annurev.an.14.100185.002241","article-title":"Sexual dimorphism","volume":"14","author":"DW Frayer","year":"1985","journal-title":"Annu Rev Anthropol"},{"issue":"4","key":"pone.0345236.ref004","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1086\/416458","article-title":"Ecological causes for the evolution of sexual dimorphism: a review of the evidence","volume":"64","author":"R Shine","year":"1989","journal-title":"Q Rev Biol"},{"key":"pone.0345236.ref005","volume-title":"Encyclopedia of animal behavior","author":"MD Breed","year":"2010"},{"key":"pone.0345236.ref006","first-page":"136","article-title":"Parental investment and sexual selection.","volume-title":"Sexual selection and the descent of man, 1871-1971","author":"RL Trivers","year":"1972"},{"issue":"3","key":"pone.0345236.ref007","doi-asserted-by":"crossref","first-page":"450","DOI":"10.2307\/1382127","article-title":"Sexual dimorphism in the greater one-horned rhinoceros (Rhinoceros unicornis","volume":"72","author":"E Dinerstein","year":"1991","journal-title":"). J Mammal"},{"issue":"1","key":"pone.0345236.ref008","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1111\/j.1095-8312.2008.01035.x","article-title":"The evolution of sexual size dimorphism in cottontail rabbits (Sylvilagus, Leporidae","volume":"95","author":"CM Davis","year":"2008","journal-title":"). Biol J Linn Soc Lond"},{"key":"pone.0345236.ref009","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.anbehav.2017.03.030","article-title":"Role of fecundity selection on the evolution of sexual size dimorphism in mammals","volume":"128","author":"M Cassini","year":"2017","journal-title":"Anim Behav"},{"issue":"1","key":"pone.0345236.ref010","doi-asserted-by":"crossref","first-page":"1872","DOI":"10.1038\/s41467-024-45739-5","article-title":"New estimates indicate that males are not larger than females in most mammal species","volume":"15","author":"KJ Tombak","year":"2024","journal-title":"Nat Commun"},{"key":"pone.0345236.ref011","first-page":"402","article-title":"Sexual dimorphisms and breeding systems in pinnipeds, ungulates, primates, and humans.","volume-title":"Evolutionary biology and human social behavior: an anthropological perspective","author":"RD Alexander","year":"1979"},{"issue":"1285","key":"pone.0345236.ref012","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1098\/rspb.1989.0027","article-title":"Mammalian mating systems","volume":"236","author":"TH Clutton-Brock","year":"1989","journal-title":"Proc R Soc Lond B Biol Sci"},{"issue":"5","key":"pone.0345236.ref013","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1016\/0169-5347(89)90212-7","article-title":"The evolution of sexual dimorphism in animals: hypotheses and tests","volume":"4","author":"AV Hedrick","year":"1989","journal-title":"Trends Ecol Evol"},{"key":"pone.0345236.ref014","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1086\/596461","article-title":"Female mate choice in mammals","volume":"84","author":"T Clutton-Brock","year":"2009","journal-title":"Q Rev Biol."},{"issue":"1","key":"pone.0345236.ref015","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1111\/j.1420-9101.2008.01633.x","article-title":"Male-male competition, female mate choice and their interaction: determining total sexual selection","volume":"22","author":"J Hunt","year":"2009","journal-title":"J Evol Biol"},{"key":"pone.0345236.ref016","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1016\/j.anbehav.2018.02.004","article-title":"While males fight, females choose: male phenotypic quality informs female mate choice in mammals","volume":"138","author":"DL Morina","year":"2018","journal-title":"Anim Behav"},{"key":"pone.0345236.ref017","volume-title":"The differences between the sexes","author":"RV Short","year":"1994"},{"key":"pone.0345236.ref018","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1002\/(SICI)1098-2345(1997)43:1<1::AID-AJP1>3.0.CO;2-0","article-title":"Relative growth, ontogeny, and sexual dimorphism in gorilla (Gorilla gorilla gorilla and G. g. beringei): evolutionary and ecological considerations","volume":"43","author":"AB Taylor","year":"1997","journal-title":"Am J Primatol"},{"issue":"5585","key":"pone.0345236.ref019","doi-asserted-by":"crossref","first-page":"1339","DOI":"10.1126\/science.1073257","article-title":"Sexual selection, temperature, and the lion\u2019s mane","volume":"297","author":"PM West","year":"2002","journal-title":"Science"},{"issue":"2","key":"pone.0345236.ref020","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1086\/409310","article-title":"Mammals in which females are larger than males","volume":"51","author":"K Ralls","year":"1976","journal-title":"Q Rev Biol"},{"issue":"981","key":"pone.0345236.ref021","doi-asserted-by":"crossref","first-page":"917","DOI":"10.1086\/283223","article-title":"Sexual dimorphism in mammals: avian models and unanswered questions","volume":"111","author":"K Ralls","year":"1977","journal-title":"Am Nat"},{"issue":"3","key":"pone.0345236.ref022","doi-asserted-by":"crossref","first-page":"540","DOI":"10.1086\/286004","article-title":"A comparative analysis of allometry for sexual size dimorphism: assessing Rensch\u2019s rule","volume":"149","author":"E Abouheif","year":"1997","journal-title":"Am Nat"},{"key":"pone.0345236.ref023","article-title":"Primate ecology and social organization","volume":"183","author":"TH Clutton-Brock","year":"1977","journal-title":"J Zool"},{"issue":"1","key":"pone.0345236.ref024","doi-asserted-by":"crossref","first-page":"40","DOI":"10.1007\/s004420050023","article-title":"The relationship between ecological segregation and sexual body size dimorphism in large herbivores","volume":"124","author":"A Mysterud","year":"2000","journal-title":"Oecologia"},{"key":"pone.0345236.ref025","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1002\/ajpa.10011","article-title":"Sexual dimorphism in primate evolution","author":"JM Plavcan","year":"2001","journal-title":"Am J Phys Anthropol"},{"issue":"6","key":"pone.0345236.ref026","doi-asserted-by":"crossref","first-page":"1276","DOI":"10.1111\/j.0014-3820.2002.tb01438.x","article-title":"The origins of sexual dimorphism in body size in ungulates","volume":"56","author":"FJ P\u00e9rez-Barber\u00eda","year":"2002","journal-title":"Evolution"},{"issue":"1","key":"pone.0345236.ref027","doi-asserted-by":"crossref","first-page":"164","DOI":"10.1671\/0272-4634(2002)022[0164:SDSBAI]2.0.CO;2","article-title":"Sexual dimorphism, social behavior, and intrasexual competition in large Pleistocene carnivorans","volume":"22","author":"B Van Valkenburgh","year":"2002","journal-title":"J Vertebr Paleontol"},{"key":"pone.0345236.ref028","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1111\/j.1096-3642.1946.tb00095.x","article-title":"A Sexual difference in the skulls of Asiatic rhinoceroses","volume":"115","author":"RI Pocock","year":"1946","journal-title":"Proc Zool Soc Lond"},{"key":"pone.0345236.ref029","doi-asserted-by":"crossref","first-page":"106288","DOI":"10.1016\/j.rvsc.2026.106288","article-title":"Skull morphometry and sexual dimorphism in the Kyrgyz horse: a multivariate size-corrected analysis","volume":"210","author":"HH Ar\u0131","year":"2026","journal-title":"Res Vet Sci"},{"key":"pone.0345236.ref030","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1017\/S2078633614000514","article-title":"No functional sexual dimorphism in Minorcan horse assessed by geometric morphometric methods","volume":"56","author":"PM Par\u00e9s-Casanova","year":"2015","journal-title":"Anim Genet Res"},{"key":"pone.0345236.ref031","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1590\/1809-4392202101382","article-title":"Ontogenetic skull variation in an Amazonian population of lowland tapir, Tapirus terrestris (Mammalia: Perissodactyla) in the department of Loreto, Peru","volume":"51","author":"RR Rojas","year":"2021","journal-title":"Acta Amaz"},{"key":"pone.0345236.ref032","doi-asserted-by":"crossref","first-page":"166","DOI":"10.1644\/863.1","article-title":"Tapirus pinchaque (Perissodactyla: Tapiridae","volume":"42","author":"M Padilla","year":"2010","journal-title":"Mamm Species"},{"key":"pone.0345236.ref033","doi-asserted-by":"crossref","DOI":"10.1017\/CBO9780511565441","author":"RN Owen-Smith","year":"1988","journal-title":"Megaherbivores: the influence of very large body size on ecology. Cambridge: Cambridge University Press"},{"issue":"13","key":"pone.0345236.ref034","doi-asserted-by":"crossref","first-page":"5223","DOI":"10.1073\/pnas.87.13.5223","article-title":"The earliest fossil evidence for sexual dimorphism in primates","volume":"87","author":"L Krishtalka","year":"1990","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pone.0345236.ref035","first-page":"185","article-title":"Intraspecific variation and sexual dimorphism in cranial and dental variables among higher primates and their bearing on the hominid fossil record","volume":"174","author":"BA Wood","year":"1991","journal-title":"J Anat"},{"issue":"3","key":"pone.0345236.ref036","doi-asserted-by":"crossref","first-page":"327","DOI":"10.1006\/jhev.2000.0423","article-title":"Inferring social behavior from sexual dimorphism in the fossil record","volume":"39","author":"JM Plavcan","year":"2000","journal-title":"J Hum Evol"},{"issue":"4","key":"pone.0345236.ref037","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1017\/S0094837300025501","article-title":"Variation, sexual dimorphism, and social structure in the early Eocene horse Hyracotherium (Mammalia, Perissodactyla)","volume":"7","author":"PD Gingerich","year":"1981","journal-title":"Paleobiology"},{"key":"pone.0345236.ref038","doi-asserted-by":"crossref","first-page":"741","DOI":"10.1111\/1475-4983.00095","article-title":"Sexual dimorphism in a new species of the actinopterygian Peltopleurus from the Triassic of northern Italy","volume":"42","author":"C Lombardo","year":"1999","journal-title":"Palaeontology"},{"issue":"5","key":"pone.0345236.ref039","doi-asserted-by":"crossref","first-page":"2559","DOI":"10.1073\/pnas.96.5.2559","article-title":"Canine sexual dimorphism in Egyptian Eocene anthropoid primates: Catopithecus and Proteopithecus","volume":"96","author":"EL Simons","year":"1999","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"2","key":"pone.0345236.ref040","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0004366","article-title":"New protocetid whale from the middle\u00a0Eocene\u00a0of Pakistan: birth on land, precocial development, and sexual dimorphism","volume":"4","author":"PD Gingerich","year":"2009","journal-title":"PLoS One"},{"key":"pone.0345236.ref041","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1080\/08912960903450505","article-title":"The socio-sexual behaviour of extant archosaurs: implications for understanding dinosaur behaviour","volume":"21","author":"TE Isles","year":"2009","journal-title":"Hist Biol"},{"issue":"4","key":"pone.0345236.ref042","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1016\/j.geobios.2010.10.007","article-title":"Ontogenetic changes and sexual dimorphism in the mandible of adult woolly mammoths (Mammuthus primigenius","volume":"44","author":"DJ \u00c1lvarez-Lao","year":"2011","journal-title":"). Geobios"},{"key":"pone.0345236.ref043","doi-asserted-by":"crossref","first-page":"1381","DOI":"10.1038\/ncomms2377","article-title":"Gender identification of the Mesozoic bird Confuciusornis sanctus","volume":"4","author":"A Chinsamy","year":"2013","journal-title":"Nat Commun"},{"issue":"5","key":"pone.0345236.ref044","doi-asserted-by":"crossref","DOI":"10.1080\/02724634.2016.1180298","article-title":"Sexual dimorphism in the late Miocene mihirung Dromornis stirtoni (Aves: Dromornithidae) from the Alcoota Local Fauna of central Australia","volume":"36","author":"WD Handley","year":"2016","journal-title":"J Vertebr Paleontol"},{"key":"pone.0345236.ref045","doi-asserted-by":"crossref","first-page":"24087","DOI":"10.1038\/srep24087","article-title":"Reacquisition of the lower temporal bar in sexually dimorphic fossil lizards provides a rare case of convergent evolution","volume":"6","author":"TR Sim\u00f5es","year":"2016","journal-title":"Sci Rep"},{"key":"pone.0345236.ref046","doi-asserted-by":"crossref","first-page":"14978","DOI":"10.1038\/s41598-018-33302-4","article-title":"Separating sexual dimorphism from other morphological variation in a specimen complex of fossil marine reptiles (Reptilia, Ichthyosauriformes, Chaohusaurus","volume":"8","author":"R Motani","year":"2018","journal-title":"). Sci Rep"},{"key":"pone.0345236.ref047","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.89158","article-title":"Sexual dimorphism in dinosaurs","volume":"12","author":"SA Ludwig","year":"2023","journal-title":"Elife"},{"key":"pone.0345236.ref048","unstructured":"Rinehart LF, Lucas SG. Lateral skull angle: a new sexual dimorphism signal in temnospondyl amphibians. New Mexico Museum of Natural History and Science Bulletin. 2023;94:559\u201383."},{"key":"pone.0345236.ref049","first-page":"95","article-title":"Growth changes in Asian dinosaurs and some problems of their taxonomy","volume":"3","author":"AK Rozhdestvensky","year":"1965","journal-title":"Paleontol \u017d"},{"key":"pone.0345236.ref050","first-page":"33","article-title":"Methods to estimate sexual dimorphism from unsexed samples: a test with computer-generated samples","volume":"67","author":"K Koscinski","year":"2004","journal-title":"Anthropol Rev"},{"issue":"4","key":"pone.0345236.ref051","doi-asserted-by":"crossref","first-page":"535","DOI":"10.1111\/pala.12298","article-title":"Protracted growth impedes the detection of sexual dimorphism in non\u2010avian dinosaurs","volume":"60","author":"DWE Hone","year":"2017","journal-title":"Palaeontology"},{"issue":"3","key":"pone.0345236.ref052","doi-asserted-by":"crossref","first-page":"495","DOI":"10.1017\/pab.2016.51","article-title":"Recognizing sexual dimorphism in the fossil record: lessons from nonavian dinosaurs","volume":"43","author":"JC Mallon","year":"2017","journal-title":"Paleobiology"},{"issue":"4","key":"pone.0345236.ref053","doi-asserted-by":"crossref","first-page":"820","DOI":"10.1111\/j.1558-5646.1988.tb02501.x","article-title":"Sexual dimorphism in body size: a model and a test","volume":"42","author":"A Arak","year":"1988","journal-title":"Evol"},{"key":"pone.0345236.ref054","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1002\/ajpa.1330790113","article-title":"Comparison of t-tests for differences in sexual dimorphism between populations","volume":"79","author":"DL Greene","year":"1989","journal-title":"Am J Phys Anthropol"},{"key":"pone.0345236.ref055","first-page":"225","article-title":"Methodology and reliability of sex determination from the skeleton.","author":"J Bruzek","year":"2006","journal-title":"Forensic anthropology and medicine: complementary sciences from recovery to cause of death. Totowa, NJ: Humana Press"},{"key":"pone.0345236.ref056","doi-asserted-by":"crossref","first-page":"753","DOI":"10.1016\/j.quascirev.2009.11.022","article-title":"Highly skewed sex ratios and biased fossil deposition of moa: ancient DNA provides new insight on New Zealand\u2019s extinct megafauna","volume":"29","author":"ME Allentoft","year":"2010","journal-title":"Quat Sci Rev"},{"issue":"5","key":"pone.0345236.ref057","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1016\/j.tree.2013.01.011","article-title":"Misconceptions of sexual selection and species recognition: a response to Knell et al. and to Mendelson and Shaw","volume":"28","author":"K Padian","year":"2013","journal-title":"Trends Ecol Evol"},{"issue":"2","key":"pone.0345236.ref058","doi-asserted-by":"crossref","first-page":"20150947","DOI":"10.1098\/rsbl.2015.0947","article-title":"Ontogeny and the fossil record: what, if anything, is an adult dinosaur?","volume":"12","author":"DWE Hone","year":"2016","journal-title":"Biol Lett"},{"key":"pone.0345236.ref059","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1201\/9780429343001-14","article-title":"Phenotypic plasticity in the fossil record.","volume-title":"Phenotypic plasticity & evolution: causes, consequences, controversies","author":"AM Lister","year":"2021"},{"issue":"12","key":"pone.0345236.ref060","doi-asserted-by":"crossref","first-page":"1165","DOI":"10.1016\/j.tree.2023.08.005","article-title":"Measurement theory and paleobiology","volume":"38","author":"KL Voje","year":"2023","journal-title":"Trends Ecol Evol"},{"key":"pone.0345236.ref061","first-page":"929","article-title":"Quantitative aspects of relative growth and sexual dimorphism in Protoceratops","volume":"50","author":"P Dodson","journal-title":"J Paleontol"},{"key":"pone.0345236.ref062","doi-asserted-by":"crossref","first-page":"641","DOI":"10.1080\/02724634.2011.557112","article-title":"Evidence for sexual dimorphism in the stegosaurian dinosaur Kentrosaurus aethiopicus from the Upper Jurassic of Tanzania","volume":"31","author":"HE Barden","year":"2011","journal-title":"J Vertebr Paleontol"},{"issue":"5780","key":"pone.0345236.ref063","doi-asserted-by":"crossref","first-page":"328","DOI":"10.1038\/287328a0","article-title":"Sexual dimorphism in early anthropoids","volume":"287","author":"JG Fleagle","year":"1980","journal-title":"Nature"},{"issue":"2","key":"pone.0345236.ref064","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1073\/pnas.79.2.209","article-title":"Sexual dimorphism in Ramapithecinae","volume":"79","author":"RF Kay","year":"1982","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pone.0345236.ref065","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1007\/BF02693493","article-title":"Sivapithecus simonsi, a new species of miocene hominoid, with comments on the phylogenetic status of the ramapithecinae","volume":"3","author":"RF Kay","year":"1982","journal-title":"Int J Primatol"},{"key":"pone.0345236.ref066","first-page":"404","article-title":"The percentage limits for the distribution of range in samples from a normal population. (n \u2264 100.","volume":"24","author":"ES Pearson","year":"1932","journal-title":"). Biometrika"},{"issue":"3","key":"pone.0345236.ref067","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1002\/ajpa.1330900306","article-title":"Sexual dimorphism in large-bodied primates: the case of the subfossil lemurs","volume":"90","author":"LR Godfrey","year":"1993","journal-title":"Am J Phys Anthropol"},{"issue":"2","key":"pone.0345236.ref068","doi-asserted-by":"crossref","first-page":"191","DOI":"10.1002\/(SICI)1096-8644(199606)100:2<191::AID-AJPA3>3.0.CO;2-0","article-title":"Estimating sexual dimorphism by method-of-moments","volume":"100","author":"SC Josephson","year":"1996","journal-title":"Am J Phys Anthropol"},{"issue":"2","key":"pone.0345236.ref069","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1006\/jasc.1996.0100","article-title":"Mixture analysis and its preliminary application in archaeology","volume":"24","author":"Z Dong","year":"1997","journal-title":"J Archaeol Sci"},{"issue":"4","key":"pone.0345236.ref070","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0123503","article-title":"Evidence for sexual dimorphism in the plated dinosaur Stegosaurus mjosi (Ornithischia, Stegosauria) from the Morrison Formation (Upper Jurassic) of western USA","volume":"10","author":"ET Saitta","year":"2015","journal-title":"PLoS One"},{"issue":"4","key":"pone.0345236.ref071","doi-asserted-by":"crossref","first-page":"465","DOI":"10.1002\/ajpa.1330940403","article-title":"Comparison of four simple methods for estimating sexual dimorphism in fossils","volume":"94","author":"JM Plavcan","year":"1994","journal-title":"Am J Phys Anthropol"},{"issue":"2","key":"pone.0345236.ref072","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1093\/biolinnean\/blaa105","article-title":"An effect size statistical framework for investigating sexual dimorphism in non-avian dinosaurs and other extinct taxa","volume":"131","author":"ET Saitta","year":"2020","journal-title":"Biol J Linn Soc Lond"},{"key":"pone.0345236.ref073","author":"OC Marsh","year":"1886"},{"issue":"3","key":"pone.0345236.ref074","first-page":"506","article-title":"Prouintatherium, new uintathere genus, earliest Eocene, Hoback Formation, Wyoming, and the phylogeny of Dinocerata","volume":"32","author":"JA Dorr","year":"1958","journal-title":"J Paleontol"},{"key":"pone.0345236.ref075","first-page":"1","article-title":"Revision of the uintatheres","volume":"14","author":"W Wheeler","year":"1961","journal-title":"Bull Peabody Mus Nat Hist"},{"key":"pone.0345236.ref076","unstructured":"Schoch RM, Lucas SG. The phylogeny and classification of the Dinocerata (Mammalia, Eutheria). Bull Geol Inst Univ Uppsala. 1985;11:31\u201358."},{"key":"pone.0345236.ref077","first-page":"284","article-title":"Dinocerata.","volume-title":"Evolution of tertiary mammals of North America","author":"SG Lucas","year":"1998"},{"key":"pone.0345236.ref078","article-title":"The mammalian faunas of the Washakie Formation, Eocene age, of southern Wyoming","author":"WD Turnbull","year":"2002","journal-title":"Chicago: Field Museum of Natural History"},{"issue":"3","key":"pone.0345236.ref079","first-page":"208","article-title":"A skull of Uintatherium from Henan","volume":"19","author":"Y Tong","year":"1981","journal-title":"Ku Chi Chui Tung Wu Yu Ku Jen Lei"},{"issue":"3","key":"pone.0345236.ref080","doi-asserted-by":"crossref","DOI":"10.2992\/007.090.0305","article-title":"Nothernmost record of Dinocerata (Mammalia: Eutheria) in North America from the Middle Eocene Kishenehn formation of\u00a0","volume":"90","author":"KD Mulcahy","year":"2025","journal-title":"Annals of Carnegie Museum"},{"issue":"81","key":"pone.0345236.ref081","first-page":"580","article-title":"Notice of proboscians from the\u00a0Eocene\u00a0of Southern Wyoming","volume":"12","author":"ED Cope","year":"1871","journal-title":"Proc Am Philos Soc"},{"issue":"3408","key":"pone.0345236.ref082","doi-asserted-by":"crossref","first-page":"1171","DOI":"10.1126\/science.131.3408.1171","article-title":"The Uintatheres and the Cope-Marsh War","volume":"131","author":"WH Wheeler","year":"1960","journal-title":"Science"},{"key":"pone.0345236.ref083","unstructured":"Burger BJ. The systematic position of the saber-toothed and horned giants of the Eocene: the Uintatheres (Order Dinocerata). Annual Meeting of the Society of Vertebrate Paleontology, Dallas, TX. 2015."},{"issue":"29","key":"pone.0345236.ref084","doi-asserted-by":"crossref","first-page":"303","DOI":"10.2475\/ajs.s3-31.184.303","article-title":"On some new forms of the Dinocerata","volume":"3","author":"WB Scott","year":"1886","journal-title":"Am J Sci"},{"key":"pone.0345236.ref085","first-page":"1","article-title":"A new Paleocene uintathere and molar evolution in the Amblypoda","volume":"387","author":"GG Simpson","year":"1929","journal-title":"Am Mus Novit"},{"key":"pone.0345236.ref086","author":"GW Flora","year":"2021"},{"key":"pone.0345236.ref087","first-page":"1","article-title":"Bison bison","volume":"266","author":"M Meagher","year":"1986","journal-title":"Mamm Species"},{"issue":"3","key":"pone.0345236.ref088","doi-asserted-by":"crossref","first-page":"342","DOI":"10.2307\/3796894","article-title":"The horns and teeth as indicators of age in bison.","volume":"23","author":"WA Fuller","year":"1959","journal-title":"J Wildl Manage"},{"issue":"4","key":"pone.0345236.ref089","doi-asserted-by":"crossref","first-page":"335","DOI":"10.5735\/086.045.0415","article-title":"Sexual dimorphism in skulls of the lowland European bison, Bison bonasus bonasus","volume":"45","author":"F Kobry\u0144czuk","year":"2008","journal-title":"Ann Zool Fennici"},{"key":"pone.0345236.ref090","doi-asserted-by":"crossref","first-page":"1418","DOI":"10.1038\/s41598-025-85654-3","article-title":"Morphological patterns of the European bison (Bison bonasus) skull","volume":"15","author":"O G\u00fcndemir","year":"2025","journal-title":"Sci Rep"},{"issue":"4","key":"pone.0345236.ref091","first-page":"323","article-title":"Notice of a new species of Tinoceras","volume":"3","author":"OC Marsh","year":"1872","journal-title":"Am J Sci"},{"key":"pone.0345236.ref092","doi-asserted-by":"crossref","unstructured":"Osborn HF, Scott WB, Speir F Jr. Palaeontological report of the Princeton scientific expedition of 1877. In: Contr. Mus. Geol. Archaeol. Princeton College. 1878;1(8):1\u2013146.","DOI":"10.5962\/bhl.title.15039"},{"key":"pone.0345236.ref093","first-page":"5","article-title":"A memoir upon Loxolophodon and Uintatherium, two genera of the sub-order Dinocerata. Contributions from the E.M. Mus. Geol. Archaeol. College N","volume":"1","author":"HF Osborn","year":"1881","journal-title":"J. Contr"},{"issue":"2","key":"pone.0345236.ref094","first-page":"703","article-title":"The titanotheres of ancient Wyoming, Dakota, and Nebraska. U. S. Geol","volume":"55","author":"HF Osborn","year":"1929","journal-title":"Survey Mon"},{"key":"pone.0345236.ref095","doi-asserted-by":"crossref","first-page":"6","DOI":"10.5962\/bhl.title.5883","volume-title":"Some memories of a palaeontologist","author":"WB Scott","year":"1939"},{"key":"pone.0345236.ref096","author":"R Core Team","year":"2025"},{"key":"pone.0345236.ref097","author":"H Wickham","year":"2016"},{"key":"pone.0345236.ref098","author":"M Maechler","year":"2025"},{"key":"pone.0345236.ref099","author":"L Scrucca","year":"2023"},{"issue":"2","key":"pone.0345236.ref100","doi-asserted-by":"crossref","DOI":"10.1214\/aos\/1176344136","article-title":"Estimating the dimension of a model","volume":"6","author":"G Schwarz","year":"1978","journal-title":"Ann Statist"},{"issue":"1","key":"pone.0345236.ref101","doi-asserted-by":"crossref","first-page":"1","DOI":"10.18637\/jss.v070.i01","article-title":"missMDA: a package for handling missing values in multivariate data analysis","volume":"70","author":"J Josse","year":"2016","journal-title":"J Stat Softw"},{"key":"pone.0345236.ref102","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1002\/wics.101","article-title":"Principal component analysis","volume":"2","author":"H Abdi","year":"2010","journal-title":"Wiley Interdiscip Rev Comput Stat"},{"issue":"1","key":"pone.0345236.ref103","first-page":"1","article-title":"FactoMineR: an R package for multivariate analysis","volume":"25","author":"S Le","year":"2008","journal-title":"J Stat Softw"},{"key":"pone.0345236.ref104","article-title":"vegan:","author":"J Oksanen","year":"2026","journal-title":"Community Ecology Package"},{"key":"pone.0345236.ref105","doi-asserted-by":"crossref","first-page":"80716","DOI":"10.1109\/ACCESS.2020.2988796","article-title":"Unsupervised k-means clustering algorithm","volume":"8","author":"KP Sinaga","year":"2020","journal-title":"IEEE Access"},{"key":"pone.0345236.ref106","author":"A Kassambara","year":"2020"},{"issue":"18","key":"pone.0345236.ref107","doi-asserted-by":"crossref","first-page":"2847","DOI":"10.1093\/bioinformatics\/btw313","article-title":"Complex heatmaps reveal patterns and correlations in multidimensional genomic data","volume":"32","author":"Z Gu","year":"2016","journal-title":"Bioinformatics"},{"issue":"3","key":"pone.0345236.ref108","article-title":"Complex heatmap visualization","volume":"1","author":"Z Gu","year":"2022","journal-title":"Imeta"},{"key":"pone.0345236.ref109","author":"R Suzuki","year":"2019"},{"issue":"3","key":"pone.0345236.ref110","doi-asserted-by":"crossref","first-page":"747","DOI":"10.1139\/z87-113","article-title":"Sexual dimorphism in the cranial development of Scandinavian moose (Alces alces (L.) alces","volume":"65","author":"L Bartosiewicz","year":"1987","journal-title":"). 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Widlansky","year":"2022","journal-title":"Clim Past"},{"issue":"1","key":"pone.0345236.ref135","doi-asserted-by":"crossref","first-page":"37","DOI":"10.37520\/fi.2023.004","article-title":"Diverse fruits and seeds of the mid-Eocene Kishenehn Formation, northwestern Montana, USA, and their implications for biogeography","volume":"79","author":"MA Smith","year":"2023","journal-title":"FI"},{"issue":"1","key":"pone.0345236.ref136","doi-asserted-by":"crossref","first-page":"1","DOI":"10.2307\/1374488","article-title":"The Asiatic two-horned rhinoceros","volume":"20","author":"T Hubback","year":"1939","journal-title":"J Mammal"},{"issue":"4","key":"pone.0345236.ref137","doi-asserted-by":"crossref","first-page":"114","DOI":"10.1038\/scientificamerican0469-114","article-title":"Horns and antlers","volume":"220","author":"W Modell","year":"1969","journal-title":"Sci Am"},{"key":"pone.0345236.ref138","volume-title":"Horns, pronghorns, and antlers: evolution, morphology, physiology, and social 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Roberts","year":"1996","journal-title":"Behav"},{"issue":"1","key":"pone.0345236.ref142","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1177\/1059712307087283","article-title":"Mutual mate choice can drive costly signaling even under perfect monogamy","volume":"16","author":"PL Hooper","year":"2008","journal-title":"Adapt Behav"},{"issue":"1677","key":"pone.0345236.ref143","first-page":"4329","article-title":"Evolution of weaponry in female bovids","volume":"276","author":"T Stankowich","year":"2009","journal-title":"Proc Biol Sci"},{"issue":"3","key":"pone.0345236.ref144","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1111\/jzo.12023","article-title":"Social competition and its consequences in female mammals","volume":"289","author":"T Clutton-Brock","year":"2013","journal-title":"J Zool"},{"issue":"1","key":"pone.0345236.ref145","doi-asserted-by":"crossref","first-page":"19492","DOI":"10.1038\/s41598-020-76615-z","article-title":"Mutual mate choice and its 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