{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,18]],"date-time":"2026-02-18T13:22:09Z","timestamp":1771420929736,"version":"3.50.1"},"reference-count":67,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,1,26]],"date-time":"2026-01-26T00:00:00Z","timestamp":1769385600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,2,18]],"date-time":"2026-02-18T00:00:00Z","timestamp":1771372800000},"content-version":"vor","delay-in-days":23,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"name":"The APC\/BPC is financed by Wroclaw University of Environmental and Life Sciences (Poland)."}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["BMC Vet Res"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:sec>\n                    <jats:title>Background<\/jats:title>\n                    <jats:p>The aardwolf skulls were analyzed and compared to the spotted hyena, the brown hyena and the striped hyena, using specimens from Polish and South African collections. Addressing gaps in detailed anatomy and morphometrics, this study used extensive analyzes, encompassing 64 morphometric parameters and 7 indices, to examine the morphology and morphometrics of cranial and mandibular structures and quantify interspecific and intraspecific variation.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>The comparative analysis of hyena skulls revealed significant differences between the aardwolf and the other three species. The aardwolf consistently had the smallest and proportionally narrowest skull, characterized by high morphological stability and a wide neurocranium. The brown hyena and spotted hyena had the largest skulls; however, the brown hyena showed greater homogeneity, while the spotted hyena displayed higher absolute variability. Morphologically, the aardwolf\u2019s basilar part was cylindrical, unlike the pyramidal shape in the spotted hyena, and it possessed prominent nuchal tubercles that were reduced in the brown and spotted hyenas.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusions<\/jats:title>\n                    <jats:p>This study demonstrates that the morphological and morphometric features of the aardwolf\u2019s skull are highly specialized and fundamentally distinct from the other three hyena species, reflecting its unique dietary niche. The powerful skulls of the spotted and brown hyenas, in contrast, correlate with their roles as apex predators and scavengers.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.1186\/s12917-026-05290-5","type":"journal-article","created":{"date-parts":[[2026,1,26]],"date-time":"2026-01-26T06:03:13Z","timestamp":1769407393000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Insights into cranial anatomy and craniometry of the aardwolf (Proteles cristata) with comparisons to extant hyaenids"],"prefix":"10.1186","volume":"22","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-8048-8031","authenticated-orcid":false,"given":"Krzysztof","family":"Stegmann","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6651-9660","authenticated-orcid":false,"given":"Shaw","family":"Badenhorst","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0294-3777","authenticated-orcid":false,"given":"Wojciech","family":"Borawski","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4296-3174","authenticated-orcid":false,"given":"Dominik","family":"Poradowski","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4357-2271","authenticated-orcid":false,"given":"Maciej","family":"Janeczek","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7462-3524","authenticated-orcid":false,"given":"Anna","family":"Mucha","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1154-8736","authenticated-orcid":false,"given":"Joanna","family":"Kle\u0107kowska-Nawrot","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2026,1,26]]},"reference":[{"key":"5290_CR1","doi-asserted-by":"publisher","DOI":"10.56021\/9780801857898","volume-title":"Walker\u2019s mammals of the world","author":"RM Nowak","year":"1999","unstructured":"Nowak RM. Walker\u2019s mammals of the world. Volume I, 6th ed. Baltimore and London: Johns Hopkins University; 1999.","edition":"6"},{"key":"5290_CR2","doi-asserted-by":"crossref","unstructured":"Green DS. Proteles cristata. The IUCN Red List of Threatened Species 2015: e.T18372A45195681. 2015. https:\/\/dx.doi.org\/10.2305\/IUCN.UK.2015-2.RLTS.T18372A45195681.en. Accessed on 07 November 2024.","DOI":"10.2305\/IUCN.UK.2015-2.RLTS.T18372A45195681.en"},{"key":"5290_CR3","unstructured":"Anderson M, Mills G. Proteles cristata (Mediterranean assessment). The IUCN Red List of Threatened Species 2010: e.T18372A8164230. 2010. Accessed on 07 November 2024."},{"key":"5290_CR4","doi-asserted-by":"crossref","unstructured":"Bohm T, H\u00f6ner OR. Crocuta crocuta. The IUCN Red List of Threatened Species 2015: e.T5674A45194782. 2015. https:\/\/dx.doi.org\/10.2305\/IUCN.UK.2015-2.RLTS.T5674A45194782.en. Accessed on 07 November 2024.","DOI":"10.2305\/IUCN.UK.2015-2.RLTS.T5674A45194782.en"},{"key":"5290_CR5","doi-asserted-by":"crossref","unstructured":"Wiesel I. Parahyaena brunnea. The IUCN Red List of Threatened Species 2015: e.T10276A82344448. 2015. https:\/\/dx.doi.org\/10.2305\/IUCN.UK.2015-4.RLTS.T10276A82344448.en. Accessed on 07 November 2024.","DOI":"10.2305\/IUCN.UK.2015-4.RLTS.T10276A82344448.en"},{"key":"5290_CR6","doi-asserted-by":"crossref","unstructured":"AbiSaid M, Dloniak SMD. Hyaena hyaena. The IUCN Red List of Threatened Species 2015: e.T10274A45195080. 2015. https:\/\/dx.doi.org\/10.2305\/IUCN.UK.2015-2.RLTS.T10274A45195080.en. Accessed on 07 November 2024.","DOI":"10.2305\/IUCN.UK.2015-2.RLTS.T10274A45195080.en"},{"key":"5290_CR7","unstructured":"Jdeidi T, Masseti M, Nader I, de Smet K, Cuzin F. Hyaena hyaena (Mediterranean assessment). The IUCN red list of threatened species 2010: e.T10274A3188449. 2010.\u00a0https:\/\/www.iucnredlist.org\/species\/10274\/3188449.\u00a0Accessed 07 Nov 2024."},{"key":"5290_CR8","volume-title":"Smithers\u2019 mammals of Southern Africa. A field guide","year":"2000","unstructured":"Apps P, editor. Smithers\u2019 mammals of Southern Africa. A field guide. Cape Town, South Africa: Struik; 2000."},{"key":"5290_CR9","volume-title":"Larger mammals of Africa","author":"J Dorst","year":"1972","unstructured":"Dorst J, Dandelot P. Larger mammals of Africa. 2nd ed. London, UK: HarperCollins; 1972.","edition":"2"},{"key":"5290_CR10","volume-title":"The safari companion: A guide to watching African mammals including hoofed mammals, Carnivores, and primates","author":"RD Estes","year":"1999","unstructured":"Estes RD. The safari companion: A guide to watching African mammals including hoofed mammals, Carnivores, and primates. Vermont, USA: Chelsea Green Publishing Company; 1999."},{"key":"5290_CR11","volume-title":"Field guide to the mammals of the Kruger National park","author":"H Sch\u00fctze","year":"2002","unstructured":"Sch\u00fctze H. Field guide to the mammals of the Kruger National park. Cape Town, South Africa: Struik; 2002."},{"key":"5290_CR12","volume-title":"Field guide to the larger mammals of Africa","author":"C Stuart","year":"2006","unstructured":"Stuart C, Stuart T. Field guide to the larger mammals of Africa. 3rd ed. Cape Town, South Africa: Struik Nature; 2006.","edition":"3"},{"issue":"2","key":"5290_CR13","doi-asserted-by":"publisher","first-page":"78","DOI":"10.1111\/j.1096-3642.1997.tb01276.x","volume":"3","author":"E Atay","year":"2020","unstructured":"Atay E, Tatl\u0131 M, \u00c7etin \u0130. A study on the skull of the striped hyaena (Hyaena hyaena: Hyaenidae, Carnivora). Eurasian J Biol Chem Sci. 2020;3(2):78\u201382. https:\/\/doi.org\/10.1111\/j.1096-3642.1997.tb01276.x.","journal-title":"Eurasian J Biol Chem Sci"},{"issue":"2","key":"5290_CR14","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1111\/j.1096-3642.1997.tb01276.x","volume":"120","author":"AR Biknevicius","year":"1997","unstructured":"Biknevicius AR, Leigh SR. Patterns of growth of the mandibular corpus in spotted hyenas (Crocuta crocuta) and Cougars (Puma concolor). Zool J Linn Soc. 1997;120(2):139\u201361. https:\/\/doi.org\/10.1111\/j.1096-3642.1997.tb01276.x.","journal-title":"Zool J Linn Soc"},{"issue":"3","key":"5290_CR15","doi-asserted-by":"publisher","first-page":"273","DOI":"10.1111\/j.1469-7998.2000.tb00622.x","volume":"252","author":"WJ Binder","year":"2020","unstructured":"Binder WJ, Van Valkenburgh B. Development of bite strength and feeding behaviour in juvenile spotted hyenas (Crocuta crocuta). J Zool. 2020;252(3):273\u201383. https:\/\/doi.org\/10.1111\/j.1469-7998.2000.tb00622.x.","journal-title":"J Zool"},{"issue":"2","key":"5290_CR16","doi-asserted-by":"publisher","first-page":"133","DOI":"10.1017\/S0952836905006643","volume":"266","author":"P Christiansen","year":"2005","unstructured":"Christiansen P, Adolfssen JS. Bite forces, canine strength and skull allometry in carnivores (Mammalia, Carnivora). J Zool. 2005;266(2):133\u201351. https:\/\/doi.org\/10.1017\/S0952836905006643.","journal-title":"J Zool"},{"issue":"3","key":"5290_CR17","doi-asserted-by":"publisher","first-page":"490","DOI":"10.1666\/09062.1","volume":"37","author":"B Figueirido","year":"2011","unstructured":"Figueirido B, MacLeod N, Krieger J, De Renzi M, P\u00e9rez-Claros JA, Palmqvist P. Constraint and adaptation in the evolution of carnivoran skull shape. Paleobiology. 2011;37(3):490\u2013518. https:\/\/doi.org\/10.1666\/09062.1.","journal-title":"Paleobiology"},{"issue":"7","key":"5290_CR18","doi-asserted-by":"publisher","first-page":"1975","DOI":"10.1111\/evo.12059","volume":"67","author":"B Figueirido","year":"2013","unstructured":"Figueirido B, Tseng ZJ, Mart\u00edn-Serra A. Skull shape evolution in durophagous carnivorans. Evolution. 2013;67(7):1975\u201393. https:\/\/doi.org\/10.1111\/evo.12059.","journal-title":"Evolution"},{"issue":"1","key":"5290_CR19","doi-asserted-by":"publisher","first-page":"4","DOI":"10.1111\/j.1469-7998.1869.tb07286.x","volume":"37","author":"WH Flower","year":"1869","unstructured":"Flower WH. On the value of the characters of the base of the cranium in the classification of the order Carnivora, and on the systematic position of Bassaris and other disputed forms. Proc Zoological Soc Lond. 1869;37(1):4\u201337. https:\/\/doi.org\/10.1111\/j.1469-7998.1869.tb07286.x.","journal-title":"Proc Zoological Soc Lond"},{"issue":"1","key":"5290_CR20","doi-asserted-by":"publisher","first-page":"169","DOI":"10.1111\/j.0014-3820.2006.tb01091.x","volume":"60","author":"A Goswami","year":"2006","unstructured":"Goswami A. Morphological integration in the carnivoran skull. Evolution. 2006;60(1):169\u201383. https:\/\/doi.org\/10.1111\/j.0014-3820.2006.tb01091.x.","journal-title":"Evolution"},{"issue":"1","key":"5290_CR21","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1002\/jmor.1051430103","volume":"143","author":"RM Hunt Jr","year":"1974","unstructured":"Hunt RM Jr. The auditory bulla in carnivora: an anatomical basis for reappraisal of carnivore evolution. J Morphol. 1974;143(1):21\u201375. https:\/\/doi.org\/10.1002\/jmor.1051430103.","journal-title":"J Morphol"},{"issue":"3","key":"5290_CR22","doi-asserted-by":"publisher","first-page":"627","DOI":"10.1080\/02724634.1998.10011089","volume":"18","author":"RM Joeckel","year":"1998","unstructured":"Joeckel RM. Unique frontal sinuses in fossil and living hyaenidae (Mammalia, Carnivora): description and interpretation. J Vertebr Paleontol. 1998;18(3):627\u201339. https:\/\/doi.org\/10.1080\/02724634.1998.10011089.","journal-title":"J Vertebr Paleontol"},{"key":"5290_CR23","doi-asserted-by":"publisher","first-page":"e10541","DOI":"10.7717\/peerj.10541","volume":"8","author":"JA P\u00e9rez-Claros","year":"2020","unstructured":"P\u00e9rez-Claros JA, Coca-Ortega C. Canines and carnassials as indicators of sociality in durophagous hyaenids: analyzing the past to understand the present. PeerJ. 2020;8:e10541. https:\/\/doi.org\/10.7717\/peerj.10541.","journal-title":"PeerJ"},{"key":"5290_CR24","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1111\/j.1096-3642.1916.tb02018.x","volume":"86","author":"RI Pocock","year":"1916","unstructured":"Pocock P RI. The tympanic bulla in hyenas. Proc Zoological Soc Lond. 1916;86:303\u20137.","journal-title":"Proc Zoological Soc Lond"},{"key":"5290_CR25","doi-asserted-by":"crossref","first-page":"442","DOI":"10.1111\/j.1096-3642.1916.tb02018.x","volume":"86","author":"RI Pocock","year":"1916","unstructured":"Pocock P RI. The alisphenoid Canal in civets and hyaenas. Proc Zoological Soc Lond. 1916;86:442\u20135.","journal-title":"Proc Zoological Soc Lond"},{"issue":"2","key":"5290_CR26","doi-asserted-by":"publisher","first-page":"363","DOI":"10.1007\/s10914-023-09653-9","volume":"30","author":"J Rajmil","year":"2023","unstructured":"Rajmil J, Velazco PM, Giannini NP. Growing apart: comparative cranial ontogeny in the myrmecophagous Aardwolf (Proteles cristata) and the bone-cracking spotted hyaena (Crocuta crocuta). J Mammalian Evol. 2023;30(2):363\u201370. https:\/\/doi.org\/10.1007\/s10914-023-09653-9.","journal-title":"J Mammalian Evol"},{"key":"5290_CR27","first-page":"343","volume":"13","author":"JM Rensberger","year":"1999","unstructured":"Rensberger JM. Enamel microstructural specialization in the canine of the spotted hyena, Crocuta Crocuta. Scanning Microsc. 1999;13:343\u201361.","journal-title":"Scanning Microsc"},{"issue":"2","key":"5290_CR28","doi-asserted-by":"publisher","first-page":"246","DOI":"10.1111\/j.1095-8312.2008.01052.x","volume":"95","author":"JB Tanner","year":"2008","unstructured":"Tanner JB, Dumont ER, Sakai ST, Lundrigan BL, Holekamp KE. Of arcs and vaults: the biomechanics of bone-cracking in spotted hyenas (Crocuta crocuta). Biol J Linn Soc. 2008;95(2):246\u201355. https:\/\/doi.org\/10.1111\/j.1095-8312.2008.01052.x.","journal-title":"Biol J Linn Soc"},{"issue":"3","key":"5290_CR29","doi-asserted-by":"publisher","first-page":"353","DOI":"10.1002\/jmor.10802","volume":"271","author":"JB Tanner","year":"2010","unstructured":"Tanner JB, Zelditch ML, Lundrigan BL, Holekamp KE. Ontogenetic change in skull morphology and mechanical advantage in the spotted hyena (Crocuta crocuta). J Morphol. 2010;271(3):353\u201365. https:\/\/doi.org\/10.1002\/jmor.10802.","journal-title":"J Morphol"},{"issue":"3","key":"5290_CR30","doi-asserted-by":"publisher","first-page":"683","DOI":"10.1111\/j.1096-3642.2009.00555.x","volume":"158","author":"ZJ Tseng","year":"2010","unstructured":"Tseng ZJ, Binder WJ. Mandibular biomechanics of Crocuta crocuta, canis lupus, and the late miocene dinocrocuta gigantea (Carnivora, Mammalia). Zool J Linn Soc. 2010;158(3):683\u201396. https:\/\/doi.org\/10.1111\/j.1096-3642.2009.00555.x.","journal-title":"Zool J Linn Soc"},{"issue":"3","key":"5290_CR31","doi-asserted-by":"publisher","first-page":"540","DOI":"10.1111\/j.1095-8312.2010.01602.x","volume":"102","author":"ZJ Tseng","year":"2011","unstructured":"Tseng ZJ, Stynder D. Mosaic functionality in a transitional ecomorphology: skull biomechanics in stem hyaeninae compared to modern South African carnivorans. Biol J Linn Soc. 2011;102(3):540\u201359. https:\/\/doi.org\/10.1111\/j.1095-8312.2010.01602.x.","journal-title":"Biol J Linn Soc"},{"issue":"2","key":"5290_CR32","doi-asserted-by":"publisher","first-page":"109","DOI":"10.1071\/ZO9860109","volume":"34","author":"L Werdelin","year":"1986","unstructured":"Werdelin L. Comparison of skull shape in marsupial and placental carnivores. Australian J Zool. 1986;34(2):109\u201317. https:\/\/doi.org\/10.1071\/ZO9860109.","journal-title":"Australian J Zool"},{"key":"5290_CR33","doi-asserted-by":"publisher","first-page":"1","DOI":"10.18261\/8200374815-1991-01","volume":"30","author":"L Werdelin","year":"1991","unstructured":"Werdelin L, Solounias N. The hyaenidae: taxonomy, systematics, and evolution. Fossils Strata. 1991;30:1\u2013104. https:\/\/doi.org\/10.18261\/8200374815-1991-01.","journal-title":"Fossils Strata"},{"issue":"5","key":"5290_CR34","doi-asserted-by":"publisher","first-page":"1251","DOI":"10.1111\/j.1558-5646.2007.00101.x","volume":"61","author":"S Wroe","year":"2007","unstructured":"Wroe S, Milne N. Convergence and remarkably consistent constraint in the evolution of carnivore skull shape. Evolution. 2007;61(5):1251\u201360. https:\/\/doi.org\/10.1111\/j.1558-5646.2007.00101.x.","journal-title":"Evolution"},{"key":"5290_CR35","unstructured":"Gasse H. Nomina Anatomica Veterinaria. International Committee On Veterinary Gross Anatomical Nomenclature. 6th ed. Hannover: Chairman Committee; 2017. p. 73\u2013147."},{"key":"5290_CR36","volume-title":"Lehrbuch der anatomie der haustiere","author":"R Nickel","year":"2004","unstructured":"Nickel R, Schummer A, Seiferle E. Lehrbuch der anatomie der haustiere. Volume 1. Stuttgart: Parey; 2004."},{"key":"5290_CR37","unstructured":"von den Driesch A. A guide to the measurement of animal bones from archaeological sites. Volume 1. Peabody Museum Bulletins; 1976."},{"key":"5290_CR38","first-page":"125","volume":"Abt7","author":"JU Duerst","year":"1926","unstructured":"Duerst JU. Vergleichende untersuchungsmethoden am Skelett Bei S\u00e4ugern. Handbuch Der Biologischen Arbeitsmethoden. 1926;Abt7:125\u2013530.","journal-title":"Handbuch Der Biologischen Arbeitsmethoden"},{"issue":"6","key":"5290_CR39","doi-asserted-by":"publisher","first-page":"563","DOI":"10.1016\/S0940-9602(97)80022-2","volume":"179","author":"V Onar","year":"1997","unstructured":"Onar V, Mutu\u015f R, Kahvecio\u011flu KO. Morphometric analysis of the foramen magnum in German shepherd dogs (Alsatians). Annals Anat. 1997;179(6):563\u20138. https:\/\/doi.org\/10.1016\/S0940-9602(97)80022-2.","journal-title":"Annals Anat"},{"key":"5290_CR40","doi-asserted-by":"publisher","first-page":"109704","DOI":"10.1016\/j.palaeo.2020.109704","volume":"547","author":"MJ Powers","year":"2020","unstructured":"Powers MJ, Sullivan C, Currie PJ. Re-examining ratio based premaxillary and maxillary characters in eudromaeosauria (Dinosauria: Theropoda): divergent trends in snout morphology between Asian and North American taxa. Palaeogeogr Palaeoclimatol Palaeoecol. 2020;547:109704. https:\/\/doi.org\/10.1016\/j.palaeo.2020.109704.","journal-title":"Palaeogeogr Palaeoclimatol Palaeoecol"},{"issue":"1","key":"5290_CR41","first-page":"5","volume":"4","author":"D Chessel","year":"2004","unstructured":"Chessel D, Dufour AB, Thioulouse J. The ade4 package-I-One-table methods. R News. 2004;4(1):5\u201310.","journal-title":"R News"},{"key":"5290_CR42","doi-asserted-by":"publisher","first-page":"1","DOI":"10.18637\/jss.v022.i04","volume":"22","author":"S Dray","year":"2007","unstructured":"Dray S, Dufour AB. The ade4 package: implementing the duality diagram for ecologists. J Stat Softw. 2007;22:1\u201320.","journal-title":"J Stat Softw"},{"key":"5290_CR43","unstructured":"Kassambara A, Mundt F. Factoextra: extract and visualize the results of multivariate data analyses. 2020. https:\/\/CRAN.R-project.org\/package=factoextra"},{"key":"5290_CR44","unstructured":"R Core Team. R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria. 2024. https:\/\/www.R-project.org\/"},{"issue":"2","key":"5290_CR45","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1890\/0012-9658(2007)88[347:bfaeat]2.0.co;2","volume":"88","author":"P Christiansen","year":"2007","unstructured":"Christiansen P, Wroe S. Bite forces and evolutionary adaptations to feeding ecology in carnivores. Ecology. 2007;88(2):347\u201358. https:\/\/doi.org\/10.1890\/0012-9658(2007)88[347:bfaeat]2.0.co;2.","journal-title":"Ecology"},{"key":"5290_CR46","unstructured":"Ewer RF. The carnivores. Ithaca, New York, USA: Cornell University Press; 1973."},{"issue":"9","key":"5290_CR47","doi-asserted-by":"publisher","first-page":"1367","DOI":"10.3390\/ani14091367","volume":"14","author":"K R\u00f3\u017cycka","year":"2024","unstructured":"R\u00f3\u017cycka K, Skibniewska E, Rajkowski \u0141, Skibniewski M. Craniometric characteristics of selected carnivora species kept in captivity in relation to bite force and bending strength of the upper canines. Animals. 2024;14(9):1367. https:\/\/doi.org\/10.3390\/ani14091367.","journal-title":"Animals"},{"key":"5290_CR48","doi-asserted-by":"crossref","unstructured":"Van Valkenburgh B. Carnivore dental adaptations and diet: a study of trophic diversity within guilds. In: Boston MA, editor. Carnivore behavior, ecology, and evolution. 1st ed. New York: Springer; 1989. p. 410\u201336.","DOI":"10.1007\/978-1-4757-4716-4_16"},{"issue":"1563","key":"5290_CR49","doi-asserted-by":"publisher","first-page":"619","DOI":"10.1098\/rspb.2004.2986","volume":"272","author":"S Wroe","year":"2005","unstructured":"Wroe S, McHenry C, Thomason J. Bite club: comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa. Proc Royal Soc B: Biol Sci. 2005;272(1563):619\u201325. https:\/\/doi.org\/10.1098\/rspb.2004.2986.","journal-title":"Proc Royal Soc B: Biol Sci"},{"issue":"4","key":"5290_CR50","doi-asserted-by":"publisher","first-page":"423","DOI":"10.1111\/j.1469-7998.2006.00286.x","volume":"272","author":"P Christiansen","year":"2007","unstructured":"Christiansen P. Evolutionary implications of bite mechanics and feeding ecology in bears. J Zool. 2007;272(4):423\u201343. https:\/\/doi.org\/10.1111\/j.1469-7998.2006.00286.x.","journal-title":"J Zool"},{"issue":"7","key":"5290_CR51","doi-asserted-by":"publisher","first-page":"896","DOI":"10.1002\/jmor.10643","volume":"269","author":"P Christiansen","year":"2008","unstructured":"Christiansen P. Feeding ecology and morphology of the upper canines in bears (Carnivora: Ursidae). J Morphol. 2008;269(7):896\u2013908. https:\/\/doi.org\/10.1002\/jmor.10643.","journal-title":"J Morphol"},{"issue":"2","key":"5290_CR52","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1002\/ar.24748","volume":"305","author":"A Hartstone-Rose","year":"2022","unstructured":"Hartstone-Rose A, Dickinson E, Deutsch AR, Worden N, Hirschkorn A. Masticatory muscle architectural correlates of dietary diversity in Canidae, Ursidae, and across the order carnivora. Anat Rec. 2022;305(2):477\u201397. https:\/\/doi.org\/10.1002\/ar.24748.","journal-title":"Anat Rec"},{"issue":"1","key":"5290_CR53","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1017\/S0952836904004856","volume":"263","author":"T Sacco","year":"2004","unstructured":"Sacco T, Van Valkenburgh B. Ecomorphological indicators of feeding behaviour in the bears (Carnivora: Ursidae). J Zool. 2004;263(1):41\u201354. https:\/\/doi.org\/10.1017\/S0952836904004856.","journal-title":"J Zool"},{"issue":"2","key":"5290_CR54","doi-asserted-by":"publisher","first-page":"326","DOI":"10.1097\/00001665-200603000-00020","volume":"17","author":"N Acer","year":"2006","unstructured":"Acer N, Sahin B, Ekinci N, Erg\u00fcr H, Basaloglu H. Relation between intracranial volume and the surface area of the foramen magnum. J Craniofac Surg. 2006;17(2):326\u201330. https:\/\/doi.org\/10.1097\/00001665-200603000-00020.","journal-title":"J Craniofac Surg"},{"issue":"7","key":"5290_CR55","doi-asserted-by":"publisher","first-page":"e2807","DOI":"10.1371\/journal.pone.0002807","volume":"3","author":"P Christiansen","year":"2008","unstructured":"Christiansen P. Evolution of skull and mandible shape in cats (Carnivora: Felidae). PLoS ONE. 2008;3(7):e2807. https:\/\/doi.org\/10.1371\/journal.pone.0002807.","journal-title":"PLoS ONE"},{"issue":"4","key":"5290_CR56","doi-asserted-by":"publisher","first-page":"784","DOI":"10.1111\/j.1095-8312.2008.01169.x","volume":"96","author":"J Meachen-Samuels","year":"2009","unstructured":"Meachen-Samuels J, Van Valkenburgh B. Craniodental indicators of prey size preference in the felidae. Biol J Linn Soc. 2009;96(4):784\u201399. https:\/\/doi.org\/10.1111\/j.1095-8312.2008.01169.x.","journal-title":"Biol J Linn Soc"},{"issue":"6","key":"5290_CR57","doi-asserted-by":"publisher","first-page":"878","DOI":"10.1093\/sysbio\/syt053","volume":"62","author":"P Piras","year":"2013","unstructured":"Piras P, Maiorino L, Teresi L, Meloro C, Lucci F, Kotsakis T, Raia P. Bite of the cats: relationships between functional integration and mechanical performance as revealed by mandible geometry. Syst Biol. 2013;62(6):878\u2013900. https:\/\/doi.org\/10.1093\/sysbio\/syt053.","journal-title":"Syst Biol"},{"issue":"2","key":"5290_CR58","doi-asserted-by":"publisher","first-page":"414","DOI":"10.1111\/j.1096-3642.2010.00636.x","volume":"161","author":"FL Sicuro","year":"2011","unstructured":"Sicuro FL, Oliveira LFB. Skull morphology and functionality of extant felidae (Mammalia: Carnivora): a phylogenetic and evolutionary perspective. Zool J Linn Soc. 2011;161(2):414\u201362. https:\/\/doi.org\/10.1111\/j.1096-3642.2010.00636.x.","journal-title":"Zool J Linn Soc"},{"key":"5290_CR59","volume-title":"The big cats and their fossil relatives: an illustrated guide to their evolution and natural history","author":"A Turner","year":"1997","unstructured":"Turner A. The big cats and their fossil relatives: an illustrated guide to their evolution and natural history. New York: Columbia University; 1997."},{"issue":"7","key":"5290_CR60","doi-asserted-by":"publisher","first-page":"951","DOI":"10.1002\/ar.23355","volume":"299","author":"F Penrose","year":"2016","unstructured":"Penrose F, Kemp GJ, Jeffery N. Scaling and accommodation of jaw adductor muscles in Canidae. Anat Rec. 2016;299(7):951\u201366. https:\/\/doi.org\/10.1002\/ar.23355.","journal-title":"Anat Rec"},{"key":"5290_CR61","volume-title":"Dogs: their fossil relatives and evolutionary history","author":"X Wang","year":"2010","unstructured":"Wang X. Dogs: their fossil relatives and evolutionary history. New York: Columbia University; 2010."},{"key":"5290_CR62","unstructured":"Okarma H, Schmidt K. Ry\u015b\u2013monografia przyrodniczo-\u0142owiecka. Wyd., Edukacyjno-Naukowe. H2O, Krak\u00f3w; 2013."},{"key":"5290_CR63","doi-asserted-by":"publisher","unstructured":"Eiting TP, Perot JB, Dumont ER. How much does nasal cavity morphology matter? Patterns and rates of olfactory airflow in phyllostomid bats. Proceedings of the Royal Society B: Biological Sciences. 2015;282(1800):20142161. https:\/\/doi.org\/10.1098\/rspb.2014.2161","DOI":"10.1098\/rspb.2014.2161"},{"issue":"8","key":"5290_CR64","doi-asserted-by":"publisher","first-page":"2463","DOI":"10.3390\/ani11082463","volume":"11","author":"A Kokoci\u0144ska-Kusiak","year":"2021","unstructured":"Kokoci\u0144ska-Kusiak A, Woszczy\u0142o M, Zybala M, Maciocha J, Bar\u0142owska K, Dzi\u0119cio\u0142 M. Canine olfaction: physiology, behavior, and possibilities for practical applications. Animals. 2021;11(8):2463. https:\/\/doi.org\/10.3390\/ani11082463.","journal-title":"Animals"},{"issue":"3","key":"5290_CR65","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1093\/molbev\/msaf035","volume":"42","author":"A Mouton","year":"2025","unstructured":"Mouton A, Bird DJ, Li G, Craven BA, Levine JM, Morselli M, et al. Genetic and anatomical determinants of olfaction in dogs and wild Canids. Mol Biol Evol. 2025;42(3):1\u201314. https:\/\/doi.org\/10.1093\/molbev\/msaf035.","journal-title":"Mol Biol Evol"},{"issue":"4","key":"5290_CR66","doi-asserted-by":"publisher","first-page":"331","DOI":"10.1037\/0735-7036.116.4.331","volume":"116","author":"CM Drea","year":"2002","unstructured":"Drea CM, Vignieri SN, Cunningham SB, Glickman SE. Responses to olfactory stimuli in spotted hyenas (Crocuta crocuta): I. Investigation of environmental odors and the function of rolling. J Comp Psychol. 2002;116(4):331\u201341. https:\/\/doi.org\/10.1037\/0735-7036.116.4.331.","journal-title":"J Comp Psychol"},{"issue":"4","key":"5290_CR67","doi-asserted-by":"publisher","first-page":"342","DOI":"10.1037\/0735-7036.116.4.342","volume":"116","author":"CM Drea","year":"2002","unstructured":"Drea CM, Vignieri SN, Kim HS, Weldele ML, Glickman SE. Responses to olfactory stimuli in spotted hyenas (Crocuta crocuta): II. Discrimination of conspecific scent. J Comp Psychol. 2002;116(4):342\u20139. https:\/\/doi.org\/10.1037\/0735-7036.116.4.342.","journal-title":"J Comp Psychol"}],"container-title":["BMC Veterinary Research"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s12917-026-05290-5","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12917-026-05290-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12917-026-05290-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,18]],"date-time":"2026-02-18T12:52:25Z","timestamp":1771419145000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s12917-026-05290-5"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1,26]]},"references-count":67,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["5290"],"URL":"https:\/\/doi.org\/10.1186\/s12917-026-05290-5","relation":{},"ISSN":["1746-6148"],"issn-type":[{"value":"1746-6148","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,1,26]]},"assertion":[{"value":"6 August 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"5 January 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"26 January 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"According to Polish and European law, studies on tissues obtained post-mortem do not require the approval of the Ethics Committee (Journal of Laws of the Republic of Poland, the Act of January 15, 2015, on the protection of animals used for scientific or educational purposes (Directive 2010\/63\/EU of the European Parliament and of the Council of 22 September 2010 on the protection of animals used for scientific purposes). No ethical approval is required to study animal skeletons in South Africa. All specimens analyzed in this study were obtained from existing institutional collections housed at the Department of Biostructure and Animal Physiology, Faculty of Veterinary Medicine, Wroc\u0142aw University of Environmental and Life Sciences (WUELS, Poland), and the Evolutionary Studies Institute, University of the Witwatersrand (ESIUW, South Africa). According to Polish and European legislation, studies using post-mortem animal material do not require ethics committee approval (Act of 15 January 2015 on the protection of animals used for scientific or educational purposes, implementing Directive 2010\/63\/EU of the European Parliament and of the Council). Similarly, in South Africa, no ethical approval is required for research involving existing skeletal specimens. The study was based on 12 adult skulls representing four hyena species: aardwolf (Proteles cristata, n = 3), spotted hyena (Crocuta crocuta, n = 5), striped hyena (Hyaena hyaena, n = 1), and brown hyena (Parahyaena brunnea, n = 3). The accession codes of the specimens examined were:\n                      \n                      Permission to access and study the collections was granted by the curators of both institutions.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"Not applicable.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare no competing interests.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"114"}}