{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T07:51:06Z","timestamp":1791532266300,"version":"4.3.4"},"reference-count":60,"publisher":"Wiley","license":[{"start":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T00:00:00Z","timestamp":1791504000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"},{"start":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T00:00:00Z","timestamp":1791504000000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"content-domain":{"domain":["anatomypubs.onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["The Anatomical Record"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    Cranial bones of squamate reptiles diverge in shape and structure across lineages, while preserving the basic skull plan. Therefore, variation in cranial morphology of squamates is key to understanding phylogenetic relationships, species delimitation, and lifestyles, which ultimately contribute to insights into their evolution and ecology. Nevertheless, although Gymnophthalmidae is the most diverse endemic non\u2010ophidian squamate family in the Neotropics, studies on comparative cranial osteology within this lineage remain scarce. Herein, we thoroughly describe and compare the external skull anatomy of six species of the Andean genus\n                    <jats:italic>Riama<\/jats:italic>\n                    from Colombia and Ecuador (\n                    <jats:italic>R. columbiana<\/jats:italic>\n                    ,\n                    <jats:italic>R. simotera<\/jats:italic>\n                    ,\n                    <jats:italic>R. striata<\/jats:italic>\n                    , and three undescribed taxa), aiming to provide phenotypic evidence for phylogenetic, taxonomic, and functional analyses. We examined 12 cleared\u2010and\u2010stained skulls of adult specimens (one male and one female per species) to preliminarily assess inter\u2010 and intraspecific variation within\n                    <jats:italic>Riama<\/jats:italic>\n                    . Interspecific variation occurs in the degree of contact between nasals (and consequently in the articulation of the nasal process of the premaxilla), and in the absence\/presence of the posterior process of the jugal, among others. An anterior process of the ventral crest of the vomer in female\n                    <jats:italic>R. columbiana<\/jats:italic>\n                    is seemingly unique among gymnophthalmids. The number of tooth positions and teeth morphology vary both among sexes and species. Preliminary evidence implies sexual dimorphism and bilateral variation in some characters. Our findings suggest that variation in skull anatomy within\n                    <jats:italic>Riama<\/jats:italic>\n                    can inform species delimitation, and phylogenetic and functional analyses. Finally, we discuss variation in cranial morphology among species of\n                    <jats:italic>Riama<\/jats:italic>\n                    within a phylogenetic context.\n                  <\/jats:p>","DOI":"10.1002\/ar.70367","type":"journal-article","created":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T07:33:07Z","timestamp":1791531187000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Cranial osteology of Neotropical squamate reptiles of the genus\n                    <i>Riama<\/i>\n                    (Gymnophthalmidae: Cercosaurinae)"],"prefix":"10.1002","author":[{"ORCID":"https:\/\/orcid.org\/0009-0000-7201-0716","authenticated-orcid":false,"given":"Yuly A.","family":"Vega\u2010Rangel","sequence":"first","affiliation":[{"name":"Universidad Nacional de Colombia, Sede de La Paz  La Paz Cesar Colombia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-9049-5845","authenticated-orcid":false,"given":"Sandy","family":"Escobar\u2010Maestre","sequence":"additional","affiliation":[{"name":"Universidad Nacional de Colombia, Sede de La Paz  La Paz Cesar Colombia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5978-9312","authenticated-orcid":false,"given":"Teddy","family":"Angarita\u2010Sierra","sequence":"additional","affiliation":[{"name":"Universidad Nacional de Colombia, Sede de La Paz  La Paz Cesar Colombia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4253-1101","authenticated-orcid":false,"given":"Santiago J.","family":"S\u00e1nchez\u2010Pacheco","sequence":"additional","affiliation":[{"name":"Universidad Nacional de Colombia, Sede de La Paz  La Paz Cesar Colombia"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,10,9]]},"reference":[{"issue":"3","key":"e_1_2_10_2_1","first-page":"1","article-title":"A new Telmatobius and new teiid lizards from South America","volume":"9","author":"Andersson L. G.","year":"1914","journal-title":"Arkiv f\u00f6r Zoologi"},{"key":"e_1_2_10_3_1","volume-title":"Reptiles of Ecuador: Life in the middle of the world","author":"Arteaga A.","year":"2024"},{"key":"e_1_2_10_4_1","doi-asserted-by":"publisher","DOI":"10.1046\/j.1096-3642.2003.00077.x"},{"key":"e_1_2_10_5_1","volume-title":"Catalogue of the lizards in the British museum (natural history)","author":"Boulenger G. A.","year":"1885"},{"issue":"1","key":"e_1_2_10_6_1","first-page":"77","article-title":"First report on additions to the lizard collection in the British museum (natural history)","volume":"1890","author":"Boulenger G. A.","year":"1890","journal-title":"Proceedings of the Zoological Society of London"},{"key":"e_1_2_10_7_1","doi-asserted-by":"publisher","DOI":"10.1080\/10635150490445797"},{"key":"e_1_2_10_8_1","doi-asserted-by":"publisher","DOI":"10.1206\/310.1"},{"issue":"2","key":"e_1_2_10_9_1","first-page":"159","article-title":"Report on the reptiles brought by Professor James Orton from the middle and upper Amazon, and western Peru","volume":"8","author":"Cope E.","year":"1875","journal-title":"Journal of the Academy of Natural Sciences of Philadelphia"},{"key":"e_1_2_10_10_1","doi-asserted-by":"publisher","DOI":"10.2994\/SAJH-D-13-00022.1"},{"key":"e_1_2_10_11_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1096-3642.2005.00145.x"},{"key":"e_1_2_10_12_1","first-page":"1","volume-title":"Biology of the Reptilia, Vol. 20 morphology H: The skull of Lepidosauria","author":"Evans S. E.","year":"2008"},{"key":"e_1_2_10_13_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.25572"},{"key":"e_1_2_10_14_1","doi-asserted-by":"publisher","DOI":"10.5962\/bhl.title.4683"},{"key":"e_1_2_10_15_1","doi-asserted-by":"publisher","DOI":"10.1111\/cla.12150"},{"key":"e_1_2_10_16_1","unstructured":"Gray J. A.(2018).Skull evolution in the Australian dragon lizards[PhD thesis University of Adelaide]."},{"key":"e_1_2_10_17_1","doi-asserted-by":"publisher","DOI":"10.1080\/00222933809512291"},{"key":"e_1_2_10_18_1","volume-title":"Catalogue of the specimens of lizards in the collection of the British museum","author":"Gray J. E.","year":"1845"},{"key":"e_1_2_10_19_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1096-3642.1851.tb01128.x"},{"key":"e_1_2_10_20_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7998.1858.tb06398.x"},{"key":"e_1_2_10_21_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1463-6395.2008.00378.x"},{"key":"e_1_2_10_22_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.24038"},{"key":"e_1_2_10_23_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7580.2012.01549.x"},{"key":"e_1_2_10_24_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.25530"},{"key":"e_1_2_10_25_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.24246"},{"key":"e_1_2_10_26_1","first-page":"27","article-title":"A reappraisal of the phylogeny of lizards of the family Gymnophthalmidae (Sauria, Scincomorpha)","volume":"12","author":"Hoyos J. M.","year":"1998","journal-title":"Revista Espa\u00f1ola de Herpetolog\u00eda"},{"key":"e_1_2_10_27_1","doi-asserted-by":"publisher","DOI":"10.2307\/1466981"},{"key":"e_1_2_10_28_1","doi-asserted-by":"publisher","DOI":"10.1111\/zoj.12241"},{"key":"e_1_2_10_29_1","first-page":"53","article-title":"Osteolog\u00eda craneal de Pantodactylus schreibersii schreibersii (Wiegmann, 1834) y su contribuci\u00f3n a la discusi\u00f3n de Gymnophthalmidae (Reptilia)","volume":"23","author":"L\u00f3pez A. M.","year":"1995","journal-title":"Anales del Museo de Historia Natural de Valpara\u00edso"},{"key":"e_1_2_10_30_1","doi-asserted-by":"publisher","DOI":"10.11606\/0031-1049.1974.27.p179-213"},{"key":"e_1_2_10_31_1","unstructured":"McKee H.(2021).Cranial osteology and comparison of Oreosaurus luctuosus and Oreosaurus rhodogaster[Spring Honors Capstone Projects University of Texas at Arlington].https:\/\/mavmatrix.uta.edu\/honors_spring2021\/61"},{"key":"e_1_2_10_32_1","volume-title":"Versuch eines Systems der Amphibien. Tentamen Systematis Amphibiorum","author":"Merrem B.","year":"1820"},{"issue":"3","key":"e_1_2_10_33_1","first-page":"215","article-title":"Cranial anatomy of Euspondylus acutirostris (Squamata: Gymnophthalmidae) and its placement in a modern phylogenetic hypothesis","volume":"9","author":"Montero R.","year":"2002","journal-title":"Russian Journal of Herpetology"},{"key":"e_1_2_10_34_1","doi-asserted-by":"publisher","DOI":"10.1080\/00222937908679832"},{"key":"e_1_2_10_35_1","first-page":"105","article-title":"Sesamoid elements in lizards","volume":"23","author":"Otero T.","year":"2013","journal-title":"Herpetological Journal"},{"key":"e_1_2_10_36_1","doi-asserted-by":"publisher","DOI":"10.1006\/bijl.2001.0580"},{"key":"e_1_2_10_37_1","first-page":"165","article-title":"\u00dcber Cercosaura und die mit dieser Gattung verwandten Eidechsen aus S\u00fcdamerica","volume":"1862","author":"Peters W.","year":"1862","journal-title":"Abhandlungen der K\u00f6niglichen Akademie der Wissenschaften zu Berlin"},{"key":"e_1_2_10_38_1","doi-asserted-by":"publisher","DOI":"10.2307\/1444132"},{"key":"e_1_2_10_39_1","doi-asserted-by":"publisher","DOI":"10.11606\/0031-1049.1984.35.p361-366"},{"issue":"19","key":"e_1_2_10_40_1","first-page":"285","article-title":"Herpetofauna das dunas interiores do Rio S\u00e3o Francisco, Bahia, Brasil. I. Introdu\u00e7\u00e3o \u00e0 \u00e1rea e descri\u00e7\u00e3o de um novo g\u00eanero de Microteiideos (Calyptommatus) com notas sobre sua ecologia, distribui\u00e7\u00e3o e especia\u00e7\u00e3o (Sauria, Teiidae)","volume":"37","author":"Rodrigues M. T.","year":"1991","journal-title":"Papeis Avulsos de Zoologia"},{"key":"e_1_2_10_41_1","doi-asserted-by":"publisher","DOI":"10.11646\/zootaxa.1873.1.5"},{"key":"e_1_2_10_42_1","doi-asserted-by":"publisher","DOI":"10.1206\/0003-0082(2007)496[1:ANGOML]2.0.CO;2"},{"key":"e_1_2_10_43_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.10878"},{"key":"e_1_2_10_44_1","doi-asserted-by":"publisher","DOI":"10.1655\/09-033.1"},{"key":"e_1_2_10_45_1","doi-asserted-by":"publisher","DOI":"10.1590\/S0031-10492010000200001"},{"key":"e_1_2_10_46_1","doi-asserted-by":"publisher","DOI":"10.11865\/zs.2024401"},{"key":"e_1_2_10_47_1","first-page":"11","article-title":"Notes on the occurrence of Riama simotera (Squamata, Gymnophthalmidae) in Colombia","volume":"113","author":"S\u00e1nchez\u2010Pacheco S. J.","year":"2010","journal-title":"Herpetological Bulletin"},{"key":"e_1_2_10_48_1","doi-asserted-by":"publisher","DOI":"10.1111\/cla.12203"},{"key":"e_1_2_10_49_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.10288"},{"key":"e_1_2_10_50_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1096-3642.2007.00364.x"},{"key":"e_1_2_10_51_1","first-page":"107","article-title":"Revised procedures for staining and clearing small fishes and other vertebrates for bone and cartilage study","volume":"9","author":"Taylor W. R.","year":"1985","journal-title":"Cybium"},{"key":"e_1_2_10_52_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2016.03.006"},{"key":"e_1_2_10_53_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2013.03.013"},{"key":"e_1_2_10_54_1","doi-asserted-by":"publisher","DOI":"10.22201\/fc.25942158e.2022.2.422"},{"key":"e_1_2_10_55_1","unstructured":"Uetz P. Freed P. Aguilar R. Reyes F. Kudera J. &Ho\u0161ek J.(2026).The Reptile Database.http:\/\/www.reptile-database.org"},{"key":"e_1_2_10_56_1","doi-asserted-by":"publisher","DOI":"10.1093\/zoolinnean\/zlz124"},{"key":"e_1_2_10_57_1","doi-asserted-by":"publisher","DOI":"10.1093\/biolinnean\/blae080"},{"key":"e_1_2_10_58_1","doi-asserted-by":"publisher","DOI":"10.1163\/15685381-00003085"},{"key":"e_1_2_10_59_1","doi-asserted-by":"publisher","DOI":"10.5962\/bhl.part.7963"},{"key":"e_1_2_10_60_1","volume-title":"Nat\u00fcrliches System der Amphibien: Mit vorangehender Classification der S\u00e4ugethiere und V\u00f6gel; ein Beitrag zur vergleichenden Zoologie","author":"Wagler J. G.","year":"1830"},{"key":"e_1_2_10_61_1","doi-asserted-by":"publisher","DOI":"10.7717\/peerj.12523"}],"container-title":["The Anatomical Record"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/anatomypubs.onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/ar.70367","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/anatomypubs.onlinelibrary.wiley.com\/doi\/full-xml\/10.1002\/ar.70367","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/anatomypubs.onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/ar.70367","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T07:33:12Z","timestamp":1791531192000},"score":1,"resource":{"primary":{"URL":"https:\/\/anatomypubs.onlinelibrary.wiley.com\/doi\/10.1002\/ar.70367"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10,9]]},"references-count":60,"alternative-id":["10.1002\/ar.70367"],"URL":"https:\/\/doi.org\/10.1002\/ar.70367","archive":["Portico"],"relation":{},"ISSN":["1932-8486","1932-8494"],"issn-type":[{"value":"1932-8486","type":"print"},{"value":"1932-8494","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,9]]},"assertion":[{"value":"2026-04-21","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-09-15","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-10-09","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"ar.70367"}}