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(Northeastern France)","volume":"26","author":"P Godefroit","year":"1997","journal-title":"Palaeovertebrata"},{"key":"ref15","first-page":"64A","article-title":"Were there venomous theropods?","volume":"20","author":"RA Rodriguez-de la Rosa","year":"2000","journal-title":"J Vert Paleontol"},{"key":"ref16","doi-asserted-by":"crossref","first-page":"547","DOI":"10.1671\/0272-4634(2007)27[547:CGMFAR]2.0.CO;2","article-title":"Canine grooves: morphology, function, and relevance to venom","volume":"27","author":"KE Folinsbee","year":"2007","journal-title":"J Vert Paleontol"},{"key":"ref17","doi-asserted-by":"crossref","first-page":"409","DOI":"10.1134\/S0031030108040102","article-title":"The first Whaitsiid (Therocephalia, Theromorpha) from the terminal Permian of eastern Europe","volume":"42","author":"MF Ivakhnenko","year":"2008","journal-title":"Paleontol J"},{"key":"ref18","doi-asserted-by":"crossref","first-page":"669","DOI":"10.4202\/app.2009.0045","article-title":"The earliest known 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of Karroo Fossil Reptiles","volume":"41","author":"R Broom","year":"1931","journal-title":"Rec Albany Mus"},{"key":"ref30","article-title":"The mammal-like reptiles of South Africa and the origin of mammals","author":"R Broom","year":"1932"},{"key":"ref31","doi-asserted-by":"crossref","first-page":"20","DOI":"10.2307\/1564416","article-title":"Observations of Tooth Surface Morphology in Snakes","volume":"19","author":"RH Vaeth","year":"1985","journal-title":"J Herpetol"},{"key":"ref32","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1671\/0272-4634(2007)27[541:TCMATI]2.0.CO;2","article-title":"The comparative method and the inference of venom-delivery systems in fossil mammals","volume":"27","author":"CM Orr","year":"2007","journal-title":"J Vert Paleontol"},{"key":"ref33","doi-asserted-by":"crossref","first-page":"13022","DOI":"10.1073\/pnas.1402905111","article-title":"New high-resolution computed tomography data of the Taung partial cranium and endocast and their bearing on metopism and hominin brain evolution","volume":"111","author":"RL Holloway","year":"2014","journal-title":"Proc Natl Acad Sci USA"},{"key":"ref34","doi-asserted-by":"crossref","first-page":"25604","DOI":"10.1038\/srep25604","article-title":"Palaeoneurological clues to the evolution of defining mammalian soft tissue traits","volume":"6","author":"J Benoit","year":"2016","journal-title":"Sci Rep"},{"issue":"11\u201312","key":"ref35","first-page":"69","article-title":"Physiological implications of the abnormal absence of the parietal foramen in a Late Permian cynodont (Therapsida)","volume":"102","author":"J Benoit","year":"2005","journal-title":"Sci Nat"},{"issue":"6","key":"ref36","doi-asserted-by":"crossref","first-page":"e64978","DOI":"10.1371\/journal.pone.0064978","article-title":"Synchrotron Reveals Early Triassic Odd Couple:Injured Amphibian and Aestivating Therapsid Share Burrow","volume":"8","author":"V Fernandez","year":"2013","journal-title":"PLoS 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ISE-MeshTools, a 3D interactive fossil reconstruction freeware. 12th Annual Meeting of EAVP, Torino, Italy; 2014."},{"key":"ref38","first-page":"1","article-title":"Review of the Pelycosauria","volume":"28","author":"AS Romer","year":"1940","journal-title":"Geol Soc Am Spec Pap"},{"key":"ref39","article-title":"Vertebrate Paleontology","author":"AS Romer","year":"1945"},{"key":"ref40","first-page":"9","article-title":"International Colloquium on the Evolution of Lower and Non Specialized Mammals. 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The comparative and functional morphology of the early Therocephalia (Amniota: Therapsida). Ph. D. Thesis, University of Stellenbosch. 1987. Available from: <ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"http:\/\/scholar.sun.ac.za\/handle\/10019.1\/65930\" xlink:type=\"simple\">http:\/\/scholar.sun.ac.za\/handle\/10019.1\/65930<\/ext-link>"},{"key":"ref53","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1111\/j.1469-7998.1961.tb06083.x","article-title":"Notes on tooth replacement in the Nile crocodile <italic>Crocodilus niloticus<\/italic>","volume":"136","author":"DFG Poole","year":"1961","journal-title":"Proc Zool Soc Lond"},{"key":"ref54","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1002\/jmor.1052180208","article-title":"Heterodonty and patterns of tooth replacement in <italic>Crocodylus niloticus<\/italic>","volume":"218","author":"JA Kieser","year":"1993","journal-title":"J Morphol"},{"key":"ref55","first-page":"152","article-title":"Chemical Signals in Vertebrates","volume":"10","author":"SJ Rehorek","year":"2005"},{"key":"ref56","doi-asserted-by":"crossref","first-page":"1433","DOI":"10.1002\/ar.21046","article-title":"The Orbitofacial Glands of Bats: An Investigation of the Potential Correlation of Gland Structure with Social Organization","volume":"293","author":"SJ Rehorek","year":"2010","journal-title":"Anat Rec"},{"key":"ref57","first-page":"35","article-title":"Evidence for salt glands in the Triassic reptile <italic>Diademodon<\/italic> (Therapsida; Cynodontia)","volume":"22","author":"FE Grine","year":"1979","journal-title":"Pal Afr"},{"key":"ref58","article-title":"Ost\u00e9ologie","volume":"1","author":"R Barone","year":"1999"},{"key":"ref59","doi-asserted-by":"crossref","first-page":"1019","DOI":"10.1086\/425185","article-title":"The Evolution of Endothermy in Terrestrial Vertebrates: Who? 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