334 lines
87 KiB
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334 lines
87 KiB
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<document ID-DOI="http://dx.doi.org/10.3897/vz.73.e101372" ID-Pensoft-Pub="2625-8498-73-717" ID-Pensoft-UUID="318B657A15AB5708B3C35FC1A82B4945" ID-ZooBank="8F3D5EDA2F184E5CA53E2F7741FF1339" ModsDocID="2625-8498-73-717" checkinTime="1695893367890" checkinUser="pensoft" docAuthor="Szyndlar, Zbigniew & Georgalis, Georgios L." docDate="2023" docId="4C442FBE5F3D63DAC3405CF3AF0F26F9" docLanguage="en" docName="VertZool 73: 717-886" docOrigin="Vertebrate Zoology 73" docPubDate="2023-09-27" docSource="http://dx.doi.org/10.3897/vz.73.e101372" docTitle="Constrictores Duméril & Bibron 1844" docType="treatment" docVersion="1" id="318B657A15AB5708B3C35FC1A82B4945" lastPageNumber="717" masterDocId="318B657A15AB5708B3C35FC1A82B4945" masterDocTitle="An illustrated atlas of the vertebral morphology of extant non-caenophidian snakes, with special emphasis on the cloacal and caudal portions of the column" masterLastPageNumber="886" masterPageNumber="717" pageNumber="717" updateTime="1695893367890" updateUser="pensoft">
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<mods:title id="7B153387617C750DE0D066FE4A9B9B36">An illustrated atlas of the vertebral morphology of extant non-caenophidian snakes, with special emphasis on the cloacal and caudal portions of the column</mods:title>
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<mods:namePart id="95C4DC19CB66B0009F7295D1822B6E87">Szyndlar, Zbigniew</mods:namePart>
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<mods:affiliation id="16F3E43FD74BFEAA328D8CD759D48305">Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Slawkowska 17, 31 - 016 Krakow, Poland</mods:affiliation>
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<mods:namePart id="328F2777A4B3126A891CF98007993E87">Georgalis, Georgios L.</mods:namePart>
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<mods:affiliation id="CCD7B943AC92900F9B96E1D4D4616977">Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Slawkowska 17, 31 - 016 Krakow, Poland</mods:affiliation>
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<mods:nameIdentifier id="50B88220ACF051C8EC96BA56FB964615" type="email">dimetrodon82@gmail.com</mods:nameIdentifier>
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<mods:title id="788F8BEF312796C2C0CE483D1B33FC47">Vertebrate Zoology</mods:title>
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<mods:date id="B08986E0711C05929E7371FDEAED1426">2023</mods:date>
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<mods:identifier id="DE834FF041EE6FC6D158FC39A0305B1A" type="DOI">http://dx.doi.org/10.3897/vz.73.e101372</mods:identifier>
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<paragraph id="0DDE521B4B02998E94C6937DD153F1F1" pageId="0" pageNumber="717">
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<taxonomicName id="76A8F1AD7389874FD2185352DC3DD9F3" authority="Duméril & Bibron, 1844" authorityName="Duméril & Bibron" authorityYear="1844" higherTaxonomySource="treatment-meta" infraOrder="Scolecophidia" kingdom="Animalia" lsidName="" order="Squamata" pageId="0" pageNumber="717" rank="infraOrder">
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Scolecophidia
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& Bibron, 1844
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<subSubSection id="B2CDD1C9ED2348AB70ED6F1F37DFAB7D" pageId="0" pageNumber="717" type="general information">
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<paragraph id="6F7CC1DD5B569806259D9C7D333D3F85" pageId="0" pageNumber="717">General information.</paragraph>
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Commonly called "worm snakes" (as their name in Greek literally translates [σκώληξ +
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<normalizedToken id="86118C8F5CF5FD0E0B5C8BDA94463320" originalValue="ὀφίδια">ὀφίδια</normalizedToken>
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]) or "blind snakes",
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<taxonomicName id="535487CD1958DE118E8158C88FD23958" infraOrder="Scolecophidia" kingdom="Animalia" lsidName="" order="Squamata" pageId="0" pageNumber="717" rank="infraOrder">Scolecophidia</taxonomicName>
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was for a long time considered a monophyletic assemblage of basal snakes (e.g.,
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<bibRefCitation id="A50FB957F916B68763BD715C857F2779" author="Underwood, G" journalOrPublisher="British Museum (Natural History), London" pageId="0" pageNumber="717" refId="B634" refString="Underwood, G, 1967. A Contribution to the Classification of Snakes. British Museum (Natural History), London" title="A Contribution to the Classification of Snakes." year="1967">Underwood 1967</bibRefCitation>
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;
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<bibRefCitation id="073C41C9EC5F50AED035C4A73359FA5A" author="Rage, J-C" editor="Wellnhofer, P" journalOrPublisher="Gustav Fischer, Stuttgart, New York" pageId="0" pageNumber="717" pagination="1 - 80" refId="B477" refString="Rage, J-C, 1984. Serpentes. In: Wellnhofer, P, Ed., Encyclopedia of Paleoherpetology, Part 11. Gustav Fischer, Stuttgart, New York: 1 - 80" title="Serpentes." volumeTitle="Encyclopedia of Paleoherpetology, Part 11." year="1984">Rage 1984</bibRefCitation>
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). Certain 19th century workers even included them within lizards and not snakes (e.g.,
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<bibRefCitation id="85DECA6FD73E592F0FB5DD2F28C0C380" DOI="https://doi.org/10.5962/bhl.title.4058" author="Bonaparte, CL" journalOrPublisher="Memorie della Reale Accademia delle Scienze di Torino (Serie II)" pageId="0" pageNumber="717" pagination="385 - 456" refId="B70" refString="Bonaparte, CL, 1839. Amphibia Europaea. Ad systema nostrum vertebratorum ordinata. Typographica regia. Memorie della Reale Accademia delle Scienze di Torino (Serie II) 2: 385 - 456, DOI: https://doi.org/10.5962/bhl.title.4058" title="Amphibia Europaea. Ad systema nostrum vertebratorum ordinata. Typographica regia." url="https://doi.org/10.5962/bhl.title.4058" volume="2" year="1839">Bonaparte 1839</bibRefCitation>
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;
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<bibRefCitation id="3A0BED40EC4FC909374E2369585FB65E" author="Gray, JE" journalOrPublisher="British Museum (Natural History), London" pageId="0" pageNumber="717" refId="B224" refString="Gray, JE, 1845. Catalogue of the Specimens of Lizards in the Collection of the British Museum. British Museum (Natural History), London" title="Catalogue of the Specimens of Lizards in the Collection of the British Museum." year="1845">Gray 1845</bibRefCitation>
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). Notably, a similar view was held also more than one century after, when
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<bibRefCitation id="F9FC1BD8D26574E683965771C7CB5D48" author="McDowell, SB" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="0" pageNumber="717" pagination="1 - 145" refId="B381" refString="McDowell, SB, Bogert, CM, 1954. The systematic position of Lanthanotus and the affinities of the anguinomorphan lizards. Bulletin of the American Museum of Natural History 105: 1 - 145" title="The systematic position of Lanthanotus and the affinities of the anguinomorphan lizards." volume="105" year="1954">McDowell and Bogert (1954)</bibRefCitation>
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suggested that
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<taxonomicName id="F392842EDDF704F1E8FF836FF0F323A3" infraOrder="Scolecophidia" kingdom="Animalia" lsidName="" order="Squamata" pageId="0" pageNumber="717" rank="infraOrder">Scolecophidia</taxonomicName>
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do not belong to snakes, and instead represent a lineage of limbless lizards that became convergent to snakes, a view accepted also by few others (
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<bibRefCitation id="0BF6B533AC2B8AD5F5DCA4AD50E7BA6B" DOI="https://doi.org/10.1071/ZO9600181" author="Robb, J" journalOrPublisher="Australian Journal of Zoology" pageId="0" pageNumber="717" pagination="181 - 216" refId="B506" refString="Robb, J, 1960. The internal anatomy of Typhlops Schneider (Repilia). Australian Journal of Zoology 8: 181 - 216, DOI: https://doi.org/10.1071/ZO9600181" title="The internal anatomy of Typhlops Schneider (Repilia)." url="https://doi.org/10.1071/ZO9600181" volume="8" year="1960">Robb 1960</bibRefCitation>
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;
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<bibRefCitation id="50C8C6E6A9388886D49A05049E0059FE" DOI="https://doi.org/10.2307/1441214" author="Goin, C" journalOrPublisher="Freeman, San Francisco, CA" pageId="0" pageNumber="717" refId="B211" refString="Goin, C, Goin, O, 1962. Introduction to Herpetology. W. H. Freeman, San Francisco, CA, DOI: https://doi.org/10.2307/1441214" title="Introduction to Herpetology. W. H." url="https://doi.org/10.2307/1441214" year="1962">Goin and Goin 1962</bibRefCitation>
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)! Nevertheless, the true ophidian nature of scolecophidians is currently universally accepted, though their monophyly has been seriously challenged, primarily based on molecular data (
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<bibRefCitation id="4E70267FA84F7172B2E99CB67CA0341C" DOI="https://doi.org/10.1016/j.crvi.2005.10.001" author="Vidal, N" journalOrPublisher="Comptes Rendus Biologies" pageId="0" pageNumber="717" pagination="1000 - 1008" refId="B643" refString="Vidal, N, Hedges, BS, 2005. The phylogeny of squamate reptiles (lizards, snakes, and amphisbaenians) inferred from nine nuclear protein-coding genes. Comptes Rendus Biologies 328: 1000 - 1008, DOI: https://doi.org/10.1016/j.crvi.2005.10.001" title="The phylogeny of squamate reptiles (lizards, snakes, and amphisbaenians) inferred from nine nuclear protein-coding genes." url="https://doi.org/10.1016/j.crvi.2005.10.001" volume="328" year="2005">Vidal and Hedges 2005</bibRefCitation>
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<bibRefCitation id="926026AD5876E83CA04D09CDC7F1EA10" DOI="https://doi.org/10.1080/10635150802166053" author="Wiens, JJ" journalOrPublisher="Systematic Biology" pageId="0" pageNumber="717" pagination="420 - 431" refId="B679" refString="Wiens, JJ, Kuczynski, CA, Smith, SA, Mulcahy, DG, Sites, JW Jr., Townsend, TM, Reeder, TW, 2008. Branch lengths, support, and congruence: Testing the phylogenomic approach with 20 nuclear loci in snakes. Systematic Biology 57: 420 - 431, DOI: https://doi.org/10.1080/10635150802166053" title="Branch lengths, support, and congruence: Testing the phylogenomic approach with 20 nuclear loci in snakes." url="https://doi.org/10.1080/10635150802166053" volume="57" year="2008">Wiens et al. 2008</bibRefCitation>
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;
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<bibRefCitation id="DA7131E965484CAFB009860895DE212F" DOI="https://doi.org/10.1098/rsbl.2010.0220" author="Vidal, N" journalOrPublisher="Biology Letters" pageId="0" pageNumber="717" pagination="1 - 4" refId="B645" refString="Vidal, N, Marin, J, Morini, M, Donnellan, S, Branch, WR, Thomas, R, Vences, M, Wynn, A, Cruaud, C, Hedges, BS, 2010. Blindsnake evolutionary tree reveals long history on Gondwana. Biology Letters 2010: 1 - 4, DOI: https://doi.org/10.1098/rsbl.2010.0220" title="Blindsnake evolutionary tree reveals long history on Gondwana." url="https://doi.org/10.1098/rsbl.2010.0220" volume="2010" year="2010">Vidal et al. 2010</bibRefCitation>
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<bibRefCitation id="248DDF7220EFB891CF25B6BE000BF04D" DOI="https://doi.org/10.1111/j.1558-5646.2011.01437.x" author="Pyron, RA" journalOrPublisher="Evolution" pageId="0" pageNumber="717" pagination="163 - 178" refId="B467" refString="Pyron, RA, Burbrink, FT, 2012. Extinction, ecological opportunity, and the origins of global snake diversity. Evolution 66: 163 - 178, DOI: https://doi.org/10.1111/j.1558-5646.2011.01437.x" title="Extinction, ecological opportunity, and the origins of global snake diversity." url="https://doi.org/10.1111/j.1558-5646.2011.01437.x" volume="66" year="2012">Pyron and Burbrink 2012</bibRefCitation>
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<bibRefCitation id="3FE70E2362B6463868E3CAAB4A172327" DOI="https://doi.org/10.1186/1471-2148-13-93" author="Pyron, RA" journalOrPublisher="Evolution" pageId="0" pageNumber="717" refId="B468" refString="Pyron, RA, Burbrink, FT, Wiens, JJ, 2013. A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology 13: 93. https://doi.org/10.1186/1471-2148-13-93" title="A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology 13: 93." url="https://doi.org/10.1186/1471-2148-13-93" year="2013">Pyron et al. 2013</bibRefCitation>
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<bibRefCitation id="4A5D5DCDB0E09C5422E1C87C0B503674" DOI="https://doi.org/10.1186/s12862-015-0358-5" author="Hsiang, AY" journalOrPublisher="Journal of Anatomy" pageId="0" pageNumber="717" refId="B287" refString="Hsiang, AY, Field, DJ, Webster, TH, Behlke, ADB, Davis, MB, Racicot, RA, Gauthier, JA, 2015. The origin of snakes: Revealing the ecology, behavior, and evolutionary history of early snakes using genomics, phenomics, and the fossil record. BMC Evolutionary Biology 15: 87. https://doi.org/10.1186/s12862-015-0358-5" title="The origin of snakes: Revealing the ecology, behavior, and evolutionary history of early snakes using genomics, phenomics, and the fossil record. BMC Evolutionary Biology 15: 87." url="https://doi.org/10.1186/s12862-015-0358-5" year="2015">Hsiang et al. 2015</bibRefCitation>
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<bibRefCitation id="8BC8F61C4586DAE53539AB47CF6BD390" DOI="https://doi.org/10.1371/journal.pone.0118199" author="Reeder, TW" journalOrPublisher="Biological Journal of the Linnean Society" pageId="0" pageNumber="717" refId="B494" refString="Reeder, TW, Townsend, TM, Mulcahy, DG, Noonan, BP, Wood, PL Jr, Sites, JW Jr, Wiens, JJ, 2015. Integrated analyses resolve conflicts over squamate reptile phylogeny and reveal unexpected placements for fossil taxa. PLoS ONE 10: e0118199. https://doi.org/10.1371/journal.pone.0118199" title="Integrated analyses resolve conflicts over squamate reptile phylogeny and reveal unexpected placements for fossil taxa. PLoS ONE 10: e 0118199." url="https://doi.org/10.1371/journal.pone.0118199" year="2015">Reeder et al. 2015</bibRefCitation>
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<bibRefCitation id="B9C11F7A9ADEF5DC3DB1E38F3D22A0FC" DOI="https://doi.org/10.1016/j.ympev.2015.10.009" author="Zheng, Y" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="717" pagination="537 - 547" refId="B696" refString="Zheng, Y, Wiens, JJ, 2016. Combining phylogenomic and supermatrix approaches, and a time-calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52 genes and 4162 species. Molecular Phylogenetics and Evolution 94: 537 - 547, DOI: https://doi.org/10.1016/j.ympev.2015.10.009" title="Combining phylogenomic and supermatrix approaches, and a time-calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52 genes and 4162 species." url="https://doi.org/10.1016/j.ympev.2015.10.009" volume="94" year="2016">Zheng and Wiens 2016</bibRefCitation>
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<bibRefCitation id="40269142625362827A188D194CC72B07" DOI="https://doi.org/10.1098/rsbl.2017.0393" author="Streicher, JW" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="717" refId="B597" refString="Streicher, JW, Wiens, JJ, 2017. Phylogenomic analyses of more than 4000 nuclear loci resolve the origin of snakes among lizard families. Biology Letters 13: 20170393. https://doi.org/10.1098/rsbl.2017.0393" title="Phylogenomic analyses of more than 4000 nuclear loci resolve the origin of snakes among lizard families. Biology Letters 13: 20170393." url="https://doi.org/10.1098/rsbl.2017.0393" year="2017">Streicher and Wiens 2017</bibRefCitation>
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<bibRefCitation id="4310C1F564BC505AD73C9DEE9E91C44A" DOI="https://doi.org/10.1111/jeb.13373" author="Miralles, A" journalOrPublisher="Journal of Evolutionary Biology" pageId="0" pageNumber="717" pagination="1782 - 1793" refId="B387" refString="Miralles, A, Marin, J, Markus, D, Herrel, A, Hedges, BS, Vidal, N, 2018. Molecular evidence for the paraphyly of Scolecophidia and its evolutionary implications. Journal of Evolutionary Biology 31: 1782 - 1793, DOI: https://doi.org/10.1111/jeb.13373" title="Molecular evidence for the paraphyly of Scolecophidia and its evolutionary implications." url="https://doi.org/10.1111/jeb.13373" volume="31" year="2018">Miralles et al. 2018</bibRefCitation>
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;
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<bibRefCitation id="AB0B2B9DDC79CE21FE13A197D0D4E660" DOI="https://doi.org/10.1093/sysbio/syz062" author="Burbrink, FT" journalOrPublisher="Systematic Biology" pageId="0" pageNumber="717" pagination="502 - 520" refId="B101" refString="Burbrink, FT, Grazziotin, FG, Pyron, RA, Cundall, D, Donnellan, S, Irish, F, Keogh, JS, Kraus, F, Murphy, RW, Noonan, B, Raxworthy, CJ, Ruane, S, Lemmon, AR, Lemmon, EM, Zaher, H, 2020. Interrogating genomic-scale data for Squamata (lizards, snakes, and amphisbaenians) shows no support for key traditional morphological relationships. Systematic Biology 69: 502 - 520, DOI: https://doi.org/10.1093/sysbio/syz062" title="Interrogating genomic-scale data for Squamata (lizards, snakes, and amphisbaenians) shows no support for key traditional morphological relationships." url="https://doi.org/10.1093/sysbio/syz062" volume="69" year="2020">Burbrink et al. 2020</bibRefCitation>
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), as well as total evidence analyses (
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<bibRefCitation id="821FF88C8C29F8C7ECD3D4503EB4D693" DOI="https://doi.org/10.1093/zoolinnean/zlac001" author="Zaher, H" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="0" pageNumber="717" pagination="656 - 697" refId="B694" refString="Zaher, H, Mohabey, DM, Grazziotin, FG, Mantilla, JAW, 2023. The skull of Sanajeh indicus, a Cretaceous snake with an upper temporal bar, and the origin of ophidian wide-gaped feeding. Zoological Journal of the Linnean Society 197: 656 - 697, DOI: https://doi.org/10.1093/zoolinnean/zlac001" title="The skull of Sanajeh indicus, a Cretaceous snake with an upper temporal bar, and the origin of ophidian wide-gaped feeding." url="https://doi.org/10.1093/zoolinnean/zlac001" volume="197" year="2023">Zaher et al. 2023</bibRefCitation>
|
||
). More specifically, recent analyses have suggested that anomalepidids either represent the sister group of alethinophidians, being thus not sharing an exclusive common ancestor with the remaining scolecophidians, or lie more basal to leptotyphlopids and typhlopoids (
|
||
<bibRefCitation id="7B9B71C75AC8D2621947BD726DF111D0" DOI="https://doi.org/10.1080/10635150802166053" author="Wiens, JJ" journalOrPublisher="Systematic Biology" pageId="0" pageNumber="717" pagination="420 - 431" refId="B679" refString="Wiens, JJ, Kuczynski, CA, Smith, SA, Mulcahy, DG, Sites, JW Jr., Townsend, TM, Reeder, TW, 2008. Branch lengths, support, and congruence: Testing the phylogenomic approach with 20 nuclear loci in snakes. Systematic Biology 57: 420 - 431, DOI: https://doi.org/10.1080/10635150802166053" title="Branch lengths, support, and congruence: Testing the phylogenomic approach with 20 nuclear loci in snakes." url="https://doi.org/10.1080/10635150802166053" volume="57" year="2008">Wiens et al. 2008</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="A182528513D6FE2C143EAB458D079286" DOI="https://doi.org/10.1111/j.1558-5646.2011.01437.x" author="Pyron, RA" journalOrPublisher="Evolution" pageId="0" pageNumber="717" pagination="163 - 178" refId="B467" refString="Pyron, RA, Burbrink, FT, 2012. Extinction, ecological opportunity, and the origins of global snake diversity. Evolution 66: 163 - 178, DOI: https://doi.org/10.1111/j.1558-5646.2011.01437.x" title="Extinction, ecological opportunity, and the origins of global snake diversity." url="https://doi.org/10.1111/j.1558-5646.2011.01437.x" volume="66" year="2012">Pyron and Burbrink 2012</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="C09DE71E0DD0C1800FF403CE63C84552" DOI="https://doi.org/10.1111/jeb.13373" author="Miralles, A" journalOrPublisher="Journal of Evolutionary Biology" pageId="0" pageNumber="717" pagination="1782 - 1793" refId="B387" refString="Miralles, A, Marin, J, Markus, D, Herrel, A, Hedges, BS, Vidal, N, 2018. Molecular evidence for the paraphyly of Scolecophidia and its evolutionary implications. Journal of Evolutionary Biology 31: 1782 - 1793, DOI: https://doi.org/10.1111/jeb.13373" title="Molecular evidence for the paraphyly of Scolecophidia and its evolutionary implications." url="https://doi.org/10.1111/jeb.13373" volume="31" year="2018">Miralles et al. 2018</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="A49E6B174C9D5886A9881882CBB0D230" DOI="https://doi.org/10.1093/sysbio/syz062" author="Burbrink, FT" journalOrPublisher="Systematic Biology" pageId="0" pageNumber="717" pagination="502 - 520" refId="B101" refString="Burbrink, FT, Grazziotin, FG, Pyron, RA, Cundall, D, Donnellan, S, Irish, F, Keogh, JS, Kraus, F, Murphy, RW, Noonan, B, Raxworthy, CJ, Ruane, S, Lemmon, AR, Lemmon, EM, Zaher, H, 2020. Interrogating genomic-scale data for Squamata (lizards, snakes, and amphisbaenians) shows no support for key traditional morphological relationships. Systematic Biology 69: 502 - 520, DOI: https://doi.org/10.1093/sysbio/syz062" title="Interrogating genomic-scale data for Squamata (lizards, snakes, and amphisbaenians) shows no support for key traditional morphological relationships." url="https://doi.org/10.1093/sysbio/syz062" volume="69" year="2020">Burbrink et al. 2020</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="1D2D10570EC52D42710444EE726B241D" DOI="https://doi.org/10.1093/zoolinnean/zlac001" author="Zaher, H" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="0" pageNumber="717" pagination="656 - 697" refId="B694" refString="Zaher, H, Mohabey, DM, Grazziotin, FG, Mantilla, JAW, 2023. The skull of Sanajeh indicus, a Cretaceous snake with an upper temporal bar, and the origin of ophidian wide-gaped feeding. Zoological Journal of the Linnean Society 197: 656 - 697, DOI: https://doi.org/10.1093/zoolinnean/zlac001" title="The skull of Sanajeh indicus, a Cretaceous snake with an upper temporal bar, and the origin of ophidian wide-gaped feeding." url="https://doi.org/10.1093/zoolinnean/zlac001" volume="197" year="2023">Zaher et al. 2023</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="53EC97C6F590C834EE8742C5BE3DA559" pageId="0" pageNumber="717">
|
||
The fossil record of scolecophidians is poor, consisting exclusively of vertebrae spanning across a small number of Cenozoic localities (
|
||
<bibRefCitation id="32AA69D3215E61BD4E8FC7F92955FAC3" DOI="https://doi.org/10.3989/egeol.91473-4422" author="Szyndlar, Z" journalOrPublisher="Estudios Geologicos" pageId="0" pageNumber="717" pagination="103 - 126" refId="B608" refString="Szyndlar, Z, 1991a. A review of Neogene and Quaternary snakes of Central and Eastern Europe. Part I: Scolecophidia, Boidae, Colubrinae. Estudios Geologicos 47: 103 - 126, DOI: https://doi.org/10.3989/egeol.91473-4422" title="A review of Neogene and Quaternary snakes of Central and Eastern Europe. Part I: Scolecophidia, Boidae, Colubrinae." url="https://doi.org/10.3989/egeol.91473-4422" volume="47" year="1991 a">Szyndlar 1991a</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="7867ABABACE2D3FA727FB5AC4E4224FC" DOI="https://doi.org/10.1016/j.geobios.2012.10.016" author="Mead, JI" journalOrPublisher="Geobios" pageId="0" pageNumber="717" pagination="225 - 231" refId="B382" refString="Mead, JI, 2013. Scolecophidia (Serpentes) of the Late Oligocene and Early Miocene, North America, and a fossil history overview. Geobios 46: 225 - 231, DOI: https://doi.org/10.1016/j.geobios.2012.10.016" title="Scolecophidia (Serpentes) of the Late Oligocene and Early Miocene, North America, and a fossil history overview." url="https://doi.org/10.1016/j.geobios.2012.10.016" volume="46" year="2013">Mead 2013</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="A2DBE41DEB074CD1DDD459302CF45247" DOI="https://doi.org/10.1080/08912963.2016.1234619" author="Georgalis, GL" journalOrPublisher="Historical Biology" pageId="0" pageNumber="717" pagination="730 - 742" refId="B206" refString="Georgalis, GL, Villa, A, Delfino, M, 2017. Fossil lizards and snakes from Ano Metochi - A diverse squamate fauna from the latest Miocene of northern Greece. Historical Biology 29: 730 - 742, DOI: https://doi.org/10.1080/08912963.2016.1234619" title="Fossil lizards and snakes from Ano Metochi - A diverse squamate fauna from the latest Miocene of northern Greece." url="https://doi.org/10.1080/08912963.2016.1234619" volume="29" year="2017">Georgalis et al. 2017</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="01A36169ACA88847DBAE8275E51C584B" DOI="https://doi.org/10.1017/9781108938891.007" author="Ivanov, M" editor="Gower, D" journalOrPublisher="Cambridge University Press, Cambridge" pageId="0" pageNumber="717" pagination="85 - 110" refId="B297" refString="Ivanov, M, 2022. Miocene Snakes of Eurasia: A review of the evolution of snake communities. In: Gower, D, Zaher, H, Eds., The Origin and Early Evolution of Snakes. Cambridge University Press, Cambridge: 85 - 110, DOI: https://doi.org/10.1017/9781108938891.007" title="Miocene Snakes of Eurasia: A review of the evolution of snake communities." url="https://doi.org/10.1017/9781108938891.007" volumeTitle="The Origin and Early Evolution of Snakes." year="2022">Ivanov 2022</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="1F43B78C9C6A8B78439E7D763FD0C142" DOI="https://doi.org/10.1017/9781108938891.006" author="Smith, KT" editor="Gower, D" journalOrPublisher="Cambridge University Press, Cambridge" pageId="0" pageNumber="717" pagination="55 - 84" refId="B571" refString="Smith, KT, Georgalis, GL, 2022. The diversity and distribution of Palaeogene snakes - A review, with comments on vertebral sufficiency. In: Gower, D, Zaher, H, Eds., , The Origin and Early Evolution of Snakes. Cambridge University Press, Cambridge: 55 - 84, DOI: https://doi.org/10.1017/9781108938891.006" title="The diversity and distribution of Palaeogene snakes - A review, with comments on vertebral sufficiency." url="https://doi.org/10.1017/9781108938891.006" volumeTitle=", The Origin and Early Evolution of Snakes." year="2022">Smith and Georgalis 2022</bibRefCitation>
|
||
) and a potential Cretaceous occurrence (
|
||
<bibRefCitation id="2BA05CB77061D0BA77D54EA2E7B9942A" DOI="https://doi.org/10.1016/j.isci.2020.101834" author="Fachini, ST" journalOrPublisher="Cuadernos de Herpetologia" pageId="0" pageNumber="717" refId="B170" refString="Fachini, ST, Onary, S, Palci, A, Lee, MSY, Bronzati, M, Hsiou, AS, 2020. Cretaceous blind snake from Brazil fills major gap in snake evolution. iScience 23: 101834. https://doi.org/10.1016/j.isci.2020.101834" title="Cretaceous blind snake from Brazil fills major gap in snake evolution. iScience 23: 101834." url="https://doi.org/10.1016/j.isci.2020.101834" year="2020">Fachini et al. 2020</bibRefCitation>
|
||
; but see
|
||
<bibRefCitation id="3AABA0034A4695C6922CEC491C418402" DOI="https://doi.org/10.1017/9781108938891.005" author="Head, JJ" editor="Gower, D" journalOrPublisher="Cambridge University Press, Cambridge" pageId="0" pageNumber="717" pagination="26 - 54" refId="B248" refString="Head, JJ, Howard, AFC, Mueller, J, 2022. The first 80 million years of snake evolution: The Mesozoic fossil record of snakes and its implications for origin hypotheses, biogeography, and mass extinction. In: Gower, D, Zaher, H, Eds., The Origin and Early Evolution of Snakes. Cambridge University Press, Cambridge: 26 - 54, DOI: https://doi.org/10.1017/9781108938891.005" title="The first 80 million years of snake evolution: The Mesozoic fossil record of snakes and its implications for origin hypotheses, biogeography, and mass extinction." url="https://doi.org/10.1017/9781108938891.005" volumeTitle="The Origin and Early Evolution of Snakes." year="2022">Head et al. 2022</bibRefCitation>
|
||
for an alternative interpretation of the latter Cretaceous form). In the vast majority of cases, scolecophidian fossil vertebrae cannot be referred more precisely to the family level, and this is especially the case for the Paleogene and Neogene remains (
|
||
<bibRefCitation id="F94EF49B206684EF38AE9182017BFE36" DOI="https://doi.org/10.3989/egeol.91473-4422" author="Szyndlar, Z" journalOrPublisher="Estudios Geologicos" pageId="0" pageNumber="717" pagination="103 - 126" refId="B608" refString="Szyndlar, Z, 1991a. A review of Neogene and Quaternary snakes of Central and Eastern Europe. Part I: Scolecophidia, Boidae, Colubrinae. Estudios Geologicos 47: 103 - 126, DOI: https://doi.org/10.3989/egeol.91473-4422" title="A review of Neogene and Quaternary snakes of Central and Eastern Europe. Part I: Scolecophidia, Boidae, Colubrinae." url="https://doi.org/10.3989/egeol.91473-4422" volume="47" year="1991 a">Szyndlar 1991a</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="F1E4D1DE72A242C13D7EB11D6BBDA4B3" DOI="https://doi.org/10.1016/j.geobios.2012.10.016" author="Mead, JI" journalOrPublisher="Geobios" pageId="0" pageNumber="717" pagination="225 - 231" refId="B382" refString="Mead, JI, 2013. Scolecophidia (Serpentes) of the Late Oligocene and Early Miocene, North America, and a fossil history overview. Geobios 46: 225 - 231, DOI: https://doi.org/10.1016/j.geobios.2012.10.016" title="Scolecophidia (Serpentes) of the Late Oligocene and Early Miocene, North America, and a fossil history overview." url="https://doi.org/10.1016/j.geobios.2012.10.016" volume="46" year="2013">Mead 2013</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="98C53743E0178194015D81972B2E1C52" DOI="https://doi.org/10.1080/08912963.2016.1234619" author="Georgalis, GL" journalOrPublisher="Historical Biology" pageId="0" pageNumber="717" pagination="730 - 742" refId="B206" refString="Georgalis, GL, Villa, A, Delfino, M, 2017. Fossil lizards and snakes from Ano Metochi - A diverse squamate fauna from the latest Miocene of northern Greece. Historical Biology 29: 730 - 742, DOI: https://doi.org/10.1080/08912963.2016.1234619" title="Fossil lizards and snakes from Ano Metochi - A diverse squamate fauna from the latest Miocene of northern Greece." url="https://doi.org/10.1080/08912963.2016.1234619" volume="29" year="2017">Georgalis et al. 2017</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="20B438F86B4B1CED8B1F951668B6BC8C" DOI="https://doi.org/10.1017/9781108938891.007" author="Ivanov, M" editor="Gower, D" journalOrPublisher="Cambridge University Press, Cambridge" pageId="0" pageNumber="717" pagination="85 - 110" refId="B297" refString="Ivanov, M, 2022. Miocene Snakes of Eurasia: A review of the evolution of snake communities. In: Gower, D, Zaher, H, Eds., The Origin and Early Evolution of Snakes. Cambridge University Press, Cambridge: 85 - 110, DOI: https://doi.org/10.1017/9781108938891.007" title="Miocene Snakes of Eurasia: A review of the evolution of snake communities." url="https://doi.org/10.1017/9781108938891.007" volumeTitle="The Origin and Early Evolution of Snakes." year="2022">Ivanov 2022</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="8630D8835BF366B769CF72EA9B1499B0" DOI="https://doi.org/10.1017/9781108938891.006" author="Smith, KT" editor="Gower, D" journalOrPublisher="Cambridge University Press, Cambridge" pageId="0" pageNumber="717" pagination="55 - 84" refId="B571" refString="Smith, KT, Georgalis, GL, 2022. The diversity and distribution of Palaeogene snakes - A review, with comments on vertebral sufficiency. In: Gower, D, Zaher, H, Eds., , The Origin and Early Evolution of Snakes. Cambridge University Press, Cambridge: 55 - 84, DOI: https://doi.org/10.1017/9781108938891.006" title="The diversity and distribution of Palaeogene snakes - A review, with comments on vertebral sufficiency." url="https://doi.org/10.1017/9781108938891.006" volumeTitle=", The Origin and Early Evolution of Snakes." year="2022">Smith and Georgalis 2022</bibRefCitation>
|
||
). Nevertheless, some Quaternary fossil vertebrae have been identified to the family or even the genus level, aided mainly by a geographic rationale (e.g.,
|
||
<bibRefCitation id="57FFD2E2B6B240A7C25FB228F80F96FD" DOI="http://dx.doi.org/10.1016/j.crpv.2014.10.005" author="Bochaton, C" journalOrPublisher="Comptes Rendus Palevol" pageId="0" pageNumber="717" pagination="101 - 110" refId="B60" refString="Bochaton, C, Grouard, S, Cornette, R, Ineich, I, Lenoble, A, Tresset, A, Bailon, S, 2015. Fossil and subfossil herpetofauna from Cadet 2 Cave (Marie-Galante, Guadeloupe Islands, F. W. I.): Evolution of an insular herpetofauna since the Late Pleistocene. Comptes Rendus Palevol 14: 101 - 110, DOI: http://dx.doi.org/10.1016/j.crpv.2014.10.005" title="Fossil and subfossil herpetofauna from Cadet 2 Cave (Marie-Galante, Guadeloupe Islands, F. W. I.): Evolution of an insular herpetofauna since the Late Pleistocene." url="http://dx.doi.org/10.1016/j.crpv.2014.10.005" volume="14" year="2015">Bochaton et al. 2015</bibRefCitation>
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;
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<bibRefCitation id="8270FBF5346FD4645D10A73C0AFFAA67" DOI="https://doi.org/10.4202/app.00766.2020" author="Syromyatnikova, E" journalOrPublisher="Acta Palaeontologica Polonica" pageId="0" pageNumber="717" pagination="395 - 407" refId="B603" refString="Syromyatnikova, E, Aranda, E, Fiol Gonzalez, S, 2021. First insight into the diversity of snakes in the Pleistocene of Cuba. Acta Palaeontologica Polonica 66: 395 - 407, DOI: https://doi.org/10.4202/app.00766.2020" title="First insight into the diversity of snakes in the Pleistocene of Cuba." url="https://doi.org/10.4202/app.00766.2020" volume="66" year="2021">Syromyatnikova et al. 2021</bibRefCitation>
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;
|
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<bibRefCitation id="CA24352F7C0FBD3D220350A247233FE3" DOI="https://doi.org/10.1080/08912963.2023.2242369" author="Peralta, MJ" journalOrPublisher="Universita di Torino" pageId="0" pageNumber="717" refId="B438" refString="Peralta, MJ, Ferrero, BS, 2023. First Quaternary fossil record of a blind snake (Scolecophidia, Serpentes) from South America (Argentina). Historical Biology https://doi.org/10.1080/08912963.2023.2242369" title="First Quaternary fossil record of a blind snake (Scolecophidia, Serpentes) from South America (Argentina). Historical Biology" url="https://doi.org/10.1080/08912963.2023.2242369" year="2023">Peralta and Ferrero 2023</bibRefCitation>
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;
|
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<bibRefCitation id="B26E0BC522B43E803793D11C0186B684" DOI="https://doi.org/10.1016/j.geobios.2023.03.005" author="Ramello, G" journalOrPublisher="Cretaceous Research" pageId="0" pageNumber="717" refId="B491" refString="Ramello, G, Delfino, M, Mori, E, Viviano, A, Pavia, G, Carnevale, G, Pavia, M, 2023. Holocene vertebrate assemblages provide the first evidence for the presence of the barn owl (Tytonidae, Tyto alba) on Socotra Island (Yemen). Geobios https://doi.org/10.1016/j.geobios.2023.03.005" title="Holocene vertebrate assemblages provide the first evidence for the presence of the barn owl (Tytonidae, Tyto alba) on Socotra Island (Yemen). Geobios" url="https://doi.org/10.1016/j.geobios.2023.03.005" year="2023">Ramello et al. 2023</bibRefCitation>
|
||
). Despite this very poor fossil record, divergence date estimates suggest that scolecophidians have split from other snakes around the Early Cretaceous (
|
||
<bibRefCitation id="428590F49FE3B1CDE069700F3B6145C0" DOI="https://doi.org/10.1111/j.1558-5646.2011.01437.x" author="Pyron, RA" journalOrPublisher="Evolution" pageId="0" pageNumber="717" pagination="163 - 178" refId="B467" refString="Pyron, RA, Burbrink, FT, 2012. Extinction, ecological opportunity, and the origins of global snake diversity. Evolution 66: 163 - 178, DOI: https://doi.org/10.1111/j.1558-5646.2011.01437.x" title="Extinction, ecological opportunity, and the origins of global snake diversity." url="https://doi.org/10.1111/j.1558-5646.2011.01437.x" volume="66" year="2012">Pyron and Burbrink 2012</bibRefCitation>
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;
|
||
<bibRefCitation id="7C240D12F41EB0E84A71CAE30B4735A7" DOI="https://doi.org/10.1016/j.ympev.2015.10.009" author="Zheng, Y" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="717" pagination="537 - 547" refId="B696" refString="Zheng, Y, Wiens, JJ, 2016. Combining phylogenomic and supermatrix approaches, and a time-calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52 genes and 4162 species. Molecular Phylogenetics and Evolution 94: 537 - 547, DOI: https://doi.org/10.1016/j.ympev.2015.10.009" title="Combining phylogenomic and supermatrix approaches, and a time-calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52 genes and 4162 species." url="https://doi.org/10.1016/j.ympev.2015.10.009" volume="94" year="2016">Zheng and Wiens 2016</bibRefCitation>
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||
;
|
||
<bibRefCitation id="5D50C46139CE9EE8BB1E9C8A407C38F4" DOI="https://doi.org/10.1111/jeb.13373" author="Miralles, A" journalOrPublisher="Journal of Evolutionary Biology" pageId="0" pageNumber="717" pagination="1782 - 1793" refId="B387" refString="Miralles, A, Marin, J, Markus, D, Herrel, A, Hedges, BS, Vidal, N, 2018. Molecular evidence for the paraphyly of Scolecophidia and its evolutionary implications. Journal of Evolutionary Biology 31: 1782 - 1793, DOI: https://doi.org/10.1111/jeb.13373" title="Molecular evidence for the paraphyly of Scolecophidia and its evolutionary implications." url="https://doi.org/10.1111/jeb.13373" volume="31" year="2018">Miralles et al. 2018</bibRefCitation>
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;
|
||
<bibRefCitation id="F65C2304B9A21CD5E89B42941147E420" DOI="https://doi.org/10.1016/j.isci.2020.101834" author="Fachini, ST" journalOrPublisher="Cuadernos de Herpetologia" pageId="0" pageNumber="717" refId="B170" refString="Fachini, ST, Onary, S, Palci, A, Lee, MSY, Bronzati, M, Hsiou, AS, 2020. Cretaceous blind snake from Brazil fills major gap in snake evolution. iScience 23: 101834. https://doi.org/10.1016/j.isci.2020.101834" title="Cretaceous blind snake from Brazil fills major gap in snake evolution. iScience 23: 101834." url="https://doi.org/10.1016/j.isci.2020.101834" year="2020">Fachini et al. 2020</bibRefCitation>
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).
|
||
</paragraph>
|
||
<paragraph id="746F4B7D64B0A5F9DAE9F16B422FB101" pageId="0" pageNumber="717">
|
||
A considerable amount of skeletal studies in scolecophidians has focused primarily on their cranial anatomy already since the 19th century (e.g.,
|
||
<bibRefCitation id="001A317CC7F9088A62FB5B9F395ABA8E" author="Mueller, J" journalOrPublisher="Zeitschrift fuer Physiologie" pageId="0" pageNumber="717" pagination="190 - 275" refId="B394" refString="Mueller, J, 1831. Beitraege zur Anatomie und Naturgeschichte der Amphibien. Zeitschrift fuer Physiologie 4: 190 - 275" title="Beitraege zur Anatomie und Naturgeschichte der Amphibien." volume="4" year="1831">
|
||
<normalizedToken id="51F7DC7D639220F67BD85D02DFC68828" originalValue="Müller">Mueller</normalizedToken>
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||
1831
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="F6B92110A629AF6F114D6A5EDE9360CC" DOI="https://doi.org/10.5962/bhl.title.45246" author="Jan, G" journalOrPublisher="Bailliere et fils, Paris" pageId="0" pageNumber="717" refId="B307" refString="Jan, G, Sordelli, F, 1860-1866. Iconographie generale des ophidiens. Tome premier (livrais 1 a 17), contenant cent deux planches. Georges Jan & Ferdinand Sordelli, Milan. https://doi.org/10.5962/bhl.title.45246" title="Iconographie generale des ophidiens. Tome premier (livrais 1 a 17), contenant cent deux planches. Georges Jan & Ferdinand Sordelli, Milan." url="https://doi.org/10.5962/bhl.title.45246" year="1860 - 1866">Jan and Sordelli 1860-1866</bibRefCitation>
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;
|
||
<bibRefCitation id="54C7D06765A8BA3A98948786FDA55109" author="Boulenger, GΑ" journalOrPublisher="Trustees of the British Museum (Natural History), London" pageId="0" pageNumber="717" refId="B83" refString="Boulenger, GΑ, 1893. Catalogue of the Snakes in the British Museum (Natural History). Volume 1, Containing the Families Typhlopidae, Glauconidae, Boidae, Ilysiidae, Uropeltidae, Xenopeltidae, and Colubridae Aglyphae, part. Trustees of the British Museum (Natural History), London" title="Catalogue of the Snakes in the British Museum (Natural History). Volume 1, Containing the Families Typhlopidae, Glauconidae, Boidae, Ilysiidae, Uropeltidae, Xenopeltidae, and Colubridae Aglyphae, part." year="1893">Boulenger 1893</bibRefCitation>
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;
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<bibRefCitation id="1E4AE0AA6A63B0A9B47BE3DC1E0E02E4" author="Waite, ER" journalOrPublisher="Records of the South Australian Museum" pageId="0" pageNumber="717" pagination="1 - 38" refId="B659" refString="Waite, ER, 1918b. Review of the Australian blind snakes (Family Typhlopidae). Records of the South Australian Museum 1: 1 - 38" title="Review of the Australian blind snakes (Family Typhlopidae)." volume="1" year="1918 b">Waite 1918b</bibRefCitation>
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<bibRefCitation id="320500BC43D5373BFF8A166A683B96C7" DOI="https://doi.org/10.1007/BF03049353" author="Mahendra, BC" journalOrPublisher="Proceedings of the Indian Academy of Sciences" pageId="0" pageNumber="717" pagination="128 - 142" refId="B362" refString="Mahendra, BC, 1936. Contributions to the osteology of the Ophidia. I. The endoskeleton of the so-called "blind snake", Typhlops braminus Daud. Proceedings of the Indian Academy of Sciences 3: 128 - 142, DOI: https://doi.org/10.1007/BF03049353" title="Contributions to the osteology of the Ophidia. I. The endoskeleton of the so-called " blind snake ", Typhlops braminus Daud." url="https://doi.org/10.1007/BF03049353" volume="3" year="1936">Mahendra 1936</bibRefCitation>
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<bibRefCitation id="3B1ABACC51398AAF9EB56C053DD226CE" author="Dunn, ER" journalOrPublisher="Bulletin of the Museum of Comparative Zoology" pageId="0" pageNumber="717" pagination="509 - 526" refId="B160" refString="Dunn, ER, 1941. Notes on the snake genus Anomalepis. Bulletin of the Museum of Comparative Zoology 87: 509 - 526" title="Notes on the snake genus Anomalepis." volume="87" year="1941">Dunn 1941</bibRefCitation>
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<bibRefCitation id="643E54161EF8B07E4C2440799FBC074C" DOI="https://doi.org/10.1002/jmor.1050740203" author="Dunn, ER" journalOrPublisher="Journal of Morphology" pageId="0" pageNumber="717" pagination="287 - 294" refId="B164" refString="Dunn, ER, Tihen, JA, 1944. The skeletal anatomy of Liotyphlops albirostris. Journal of Morphology 74: 287 - 294, DOI: https://doi.org/10.1002/jmor.1050740203" title="The skeletal anatomy of Liotyphlops albirostris." url="https://doi.org/10.1002/jmor.1050740203" volume="74" year="1944">Dunn and Tihen 1944</bibRefCitation>
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<bibRefCitation id="06BF0C3949AD3AA21631C4E659EC2857" DOI="https://doi.org/10.5962/bhl.title.50341" author="List, JC" journalOrPublisher="Illinois Biological Monographs" pageId="0" pageNumber="717" pagination="1 - 112" refId="B352" refString="List, JC, 1966. Comparative osteology of the snake families Typhlopidae and Leptotyphlopidae. Illinois Biological Monographs 36: 1 - 112, DOI: https://doi.org/10.5962/bhl.title.50341" title="Comparative osteology of the snake families Typhlopidae and Leptotyphlopidae." url="https://doi.org/10.5962/bhl.title.50341" volume="36" year="1966">List 1966</bibRefCitation>
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<bibRefCitation id="CEE277FB245A35401AC3A942321CC61C" DOI="https://doi.org/10.1670/206-5.1" author="Wallach, V" journalOrPublisher="Journal of Herpetology" pageId="0" pageNumber="717" pagination="690 - 702" refId="B665" refString="Wallach, V, Brown, RM, Diesmos, AC, Gee, GVA, 2007. An enigmatic new species of blind snake from Luzon Island, Northern Philippines, with a synopsis of the genus Acutotyphlops (Serpentes: Typhlopidae). Journal of Herpetology 41: 690 - 702, DOI: https://doi.org/10.1670/206-5.1" title="An enigmatic new species of blind snake from Luzon Island, Northern Philippines, with a synopsis of the genus Acutotyphlops (Serpentes: Typhlopidae)." url="https://doi.org/10.1670/206-5.1" volume="41" year="2007">Wallach et al. 2007</bibRefCitation>
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<bibRefCitation id="B66DA96DB8AF54FBA263391A03F5D9E2" DOI="https://doi.org/10.11646/zootaxa.2255.1.1" author="Broadley, DG" journalOrPublisher="Zootaxa" pageId="0" pageNumber="717" pagination="1 - 100" refId="B96" refString="Broadley, DG, Wallach, V, 2009. A review of the eastern and southern African blind-snakes (Serpentes: Typhlopidae), excluding Letheobia Cope, with the description of two new genera and a new species. Zootaxa 2255: 1 - 100, DOI: https://doi.org/10.11646/zootaxa.2255.1.1" title="A review of the eastern and southern African blind-snakes (Serpentes: Typhlopidae), excluding Letheobia Cope, with the description of two new genera and a new species." url="https://doi.org/10.11646/zootaxa.2255.1.1" volume="2255" year="2009">Broadley and Wallach 2009</bibRefCitation>
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<bibRefCitation id="C55D55BEF96AC50F41CB9DEDCA6F49F0" DOI="https://doi.org/10.1002/jmor.10703" author="Rieppel, O" journalOrPublisher="Journal of Morphology" pageId="0" pageNumber="717" pagination="536 - 557" refId="B502" refString="Rieppel, O, Kley, NJ, Maisano, JA, 2009. Morphology of the skull of the white-nosed blindsnake, Liotyphlops albirostris (Scolecophidia: Anomalepididae). Journal of Morphology 270: 536 - 557, DOI: https://doi.org/10.1002/jmor.10703" title="Morphology of the skull of the white-nosed blindsnake, Liotyphlops albirostris (Scolecophidia: Anomalepididae)." url="https://doi.org/10.1002/jmor.10703" volume="270" year="2009">Rieppel et al. 2009</bibRefCitation>
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<bibRefCitation id="C01E2F332842048513C223D51345AAC7" DOI="https://doi.org/10.1002/ar.23191" author="Pinto, RR" journalOrPublisher="Anatomical Record" pageId="0" pageNumber="717" pagination="1722 - 1747" refId="B462" refString="Pinto, RR, Martins, AR, Curcio, F, Ramos, LO, 2015. Osteology and cartilaginous elements of Trilepida salgueiroi (Amaral, 1954) (Scolecophidia: Leptotyphopidae). Anatomical Record 298: 1722 - 1747, DOI: https://doi.org/10.1002/ar.23191" title="Osteology and cartilaginous elements of Trilepida salgueiroi (Amaral, 1954) (Scolecophidia: Leptotyphopidae)." url="https://doi.org/10.1002/ar.23191" volume="298" year="2015">Pinto et al. 2015</bibRefCitation>
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<bibRefCitation id="C9BF380C81E41A04B0818F513ED3806E" DOI="https://doi.org/10.11646/zootaxa.4299.1.3" author="Kraus, F" journalOrPublisher="Zootaxa" pageId="0" pageNumber="717" pagination="75 - 94" refId="B326" refString="Kraus, F, 2017. New species of blindsnakes (Squamata: Gerrhopilidae) from the offshore islands of Papua New Guinea. Zootaxa 4299: 75 - 94, DOI: https://doi.org/10.11646/zootaxa.4299.1.3" title="New species of blindsnakes (Squamata: Gerrhopilidae) from the offshore islands of Papua New Guinea." url="https://doi.org/10.11646/zootaxa.4299.1.3" volume="4299" year="2017">Kraus 2017</bibRefCitation>
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<bibRefCitation id="87FAE95E3BE9062542FFCD533BC752A8" DOI="https://doi.org/10.1111/joa.12952" author="Chretien, J" journalOrPublisher="Journal of Anatomy" pageId="0" pageNumber="717" pagination="637 - 655" refId="B115" refString="Chretien, J, Wang-Claypool, CY, Glaw, F, Scherz, MD, 2019. The bizarre skull of Xenotyphlops sheds light on synapomorphies of Typhlopoidea. Journal of Anatomy 234: 637 - 655, DOI: https://doi.org/10.1111/joa.12952" title="The bizarre skull of Xenotyphlops sheds light on synapomorphies of Typhlopoidea." url="https://doi.org/10.1111/joa.12952" volume="234" year="2019">Chretien et al. 2019</bibRefCitation>
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<bibRefCitation id="8617A716C1D2C44CFFD988FB23669368" DOI="https://doi.org/10.1643/CH-19-181" author="Marra Santos, FJ" journalOrPublisher="Copeia" pageId="0" pageNumber="717" pagination="239 - 243" refId="B366" refString="Marra Santos, FJ, Reis, RE, 2019. Redescription of the blind snake Anomalepis colombia (Serpentes: Anomalepididae) using high-resolution X-ray computed tomography. Copeia 2019: 239 - 243, DOI: https://doi.org/10.1643/CH-19-181" title="Redescription of the blind snake Anomalepis colombia (Serpentes: Anomalepididae) using high-resolution X-ray computed tomography." url="https://doi.org/10.1643/CH-19-181" volume="2019" year="2019">Marra Santos and Reis 2019</bibRefCitation>
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<bibRefCitation id="6AE920B0E7725A5E4CB07CA2385D4AD4" DOI="https://doi.org/10.1002/ar.24730" author="Linares-Vargas, CA" journalOrPublisher="Anatomical Record" pageId="0" pageNumber="717" pagination="2264 - 2278" refId="B348" refString="Linares-Vargas, CA, Bolivar-Garcia, W, Herrera-Martinez, A, Osorio-Dominguez, D, Ospina, OE, Thomas, R, Daza, JD, 2021. The status of the anomalepidid snake Liotyphlops albirostris and the revalidation of three taxa based on morphology and ecological niche models. Anatomical Record 304: 2264 - 2278, DOI: https://doi.org/10.1002/ar.24730" title="The status of the anomalepidid snake Liotyphlops albirostris and the revalidation of three taxa based on morphology and ecological niche models." url="https://doi.org/10.1002/ar.24730" volume="304" year="2021">Linares-Vargas et al. 2021</bibRefCitation>
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<bibRefCitation id="1584EC6878576FEC1A3EA6094F57B8A0" DOI="https://doi.org/10.1002/ar.24591" author="Lira, I" journalOrPublisher="Anatomical Record" pageId="0" pageNumber="717" pagination="2198 - 2214" refId="B351" refString="Lira, I, Martins, A, 2021. Digging into blindsnakes' morphology: Description of the skull, lower jaw, and cervical vertebrae of two Amerotyphlops (Hedges et al., 2014) (Serpentes, Typhlopidae) with comments on the typhlopoidean skull morphological diversity. Anatomical Record 304: 2198 - 2214, DOI: https://doi.org/10.1002/ar.24591" title="Digging into blindsnakes' morphology: Description of the skull, lower jaw, and cervical vertebrae of two Amerotyphlops (Hedges et al., 2014) (Serpentes, Typhlopidae) with comments on the typhlopoidean skull morphological diversity." url="https://doi.org/10.1002/ar.24591" volume="304" year="2021">Lira and Martins 2021</bibRefCitation>
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;
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<bibRefCitation id="DB60E0D9250EA5994E70DFE095D69EA9" DOI="https://doi.org/10.1093/zoolinnean/zlac050" author="Chuliver, M" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="0" pageNumber="717" pagination="698 - 718" refId="B116" refString="Chuliver, M, Scanferla, A, Koch, C, 2023. Ontogeny of the skull of the blind snake Amerotyphlops brongersmianus (Serpentes: Typhlopidae) brings new insights on snake cranial evolution. Zoological Journal of the Linnean Society 197: 698 - 718, DOI: https://doi.org/10.1093/zoolinnean/zlac050" title="Ontogeny of the skull of the blind snake Amerotyphlops brongersmianus (Serpentes: Typhlopidae) brings new insights on snake cranial evolution." url="https://doi.org/10.1093/zoolinnean/zlac050" volume="197" year="2023">Chuliver et al. 2023</bibRefCitation>
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;
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<bibRefCitation id="F1FD6F5C505B6DD49F4CC4F80105B507" DOI="https://doi.org/10.1093/zoolinnean/zlac059" author="Graboski, R" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="0" pageNumber="717" pagination="719 - 751" refId="B218" refString="Graboski, R, Arredondo, JC, Grazziotin, FG, Guerra-Fuentes, RA, da Silva, AAA, Prudente, AC, Pinto, RR, Rodrigues, MT, Bonatto, SL, Zaher, H, 2023. Revealing the cryptic diversity of the widespread and poorly known South American blind snake genus Amerotyphlops (Typhlopidae: Scolecophidia) through integrative taxonomy. Zoological Journal of the Linnean Society 197: 719 - 751, DOI: https://doi.org/10.1093/zoolinnean/zlac059" title="Revealing the cryptic diversity of the widespread and poorly known South American blind snake genus Amerotyphlops (Typhlopidae: Scolecophidia) through integrative taxonomy." url="https://doi.org/10.1093/zoolinnean/zlac059" volume="197" year="2023">Graboski et al. 2023</bibRefCitation>
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;
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<bibRefCitation id="855C879EA7C37838EA732BCE6B7CDBFE" DOI="https://doi.org/10.3897/vz.73.e98170" author="Martins, A" journalOrPublisher="Vertebrate Zoology" pageId="0" pageNumber="717" pagination="345 - 366" refId="B368" refString="Martins, A, Folly, M, Ferreira, GN, Garcia da Silva, AS, Koch, C, Fouquet, A, Machado, A, Lopes, RT, Pinto, R, Rodrigues, MT, Passos, P, 2023. An evolutionary paradox on threadsnakes: Phenotypic and molecular evidence reveal a new and remarkably polymorphic species of Siagonodon (Serpentes: Leptotyphlopidae: Epictinae) from Amazonia. Vertebrate Zoology 73: 345 - 366, DOI: https://doi.org/10.3897/vz.73.e98170" title="An evolutionary paradox on threadsnakes: Phenotypic and molecular evidence reveal a new and remarkably polymorphic species of Siagonodon (Serpentes: Leptotyphlopidae: Epictinae) from Amazonia." url="https://doi.org/10.3897/vz.73.e98170" volume="73" year="2023">Martins et al. 2023</bibRefCitation>
|
||
) revealing indeed important differences among the different families. In contrast, their vertebral morphology has received very little attention (e.g.,
|
||
<bibRefCitation id="1BC15B34C3FDD928E87CFB4A8E37FC8E" DOI="https://doi.org/10.1111/j.1096-3642.1933.tb01595.x" author="Mookerjee, HK" journalOrPublisher="Proceedings of the Zoological Society of London" pageId="0" pageNumber="717" refId="B391" refString="Mookerjee, HK, Das, GM, 1933. On the peculiar apertures in the vertebral centra of Typhlops braminus. Proceedings of the Zoological Society of London, Part I: 283. https://doi.org/10.1111/j.1096-3642.1933.tb01595.x" title="On the peculiar apertures in the vertebral centra of Typhlops braminus. Proceedings of the Zoological Society of London, Part I: 283." url="https://doi.org/10.1111/j.1096-3642.1933.tb01595.x" year="1933">Mookerjee and Das 1933</bibRefCitation>
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;
|
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<bibRefCitation id="AE9AAE2A472AF767168A93D08E322C8A" DOI="https://doi.org/10.1007/BF03049353" author="Mahendra, BC" journalOrPublisher="Proceedings of the Indian Academy of Sciences" pageId="0" pageNumber="717" pagination="128 - 142" refId="B362" refString="Mahendra, BC, 1936. Contributions to the osteology of the Ophidia. I. The endoskeleton of the so-called "blind snake", Typhlops braminus Daud. Proceedings of the Indian Academy of Sciences 3: 128 - 142, DOI: https://doi.org/10.1007/BF03049353" title="Contributions to the osteology of the Ophidia. I. The endoskeleton of the so-called " blind snake ", Typhlops braminus Daud." url="https://doi.org/10.1007/BF03049353" volume="3" year="1936">Mahendra 1936</bibRefCitation>
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;
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<bibRefCitation id="4F8D3FD90FE4F1DBD3C48A17CD66976E" DOI="https://doi.org/10.5962/bhl.title.50341" author="List, JC" journalOrPublisher="Illinois Biological Monographs" pageId="0" pageNumber="717" pagination="1 - 112" refId="B352" refString="List, JC, 1966. Comparative osteology of the snake families Typhlopidae and Leptotyphlopidae. Illinois Biological Monographs 36: 1 - 112, DOI: https://doi.org/10.5962/bhl.title.50341" title="Comparative osteology of the snake families Typhlopidae and Leptotyphlopidae." url="https://doi.org/10.5962/bhl.title.50341" volume="36" year="1966">List 1966</bibRefCitation>
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;
|
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<bibRefCitation id="47F93D7A218B9FB0FBA2FB4C0B28E15A" author="Fabrezi, M" journalOrPublisher="Cuadernos de Herpetologia" pageId="0" pageNumber="717" pagination="1 - 20" refId="B169" refString="Fabrezi, M, Marcus, A, Scrocchi, G, 1985. Contribucion al conocimento de los Leptotyphlopidae de Argentina I. Leptotyphlops weyrauchi y Leptotyphlops albipuncta. Cuadernos de Herpetologia 1: 1 - 20" title="Contribucion al conocimento de los Leptotyphlopidae de Argentina I. Leptotyphlops weyrauchi y Leptotyphlops albipuncta." volume="1" year="1985">Fabrezi et al. 1985</bibRefCitation>
|
||
), though there seems to be growing interest recently, substantially aided by the usage of non-invasive technologies, such as
|
||
<normalizedToken id="BC11E68EAAAF4AB5D005E94D5B5D33EF" originalValue="μCT">μCT</normalizedToken>
|
||
scanning (e.g.,
|
||
<bibRefCitation id="60749E3FD35E8584B9305480A51835FD" DOI="https://doi.org/10.1111/jzs.12316" author="Martins, AR" journalOrPublisher="Journal of Zoological Systematics and Evolutionary Research" pageId="0" pageNumber="717" pagination="840 - 863" refId="B371" refString="Martins, AR, Koch, C, Pinto, R, Folly, M, Fouquet, A, Passos, P, 2019. From the inside out: Discovery of a new genus of threadsnakes based on anatomical and molecular data, with discussion of the leptotyphlopid hemipenial morphology. Journal of Zoological Systematics and Evolutionary Research 57: 840 - 863, DOI: https://doi.org/10.1111/jzs.12316" title="From the inside out: Discovery of a new genus of threadsnakes based on anatomical and molecular data, with discussion of the leptotyphlopid hemipenial morphology." url="https://doi.org/10.1111/jzs.12316" volume="57" year="2019">Martins et al. 2019</bibRefCitation>
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,
|
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<bibRefCitation id="6512BEF4B1441A5AEAB40170F725F93F" DOI="https://doi.org/10.1002/ar.24716" author="Martins, A" journalOrPublisher="Anatomical Record" pageId="0" pageNumber="717" pagination="2118 - 2148" refId="B369" refString="Martins, A, Koch, C, Joshi, M, Pinto, R, Machado, A, Lopes, R, Passos, P, 2021a. Evolutionary treasures hidden in the West Indies: Comparative osteology and visceral morphology reveals intricate miniaturization in the insular genera Mitophis Hedges, Adalsteinsson, & Branch, 2009 and Tetracheilostoma Jan, 1861 (Leptotyphlopidae: Epictinae: Tetracheilostomina). Anatomical Record 304: 2118 - 2148, DOI: https://doi.org/10.1002/ar.24716" title="Evolutionary treasures hidden in the West Indies: Comparative osteology and visceral morphology reveals intricate miniaturization in the insular genera Mitophis Hedges, Adalsteinsson, & Branch, 2009 and Tetracheilostoma Jan, 1861 (Leptotyphlopidae: Epictinae: Tetracheilostomina)." url="https://doi.org/10.1002/ar.24716" volume="304" year="2021 a">2021a</bibRefCitation>
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,
|
||
<bibRefCitation id="0A7891673A567F7B306719096E561C01" DOI="https://doi.org/10.1002/ar.24747" author="Martins, A" journalOrPublisher="Anatomical Record" pageId="0" pageNumber="717" pagination="2149 - 2182" refId="B370" refString="Martins, A, Koch, C, Joshi, M, Pinto, R, Passos, P, 2021b. Picking up the threads: Comparative osteology and associated cartilaginous elements for members of the genus Trilepida Hedges, 2011 (Serpentes, Leptotyphlopidae) with new insights on the Epictinae systematics. Anatomical Record 304: 2149 - 2182, DOI: https://doi.org/10.1002/ar.24747" title="Picking up the threads: Comparative osteology and associated cartilaginous elements for members of the genus Trilepida Hedges, 2011 (Serpentes, Leptotyphlopidae) with new insights on the Epictinae systematics." url="https://doi.org/10.1002/ar.24747" volume="304" year="2021 b">2021b</bibRefCitation>
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;
|
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<bibRefCitation id="0F0E3A1B528B5E84AC22C85CDF5F6B47" DOI="https://doi.org/10.1002/ar.24686" author="Herrel, A" journalOrPublisher="Anatomical Record" pageId="0" pageNumber="717" pagination="2292 - 2302" refId="B260" refString="Herrel, A, Lowie, A, Miralles, A, Gaucher, P, Kley, NJ, Measey, J, Tolley, KA, 2021. Burrowing in blindsnakes: A preliminary analysis of burrowing forces and consequences for the evolution of morphology. Anatomical Record 304: 2292 - 2302, DOI: https://doi.org/10.1002/ar.24686" title="Burrowing in blindsnakes: A preliminary analysis of burrowing forces and consequences for the evolution of morphology." url="https://doi.org/10.1002/ar.24686" volume="304" year="2021">Herrel et al. 2021</bibRefCitation>
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;
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<bibRefCitation id="D2D3050ACE1BF93C8B2BEECC172CCC8D" DOI="https://doi.org/10.7717/peerj.7411" author="Koch, C" journalOrPublisher="Anatomical Record" pageId="0" pageNumber="717" refId="B324" refString="Koch, C, Martins, A, Schweiger, S, 2019. A century of waiting: Description of a new Epictia Gray, 1845 (Serpentes: Leptotyphlopidae) based on specimens housed for more than 100 years in the collection of the Natural History Museum Vienna (NMW). PeerJ 7: e7411. https://doi.org/10.7717/peerj.7411" title="A century of waiting: Description of a new Epictia Gray, 1845 (Serpentes: Leptotyphlopidae) based on specimens housed for more than 100 years in the collection of the Natural History Museum Vienna (NMW). PeerJ 7: e 7411." url="https://doi.org/10.7717/peerj.7411" year="2019">Koch et al. 2019</bibRefCitation>
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,
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||
<bibRefCitation id="A7A23070F67C9709AE9518AD68ED3E3A" DOI="https://doi.org/10.1002/ar.24676" author="Koch, C" journalOrPublisher="Anatomical Record" pageId="0" pageNumber="717" pagination="2183 - 2197" refId="B323" refString="Koch, C, Martins, A, Joshi, M, Pinto, RR, Passos, P, 2021. Osteology of the enigmatic threadsnake species Epictia unicolor and Trilepida guayaquilensis (Serpentes, Leptotyphlopidae) with generic insights. Anatomical Record 304: 2183 - 2197, DOI: https://doi.org/10.1002/ar.24676" title="Osteology of the enigmatic threadsnake species Epictia unicolor and Trilepida guayaquilensis (Serpentes, Leptotyphlopidae) with generic insights." url="https://doi.org/10.1002/ar.24676" volume="304" year="2021">2021</bibRefCitation>
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||
;
|
||
<bibRefCitation id="CAEF2AEB49FF640729CCC7A2324A9F5A" DOI="https://doi.org/10.1002/ar.24591" author="Lira, I" journalOrPublisher="Anatomical Record" pageId="0" pageNumber="717" pagination="2198 - 2214" refId="B351" refString="Lira, I, Martins, A, 2021. Digging into blindsnakes' morphology: Description of the skull, lower jaw, and cervical vertebrae of two Amerotyphlops (Hedges et al., 2014) (Serpentes, Typhlopidae) with comments on the typhlopoidean skull morphological diversity. Anatomical Record 304: 2198 - 2214, DOI: https://doi.org/10.1002/ar.24591" title="Digging into blindsnakes' morphology: Description of the skull, lower jaw, and cervical vertebrae of two Amerotyphlops (Hedges et al., 2014) (Serpentes, Typhlopidae) with comments on the typhlopoidean skull morphological diversity." url="https://doi.org/10.1002/ar.24591" volume="304" year="2021">Lira and Martins 2021</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="5147886E00858626DA47C271EBCDB618" pageId="0" pageNumber="717">All scolecophidians (leptotyphlopids, typhlopoids, and anomalepidids) seem to display a very simple and relatively homogenous vertebral morphology, which at most times renders it almost impossible to differentiate members of particular families based on postcranial osteology. They are all characterized by an elongate centrum, depressed cotyle and condyle, depressed neural arch, presence of relatively long prezygapophyseal accessory processes, the direction of the major axis of the prezygapophyseal articular facets approximating the direction of the major axis of the vertebra, absent or very shallow median notch of the neural arch, absent haemal keels in mid- and posterior trunk vertebrae, vestigial neural spine (present only in the anterior trunk vertebrae) shifted posteriorly, absence of any subcentral structures in the cloacal and caudal portion of the column, and very low number of caudal vertebrae.</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="4B1A1FC410C7A4CD0E87F81F60ACA8AC" pageId="0" pageNumber="717" type="vertebral distinction among scolecophidian families">
|
||
<paragraph id="11A478AF8B205FE70DBF5C7A1678209F" pageId="0" pageNumber="717">Vertebral distinction among scolecophidian families.</paragraph>
|
||
<paragraph id="F9EAF0DE1FEE3B5B54E6067BC25F9CB6" pageId="0" pageNumber="717">Despite the morphological homogeneity of scolecophidian families, certain features have been addressed in the literature to distinguish them or of possessing potential diagnostic utility for family level determination. These are summarized and assessed in this section.</paragraph>
|
||
<paragraph id="A32EC43890DD54518907C315E70E09CF" pageId="0" pageNumber="717">
|
||
<bibRefCitation id="B552734E9DC1CDC8720B415E5B549786" DOI="https://doi.org/10.1111/j.1096-3642.1933.tb01595.x" author="Mookerjee, HK" journalOrPublisher="Proceedings of the Zoological Society of London" pageId="0" pageNumber="717" refId="B391" refString="Mookerjee, HK, Das, GM, 1933. On the peculiar apertures in the vertebral centra of Typhlops braminus. Proceedings of the Zoological Society of London, Part I: 283. https://doi.org/10.1111/j.1096-3642.1933.tb01595.x" title="On the peculiar apertures in the vertebral centra of Typhlops braminus. Proceedings of the Zoological Society of London, Part I: 283." url="https://doi.org/10.1111/j.1096-3642.1933.tb01595.x" year="1933">Mookerjee and Das (1933)</bibRefCitation>
|
||
and
|
||
<bibRefCitation id="2AF8D8DFB6CC4257CC55F72C8719E178" author="Mahendra, BC" journalOrPublisher="Current Science" pageId="0" pageNumber="717" pagination="320 - 322" refId="B361" refString="Mahendra, BC, 1935. The sub-central foramina of the Squamata. Current Science 4: 320 - 322" title="The sub-central foramina of the Squamata." volume="4" year="1935">Mahendra (1935)</bibRefCitation>
|
||
highlighted the presence of a single subcentral foramen in trunk vertebrae of typhlopids, with the latter author regarding it as unique among extant snakes. Since then, the diagnostic utility of the subcentral foramen in typhlopids has been repeatedly highlighted (
|
||
<bibRefCitation id="32739DCC5F4AB12D62EFDF8F994EF26D" DOI="https://doi.org/10.5962/bhl.title.50341" author="List, JC" journalOrPublisher="Illinois Biological Monographs" pageId="0" pageNumber="717" pagination="1 - 112" refId="B352" refString="List, JC, 1966. Comparative osteology of the snake families Typhlopidae and Leptotyphlopidae. Illinois Biological Monographs 36: 1 - 112, DOI: https://doi.org/10.5962/bhl.title.50341" title="Comparative osteology of the snake families Typhlopidae and Leptotyphlopidae." url="https://doi.org/10.5962/bhl.title.50341" volume="36" year="1966">List 1966</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="47BDA0DEB52D4C95AF203EC4D6E05AC8" DOI="https://doi.org/10.1017/S1464793102005924" author="Lee, M" journalOrPublisher="Biological Reviews" pageId="0" pageNumber="717" pagination="333 - 402" refId="B346" refString="Lee, M, Scanlon, JD, 2002. Snake phylogeny based on osteology, soft anatomy and behaviour. Biological Reviews 77: 333 - 402, DOI: https://doi.org/10.1017/S1464793102005924" title="Snake phylogeny based on osteology, soft anatomy and behaviour." url="https://doi.org/10.1017/S1464793102005924" volume="77" year="2002">Lee and Scanlon 2002</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="6919563E0310489F2D9CA98667DC8FF9" author="Wallach, V" journalOrPublisher="Hamadryad" pageId="0" pageNumber="717" pagination="34 - 61" refId="B663" refString="Wallach, V, 2009. Ramphotyphlops braminus (Daudin): A synopsis of morphology, taxonomy, nomenclature and distribution (Serpentes: Typhlopidae). Hamadryad 34: 34 - 61" title="Ramphotyphlops braminus (Daudin): A synopsis of morphology, taxonomy, nomenclature and distribution (Serpentes: Typhlopidae)." volume="34" year="2009">Wallach 2009</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="2F1318899B8148C0B4B8A7E1B54A0EE2" author="Wallach, V" journalOrPublisher="Podarcis" pageId="0" pageNumber="717" pagination="4 - 12" refId="B664" refString="Wallach, V, 2020. How to easily identify the flowerpot blindsnake, Indotyphlops braminus (Daudin, 1803), with proposal of a new genus (Serpentes: Typhlopidae). Podarcis 11: 4 - 12" title="How to easily identify the flowerpot blindsnake, Indotyphlops braminus (Daudin, 1803), with proposal of a new genus (Serpentes: Typhlopidae)." volume="11" year="2020">2020</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="1FF2E3D67DAF53665A506DE47F6B4761" DOI="https://doi.org/10.1016/j.isci.2020.101834" author="Fachini, ST" journalOrPublisher="Cuadernos de Herpetologia" pageId="0" pageNumber="717" refId="B170" refString="Fachini, ST, Onary, S, Palci, A, Lee, MSY, Bronzati, M, Hsiou, AS, 2020. Cretaceous blind snake from Brazil fills major gap in snake evolution. iScience 23: 101834. https://doi.org/10.1016/j.isci.2020.101834" title="Cretaceous blind snake from Brazil fills major gap in snake evolution. iScience 23: 101834." url="https://doi.org/10.1016/j.isci.2020.101834" year="2020">Fachini et al. 2020</bibRefCitation>
|
||
).
|
||
<bibRefCitation id="CDE3F9216CE242EFE3414ED451602B6F" author="Wallach, V" journalOrPublisher="Podarcis" pageId="0" pageNumber="717" pagination="4 - 12" refId="B664" refString="Wallach, V, 2020. How to easily identify the flowerpot blindsnake, Indotyphlops braminus (Daudin, 1803), with proposal of a new genus (Serpentes: Typhlopidae). Podarcis 11: 4 - 12" title="How to easily identify the flowerpot blindsnake, Indotyphlops braminus (Daudin, 1803), with proposal of a new genus (Serpentes: Typhlopidae)." volume="11" year="2020">Wallach (2020)</bibRefCitation>
|
||
even suggested that this foramen was unusual among typhlopids and could be used as a characteristic of
|
||
<taxonomicName id="210309D5D1BD8D98A23008E0125C3642" baseAuthorityName="Daudin" baseAuthorityYear="1803" class="Reptilia" family="Typhlopidae" genus="Indotyphlops" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Indotyphlops braminus" order="Squamata" pageId="0" pageNumber="717" phylum="Chordata" rank="species" species="braminus">
|
||
<emphasis id="2E7276E1353C838049A2C1CCDC6F5886" italics="true" pageId="0" pageNumber="717">Indotyphlops braminus</emphasis>
|
||
</taxonomicName>
|
||
(Daudin, 1803).
|
||
<bibRefCitation id="6F898A3DDF26621FA3742AC1382C0E45" DOI="https://doi.org/10.1016/j.isci.2020.101834" author="Fachini, ST" journalOrPublisher="Cuadernos de Herpetologia" pageId="0" pageNumber="717" refId="B170" refString="Fachini, ST, Onary, S, Palci, A, Lee, MSY, Bronzati, M, Hsiou, AS, 2020. Cretaceous blind snake from Brazil fills major gap in snake evolution. iScience 23: 101834. https://doi.org/10.1016/j.isci.2020.101834" title="Cretaceous blind snake from Brazil fills major gap in snake evolution. iScience 23: 101834." url="https://doi.org/10.1016/j.isci.2020.101834" year="2020">Fachini et al. (2020)</bibRefCitation>
|
||
regarded "the presence of asymmetrical subcentral foramina" as diagnostic of scolecophidians. We acknowledge the presence of such single large subcentral foramen in typhlopids, which, as it can be seen in the figures, it is not constantly present in all vertebrae and in all species. In our leptotyphlopid sample, this feature was not observed, but, judging from the published literature, we have to note that these foramina can be present in all trunk and cloacal vertebrae of certain leptotyphlopid taxa (e.g.,
|
||
<taxonomicName id="49F981B99F602471C92708542950BCE1" authorityName="Hedges" authorityYear="2011" class="Amphibia" family="Leptotyphlopidae" genus="Trilepida" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trilepida" order="Squamata" pageId="0" pageNumber="717" phylum="Chordata" rank="genus">
|
||
<emphasis id="AC7E56CD1998FF7E2896AEFBEF3A9779" italics="true" pageId="0" pageNumber="717">Trilepida</emphasis>
|
||
</taxonomicName>
|
||
Hedges, 2011; see
|
||
<bibRefCitation id="1780A410A6E36A8DAEC22A719B980CD2" DOI="https://doi.org/10.1002/ar.23191" author="Pinto, RR" journalOrPublisher="Anatomical Record" pageId="0" pageNumber="717" pagination="1722 - 1747" refId="B462" refString="Pinto, RR, Martins, AR, Curcio, F, Ramos, LO, 2015. Osteology and cartilaginous elements of Trilepida salgueiroi (Amaral, 1954) (Scolecophidia: Leptotyphopidae). Anatomical Record 298: 1722 - 1747, DOI: https://doi.org/10.1002/ar.23191" title="Osteology and cartilaginous elements of Trilepida salgueiroi (Amaral, 1954) (Scolecophidia: Leptotyphopidae)." url="https://doi.org/10.1002/ar.23191" volume="298" year="2015">Pinto et al. 2015</bibRefCitation>
|
||
) as well as in anomalepidids, where they are also intracolumnarily variable (i.e., present in the anterior trunk vertebrae of
|
||
<taxonomicName id="F5B8FF51234F2CD84D967251A38474E6" class="Reptilia" family="Anomalepididae" genus="Anomalepis" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Anomalepis" order="Squamata" pageId="0" pageNumber="717" phylum="Chordata" rank="genus">
|
||
<emphasis id="53E9179CA0DF96475900A1F1CA1F5912" italics="true" pageId="0" pageNumber="717">Anomalepis</emphasis>
|
||
</taxonomicName>
|
||
Jan, 1860 in
|
||
<bibRefCitation id="9E39B7D6E799C30533FEFB860620B745" DOI="https://doi.org/10.5962/bhl.title.45246" author="Jan, G" journalOrPublisher="Bailliere et fils, Paris" pageId="0" pageNumber="717" refId="B307" refString="Jan, G, Sordelli, F, 1860-1866. Iconographie generale des ophidiens. Tome premier (livrais 1 a 17), contenant cent deux planches. Georges Jan & Ferdinand Sordelli, Milan. https://doi.org/10.5962/bhl.title.45246" title="Iconographie generale des ophidiens. Tome premier (livrais 1 a 17), contenant cent deux planches. Georges Jan & Ferdinand Sordelli, Milan." url="https://doi.org/10.5962/bhl.title.45246" year="1860 - 1866">Jan and Sordelli 1860-1866</bibRefCitation>
|
||
but absent posteriorly in this taxon [
|
||
<bibRefCitation id="6B63B1D55E2A79122367ADD0014FBD77" author="Dunn, ER" journalOrPublisher="Bulletin of the Museum of Comparative Zoology" pageId="0" pageNumber="717" pagination="509 - 526" refId="B160" refString="Dunn, ER, 1941. Notes on the snake genus Anomalepis. Bulletin of the Museum of Comparative Zoology 87: 509 - 526" title="Notes on the snake genus Anomalepis." volume="87" year="1941">Dunn 1941</bibRefCitation>
|
||
] and in the posterior trunk vertebrae of
|
||
<taxonomicName id="FE1E0D99E380FB6D110E3C948650BA35" class="Reptilia" family="Anomalepididae" genus="Liotyphlops" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Liotyphlops" order="Squamata" pageId="0" pageNumber="717" phylum="Chordata" rank="genus">
|
||
<emphasis id="EC6F83889B403071F2260B95B4B30CFC" italics="true" pageId="0" pageNumber="717">Liotyphlops</emphasis>
|
||
</taxonomicName>
|
||
Peters, 1881 [
|
||
<bibRefCitation id="60CE80AF00F7955AD5540D576AD3D91F" DOI="https://doi.org/10.1002/jmor.1050740203" author="Dunn, ER" journalOrPublisher="Journal of Morphology" pageId="0" pageNumber="717" pagination="287 - 294" refId="B164" refString="Dunn, ER, Tihen, JA, 1944. The skeletal anatomy of Liotyphlops albirostris. Journal of Morphology 74: 287 - 294, DOI: https://doi.org/10.1002/jmor.1050740203" title="The skeletal anatomy of Liotyphlops albirostris." url="https://doi.org/10.1002/jmor.1050740203" volume="74" year="1944">Dunn and Tihen 1944</bibRefCitation>
|
||
]). It should be noted also that such foramina can be variably present in aniliids, uropeltids, and calabariids as well (see the respective entries below). Certain fossorial lizards also possess distinctive subcentral foramina in their vertebrae (e.g., amphisbaenians, see
|
||
<bibRefCitation id="18A9E8074D57CDD613E351CA9BD08B64" DOI="https://doi.org/10.1016/j.annpal.2018.03.002" author="Georgalis, GL" journalOrPublisher="Annales de Paleontologie" pageId="0" pageNumber="717" pagination="155 - 159" refId="B207" refString="Georgalis, GL, Villa, A, Delfino, M, 2018. The last amphisbaenian (Squamata) from continental Eastern Europe. Annales de Paleontologie 104: 155 - 159, DOI: https://doi.org/10.1016/j.annpal.2018.03.002" title="The last amphisbaenian (Squamata) from continental Eastern Europe." url="https://doi.org/10.1016/j.annpal.2018.03.002" volume="104" year="2018">Georgalis et al. 2018</bibRefCitation>
|
||
and
|
||
<bibRefCitation id="28CAC0165D59547B173339EC4AE7CE43" DOI="https://doi.org/10.1126/sciadv.aat5042" author="Xing, L" journalOrPublisher="Proceedings of the Zoological Society of London" pageId="0" pageNumber="717" refId="B686" refString="Xing, L, Caldwell, MW, Chen, R, Nydam, RL, Palci, A, Simoes, TR, McKellar, RC, Lee, MSY, Liu, Y, Shi, H, Wang, K, Bai, M, 2018. A mid-Cretaceous embryonic-to-neonate snake in amber from Myanmar. Science Advances 4: eaat5042. https://doi.org/10.1126/sciadv.aat5042" title="A mid-Cretaceous embryonic-to-neonate snake in amber from Myanmar. Science Advances 4: eaat 5042." url="https://doi.org/10.1126/sciadv.aat5042" year="2018">Xing et al. 2018</bibRefCitation>
|
||
; dibamids, see figs in
|
||
<bibRefCitation id="68F11EE557E7C7D7D757AD751C789E41" DOI="https://doi.org/10.1126/sciadv.aat5042" author="Xing, L" journalOrPublisher="Proceedings of the Zoological Society of London" pageId="0" pageNumber="717" refId="B686" refString="Xing, L, Caldwell, MW, Chen, R, Nydam, RL, Palci, A, Simoes, TR, McKellar, RC, Lee, MSY, Liu, Y, Shi, H, Wang, K, Bai, M, 2018. A mid-Cretaceous embryonic-to-neonate snake in amber from Myanmar. Science Advances 4: eaat5042. https://doi.org/10.1126/sciadv.aat5042" title="A mid-Cretaceous embryonic-to-neonate snake in amber from Myanmar. Science Advances 4: eaat 5042." url="https://doi.org/10.1126/sciadv.aat5042" year="2018">Xing et al. 2018</bibRefCitation>
|
||
). This feature should therefore be more properly and quantitatively assessed across many different scolecophidian genera, before it can obtain any diagnostic utility for taxonomic determinations.
|
||
</paragraph>
|
||
<paragraph id="D0DDC64CF6C827073588ED59513349C6" pageId="0" pageNumber="717">
|
||
In his monographic treatise,
|
||
<bibRefCitation id="9C0A641A8D48D8BBBB870E6D3645D498" DOI="https://doi.org/10.5962/bhl.title.50341" author="List, JC" journalOrPublisher="Illinois Biological Monographs" pageId="0" pageNumber="717" pagination="1 - 112" refId="B352" refString="List, JC, 1966. Comparative osteology of the snake families Typhlopidae and Leptotyphlopidae. Illinois Biological Monographs 36: 1 - 112, DOI: https://doi.org/10.5962/bhl.title.50341" title="Comparative osteology of the snake families Typhlopidae and Leptotyphlopidae." url="https://doi.org/10.5962/bhl.title.50341" volume="36" year="1966">List (1966)</bibRefCitation>
|
||
noted that some species of
|
||
<taxonomicName id="A62129D7F90DAA37418A60D3716474AA" class="Reptilia" family="Typhlopidae" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="" order="Squamata" pageId="0" pageNumber="717" phylum="Chordata" rank="family">Typhlopidae</taxonomicName>
|
||
possessed a peculiar rodlike ossification associated with the fused terminal vertebrae (not occurring in other snakes), which he named as urostyle, but besides he did not find any identifying vertebral characters.
|
||
</paragraph>
|
||
<paragraph id="E91765F60539B17FBCAB491D552D6E9A" pageId="0" pageNumber="717">
|
||
<bibRefCitation id="A89A8D2FBFADA341815F9D6F9EECF7B9" DOI="https://doi.org/10.5962/bhl.title.50341" author="List, JC" journalOrPublisher="Illinois Biological Monographs" pageId="0" pageNumber="717" pagination="1 - 112" refId="B352" refString="List, JC, 1966. Comparative osteology of the snake families Typhlopidae and Leptotyphlopidae. Illinois Biological Monographs 36: 1 - 112, DOI: https://doi.org/10.5962/bhl.title.50341" title="Comparative osteology of the snake families Typhlopidae and Leptotyphlopidae." url="https://doi.org/10.5962/bhl.title.50341" volume="36" year="1966">List (1966)</bibRefCitation>
|
||
also stated that zygosphenes and zygantra of leptotyphlopids are well dorsal to the level of the zygapophyseal joint.
|
||
<bibRefCitation id="97E9E668E79E3F7FDEF99542C8DA15B7" author="Holman, JA" journalOrPublisher="Indiana University Press, Bloomington and Indianapolis, IN" pageId="0" pageNumber="717" refId="B281" refString="Holman, JA, 2000. Fossil Snakes of North America: Origin, Evolution, Distribution, Paleoecology. Indiana University Press, Bloomington and Indianapolis, IN" title="Fossil Snakes of North America: Origin, Evolution, Distribution, Paleoecology." year="2000">Holman (2000)</bibRefCitation>
|
||
stated that vertebrae of leptotyphlopids are more elongate than those of typhlopids. We consider that these features are variable and more widespread across scolecophidians, and so they cannot be used for determination.
|
||
</paragraph>
|
||
<paragraph id="60B0134B8FB5422CF70578597B73A458" pageId="0" pageNumber="717">
|
||
One interesting feature that was addressed by
|
||
<bibRefCitation id="96E8E767F919E9BD6AED9FC2B475A16F" DOI="https://doi.org/10.1002/ar.23191" author="Pinto, RR" journalOrPublisher="Anatomical Record" pageId="0" pageNumber="717" pagination="1722 - 1747" refId="B462" refString="Pinto, RR, Martins, AR, Curcio, F, Ramos, LO, 2015. Osteology and cartilaginous elements of Trilepida salgueiroi (Amaral, 1954) (Scolecophidia: Leptotyphopidae). Anatomical Record 298: 1722 - 1747, DOI: https://doi.org/10.1002/ar.23191" title="Osteology and cartilaginous elements of Trilepida salgueiroi (Amaral, 1954) (Scolecophidia: Leptotyphopidae)." url="https://doi.org/10.1002/ar.23191" volume="298" year="2015">Pinto et al. (2015)</bibRefCitation>
|
||
is that leptotyphlopids seem to possess a higher number of caudal vertebrae than typhlopids and anomalepidids, a fact further highlighted subsequently by
|
||
<bibRefCitation id="C70AC05C12695845AE3C2CDAD2079C02" DOI="https://doi.org/10.1002/ar.24716" author="Martins, A" journalOrPublisher="Anatomical Record" pageId="0" pageNumber="717" pagination="2118 - 2148" refId="B369" refString="Martins, A, Koch, C, Joshi, M, Pinto, R, Machado, A, Lopes, R, Passos, P, 2021a. Evolutionary treasures hidden in the West Indies: Comparative osteology and visceral morphology reveals intricate miniaturization in the insular genera Mitophis Hedges, Adalsteinsson, & Branch, 2009 and Tetracheilostoma Jan, 1861 (Leptotyphlopidae: Epictinae: Tetracheilostomina). Anatomical Record 304: 2118 - 2148, DOI: https://doi.org/10.1002/ar.24716" title="Evolutionary treasures hidden in the West Indies: Comparative osteology and visceral morphology reveals intricate miniaturization in the insular genera Mitophis Hedges, Adalsteinsson, & Branch, 2009 and Tetracheilostoma Jan, 1861 (Leptotyphlopidae: Epictinae: Tetracheilostomina)." url="https://doi.org/10.1002/ar.24716" volume="304" year="2021 a">Martins et al. (2021a)</bibRefCitation>
|
||
. Based on our observations and the extensive literature overview presented below, we here tentatively concur with this statement, but we have to emphasize that data on caudal vertebral counts are missing from most typhlopid and leptotyphlopid species, in particular the African taxa. It would be interesting indeed to see how this number of caudal vertebrae ranges in the genera
|
||
<taxonomicName id="E4D5585FAD089400CA3BF3713F412E61" authorityName="Orejas-Miranda, Roux-Esteve & Guibe" authorityYear="1970" class="Reptilia" family="Leptotyphlopidae" genus="Rhinoleptus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Rhinoleptus" order="Squamata" pageId="0" pageNumber="717" phylum="Chordata" rank="genus">
|
||
<emphasis id="2C347ABEA8B2E02A53AA3316EE88D953" italics="true" pageId="0" pageNumber="717">Rhinoleptus</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<taxonomicName id="0A0574DB69F7E6B619547571D6380F3C" class="Reptilia" family="Leptotyphlopidae" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="" order="Squamata" pageId="0" pageNumber="717" phylum="Chordata" rank="family">Leptotyphlopidae</taxonomicName>
|
||
) and
|
||
<taxonomicName id="DD9FF72C961342AF305BFCA9F7CBDBD7" class="Squamata" family="Gerrhopilidae" genus="Letheobia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Letheobia" order="Squamata" pageId="0" pageNumber="717" phylum="Chordata" rank="genus">
|
||
<emphasis id="5CEA0D317E12A2EDEB380E65B80B06BB" italics="true" pageId="0" pageNumber="717">Letheobia</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<taxonomicName id="AE2D9B5076BBE65E41BC0C401ADC6725" class="Reptilia" family="Typhlopidae" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="" order="Squamata" pageId="0" pageNumber="717" phylum="Chordata" rank="family">Typhlopidae</taxonomicName>
|
||
), which possess the highest total vertebral counts among all extant snakes (see below).
|
||
</paragraph>
|
||
<paragraph id="EE1FE6654AC22DD006A7D2CF6A838D11" pageId="0" pageNumber="717">
|
||
<bibRefCitation id="A8CA0F914DB49AD32DA46DA1393E9808" DOI="https://doi.org/10.1016/j.isci.2020.101834" author="Fachini, ST" journalOrPublisher="Cuadernos de Herpetologia" pageId="0" pageNumber="717" refId="B170" refString="Fachini, ST, Onary, S, Palci, A, Lee, MSY, Bronzati, M, Hsiou, AS, 2020. Cretaceous blind snake from Brazil fills major gap in snake evolution. iScience 23: 101834. https://doi.org/10.1016/j.isci.2020.101834" title="Cretaceous blind snake from Brazil fills major gap in snake evolution. iScience 23: 101834." url="https://doi.org/10.1016/j.isci.2020.101834" year="2020">Fachini et al. (2020)</bibRefCitation>
|
||
further highlighted two additional characters, the "high position of the paradiapophyses (synapophyses in their terminology), which are located dorsal to the ventral margin of the cotyle" and the "smooth ventral margin of the centrum" as characteristic of typhlopids (and scolecophidians in general), considering these features as absent in all other snakes and therefore exclusively present for scolecophidians. We confirm the former feature as present (and sometimes distinct) in scolecophidians, but we have to note that this is also (intracolumnarily) variable present in few vertebrae of aniliids and uropeltids. As for the latter feature, we confirm that typhlopids (and scolecophidians as a whole) all possess a smooth centrum devoid of subcentral structures in their trunk vertebrae.
|
||
</paragraph>
|
||
<paragraph id="1D375473A88F659E393315CF1BFB048B" pageId="0" pageNumber="717">
|
||
Finally, one feature that was highlighted by
|
||
<bibRefCitation id="E4CDF9454CB56490B80EB07A1B382CF1" DOI="https://doi.org/10.1002/ar.24686" author="Herrel, A" journalOrPublisher="Anatomical Record" pageId="0" pageNumber="717" pagination="2292 - 2302" refId="B260" refString="Herrel, A, Lowie, A, Miralles, A, Gaucher, P, Kley, NJ, Measey, J, Tolley, KA, 2021. Burrowing in blindsnakes: A preliminary analysis of burrowing forces and consequences for the evolution of morphology. Anatomical Record 304: 2292 - 2302, DOI: https://doi.org/10.1002/ar.24686" title="Burrowing in blindsnakes: A preliminary analysis of burrowing forces and consequences for the evolution of morphology." url="https://doi.org/10.1002/ar.24686" volume="304" year="2021">Herrel et al. (2021)</bibRefCitation>
|
||
is that (at least certain) typhlopids possess more robust cotyles, condyles, prezygapophyses, and postzygapophyses in their anterior trunk vertebrae, compared to those of leptotyphlopids and anomalepidids, which was explained as an adaptation for higher pushing forces during burrowing of the former group. This would of course prove to be of importance for taxonomic determination but its utility should be further quantified and studied across a high number of taxa.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |