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<mods:title id="B96130A876C148FE7915D03097B07469">Squamates and amphibians from the Natufian cemetery of Raqefet Cave, Israel: taphonomy, paleoenvironments and paleoclimate</mods:title>
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<mods:namePart id="E624C7DC71C97AF3EFD2718FDA3EBE40">Ma‛ayan Lev, Dani Nadel, Mina Weinstein-Evron</mods:namePart>
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<mods:title id="54227F8FC199E26F4E69121B4440048C">An International Journal of Paleobiology</mods:title>
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<treatment id="03BD87E0FFB340296E07C329B68B1336" ID-DOI="http://doi.org/10.5281/zenodo.7542136" ID-GBIF-Taxon="208979637" ID-Zenodo-Dep="7542136" LSID="urn:lsid:plazi:treatment:03BD87E0FFB340296E07C329B68B1336" httpUri="http://treatment.plazi.org/id/03BD87E0FFB340296E07C329B68B1336" lastPageNumber="2406" pageId="12" pageNumber="2406">
<subSubSection id="C30E657DFFB340296E07C329B7171467" box="[112,535,598,621]" pageId="12" pageNumber="2406" type="nomenclature">
<paragraph id="8BAB36F6FFB340296E07C329B7171467" blockId="12.[112,535,598,621]" box="[112,535,598,621]" pageId="12" pageNumber="2406">
<heading id="D0E3819AFFB340296E07C329B7171467" bold="true" box="[112,535,598,621]" fontSize="10" level="6" pageId="12" pageNumber="2406" reason="2">
<taxonomicName id="4C144D75FFB340296E07C329B7171467" ID-CoL="33SV2" authority="(Werner, 1938)" baseAuthorityName="Werner" baseAuthorityYear="1938" box="[112,535,598,621]" class="Reptilia" family="Viperidae" genus="Daboia" isUncertain="true" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="2406" phylum="Chordata" rank="species" species="palaestinae">
<emphasis id="B960EAE4FFB340296E07C329B7171467" bold="true" box="[112,535,598,621]" pageId="12" pageNumber="2406">
Daboia
<emphasis id="B960EAE4FFB340296EBFC329B5E21467" bold="true" box="[200,226,598,621]" italics="true" pageId="12" pageNumber="2406">cf.</emphasis>
Palaestinae
<emphasis id="B960EAE4FFB340296F04C329B7171467" bold="true" box="[371,535,598,621]" italics="true" pageId="12" pageNumber="2406">(Werner, 1938)</emphasis>
</emphasis>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C30E657DFFB340296EE7C3FDB0B61493" pageId="12" pageNumber="2406" type="description">
<paragraph id="8BAB36F6FFB340296EE7C3FDB7561505" blockId="12.[112,798,642,1692]" pageId="12" pageNumber="2406">
Vipers are common in the assemblage (NISP = 116, 11%) and were identified solely by vertebrae (
<figureCitation id="132F2A73FFB340296FA9C3DFB73414BD" box="[478,564,672,695]" captionStart="Figure 6" captionStartId="11.[112,167,1063,1083]" captionTargetBox="[285,1340,151,1034]" captionTargetId="figure-507@11.[285,1341,143,1035]" captionTargetPageId="11" captionText="Figure 6. Snake bones from Raqefet Cave. Eryx jaculus (a) trunk vertebra, dorsal, ventral, anterior, posterior and lateral views; Hemorrhois nummifer (b) trunk vertebra, dorsal, ventral,anterior,posterior and lateral views; Dolichophis jugularis (c) trunk vertebra dorsal,ventral, anterior, posterior and lateral views; Malpolon insignitus (d) trunk vertebra dorsal, ventral, anterior, posterior and lateral views; Daboia cf. palaestinae (e) vertebra dorsal, ventral, anterior, posterior and lateral views." figureDoi="http://doi.org/10.5281/zenodo.7542253" httpUri="https://zenodo.org/record/7542253/files/figure.png" pageId="12" pageNumber="2406">Figure 6</figureCitation>
(e)). The trunk vertebrae of the
<taxonomicName id="4C144D75FFB340296E94C3C2B44714DE" authorityName="Oppel" authorityYear="1811" box="[227,327,701,724]" class="Reptilia" family="Viperidae" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="2406" phylum="Chordata" rank="family">Viperidae</taxonomicName>
family are characterised by the presence of straight hypapophyses throughout the precloacal region and posteriorly depressed neural arches (
<bibRefCitation id="EF854B07FFB340296FDEC387B7481505" author="Szyndlar Z." box="[425,584,760,783]" journalOrPublisher="Estudios Geologicos." pageId="12" pageNumber="2406" pagination="237 - 266" part="47" refId="ref20759" refString="Szyndlar Z. 1991 b. A review of Neogene and Quaternary snakes of Central and Eastern Europe. Part II: natricinae, Elapidae. Viperidae. Estudios Geologicos. 47: 237 - 266." title="A review of Neogene and Quaternary snakes of Central and Eastern Europe. Part II: natricinae, Elapidae. Viperidae" type="journal article" year="1991">Szyndlar 1991b</bibRefCitation>
).
</paragraph>
<paragraph id="8BAB36F6FFB340296EE7C26AB7A0137D" blockId="12.[112,798,642,1692]" pageId="12" pageNumber="2406">The vertebra is characterised by a large oval cotyle that is slightly ventrodorsally flattened. The condyle is large and oval. In dorsal view the vertebra is longer than wide with a well-marked interzygapophyseal constriction. The neural spine is relatively strong, high and long. The zygosphene is wide, and its anterior border is slightly concave. The prezygapophyseal particular facets are elongated and subtriangular. The prezygapophyses are long and pointed posterolaterally. In ventral view, the centrum has a slightly rounded triangular shape. The paradiapophyses are well-developed and protrude anteriorly from the centrum. The haemal keel is well-defined and expands posteriorly past the condyle and has two small subcentral foramen on each side. In anterior view, the zygophene is wide, slightly convex dorsally and concave ventrally. The neural canal is narrow, and its anterior edge is U-shaped. There are two small paracotylar foramina. The prezygapophyses extend dorsally and the prezygapophyseal accessory process is blunt and project dorsolaterally. In posterior view, the neural canal is wide with a U-shaped posterior edge. The postzygapophyses are laterally tilted. In lateral view, the neural arch starts to rise dorsally on the anterior edge, post the zygophene it is rectangular in shape. The parapophyses protrude anteroventrally past the cotyle.</paragraph>
<paragraph id="8BAB36F6FFB340296EE7C402B4771096" blockId="12.[112,798,642,1692]" pageId="12" pageNumber="2406">
The specimens most resemble the
<taxonomicName id="4C144D75FFB340296C76C402B7C6139E" baseAuthorityName="Werner" baseAuthorityYear="1938" box="[513,710,1405,1428]" class="Reptilia" family="Viperidae" genus="Daboia" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="2406" phylum="Chordata" rank="species" species="palaestinae">
<emphasis id="B960EAE4FFB340296C76C402B7C6139E" box="[513,710,1405,1428]" italics="true" pageId="12" pageNumber="2406">Daboia palaestinae</emphasis>
</taxonomicName>
species, the most common viperid species inhabiting the Mediterranean climate zone of the southern Levant. However, other similar viperids cannot be ruled out. Assuming that we deal with
<taxonomicName id="4C144D75FFB340296CFFC4ABB61713E6" baseAuthorityName="Werner" baseAuthorityYear="1938" box="[648,791,1492,1516]" class="Reptilia" family="Viperidae" genus="Daboia" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="2406" phylum="Chordata" rank="species" species="palaestinae">
<emphasis id="B960EAE4FFB340296CFFC4ABB61713E6" box="[648,791,1492,1516]" italics="true" pageId="12" pageNumber="2406">D. palaestinae</emphasis>
</taxonomicName>
, the species is mostly common in Mediterranean habitats: Mediterranean maquis, Mediterranean alluvial valleys, the Mediterranean coastal plain and the Mediterranean park forest (supplementary
<figureCitation id="132F2A73FFB340296F61C735B471106B" box="[278,369,1610,1633]" captionStart="Figure 2" captionStartId="3.[112,167,1296,1316]" captionTargetBox="[261,1369,151,1264]" captionTargetId="figure-387@3.[253,1373,143,1267]" captionTargetPageId="3" captionText="Figure 2. Plan of Raqefet Cave (a) provenance of the faunal samples, with the different contexts presented in this study labelled in red (b)." figureDoi="http://doi.org/10.5281/zenodo.7542243" httpUri="https://zenodo.org/record/7542243/files/figure.png" pageId="12" pageNumber="2406">
Figure
<date id="FFAA1036FFB340296F29C735B471106B" box="[350,369,1610,1633]" pageId="12" pageNumber="2406">2i</date>
</figureCitation>
). This species was suggested to be present at two Epipaleolithic sites in Israel: EWT (
<bibRefCitation id="EF854B07FFB340296C37C717B7E21075" author="Lev M &amp; Weinstein-Evron M &amp; Yeshurun R." box="[576,738,1640,1663]" journalOrPublisher="Sci Rep." pageId="12" pageNumber="2406" pagination="9373" part="10" refId="ref18820" refString="Lev M, Weinstein-Evron M, Yeshurun R. 2020. Squamate bone taphonomy: a new experimental framework and its application to the Natufian zooarchaeological record. Sci Rep. 10 (1): 9373. doi: 10.1038 / s 41598 - 020 - 66301 - 5." title="Squamate bone taphonomy: a new experimental framework and its application to the Natufian zooarchaeological record" type="journal article" year="2020">Lev et al. 2020</bibRefCitation>
) and Eynan (
<bibRefCitation id="EF854B07FFB340296EC8C7FAB4691096" author="Biton R &amp; Bailon S &amp; Birkenfeld M &amp; Bridault A &amp; Khalaily H &amp; Valla FR &amp; Rabinovich R." box="[191,361,1669,1692]" journalOrPublisher="PLoS One." pageId="12" pageNumber="2406" pagination="0247283" part="16" refId="ref16614" refString="Biton R, Bailon S, Birkenfeld M, Bridault A, Khalaily H, Valla FR, Rabinovich R. 2021. The anurans and squamates assemblage from Final Natufian Eynan (Ain Mallaha, Israel) with an emphasis on snake-human interactions. PLoS One. 16 (2): e 0247283. doi: 10.1371 / journal. pone. 0247283." title="The anurans and squamates assemblage from Final Natufian Eynan (Ain Mallaha, Israel) with an emphasis on snake-human interactions" type="journal article" year="2021">Biton et al. 2021</bibRefCitation>
).
</paragraph>
<paragraph id="8BAB36F6FFB340296E07C7B1B75E10EF" blockId="12.[112,606,1742,1765]" box="[112,606,1742,1765]" pageId="12" pageNumber="2406">
<emphasis id="B960EAE4FFB340296E07C7B1B75E10EF" bold="true" box="[112,606,1742,1765]" italics="true" pageId="12" pageNumber="2406">Intra-site analysis and comparison with EWT</emphasis>
</paragraph>
<paragraph id="8BAB36F6FFB340296E07C785B0B61493" blockId="12.[112,797,1786,2015]" lastBlockId="12.[829,1514,143,1340]" pageId="12" pageNumber="2406">
The intra-site comparison was conducted using NISP. The most commonly identified species in the assemblage is the Large Whip Snake (
<taxonomicName id="4C144D75FFB340296ECCC64AB48F1146" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[187,399,1845,1868]" class="Reptilia" family="Colubridae" genus="Dolichophis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="2406" phylum="Chordata" rank="species" species="jugularis">
<emphasis id="B960EAE4FFB340296ECCC64AB48F1146" box="[187,399,1845,1868]" italics="true" pageId="12" pageNumber="2406">Dolichophis jugularis</emphasis>
</taxonomicName>
, 16% of the total assemblage), followed by the European Glass Lizard (
<taxonomicName id="4C144D75FFB340296FADC62DB79E1163" baseAuthorityName="Pallas" baseAuthorityYear="1775" box="[474,670,1874,1897]" class="Reptilia" family="Anguidae" genus="Pseudopus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="2406" phylum="Chordata" rank="species" species="apodus">
<emphasis id="B960EAE4FFB340296FADC62DB79E1163" box="[474,670,1874,1897]" italics="true" pageId="12" pageNumber="2406">Pseudopus apodus</emphasis>
</taxonomicName>
, 13%), the Common Viper (
<taxonomicName id="4C144D75FFB340296F58C60FB71E118D" baseAuthorityName="Werner" baseAuthorityYear="1938" box="[303,542,1904,1927]" class="Reptilia" family="Viperidae" genus="Daboia" isUncertain="true" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="2406" phylum="Chordata" rank="species" species="palaestinae">
<emphasis id="B960EAE4FFB340296F58C60FB47A118D" box="[303,378,1904,1927]" italics="true" pageId="12" pageNumber="2406">Daboia</emphasis>
cf.
<emphasis id="B960EAE4FFB340296FDDC60FB71E118D" box="[426,542,1904,1927]" italics="true" pageId="12" pageNumber="2406">palaestinae</emphasis>
</taxonomicName>
, 11%) and the Eastern Montpellier Snake (
<taxonomicName id="4C144D75FFB340296F4BC6F2B70411A9" baseAuthorityName="Geoffroy De St-hilaire" baseAuthorityYear="1827" box="[316,516,1932,1956]" class="Reptilia" family="Lamprophiidae" genus="Malpolon" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="2406" phylum="Chordata" rank="species" species="insignitus">
<emphasis id="B960EAE4FFB340296F4BC6F2B70411A9" box="[316,516,1932,1956]" italics="true" pageId="12" pageNumber="2406">Malpolon insignitus</emphasis>
</taxonomicName>
, 7%). The taxonomic composition displays some intra-site differences (
<tableCitation id="C696034DFFB340296C32C6D5B79111CB" box="[581,657,1962,1985]" captionStart="Table 1" captionStartId="8.[112,160,143,163]" captionTargetPageId="8" captionText="Table 1. Taxonomic composition of Raqefet Cave squamate assemblage (NISP = 1072) by context." httpUri="http://table.plazi.org/id/DF6B667EFFB7402D6E07C1F0B44716B1" pageId="12" pageNumber="2406" tableUuid="DF6B667EFFB7402D6E07C1F0B44716B1">Table 1</tableCitation>
). Taxonomic diversity varies between the different contexts of the site (
<figureCitation id="132F2A73FFB340296CB0C6B7B61C11D5" box="[711,796,1992,2015]" captionStart="Figure 7" captionStartId="13.[112,167,1778,1798]" captionTargetBox="[134,786,147,1741]" captionTargetId="figure-349@13.[122,787,143,1749]" captionTargetPageId="13" captionText="Figure 7. Intra- and inter-site comparisons between Raqefet Cave (RAQ, shaded) and EWT contexts. The supposed ‘least anthropogenic̍ archaeological context in each site is denoted by arrows: (a) taxonomic evenness (Simpson̍s index), data from Table 1; (b) centrum length of the vertebrae (reflecting animal body size); (c) the protruding part index (measuring bone fragmentation)." figureDoi="http://doi.org/10.5281/zenodo.7542255" httpUri="https://zenodo.org/record/7542255/files/figure.png" pageId="12" pageNumber="2406">Figure 7</figureCitation>
(a)), with the highest evenness at the B5 niche (Simpson’s index of 0.8) and the lowest at bedrock basin C-XXIII (Simpson’s index of 0.69). The B5 niche taxonomic richness (NTaxa = 8) is similar to Locus 1, both are higher than Locus 3 (NTaxa = 5) and bedrock basin C-XXIII (NTaxa = 6). The most commonly identified species in the Natufian grave deposits (Loci 1 and 3; both are very similar) are the Large Whip Snake and the European Glass Lizard (
<tableCitation id="C696034DFFB340296BE7C040B0DB175C" box="[1424,1499,319,342]" captionStart="Table 1" captionStartId="8.[112,160,143,163]" captionTargetPageId="8" captionText="Table 1. Taxonomic composition of Raqefet Cave squamate assemblage (NISP = 1072) by context." httpUri="http://table.plazi.org/id/DF6B667EFFB7402D6E07C1F0B44716B1" pageId="12" pageNumber="2406" tableUuid="DF6B667EFFB7402D6E07C1F0B44716B1">Table 1</tableCitation>
), followed by the viper, the Eastern Montpellier Snake, and the Rough-tail Rock
<taxonomicName id="4C144D75FFB340296A7DC005B157179B" authorityName="Daudin" authorityYear="1802" box="[1034,1111,378,401]" class="Reptilia" family="Agamidae" genus="Agama" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="2406" phylum="Chordata" rank="genus">Agama</taxonomicName>
. The two other contexts present a different picture. The European Glass Lizard is by far the most common species in bedrock basin C-XXIII, possibly representing a whole specimen (evident by the representation of almost all body parts). The Glass Lizard is followed by the Eastern Montpellier Snake, the Rough-tail Rock
<taxonomicName id="4C144D75FFB340296A1DC372B1B4142E" authorityName="Daudin" authorityYear="1802" box="[1130,1204,525,548]" class="Reptilia" family="Agamidae" genus="Agama" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="2406" phylum="Chordata" rank="genus">Agama</taxonomicName>
, the Large Whip Snake, and the viper. In contrast, in the B5 Niche, the most common species is the Rough-tail Rock
<taxonomicName id="4C144D75FFB340296A79C338B1561454" authorityName="Daudin" authorityYear="1802" box="[1038,1110,583,606]" class="Reptilia" family="Agamidae" genus="Agama" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="2406" phylum="Chordata" rank="genus">Agama</taxonomicName>
(still only 13% of 65 NISP), followed by a few specimens of the European Glass Lizard, the Large Whip Snake, the Eastern Montpellier Snake and the common viper.
</paragraph>
</subSubSection>
<subSubSection id="C30E657DFFB340296D2AC3E0B68B1336" pageId="12" pageNumber="2406" type="diagnosis">
<paragraph id="8BAB36F6FFB340296D2AC3E0B16F15D6" blockId="12.[829,1514,143,1340]" pageId="12" pageNumber="2406">
We compare our results to the LN of the contemporaneous record from EWT,
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to the west. Similar squamate and amphibian taxa were identified in the EWT assemblage, but their relative abundances differed. The most commonly identified species in the assemblage as a whole was the European Glass Lizard, followed by the Large Whip Snake and the Eastern Montpellier Snake. Unlike the Raqefet assemblage, the viper, the third most common species in Raqefet Cave, is rare in the EWT assemblage (2% of the total assemblage;
<bibRefCitation id="EF854B07FFB340296DCBC2F5B15315AB" author="Lev M &amp; Weinstein-Evron M &amp; Yeshurun R." box="[956,1107,906,929]" journalOrPublisher="Sci Rep." pageId="12" pageNumber="2406" pagination="9373" part="10" refId="ref18820" refString="Lev M, Weinstein-Evron M, Yeshurun R. 2020. Squamate bone taphonomy: a new experimental framework and its application to the Natufian zooarchaeological record. Sci Rep. 10 (1): 9373. doi: 10.1038 / s 41598 - 020 - 66301 - 5." title="Squamate bone taphonomy: a new experimental framework and its application to the Natufian zooarchaeological record" type="journal article" year="2020">Lev et al. 2020</bibRefCitation>
). Here, too, a sample coming from the least anthropogenic context (Loc. 25) displayed the highest taxonomic evenness (
<figureCitation id="132F2A73FFB340296D98C2BAB14515D6" box="[1007,1093,965,988]" captionStart="Figure 7" captionStartId="13.[112,167,1778,1798]" captionTargetBox="[134,786,147,1741]" captionTargetId="figure-349@13.[122,787,143,1749]" captionTargetPageId="13" captionText="Figure 7. Intra- and inter-site comparisons between Raqefet Cave (RAQ, shaded) and EWT contexts. The supposed ‘least anthropogenic̍ archaeological context in each site is denoted by arrows: (a) taxonomic evenness (Simpson̍s index), data from Table 1; (b) centrum length of the vertebrae (reflecting animal body size); (c) the protruding part index (measuring bone fragmentation)." figureDoi="http://doi.org/10.5281/zenodo.7542255" httpUri="https://zenodo.org/record/7542255/files/figure.png" pageId="12" pageNumber="2406">Figure 7</figureCitation>
(a)).
</paragraph>
<paragraph id="8BAB36F6FFB340296D2AC29DB68B1336" blockId="12.[829,1514,143,1340]" pageId="12" pageNumber="2406">
There is also a clear body-size difference among contexts, measured by the centrum length of the trunk vertebrae (
<figureCitation id="132F2A73FFB340296B19C280B0C6121C" box="[1390,1478,1023,1046]" captionStart="Figure 7" captionStartId="13.[112,167,1778,1798]" captionTargetBox="[134,786,147,1741]" captionTargetId="figure-349@13.[122,787,143,1749]" captionTargetPageId="13" captionText="Figure 7. Intra- and inter-site comparisons between Raqefet Cave (RAQ, shaded) and EWT contexts. The supposed ‘least anthropogenic̍ archaeological context in each site is denoted by arrows: (a) taxonomic evenness (Simpson̍s index), data from Table 1; (b) centrum length of the vertebrae (reflecting animal body size); (c) the protruding part index (measuring bone fragmentation)." figureDoi="http://doi.org/10.5281/zenodo.7542255" httpUri="https://zenodo.org/record/7542255/files/figure.png" pageId="12" pageNumber="2406">Figure 7</figureCitation>
(b); one-way ANOVA, F = 4.78, p &lt;0.05). The mean centrum length of the vertebrae of B5 Niche is considerably smaller than all other samples (Tukey’s Q p &lt;0.05) and especially compared to Mortar C-XXIII. This pattern is again repeated at EWT where the Loc. 25 sample presents the smallest animals (
<figureCitation id="132F2A73FFB340296ABAC5EDB02412A3" box="[1229,1316,1170,1193]" captionStart="Figure 7" captionStartId="13.[112,167,1778,1798]" captionTargetBox="[134,786,147,1741]" captionTargetId="figure-349@13.[122,787,143,1749]" captionTargetPageId="13" captionText="Figure 7. Intra- and inter-site comparisons between Raqefet Cave (RAQ, shaded) and EWT contexts. The supposed ‘least anthropogenic̍ archaeological context in each site is denoted by arrows: (a) taxonomic evenness (Simpson̍s index), data from Table 1; (b) centrum length of the vertebrae (reflecting animal body size); (c) the protruding part index (measuring bone fragmentation)." figureDoi="http://doi.org/10.5281/zenodo.7542255" httpUri="https://zenodo.org/record/7542255/files/figure.png" pageId="12" pageNumber="2406">Figure 7</figureCitation>
(b)). Generally, the Raqefet Cave samples contain smaller-bodied squamates compared to the EWT samples, with the exception of Mortar C-XXIII that displays similarly large-bodied reptiles to the domestic contexts of EWT (Inside and Outside the human dwelling and Locus 67) and to the LN.
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