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<mods:title id="2B559F9A7ECAA10E619908D1425CAFC7">Miocene and Pliocene amphibians from Hambach (Germany): New evidence for a late Neogene refuge in northwestern Europe</mods:title>
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<taxonomicName id="4C5CECF0D074FFA8FB8A9AE3FBAD74E6" ID-CoL="PW" ID-ENA="8342" authorityName="Fischer" authorityYear="1813" box="[1068,1139,721,746]" class="Amphibia" kingdom="Animalia" order="Anura" pageId="32" pageNumber="33" phylum="Chordata" rank="order">Anura</taxonomicName>
indet.
</paragraph>
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<paragraph id="8BE39773D074FFA9FCEE9AC9FE3273F1" blockId="32.[840,1441,763,1908]" lastBlockId="33.[192,792,1252,1661]" lastPageId="33" lastPageNumber="34" pageId="32" pageNumber="33">
<emphasis id="B9284B61D074FFA8FCEE9AC9FC6A7518" bold="true" box="[840,948,763,788]" pageId="32" pageNumber="33">Material.</emphasis>
Hambach 6C: four premaxillae (IPB-HaH 2268/2270, IPB-HaH 2299); 51 fragments of maxilla (IPB-HaH 2233/2265, IPB-HaH 2282/2298, IPB-HaH 2300); 17 trunk vertebrae (IPB-HaH 2089/2097, IPB-HaH 2144/2145, IPB-HaH 2157/ 2158, IPB-HaH 2181/2183, IPB-HaH 2192); seven sacral vertebrae (IPB-HaH 2087, IPB-HaH 2206/ 2208, IPB-HaH 2217/2218, IPB-HaH 2225); 12 urostyles (IPB-HaH 2088, IPB-HaH 2197/2198, IPB-HaH 2201/2205, IPB-HaH 2216, IPB-HaH 2226/2227, IPB-HaH 2306); three coracoids (IPB-HaH 2313/2314, IPB-HaH 2378); nine humeri (IPB-HaH 2063, IPB-HaH 2303/2304, IPB-HaH 2311/ 2312, IPB-HaH 2317/2318, IPB-HaH 2331); five radioulnae (IPB-HaH 2301/2302, IPB-HaH 2309, IPB-HaH 2330, IPB-HaH 2335); two ilia (IPB-HaH 2319/2320); one ischium (IPB-HaH 2322); one femur (IPB-HaH 2326); 14 tibiofibulae (IPB-HaH 2054, IPB-HaH 2062, IPB-HaH 2064/2068, IPB-HaH 2305, IPB-HaH 2310, IPB-HaH 2327, IPB-HaH 2332/2334, IPB-HaH 2400); four indeterminate elements (IPB-HaH 2308, IPB-HaH 2315, IPB-HaH 2328, IPB-HaH 2398). Hambach 11: 29 maxillae (IPB-HaR 2044/2070, IPB-HaR 2177/ 2178); one angular (IPB-HaR 2114); two atlases (IPB-HaR 2025, IPB-HaR 2035); five trunk vertebrae (IPB-HaR 2024; IPB-HaR 2027/2029; IPB-HaR 2037); two sacral vertebrae (IPB-HaR 2026, IPB-HaR 2036); three urostyles (IPB-HaR 2038/ 2040); two coracoids (IPB-HaR 2125/2126); two scapulae (IPB-HaR 2072/2073); 10 humeri (IPB-HaR 2089/2095, IPB-HaR 2108/2109, IPB-HaR 2153); 16 radioulnae (IPB-HaR 2074/2079, IPB-HaR 2107, IPB-HaR 2123/2124, IPB-HaR 2128, IPB-HaR 2155/2157, IPB-HaR 2180/2182); four ilia (IPB-HaR 2085/2086, IPB-HaR 2104/2105); one ischium (IPB-HaR 2106); three femurs (IPB-HaR 2133/2134, IPB-HaR 2141); 25 tibiofibulae (IPB-HaR 2080/2082, IPB-HaR 2118/2122, IPB-HaR 2135/2140, IPB-HaR 2158/2168); one indeterminate element (IPB-HaR 2169). Hambach 11C: three maxillae (IPB-HaR 2422/2424); one ornamented bone fragment (IPB-HaR 2430); four tibiofibulae (IPB-HaR 2410/2413); one indeterminate element (IPB-HaR 2409).
</paragraph>
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<emphasis id="B9284B61D075FFA9FB879A9AFB1974CE" bold="true" box="[1057,1223,680,706]" pageId="33" pageNumber="34">DISCUSSION</emphasis>
</heading>
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<paragraph id="8BE39773D075FFA9FCEE9AEBFAF3751E" blockId="33.[840,1328,729,786]" pageId="33" pageNumber="34">
<emphasis id="B9284B61D075FFA9FCEE9AEBFAF3751E" bold="true" pageId="33" pageNumber="34">The Middle Miocene and Late Pliocene Amphibians from Hambach Compared</emphasis>
</paragraph>
<paragraph id="8BE39773D075FFAAFC249B1BFCDD705A" blockId="33.[840,1441,808,1890]" lastBlockId="34.[192,793,1149,1910]" lastPageId="34" lastPageNumber="35" pageId="33" pageNumber="34">
Deposition of the two fossiliferous levels in Hambach, stratum 6C (with site Hambach 6C) and stratum 11 (with sites Hambach 11 and 11C), is separated by about a dozen Myr and happened in very different global climatic contexts (the MCO vs a period of climate deterioration preceding the onset of Quaternary glacial cycles). This is clearly reflected in the faunal composition, with a far higher diversity in the Middle Miocene association compared to the Late Pliocene one and the presence of several taxa linked with subtropical conditions in the former (e.g., among amphibians and reptiles: cryptobranchids,
<emphasis id="B9284B61D075FFA9FBDF9C9AFAC472CE" box="[1145,1306,1192,1218]" italics="true" pageId="33" pageNumber="34">
<taxonomicName id="4C5CECF0D075FFA9FBDF9C9AFAC872CE" authorityName="Peters" authorityYear="1868" box="[1145,1302,1192,1218]" class="Testudines" family="Chelydridae" genus="Chelydropsis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">Chelydropsis</taxonomicName>
,
</emphasis>
<taxonomicName id="4C5CECF0D075FFA9FA849C9BFBC772EE" authority="Pomel, 1847" authorityName="Pomel" authorityYear="1847" class="Crocodylia" family="Alligatoridae" genus="Diplocynodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D075FFA9FA849C9BFCAA72EE" italics="true" pageId="33" pageNumber="34">Diplocynodon</emphasis>
<bibRefCitation id="EFCDEA82D075FFA9FCDB9CFBFBC772EE" author="Pomel, A." box="[893,1049,1225,1250]" pageId="33" pageNumber="34" pagination="378 - 385" refId="ref35448" refString="Pomel, A. 1847. Note sur des animaux fossiles decouverts dans le departement de l'Allier. Bulletin de la Societe geologique de France, 4: 378 - 385." type="journal article" year="1847">Pomel, 1847</bibRefCitation>
</taxonomicName>
, cobras). Based on our observations and identifications (
<tableCitation id="C6DEA2C8D075FFA9FBDF9CDBFB0B730E" box="[1145,1237,1257,1282]" captionStart="TABLE 1" captionStartId="34.[192,272,161,184]" captionTargetBox="[327,1293,207,1085]" captionText="TABLE 1. Distribution of amphibian taxa identified in the fossiliferous localities within the Hambach mine." pageId="33" pageNumber="34">Table 1</tableCitation>
), the amphibian component of the Hambach faunas matches with this pattern, all in all. The Middle Miocene fossil assemblage from Hambach 6C includes at least six different urodeles and six anurans. The Hambach 11 and 11C combined assemblage, on the other hand, listed only half the number of urodeles, even though the total of recognized anurans amounts to the same value. The remains attributed to the indeterminate cryptobranchid and to
<taxonomicName id="4C5CECF0D075FFA9FAF49E3AFC5D704E" authorityName="Pomel" authorityYear="1853" class="Amphibia" family="Salamandridae" genus="Chelotriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D075FFA9FAF49E3AFC5D704E" italics="true" pageId="33" pageNumber="34">Chelotriton</emphasis>
</taxonomicName>
stand as evidence of the already-mentioned “tropical” elements present during the Middle Miocene. In agreement with crocodylians (
<bibRefCitation id="EFCDEA82D075FFA9FAB09E5BFC5970AE" author="Mors, T &amp; von der Hocht, F. &amp; Wutzler, B." pageId="33" pageNumber="34" pagination="145 - 170" refId="ref35109" refString="Mors, T, von der Hocht, F., and Wutzler, B. 2000. Die erste Wirbeltierfauna aus der miozanen Braunkohle der Niederrheinischen Bucht (Ville-Schichten, Tagebau Hambach). Palaontologische Zeitschrift, 74: 145 - 170. https: // doi. org / 10.1007 / BF 02987958" type="journal article" year="2000">Mörs et al., 2000</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D075FFA9FC309EBBFBFC70AE" author="Mors, T." box="[918,1058,1673,1698]" pageId="33" pageNumber="34" pagination="177 - 183" refId="ref34899" refString="Mors, T. 2002. Biostratigraphy and paleoecology of continental Tertiary vertebrate faunas in the Lower Rhine Embayment (NW-Germany). Netherlands Journal of Geosciences - Geologie en Mijnbouw, 81: 177 - 183. https: // doi. org / 10.1017 / S 0016774600022411" type="journal article" year="2002">Mörs, 2002</bibRefCitation>
) and cobras (
<bibRefCitation id="EFCDEA82D075FFA9FB6B9EB6FC5470CE" author="Cernansky, A. &amp; Szyndlar, Z. &amp; Mors, T." pageId="33" pageNumber="34" pagination="329 - 354" refId="ref31064" refString="Cernansky, A., Szyndlar, Z., and Mors, T. 2017. Fossil squamate faunas from the Neogene of Hambach (northwestern Germany). Palaeobiodiversity and Palaeoenvironments, 97: 329 - 354. https: // doi. org / 10.1007 / s 12549 - 016 - 0252 - 1" type="journal article" year="2017">Čerňanský et al., 2017</bibRefCitation>
), but in contrast with what happens with
<taxonomicName id="4C5CECF0D075FFA9FAC09E9AFC6F70EE" authorityName="Peters" authorityYear="1868" class="Testudines" family="Chelydridae" genus="Chelydropsis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D075FFA9FAC09E9AFC6F70EE" italics="true" pageId="33" pageNumber="34">Chelydropsis</emphasis>
</taxonomicName>
turtles (which possibly persisted in the Upper Pliocene deposits;
<bibRefCitation id="EFCDEA82D075FFA9FBD19EDBFADD710E" author="Mors, T." box="[1143,1283,1769,1794]" pageId="33" pageNumber="34" pagination="177 - 183" refId="ref34899" refString="Mors, T. 2002. Biostratigraphy and paleoecology of continental Tertiary vertebrate faunas in the Lower Rhine Embayment (NW-Germany). Netherlands Journal of Geosciences - Geologie en Mijnbouw, 81: 177 - 183. https: // doi. org / 10.1017 / S 0016774600022411" type="journal article" year="2002">Mörs, 2002</bibRefCitation>
), the two urodeles are absent in younger sites at Hambach, hinting towards a likely extirpation between
<collectionCode id="ED4D0FB6D075FFA9FAFB9F1BFA58714E" box="[1373,1414,1833,1858]" country="Brazil" name="Museu Nacional, Universidade Federal do Rio de Janeiro" pageId="33" pageNumber="34">MN</collectionCode>
5 and
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16a. Other urodeles sharing the same pattern of a Miocene presence and Pliocene absence in Hambach include
<taxonomicName id="4C5CECF0D076FFAAFE1D9CAFFD7772BA" authorityName="Macaluso, Villa, and Mors" authorityYear="2022" box="[443,681,1181,1206]" class="Amphibia" family="Proteidae" genus="Euronecturus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="grogu">
<emphasis id="B9284B61D076FFAAFE1D9CAFFD7772BA" box="[443,681,1181,1206]" italics="true" pageId="34" pageNumber="35">Euronecturus grogu</emphasis>
</taxonomicName>
and
<taxonomicName id="4C5CECF0D076FFAAFD4D9CAFFEFF72DA" class="Amphibia" family="Salamandridae" genus="Triturus" kingdom="Animalia" order="Caudata" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFD4D9CAFFF2A72DA" italics="true" pageId="34" pageNumber="35">Triturus</emphasis>
sp.
</taxonomicName>
It is interesting to note that the proteid
<emphasis id="B9284B61D076FFAAFD599C8FFED572FA" italics="true" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFD599C8FFED972FA" authorityName="Macaluso, Villa, and Mors" authorityYear="2022" class="Amphibia" family="Proteidae" genus="Euronecturus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="grogu">E. grogu</taxonomicName>
,
</emphasis>
of which Hambach 6C is the only occurrence known worldwide at the moment, seems to be replaced by
<taxonomicName id="4C5CECF0D076FFAAFED39D2FFE27733A" authorityName="Estes and Darevsky" authorityYear="1977" box="[373,505,1309,1334]" class="Amphibia" family="Proteidae" genus="Mioproteus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="34" pageNumber="35" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D076FFAAFED39D2FFE27733A" box="[373,505,1309,1334]" italics="true" pageId="34" pageNumber="35">Mioproteus</emphasis>
</taxonomicName>
, a member of a different “modern” lineage of Eurasian proteids (
<bibRefCitation id="EFCDEA82D076FFAAFD059D0FFE83737A" author="Macaluso, L. &amp; Villa, A. &amp; Mors, T." pageId="34" pageNumber="35" pagination="12585" refId="ref34411" refString="Macaluso, L., Villa, A., and Mors, T. 2022 b. A new proteid salamander (Urodela; Proteidae) from the middle Miocene of Hambach (Germany) and implications for the evolution of the family. Palaeontology, 65: e 12585. https: // doi. org / 10.1111 / pala. 12585" type="journal article" year="2022">Macaluso et al., 2022b</bibRefCitation>
). The ranid frogs
<taxonomicName id="4C5CECF0D076FFAAFD949D6FFD3E737A" box="[562,736,1373,1398]" class="Amphibia" family="Ranidae" genus="Pelophylax" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFD949D6FFD6A737A" box="[562,692,1373,1398]" italics="true" pageId="34" pageNumber="35">Pelophylax</emphasis>
sp.
</taxonomicName>
and
<taxonomicName id="4C5CECF0D076FFAAFF669D4FFEF5739A" box="[192,299,1405,1430]" class="Amphibia" family="Ranidae" genus="Rana" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFF669D4FFEDE739A" box="[192,256,1405,1430]" italics="true" pageId="34" pageNumber="35">Rana</emphasis>
sp.
</taxonomicName>
are also recognized in Hambach 6C only, but the presence of ranids in the Upper Pliocene as well is attested by few remains from Hambach 11. At least one of these remains, the ilium IPB-HaR 2087, may even display green frog features, and thus the absence of either one or the two ranids from the younger assemblage may be artifactual.
</paragraph>
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<emphasis id="B9284B61D075FFA9FF669C2FFE997239" bold="true" box="[192,327,1053,1077]" pageId="33" pageNumber="34">FIGURE 23.</emphasis>
Reptiles from Hambach: right squamosal (IPB-HaH 2340) of
<taxonomicName id="4C5CECF0D075FFA9FE339C09FD72725E" authorityName="Cernansky" authorityYear="2010" box="[405,684,1083,1106]" class="Squamata" family="Chamaeleonidae" genus="Chamaeleo" isUncertain="true" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="andrusovi">
<emphasis id="B9284B61D075FFA9FE339C09FDCE725E" box="[405,528,1083,1106]" italics="true" pageId="33" pageNumber="34">Chamaeleo</emphasis>
aff.
<emphasis id="B9284B61D075FFA9FDE29C09FD72725E" box="[580,684,1083,1106]" italics="true" pageId="33" pageNumber="34">andrusovi</emphasis>
</taxonomicName>
in lateral (A) and medial (B) views; left frontal (IPB-HaR 2171) of
<taxonomicName id="4C5CECF0D075FFA9FF669C44FE027281" baseAuthorityName="Kormos" baseAuthorityYear="1911" box="[192,476,1141,1165]" class="Squamata" family="Anguidae" genus="Pseudopus" isUncertain="true" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="pannonicus">
<emphasis id="B9284B61D075FFA9FF669C44FEE67281" box="[192,312,1142,1165]" italics="true" pageId="33" pageNumber="34">Pseudopus</emphasis>
cf.
<emphasis id="B9284B61D075FFA9FEC49C44FE027281" box="[354,476,1142,1165]" italics="true" pageId="33" pageNumber="34">pannonicus</emphasis>
</taxonomicName>
in dorsal (C) and ventral (D) views. Scale bars equal 1 mm.
</paragraph>
</caption>
<subSubSection id="C346C4F8D075FFA9FF669E36FDCE7071" pageId="33" pageNumber="34" type="discussion">
<paragraph id="8BE39773D075FFA9FF669E36FDCE7071" blockId="33.[192,792,1252,1661]" pageId="33" pageNumber="34">
<emphasis id="B9284B61D075FFA9FF669E36FEE67011" bold="true" box="[192,312,1540,1565]" pageId="33" pageNumber="34">Remarks.</emphasis>
Numerous fossil elements can be assigned to anurans, but are either too poorly preserved or not taxonomically significant enough for a more refined identification.
</paragraph>
</subSubSection>
<subSection id="E2D38E98D075FFA9FF669EA6FA5B748E" pageId="33" pageNumber="34" type="multiple">
<paragraph id="8BE39773D075FFA9FF669EA6FEF970A1" blockId="33.[192,295,1684,1709]" box="[192,295,1684,1709]" pageId="33" pageNumber="34">
<emphasis id="B9284B61D075FFA9FF669EA6FEF970A1" bold="true" box="[192,295,1684,1709]" pageId="33" pageNumber="34">Reptiles</emphasis>
</paragraph>
<paragraph id="8BE39773D075FFA9FF5C9EF6FA5B748E" blockId="33.[192,793,1732,1885]" lastBlockId="33.[840,1441,168,642]" pageId="33" pageNumber="34">
While studying the amphibian material, we chanced upon a few remains of reptiles that were accidentally not included in Čerňanský et al.s (2017) work. Most of these are fragments of poorly diagnostic bones, but at least
<specimenCount id="9D5A5CFAD075FFA9FD959F76FD387151" box="[563,742,1860,1885]" count="2" pageId="33" pageNumber="34" type="generic">two specimens</specimenCount>
are worth mentioning and figuring. IPB-HaH 2340 (
<figureCitation id="13678BF6D075FFA9FAD6989BFC4876EE" captionStart="FIGURE 23" captionStartId="33.[192,283,1053,1076]" captionTargetBox="[194,789,182,1029]" captionTargetId="figure-538@33.[190,790,172,1034]" captionTargetPageId="33" captionText="FIGURE 23. Reptiles from Hambach: right squamosal (IPB-HaH 2340) of Chamaeleo aff. andrusovi in lateral (A) and medial (B) views; left frontal (IPB-HaR 2171) of Pseudopus cf. pannonicus in dorsal (C) and ventral (D) views. Scale bars equal 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10962343" httpUri="https://zenodo.org/record/10962343/files/figure.png" pageId="33" pageNumber="34">Figure 23</figureCitation>
A-B), from Hambach 6C, is probably a fragment of the posterodorsal process of a right jugal. The ornamentation recalls
<taxonomicName id="4C5CECF0D075FFA9FB30993AFBC9774E" authority="Cernansky, 2010" authorityName="Cernansky" authorityYear="2010" class="Squamata" family="Chamaeleonidae" genus="Chamaeleo" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="andrusovi">
<emphasis id="B9284B61D075FFA9FB30993AFA7E772E" box="[1174,1440,264,290]" italics="true" pageId="33" pageNumber="34">Chamaeleo andrusovi</emphasis>
<bibRefCitation id="EFCDEA82D075FFA9FCEE9916FBC9774E" author="Cernansky, A." box="[840,1047,292,322]" pageId="33" pageNumber="34" pagination="605 - 613" refId="ref31010" refString="Cernansky, A. 2010. A revision of chamaeleonids from the Lower Miocene of the Czech Republic with description of a new species of Chamaeleo (Squamata, Chamaeleonidae). Geobios, 43: 605 - 613. https: // doi. org / 10.1016 / j. geobios. 2010.04.001" type="journal article" year="2010">Čerňanský, 2010</bibRefCitation>
</taxonomicName>
, even though the topotypic jugals of this species show two rows of protuberances instead of just one as in the case of the specimen from Hambach. It is here attributed to
<taxonomicName id="4C5CECF0D075FFA9FABE99BAFC3277CE" authorityName="Cernansky" authorityYear="2010" class="Squamata" family="Chamaeleonidae" genus="Chamaeleo" isUncertain="true" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="andrusovi">
<emphasis id="B9284B61D075FFA9FABE99BAFA7E77AD" box="[1304,1440,392,417]" italics="true" pageId="33" pageNumber="34">Chamaeleo</emphasis>
aff.
<emphasis id="B9284B61D075FFA9FCDE999BFC3277CE" box="[888,1004,425,450]" italics="true" pageId="33" pageNumber="34">andrusovi</emphasis>
</taxonomicName>
, similar to the squamosal fragment described by
<bibRefCitation id="EFCDEA82D075FFA9FC5199F6FAFB77EE" author="Cernansky, A. &amp; Szyndlar, Z. &amp; Mors, T." box="[1015,1317,452,482]" pageId="33" pageNumber="34" pagination="329 - 354" refId="ref31064" refString="Cernansky, A., Szyndlar, Z., and Mors, T. 2017. Fossil squamate faunas from the Neogene of Hambach (northwestern Germany). Palaeobiodiversity and Palaeoenvironments, 97: 329 - 354. https: // doi. org / 10.1007 / s 12549 - 016 - 0252 - 1" type="journal article" year="2017">Čerňanský et al. (2017)</bibRefCitation>
from the same site. IPB-HaR 2171 (
<figureCitation id="13678BF6D075FFA9FB0799DBFAC1740E" box="[1185,1311,488,514]" captionStart="FIGURE 23" captionStartId="33.[192,283,1053,1076]" captionTargetBox="[194,789,182,1029]" captionTargetId="figure-538@33.[190,790,172,1034]" captionTargetPageId="33" captionText="FIGURE 23. Reptiles from Hambach: right squamosal (IPB-HaH 2340) of Chamaeleo aff. andrusovi in lateral (A) and medial (B) views; left frontal (IPB-HaR 2171) of Pseudopus cf. pannonicus in dorsal (C) and ventral (D) views. Scale bars equal 1 mm." figureDoi="http://doi.org/10.5281/zenodo.10962343" httpUri="https://zenodo.org/record/10962343/files/figure.png" pageId="33" pageNumber="34">Figure 23</figureCitation>
C-D), from Hambach 11, can be associated to the same taxon as the nasal assigned by
<bibRefCitation id="EFCDEA82D075FFA9FB259A16FA41744E" author="Cernansky, A. &amp; Szyndlar, Z. &amp; Mors, T." box="[1155,1439,548,578]" pageId="33" pageNumber="34" pagination="329 - 354" refId="ref31064" refString="Cernansky, A., Szyndlar, Z., and Mors, T. 2017. Fossil squamate faunas from the Neogene of Hambach (northwestern Germany). Palaeobiodiversity and Palaeoenvironments, 97: 329 - 354. https: // doi. org / 10.1007 / s 12549 - 016 - 0252 - 1" type="journal article" year="2017">Čerňanský et al. (2017)</bibRefCitation>
to
<taxonomicName id="4C5CECF0D075FFA9FCD49A7BFA42746E" authority="(Kormos, 1911)" baseAuthorityName="Kormos" baseAuthorityYear="1911" box="[882,1436,584,610]" class="Squamata" family="Anguidae" genus="Pseudopus" isUncertain="true" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="pannonicus">
<emphasis id="B9284B61D075FFA9FCD49A7BFC29746E" box="[882,1015,585,610]" italics="true" pageId="33" pageNumber="34">Pseudopus</emphasis>
cf.
<emphasis id="B9284B61D075FFA9FB9D9A7BFB1D746E" box="[1083,1219,585,610]" italics="true" pageId="33" pageNumber="34">pannonicus</emphasis>
(
<bibRefCitation id="EFCDEA82D075FFA9FB7A9A7BFA4B746E" author="Kormos, T." box="[1244,1429,585,610]" pageId="33" pageNumber="34" pagination="48 - 64" refId="ref33803" refString="Kormos, T. 1911. A Polgardi pliocen csontlelet. Foldtani Kozlony, 41: 48 - 64." type="journal article" year="1911">Kormos, 1911</bibRefCitation>
)
</taxonomicName>
, because they share a similar dermal sculpturing.
</paragraph>
</subSection>
<caption id="DF23C7FBD076FFAAFF669893FAC876B4" box="[192,1302,161,184]" pageId="34" pageNumber="35" startId="34.[192,272,161,184]" targetBox="[327,1293,207,1085]" targetIsTable="true" targetPageId="34" targetType="table">
<paragraph id="8BE39773D076FFAAFF669893FAC876B4" blockId="34.[192,1302,161,184]" box="[192,1302,161,184]" pageId="34" pageNumber="35">
<emphasis id="B9284B61D076FFAAFF669893FEF776B4" bold="true" box="[192,297,161,184]" pageId="34" pageNumber="35">TABLE 1.</emphasis>
Distribution of amphibian taxa identified in the fossiliferous localities within the Hambach mine.
</paragraph>
</caption>
<paragraph id="8BE39773D076FFAAFD0798FDFB137231" pageId="34" pageNumber="35">
<table id="F95C65D3D0760077FEE198FDFAD37231" box="[327,1293,207,1085]" gridcols="4" gridrows="24" pageId="34" pageNumber="35">
<tr id="356C9531D0760077FEE198FDFAD376EF" box="[327,1293,207,227]" gridrow="0" pageId="34" pageNumber="35" rowspan-0="1">
<th id="76BDFC4DD0760077FD0798FDFCE176EF" box="[673,831,207,227]" gridcol="1" gridrow="0" pageId="34" pageNumber="35">Middle Miocene</th>
<th id="76BDFC4DD0760077FC1798FDFAD376EF" box="[945,1293,207,227]" colspan="2" colspanRight="1" gridcol="2" gridrow="0" pageId="34" pageNumber="35">Late Pliocene</th>
</tr>
<tr id="356C9531D0760077FEE198C3FAD37709" box="[327,1293,241,261]" gridrow="1" pageId="34" pageNumber="35" rowspan-0="1">
<td id="76BDFC4DD0760077FD0798C3FCE17709" box="[673,831,241,261]" gridcol="1" gridrow="1" pageId="34" pageNumber="35">Hambach 6C</td>
<td id="76BDFC4DD0760077FC1798C3FBF37709" box="[945,1069,241,261]" gridcol="2" gridrow="1" pageId="34" pageNumber="35">Hambach 11</td>
<td id="76BDFC4DD0760077FB2798C3FAD37709" box="[1153,1293,241,261]" gridcol="3" gridrow="1" pageId="34" pageNumber="35">
<emphasis id="B9284B61D076FFAAFB2798C3FAD37709" bold="true" box="[1153,1293,241,261]" pageId="34" pageNumber="35">Hambach 11C</emphasis>
</td>
</tr>
<tr id="356C9531D0760077FEE1992BFAD37721" box="[327,1293,281,301]" gridrow="2" pageId="34" pageNumber="35" rowspan-2="1" rowspan-3="1">
<th id="76BDFC4DD0760077FEE1992BFDB77721" box="[327,617,281,301]" gridcol="0" gridrow="2" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFEE1992BFE317721" authorityName="Fitzinger" authorityYear="1826" box="[327,495,281,301]" class="Amphibia" family="Cryptobranchidae" kingdom="Animalia" order="Caudata" pageId="34" pageNumber="35" phylum="Chordata" rank="family">Cryptobranchidae</taxonomicName>
indet.
</th>
<td id="76BDFC4DD0760077FD07992BFCE17721" box="[673,831,281,301]" gridcol="1" gridrow="2" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE1990CFAD3775F" box="[327,1293,318,339]" gridrow="3" pageId="34" pageNumber="35">
<th id="76BDFC4DD0760077FEE1990CFDB7775F" box="[327,617,318,339]" gridcol="0" gridrow="3" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFEE1990DFD80775F" baseAuthorityName="Vasilyan and Yanenko" baseAuthorityYear="2020" box="[327,606,318,339]" class="Amphibia" family="Batrachosauroididae" genus="Palaeoproteus" higherTaxonomySource="GBIF" isUncertain="true" kingdom="Animalia" order="Caudata" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="miocenicus">
<emphasis id="B9284B61D076FFAAFEE1990DFE0E775F" box="[327,464,319,339]" italics="true" pageId="34" pageNumber="35">Palaeoproteus</emphasis>
cf.
<emphasis id="B9284B61D076FFAAFE55990DFD80775F" box="[499,606,319,339]" italics="true" pageId="34" pageNumber="35">miocenicus</emphasis>
</taxonomicName>
</th>
<td id="76BDFC4DD0760077FD07990CFCE1775F" box="[673,831,318,339]" gridcol="1" gridrow="3" pageId="34" pageNumber="35">x</td>
<td id="76BDFC4DD0760077FC17990CFBF3775F" box="[945,1069,318,339]" gridcol="2" gridrow="3" pageId="34" pageNumber="35">x</td>
<td id="76BDFC4DD0760077FB27990CFAD3775F" box="[1153,1293,318,339]" gridcol="3" gridrow="3" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE19956FAD37774" box="[327,1293,356,376]" gridrow="4" pageId="34" pageNumber="35" rowspan-2="1" rowspan-3="1">
<th id="76BDFC4DD0760077FEE19956FDB77774" box="[327,617,356,376]" gridcol="0" gridrow="4" pageId="34" pageNumber="35">
?
<taxonomicName id="4C5CECF0D076FFAAFEF59956FDB77774" baseAuthorityName="Vasilyan and Yanenko" baseAuthorityYear="2020" box="[339,617,356,376]" class="Amphibia" family="Batrachosauroididae" genus="Palaeoproteus" higherTaxonomySource="GBIF" isUncertain="true" kingdom="Animalia" order="Caudata" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="miocenicus">
<emphasis id="B9284B61D076FFAAFEF59956FE027774" box="[339,476,356,376]" italics="true" pageId="34" pageNumber="35">Palaeoproteus</emphasis>
cf.
<emphasis id="B9284B61D076FFAAFE589956FDB77774" box="[510,617,356,376]" italics="true" pageId="34" pageNumber="35">miocenicus</emphasis>
</taxonomicName>
</th>
<td id="76BDFC4DD0760077FD079956FCE17774" box="[673,831,356,376]" gridcol="1" gridrow="4" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE199BBFAD37791" box="[327,1293,393,413]" gridrow="5" pageId="34" pageNumber="35" rowspan-2="1" rowspan-3="1">
<th id="76BDFC4DD0760077FEE199BBFDB77791" box="[327,617,393,413]" gridcol="0" gridrow="5" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFEE199BBFDDC7791" authorityName="Macaluso, Villa, and Mors" authorityYear="2022" box="[327,514,393,413]" class="Amphibia" family="Proteidae" genus="Euronecturus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="grogu">
<emphasis id="B9284B61D076FFAAFEE199BBFDDC7791" box="[327,514,393,413]" italics="true" pageId="34" pageNumber="35">Euronecturus grogu</emphasis>
</taxonomicName>
</th>
<td id="76BDFC4DD0760077FD0799BBFCE17791" box="[673,831,393,413]" gridcol="1" gridrow="5" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE1999CFAD377CF" box="[327,1293,430,451]" gridrow="6" pageId="34" pageNumber="35" rowspan-1="1">
<th id="76BDFC4DD0760077FEE1999CFDB777CF" box="[327,617,430,451]" gridcol="0" gridrow="6" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFEE1999DFDD777CF" box="[327,521,430,451]" class="Amphibia" family="Proteidae" genus="Mioproteus" higherTaxonomySource="GBIF" isUncertain="true" kingdom="Animalia" order="Caudata" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="wezei">
<emphasis id="B9284B61D076FFAAFEE1999DFE6E77CF" box="[327,432,431,451]" italics="true" pageId="34" pageNumber="35">Mioproteus</emphasis>
cf.
<emphasis id="B9284B61D076FFAAFE75999DFDD777CF" box="[467,521,431,451]" italics="true" pageId="34" pageNumber="35">wezei</emphasis>
</taxonomicName>
</th>
<td id="76BDFC4DD0760077FC17999CFBF377CF" box="[945,1069,430,451]" gridcol="2" gridrow="6" pageId="34" pageNumber="35">x</td>
<td id="76BDFC4DD0760077FB27999CFAD377CF" box="[1153,1293,430,451]" gridcol="3" gridrow="6" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE199E6FAD377E4" box="[327,1293,468,488]" gridrow="7" pageId="34" pageNumber="35" rowspan-2="1" rowspan-3="1">
<th id="76BDFC4DD0760077FEE199E6FDB777E4" box="[327,617,468,488]" gridcol="0" gridrow="7" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFEE199E6FE0F77E4" box="[327,465,468,488]" class="Amphibia" family="Salamandridae" genus="Chelotriton" kingdom="Animalia" order="Caudata" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFEE199E6FE7077E4" box="[327,430,468,488]" italics="true" pageId="34" pageNumber="35">Chelotriton</emphasis>
sp.
</taxonomicName>
</th>
<td id="76BDFC4DD0760077FD0799E6FCE177E4" box="[673,831,468,488]" gridcol="1" gridrow="7" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE199CBFAD37401" box="[327,1293,505,525]" gridrow="8" pageId="34" pageNumber="35" rowspan-2="1" rowspan-3="1">
<th id="76BDFC4DD0760077FEE199CBFDB77401" box="[327,617,505,525]" gridcol="0" gridrow="8" pageId="34" pageNumber="35">
?
<taxonomicName id="4C5CECF0D076FFAAFEF599CBFE037401" box="[339,477,505,525]" class="Amphibia" family="Salamandridae" genus="Chelotriton" kingdom="Animalia" order="Caudata" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFEF599CBFE647401" box="[339,442,505,525]" italics="true" pageId="34" pageNumber="35">Chelotriton</emphasis>
sp.
</taxonomicName>
</th>
<td id="76BDFC4DD0760077FD0799CBFCE17401" box="[673,831,505,525]" gridcol="1" gridrow="8" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE19A2DFAD3743F" box="[327,1293,543,563]" gridrow="9" pageId="34" pageNumber="35" rowspan-3="1">
<th id="76BDFC4DD0760077FEE19A2DFDB7743F" box="[327,617,543,563]" gridcol="0" gridrow="9" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFEE19A2DFE12743F" box="[327,460,543,563]" class="Amphibia" family="Salamandridae" genus="Lissotriton" kingdom="Animalia" order="Caudata" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFEE19A2DFE76743F" box="[327,424,543,563]" italics="true" pageId="34" pageNumber="35">Lissotriton</emphasis>
sp.
</taxonomicName>
</th>
<td id="76BDFC4DD0760077FD079A2DFCE1743F" box="[673,831,543,563]" gridcol="1" gridrow="9" pageId="34" pageNumber="35">x</td>
<td id="76BDFC4DD0760077FC179A2DFBF3743F" box="[945,1069,543,563]" gridcol="2" gridrow="9" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE19A76FAD37454" box="[327,1293,580,600]" gridrow="10" pageId="34" pageNumber="35" rowspan-2="1" rowspan-3="1">
<th id="76BDFC4DD0760077FEE19A76FDB77454" box="[327,617,580,600]" gridcol="0" gridrow="10" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFEE19A76FE6F7454" box="[327,433,580,600]" class="Amphibia" family="Salamandridae" genus="Triturus" kingdom="Animalia" order="Caudata" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFEE19A76FE507454" box="[327,398,580,600]" italics="true" pageId="34" pageNumber="35">Triturus</emphasis>
sp.
</taxonomicName>
</th>
<td id="76BDFC4DD0760077FD079A76FCE17454" box="[673,831,580,600]" gridcol="1" gridrow="10" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE19A5BFAD37471" box="[327,1293,617,637]" gridrow="11" pageId="34" pageNumber="35" rowspan-2="1" rowspan-3="1">
<th id="76BDFC4DD0760077FEE19A5BFDB77471" box="[327,617,617,637]" gridcol="0" gridrow="11" pageId="34" pageNumber="35">Urodela indet.</th>
<td id="76BDFC4DD0760077FD079A5BFCE17471" box="[673,831,617,637]" gridcol="1" gridrow="11" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE19ABDFAD374AF" box="[327,1293,655,675]" gridrow="12" pageId="34" pageNumber="35">
<th id="76BDFC4DD0760077FEE19ABDFDB774AF" box="[327,617,655,675]" gridcol="0" gridrow="12" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFEE19ABDFE6E74AF" box="[327,432,655,675]" class="Amphibia" family="Alytidae" genus="Latonia" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFEE19ABDFE5374AF" box="[327,397,655,675]" italics="true" pageId="34" pageNumber="35">Latonia</emphasis>
sp.
</taxonomicName>
</th>
<td id="76BDFC4DD0760077FD079ABDFCE174AF" box="[673,831,655,675]" gridcol="1" gridrow="12" pageId="34" pageNumber="35">x</td>
<td id="76BDFC4DD0760077FC179ABDFBF374AF" box="[945,1069,655,675]" gridcol="2" gridrow="12" pageId="34" pageNumber="35">x</td>
<td id="76BDFC4DD0760077FB279ABDFAD374AF" box="[1153,1293,655,675]" gridcol="3" gridrow="12" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE19A86FAD374C4" box="[327,1293,692,712]" gridrow="13" pageId="34" pageNumber="35" rowspan-1="1">
<th id="76BDFC4DD0760077FEE19A86FDB774C4" box="[327,617,692,712]" gridcol="0" gridrow="13" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFEE19A86FD9674C4" box="[327,584,692,712]" class="Amphibia" family="Palaeobatrachidae" genus="Palaeobatrachus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="eurydices">
<emphasis id="B9284B61D076FFAAFEE19A86FD9674C4" box="[327,584,692,712]" italics="true" pageId="34" pageNumber="35">Palaeobatrachus eurydices</emphasis>
</taxonomicName>
</th>
<td id="76BDFC4DD0760077FC179A86FBF374C4" box="[945,1069,692,712]" gridcol="2" gridrow="13" pageId="34" pageNumber="35">x</td>
<td id="76BDFC4DD0760077FB279A86FAD374C4" box="[1153,1293,692,712]" gridcol="3" gridrow="13" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE19AEBFAD374E1" box="[327,1293,729,749]" gridrow="14" pageId="34" pageNumber="35" rowspan-2="1" rowspan-3="1">
<th id="76BDFC4DD0760077FEE19AEBFDB774E1" box="[327,617,729,749]" gridcol="0" gridrow="14" pageId="34" pageNumber="35">
?
<taxonomicName id="4C5CECF0D076FFAAFEF69AEBFDDA74E1" authorityName="Cope" authorityYear="1865" box="[336,516,729,749]" class="Amphibia" family="Palaeobatrachidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="family">Palaeobatrachidae</taxonomicName>
indet.
</th>
<td id="76BDFC4DD0760077FD079AEBFCE174E1" box="[673,831,729,749]" gridcol="1" gridrow="14" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE19ACCFAD3751F" box="[327,1293,766,787]" gridrow="15" pageId="34" pageNumber="35" rowspan-1="1" rowspan-3="1">
<th id="76BDFC4DD0760077FEE19ACCFDB7751F" box="[327,617,766,787]" gridcol="0" gridrow="15" pageId="34" pageNumber="35">
cf.
<taxonomicName id="4C5CECF0D076FFAAFEC59ACDFE23751F" box="[355,509,767,787]" class="Amphibia" family="Pelobatidae" genus="Eopelobates" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFEC59ACDFE04751F" box="[355,474,767,787]" italics="true" pageId="34" pageNumber="35">Eopelobates</emphasis>
sp.
</taxonomicName>
</th>
<td id="76BDFC4DD0760077FC179ACCFBF3751F" box="[945,1069,766,787]" gridcol="2" gridrow="15" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE19B16FAD37534" box="[327,1293,804,824]" gridrow="16" pageId="34" pageNumber="35" rowspan-1="1" rowspan-2="1">
<th id="76BDFC4DD0760077FEE19B16FDB77534" box="[327,617,804,824]" gridcol="0" gridrow="16" pageId="34" pageNumber="35">
?
<taxonomicName id="4C5CECF0D076FFAAFEF59B16FE337534" box="[339,493,804,824]" class="Amphibia" family="Pelobatidae" genus="Eopelobates" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFEF59B16FE147534" box="[339,458,804,824]" italics="true" pageId="34" pageNumber="35">Eopelobates</emphasis>
sp.
</taxonomicName>
</th>
<td id="76BDFC4DD0760077FB279B16FAD37534" box="[1153,1293,804,824]" gridcol="3" gridrow="16" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE19B7BFAD37551" box="[327,1293,841,861]" gridrow="17" pageId="34" pageNumber="35" rowspan-2="1" rowspan-3="1">
<th id="76BDFC4DD0760077FEE19B7BFDB77551" box="[327,617,841,861]" gridcol="0" gridrow="17" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFEE19B7BFE697551" authorityName="Bonaparte" authorityYear="1850" box="[327,439,841,861]" class="Amphibia" family="Pelobatidae" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="family">Pelobatidae</taxonomicName>
indet.
</th>
<td id="76BDFC4DD0760077FD079B7BFCE17551" box="[673,831,841,861]" gridcol="1" gridrow="17" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE19B5DFAD3758F" box="[327,1293,879,899]" gridrow="18" pageId="34" pageNumber="35" rowspan-3="1">
<th id="76BDFC4DD0760077FEE19B5DFDB7758F" box="[327,617,879,899]" gridcol="0" gridrow="18" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFEE19B5DFE4B758F" box="[327,405,879,899]" class="Amphibia" family="Hylidae" genus="Hyla" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFEE19B5DFEAC758F" box="[327,370,879,899]" italics="true" pageId="34" pageNumber="35">Hyla</emphasis>
sp.
</taxonomicName>
</th>
<td id="76BDFC4DD0760077FD079B5DFCE1758F" box="[673,831,879,899]" gridcol="1" gridrow="18" pageId="34" pageNumber="35">x</td>
<td id="76BDFC4DD0760077FC179B5DFBF3758F" box="[945,1069,879,899]" gridcol="2" gridrow="18" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE19BA6FAD375A4" box="[327,1293,916,936]" gridrow="19" pageId="34" pageNumber="35" rowspan-1="1" rowspan-3="1">
<th id="76BDFC4DD0760077FEE19BA6FDB775A4" box="[327,617,916,936]" gridcol="0" gridrow="19" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFEE19BA6FEAD75A4" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[327,371,916,936]" class="Amphibia" family="Bufonidae" genus="Bufo" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D076FFAAFEE19BA6FEAD75A4" box="[327,371,916,936]" italics="true" pageId="34" pageNumber="35">Bufo</emphasis>
</taxonomicName>
gr.
<emphasis id="B9284B61D076FFAAFE309BA6FE6175A4" box="[406,447,916,936]" italics="true" pageId="34" pageNumber="35">bufo</emphasis>
</th>
<td id="76BDFC4DD0760077FC179BA6FBF375A4" box="[945,1069,916,936]" gridcol="2" gridrow="19" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE19B8BFAD375C1" box="[327,1293,953,973]" gridrow="20" pageId="34" pageNumber="35" rowspan-2="1" rowspan-3="1">
<th id="76BDFC4DD0760077FEE19B8BFDB775C1" box="[327,617,953,973]" gridcol="0" gridrow="20" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFEE19B8BFE0D75C1" box="[327,467,953,973]" class="Amphibia" family="Ranidae" genus="Pelophylax" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFEE19B8BFE7175C1" box="[327,431,953,973]" italics="true" pageId="34" pageNumber="35">Pelophylax</emphasis>
sp.
</taxonomicName>
</th>
<td id="76BDFC4DD0760077FD079B8BFCE175C1" box="[673,831,953,973]" gridcol="1" gridrow="20" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE19BEDFAD375FF" box="[327,1293,991,1011]" gridrow="21" pageId="34" pageNumber="35" rowspan-2="1" rowspan-3="1">
<th id="76BDFC4DD0760077FEE19BEDFDB775FF" box="[327,617,991,1011]" gridcol="0" gridrow="21" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFEE19BEDFE4375FF" box="[327,413,991,1011]" class="Amphibia" family="Ranidae" genus="Rana" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFEE19BEDFEA475FF" box="[327,378,991,1011]" italics="true" pageId="34" pageNumber="35">Rana</emphasis>
sp.
</taxonomicName>
</th>
<td id="76BDFC4DD0760077FD079BEDFCE175FF" box="[673,831,991,1011]" gridcol="1" gridrow="21" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE19C36FAD37214" box="[327,1293,1028,1048]" gridrow="22" pageId="34" pageNumber="35" rowspan-3="1">
<th id="76BDFC4DD0760077FEE19C36FDB77214" box="[327,617,1028,1048]" gridcol="0" gridrow="22" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFEE19C36FE487214" authorityName="Batsch" authorityYear="1796" box="[327,406,1028,1048]" class="Amphibia" family="Ranidae" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="family">Ranidae</taxonomicName>
indet.
</th>
<td id="76BDFC4DD0760077FD079C36FCE17214" box="[673,831,1028,1048]" gridcol="1" gridrow="22" pageId="34" pageNumber="35">x</td>
<td id="76BDFC4DD0760077FC179C36FBF37214" box="[945,1069,1028,1048]" gridcol="2" gridrow="22" pageId="34" pageNumber="35">x</td>
</tr>
<tr id="356C9531D0760077FEE19C1BFAD37231" box="[327,1293,1065,1085]" gridrow="23" pageId="34" pageNumber="35">
<th id="76BDFC4DD0760077FEE19C1BFDB77231" box="[327,617,1065,1085]" gridcol="0" gridrow="23" pageId="34" pageNumber="35">
<taxonomicName id="4C5CECF0D076FFAAFEE19C1BFE5E7231" authorityName="Fischer" authorityYear="1813" box="[327,384,1065,1085]" class="Amphibia" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="order">Anura</taxonomicName>
indet.
</th>
<td id="76BDFC4DD0760077FD079C1BFCE17231" box="[673,831,1065,1085]" gridcol="1" gridrow="23" pageId="34" pageNumber="35">x</td>
<td id="76BDFC4DD0760077FC179C1BFBF37231" box="[945,1069,1065,1085]" gridcol="2" gridrow="23" pageId="34" pageNumber="35">x</td>
<td id="76BDFC4DD0760077FB279C1BFAD37231" box="[1153,1293,1065,1085]" gridcol="3" gridrow="23" pageId="34" pageNumber="35">x</td>
</tr>
</table>
</paragraph>
<paragraph id="8BE39773D076FFAAFF5C9E6FFC00717A" blockId="34.[192,793,1149,1910]" lastBlockId="34.[840,1441,1149,1910]" pageId="34" pageNumber="35">
Nevertheless, there are also amphibians clearly persisting in Hambach in both Middle Miocene and Late Pliocene times. These include at least two urodeles,
<taxonomicName id="4C5CECF0D076FFAAFE159E8FFCC670DA" baseAuthorityName="Vasilyan and Yanenko" baseAuthorityYear="2020" box="[435,792,1725,1750]" class="Amphibia" family="Batrachosauroididae" genus="Palaeoproteus" higherTaxonomySource="GBIF" isUncertain="true" kingdom="Animalia" order="Caudata" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="miocenicus">
<emphasis id="B9284B61D076FFAAFE159E8FFD8170DA" box="[435,607,1725,1750]" italics="true" pageId="34" pageNumber="35">Palaeoproteus</emphasis>
cf.
<emphasis id="B9284B61D076FFAAFD359E8FFCC670DA" box="[659,792,1725,1750]" italics="true" pageId="34" pageNumber="35">miocenicus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C5CECF0D076FFAAFF5E9EEFFE7A70FA" box="[248,420,1757,1782]" class="Amphibia" family="Salamandridae" genus="Lissotriton" kingdom="Animalia" order="Caudata" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFF5E9EEFFEAF70FA" box="[248,369,1757,1782]" italics="true" pageId="34" pageNumber="35">Lissotriton</emphasis>
sp.
</taxonomicName>
, and two anurans,
<taxonomicName id="4C5CECF0D076FFAAFD349EEFFCC770FA" box="[658,793,1757,1782]" class="Amphibia" family="Alytidae" genus="Latonia" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFD349EEFFD3770FA" box="[658,745,1757,1782]" italics="true" pageId="34" pageNumber="35">Latonia</emphasis>
sp.
</taxonomicName>
and
<taxonomicName id="4C5CECF0D076FFAAFF5C9ECFFEBF711A" box="[250,353,1789,1814]" class="Amphibia" family="Hylidae" genus="Hyla" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFF5C9ECFFEF1711A" box="[250,303,1789,1814]" italics="true" pageId="34" pageNumber="35">Hyla</emphasis>
sp.
</taxonomicName>
As mentioned, ranids can also be added to this list, even though detailed identification of the Pliocene remains at specific rank is prevented. Among frogs, palaeobatrachids and pelobatids show the reverse situation, with Late Pliocene remains that can be identified at species
<emphasis id="B9284B61D076FFAAFCEE9C8FFC3572DB" box="[840,1003,1213,1239]" italics="true" pageId="34" pageNumber="35">
(
<taxonomicName id="4C5CECF0D076FFAAFCF79C8FFC3C72DB" box="[849,994,1213,1239]" class="Amphibia" family="Palaeobatrachidae" genus="Palaeobatrachus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="eurydices">P. eurydices</taxonomicName>
)
</emphasis>
or, possibly, genus (cf.
<taxonomicName id="4C5CECF0D076FFAAFAAA9C8FFCB272FA" class="Amphibia" family="Pelobatidae" genus="Eopelobates" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFAAA9C8FFA7E72DA" box="[1292,1440,1213,1238]" italics="true" pageId="34" pageNumber="35">Eopelobates</emphasis>
sp.
</taxonomicName>
and?
<taxonomicName id="4C5CECF0D076FFAAFC699CEFFB4372FA" box="[975,1181,1245,1270]" class="Amphibia" family="Pelobatidae" genus="Eopelobates" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D076FFAAFC699CEFFBBD72FA" box="[975,1123,1245,1270]" italics="true" pageId="34" pageNumber="35">Eopelobates</emphasis>
sp.
</taxonomicName>
) ranks respectively, whereas only the general presence of the families can be reported for the Middle Miocene (even in a tentative way in the case of palaeobatrachids). On the opposed side, the toad
<taxonomicName id="4C5CECF0D076FFAAFB2F9D6FFB1E737A" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[1161,1216,1373,1398]" class="Amphibia" family="Bufonidae" genus="Bufo" kingdom="Animalia" order="Anura" pageId="34" pageNumber="35" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D076FFAAFB2F9D6FFB1E737A" box="[1161,1216,1373,1398]" italics="true" pageId="34" pageNumber="35">Bufo</emphasis>
</taxonomicName>
gr.
<emphasis id="B9284B61D076FFAAFB4B9D6FFAFF737A" box="[1261,1313,1373,1398]" italics="true" pageId="34" pageNumber="35">bufo</emphasis>
is the only anuran whose presence is certified just for the Late Pliocene in Hambach. Therefore, considering the uncertainty surrounding the possible persistence of some taxa from the Middle Miocene, the composition of the Late Pliocene Hambach 11 and 11C amphibian assemblage is certainly impoverished compared to the older one from Hambach 6C, apparently mainly as far as the urodele component is concerned, but it appears to exhibit an overall similarity and continuity. This differs from other tetrapod groups, which are sporadically represented by the same or related taxa in both Miocene and Pliocene levels (e.g., some petauristid rodents and possibly the already-mentioned
<taxonomicName id="4C5CECF0D076FFAAFB779F2FFAB4713B" authorityName="Peters" authorityYear="1868" box="[1233,1386,1821,1847]" class="Testudines" family="Chelydridae" genus="Chelydropsis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D076FFAAFB779F2FFAB4713B" box="[1233,1386,1821,1847]" italics="true" pageId="34" pageNumber="35">Chelydropsis</emphasis>
</taxonomicName>
turtles;
<bibRefCitation id="EFCDEA82D076FFAAFC279F0FFB91715A" author="Mors, T &amp; von der Hocht, F. &amp; Wutzler, B." box="[897,1103,1853,1878]" pageId="34" pageNumber="35" pagination="145 - 170" refId="ref35109" refString="Mors, T, von der Hocht, F., and Wutzler, B. 2000. Die erste Wirbeltierfauna aus der miozanen Braunkohle der Niederrheinischen Bucht (Ville-Schichten, Tagebau Hambach). Palaontologische Zeitschrift, 74: 145 - 170. https: // doi. org / 10.1007 / BF 02987958" type="journal article" year="2000">Mörs et al., 2000</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D076FFAAFBFD9F0FFB3B715A" author="Mors, T." box="[1115,1253,1853,1878]" pageId="34" pageNumber="35" pagination="177 - 183" refId="ref34899" refString="Mors, T. 2002. Biostratigraphy and paleoecology of continental Tertiary vertebrate faunas in the Lower Rhine Embayment (NW-Germany). Netherlands Journal of Geosciences - Geologie en Mijnbouw, 81: 177 - 183. https: // doi. org / 10.1017 / S 0016774600022411" type="journal article" year="2002">Mörs, 2002</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D076FFAAFB579F0FFC0A717A" author="Van Laere, G. &amp; Mors, T." pageId="34" pageNumber="35" pagination="223 - 241" refId="ref38874" refString="Van Laere, G. and Mors, T. 2023. Beavers and flying squirrels (Rodentia: Castoridae, Pteromyini) from the Late Pliocene of Hambach 11 C, Germany. Geodiversitas, 45: 223 - 241. https: // doi. org / 10.5252 / geodiversitas 2023 v 45 a 7" type="journal article" year="2023">
<collectingRegion id="49985991D076FFAAFB579F0FFAC0715A" box="[1265,1310,1853,1878]" country="Turkey" name="Van" pageId="34" pageNumber="35">Van</collectingRegion>
Laere and Mörs, 2023
</bibRefCitation>
).
</paragraph>
<paragraph id="8BE39773D077FFABFF5C989BFE8575CE" blockId="35.[192,793,169,962]" pageId="35" pageNumber="36">
In terms of abundance, the highest numbers of remains identified per taxon in the Miocene assemblage are those referred to
<taxonomicName id="4C5CECF0D077FFABFDCE98DAFCC7770E" box="[616,793,232,258]" class="Amphibia" family="Salamandridae" genus="Chelotriton" kingdom="Animalia" order="Caudata" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D077FFABFDCE98DAFD37770D" box="[616,745,232,257]" italics="true" pageId="35" pageNumber="36">Chelotriton</emphasis>
sp.
</taxonomicName>
(57 remains) and
<taxonomicName id="4C5CECF0D077FFABFE31993BFDC2772E" box="[407,540,265,290]" class="Amphibia" family="Alytidae" genus="Latonia" kingdom="Animalia" order="Anura" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D077FFABFE31993BFE30772E" box="[407,494,265,290]" italics="true" pageId="35" pageNumber="36">Latonia</emphasis>
sp.
</taxonomicName>
(37 remains). These are, therefore, the taxa dominating the oldest Hambach oryctocenosis.
<taxonomicName id="4C5CECF0D077FFABFE12997BFDD5776E" authorityName="von Meyer" authorityYear="1843" box="[436,523,329,354]" class="Amphibia" family="Alytidae" genus="Latonia" kingdom="Animalia" order="Anura" pageId="35" pageNumber="36" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D077FFABFE12997BFDD5776E" box="[436,523,329,354]" italics="true" pageId="35" pageNumber="36">Latonia</emphasis>
</taxonomicName>
persisted as dominant also in the Pliocene with 22 remains (18 from Hambach 11 and six from Hambach 11C), following
<taxonomicName id="4C5CECF0D077FFABFCA499BBFEEC77CE" class="Amphibia" family="Palaeobatrachidae" genus="Palaeobatrachus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="eurydices">
<emphasis id="B9284B61D077FFABFCA499BBFEEC77CE" italics="true" pageId="35" pageNumber="36">P. eurydices</emphasis>
</taxonomicName>
represented by 27 remains (23 and four from Hambach 11 and 11C, respectively). As already mentioned,
<taxonomicName id="4C5CECF0D077FFABFE0F99DAFDF4740D" authorityName="Pomel" authorityYear="1853" box="[425,554,488,513]" class="Amphibia" family="Salamandridae" genus="Chelotriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="35" pageNumber="36" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D077FFABFE0F99DAFDF4740D" box="[425,554,488,513]" italics="true" pageId="35" pageNumber="36">Chelotriton</emphasis>
</taxonomicName>
is not present in the Pliocene 11 and 11C sites, in agreement with a general European trend of extinction of this newt. The only Pliocene remains referred to this genus come in fact from the Upper Pliocene of southern
<collectingCountry id="F34BD7E3D077FFABFF669ABBFECD74AE" box="[192,275,649,674]" name="France" pageId="35" pageNumber="36">France</collectingCountry>
(
<bibRefCitation id="EFCDEA82D077FFABFE879ABBFE6574AE" author="Bailon, S." box="[289,443,649,674]" pageId="35" pageNumber="36" pagination="7 - 28" refId="ref29771" refString="Bailon, S. 1989. Les amphibiens et les reptiles du Pliocene Superieur de Balaruc II (Herault, France). Palaeovertebrata, 19: 7 - 28." type="journal article" year="1989">Bailon, 1989</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFE609ABBFD0774AE" author="Macaluso, L. &amp; Mannion, P. D. &amp; Evans, S. E. &amp; Carnevale, G. &amp; Monti, S. &amp; Marchitelli, D. &amp; Delfino, M." box="[454,729,649,674]" pageId="35" pageNumber="36" pagination="220935" refId="ref34329" refString="Macaluso, L., Mannion, P. D., Evans, S. E., Carnevale, G., Monti, S., Marchitelli, D., and Delfino, M. 2022 a. Biogeographic history of Palearctic caudates revealed by a critical appraisal of their fossil record quality and spatio-temporal distribution. Royal Society Open Science, 9: 220935. https: // doi. org / 10.1098 / rsos. 220935" type="journal article" year="2022">Macaluso et al., 2022a</bibRefCitation>
). The most-represented urodele in the Upper Pliocene of Hambach is
<taxonomicName id="4C5CECF0D077FFABFEFC9AFBFD8C74EE" box="[346,594,712,738]" class="Amphibia" family="Proteidae" genus="Mioproteus" higherTaxonomySource="GBIF" isUncertain="true" kingdom="Animalia" order="Caudata" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="wezei">
<emphasis id="B9284B61D077FFABFEFC9AFBFE0074EE" box="[346,478,713,738]" italics="true" pageId="35" pageNumber="36">Mioproteus</emphasis>
cf.
<emphasis id="B9284B61D077FFABFDA89AFBFD8C74EE" box="[526,594,713,738]" italics="true" pageId="35" pageNumber="36">wezei</emphasis>
</taxonomicName>
, represented by 15 remains (nine and six, respectively, from Hambach 11 and 11C). Other taxa from both the Miocene and Pliocene assemblages are only represented by very few, less than 10 remains each (except for
<taxonomicName id="4C5CECF0D077FFABFE2F9B5BFDCC758E" box="[393,530,873,898]" class="Amphibia" family="Salamandridae" genus="Triturus" kingdom="Animalia" order="Caudata" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D077FFABFE2F9B5BFE3F758E" box="[393,481,873,898]" italics="true" pageId="35" pageNumber="36">Triturus</emphasis>
sp.
</taxonomicName>
, with 12). Thus, these represent only very minor components of the fossil associations.
</paragraph>
<paragraph id="8BE39773D077FFABFF669BEBFDA3721E" blockId="35.[192,728,984,1042]" pageId="35" pageNumber="36">
<emphasis id="B9284B61D077FFABFF669BEBFDA3721E" bold="true" pageId="35" pageNumber="36">Palaeobiogeographical Significance of the Hambach Amphibian Assemblages</emphasis>
</paragraph>
<paragraph id="8BE39773D077FFABFF5C9C1BFD96736E" blockId="35.[192,793,1064,1890]" pageId="35" pageNumber="36">
The amphibian fossil record from Hambach already proved to host unexpected surprises with the discovery of
<taxonomicName id="4C5CECF0D077FFABFE239C5BFDAE728E" authorityName="Macaluso, Villa, and Mors" authorityYear="2022" box="[389,624,1129,1154]" class="Amphibia" family="Proteidae" genus="Euronecturus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="grogu">
<emphasis id="B9284B61D077FFABFE239C5BFDAE728E" box="[389,624,1129,1154]" italics="true" pageId="35" pageNumber="36">Euronecturus grogu</emphasis>
</taxonomicName>
, an enigmatic proteid (see
<bibRefCitation id="EFCDEA82D077FFABFEF89CBBFDA272AE" author="Macaluso, L. &amp; Villa, A. &amp; Mors, T." box="[350,636,1161,1186]" pageId="35" pageNumber="36" pagination="12585" refId="ref34411" refString="Macaluso, L., Villa, A., and Mors, T. 2022 b. A new proteid salamander (Urodela; Proteidae) from the middle Miocene of Hambach (Germany) and implications for the evolution of the family. Palaeontology, 65: e 12585. https: // doi. org / 10.1111 / pala. 12585" type="journal article" year="2022">Macaluso et al., 2022b</bibRefCitation>
) that is only known from its
<typeStatus id="54E729D1D077FFABFEDE9C9BFE7472CE" box="[376,426,1193,1218]" pageId="35" pageNumber="36">type</typeStatus>
locality of Hambach 6C at the moment. The extended study of the whole amphibian record revealed that this is not the only noteworthy taxon preserved in the site, and that significant findings were hidden in both the Miocene and Pliocene assemblages.
</paragraph>
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Cryptobranchids are in general poorly represented in the fossil record, even though a particular concentration of Neogene cryptobranchid-bearing sites is found in central Europe, and more specifically in Southern
<collectingCountry id="F34BD7E3D077FFABFE3F9DDAFDD9700E" box="[409,519,1512,1538]" name="Germany" pageId="35" pageNumber="36">Germany</collectingCountry>
(
<bibRefCitation id="EFCDEA82D077FFABFDBF9DDBFCD1700E" author="Bohme, M. &amp; Vasilyan, D. &amp; Winklhofer, M." box="[537,783,1513,1538]" pageId="35" pageNumber="36" pagination="64 - 72" refId="ref30639" refString="Bohme, M., Vasilyan, D., and Winklhofer, M. 2012. Habitat tracking, range dynamics and palaeoclimatic significance of Eurasian giant salamanders (Cryptobranchidae) - indications for elevated Central Asian humidity during Cenozoic global warm periods. Palaeogeography, Palaeoclimatology, Palaeoecology, 342 - 343: 64 - 72. https: // doi. org / 10.1016 / j. palaeo. 2012.04.032" type="journal article" year="2012">Böhme et al., 2012</bibRefCitation>
). Still, they are almost unknown from the European early Middle Miocene, even though their presence in Lower, upper Middle and Upper Miocene sites (
<bibRefCitation id="EFCDEA82D077FFABFF609E5BFE51708E" author="Westphal, F." box="[198,399,1641,1666]" pageId="35" pageNumber="36" pagination="20 - 92" refId="ref40178" refString="Westphal, F. 1958. Die Tertiaren und rezenten Eurasiatischen Riesensalamander. Palaeontolographica Abteilung A, 110: 20 - 92." type="journal article" year="1958">Westphal, 1958</bibRefCitation>
,
<bibRefCitation id="EFCDEA82D077FFABFE029E5BFE3D708E" author="Westphal, F." box="[420,483,1641,1666]" pageId="35" pageNumber="36" pagination="253 - 260" refId="ref40237" refString="Westphal, F. 1970. Neue Riesensalamander-Funde (Andrias, Amphibia) aus der Oberen Susswassermolasse von Wartenberg in Bayern. Mitteilungen der Bayerischen Staatssammlung fur Palaontologie und historische Geologie, 10: 253 - 260." type="journal article" year="1970">1970</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFE519E5BFD6F708E" author="Bottcher, R." box="[503,689,1641,1666]" pageId="35" pageNumber="36" pagination="1 - 38" refId="ref30815" refString="Bottcher, R. 1987. Neue Funde von Andrias scheuchzeri (Cryptobranchidae, Amphibia) aus der suddeutschen Molasse. Stuttgarter Beitrage zur Naturkunde B, 131: 1 - 38." type="journal article" year="1987">Böttcher, 1987</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFD639E5BFF2170AE" author="Miklas, P. M." pageId="35" pageNumber="36" pagination="161 - 211" refId="ref34749" refString="Miklas, P. M. 2002. Die Amphibienfauna (Amphibia: Caudata, Anura) der obermiozanen Fundstelle Gotzendorf an der Leitha (sudliches Wiener Becken, Niederosterreich). Annalen des Naturhistorischen Museums in Wien, 103 A: 161 - 211." type="journal article" year="2002">Miklas, 2002</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFEAD9EBBFE7170AE" author="Bohme, M." box="[267,431,1673,1698]" pageId="35" pageNumber="36" pagination="389 - 401" refId="ref30441" refString="Bohme, M. 2003. Miocene Climatic Optimum: evidence from lower vertebrates of central Europe. Palaeogeography, Palaeoclimatology, Palaeoecology, 195: 389 - 401. https: // doi. org / 10.1016 / S 0031 - 0182 (03) 00367 - 5" type="journal article" year="2003">Böhme, 2003</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFE1D9EBAFD7870AE" author="Kvacek, Z. &amp; Bohme, M. &amp; Dvorak, Z. &amp; Konzalova, M. &amp; Mach, K. &amp; Prokop, J. &amp; Rajchl, M." box="[443,678,1672,1698]" pageId="35" pageNumber="36" pagination="1 - 40" refId="ref33850" refString="Kvacek, Z., Bohme, M., Dvorak, Z., Konzalova, M., Mach, K., Prokop, J., and Rajchl, M. 2004. Early Miocene freshwater and swamp ecosystems of the Most Basin (northern Bohemia) with particular reference to the Bilina Mine section. Journal of the Czech Geological Society, 49: 1 - 40." type="journal article" year="2004">Kvaček et al., 2004</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFD149EBAFF2170CE" author="Tempfer, P. M." pageId="35" pageNumber="36" pagination="257 - 268" refId="ref38328" refString="Tempfer, P. M. 2004. Andrias scheuchzeri (Caudata: Cryptobranchidae) from the Upper Miocene (MN 7 / 8) locality Mataschen / Styria. Joannea Geologie &amp; Palaontologie, 5: 257 - 268." type="journal article" year="2004">Tempfer, 2004</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFEA99E9BFDDD70CE" author="Bohme, M. &amp; Vasilyan, D. &amp; Winklhofer, M." box="[271,515,1705,1730]" pageId="35" pageNumber="36" pagination="64 - 72" refId="ref30639" refString="Bohme, M., Vasilyan, D., and Winklhofer, M. 2012. Habitat tracking, range dynamics and palaeoclimatic significance of Eurasian giant salamanders (Cryptobranchidae) - indications for elevated Central Asian humidity during Cenozoic global warm periods. Palaeogeography, Palaeoclimatology, Palaeoecology, 342 - 343: 64 - 72. https: // doi. org / 10.1016 / j. palaeo. 2012.04.032" type="journal article" year="2012">Böhme et al., 2012</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFDB59E9BFCCB70CE" author="Vasilyan, D. &amp; Bohme, M. &amp; Chkhikvadze, V. M. &amp; Semenov, Y. A. &amp; Joyce, W. G." box="[531,789,1705,1730]" pageId="35" pageNumber="36" pagination="301 - 318" refId="ref38977" refString="Vasilyan, D., Bohme, M., Chkhikvadze, V. M., Semenov, Y. A., and Joyce, W. G. 2013. A new giant salamander (Urodela, Pancryptobrancha) from the Miocene of eastern Europe (Grytsiv, Ukraine). Journal of Vertebrate Paleontology, 33: 301 - 318. https: // doi. org / 10.1080 / 02724634.2013.722151" type="journal article" year="2013">Vasilyan et al., 2013</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFF669EFAFE8E70EE" author="Sach, V. J." box="[192,336,1736,1762]" pageId="35" pageNumber="36" pagination="1 - 115" refId="ref36429" refString="Sach, V. J. 2016. Fossilienkatalog der Miozan-Molasse in Sudwestdeutschland (Obere Susswassermolasse, Brackwassermolasse, Obere Meeresmolasse, Untere Susswassermolasse). Documenta Naturae, SB 70: 1 - 115." type="journal article" year="2016">Sach, 2016</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFEC69EFAFDA170EE" author="Georgalis, G. L. &amp; Villa, A. &amp; Ivanov, M. &amp; Vasilyan, D. &amp; Delfino, M." box="[352,639,1736,1762]" pageId="35" pageNumber="36" pagination="1 - 99" refId="ref32530" refString="Georgalis, G. L., Villa, A., Ivanov, M., Vasilyan, D., and Delfino, M. 2019 b. Fossil amphibians and reptiles from the Neogene locality of Maramena (Greece), the most diverse European herpetofauna at the Miocene / Pliocene transition boundary. Palaeontologia Electronica, 22.3.68: 1 - 99. https: // doi. org / 10.26879 / 908" type="book chapter" year="2019">Georgalis et al., 2019b</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFD299EFAFEF4710E" author="Szentesi, Z. &amp; Sebe, K. &amp; Szabo, M." pageId="35" pageNumber="36" pagination="353 - 366" refId="ref38136" refString="Szentesi, Z., Sebe, K., and Szabo, M. 2020. Giant salamander from the Miocene of the Mecsek mountains (Pecs-Danitzpuszta, southwestern Hungary). PalZ, 94: 353 - 366. https: // doi. org / 10.1007 / s 12542 - 019 - 00499 - 2" type="journal article" year="2020">Szentesi et al., 2020</bibRefCitation>
) suggests their continuous persistence in the continent. As a matter of fact, Hambach 6C is currently the only known occurrence of a cryptobranchid from the early Middle Miocene time frame in Europe, and even adds to a limited Eurasian Langhian record that further includes only few localities in
<collectingCountry id="F34BD7E3D077FFABFC5098DBFB5F770E" box="[1014,1153,233,258]" name="Kazakhstan" pageId="35" pageNumber="36">Kazakhstan</collectingCountry>
(
<bibRefCitation id="EFCDEA82D077FFABFB0798DAFA42770E" author="Chkhikvadze, V. M." box="[1185,1436,232,258]" pageId="35" pageNumber="36" pagination="63 - 67" refId="ref31121" refString="Chkhikvadze, V. M. 1982. On the findings of fossil Cryptobranchidae in the USSR and Mongolia. Vertebrata Hungarica, 21: 63 - 67." type="journal article" year="1982">Chkhikvadze, 1982</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFCEE993BFBB4772E" author="Bohme, M. &amp; Vasilyan, D. &amp; Winklhofer, M." box="[840,1130,265,290]" pageId="35" pageNumber="36" pagination="64 - 72" refId="ref30639" refString="Bohme, M., Vasilyan, D., and Winklhofer, M. 2012. Habitat tracking, range dynamics and palaeoclimatic significance of Eurasian giant salamanders (Cryptobranchidae) - indications for elevated Central Asian humidity during Cenozoic global warm periods. Palaeogeography, Palaeoclimatology, Palaeoecology, 342 - 343: 64 - 72. https: // doi. org / 10.1016 / j. palaeo. 2012.04.032" type="journal article" year="2012">Böhme et al., 2012</bibRefCitation>
). Cryptobranchids later declined in Europe during the Pliocene, with their last occurrence at Willershausen, in central
<collectingCountry id="F34BD7E3D077FFABFACD997AFC57778E" name="Germany" pageId="35" pageNumber="36">Germany</collectingCountry>
(
<bibRefCitation id="EFCDEA82D077FFABFC3D995BFB83778E" author="Westphal, F." box="[923,1117,361,386]" pageId="35" pageNumber="36" pagination="67 - 73" refId="ref40201" refString="Westphal, F. 1967. Erster Nachweis des Riesensalamanders (Andrias, Urodela, Amphibia) im europaischen Jungpliozan. Neues Jahrbuch fur Geologie und Palaontologie - Monatshefte, 1967: 67 - 73." type="journal article" year="1967">Westphal, 1967</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFBCA995BFA80778E" author="Bohme, M. &amp; Vasilyan, D. &amp; Winklhofer, M." box="[1132,1374,361,386]" pageId="35" pageNumber="36" pagination="64 - 72" refId="ref30639" refString="Bohme, M., Vasilyan, D., and Winklhofer, M. 2012. Habitat tracking, range dynamics and palaeoclimatic significance of Eurasian giant salamanders (Cryptobranchidae) - indications for elevated Central Asian humidity during Cenozoic global warm periods. Palaeogeography, Palaeoclimatology, Palaeoecology, 342 - 343: 64 - 72. https: // doi. org / 10.1016 / j. palaeo. 2012.04.032" type="journal article" year="2012">Böhme et al., 2012</bibRefCitation>
). The age of Willershausen was reported as Zanclean by
<bibRefCitation id="EFCDEA82D077FFABFCEE999BFBEA77CE" author="Bohme, M. &amp; Vasilyan, D. &amp; Winklhofer, M." box="[840,1076,425,450]" pageId="35" pageNumber="36" pagination="64 - 72" refId="ref30639" refString="Bohme, M., Vasilyan, D., and Winklhofer, M. 2012. Habitat tracking, range dynamics and palaeoclimatic significance of Eurasian giant salamanders (Cryptobranchidae) - indications for elevated Central Asian humidity during Cenozoic global warm periods. Palaeogeography, Palaeoclimatology, Palaeoecology, 342 - 343: 64 - 72. https: // doi. org / 10.1016 / j. palaeo. 2012.04.032" type="journal article" year="2012">Böhme et al. (2012</bibRefCitation>
, table 1), but it is now considered as Piacenzian (
<bibRefCitation id="EFCDEA82D077FFABFB9E99FAFAFD77EE" author="Kolibac, J. &amp; Adroit, B. &amp; Groning, E. &amp; Brauckmann, C. &amp; Wappler, T." box="[1080,1315,456,482]" pageId="35" pageNumber="36" pagination="681 - 689" refId="ref33729" refString="Kolibac, J., Adroit, B., Groning, E., Brauckmann, C., and Wappler, T. 2016. First record of the family Trogossitidae (Insecta, Coleoptera) in the Late Pliocene deposits of Willershausen (Germany). PalZ, 90: 681 - 689. https: // doi. org / 10.1007 / s 12542 - 016 - 0316 - 6" type="journal article" year="2016">Kolibáč et al., 2016</bibRefCitation>
, and reference therein). Adding to being the only data currently available on European early Middle Miocene cryptobranchids, Hambach 6C represents also the westernmost point ever reached by these urodeles in Eurasia, based on our current knowledge. Only the late Oligocene occurrence at Rott (
<bibRefCitation id="EFCDEA82D077FFABFCE89A9BFBD074CE" author="Westphal, F." box="[846,1038,681,706]" pageId="35" pageNumber="36" pagination="20 - 92" refId="ref40178" refString="Westphal, F. 1958. Die Tertiaren und rezenten Eurasiatischen Riesensalamander. Palaeontolographica Abteilung A, 110: 20 - 92." type="journal article" year="1958">Westphal, 1958</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFBBC9A9BFB7D74CE" author="Mors, T." box="[1050,1187,681,706]" pageId="35" pageNumber="36" pagination="177 - 183" refId="ref34899" refString="Mors, T. 2002. Biostratigraphy and paleoecology of continental Tertiary vertebrate faunas in the Lower Rhine Embayment (NW-Germany). Netherlands Journal of Geosciences - Geologie en Mijnbouw, 81: 177 - 183. https: // doi. org / 10.1017 / S 0016774600022411" type="journal article" year="2002">Mörs, 2002</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFB099A9BFA4874CE" author="Bohme, M. &amp; Vasilyan, D. &amp; Winklhofer, M." box="[1199,1430,681,706]" pageId="35" pageNumber="36" pagination="64 - 72" refId="ref30639" refString="Bohme, M., Vasilyan, D., and Winklhofer, M. 2012. Habitat tracking, range dynamics and palaeoclimatic significance of Eurasian giant salamanders (Cryptobranchidae) - indications for elevated Central Asian humidity during Cenozoic global warm periods. Palaeogeography, Palaeoclimatology, Palaeoecology, 342 - 343: 64 - 72. https: // doi. org / 10.1016 / j. palaeo. 2012.04.032" type="journal article" year="2012">Böhme et al., 2012</bibRefCitation>
), close but located slightly eastward to Hambach, further supports the past existence of cryptobranchid populations west of the Rhine Graben.
</paragraph>
<paragraph id="8BE39773D077FFACFC249B1AFEE874F3" blockId="35.[840,1441,169,1889]" lastBlockId="36.[192,793,166,1887]" lastPageId="36" lastPageNumber="37" pageId="35" pageNumber="36">
One of the most unexpected findings from Hambach is the batrachosauroidid
<taxonomicName id="4C5CECF0D077FFABFB4B9B7BFA47756E" baseAuthorityName="Vasilyan and Yanenko" baseAuthorityYear="2020" box="[1261,1433,841,866]" class="Amphibia" family="Batrachosauroididae" genus="Palaeoproteus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="35" pageNumber="36" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D077FFABFB4B9B7BFA47756E" box="[1261,1433,841,866]" italics="true" pageId="35" pageNumber="36">Palaeoproteus</emphasis>
</taxonomicName>
. Batrachosauroidids are enigmatic urodeles, which are mainly known from the Cretaceous and Palaeogene of North America (
<bibRefCitation id="EFCDEA82D077FFABFB319B9AFC5975EE" author="Gardner, J. D. &amp; DeMar, D. G., Jr." pageId="35" pageNumber="36" pagination="459 - 515" refId="ref32210" refString="Gardner, J. D. and DeMar, D. G., Jr. 2013. Mesozoic and Palaeocene lissamphibian assemblages of North America: a comprehensive review. Palaeobiodiversity and Palaeoenvironments, 93: 459 - 515. https: // doi. org / 10.1007 / s 12549 - 013 - 0130 - z" type="journal article" year="2013">Gardner and DeMar, 2013</bibRefCitation>
, and reference therein). They persisted in the
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throughout the Miocene, with a handful of occurrences in
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(
<bibRefCitation id="EFCDEA82D077FFABFBD89C3BFAC2722E" author="Naylor, B. G." box="[1150,1308,1033,1058]" pageId="35" pageNumber="36" pagination="1 - 14" refId="ref35164" refString="Naylor, B. G. 1981. A new salamander of the family Batrachosauroididae from the Late Miocene of North America, with notes on other batrachosauroidids. PaleoBios, 39: 1 - 14." type="journal article" year="1981">Naylor, 1981</bibRefCitation>
),
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(
<bibRefCitation id="EFCDEA82D077FFABFCE89C1BFB4A724E" author="Weems, R. E. &amp; George, R. A." box="[846,1172,1064,1090]" pageId="35" pageNumber="36" pagination="570 - 588" refId="ref40084" refString="Weems, R. E. and George, R. A. 2013. Amphibians and nonmarine turtles from the Miocene Calvert Formation of Delaware, Maryland, and Virginia (USA). Journal of Paleontology, 87: 570 - 588. https: // doi. org / 10.1666 / 12 - 071" type="journal article" year="2013">Weems and George, 2013</bibRefCitation>
),
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(
<bibRefCitation id="EFCDEA82D077FFABFAAE9C1BFA42724E" author="Estes, R." box="[1288,1436,1065,1090]" pageId="35" pageNumber="36" pagination="234 - 256" refId="ref31673" refString="Estes, R. 1963. Early Miocene salamanders and lizards from Florida. Quarterly Journal of the Florida Academy of Sciences, 26: 234 - 256." type="journal article" year="1963">Estes, 1963</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFCEE9C7BFC2E726E" author="Bryant, D. J." box="[840,1008,1097,1122]" pageId="35" pageNumber="36" pagination="472 - 489" refId="ref30963" refString="Bryant, D. J. 1991. New early Barstovian (middle Miocene) vertebrates from the upper Torreya Formation, eastern Florida panhandle. Journal of Vertebrate Paleontology, 11: 472 - 489. https: // doi. org / 10.1080 / 02724634.1991.10011416" type="journal article" year="1991">Bryant, 1991</bibRefCitation>
),
<collectingRegion id="49985991D077FFABFBAA9C7BFBA0726E" box="[1036,1150,1097,1122]" country="United States of America" name="Louisiana" pageId="35" pageNumber="36">Louisiana</collectingRegion>
(
<bibRefCitation id="EFCDEA82D077FFABFB319C7BFA86726E" author="Williams, M. J." box="[1175,1368,1097,1122]" pageId="35" pageNumber="36" refId="ref40275" refString="Williams, M. J. 2009. Miocene herpetofaunas from the central Gulf Coast USA: their paleoecology, biogeography, and biostratigraphy. Unpublished PhD Thesis, Louisiana State University and Agricultural and Mechanical College, Baton Rouge, Lousiana, U. S. A." type="book" year="2009">Williams, 2009</bibRefCitation>
), and
<collectingRegion id="49985991D077FFABFCEE9C5BFC50728E" box="[840,910,1129,1154]" country="United States of America" name="Texas" pageId="35" pageNumber="36">Texas</collectingRegion>
(
<bibRefCitation id="EFCDEA82D077FFABFC3B9C5BFB61728E" author="Taylor, E. H. &amp; Hesse, C. J." box="[925,1215,1129,1154]" pageId="35" pageNumber="36" pagination="185 - 193" refId="ref38286" refString="Taylor, E. H. and Hesse, C. J. 1943. A new salamander from the upper Miocene beds of San Jacinto County, Texas. The American Journal of Science, 241: 185 - 193." type="journal article" year="1943">Taylor and Hesse, 1943</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFB6D9C5AFA42728E" author="Auffenberg, W." box="[1227,1436,1128,1154]" pageId="35" pageNumber="36" pagination="169 - 176" refId="ref29635" refString="Auffenberg, W. 1958. A new family of Miocene salamanders from the Texas coastal plain. Quarterly Journal of the Florida Academy of Sciences, 21: 169 - 176." type="journal article" year="1958">Auffenberg, 1958</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFCEE9CBBFB1272AE" author="Hinderstein, B. &amp; Boyce, J." box="[840,1228,1161,1186]" pageId="35" pageNumber="36" pagination="369 - 372" refId="ref33182" refString="Hinderstein, B. and Boyce, J. 1977. The Miocene salamander Batrachosauroides dissimulans (Amphibia, Urodela) from East Texas. Journal of Herpetology, 11: 369 - 372." type="journal article" year="1977">Hinderstein and Boyce, 1977</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFB439CBBFA4272AE" author="Holman, J. A." box="[1253,1436,1161,1186]" pageId="35" pageNumber="36" pagination="391 - 403" refId="ref33217" refString="Holman, J. A. 1977. Amphibians and reptiles from the Gulf Coast Miocene of Texas. Herpetologica, 33: 391 - 403." type="journal article" year="1977">Holman, 1977</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFCEE9C9BFC2D72CE" author="Albright, L. B." box="[840,1011,1193,1218]" pageId="35" pageNumber="36" pagination="1131 - 1145" refId="ref29584" refString="Albright, L. B. 1994. Lower vertebrates from an Arikareean (earliest Miocene) fauna near the Toledo Bend Dam, Newton County, Texas. Journal of Paleontology, 68: 1131 - 1145. https: // doi. org / 10.1017 / S 002233600002672 X" type="journal article" year="1994">Albright, 1994</bibRefCitation>
). They are also known from Europe, even though their record on this continent is far poorer. They were possibly present already during the Cretaceous (
<bibRefCitation id="EFCDEA82D077FFABFBA89D3AFB62732D" author="Duffaud, S." box="[1038,1212,1288,1313]" pageId="35" pageNumber="36" refId="ref31584" refString="Duffaud, S. 1995. A Batrachosauroididae (Amphibia, Caudata) from the late Cretaceous of Champ-Garimond (Southern France). DGF Online Series, available at https: // www. 2 dgf. dk / dgf-online-series" type="url" year="1995">Duffaud, 1995</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFB6F9D3AFC59734D" author="Evans, S. E. &amp; Milner, A. R." pageId="35" pageNumber="36" pagination="627 - 646" refId="ref31837" refString="Evans, S. E. and Milner, A. R. 1996. A metamorphosed salamander from the early Cretaceous of Las Hoyas, Spain. Philosophical Transactions of The Royal Society B Biological Sciences, 351: 627 - 646." type="journal article" year="1996">Evans and Milner, 1996</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFC309D1AFB31734D" author="Evans, S. E. &amp; McGowan, G. J." box="[918,1263,1320,1346]" pageId="35" pageNumber="36" pagination="103 - 119" refId="ref31800" refString="Evans, S. E. and McGowan, G. J. 2002. Lissamphibian remains from the Purbeck Limestone Group, Southern England. Special Papers in Palaeontology, 68: 103 - 119." type="journal article" year="2002">Evans and McGowan, 2002</bibRefCitation>
), and then are seldomly recovered in the Palaeocene (
<bibRefCitation id="EFCDEA82D077FFABFA9E9D7AFC65738D" author="Estes, R. &amp; Hecht, M. &amp; Hoffstetter, R." pageId="35" pageNumber="36" pagination="1 - 25" refId="ref31766" refString="Estes, R., Hecht, M., and Hoffstetter, R. 1967. Paleocene amphibians from Cernay, France. American Museum Novitates, 2295: 1 - 25." type="journal article" year="1967">Estes et al., 1967</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFC699D5AFAF5738D" author="Groessens-Van Dyck, M. C." box="[975,1323,1384,1410]" pageId="35" pageNumber="36" pagination="87 - 100" refId="ref32935" refString="Groessens-Van Dyck, M. C. 1981. Etude des amphibiens du Montien continental de Hainin. Bulletin de la Societe Belge de Geologie / Bulletin van de Belgische Vereniging voor Geologie, 90: 87 - 100." type="journal article" year="1981">Groessens-Van Dyck, 1981</bibRefCitation>
), Eocene (
<bibRefCitation id="EFCDEA82D077FFABFCE99DBAFC3673AD" author="Herre, W." box="[847,1000,1416,1441]" pageId="35" pageNumber="36" pagination="1 - 85" refId="ref32975" refString="Herre, W. 1935. Die Schwanzlurche der mitteleocanen (oberlutetischen) Braunkohle des Geiseltales und die Phylogenie der Urodelen unter Einschluss der fossilen Formen. Zoologica - Original-Abhandlungen aus dem Gesamtgebiete der Zoologie, 87: 1 - 85." type="journal article" year="1935">Herre, 1935</bibRefCitation>
), and Upper Miocene (
<bibRefCitation id="EFCDEA82D077FFABFAA49DBAFBDB73CD" author="Vasilyan, D. &amp; Yanenko, V." pageId="35" pageNumber="36" pagination="2733" refId="ref38931" refString="Vasilyan, D. and Yanenko, V. 2020. The last Palaeoproteus (Urodela: Batrachosauroididae) of Europe. Scientific Reports, 10: 2733. https: // doi. org / 10.1038 / s 41598 - 020 - 59255 - 1" type="journal article" year="2020">Vasilyan and Yanenko, 2020</bibRefCitation>
). A single record from the Lower Miocene is also reported (
<bibRefCitation id="EFCDEA82D077FFABFBDC9DFAFAC173ED" author="Bohme, M." box="[1146,1311,1480,1505]" pageId="35" pageNumber="36" pagination="389 - 401" refId="ref30441" refString="Bohme, M. 2003. Miocene Climatic Optimum: evidence from lower vertebrates of central Europe. Palaeogeography, Palaeoclimatology, Palaeoecology, 195: 389 - 401. https: // doi. org / 10.1016 / S 0031 - 0182 (03) 00367 - 5" type="journal article" year="2003">Böhme, 2003</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D077FFABFA8C9DFAFC69700D" author="Kvacek, Z. &amp; Bohme, M. &amp; Dvorak, Z. &amp; Konzalova, M. &amp; Mach, K. &amp; Prokop, J. &amp; Rajchl, M." pageId="35" pageNumber="36" pagination="1 - 40" refId="ref33850" refString="Kvacek, Z., Bohme, M., Dvorak, Z., Konzalova, M., Mach, K., Prokop, J., and Rajchl, M. 2004. Early Miocene freshwater and swamp ecosystems of the Most Basin (northern Bohemia) with particular reference to the Bilina Mine section. Journal of the Czech Geological Society, 49: 1 - 40." type="journal article" year="2004">Kvaček et al., 2004</bibRefCitation>
), but detailed descriptions and identification of these fossils remain unpublished so far. Similar to the cryptobranchids, the Hambach 6C
<taxonomicName id="4C5CECF0D077FFABFCEE9E7AFC2A706D" baseAuthorityName="Vasilyan and Yanenko" baseAuthorityYear="2020" box="[840,1012,1608,1633]" class="Amphibia" family="Batrachosauroididae" genus="Palaeoproteus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="35" pageNumber="36" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D077FFABFCEE9E7AFC2A706D" box="[840,1012,1608,1633]" italics="true" pageId="35" pageNumber="36">Palaeoproteus</emphasis>
</taxonomicName>
remains are the only batrachosauroidid fossils currently known from the Middle Miocene in Europe. Adding to this, those coming from Hambach 11 and 11C further stand out as the youngest occurrence of this clade worldwide, as well as the only one postdating the Miocene/Pliocene transition. Moreover, Hambach lists among the northernmost sites reached by these urodeles in post-Palaeogene times, being located even slightly more northward than Hrytsiv in
<collectingCountry id="F34BD7E3D070FFACFD1A9894FCC676B3" box="[700,792,166,191]" name="Ukraine" pageId="36" pageNumber="37">Ukraine</collectingCountry>
(
<bibRefCitation id="EFCDEA82D070FFACFF6098F4FDC276D3" author="Vasilyan, D. &amp; Yanenko, V." box="[198,540,198,223]" pageId="36" pageNumber="37" pagination="2733" refId="ref38931" refString="Vasilyan, D. and Yanenko, V. 2020. The last Palaeoproteus (Urodela: Batrachosauroididae) of Europe. Scientific Reports, 10: 2733. https: // doi. org / 10.1038 / s 41598 - 020 - 59255 - 1" type="journal article" year="2020">Vasilyan and Yanenko, 2020</bibRefCitation>
) and Ahníkov/Merkur North in
<collectingCountry id="F34BD7E3D070FFACFE8898D4FE3076F3" box="[302,494,230,256]" name="Czech Republic" pageId="36" pageNumber="37">Czech Republic</collectingCountry>
(where the presence of these amphibians can be seen as unconfirmed, for the time being;
<bibRefCitation id="EFCDEA82D070FFACFED19914FDC77733" author="Bohme, M." box="[375,537,294,319]" pageId="36" pageNumber="37" pagination="389 - 401" refId="ref30441" refString="Bohme, M. 2003. Miocene Climatic Optimum: evidence from lower vertebrates of central Europe. Palaeogeography, Palaeoclimatology, Palaeoecology, 195: 389 - 401. https: // doi. org / 10.1016 / S 0031 - 0182 (03) 00367 - 5" type="journal article" year="2003">Böhme, 2003</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFD829914FCD17733" author="Kvacek, Z. &amp; Bohme, M. &amp; Dvorak, Z. &amp; Konzalova, M. &amp; Mach, K. &amp; Prokop, J. &amp; Rajchl, M." box="[548,783,294,319]" pageId="36" pageNumber="37" pagination="1 - 40" refId="ref33850" refString="Kvacek, Z., Bohme, M., Dvorak, Z., Konzalova, M., Mach, K., Prokop, J., and Rajchl, M. 2004. Early Miocene freshwater and swamp ecosystems of the Most Basin (northern Bohemia) with particular reference to the Bilina Mine section. Journal of the Czech Geological Society, 49: 1 - 40." type="journal article" year="2004">Kvaček et al., 2004</bibRefCitation>
). Both the Miocene and Pliocene Hambach material is here referred to the European genus
<taxonomicName id="4C5CECF0D070FFACFD309954FF2C7793" baseAuthorityName="Vasilyan and Yanenko" baseAuthorityYear="2020" class="Amphibia" family="Batrachosauroididae" genus="Palaeoproteus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D070FFACFD309954FF2C7793" italics="true" pageId="36" pageNumber="37">Palaeoproteus</emphasis>
</taxonomicName>
, and even to a taxon related to the species
<emphasis id="B9284B61D070FFACFCA499B4FE9377B3" italics="true" pageId="36" pageNumber="37">
<taxonomicName id="4C5CECF0D070FFACFCA499B4FE9777B3" baseAuthorityName="Vasilyan and Yanenko" baseAuthorityYear="2020" class="Amphibia" family="Batrachosauroididae" genus="Palaeoproteus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="species" species="miocenicus">P. miocenicus</taxonomicName>
.
</emphasis>
Even though a confident identification of the fossils with this Late Miocene species is currently prevented, Hambach is evidence that either one or two morphologically similar forms (depending on whether occurrences from Hambach 6C and Hambach 11 and 11C are conspecific or not) were already present in Europe several Myr before the oldest occurrence reported by
<bibRefCitation id="EFCDEA82D070FFACFDD69AB4FE5674B3" author="Vasilyan, D. &amp; Yanenko, V." pageId="36" pageNumber="37" pagination="2733" refId="ref38931" refString="Vasilyan, D. and Yanenko, V. 2020. The last Palaeoproteus (Urodela: Batrachosauroididae) of Europe. Scientific Reports, 10: 2733. https: // doi. org / 10.1038 / s 41598 - 020 - 59255 - 1" type="journal article" year="2020">Vasilyan and Yanenko (2020)</bibRefCitation>
and successively persisted at least up to slightly before the Plio-Pleistocene boundary.
</paragraph>
<paragraph id="8BE39773D070FFACFF5C9B34FE587153" blockId="36.[192,793,166,1887]" pageId="36" pageNumber="37">
<taxonomicName id="4C5CECF0D070FFACFF5C9B34FE467513" authorityName="Macaluso, Villa &amp; Mörs" authorityYear="2021" box="[250,408,774,799]" class="Amphibia" family="Proteidae" genus="Euronecturus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D070FFACFF5C9B34FE467513" box="[250,408,774,799]" italics="true" pageId="36" pageNumber="37">Euronecturus</emphasis>
</taxonomicName>
is not the only proteid found in Hambach:
<taxonomicName id="4C5CECF0D070FFACFEE79B14FE1B7533" authorityName="Estes and Darevsky" authorityYear="1977" box="[321,453,806,831]" class="Amphibia" family="Proteidae" genus="Mioproteus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D070FFACFEE79B14FE1B7533" box="[321,453,806,831]" italics="true" pageId="36" pageNumber="37">Mioproteus</emphasis>
</taxonomicName>
, a possible representative of a modern European clade including also the extant
<taxonomicName id="4C5CECF0D070FFACFF669B54FD987573" authority="(Macaluso et al., 2022 b)" baseAuthorityName="Macaluso" baseAuthorityYear="2022" box="[192,582,870,895]" class="Amphibia" family="Proteidae" genus="Proteus" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D070FFACFF669B54FEC27573" box="[192,284,870,895]" italics="true" pageId="36" pageNumber="37">Proteus</emphasis>
(
<bibRefCitation id="EFCDEA82D070FFACFE8F9B54FDE07573" author="Macaluso, L. &amp; Villa, A. &amp; Mors, T." box="[297,574,870,895]" pageId="36" pageNumber="37" pagination="12585" refId="ref34411" refString="Macaluso, L., Villa, A., and Mors, T. 2022 b. A new proteid salamander (Urodela; Proteidae) from the middle Miocene of Hambach (Germany) and implications for the evolution of the family. Palaeontology, 65: e 12585. https: // doi. org / 10.1111 / pala. 12585" type="journal article" year="2022">Macaluso et al., 2022b</bibRefCitation>
)
</taxonomicName>
, actually replaced it in the youngest level. Several other occurrences of
<taxonomicName id="4C5CECF0D070FFACFF7B9B94FEBF75B3" authorityName="Estes and Darevsky" authorityYear="1977" box="[221,353,934,959]" class="Amphibia" family="Proteidae" genus="Mioproteus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D070FFACFF7B9B94FEBF75B3" box="[221,353,934,959]" italics="true" pageId="36" pageNumber="37">Mioproteus</emphasis>
</taxonomicName>
are known from
<collectingCountry id="F34BD7E3D070FFACFD829B94FD4C75CC" box="[548,658,934,960]" name="Germany" pageId="36" pageNumber="37">Germany</collectingCountry>
throughout the Miocene (
<bibRefCitation id="EFCDEA82D070FFACFE269BF4FD9875D3" author="Schleich, H. H." box="[384,582,966,992]" pageId="36" pageNumber="37" pagination="67 - 149" refId="ref36773" refString="Schleich, H. H. 1985. Zur Verbreitung tertiarer und quartarer Reptilien und Amphibien. I. Suddeutschland. Munchner Geowissenschaftliche Abhandlungen Reihe A, 4: 67 - 149." type="journal article" year="1985">Schleich, 1985</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFDC49BF4FCCA75D3" author="Bohme, M." box="[610,788,966,991]" pageId="36" pageNumber="37" pagination="389 - 401" refId="ref30441" refString="Bohme, M. 2003. Miocene Climatic Optimum: evidence from lower vertebrates of central Europe. Palaeogeography, Palaeoclimatology, Palaeoecology, 195: 389 - 401. https: // doi. org / 10.1016 / S 0031 - 0182 (03) 00367 - 5" type="journal article" year="2003">Böhme, 2003</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFF669BD4FE6F75F3" author="Bottcher, R. &amp; Heizmann, E. P. J. &amp; Rasser, M. W. &amp; Ziegler, R." box="[192,433,998,1023]" pageId="36" pageNumber="37" pagination="237 - 260" refId="ref30874" refString="Bottcher, R., Heizmann, E. P. J., Rasser, M. W., and Ziegler, R. 2009. Biostratigraphy and palaeoecology of a Middle Miocene (Karpathian, MN 5) fauna from the northern margin of the North Alpine Foreland Basin (Oggenhausen 2, SW' Germany). Neues Jahrbuch fur Geologie und Palaontologie, 254: 237 - 260. https: // doi. org / 10.1127 / 0077 - 7749 / 2009 / 0011" type="journal article" year="2009">Böttcher et al., 2009</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFE1A9BD4FD4E75F3" author="Prieto, J. &amp; Bohme, M. &amp; Maurer, H. &amp; Heissig, K. &amp; Abdul Aziz, H." box="[444,656,998,1023]" pageId="36" pageNumber="37" pagination="1767 - 1791" refId="ref35524" refString="Prieto, J., Bohme, M., Maurer, H., Heissig, K., and Abdul Aziz, H. 2009. Biostratigraphy and sedimentology of the Fluviatile Untere Serie (Early and Middle Miocene) in the central part of the North Alpine Foreland Basin: implications for palaeoenvironment and climate. International Journal of Earth Sciences, 98: 1767 - 1791. https: // doi. org / 10.1007 / s 00531 - 008 - 0331 - 2" type="journal article" year="2009">Prieto et al., 2009</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFD3D9BD4FE917213" author="Abdul Aziz, H. &amp; Bohme, M. &amp; Rocholl, A. &amp; Prieto, J. &amp; Wijbrans, J. R. &amp; Bachtadse, V. &amp; Ulbig, A." journalOrPublisher="International Journal of Earth Sciences" pageId="36" pageNumber="37" pagination="1859 - 1886" part="99" refId="ref29490" refString="Abdul Aziz, H., Bohme, M., Rocholl, A., Prieto, J., Wijbrans, J. R., Bachtadse, V., and Ulbig, A. 2010. Integrated stratigraphy and 40 Ar / 39 Ar chronology of the early to middle Miocene Upper Freshwater Molasse in western Bavaria (Germany). International Journal of Earth Sciences, 99: 1859 - 1886. https: // doi. org / 10.1007 / s 00531 - 009 - 0475 - 8" title="Integrated stratigraphy and 40 Ar / 39 Ar chronology of the early to middle Miocene Upper Freshwater Molasse in western Bavaria (Germany)" type="journal article" year="2010">Abdul Aziz et al., 2010</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFEFB9C34FE327213" author="Sach, V. J." box="[349,492,1030,1056]" pageId="36" pageNumber="37" pagination="1 - 115" refId="ref36429" refString="Sach, V. J. 2016. Fossilienkatalog der Miozan-Molasse in Sudwestdeutschland (Obere Susswassermolasse, Brackwassermolasse, Obere Meeresmolasse, Untere Susswassermolasse). Documenta Naturae, SB 70: 1 - 115." type="journal article" year="2016">Sach, 2016</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFE5C9C34FCCB7213" author="Macaluso, L. &amp; Mannion, P. D. &amp; Evans, S. E. &amp; Carnevale, G. &amp; Monti, S. &amp; Marchitelli, D. &amp; Delfino, M." box="[506,789,1030,1055]" pageId="36" pageNumber="37" pagination="220935" refId="ref34329" refString="Macaluso, L., Mannion, P. D., Evans, S. E., Carnevale, G., Monti, S., Marchitelli, D., and Delfino, M. 2022 a. Biogeographic history of Palearctic caudates revealed by a critical appraisal of their fossil record quality and spatio-temporal distribution. Royal Society Open Science, 9: 220935. https: // doi. org / 10.1098 / rsos. 220935" type="journal article" year="2022">Macaluso et al., 2022a</bibRefCitation>
,
<bibRefCitation id="EFCDEA82D070FFACFF669C14FECE7233" author="Macaluso, L. &amp; Villa, A. &amp; Mors, T." box="[192,272,1062,1087]" pageId="36" pageNumber="37" pagination="12585" refId="ref34411" refString="Macaluso, L., Villa, A., and Mors, T. 2022 b. A new proteid salamander (Urodela; Proteidae) from the middle Miocene of Hambach (Germany) and implications for the evolution of the family. Palaeontology, 65: e 12585. https: // doi. org / 10.1111 / pala. 12585" type="journal article" year="2022">2022b</bibRefCitation>
), including fossils referred to both
<taxonomicName id="4C5CECF0D070FFACFD059C14FF237253" authorityName="Estes and Darevsky" authorityYear="1977" class="Amphibia" family="Proteidae" genus="Mioproteus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="species" species="caucasicus">
<emphasis id="B9284B61D070FFACFD059C14FF237253" italics="true" pageId="36" pageNumber="37">M. caucasicus</emphasis>
</taxonomicName>
and indeterminate species (even though most of these were never properly published). Its absence in Hambach 6C, thus, appears puzzling per se, especially when paired with the presence of
<taxonomicName id="4C5CECF0D070FFACFF669CF4FE8072D3" authorityName="Macaluso, Villa &amp; Mörs" authorityYear="2021" box="[192,350,1222,1247]" class="Amphibia" family="Proteidae" genus="Euronecturus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D070FFACFF669CF4FE8072D3" box="[192,350,1222,1247]" italics="true" pageId="36" pageNumber="37">Euronecturus</emphasis>
</taxonomicName>
. Some sort of competitive exclusion cannot be discounted, even though only treated as speculative at the moment due to the very poor information we have about the latter. Later on,
<taxonomicName id="4C5CECF0D070FFACFD429D14FEC77353" authorityName="Estes and Darevsky" authorityYear="1977" class="Amphibia" family="Proteidae" genus="Mioproteus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D070FFACFD429D14FEC77353" italics="true" pageId="36" pageNumber="37">Mioproteus</emphasis>
</taxonomicName>
survived up to the late Early Pleistocene (
<bibRefCitation id="EFCDEA82D070FFACFF609D54FDE57373" author="Mlynarski, M. &amp; Szyndlar, Z." box="[198,571,1382,1408]" pageId="36" pageNumber="37" pagination="69 - 88" refId="ref34826" refString="Mlynarski, M. and Szyndlar, Z. 1989. Plazy i gady. Folia Quaternaria, 59 - 60: 69 - 88." type="journal article" year="1989">Młynarski and Szyndlar, 1989</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFDEB9D54FCCA7373" author="Averianov, A. O." box="[589,788,1382,1407]" pageId="36" pageNumber="37" pagination="43 - 46" refId="ref29733" refString="Averianov, A. O. 2001. New records of proteid salamanders (Amphibia, Caudata) from the Pliocene of Ukraine and lower Pleistocene of Moldavia. Vestnik Zoologii, 35: 43 - 46." type="journal article" year="2001">Averianov, 2001</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFF669DB4FEBE7393" author="Ivanov, M." box="[192,352,1414,1439]" pageId="36" pageNumber="37" pagination="297 - 320" refId="ref33419" refString="Ivanov, M. 2007. Herpetological assemblages from the Pliocene to middle Pleistocene in Central Europe: palaeoecological significance. Geodiversitas, 29: 297 - 320." type="journal article" year="2007">Ivanov, 2007</bibRefCitation>
), but all findings postdating the Miocene/Pliocene transition are either unassigned to species (
<bibRefCitation id="EFCDEA82D070FFACFE8E9DF4FE3273D3" author="Averianov, A. O." box="[296,492,1478,1503]" pageId="36" pageNumber="37" pagination="43 - 46" refId="ref29733" refString="Averianov, A. O. 2001. New records of proteid salamanders (Amphibia, Caudata) from the Pliocene of Ukraine and lower Pleistocene of Moldavia. Vestnik Zoologii, 35: 43 - 46." type="journal article" year="2001">Averianov, 2001</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFE519DF4FD7473D3" author="Ratnikov, V. Yu." box="[503,682,1478,1503]" pageId="36" pageNumber="37" pagination="59 - 66" refId="ref35889" refString="Ratnikov, V. Yu. 2010. A review of tailed amphibian remains from late Cenozoic sediments of the East European Plain. Russian Journal of Herpetology, 17: 59 - 66. https: // doi. org / 10.30906 / 1026 - 2296 - 2010 - 17 - 1 - 59 - 66" type="journal article" year="2010">Ratnikov, 2010</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFD139DF4FE9373F3" author="Vasilyan, D. &amp; Zazhigin, V. S. &amp; Bohme, M." pageId="36" pageNumber="37" pagination="3025" refId="ref39119" refString="Vasilyan, D., Zazhigin, V. S., and Bohme, M. 2017. Neogene amphibians and reptiles (Caudata, Anura, Gekkota, Lacertilia, and Testudines) from the south of Western Siberia, Russia, and Northeastern Kazakhstan. PeerJ, 5: e 3025. https: // doi. org / 10.7717 / peerj. 3025" type="journal article" year="2017">Vasilyan et al., 2017</bibRefCitation>
) or referred to
<taxonomicName id="4C5CECF0D070FFACFDAE9DD4FDAA73F3" box="[520,628,1510,1535]" class="Amphibia" family="Proteidae" genus="Mioproteus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="species" species="wezei">
<emphasis id="B9284B61D070FFACFDAE9DD4FDAA73F3" box="[520,628,1510,1535]" italics="true" pageId="36" pageNumber="37">M. wezei</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFCDEA82D070FFACFD229DD4FEEE7013" author="Mlynarski, M. &amp; Szyndlar, Z. &amp; Estes, R. &amp; Sanchiz B." pageId="36" pageNumber="37" pagination="209 - 229" refId="ref34852" refString="Mlynarski, M., Szyndlar, Z., Estes, R., and Sanchiz B. 1984. Amphibians and reptiles from the Pliocene locality of Weze II near Dzialoszyn (Poland). Acta Palaeontologica Polonica, 29: 209 - 229." type="journal article" year="1984">Młynarski et al., 1984</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFEE49E34FD6B7013" author="Mlynarski, M. &amp; Szyndlar, Z." box="[322,693,1542,1568]" pageId="36" pageNumber="37" pagination="69 - 88" refId="ref34826" refString="Mlynarski, M. and Szyndlar, Z. 1989. Plazy i gady. Folia Quaternaria, 59 - 60: 69 - 88." type="journal article" year="1989">Młynarski and Szyndlar, 1989</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFD619E34FF217033" author="Bailon, S." pageId="36" pageNumber="37" pagination="1 - 8" refId="ref29841" refString="Bailon, S. 1995. Presence de Mioproteus wezei (Amphibia, Proteidae) dans le Pliocene superieur (MN 16) d'Europe de l'Ouest. Neues Jahrbuch fur Geologie und Palaontologie, Monatshefte, 1995: 1 - 8. https: // doi. org / 10.1127 / njgpm / 1995 / 1995 / 1" type="journal article" year="1995">Bailon, 1995</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFEB79E14FE077033" author="Averianov, A. O." box="[273,473,1574,1599]" pageId="36" pageNumber="37" pagination="43 - 46" refId="ref29733" refString="Averianov, A. O. 2001. New records of proteid salamanders (Amphibia, Caudata) from the Pliocene of Ukraine and lower Pleistocene of Moldavia. Vestnik Zoologii, 35: 43 - 46." type="journal article" year="2001">Averianov, 2001</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFE4D9E14FD537033" author="Ivanov, M." box="[491,653,1574,1599]" pageId="36" pageNumber="37" pagination="297 - 320" refId="ref33419" refString="Ivanov, M. 2007. Herpetological assemblages from the Pliocene to middle Pleistocene in Central Europe: palaeoecological significance. Geodiversitas, 29: 297 - 320." type="journal article" year="2007">Ivanov, 2007</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFD399E14FE7F7053" author="Syromyatnikova, E. &amp; Tesakov, A. &amp; Frolov, P. &amp; Titov, V." pageId="36" pageNumber="37" pagination="125 - 128" refId="ref38008" refString="Syromyatnikova, E., Tesakov, A., Frolov, P., and Titov, V. 2021. A Pliocene Mioproteus (Urodela: Proteidae) from the Taman Peninsula (Russia). Russian Journal of Herpetology, 28: 125 - 128. https: // doi. org / 10.30906 / 1026 - 2296 - 2021 - 28 - 2 - 125 - 128" type="journal article" year="2021">Syromyatnikova et al., 2021</bibRefCitation>
,
<bibRefCitation id="EFCDEA82D070FFACFE089E74FD187053" author="Macaluso, L. &amp; Mannion, P. D. &amp; Evans, S. E. &amp; Carnevale, G. &amp; Monti, S. &amp; Marchitelli, D. &amp; Delfino, M." box="[430,710,1606,1631]" pageId="36" pageNumber="37" pagination="220935" refId="ref34329" refString="Macaluso, L., Mannion, P. D., Evans, S. E., Carnevale, G., Monti, S., Marchitelli, D., and Delfino, M. 2022 a. Biogeographic history of Palearctic caudates revealed by a critical appraisal of their fossil record quality and spatio-temporal distribution. Royal Society Open Science, 9: 220935. https: // doi. org / 10.1098 / rsos. 220935" type="journal article" year="2022">Macaluso et al., 2022a</bibRefCitation>
). If our identification is correct, the Upper Pliocene occurrence of
<taxonomicName id="4C5CECF0D070FFACFE899EB4FDF47093" box="[303,554,1670,1695]" class="Amphibia" family="Proteidae" genus="Mioproteus" higherTaxonomySource="GBIF" isUncertain="true" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="species" species="wezei">
<emphasis id="B9284B61D070FFACFE899EB4FE6D7093" box="[303,435,1670,1695]" italics="true" pageId="36" pageNumber="37">Mioproteus</emphasis>
cf.
<emphasis id="B9284B61D070FFACFE409EB4FDF47093" box="[486,554,1670,1695]" italics="true" pageId="36" pageNumber="37">wezei</emphasis>
</taxonomicName>
in Hambach would agree with this pattern. Furthermore, this is currently one of the westernmost records of
<taxonomicName id="4C5CECF0D070FFACFD189EF4FF2C70F3" authorityName="Estes and Darevsky" authorityYear="1977" class="Amphibia" family="Proteidae" genus="Mioproteus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D070FFACFD189EF4FF2C70F3" italics="true" pageId="36" pageNumber="37">Mioproteus</emphasis>
</taxonomicName>
, with only
<taxonomicName id="4C5CECF0D070FFACFED29ED4FE0170F3" box="[372,479,1766,1791]" class="Amphibia" family="Proteidae" genus="Mioproteus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="species" species="wezei">
<emphasis id="B9284B61D070FFACFED29ED4FE0170F3" box="[372,479,1766,1791]" italics="true" pageId="36" pageNumber="37">M. wezei</emphasis>
</taxonomicName>
from Balaruc II, in
<collectingCountry id="F34BD7E3D070FFACFD639ED4FCC670F3" box="[709,792,1766,1791]" name="France" pageId="36" pageNumber="37">France</collectingCountry>
(
<bibRefCitation id="EFCDEA82D070FFACFF619F34FEB67113" author="Bailon, S." box="[199,360,1798,1823]" pageId="36" pageNumber="37" pagination="1 - 8" refId="ref29841" refString="Bailon, S. 1995. Presence de Mioproteus wezei (Amphibia, Proteidae) dans le Pliocene superieur (MN 16) d'Europe de l'Ouest. Neues Jahrbuch fur Geologie und Palaontologie, Monatshefte, 1995: 1 - 8. https: // doi. org / 10.1127 / njgpm / 1995 / 1995 / 1" type="journal article" year="1995">Bailon, 1995</bibRefCitation>
), located in an even more western position (even though in a far more southern, Mediterranean area).
</paragraph>
<paragraph id="8BE39773D070FFADFC249894FE61778E" blockId="36.[840,1441,166,1887]" lastBlockId="37.[192,793,169,1890]" lastPageId="37" lastPageNumber="38" pageId="36" pageNumber="37">
Among salamandrids,
<taxonomicName id="4C5CECF0D070FFACFB369894FACF76B3" authorityName="Pomel" authorityYear="1853" box="[1168,1297,166,191]" class="Amphibia" family="Salamandridae" genus="Chelotriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D070FFACFB369894FACF76B3" box="[1168,1297,166,191]" italics="true" pageId="36" pageNumber="37">Chelotriton</emphasis>
</taxonomicName>
is the most represented in Hambach 6C and also very common in Europe, and even in
<collectingCountry id="F34BD7E3D070FFACFB0F98D4FAC7770C" box="[1193,1305,230,256]" name="Germany" pageId="36" pageNumber="37">Germany</collectingCountry>
, during the Miocene. Hambach lists among the northernmost Miocene occurrences reached by the genus on the continent, together with the slightly more southern remains from Echzell (
<bibRefCitation id="EFCDEA82D070FFACFBE89954FA9A7773" author="Vasilyan, D. &amp; Cernansky, A. &amp; Szyndlar, Z. &amp; Mors, T." box="[1102,1348,358,383]" pageId="36" pageNumber="37" pagination="99 - 145" refId="ref39186" refString="Vasilyan, D., Cernansky, A., Szyndlar, Z., and Mors, T. 2022. Amphibian and reptilian fauna from the early Miocene of Echzell, Germany. Fossil Record, 25: 99 - 145. https: // doi. org / 10.3897 / fr. 25.83781" type="journal article" year="2022">Vasilyan et al., 2022</bibRefCitation>
). These large salamandrids apparently decreased after the Messinian Salinity Crisis. They are reported from the post-evaporitic succession of Moncucco Torinese in
<collectingCountry id="F34BD7E3D070FFACFC0F99D4FC0777F3" box="[937,985,486,511]" name="Italy" pageId="36" pageNumber="37">Italy</collectingCountry>
(
<bibRefCitation id="EFCDEA82D070FFACFC4C99D4FAD977F3" author="Colombero, S. &amp; Alba, D. M. &amp; D'Amico, C. &amp; Delfino, M. &amp; Esu, D. &amp; Giuntelli, P. &amp; Harzhauser, M. &amp; Mazza, P. P. A. &amp; Mosca, M. &amp; Neubauer, T. A. &amp; Pavia, G. &amp; Pavia, M. &amp; Villa, A. &amp; Carnevale, G." box="[1002,1287,486,512]" pageId="36" pageNumber="37" pagination="1 - 66" refId="ref31150" refString="Colombero, S., Alba, D. M., D'Amico, C., Delfino, M., Esu, D., Giuntelli, P., Harzhauser, M., Mazza, P. P. A., Mosca, M., Neubauer, T. A., Pavia, G., Pavia, M., Villa, A., and Carnevale, G. 2017. Late Messinian mollusks and vertebrates from Moncucco Torinese, north-western Italy. Paleoecological and paleoclimatological implications. Palaeontologia Electronica, 20.1.10 A: 1 - 66. https: // doi. org / 10.26879 / 658" type="book chapter" year="2017">Colombero et al., 2017</bibRefCitation>
), and subsequently the only published occurrence is from the Upper Pliocene of Balaruc II in
<collectingCountry id="F34BD7E3D070FFACFB429A14FAE97433" box="[1252,1335,550,575]" name="France" pageId="36" pageNumber="37">France</collectingCountry>
(
<bibRefCitation id="EFCDEA82D070FFACFAEA9A14FC577453" author="Bailon, S." pageId="36" pageNumber="37" pagination="7 - 28" refId="ref29771" refString="Bailon, S. 1989. Les amphibiens et les reptiles du Pliocene Superieur de Balaruc II (Herault, France). Palaeovertebrata, 19: 7 - 28." type="journal article" year="1989">Bailon, 1989</bibRefCitation>
). Their absence in Hambach 11 and 11C agrees with this observed pattern of extirpation from northern Europe possibly somewhen in the Late Miocene, and in any case before the Late Pliocene (
<bibRefCitation id="EFCDEA82D070FFACFC669AF4FB3F74D3" author="Macaluso, L. &amp; Mannion, P. D. &amp; Evans, S. E. &amp; Carnevale, G. &amp; Monti, S. &amp; Marchitelli, D. &amp; Delfino, M." box="[960,1249,710,735]" pageId="36" pageNumber="37" pagination="220935" refId="ref34329" refString="Macaluso, L., Mannion, P. D., Evans, S. E., Carnevale, G., Monti, S., Marchitelli, D., and Delfino, M. 2022 a. Biogeographic history of Palearctic caudates revealed by a critical appraisal of their fossil record quality and spatio-temporal distribution. Royal Society Open Science, 9: 220935. https: // doi. org / 10.1098 / rsos. 220935" type="journal article" year="2022">Macaluso et al., 2022a</bibRefCitation>
).
<taxonomicName id="4C5CECF0D070FFACFB519AF4FA9174D3" authorityName="Rafinesque" authorityYear="1815" box="[1271,1359,710,735]" class="Amphibia" family="Salamandridae" genus="Triturus" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D070FFACFB519AF4FA9174D3" box="[1271,1359,710,735]" italics="true" pageId="36" pageNumber="37">Triturus</emphasis>
</taxonomicName>
newts also are represented in Hambach 6C only in our material, even though, in contrast with
<taxonomicName id="4C5CECF0D070FFACFABE9B34FA477513" authorityName="Pomel" authorityYear="1853" box="[1304,1433,774,799]" class="Amphibia" family="Salamandridae" genus="Chelotriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D070FFACFABE9B34FA477513" box="[1304,1433,774,799]" italics="true" pageId="36" pageNumber="37">Chelotriton</emphasis>
</taxonomicName>
, their presence as far north as central
<collectingCountry id="F34BD7E3D070FFACFA959B14FA7E754C" box="[1331,1440,806,832]" name="Germany" pageId="36" pageNumber="37">Germany</collectingCountry>
(
<bibRefCitation id="EFCDEA82D070FFACFCE99B74FBDB7553" author="Holman, J. A." box="[847,1029,838,863]" pageId="36" pageNumber="37" pagination="1 - 254" refId="ref33244" refString="Holman, J. A. 1998. Pleistocene amphibians and reptiles in Britain and Europe. Oxford monographs on geology and geophysics, 38: 1 - 254." type="journal article" year="1998">Holman, 1998</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFBBD9B74FB177553" author="Bohme, G." box="[1051,1225,838,863]" pageId="36" pageNumber="37" pagination="79 - 97" refId="ref30380" refString="Bohme, G. 2000. Fossile Amphibien und Reptilien im Quartar Thuringens. Veroffentlichungen Naturkundemuseum Erfurt, 19 / 2000: 79 - 97." type="journal article" year="2000">Böhme, 2000</bibRefCitation>
,
<bibRefCitation id="EFCDEA82D070FFACFB789B74FAC27553" author="Bohme, M." box="[1246,1308,838,863]" pageId="36" pageNumber="37" pagination="1141 - 1158" refId="ref30604" refString="Bohme, M. 2020. New results on amphibians and reptiles from the Early Pleistocene site of Untermassfeld - bioclimatic analysis of the herpetofauna. Monographien des RGZM, 40: 1141 - 1158" type="journal article" year="2020">2020</bibRefCitation>
) and
<collectingCountry id="F34BD7E3D070FFACFAD49B74FC097573" name="Netherlands" pageId="36" pageNumber="37">The Netherlands</collectingCountry>
(
<bibRefCitation id="EFCDEA82D070FFACFC4A9B54FB117573" author="Villa, A. &amp; Blain, H. - A. &amp; van den Hoek Ostende, L. W. &amp; Delfino, M." box="[1004,1231,870,895]" pageId="36" pageNumber="37" pagination="203 - 219" refId="ref39874" refString="Villa, A., Blain, H. - A., van den Hoek Ostende, L. W., and Delfino, M. 2018 c. Fossil amphibians and reptiles from Tegelen (Province of Limburg) and the early Pleistocene palaeoclimate of The Netherlands. Quaternary Science Reviews, 187: 203 - 219. https: // doi. org / 10.1016 / j. quascirev. 2018.03.020" type="journal article" year="2018">Villa et al., 2018c</bibRefCitation>
) in northwestern Europe is known up to the Quaternary. The genus still persists nowadays in the area of Hambach with the species
<taxonomicName id="4C5CECF0D070FFACFC7D9BF4FB8275D3" baseAuthorityName="Laurenti" baseAuthorityYear="1768" box="[987,1116,966,991]" class="Amphibia" family="Salamandridae" genus="Triturus" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="species" species="cristatus">
<emphasis id="B9284B61D070FFACFC7D9BF4FB8275D3" box="[987,1116,966,991]" italics="true" pageId="36" pageNumber="37">T. cristatus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFCDEA82D070FFACFBCA9BF4FA9075D3" author="Sillero, N. &amp; Campos, J. &amp; Bonardi, A. &amp; Corti, C. &amp; Creemers, R. &amp; Crochet, P. - A. &amp; Crnobrnja Isailovic, J. &amp; Denoel, M. &amp; Ficetola, G. F. &amp; Goncalves, J. &amp; Kuzmin, S. &amp; Lymberakis, P. &amp; de Pous, P. &amp; Rodriguez, A. &amp; Sindaco, R. &amp; Speybroeck, J. &amp; Toxopeus, B. &amp; Vieites, D. R. &amp; Vences, M." box="[1132,1358,966,992]" pageId="36" pageNumber="37" pagination="1 - 31" refId="ref37047" refString="Sillero, N., Campos, J., Bonardi, A., Corti, C., Creemers, R., Crochet, P. - A., Crnobrnja Isailovic, J., Denoel, M., Ficetola, G. F., Goncalves, J., Kuzmin, S., Lymberakis, P., de Pous, P., Rodriguez, A., Sindaco, R., Speybroeck, J., Toxopeus, B., Vieites, D. R., and Vences, M. 2014. Updated distribution and biogeography of amphibians and reptiles of Europe. Amphibia-Reptilia, 35: 1 - 31. https: // doi. org / 10.1163 / 15685381 - 00002935" type="journal article" year="2014">Sillero et al., 2014</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFAFA9BF4FBEF75F3" author="Speybroeck, J. &amp; Beukema, W. &amp; Bok, B. &amp; Van der Voort, J." pageId="36" pageNumber="37" refId="ref37342" refString="Speybroeck, J., Beukema, W., Bok, B., and Van der Voort, J. 2016. Field guide to the amphibians and reptiles of Britain and Europe. Bloomsbury Publishing, London." type="book" year="2016">Speybroeck et al., 2016</bibRefCitation>
). Few published
<taxonomicName id="4C5CECF0D070FFACFB5D9BD4FA8D75F3" authorityName="Rafinesque" authorityYear="1815" box="[1275,1363,998,1023]" class="Amphibia" family="Salamandridae" genus="Triturus" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D070FFACFB5D9BD4FA8D75F3" box="[1275,1363,998,1023]" italics="true" pageId="36" pageNumber="37">Triturus</emphasis>
</taxonomicName>
dated back to
<collectionCode id="ED4D0FB6D070FFACFC029C34FC137213" box="[932,973,1030,1055]" country="Brazil" name="Museu Nacional, Universidade Federal do Rio de Janeiro" pageId="36" pageNumber="37">MN</collectionCode>
5 are also reported in Europe only from southern
<collectingCountry id="F34BD7E3D070FFACFC119C14FBFA724C" box="[951,1060,1062,1088]" name="Germany" pageId="36" pageNumber="37">Germany</collectingCountry>
(
<bibRefCitation id="EFCDEA82D070FFACFB979C14FAF97233" author="Bottcher, R. &amp; Heizmann, E. P. J. &amp; Rasser, M. W. &amp; Ziegler, R." box="[1073,1319,1062,1087]" pageId="36" pageNumber="37" pagination="237 - 260" refId="ref30874" refString="Bottcher, R., Heizmann, E. P. J., Rasser, M. W., and Ziegler, R. 2009. Biostratigraphy and palaeoecology of a Middle Miocene (Karpathian, MN 5) fauna from the northern margin of the North Alpine Foreland Basin (Oggenhausen 2, SW' Germany). Neues Jahrbuch fur Geologie und Palaontologie, 254: 237 - 260. https: // doi. org / 10.1127 / 0077 - 7749 / 2009 / 0011" type="journal article" year="2009">Böttcher et al., 2009</bibRefCitation>
), but other occurrences from
<collectingCountry id="F34BD7E3D070FFACFB8C9C74FB357253" box="[1066,1259,1094,1120]" name="Czech Republic" pageId="36" pageNumber="37">Czech Republic</collectingCountry>
and
<collectingCountry id="F34BD7E3D070FFACFA949C74FA7E726C" box="[1330,1440,1094,1120]" name="Germany" pageId="36" pageNumber="37">Germany</collectingCountry>
were mentioned by
<bibRefCitation id="EFCDEA82D070FFACFB9C9C54FB397273" author="Bohme, M." box="[1082,1255,1126,1151]" pageId="36" pageNumber="37" pagination="389 - 401" refId="ref30441" refString="Bohme, M. 2003. Miocene Climatic Optimum: evidence from lower vertebrates of central Europe. Palaeogeography, Palaeoclimatology, Palaeoecology, 195: 389 - 401. https: // doi. org / 10.1016 / S 0031 - 0182 (03) 00367 - 5" type="journal article" year="2003">Böhme (2003)</bibRefCitation>
without figures or detailed descriptions. The Hambach specimens add, thus, to the rather poor knowledge of these newts at the Early/Middle Miocene transition. Further significance can be given to the Miocene
<taxonomicName id="4C5CECF0D070FFACFAD39CD4FC417313" authorityName="Bell" authorityYear="1839" class="Amphibia" family="Salamandridae" genus="Lissotriton" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D070FFACFAD39CD4FC417313" italics="true" pageId="36" pageNumber="37">Lissotriton</emphasis>
</taxonomicName>
remains from Hambach 6C, given that these stand out among the oldest occurrences of the genus. Older remains were found in two upper Oligocene sites in southern
<collectingCountry id="F34BD7E3D070FFACFB029D54FB297373" box="[1188,1271,1382,1407]" name="France" pageId="36" pageNumber="37">France</collectingCountry>
(
<bibRefCitation id="EFCDEA82D070FFACFAAF9D54FC1F7393" author="Macaluso, L. &amp; Mannion, P. D. &amp; Evans, S. E. &amp; Carnevale, G. &amp; Monti, S. &amp; Marchitelli, D. &amp; Delfino, M." pageId="36" pageNumber="37" pagination="220935" refId="ref34329" refString="Macaluso, L., Mannion, P. D., Evans, S. E., Carnevale, G., Monti, S., Marchitelli, D., and Delfino, M. 2022 a. Biogeographic history of Palearctic caudates revealed by a critical appraisal of their fossil record quality and spatio-temporal distribution. Royal Society Open Science, 9: 220935. https: // doi. org / 10.1098 / rsos. 220935" type="journal article" year="2022">Macaluso et al., 2022a</bibRefCitation>
) and southern
<collectingCountry id="F34BD7E3D070FFACFBD19DB4FB3B73AC" box="[1143,1253,1414,1440]" name="Germany" pageId="36" pageNumber="37">Germany</collectingCountry>
(
<bibRefCitation id="EFCDEA82D070FFACFB539DB4FA497393" author="Bohme, M." box="[1269,1431,1414,1439]" pageId="36" pageNumber="37" pagination="161 - 183" refId="ref30490" refString="Bohme, M. 2008. Ectothermic vertebrates (Teleostei, Allocaudata, Urodela, Anura, Testudines, Choristodera, Crocodylia, Squamata) from the Upper Oligocene of Oberleichtersbach (Northern Bavaria, Germany). Courier Forschungsinstitut Senckenberg, 260: 161 - 183." type="journal article" year="2008">Böhme 2008</bibRefCitation>
), as well as in the Lower Miocene of southern
<collectingCountry id="F34BD7E3D070FFACFCEE9DF4FC4573D3" box="[840,923,1478,1503]" name="France" pageId="36" pageNumber="37">France</collectingCountry>
(two sites;
<bibRefCitation id="EFCDEA82D070FFACFB8C9DF4FA8D73D3" author="Hossini, S. &amp; Rage, J. - C." box="[1066,1363,1478,1503]" pageId="36" pageNumber="37" pagination="223 - 231" refId="ref33317" refString="Hossini, S. and Rage, J. - C. 2000. Palaeobatrachid frogs from the earliest Miocene (Agenian) of France, with description of a new species. Geobios, 33: 223 - 231. https: // doi. org / 10.1016 / S 0016 - 6995 (00) 80019 - 4" type="journal article" year="2000">Rage and Hossini, 2000</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFAC69DF4FBF973F3" author="Rage, J. - C. &amp; Bailon, S." pageId="36" pageNumber="37" pagination="413 - 441" refId="ref35719" refString="Rage, J. - C. and Bailon, S. 2005. Amphibians and squamate reptiles from the late early Miocene (MN 4) of Beon 1 (Montreal-du-Gers, southwestern France). Geodiversitas, 27: 413 - 441." type="journal article" year="2005">Rage and Bailon, 2005</bibRefCitation>
), western
<collectingCountry id="F34BD7E3D070FFACFB0B9DD4FAC4700C" box="[1197,1306,1510,1536]" name="Germany" pageId="36" pageNumber="37">Germany</collectingCountry>
(one site;
<bibRefCitation id="EFCDEA82D070FFACFCEE9E34FB9E7013" author="Vasilyan, D. &amp; Cernansky, A. &amp; Szyndlar, Z. &amp; Mors, T." box="[840,1088,1542,1567]" pageId="36" pageNumber="37" pagination="99 - 145" refId="ref39186" refString="Vasilyan, D., Cernansky, A., Szyndlar, Z., and Mors, T. 2022. Amphibian and reptilian fauna from the early Miocene of Echzell, Germany. Fossil Record, 25: 99 - 145. https: // doi. org / 10.3897 / fr. 25.83781" type="journal article" year="2022">Vasilyan et al., 2022</bibRefCitation>
), and southern
<collectingCountry id="F34BD7E3D070FFACFB5E9E34FABB702C" box="[1272,1381,1542,1568]" name="Germany" pageId="36" pageNumber="37">Germany</collectingCountry>
(two sites;
<bibRefCitation id="EFCDEA82D070FFACFC379E14FB527033" author="Bottcher, R. &amp; Heizmann, E. P. J. &amp; Rasser, M. W. &amp; Ziegler, R." box="[913,1164,1574,1599]" pageId="36" pageNumber="37" pagination="237 - 260" refId="ref30874" refString="Bottcher, R., Heizmann, E. P. J., Rasser, M. W., and Ziegler, R. 2009. Biostratigraphy and palaeoecology of a Middle Miocene (Karpathian, MN 5) fauna from the northern margin of the North Alpine Foreland Basin (Oggenhausen 2, SW' Germany). Neues Jahrbuch fur Geologie und Palaontologie, 254: 237 - 260. https: // doi. org / 10.1127 / 0077 - 7749 / 2009 / 0011" type="journal article" year="2009">Böttcher et al., 2009</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D070FFACFB3C9E14FA9C7033" author="Bohme, M." box="[1178,1346,1574,1599]" pageId="36" pageNumber="37" pagination="3 - 41" refId="ref30538" refString="Bohme, M. 2010. Ectothermic vertebrates (Actinopterygii, Allocaudata, Urodela, Anura, Crocodylia, Squamata) from the Miocene of Sandelzhausen (Germany, Bavaria) and their implications for environment reconstruction and palaeoclimate. Palaontologische Zeitschrift, 84: 3 - 41. https: // doi. org / 10.1007 / s 12542 - 010 - 0050 - 4" type="journal article" year="2010">Böhme, 2010</bibRefCitation>
).
<bibRefCitation id="EFCDEA82D070FFACFAF09E14FBFA7053" author="Macaluso, L. &amp; Mannion, P. D. &amp; Evans, S. E. &amp; Carnevale, G. &amp; Monti, S. &amp; Marchitelli, D. &amp; Delfino, M." pageId="36" pageNumber="37" pagination="220935" refId="ref34329" refString="Macaluso, L., Mannion, P. D., Evans, S. E., Carnevale, G., Monti, S., Marchitelli, D., and Delfino, M. 2022 a. Biogeographic history of Palearctic caudates revealed by a critical appraisal of their fossil record quality and spatio-temporal distribution. Royal Society Open Science, 9: 220935. https: // doi. org / 10.1098 / rsos. 220935" type="journal article" year="2022">Macaluso et al. (2022a)</bibRefCitation>
further mentioned possible
<taxonomicName id="4C5CECF0D070FFACFAD39E74FC417073" authorityName="Bell" authorityYear="1839" class="Amphibia" family="Salamandridae" genus="Lissotriton" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D070FFACFAD39E74FC417073" italics="true" pageId="36" pageNumber="37">Lissotriton</emphasis>
</taxonomicName>
identity for lower Eocene vertebrae from
<collectingCountry id="F34BD7E3D070FFACFCEE9EB4FC457093" box="[840,923,1670,1695]" name="France" pageId="36" pageNumber="37">France</collectingCountry>
(
<bibRefCitation id="EFCDEA82D070FFACFC0D9EB4FB587093" author="Auge, M. &amp; Duffaud, S. &amp; Lapparent De Broin, F. &amp; Rage, J. - C. &amp; Vasse, D." box="[939,1158,1670,1696]" pageId="36" pageNumber="37" pagination="23 - 33" refId="ref29668" refString="Auge, M., Duffaud, S., Lapparent De Broin, F., Rage, J. - C., and Vasse, D. 1997. Les amphibiens et les reptiles de Premontre (Cuisien, Bassin parisien): une herpetofaune de reference pour l'Eocene inferieur. Geologie de la France, 1: 23 - 33." type="journal article" year="1997">Augé et al., 1997</bibRefCitation>
), but this has yet to be confirmed. Several other localities yielding Early and Middle Miocene
<taxonomicName id="4C5CECF0D070FFACFBF59EF4FB1270D3" authorityName="Bell" authorityYear="1839" box="[1107,1228,1734,1759]" class="Amphibia" family="Salamandridae" genus="Lissotriton" kingdom="Animalia" order="Caudata" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D070FFACFBF59EF4FB1270D3" box="[1107,1228,1734,1759]" italics="true" pageId="36" pageNumber="37">Lissotriton</emphasis>
</taxonomicName>
remains in
<collectingCountry id="F34BD7E3D070FFACFACD9EF4FC5770F3" name="Germany" pageId="36" pageNumber="37">Germany</collectingCountry>
(possibly going back even to the Oligocene/ Miocene transition) were listed by
<bibRefCitation id="EFCDEA82D070FFACFB4A9F34FA437113" author="Bohme, M." box="[1260,1437,1798,1823]" pageId="36" pageNumber="37" pagination="389 - 401" refId="ref30441" refString="Bohme, M. 2003. Miocene Climatic Optimum: evidence from lower vertebrates of central Europe. Palaeogeography, Palaeoclimatology, Palaeoecology, 195: 389 - 401. https: // doi. org / 10.1016 / S 0031 - 0182 (03) 00367 - 5" type="journal article" year="2003">Böhme (2003)</bibRefCitation>
, but these fossils still await proper publication. On the other hand, the very few vertebrae from Hambach 11 represent the first Pliocene
<taxonomicName id="4C5CECF0D071FFADFD39989BFCC676CE" authorityName="Bell" authorityYear="1839" box="[671,792,169,194]" class="Amphibia" family="Salamandridae" genus="Lissotriton" kingdom="Animalia" order="Caudata" pageId="37" pageNumber="38" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D071FFADFD39989BFCC676CE" box="[671,792,169,194]" italics="true" pageId="37" pageNumber="38">Lissotriton</emphasis>
</taxonomicName>
remains found in
<collectingCountry id="F34BD7E3D071FFADFE2D98FAFE2576EE" box="[395,507,200,226]" name="Germany" pageId="37" pageNumber="38">Germany</collectingCountry>
, where otherwise occurrences in the country date to either Miocene or Quaternary. Very few further Late Pliocene remains come from
<collectingCountry id="F34BD7E3D071FFADFEE1991BFE44774E" box="[327,410,297,322]" name="Poland" pageId="37" pageNumber="38">Poland</collectingCountry>
(
<bibRefCitation id="EFCDEA82D071FFADFE0E991AFCCA774E" author="Sanchiz, B. &amp; Mlynarski, M." box="[424,788,296,322]" pageId="37" pageNumber="38" pagination="175 - 188" refId="ref36551" refString="Sanchiz, B., and Mlynarski, M. 1979 b. Pliocene salamandrids (Amphibia, Caudata) from Poland. Acta Zoologica Cracoviensia, 24: 175 - 188." type="journal article" year="1979">Sanchiz and Młynarski, 1979b</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D071FFADFF66997BFDFD776E" author="Mlynarski, M. &amp; Szyndlar, Z." box="[192,547,328,354]" pageId="37" pageNumber="38" pagination="69 - 88" refId="ref34826" refString="Mlynarski, M. and Szyndlar, Z. 1989. Plazy i gady. Folia Quaternaria, 59 - 60: 69 - 88." type="journal article" year="1989">Młynarski and Szyndlar, 1989</bibRefCitation>
) and
<collectingCountry id="F34BD7E3D071FFADFDCB997AFD73776E" box="[621,685,328,354]" name="Spain" pageId="37" pageNumber="38">Spain</collectingCountry>
(
<bibRefCitation id="EFCDEA82D071FFADFD66997AFE6B778E" author="Gomez de Soler, B. &amp; Campeny Vall-Llosera, G. &amp; Van der Made, J. &amp; Oms, O. &amp; Agusti, J. &amp; Sala, R. &amp; Blain, H. A. &amp; Burjachs, F. &amp; Claude, J. &amp; Garcia Catalan, S. &amp; Riba, D. &amp; Rosillo, R." pageId="37" pageNumber="38" pagination="1 - 17" refId="ref32756" refString="Gomez de Soler, B., Campeny Vall-Llosera, G., Van der Made, J., Oms, O., Agusti, J., Sala, R., Blain, H. A., Burjachs, F., Claude, J., Garcia Catalan, S., Riba, D., and Rosillo, R. 2012. A new key locality for the Pliocene vertebrate record of Europe: the Camp dels Ninots maar (NE Spain). Geologica Acta, 10: 1 - 17. https: // doi. org / 10.1344 / 105.000001702" type="journal article" year="2012">Gómez de Soler et al., 2012</bibRefCitation>
).
</paragraph>
<paragraph id="8BE39773D071FFADFF5C99BBFEF3736E" blockId="37.[192,793,169,1890]" pageId="37" pageNumber="38">
Anurans from Hambach also include very common taxa in Europe during corresponding time frames, as well as unexpected occurrences. The most abundant anuran in Hambach,
<taxonomicName id="4C5CECF0D071FFADFD3199DBFD30740E" authorityName="von Meyer" authorityYear="1843" box="[663,750,489,514]" class="Amphibia" family="Alytidae" genus="Latonia" kingdom="Animalia" order="Anura" pageId="37" pageNumber="38" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D071FFADFD3199DBFD30740E" box="[663,750,489,514]" italics="true" pageId="37" pageNumber="38">Latonia</emphasis>
</taxonomicName>
, is very common all over Europe, including
<collectingCountry id="F34BD7E3D071FFADFD039A3AFCCB742E" box="[677,789,520,546]" name="Germany" pageId="37" pageNumber="38">Germany</collectingCountry>
, during most of the Neogene (
<bibRefCitation id="EFCDEA82D071FFADFDBB9A1AFD17744E" author="Rocek, Z." box="[541,713,552,578]" pageId="37" pageNumber="38" pagination="717 - 751" refId="ref36160" refString="Rocek, Z. 1994 b. Taxonomy and distribution of tertiary discoglossids (Anura) of the genus Latonia v. Meyer, 1843. Geobios, 27: 717 - 751. https: // doi. org / 10.1016 / S 0016 - 6995 (94) 80058 - 8" type="journal article" year="1994">Roček, 1994b</bibRefCitation>
,
<bibRefCitation id="EFCDEA82D071FFADFD739A1BFCCB744E" author="Rocek, Z." box="[725,789,553,578]" pageId="37" pageNumber="38" pagination="397 - 439" refId="ref36270" refString="Rocek, Z. 2013. Mesozoic and Tertiary Anura of Laurasia. Palaeobiodiversity and Palaeoenvironments, 93: 397 - 439. https: // doi. org / 10.1007 / s 12549 - 013 - 0131 - y" type="journal article" year="2013">2013</bibRefCitation>
, and reference therein). It is far less distributed on the continent during the Late Pliocene, when only few occurrences in southern
<collectingCountry id="F34BD7E3D071FFADFDB49ABBFDBB74AE" box="[530,613,649,674]" name="France" pageId="37" pageNumber="38">France</collectingCountry>
(
<bibRefCitation id="EFCDEA82D071FFADFDD49ABBFCD374AE" author="Bailon, S." box="[626,781,649,674]" pageId="37" pageNumber="38" refId="ref29802" refString="Bailon, S. 1991. Amphibiens et reptiles du Pliocene et du Quaternaire de France et d'Espagne: mise en place et evolution des faunes. Unpublished PhD Thesis, Universite Paris VII, Paris, France." type="book" year="1991">Bailon, 1991</bibRefCitation>
), central
<collectingCountry id="F34BD7E3D071FFADFE879A9AFE5074CE" box="[289,398,680,706]" name="Germany" pageId="37" pageNumber="38">Germany</collectingCountry>
(
<bibRefCitation id="EFCDEA82D071FFADFE019A9BFD8674CE" author="Bohme, G." box="[423,600,681,706]" pageId="37" pageNumber="38" pagination="79 - 97" refId="ref30380" refString="Bohme, G. 2000. Fossile Amphibien und Reptilien im Quartar Thuringens. Veroffentlichungen Naturkundemuseum Erfurt, 19 / 2000: 79 - 97." type="journal article" year="2000">Böhme, 2000</bibRefCitation>
), northern
<collectingCountry id="F34BD7E3D071FFADFD419A9BFCC674CE" box="[743,792,681,706]" name="Italy" pageId="37" pageNumber="38">Italy</collectingCountry>
(
<bibRefCitation id="EFCDEA82D071FFADFF609AFAFE2474EE" author="Vergnaud-Grazzini, C." box="[198,506,712,738]" pageId="37" pageNumber="38" pagination="47 - 65" refId="ref39445" refString="Vergnaud-Grazzini, C. 1970. Les amphibiens fossiles du gisement d'Arondelli. Palaeontographica Italica, 66: 47 - 65." type="journal article" year="1970">Vergnaud-Grazzini, 1970</bibRefCitation>
), and southern
<collectingCountry id="F34BD7E3D071FFADFD1A9AFBFCC674EE" box="[700,792,713,738]" name="Ukraine" pageId="37" pageNumber="38">Ukraine</collectingCountry>
(
<bibRefCitation id="EFCDEA82D071FFADFF609ADBFEA4750E" author="Ratnikov, V. Yu." box="[198,378,745,770]" pageId="37" pageNumber="38" pagination="171 - 178" refId="ref35842" refString="Ratnikov, V. Yu. 2001. Pliocene anurans of east-European platform. Russian Journal of Herpetology, 8: 171 - 178. https: // doi. org / 10.30906 / 1026 - 2296 - 2001 - 8 - 3 - 171 - 178" type="journal article" year="2001">Ratnikov, 2001</bibRefCitation>
) are known. In the Pleistocene, relict occurrences of this alytid genus in Europe are only reported from northern
<collectingCountry id="F34BD7E3D071FFADFD8B9B1BFD4C754E" box="[557,658,809,834]" name="Hungary" pageId="37" pageNumber="38">Hungary</collectingCountry>
(
<bibRefCitation id="EFCDEA82D071FFADFD0E9B1AFF20756E" author="Szentesi, Z." pageId="37" pageNumber="38" pagination="141 - 158" refId="ref38079" refString="Szentesi, Z. 2019. Latonia gigantea (Lartet, 1851) remains and other herpetological observations from the middle Pleistocene Esztramos 6 locality (Aggtelek - Rudabanya Mountains, North Hungary). Fragmenta Palaeontologica Hungarica, 36: 141 - 158. http: // doi. org / 10.17111 / FragmPalHung. 2019.36.14" type="journal article" year="2019">Szentesi, 2019</bibRefCitation>
) and central
<collectingCountry id="F34BD7E3D071FFADFE3C9B7BFE15756E" box="[410,459,841,866]" name="Italy" pageId="37" pageNumber="38">Italy</collectingCountry>
(
<bibRefCitation id="EFCDEA82D071FFADFE7F9B7AFD16756E" author="Sorbelli, L. &amp; Villa, A. &amp; Gentili, S. &amp; Cherin, M. &amp; Carnevale, G. &amp; Tschopp, E. &amp; Delfino, M." box="[473,712,840,866]" pageId="37" pageNumber="38" pagination="555 - 583" refId="ref37258" refString="Sorbelli, L., Villa, A., Gentili, S., Cherin, M., Carnevale, G., Tschopp, E., and Delfino, M. 2021. The Early Pleistocene ectothermic vertebrates of Pietrafitta (Italy) and the last Western European occurrence of Latonia Meyer, 1843. Comptes Rendus Palevol, 20: 555 - 583. https: // doi. org / 10.5852 / cr-palevol 2021 v 20 a 26" type="journal article" year="2021">Sorbelli et al., 2021</bibRefCitation>
). Early Pleistocene
<taxonomicName id="4C5CECF0D071FFADFEFE9B5BFE71758E" authorityName="von Meyer" authorityYear="1843" box="[344,431,873,898]" class="Amphibia" family="Alytidae" genus="Latonia" kingdom="Animalia" order="Anura" pageId="37" pageNumber="38" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D071FFADFEFE9B5BFE71758E" box="[344,431,873,898]" italics="true" pageId="37" pageNumber="38">Latonia</emphasis>
</taxonomicName>
remains were also found in
<collectingCountry id="F34BD7E3D071FFADFF669BBBFECF75AE" box="[192,273,905,930]" name="Turkey" pageId="37" pageNumber="38">Turkey</collectingCountry>
(
<bibRefCitation id="EFCDEA82D071FFADFE869BBBFDC375AE" author="Vasilyan, D. &amp; Schneider, S. &amp; Bayraktutan, M. S. &amp; Sen, S." box="[288,541,905,930]" pageId="37" pageNumber="38" pagination="293 - 307" refId="ref39052" refString="Vasilyan, D., Schneider, S., Bayraktutan, M. S., and Sen, S. 2014. Early Pleistocene freshwater communities and rodents from the Pasinler Basin (Erzurum Province, north-eastern Turkey). Turkish Journal of Earth Sciences, 23: 293 - 307. https: // doi. org / 10.3906 / yer- 1307 - 16" type="journal article" year="2014">Vasilyan et al., 2014</bibRefCitation>
), but late Quaternary occurrences are limited to
<collectingCountry id="F34BD7E3D071FFADFD849B9BFDBD75CE" box="[546,611,937,962]" name="Israel" pageId="37" pageNumber="38">Israel</collectingCountry>
(
<bibRefCitation id="EFCDEA82D071FFADFDDA9B9BFEDC75EE" author="Biton, R. &amp; Boistel, R. &amp; Rabinovich, R. &amp; Gafny, S. &amp; Brumfeld, V. &amp; Bailon, S." pageId="37" pageNumber="38" pagination="1131 - 1145" refId="ref30095" refString="Biton, R., Boistel, R., Rabinovich, R., Gafny, S., Brumfeld, V., and Bailon, S. 2016. Osteological observations on the alytid Anura Latonia nigriventer with comments on functional morphology, biogeography, and evolutionary history. Journal of Morphology, 277: 1131 - 1145. https: // doi. org / 10.1002 / jmor. 20562" type="journal article" year="2016">Biton et al., 2016</bibRefCitation>
), where the last survivors of these frogs were recently rediscovered (
<bibRefCitation id="EFCDEA82D071FFADFE6C9BDBFD49720E" author="Biton, R. &amp; Geffen, E. &amp; Vences, M. &amp; Cohen, O. &amp; Bailon, S. &amp; Rabinovich, R. &amp; Malka, Y. &amp; Oron, T. &amp; Boistel, R. &amp; Brumfeld, V. &amp; Gafny, S." box="[458,663,1001,1026]" pageId="37" pageNumber="38" pagination="1959" refId="ref30010" refString="Biton, R., Geffen, E., Vences, M., Cohen, O., Bailon, S., Rabinovich, R., Malka, Y., Oron, T., Boistel, R., Brumfeld, V., and Gafny, S. 2013. The rediscovered Hula painted frog is a living fossil. Nature Communications, 4: 1959. https: // doi. org / 10.1038 / ncomms 2959" type="journal article" year="2013">Biton et al., 2013</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D071FFADFD079BDBFEDC722E" author="Perl, R. G. B. &amp; Gafny, S. &amp; Malka, Y. &amp; Renan, S. &amp; Woodhams, D. C. &amp; Rollins-Smith, L. &amp; Pask, J. D. &amp; Bletz, M. C. &amp; Geffen, E. &amp; Vences, M." pageId="37" pageNumber="38" pagination="11 - 37" refId="ref35309" refString="Perl, R. G. B., Gafny, S., Malka, Y., Renan, S., Woodhams, D. C., Rollins-Smith, L., Pask, J. D., Bletz, M. C., Geffen, E., and Vences, M. 2017. Natural history and conservation of the rediscovered Hula painted frog, Latonia nigriventer. Contributions to Zoology, 86: 11 - 37. https: // doi. org / 10.1163 / 18759866 - 08601002" type="journal article" year="2017">Perl et al., 2017</bibRefCitation>
). After the Miocene/Pliocene boundary, the German site of Kaltensundheim is the only place where
<taxonomicName id="4C5CECF0D071FFADFEB19C7BFEB0726E" authorityName="von Meyer" authorityYear="1843" box="[279,366,1097,1122]" class="Amphibia" family="Alytidae" genus="Latonia" kingdom="Animalia" order="Anura" pageId="37" pageNumber="38" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D071FFADFEB19C7BFEB0726E" box="[279,366,1097,1122]" italics="true" pageId="37" pageNumber="38">Latonia</emphasis>
</taxonomicName>
was reported (
<bibRefCitation id="EFCDEA82D071FFADFD959C7BFD09726E" author="Bohme, G." box="[563,727,1097,1122]" pageId="37" pageNumber="38" pagination="79 - 97" refId="ref30380" refString="Bohme, G. 2000. Fossile Amphibien und Reptilien im Quartar Thuringens. Veroffentlichungen Naturkundemuseum Erfurt, 19 / 2000: 79 - 97." type="journal article" year="2000">Böhme 2000</bibRefCitation>
) that reaches comparable northern latitudes with Hambach. However,
<bibRefCitation id="EFCDEA82D071FFADFE2F9CBBFDE872AE" author="Bohme, G." box="[393,566,1161,1186]" pageId="37" pageNumber="38" pagination="231 - 238" refId="ref30406" refString="Bohme, G. 2002. Amphibienreste aus dem Oberpliozan von Kaltensundheim (Rhon, Thuringen). Mitteilungen des Museum fur Naturkunde, Berlin, Geowissenschaftliche Reihe, 5: 231 - 238." type="journal article" year="2002">Böhme (2002)</bibRefCitation>
found no trace of
<taxonomicName id="4C5CECF0D071FFADFF669C9BFEC972CE" authorityName="von Meyer" authorityYear="1843" box="[192,279,1193,1218]" class="Amphibia" family="Alytidae" genus="Latonia" kingdom="Animalia" order="Anura" pageId="37" pageNumber="38" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D071FFADFF669C9BFEC972CE" box="[192,279,1193,1218]" italics="true" pageId="37" pageNumber="38">Latonia</emphasis>
</taxonomicName>
among the fossil material from this Thuringian site and recommended its removal from the taxon list of the assemblage. Thus, Hambach currently stands as the northernmost confirmed occurrence of
<taxonomicName id="4C5CECF0D071FFADFE279D1BFE06734E" authorityName="von Meyer" authorityYear="1843" box="[385,472,1321,1346]" class="Amphibia" family="Alytidae" genus="Latonia" kingdom="Animalia" order="Anura" pageId="37" pageNumber="38" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D071FFADFE279D1BFE06734E" box="[385,472,1321,1346]" italics="true" pageId="37" pageNumber="38">Latonia</emphasis>
</taxonomicName>
from the late Neogene onwards.
</paragraph>
<paragraph id="8BE39773D071FFADFF5C9D5BFAEC75CE" blockId="37.[192,793,169,1890]" lastBlockId="37.[840,1441,168,1890]" pageId="37" pageNumber="38">
Palaeobatrachids are almost completely absent from western Europe following the Miocene/Pliocene transition (
<bibRefCitation id="EFCDEA82D071FFADFDA39D9BFCCB73CE" author="Wuttke, M. &amp; Prikryl, T. &amp; Ratnikov, V. Yu. &amp; Dvorak, Z. &amp; Rocek, Z." box="[517,789,1449,1474]" pageId="37" pageNumber="38" pagination="367 - 395" refId="ref40324" refString="Wuttke, M., Prikryl, T., Ratnikov, V. Yu., Dvorak, Z., and Rocek, Z. 2012. Generic diversity and distributional dynamics of the Palaeobatrachidae (Amphibia: Anura). Palaeobiodiversity and Palaeoenvironments, 92: 367 - 395. https: // doi. org / 10.1007 / s 12549 - 012 - 0071 - y" type="journal article" year="2012">Wuttke et al., 2012</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D071FFADFF669DFAFEBE73EE" author="Rocek, Z." box="[192,352,1480,1506]" pageId="37" pageNumber="38" pagination="397 - 439" refId="ref36270" refString="Rocek, Z. 2013. Mesozoic and Tertiary Anura of Laurasia. Palaeobiodiversity and Palaeoenvironments, 93: 397 - 439. https: // doi. org / 10.1007 / s 12549 - 013 - 0131 - y" type="journal article" year="2013">Roček, 2013</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D071FFADFED79DFAFDBE73EE" author="Rocek, Z. &amp; Rage, J. - C. &amp; Venczel, M." box="[369,608,1480,1506]" pageId="37" pageNumber="38" pagination="1 - 151" refId="ref36384" refString="Rocek, Z., Rage, J. - C., and Venczel, M. 2021. Fossil frogs of the genus Palaeobatrachus (Amphibia: Anura). Abhandlungen der Senckenberg Gesellschaft fur Naturforschung, 575: 1 - 151." type="journal article" year="2021">Roček et al., 2021</bibRefCitation>
), with the only notable exceptions being Tegelen, in
<collectingCountry id="F34BD7E3D071FFADFD219DDBFEDE702E" name="Netherlands" pageId="37" pageNumber="38">The Netherlands</collectingCountry>
(
<bibRefCitation id="EFCDEA82D071FFADFEB29E3BFE35702E" author="Villa, A. &amp; Rocek, Z. &amp; Tschopp, E. &amp; van den Hoek Ostende, L. W. &amp; Delfino, M." box="[276,491,1545,1570]" pageId="37" pageNumber="38" pagination="1211664" refId="ref39607" refString="Villa, A., Rocek, Z., Tschopp, E., van den Hoek Ostende, L. W., and Delfino, M. 2016. Palaeobatrachus eurydices, sp. nov. (Amphibia, Anura), the last western European palaeobatrachid. Journal of Vertebrate Paleontology, 36: e 1211664. https: // doi. org / 10.1080 / 02724634.2016.1211664" type="journal article" year="2016">Villa et al., 2016</bibRefCitation>
), and Hambach 11 and 11C. These localities share the same palaeobatrachid species, which is currently not known elsewhere, and are geographically close, even though the Dutch site is younger, being Lower Pleistocene (Gelasian) in age. Other Pliocene and Quaternary occurrences of palaeobatrachids are all located far eastward. The closest one in geographical terms to Tegelen and Hambach, and only other post-Miocene site with palaeobatrachids in
<collectingCountry id="F34BD7E3D071FFADFD249F1AFD2C714E" box="[642,754,1832,1858]" name="Germany" pageId="37" pageNumber="38">Germany</collectingCountry>
, is possibly Voigtstedt, a Chibanian locality in
<collectingRegion id="49985991D071FFADFCEE989BFC6576CE" box="[840,955,169,194]" country="Germany" name="Thueringen" pageId="37" pageNumber="38">Thuringia</collectingRegion>
, from which
<taxonomicName id="4C5CECF0D071FFADFBCF989BFBCB76EE" authority="(Fejervary, 1917)" baseAuthorityName="Fejervary" baseAuthorityYear="1917" class="Amphibia" family="Palaeobatrachidae" genus="Palaeobatrachus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="37" pageNumber="38" phylum="Chordata" rank="species" species="langhae">
<emphasis id="B9284B61D071FFADFBCF989BFA7E76CE" box="[1129,1440,169,194]" italics="true" pageId="37" pageNumber="38">Palaeobatrachus langhae</emphasis>
(
<bibRefCitation id="EFCDEA82D071FFADFCE898FAFBD176EE" author="Fejervary, G. J." box="[846,1039,200,226]" pageId="37" pageNumber="38" pagination="141 - 199" refId="ref31988" refString="Fejervary, G. J. 1917. Anoures fossiles des couches preglaciaires de Puspokfurdo en Hongrie. Foldtani Kozlony, 47: 141 - 199." type="journal article" year="1917">Fejérváry, 1917</bibRefCitation>
)
</taxonomicName>
was reported by
<bibRefCitation id="EFCDEA82D071FFADFB5298FBFA7E76EE" author="Kretzoi, M." box="[1268,1440,201,226]" pageId="37" pageNumber="38" pagination="325 - 333" refId="ref33823" refString="Kretzoi, M. 1965. Die Amphibien aus der altpleistozanen Fundstelle Voigtstedt in Thuringen. Palaontologische Abhandlungen Abteilung A, 2: 325 - 333." type="journal article" year="1965">Kretzoi (1965)</bibRefCitation>
and later mentioned as a tentative occurrence by
<bibRefCitation id="EFCDEA82D071FFADFCEE993BFBD2772E" author="Holman, J. A." box="[840,1036,265,290]" pageId="37" pageNumber="38" pagination="1 - 254" refId="ref33244" refString="Holman, J. A. 1998. Pleistocene amphibians and reptiles in Britain and Europe. Oxford monographs on geology and geophysics, 38: 1 - 254." type="journal article" year="1998">Holman (1998)</bibRefCitation>
,
<bibRefCitation id="EFCDEA82D071FFADFB81993AFB30772E" author="Sanchiz, B." box="[1063,1262,264,290]" pageId="37" pageNumber="38" refId="ref36500" refString="Sanchiz, B. 1998. Handbuch der Palaoherpetologie. 4 - Salientia. Friedrich Pfeil, Munchen." type="book" year="1998">Sanchiz (1998)</bibRefCitation>
, and
<bibRefCitation id="EFCDEA82D071FFADFAEA993BFC47774E" author="Bohme, G." pageId="37" pageNumber="38" pagination="79 - 97" refId="ref30380" refString="Bohme, G. 2000. Fossile Amphibien und Reptilien im Quartar Thuringens. Veroffentlichungen Naturkundemuseum Erfurt, 19 / 2000: 79 - 97." type="journal article" year="2000">Böhme (2000)</bibRefCitation>
. There is no other Miocene occurrence of palaeobatrachids in the northwestern part of continental Europe surrounding Hambach and Tegelen (
<bibRefCitation id="EFCDEA82D071FFADFCE899BBFBEB77AE" author="Wuttke, M. &amp; Prikryl, T. &amp; Ratnikov, V. Yu. &amp; Dvorak, Z. &amp; Rocek, Z." box="[846,1077,393,418]" pageId="37" pageNumber="38" pagination="367 - 395" refId="ref40324" refString="Wuttke, M., Prikryl, T., Ratnikov, V. Yu., Dvorak, Z., and Rocek, Z. 2012. Generic diversity and distributional dynamics of the Palaeobatrachidae (Amphibia: Anura). Palaeobiodiversity and Palaeoenvironments, 92: 367 - 395. https: // doi. org / 10.1007 / s 12549 - 012 - 0071 - y" type="journal article" year="2012">Wuttke et al., 2012</bibRefCitation>
), and the estimated closest relatives to the Plio-Pleistocene species
<taxonomicName id="4C5CECF0D071FFADFAA9999BFA7E77CE" box="[1295,1440,425,450]" class="Amphibia" family="Palaeobatrachidae" genus="Palaeobatrachus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="37" pageNumber="38" phylum="Chordata" rank="species" species="eurydices">
<emphasis id="B9284B61D071FFADFAA9999BFA7E77CE" box="[1295,1440,425,450]" italics="true" pageId="37" pageNumber="38">P. eurydices</emphasis>
</taxonomicName>
are to be found in the Lower Miocene of southern
<collectingCountry id="F34BD7E3D071FFADFCEE99DBFC45740E" box="[840,923,489,514]" name="France" pageId="37" pageNumber="38">France</collectingCountry>
and central
<collectingCountry id="F34BD7E3D071FFADFBE999DAFB63740E" box="[1103,1213,488,514]" name="Germany" pageId="37" pageNumber="38">Germany</collectingCountry>
(
<taxonomicName id="4C5CECF0D071FFADFB7E99DBFB26742E" authority="Hossini and Rage, 2000" authorityName="Hossini and Rage" authorityYear="2000" class="Amphibia" family="Palaeobatrachidae" genus="Palaeobatrachus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="37" pageNumber="38" phylum="Chordata" rank="species" species="robustus">
<emphasis id="B9284B61D071FFADFB7E99DBFC70742E" italics="true" pageId="37" pageNumber="38">Palaeobatrachus robustus</emphasis>
<bibRefCitation id="EFCDEA82D071FFADFC189A3BFB26742E" author="Hossini, S. &amp; Rage, J. - C." box="[958,1272,521,546]" pageId="37" pageNumber="38" pagination="223 - 231" refId="ref33317" refString="Hossini, S. and Rage, J. - C. 2000. Palaeobatrachid frogs from the earliest Miocene (Agenian) of France, with description of a new species. Geobios, 33: 223 - 231. https: // doi. org / 10.1016 / S 0016 - 6995 (00) 80019 - 4" type="journal article" year="2000">Hossini and Rage, 2000</bibRefCitation>
</taxonomicName>
;
<bibRefCitation id="EFCDEA82D071FFADFAAA9A3BFC08744E" author="Hossini, S. &amp; Rage, J. - C." pageId="37" pageNumber="38" pagination="223 - 231" refId="ref33317" refString="Hossini, S. and Rage, J. - C. 2000. Palaeobatrachid frogs from the earliest Miocene (Agenian) of France, with description of a new species. Geobios, 33: 223 - 231. https: // doi. org / 10.1016 / S 0016 - 6995 (00) 80019 - 4" type="journal article" year="2000">Hossini and Rage, 2000</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D071FFADFC479A1AFBA5744E" author="Rocek, Z." box="[993,1147,552,578]" pageId="37" pageNumber="38" pagination="397 - 439" refId="ref36270" refString="Rocek, Z. 2013. Mesozoic and Tertiary Anura of Laurasia. Palaeobiodiversity and Palaeoenvironments, 93: 397 - 439. https: // doi. org / 10.1007 / s 12549 - 013 - 0131 - y" type="journal article" year="2013">Roček, 2013</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D071FFADFB219A1BFA97744E" author="Villa, A. &amp; Rocek, Z. &amp; Tschopp, E. &amp; van den Hoek Ostende, L. W. &amp; Delfino, M." box="[1159,1353,553,578]" pageId="37" pageNumber="38" pagination="1211664" refId="ref39607" refString="Villa, A., Rocek, Z., Tschopp, E., van den Hoek Ostende, L. W., and Delfino, M. 2016. Palaeobatrachus eurydices, sp. nov. (Amphibia, Anura), the last western European palaeobatrachid. Journal of Vertebrate Paleontology, 36: e 1211664. https: // doi. org / 10.1080 / 02724634.2016.1211664" type="journal article" year="2016">Villa et al., 2016</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D071FFADFAF29A1AFC0A746E" author="Rocek, Z. &amp; Rage, J. - C. &amp; Venczel, M." pageId="37" pageNumber="38" pagination="1 - 151" refId="ref36384" refString="Rocek, Z., Rage, J. - C., and Venczel, M. 2021. Fossil frogs of the genus Palaeobatrachus (Amphibia: Anura). Abhandlungen der Senckenberg Gesellschaft fur Naturforschung, 575: 1 - 151." type="journal article" year="2021">Roček et al., 2021</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D071FFADFC479A7BFB03746E" author="Vasilyan, D. &amp; Cernansky, A. &amp; Szyndlar, Z. &amp; Mors, T." box="[993,1245,585,610]" pageId="37" pageNumber="38" pagination="99 - 145" refId="ref39186" refString="Vasilyan, D., Cernansky, A., Szyndlar, Z., and Mors, T. 2022. Amphibian and reptilian fauna from the early Miocene of Echzell, Germany. Fossil Record, 25: 99 - 145. https: // doi. org / 10.3897 / fr. 25.83781" type="journal article" year="2022">Vasilyan et al., 2022</bibRefCitation>
). In this context, the fragmentary maxilla from Hambach 6C testifies that these water-dwelling frogs likely inhabited the area during the early Neogene as well, even though the preservational status prevents both a confident attribution at family rank and even more any inference on the possible persistence of the lineage leading to
<emphasis id="B9284B61D071FFADFB9F9B1BFB06754E" box="[1081,1240,809,834]" italics="true" pageId="37" pageNumber="38">
<taxonomicName id="4C5CECF0D071FFADFB9F9B1BFB0A754E" box="[1081,1236,809,834]" class="Amphibia" family="Palaeobatrachidae" genus="Palaeobatrachus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="37" pageNumber="38" phylum="Chordata" rank="species" species="eurydices">P. eurydices</taxonomicName>
.
</emphasis>
More Neogene palaeobatrachid fossils from countries such as
<collectingCountry id="F34BD7E3D071FFADFAD79B7BFC58758E" name="Belgium" pageId="37" pageNumber="38">Belgium</collectingCountry>
,
<collectingCountry id="F34BD7E3D071FFADFC319B5BFBB8758E" box="[919,1126,873,898]" name="Netherlands" pageId="37" pageNumber="38">The Netherlands</collectingCountry>
, and (northwestern)
<collectingCountry id="F34BD7E3D071FFADFACD9B5AFC5775AE" name="Germany" pageId="37" pageNumber="38">Germany</collectingCountry>
are needed to shed light on the early evolution of this possibly relictual lineage.
</paragraph>
<paragraph id="8BE39773D071FFAEFC249BFBFCC772D3" blockId="37.[840,1441,168,1890]" lastBlockId="38.[192,793,166,1887]" lastPageId="38" lastPageNumber="39" pageId="37" pageNumber="38">
A situation similar to that of palaeobatrachids arises for pelobatids from Hambach as well. Several pelobatids are reported from the Palaeogene and Neogene of Europe (e.g.,
<bibRefCitation id="EFCDEA82D071FFADFB439C1AFA43724E" author="Sanchiz, B." box="[1253,1437,1064,1090]" pageId="37" pageNumber="38" refId="ref36500" refString="Sanchiz, B. 1998. Handbuch der Palaoherpetologie. 4 - Salientia. Friedrich Pfeil, Munchen." type="book" year="1998">Sanchiz, 1998</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D071FFADFCEE9C7AFC3C726E" author="Rocek, Z." box="[840,994,1096,1122]" pageId="37" pageNumber="38" pagination="397 - 439" refId="ref36270" refString="Rocek, Z. 2013. Mesozoic and Tertiary Anura of Laurasia. Palaeobiodiversity and Palaeoenvironments, 93: 397 - 439. https: // doi. org / 10.1007 / s 12549 - 013 - 0131 - y" type="journal article" year="2013">Roček, 2013</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D071FFADFC4B9C7BFAE7726E" author="Vergilov, V. &amp; Tzankov, N." box="[1005,1337,1097,1122]" pageId="37" pageNumber="38" pagination="29 - 42" refId="ref39401" refString="Vergilov, V. and Tzankov, N. 2021. Giant pelobatid fossil larva from the middle Miocene of Bulgaria. Historia Naturalis Bulgarica, 43: 29 - 42. https: // doi. org / 10.48027 / hnb. 43.031" type="journal article" year="2021">Vergilov and Tzankov, 2021</bibRefCitation>
; and reference therein), but only few of them come from the northwestern part of the continent, where Hambach is located. Miocene pelobatids found in the area come from only three localities placed slightly eastward to the latter site: an Aquitanian indeterminate member of the family from Budenheim, near Mainz, mentioned, but neither described nor figured, by
<bibRefCitation id="EFCDEA82D071FFADFC6D9D7AFB48736D" author="Schleich, H. H." box="[971,1174,1352,1378]" pageId="37" pageNumber="38" pagination="289 - 306" refId="ref36806" refString="Schleich, H. H. 1988. Palaoherpetologische Materialien und Faunenspektren aus dem Kalktertiar des Mainzer Beckens (Oberoligozan - Untermiozan). Geologisches Jahrbuch, A 110: 289 - 306." type="journal article" year="1988">Schleich (1988)</bibRefCitation>
; several Burdigalian remains of
<taxonomicName id="4C5CECF0D071FFADFC699D5AFABF738D" authority="Venczel, 2004" authorityName="Venczel" authorityYear="2004" box="[975,1377,1384,1409]" class="Amphibia" family="Pelobatidae" genus="Pelobates" kingdom="Animalia" order="Anura" pageId="37" pageNumber="38" phylum="Chordata" rank="species" species="sanchizi">
<emphasis id="B9284B61D071FFADFC699D5AFB71738D" box="[975,1199,1384,1409]" italics="true" pageId="37" pageNumber="38">Pelobates sanchizi</emphasis>
<bibRefCitation id="EFCDEA82D071FFADFB1E9D5AFABF738D" author="Venczel, M." box="[1208,1377,1384,1409]" pageId="37" pageNumber="38" pagination="151 - 174" refId="ref39308" refString="Venczel, M. 2004. Middle Miocene anurans from the Carpathian Basin. Palaeontographica Abt. A, 271: 151 - 174. https: // doi. org / 10.1127 / pala / 271 / 2004 / 151" type="journal article" year="2004">Venczel, 2004</bibRefCitation>
</taxonomicName>
from Echzell (
<bibRefCitation id="EFCDEA82D071FFADFC099DBAFB7073AD" author="Vasilyan, D. &amp; Cernansky, A. &amp; Szyndlar, Z. &amp; Mors, T." box="[943,1198,1416,1441]" pageId="37" pageNumber="38" pagination="99 - 145" refId="ref39186" refString="Vasilyan, D., Cernansky, A., Szyndlar, Z., and Mors, T. 2022. Amphibian and reptilian fauna from the early Miocene of Echzell, Germany. Fossil Record, 25: 99 - 145. https: // doi. org / 10.3897 / fr. 25.83781" type="journal article" year="2022">Vasilyan et al., 2022</bibRefCitation>
); and a Middle Miocene premetamorphic tadpole from Climbach, near Allendorf, attributed to
<taxonomicName id="4C5CECF0D071FFADFBC69DFAFAFD73ED" box="[1120,1315,1480,1505]" class="Amphibia" family="Pelobatidae" genus="Eopelobates" kingdom="Animalia" order="Anura" pageId="37" pageNumber="38" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D071FFADFBC69DFAFB2A73ED" box="[1120,1268,1480,1505]" italics="true" pageId="37" pageNumber="38">Eopelobates</emphasis>
sp.
</taxonomicName>
(originally referred to
<taxonomicName id="4C5CECF0D071FFADFC429DDAFC5D702D" authority="Tschudi, 1838" authorityName="Tschudi" authorityYear="1838" class="Amphibia" family="Palaeobatrachidae" genus="Palaeobatrachus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="37" pageNumber="38" phylum="Chordata" rank="species" species="goldfussi">
<emphasis id="B9284B61D071FFADFC429DDAFAF4700D" box="[996,1322,1512,1537]" italics="true" pageId="37" pageNumber="38">Palaeobatrachus goldfussi</emphasis>
<bibRefCitation id="EFCDEA82D071FFADFA989DDAFC5D702D" author="Tschudi, J. J." pageId="37" pageNumber="38" refId="ref38467" refString="Tschudi, J. J. 1838. Classification der Batrachier, mit Berucksichtigung der fossilen Thiere dieser Abteilung der Reptilien. Petitpierre, Neuchatel." type="book" year="1838">Tschudi, 1838</bibRefCitation>
</taxonomicName>
by
<bibRefCitation id="EFCDEA82D071FFADFC149E3AFBA1702D" author="von Meyer, H." box="[946,1151,1544,1569]" pageId="37" pageNumber="38" pagination="123 - 182" refId="ref40026" refString="von Meyer, H. 1860. Frosche aus den Tertiar-Gebilden Deutschlands. Palaeontographica, 7: 123 - 182." type="journal article" year="1860">von Meyer, 1860</bibRefCitation>
, but later reidentified by
<bibRefCitation id="EFCDEA82D071FFADFCEE9E16FC3B704D" author="Spinar, Z. V." box="[840,997,1572,1602]" pageId="37" pageNumber="38" refId="ref37414" refString="Spinar, Z. V. 1972. Tertiary frogs from Central Europe. Academia, Prague." type="book" year="1972">Špinar, 1972</bibRefCitation>
; see
<bibRefCitation id="EFCDEA82D071FFADFB8D9E1AFB3E704D" author="Gardner, J. D." box="[1067,1248,1576,1602]" pageId="37" pageNumber="38" pagination="17 - 44" refId="ref32175" refString="Gardner, J. D. 2016. The fossil record of tadpoles. Fossil Imprint, 72: 17 - 44. https: // doi. org / 10.14446 / FI. 2016.17" type="journal article" year="2016">Gardner, 2016</bibRefCitation>
). The two fossil bones from Hambach 6C are evidence that the range of pelobatids extended further west in this part of Europe in the Middle Miocene, but it is currently impossible to figure out whether or not they were related to either the taxa living in nearby areas in Palaeogene times (e.g.,
<taxonomicName id="4C5CECF0D071FFADFB769EDAFBD7712D" authority="Spinar, 1952" authorityName="Spinar" authorityYear="1952" class="Amphibia" family="Pelobatidae" genus="Eopelobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="37" pageNumber="38" phylum="Chordata" rank="species" species="bayeri">
<emphasis id="B9284B61D071FFADFB769EDAFCB8712D" italics="true" pageId="37" pageNumber="38">Eopelobates bayeri</emphasis>
<bibRefCitation id="EFCDEA82D071FFADFCD69F36FBD7712D" author="Spinar, Z. V." box="[880,1033,1796,1826]" pageId="37" pageNumber="38" pagination="457 - 488" refId="ref37383" refString="Spinar, Z. V. 1952. Eopelobates bayeri - a new frog from the Tertiary of Bohemia. Sbornik Ustredniho ustavu geologickeho, 19: 457 - 488." type="journal article" year="1952">Špinar, 1952</bibRefCitation>
</taxonomicName>
from
<collectingCountry id="F34BD7E3D071FFADFBF39F3AFB64712D" box="[1109,1210,1800,1825]" name="Belgium" pageId="37" pageNumber="38">Belgium</collectingCountry>
,
<taxonomicName id="4C5CECF0D071FFADFB619F3AFB8E714D" authority="(Weitzel, 1938)" baseAuthorityName="Weitzel" baseAuthorityYear="1938" class="Amphibia" family="Pelobatidae" genus="Eopelobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="37" pageNumber="38" phylum="Chordata" rank="species" species="wagneri">
<emphasis id="B9284B61D071FFADFB619F3AFCAA714D" italics="true" pageId="37" pageNumber="38">Eopelobates wagneri</emphasis>
(
<bibRefCitation id="EFCDEA82D071FFADFC359F1AFB94714D" author="Weitzel, K." box="[915,1098,1832,1857]" pageId="37" pageNumber="38" pagination="42 - 46" refId="ref40141" refString="Weitzel, K. 1938. Propelodytes wagneri n. g. n. sp., ein Frosch aus dem Mittel-Eozan von Messel. Notizblatt der Hessischen Geologischen Landes-Anstalt, 19: 42 - 46." type="journal article" year="1938">Weitzel, 1938</bibRefCitation>
)
</taxonomicName>
from western
<collectingCountry id="F34BD7E3D071FFADFA8D9F1AFA43714E" box="[1323,1437,1832,1858]" name="Germany" pageId="37" pageNumber="38">Germany</collectingCountry>
;
<bibRefCitation id="EFCDEA82D071FFADFCEE9F7AFC34716D" author="Rocek, Z." box="[840,1002,1864,1889]" pageId="37" pageNumber="38" pagination="397 - 439" refId="ref36270" refString="Rocek, Z. 2013. Mesozoic and Tertiary Anura of Laurasia. Palaeobiodiversity and Palaeoenvironments, 93: 397 - 439. https: // doi. org / 10.1007 / s 12549 - 013 - 0131 - y" type="journal article" year="2013">Roček, 2013</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D071FFADFC5B9F7AFB2E716D" author="Rocek, Z. &amp; Wuttke, M. &amp; Gardner, J. D. &amp; Bhullar, B. - A. S." box="[1021,1264,1864,1889]" pageId="37" pageNumber="38" pagination="529 - 567" refId="ref36310" refString="Rocek, Z., Wuttke, M., Gardner, J. D., and Bhullar, B. - A. S. 2014. The Euro-American genus Eopelobates, and a re-definition of the family Pelobatidae (Amphibia, Anura). Palaeobiodiversity and Palaeoenvironments, 94: 529 - 567. https: // doi. org / 10.1007 / s 12549 - 014 - 0169 - 5" type="journal article" year="2014">Roček et al., 2014</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D071FFAEFAA59F7AFEDC76B3" author="Smith, K. T. &amp; Schaal, S. F. K. &amp; Habersetzer, J." lastPageId="38" lastPageNumber="39" pageId="37" pageNumber="38" refId="ref37184" refString="Smith, K. T., Schaal, S. F. K., and Habersetzer, J. 2018. Messel. An ancient greenhouse ecosystem. Senckenberg Gesellschaft fur Naturforschung, Frankfurt am Main." type="book" year="2018">Smith et al., 2018</bibRefCitation>
), the Early/Middle Miocene above-mentioned ones from western
<collectingCountry id="F34BD7E3D072FFAEFE0998F4FDC176EC" box="[431,543,198,224]" name="Germany" pageId="38" pageNumber="39">Germany</collectingCountry>
, or even those from the youngest level in Hambach. If correctly identified, the Late Pliocene
<taxonomicName id="4C5CECF0D072FFAEFE689934FDBC7713" authorityName="Parker" authorityYear="1929" box="[462,610,262,287]" class="Amphibia" family="Pelobatidae" genus="Eopelobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="38" pageNumber="39" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D072FFAEFE689934FDBC7713" box="[462,610,262,287]" italics="true" pageId="38" pageNumber="39">Eopelobates</emphasis>
</taxonomicName>
from Hambach 11 and 11C, on the other hand, shares potentially no relations with other pelobatids identified at genus rank in close-by post-Miocene sites. Known congeneric occurrences postdating the Miocene/ Pliocene transition are all located in Eastern Europe (
<bibRefCitation id="EFCDEA82D072FFAEFE8F99F4FE3777D3" author="Mlynarski, M." box="[297,489,454,479]" pageId="38" pageNumber="39" pagination="177 - 194" refId="ref34795" refString="Mlynarski, M. 1962. Notes on the amphibian and reptilian fauna of the Polish Pliocene and early Pleistocene. Acta Zoologica Cracoviensis, 7: 177 - 194." type="journal article" year="1962">Młynarski 1962</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFE5C99F4FED077F3" author="Sanchiz, B. &amp; Mlynarski, M." pageId="38" pageNumber="39" pagination="153 - 174" refId="ref36519" refString="Sanchiz, B. and Mlynarski, M. 1979 a. Remarks on the fossil anurans from the Polish Neogene. Acta Zoologica Cracoviensia, 24: 153 - 174." type="journal article" year="1979">Sanchiz and Młynarski, 1979a</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFE8799D4FD4677F3" author="Mlynarski, M. &amp; Szyndlar, Z." box="[289,664,486,512]" pageId="38" pageNumber="39" pagination="69 - 88" refId="ref34826" refString="Mlynarski, M. and Szyndlar, Z. 1989. Plazy i gady. Folia Quaternaria, 59 - 60: 69 - 88." type="journal article" year="1989">Młynarski and Szyndlar, 1989</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFD0D99D4FF217413" author="Ratnikov, V. Yu." pageId="38" pageNumber="39" pagination="171 - 178" refId="ref35842" refString="Ratnikov, V. Yu. 2001. Pliocene anurans of east-European platform. Russian Journal of Herpetology, 8: 171 - 178. https: // doi. org / 10.30906 / 1026 - 2296 - 2001 - 8 - 3 - 171 - 178" type="journal article" year="2001">Ratnikov, 2001</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFEA89A34FE607413" author="Venczel, M." box="[270,446,518,543]" pageId="38" pageNumber="39" pagination="79 - 90" refId="ref39273" refString="Venczel, M. 2001. Anurans and squamates from the Lower Pliocene (MN 14) Osztramos 1 locality (Northern Hungary). Fragmenta Palaeontologica Hungarica, 19: 79 - 90." type="journal article" year="2001">Venczel, 2001</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFE689A34FDB27413" author="Rocek, Z." box="[462,620,518,543]" pageId="38" pageNumber="39" pagination="397 - 439" refId="ref36270" refString="Rocek, Z. 2013. Mesozoic and Tertiary Anura of Laurasia. Palaeobiodiversity and Palaeoenvironments, 93: 397 - 439. https: // doi. org / 10.1007 / s 12549 - 013 - 0131 - y" type="journal article" year="2013">Roček, 2013</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFDDD9A34FF217433" author="Rocek, Z. &amp; Wuttke, M. &amp; Gardner, J. D. &amp; Bhullar, B. - A. S." pageId="38" pageNumber="39" pagination="529 - 567" refId="ref36310" refString="Rocek, Z., Wuttke, M., Gardner, J. D., and Bhullar, B. - A. S. 2014. The Euro-American genus Eopelobates, and a re-definition of the family Pelobatidae (Amphibia, Anura). Palaeobiodiversity and Palaeoenvironments, 94: 529 - 567. https: // doi. org / 10.1007 / s 12549 - 014 - 0169 - 5" type="journal article" year="2014">Roček et al., 2014</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFEA99A14FDB47433" author="Vergilov, V. &amp; Tzankov, N." box="[271,618,550,575]" pageId="38" pageNumber="39" pagination="29 - 42" refId="ref39401" refString="Vergilov, V. and Tzankov, N. 2021. Giant pelobatid fossil larva from the middle Miocene of Bulgaria. Historia Naturalis Bulgarica, 43: 29 - 42. https: // doi. org / 10.48027 / hnb. 43.031" type="journal article" year="2021">Vergilov and Tzankov, 2021</bibRefCitation>
), even though
<bibRefCitation id="EFCDEA82D072FFAEFF669A74FE3C7453" author="Rage, J. - C. &amp; Rocek, Z." box="[192,482,582,607]" pageId="38" pageNumber="39" pagination="133 - 167" refId="ref35799" refString="Rage, J. - C. and Rocek, Z. 2003. Evolution of anuran assemblages in the Tertiary and Quaternary of Europe, in the context of palaeoclimate and palaeogeography. Amphibia- Reptilia, 24: 133 - 167." type="journal article" year="2003">Rage and Roček (2003)</bibRefCitation>
,
<bibRefCitation id="EFCDEA82D072FFAEFE4B9A74FD4C7453" author="Rocek, Z." box="[493,658,582,607]" pageId="38" pageNumber="39" pagination="397 - 439" refId="ref36270" refString="Rocek, Z. 2013. Mesozoic and Tertiary Anura of Laurasia. Palaeobiodiversity and Palaeoenvironments, 93: 397 - 439. https: // doi. org / 10.1007 / s 12549 - 013 - 0131 - y" type="journal article" year="2013">Roček (2013)</bibRefCitation>
, and
<bibRefCitation id="EFCDEA82D072FFAEFD749A74FE037473" author="Vergilov, V. &amp; Tzankov, N." pageId="38" pageNumber="39" pagination="29 - 42" refId="ref39401" refString="Vergilov, V. and Tzankov, N. 2021. Giant pelobatid fossil larva from the middle Miocene of Bulgaria. Historia Naturalis Bulgarica, 43: 29 - 42. https: // doi. org / 10.48027 / hnb. 43.031" type="journal article" year="2021">Vergilov and Tzankov (2021)</bibRefCitation>
recommended caution on these occurrences due to the absence of the most diagnostic cranial elements in the respective material. Nevertheless, only
<taxonomicName id="4C5CECF0D072FFAEFE4E9AF4FD8074D3" authorityName="Wagler" authorityYear="1830" box="[488,606,710,735]" class="Amphibia" family="Pelobatidae" genus="Pelobates" kingdom="Animalia" order="Anura" pageId="38" pageNumber="39" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D072FFAEFE4E9AF4FD8074D3" box="[488,606,710,735]" italics="true" pageId="38" pageNumber="39">Pelobates</emphasis>
</taxonomicName>
is recorded in the western part of the continent in the last 5 Myr. Apart from the fossils from Hambach, the other youngest published records of
<taxonomicName id="4C5CECF0D072FFAEFDFA9B14FD2E7533" authorityName="Parker" authorityYear="1929" box="[604,752,806,831]" class="Amphibia" family="Pelobatidae" genus="Eopelobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="38" pageNumber="39" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D072FFAEFDFA9B14FD2E7533" box="[604,752,806,831]" italics="true" pageId="38" pageNumber="39">Eopelobates</emphasis>
</taxonomicName>
in western Europe are Middle Miocene in age (
<bibRefCitation id="EFCDEA82D072FFAEFF609B53FEB97573" author="Spinar, Z. V." box="[198,359,865,895]" pageId="38" pageNumber="39" refId="ref37414" refString="Spinar, Z. V. 1972. Tertiary frogs from Central Europe. Academia, Prague." type="book" year="1972">Špinar, 1972</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFED09B54FDC37573" author="Bohme, M." box="[374,541,870,895]" pageId="38" pageNumber="39" pagination="3 - 41" refId="ref30538" refString="Bohme, M. 2010. Ectothermic vertebrates (Actinopterygii, Allocaudata, Urodela, Anura, Crocodylia, Squamata) from the Miocene of Sandelzhausen (Germany, Bavaria) and their implications for environment reconstruction and palaeoclimate. Palaontologische Zeitschrift, 84: 3 - 41. https: // doi. org / 10.1007 / s 12542 - 010 - 0050 - 4" type="journal article" year="2010">Böhme, 2010</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFD8A9B54FCCA7573" author="Rocek, Z. &amp; Wuttke, M. &amp; Gardner, J. D. &amp; Bhullar, B. - A. S." box="[556,788,870,895]" pageId="38" pageNumber="39" pagination="529 - 567" refId="ref36310" refString="Rocek, Z., Wuttke, M., Gardner, J. D., and Bhullar, B. - A. S. 2014. The Euro-American genus Eopelobates, and a re-definition of the family Pelobatidae (Amphibia, Anura). Palaeobiodiversity and Palaeoenvironments, 94: 529 - 567. https: // doi. org / 10.1007 / s 12549 - 014 - 0169 - 5" type="journal article" year="2014">Roček et al., 2014</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFF669BB4FEAA7593" author="Gardner, J. D." box="[192,372,902,928]" pageId="38" pageNumber="39" pagination="17 - 44" refId="ref32175" refString="Gardner, J. D. 2016. The fossil record of tadpoles. Fossil Imprint, 72: 17 - 44. https: // doi. org / 10.14446 / FI. 2016.17" type="journal article" year="2016">Gardner, 2016</bibRefCitation>
). If our identification is correct, the possible Late Pliocene Hambach
<taxonomicName id="4C5CECF0D072FFAEFDFB9B94FD2F75B3" authorityName="Parker" authorityYear="1929" box="[605,753,934,959]" class="Amphibia" family="Pelobatidae" genus="Eopelobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="38" pageNumber="39" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D072FFAEFDFB9B94FD2F75B3" box="[605,753,934,959]" italics="true" pageId="38" pageNumber="39">Eopelobates</emphasis>
</taxonomicName>
is, thus, another unexpected late survivor from the site. Available published evidence suggest that
<taxonomicName id="4C5CECF0D072FFAEFF669C34FE8A7213" authorityName="Parker" authorityYear="1929" box="[192,340,1030,1055]" class="Amphibia" family="Pelobatidae" genus="Eopelobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="38" pageNumber="39" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D072FFAEFF669C34FE8A7213" box="[192,340,1030,1055]" italics="true" pageId="38" pageNumber="39">Eopelobates</emphasis>
</taxonomicName>
went extinct before the beginning of the Pleistocene (but see
<bibRefCitation id="EFCDEA82D072FFAEFE529C14FCCB7233" author="Rage, J. - C. &amp; Rocek, Z." box="[500,789,1062,1087]" pageId="38" pageNumber="39" pagination="133 - 167" refId="ref35799" refString="Rage, J. - C. and Rocek, Z. 2003. Evolution of anuran assemblages in the Tertiary and Quaternary of Europe, in the context of palaeoclimate and palaeogeography. Amphibia- Reptilia, 24: 133 - 167." type="journal article" year="2003">Rage and Roček, 2003</bibRefCitation>
, and
<bibRefCitation id="EFCDEA82D072FFAEFF589C74FDB97253" author="Vergilov, V. &amp; Tzankov, N." box="[254,615,1094,1119]" pageId="38" pageNumber="39" pagination="29 - 42" refId="ref39401" refString="Vergilov, V. and Tzankov, N. 2021. Giant pelobatid fossil larva from the middle Miocene of Bulgaria. Historia Naturalis Bulgarica, 43: 29 - 42. https: // doi. org / 10.48027 / hnb. 43.031" type="journal article" year="2021">Vergilov and Tzankov, 2021</bibRefCitation>
, for an older extinction date), which agrees with the fact that Quaternary sites in the area close to Hambach only yielded
<taxonomicName id="4C5CECF0D072FFAEFEB99C94FE4B72B3" authorityName="Wagler" authorityYear="1830" box="[287,405,1190,1215]" class="Amphibia" family="Pelobatidae" genus="Pelobates" kingdom="Animalia" order="Anura" pageId="38" pageNumber="39" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D072FFAEFEB99C94FE4B72B3" box="[287,405,1190,1215]" italics="true" pageId="38" pageNumber="39">Pelobates</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFCDEA82D072FFAEFE039C94FD8472B3" author="Bottcher, R." box="[421,602,1190,1215]" pageId="38" pageNumber="39" pagination="25 - 56" refId="ref30848" refString="Bottcher, R. 1991. Vertebraten. Stuttgarter Beitrage zur Naturkunde Serie B (Geologie und Palaontologie), 170: 25 - 56." type="journal article" year="1991">Böttcher, 1991</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFDCE9C94FCCA72B3" author="Holman, J. A." box="[616,788,1190,1215]" pageId="38" pageNumber="39" pagination="1 - 254" refId="ref33244" refString="Holman, J. A. 1998. Pleistocene amphibians and reptiles in Britain and Europe. Oxford monographs on geology and geophysics, 38: 1 - 254." type="journal article" year="1998">Holman, 1998</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFF669CF4FEB272D3" author="Sanchiz, B." box="[192,364,1222,1248]" pageId="38" pageNumber="39" refId="ref36500" refString="Sanchiz, B. 1998. Handbuch der Palaoherpetologie. 4 - Salientia. Friedrich Pfeil, Munchen." type="book" year="1998">Sanchiz, 1998</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFED19CF4FDEA72D3" author="Schouten, S." box="[375,564,1222,1248]" pageId="38" pageNumber="39" pagination="11 - 24" refId="ref36903" refString="Schouten, S. 2016. Een overzicht van de Pleistocene en Holocene herpetofauna (reptielen en amfibieen) van Nederland. Met aandacht voor vondsten langs de nederlandse kust. Cranium, 33: 11 - 24." type="journal article" year="2016">Schouten, 2016</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFD999CF4FCD172D3" author="Villa, A. &amp; Blain, H. - A. &amp; van den Hoek Ostende, L. W. &amp; Delfino, M." box="[575,783,1222,1247]" pageId="38" pageNumber="39" pagination="203 - 219" refId="ref39874" refString="Villa, A., Blain, H. - A., van den Hoek Ostende, L. W., and Delfino, M. 2018 c. Fossil amphibians and reptiles from Tegelen (Province of Limburg) and the early Pleistocene palaeoclimate of The Netherlands. Quaternary Science Reviews, 187: 203 - 219. https: // doi. org / 10.1016 / j. quascirev. 2018.03.020" type="journal article" year="2018">Villa et al., 2018c</bibRefCitation>
).
</paragraph>
<paragraph id="8BE39773D072FFAEFF5C9CD4FC037793" blockId="38.[192,793,166,1887]" lastBlockId="38.[840,1441,166,767]" pageId="38" pageNumber="39">
Taking into account their rather fragile skeleton, tree frogs of the genus
<taxonomicName id="4C5CECF0D072FFAEFDBD9D34FD8E7313" authorityName="Laurenti" authorityYear="1768" box="[539,592,1286,1311]" class="Amphibia" family="Hylidae" genus="Hyla" kingdom="Animalia" order="Anura" pageId="38" pageNumber="39" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D072FFAEFDBD9D34FD8E7313" box="[539,592,1286,1311]" italics="true" pageId="38" pageNumber="39">Hyla</emphasis>
</taxonomicName>
are rather common in European sites from the Miocene onwards, especially during the Quaternary. Earliest European representatives of the genus possibly entered the continent in the Early Miocene (
<bibRefCitation id="EFCDEA82D072FFAEFDD89DB4FEBA73B3" author="Sanchiz, B. &amp; Rocek, Z." pageId="38" pageNumber="39" pagination="317 - 328" refId="ref36584" refString="Sanchiz, B. and Rocek, Z. 1996. An overview of the anuran fossil record, p. 317 - 328. In Tinsley, R. C. and Kobel, H. R. (eds.), The Biology of Xenopus. Clarendon Press, Oxford, UK." type="book chapter" year="1996">Sanchiz and Roček, 1996</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFEDC9D94FD7173B3" author="Rage, J. - C. &amp; Rocek, Z." box="[378,687,1446,1471]" pageId="38" pageNumber="39" pagination="133 - 167" refId="ref35799" refString="Rage, J. - C. and Rocek, Z. 2003. Evolution of anuran assemblages in the Tertiary and Quaternary of Europe, in the context of palaeoclimate and palaeogeography. Amphibia- Reptilia, 24: 133 - 167." type="journal article" year="2003">Rage and Roček, 2003</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFD639D94FEDC73D3" author="Rocek, Z." pageId="38" pageNumber="39" pagination="397 - 439" refId="ref36270" refString="Rocek, Z. 2013. Mesozoic and Tertiary Anura of Laurasia. Palaeobiodiversity and Palaeoenvironments, 93: 397 - 439. https: // doi. org / 10.1007 / s 12549 - 013 - 0131 - y" type="journal article" year="2013">Roček, 2013</bibRefCitation>
), together with several other amphibians and reptiles (
<bibRefCitation id="EFCDEA82D072FFAEFE969DD4FD7373F3" author="Szyndlar, Z. &amp; Schleich, H. H." box="[304,685,1510,1536]" pageId="38" pageNumber="39" pagination="1 - 47" refId="ref38233" refString="Szyndlar, Z. and Schleich, H. H. 1993. Description of Miocene snakes from Petersbuch 2 with comments on the lower and middle Miocene ophidian faunas of Southern Germany. Stuttgarter Beitrage zur Naturkunde, Serie B (Geologie und Palaontologie), 192: 1 - 47." type="journal article" year="1993">Szyndlar and Schleich, 1993</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFD629DD4FF217013" author="Ivanov, M." pageId="38" pageNumber="39" pagination="559 - 588" refId="ref33377" refString="Ivanov, M. 2000. Snakes of the lower / middle Miocene transition at Vieux Collonges (Rhone, France), with comments on the colonisation of western Europe by colubroids. Geodiversitas, 22: 559 - 588." type="journal article" year="2000">Ivanov, 2000</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFEAD9E34FE2F7013" author="Delfino, M. &amp; Rage, J. - C. &amp; Rook, L." box="[267,497,1542,1567]" pageId="38" pageNumber="39" pagination="153 - 161" refId="ref31483" refString="Delfino, M., Rage, J. - C., and Rook, L. 2003. Tertiary mammal turnover phenomena: What happened to the herpetofauna? Deinsea, 10: 153 - 161." type="journal article" year="2003">Delfino et al., 2003</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFE5B9E34FCCA7013" author="Rage, J. - C. &amp; Rocek, Z." box="[509,788,1542,1567]" pageId="38" pageNumber="39" pagination="133 - 167" refId="ref35799" refString="Rage, J. - C. and Rocek, Z. 2003. Evolution of anuran assemblages in the Tertiary and Quaternary of Europe, in the context of palaeoclimate and palaeogeography. Amphibia- Reptilia, 24: 133 - 167." type="journal article" year="2003">Rage and Roček, 2003</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFF669E14FE8F7033" author="Rage, J. - C." box="[192,337,1574,1599]" pageId="38" pageNumber="39" pagination="517 - 534" refId="ref35676" refString="Rage, J. - C. 2013. Mesozoic and Cenozoic squamates of Europe. Palaeobiodiversity and Palaeoenvironments, 93: 517 - 534. https: // doi. org / 10.1007 / s 12549 - 013 - 0124 - x" type="journal article" year="2013">Rage, 2013</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFEC69E14FDB57033" author="Georgalis, G. L. &amp; Villa, A. &amp; Delfino, M." box="[352,619,1574,1600]" pageId="38" pageNumber="39" pagination="12" refId="ref32323" refString="Georgalis, G. L., Villa, A., and Delfino, M. 2016. First description of a fossil chamaeleonid from Greece and its relevance for the European biogeographic history of the group. The Science of Nature, 103: 12. https: // doi. org / 10.1007 / s 00114 - 016 - 1336 - 5" type="journal article" year="2016">Georgalis et al., 2016</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFDDC9E14FF217053" author="Ivanov, M. &amp; Ruta, M. &amp; Klembara, J. &amp; Bohme, M." pageId="38" pageNumber="39" pagination="767 - 797" refId="ref33448" refString="Ivanov, M., Ruta, M., Klembara, J., and Bohme, M. 2018. A new species of Varanus (Anguimorpha: Varanidae) from the early Miocene of the Czech Republic, and its relationships and palaeoecology. Journal of Systematic Palaeontology, 16: 767 - 797. https: // doi. org / 10.1080 / 14772019.2017.1355338" type="journal article" year="2018">Ivanov et al., 2018</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFEA89E74FE357053" author="Villa, A. &amp; Abella, J. &amp; Alba, D. M. &amp; Almecija, S. &amp; Bolet, A. &amp; Koufos, G. D. &amp; Knoll, F. &amp; Lujan, A. H. &amp; Morales, J. &amp; Robles, J. M. &amp; Sanchez, I. M. &amp; Delfino, M." box="[270,491,1606,1631]" pageId="38" pageNumber="39" pagination="0207719" refId="ref39678" refString="Villa, A., Abella, J., Alba, D. M., Almecija, S., Bolet, A., Koufos, G. D., Knoll, F., Lujan, A. H., Morales, J., Robles, J. M., Sanchez, I. M., and Delfino, M. 2018 a. Revision of Varanus marathonensis (Squamata, Varanidae) based on historical and new material: morphology, systematics, and paleobiogeography of the European monitor lizards. PLoS ONE, 13: e 0207719. https: // doi. org / 10.1371 / journal. pone. 0207719" type="journal article" year="2018">Villa et al., 2018a</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFE5C9E74FCCA7053" author="Villa, A. &amp; Delfino, M." box="[506,788,1606,1631]" pageId="38" pageNumber="39" pagination="177 - 211" refId="ref39467" refString="Villa, A. and Delfino, M. 2019. Fossil lizards and worm lizards (Reptilia, Squamata) from the Neogene and Quaternary of Europe: an overview. Swiss Journal of Palaeontology, 138: 177 - 211. https: // doi. org / 10.1007 / s 13358 - 018 - 0172 - y" type="journal article" year="2019">Villa and Delfino, 2019</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFF669E54FE067073" author="Macaluso, L. &amp; Mannion, P. D. &amp; Evans, S. E. &amp; Carnevale, G. &amp; Monti, S. &amp; Marchitelli, D. &amp; Delfino, M." box="[192,472,1638,1663]" pageId="38" pageNumber="39" pagination="220935" refId="ref34329" refString="Macaluso, L., Mannion, P. D., Evans, S. E., Carnevale, G., Monti, S., Marchitelli, D., and Delfino, M. 2022 a. Biogeographic history of Palearctic caudates revealed by a critical appraisal of their fossil record quality and spatio-temporal distribution. Royal Society Open Science, 9: 220935. https: // doi. org / 10.1098 / rsos. 220935" type="journal article" year="2022">Macaluso et al., 2022a</bibRefCitation>
). In this context, tree frogs from Hambach represent one of the oldest occurrences in
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, together with Oggenhausen
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<collectingRegion id="49985991D072FFAEFF799EF4FE1170D3" box="[223,463,1734,1759]" country="Germany" name="Baden-Wuerttemberg" pageId="38" pageNumber="39">Baden-Württemberg</collectingRegion>
(
<bibRefCitation id="EFCDEA82D072FFAEFE799EF4FD0170D3" author="Bottcher, R. &amp; Heizmann, E. P. J. &amp; Rasser, M. W. &amp; Ziegler, R." box="[479,735,1734,1759]" pageId="38" pageNumber="39" pagination="237 - 260" refId="ref30874" refString="Bottcher, R., Heizmann, E. P. J., Rasser, M. W., and Ziegler, R. 2009. Biostratigraphy and palaeoecology of a Middle Miocene (Karpathian, MN 5) fauna from the northern margin of the North Alpine Foreland Basin (Oggenhausen 2, SW' Germany). Neues Jahrbuch fur Geologie und Palaontologie, 254: 237 - 260. https: // doi. org / 10.1127 / 0077 - 7749 / 2009 / 0011" type="journal article" year="2009">Böttcher et al., 2009</bibRefCitation>
), but also the westernmost location reached in Europe by these frogs during the Neogene based on the current published knowledge (even though, again, bones of tree frogs are very fragile and maybe subject to a strong preservation bias). Hambach is also the first German site yielding Pliocene tree frog remains. Subsequently, tree frogs persisted in the area, as testified at least by the members of the
<taxonomicName id="4C5CECF0D072FFAEFA239934FC7A7733" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" class="Amphibia" family="Hylidae" genus="Hyla" kingdom="Animalia" order="Anura" pageId="38" pageNumber="39" phylum="Chordata" rank="species" species="arborea">
<emphasis id="B9284B61D072FFAEFA239934FC7A7733" italics="true" pageId="38" pageNumber="39">H. arborea</emphasis>
</taxonomicName>
group found in Tegelen (
<bibRefCitation id="EFCDEA82D072FFAEFB629914FA497733" author="Villa, A. &amp; Blain, H. - A. &amp; van den Hoek Ostende, L. W. &amp; Delfino, M." box="[1220,1431,294,319]" pageId="38" pageNumber="39" pagination="203 - 219" refId="ref39874" refString="Villa, A., Blain, H. - A., van den Hoek Ostende, L. W., and Delfino, M. 2018 c. Fossil amphibians and reptiles from Tegelen (Province of Limburg) and the early Pleistocene palaeoclimate of The Netherlands. Quaternary Science Reviews, 187: 203 - 219. https: // doi. org / 10.1016 / j. quascirev. 2018.03.020" type="journal article" year="2018">Villa et al., 2018c</bibRefCitation>
). They are widespread throughout northwestern Europe nowadays (
<bibRefCitation id="EFCDEA82D072FFAEFB8A9954FAD77773" author="Sillero, N. &amp; Campos, J. &amp; Bonardi, A. &amp; Corti, C. &amp; Creemers, R. &amp; Crochet, P. - A. &amp; Crnobrnja Isailovic, J. &amp; Denoel, M. &amp; Ficetola, G. F. &amp; Goncalves, J. &amp; Kuzmin, S. &amp; Lymberakis, P. &amp; de Pous, P. &amp; Rodriguez, A. &amp; Sindaco, R. &amp; Speybroeck, J. &amp; Toxopeus, B. &amp; Vieites, D. R. &amp; Vences, M." box="[1068,1289,358,384]" pageId="38" pageNumber="39" pagination="1 - 31" refId="ref37047" refString="Sillero, N., Campos, J., Bonardi, A., Corti, C., Creemers, R., Crochet, P. - A., Crnobrnja Isailovic, J., Denoel, M., Ficetola, G. F., Goncalves, J., Kuzmin, S., Lymberakis, P., de Pous, P., Rodriguez, A., Sindaco, R., Speybroeck, J., Toxopeus, B., Vieites, D. R., and Vences, M. 2014. Updated distribution and biogeography of amphibians and reptiles of Europe. Amphibia-Reptilia, 35: 1 - 31. https: // doi. org / 10.1163 / 15685381 - 00002935" type="journal article" year="2014">Sillero et al., 2014</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFAB39954FC0D7793" author="Speybroeck, J. &amp; Beukema, W. &amp; Bok, B. &amp; Van der Voort, J." pageId="38" pageNumber="39" refId="ref37342" refString="Speybroeck, J., Beukema, W., Bok, B., and Van der Voort, J. 2016. Field guide to the amphibians and reptiles of Britain and Europe. Bloomsbury Publishing, London." type="book" year="2016">Speybroeck et al., 2016</bibRefCitation>
).
</paragraph>
<paragraph id="8BE39773D072FFAEFC249994FB3474F3" blockId="38.[840,1441,166,767]" pageId="38" pageNumber="39">
Bufonids and ranids are also inhabitants of modern northwestern Europe, as well as common findings in Neogene and Quaternary fossil assemblages in the continent. Hambach stands out as one of the northernmost known occurrences of the common toad in Pliocene times, together with Kaltensundheim in
<collectingCountry id="F34BD7E3D072FFAEFB8C9A54FB49748C" box="[1066,1175,614,640]" name="Germany" pageId="38" pageNumber="39">Germany</collectingCountry>
(
<bibRefCitation id="EFCDEA82D072FFAEFB009A54FA977473" author="Bohme, G." box="[1190,1353,614,639]" pageId="38" pageNumber="39" pagination="231 - 238" refId="ref30406" refString="Bohme, G. 2002. Amphibienreste aus dem Oberpliozan von Kaltensundheim (Rhon, Thuringen). Mitteilungen des Museum fur Naturkunde, Berlin, Geowissenschaftliche Reihe, 5: 231 - 238." type="journal article" year="2002">Böhme, 2002</bibRefCitation>
) and at least Węże
<quantity id="4CA43A96D072FFAEFC789AB4FBD17493" box="[990,1039,646,671]" metricMagnitude="-2" metricUnit="m" metricValue="5.08" pageId="38" pageNumber="39" unit="in" value="2.0">2 in</quantity>
<collectingCountry id="F34BD7E3D072FFAEFBBD9AB4FBB07493" box="[1051,1134,646,671]" name="Poland" pageId="38" pageNumber="39">Poland</collectingCountry>
(
<bibRefCitation id="EFCDEA82D072FFAEFB279AB4FA487493" author="Mlynarski, M. &amp; Szyndlar, Z. &amp; Estes, R. &amp; Sanchiz B." box="[1153,1430,646,671]" pageId="38" pageNumber="39" pagination="209 - 229" refId="ref34852" refString="Mlynarski, M., Szyndlar, Z., Estes, R., and Sanchiz B. 1984. Amphibians and reptiles from the Pliocene locality of Weze II near Dzialoszyn (Poland). Acta Palaeontologica Polonica, 29: 209 - 229." type="journal article" year="1984">Młynarski et al., 1984</bibRefCitation>
). Both the common toad and the two ranids were found also in the Lower Pleistocene of the nearby Tegelen locality (
<bibRefCitation id="EFCDEA82D072FFAEFBAA9AD4FB3E74F3" author="Villa, A. &amp; Blain, H. - A. &amp; van den Hoek Ostende, L. W. &amp; Delfino, M." box="[1036,1248,742,767]" pageId="38" pageNumber="39" pagination="203 - 219" refId="ref39874" refString="Villa, A., Blain, H. - A., van den Hoek Ostende, L. W., and Delfino, M. 2018 c. Fossil amphibians and reptiles from Tegelen (Province of Limburg) and the early Pleistocene palaeoclimate of The Netherlands. Quaternary Science Reviews, 187: 203 - 219. https: // doi. org / 10.1016 / j. quascirev. 2018.03.020" type="journal article" year="2018">Villa et al., 2018c</bibRefCitation>
).
</paragraph>
<paragraph id="8BE39773D072FFAEFCEE9B24FB607563" blockId="38.[840,1416,790,879]" pageId="38" pageNumber="39">
<emphasis id="B9284B61D072FFAEFCEE9B24FB607563" bold="true" pageId="38" pageNumber="39">Palaeoenvironmental Insights and the Role of Northwestern Europe as a Late Neogene Refugial Area for Amphibians</emphasis>
</paragraph>
<paragraph id="8BE39773D072FFAFFC249BB4FCD976EE" blockId="38.[840,1441,902,1888]" lastBlockId="39.[192,793,169,1890]" lastPageId="39" lastPageNumber="40" pageId="38" pageNumber="39">
The Hambach palaeofaunas, and especially their amphibian components, appear rather significant for their composition, with representatives of enigmatic and poorly known taxa and unexpected occurrences both in chronological and geographical terms. The palaeoenvironments in which these palaeofaunas lived were certainly humid ones. In addition to sedimentological and palaeobotanical evidence (see Geological setting), these humid conditions are testified by the rather high diversity of amphibians and the presence of other water-related animals (e.g., fish, pond turtles,
<taxonomicName id="4C5CECF0D072FFAEFABB9CD4FC147313" authority="Laurenti, 1768" authorityName="Laurenti" authorityYear="1768" class="Squamata" family="Colubridae" genus="Natrix" kingdom="Animalia" pageId="38" pageNumber="39" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D072FFAEFABB9CD4FABC72F3" box="[1309,1378,1254,1279]" italics="true" pageId="38" pageNumber="39">Natrix</emphasis>
<bibRefCitation id="EFCDEA82D072FFAEFACD9CD4FC147313" author="Laurenti, J. N." pageId="38" pageNumber="39" refId="ref34062" refString="Laurenti, J. N. 1768. Austriaci Viennensis specimen medicum, exhibens synopsin reptilium emendatam cum experimentis circa Venena et antidota reptilium Austriacorum quod authoritate et consensu. Joannis Thomae de Trattnern, Caes. Reg. Maj. Aulae Typographi et Bibliopolae, Wien. https: // doi. org / 10.5962 / bhl. title. 5108" type="book" year="1768">Laurenti, 1768</bibRefCitation>
</taxonomicName>
snakes, dominant aquatic/semiaquatic forms among the insectivores, beavers;
<bibRefCitation id="EFCDEA82D072FFAEFAFB9D14FC597353" author="Mors, T." pageId="38" pageNumber="39" pagination="177 - 183" refId="ref34899" refString="Mors, T. 2002. Biostratigraphy and paleoecology of continental Tertiary vertebrate faunas in the Lower Rhine Embayment (NW-Germany). Netherlands Journal of Geosciences - Geologie en Mijnbouw, 81: 177 - 183. https: // doi. org / 10.1017 / S 0016774600022411" type="journal article" year="2002">Mörs, 2002</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFC339D73FB707353" author="Cernansky, A. &amp; Szyndlar, Z. &amp; Mors, T." box="[917,1198,1345,1375]" pageId="38" pageNumber="39" pagination="329 - 354" refId="ref31064" refString="Cernansky, A., Szyndlar, Z., and Mors, T. 2017. Fossil squamate faunas from the Neogene of Hambach (northwestern Germany). Palaeobiodiversity and Palaeoenvironments, 97: 329 - 354. https: // doi. org / 10.1007 / s 12549 - 016 - 0252 - 1" type="journal article" year="2017">Čerňanský et al., 2017</bibRefCitation>
). As far as amphibians are concerned, permanent water bodies are indicated by the occurrences of
<taxonomicName id="4C5CECF0D072FFAEFB529DB4FA7E7393" baseAuthorityName="Vasilyan and Yanenko" baseAuthorityYear="2020" box="[1268,1440,1414,1439]" class="Amphibia" family="Batrachosauroididae" genus="Palaeoproteus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="38" pageNumber="39" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D072FFAEFB529DB4FA7E7393" box="[1268,1440,1414,1439]" italics="true" pageId="38" pageNumber="39">Palaeoproteus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFCDEA82D072FFAEFCE99D94FC2C73B3" author="Herre, W." box="[847,1010,1446,1471]" pageId="38" pageNumber="39" pagination="1 - 85" refId="ref32975" refString="Herre, W. 1935. Die Schwanzlurche der mitteleocanen (oberlutetischen) Braunkohle des Geiseltales und die Phylogenie der Urodelen unter Einschluss der fossilen Formen. Zoologica - Original-Abhandlungen aus dem Gesamtgebiete der Zoologie, 87: 1 - 85." type="journal article" year="1935">Herre, 1935</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFBA89D94FA4973B3" author="Vasilyan, D. &amp; Yanenko, V." box="[1038,1431,1446,1471]" pageId="38" pageNumber="39" pagination="2733" refId="ref38931" refString="Vasilyan, D. and Yanenko, V. 2020. The last Palaeoproteus (Urodela: Batrachosauroididae) of Europe. Scientific Reports, 10: 2733. https: // doi. org / 10.1038 / s 41598 - 020 - 59255 - 1" type="journal article" year="2020">Vasilyan and Yanenko, 2020</bibRefCitation>
), palaeobatrachids, and, at least in the Pliocene level,
<taxonomicName id="4C5CECF0D072FFAEFC349DD4FBC873F3" authorityName="Estes and Darevsky" authorityYear="1977" box="[914,1046,1510,1535]" class="Amphibia" family="Proteidae" genus="Mioproteus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="38" pageNumber="39" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D072FFAEFC349DD4FBC873F3" box="[914,1046,1510,1535]" italics="true" pageId="38" pageNumber="39">Mioproteus</emphasis>
</taxonomicName>
(a fully aquatic taxon according to
<bibRefCitation id="EFCDEA82D072FFAEFCC19E34FB737013" author="Venczel, M. &amp; Codrea, V. A." box="[871,1197,1542,1568]" pageId="38" pageNumber="39" pagination="1508027" refId="ref39349" refString="Venczel, M. and Codrea, V. A. 2018. A new proteid salamander from the early Oligocene of Romania with notes on the paleobiogeography of Eurasian proteids. Journal of Vertebrate Paleontology, 38: e 1508027. https: // doi. org / 10.1080 / 02724634.2018.1508027" type="journal article" year="2018">Venczel and Codrea, 2018</bibRefCitation>
). The extant
<taxonomicName id="4C5CECF0D072FFAEFAEF9E34FC1A7033" baseAuthorityName="Mendelssohn and Steinitz" baseAuthorityYear="1943" class="Amphibia" family="Alytidae" genus="Latonia" kingdom="Animalia" order="Anura" pageId="38" pageNumber="39" phylum="Chordata" rank="species" species="nigriventer">
<emphasis id="B9284B61D072FFAEFAEF9E34FC1A7033" italics="true" pageId="38" pageNumber="39">Latonia nigriventer</emphasis>
</taxonomicName>
inhabits marshy areas in the Hula Valley, in
<collectingCountry id="F34BD7E3D072FFAEFC3C9E74FC057053" box="[922,987,1606,1631]" name="Israel" pageId="38" pageNumber="39">Israel</collectingCountry>
(
<bibRefCitation id="EFCDEA82D072FFAEFC4A9E74FB1B7053" author="Biton, R. &amp; Geffen, E. &amp; Vences, M. &amp; Cohen, O. &amp; Bailon, S. &amp; Rabinovich, R. &amp; Malka, Y. &amp; Oron, T. &amp; Boistel, R. &amp; Brumfeld, V. &amp; Gafny, S." box="[1004,1221,1606,1631]" pageId="38" pageNumber="39" pagination="1959" refId="ref30010" refString="Biton, R., Geffen, E., Vences, M., Cohen, O., Bailon, S., Rabinovich, R., Malka, Y., Oron, T., Boistel, R., Brumfeld, V., and Gafny, S. 2013. The rediscovered Hula painted frog is a living fossil. Nature Communications, 4: 1959. https: // doi. org / 10.1038 / ncomms 2959" type="journal article" year="2013">Biton et al., 2013</bibRefCitation>
,
<bibRefCitation id="EFCDEA82D072FFAEFB729E74FACD7053" author="Biton, R. &amp; Boistel, R. &amp; Rabinovich, R. &amp; Gafny, S. &amp; Brumfeld, V. &amp; Bailon, S." box="[1236,1299,1606,1631]" pageId="38" pageNumber="39" pagination="1131 - 1145" refId="ref30095" refString="Biton, R., Boistel, R., Rabinovich, R., Gafny, S., Brumfeld, V., and Bailon, S. 2016. Osteological observations on the alytid Anura Latonia nigriventer with comments on functional morphology, biogeography, and evolutionary history. Journal of Morphology, 277: 1131 - 1145. https: // doi. org / 10.1002 / jmor. 20562" type="journal article" year="2016">2016</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D072FFAEFA879E74FC547073" author="Perl, R. G. B. &amp; Gafny, S. &amp; Malka, Y. &amp; Renan, S. &amp; Woodhams, D. C. &amp; Rollins-Smith, L. &amp; Pask, J. D. &amp; Bletz, M. C. &amp; Geffen, E. &amp; Vences, M." pageId="38" pageNumber="39" pagination="11 - 37" refId="ref35309" refString="Perl, R. G. B., Gafny, S., Malka, Y., Renan, S., Woodhams, D. C., Rollins-Smith, L., Pask, J. D., Bletz, M. C., Geffen, E., and Vences, M. 2017. Natural history and conservation of the rediscovered Hula painted frog, Latonia nigriventer. Contributions to Zoology, 86: 11 - 37. https: // doi. org / 10.1163 / 18759866 - 08601002" type="journal article" year="2017">Perl et al., 2017</bibRefCitation>
), and the abundance of
<taxonomicName id="4C5CECF0D072FFAEFB1E9E54FAD17073" authorityName="von Meyer" authorityYear="1843" box="[1208,1295,1638,1663]" class="Amphibia" family="Alytidae" genus="Latonia" kingdom="Animalia" order="Anura" pageId="38" pageNumber="39" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D072FFAEFB1E9E54FAD17073" box="[1208,1295,1638,1663]" italics="true" pageId="38" pageNumber="39">Latonia</emphasis>
</taxonomicName>
in both the Miocene and Pliocene of Hambach hints towards persistent preference for similar swampy environments in past members of the clade as well. Pelobatids are fossorial anurans that suggest the availability of sandy soils in Hambach. Forested areas nearby are supported by the rare tree frogs, as well as
<taxonomicName id="4C5CECF0D072FFAEFC779F74FAC57153" authority="Gervais, 1849" authorityName="Gervais" authorityYear="1849" box="[977,1307,1862,1888]" class="Mammalia" family="Pliopithecidae" genus="Pliopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="38" pageNumber="39" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D072FFAEFC779F74FBBF7153" box="[977,1121,1862,1887]" italics="true" pageId="38" pageNumber="39">Pliopithecus</emphasis>
<bibRefCitation id="EFCDEA82D072FFAEFBCA9F74FAC57153" author="Gervais, P." box="[1132,1307,1862,1888]" pageId="38" pageNumber="39" pagination="699 - 700" refId="ref32606" refString="Gervais, P. 1849. Note sur une nouvelle espece de singe fossile. Comptes Rendus hebdomadaires des seances de l'Academie des sciences, 28: 699 - 700." type="journal article" year="1849">Gervais, 1849</bibRefCitation>
</taxonomicName>
, and flying squirrel remains (
<bibRefCitation id="EFCDEA82D073FFAFFE2A989BFDC976CE" author="Mors, T." box="[396,535,169,194]" pageId="39" pageNumber="40" pagination="177 - 183" refId="ref34899" refString="Mors, T. 2002. Biostratigraphy and paleoecology of continental Tertiary vertebrate faunas in the Lower Rhine Embayment (NW-Germany). Netherlands Journal of Geosciences - Geologie en Mijnbouw, 81: 177 - 183. https: // doi. org / 10.1017 / S 0016774600022411" type="journal article" year="2002">Mörs, 2002</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D073FFAFFD84989BFF2076EE" author="Van Laere, G. &amp; Mors, T." pageId="39" pageNumber="40" pagination="223 - 241" refId="ref38874" refString="Van Laere, G. and Mors, T. 2023. Beavers and flying squirrels (Rodentia: Castoridae, Pteromyini) from the Late Pliocene of Hambach 11 C, Germany. Geodiversitas, 45: 223 - 241. https: // doi. org / 10.5252 / geodiversitas 2023 v 45 a 7" type="journal article" year="2023">Van Laere and Mörs, 2023</bibRefCitation>
) for the Miocene and Pliocene, respectively.
</paragraph>
<paragraph id="8BE39773D073FFAFFF5C98DBFF2D72EE" blockId="39.[192,793,169,1890]" pageId="39" pageNumber="40">
Humidity in Hambach was particularly high during the Middle Miocene, when cryptobranchids were living in the area. As a matter of fact, these urodeles are tied to high humidity and pluviometry according to
<bibRefCitation id="EFCDEA82D073FFAFFEF1995BFD9A778E" author="Bohme, M. &amp; Vasilyan, D. &amp; Winklhofer, M." box="[343,580,361,386]" pageId="39" pageNumber="40" pagination="64 - 72" refId="ref30639" refString="Bohme, M., Vasilyan, D., and Winklhofer, M. 2012. Habitat tracking, range dynamics and palaeoclimatic significance of Eurasian giant salamanders (Cryptobranchidae) - indications for elevated Central Asian humidity during Cenozoic global warm periods. Palaeogeography, Palaeoclimatology, Palaeoecology, 342 - 343: 64 - 72. https: // doi. org / 10.1016 / j. palaeo. 2012.04.032" type="journal article" year="2012">Böhme et al. (2012)</bibRefCitation>
. A certain influx of marine conditions in an estuarine setting is highlighted by the abundant marine fish and rare cetacean remains in Hambach 6C (
<bibRefCitation id="EFCDEA82D073FFAFFD9999FBFD0F77EE" author="Mors, T." box="[575,721,457,482]" pageId="39" pageNumber="40" pagination="177 - 183" refId="ref34899" refString="Mors, T. 2002. Biostratigraphy and paleoecology of continental Tertiary vertebrate faunas in the Lower Rhine Embayment (NW-Germany). Netherlands Journal of Geosciences - Geologie en Mijnbouw, 81: 177 - 183. https: // doi. org / 10.1017 / S 0016774600022411" type="journal article" year="2002">Mörs, 2002</bibRefCitation>
). This changed at the end of the Middle Miocene (
<bibRefCitation id="EFCDEA82D073FFAFFD1F99DAFE92742E" author="Schafer, A. &amp; Utescher, T. &amp; Mors, T." pageId="39" pageNumber="40" pagination="73 - 110" refId="ref36717" refString="Schafer, A., Utescher, T., and Mors, T. 2004. Stratigraphy of the Cenozoic Lower Rhine Basin, northwestern Germany. Newsletters on Stratigraphy, 40: 73 - 110. https: // doi. org / 10.1127 / 0078 - 0421 / 2004 / 0040 - 0073" type="journal article" year="2004">Schäfer et al., 2004</bibRefCitation>
), long before the deposition of remains found in Hambach 11 and 11C, which only include freshwater fish and is devoid of other marine animals. Fish remains in Hambach also mark another change from slow currents and larger channels in Hambach 6C to more oxygenated waters with higher energy in Hambach 11 and 11C (
<bibRefCitation id="EFCDEA82D073FFAFFD759AFBFEDC750E" author="Mors, T." pageId="39" pageNumber="40" pagination="177 - 183" refId="ref34899" refString="Mors, T. 2002. Biostratigraphy and paleoecology of continental Tertiary vertebrate faunas in the Lower Rhine Embayment (NW-Germany). Netherlands Journal of Geosciences - Geologie en Mijnbouw, 81: 177 - 183. https: // doi. org / 10.1017 / S 0016774600022411" type="journal article" year="2002">Mörs, 2002</bibRefCitation>
). Moreover, another palaeoenvironmental differences highlighted by the faunal composition in the two Hambach levels lies in the temperature: the Miocene fauna is rich in thermophilic vertebrates, such as giant tortoises, crocodylians, chameleons, “Oriental vipers”, and possibly cobras (
<bibRefCitation id="EFCDEA82D073FFAFFD2F9BBBFCCA75AE" author="Mors, T." box="[649,788,905,930]" pageId="39" pageNumber="40" pagination="177 - 183" refId="ref34899" refString="Mors, T. 2002. Biostratigraphy and paleoecology of continental Tertiary vertebrate faunas in the Lower Rhine Embayment (NW-Germany). Netherlands Journal of Geosciences - Geologie en Mijnbouw, 81: 177 - 183. https: // doi. org / 10.1017 / S 0016774600022411" type="journal article" year="2002">Mörs, 2002</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D073FFAFFF669B96FE0D75CE" author="Cernansky, A. &amp; Szyndlar, Z. &amp; Mors, T." box="[192,467,932,962]" pageId="39" pageNumber="40" pagination="329 - 354" refId="ref31064" refString="Cernansky, A., Szyndlar, Z., and Mors, T. 2017. Fossil squamate faunas from the Neogene of Hambach (northwestern Germany). Palaeobiodiversity and Palaeoenvironments, 97: 329 - 354. https: // doi. org / 10.1007 / s 12549 - 016 - 0252 - 1" type="journal article" year="2017">Čerňanský et al., 2017</bibRefCitation>
), whereas these are absent in the Pliocene one. This agrees with the climate reconstruction performed for Hambach 6C based on palaeobotanical data (
<bibRefCitation id="EFCDEA82D073FFAFFDA69C3BFCC8722E" author="Utescher, T. &amp; Mosbrugger, V. &amp; Ashraf, A. R." box="[512,790,1033,1058]" pageId="39" pageNumber="40" pagination="430 - 449" refId="ref38493" refString="Utescher, T., Mosbrugger, V., and Ashraf, A. R. 2000. Terrestrial climate evolution in northwest Germany over the last 25 million years. Palaios, 15: 430 - 449. https: // doi. org / 10.2307 / 3515514" type="journal article" year="2000">Utescher et al., 2000</bibRefCitation>
,
<bibRefCitation id="EFCDEA82D073FFAFFF669C1BFEDC724E" author="Utescher, T. &amp; Mosbrugger, V. &amp; Ashraf, A. R." box="[192,258,1065,1090]" pageId="39" pageNumber="40" pagination="185 - 191" refId="ref38543" refString="Utescher, T., Mosbrugger, V., and Ashraf, A. R. 2002. Facies and paleogeography of the Tertiary of the Lower Rhine Basin - sedimentary versus climatic control. Netherlands Journal of Geosciences - Geologie en Mijnbouw, 81: 185 - 191. https: // doi. org / 10.1017 / S 0016774600022423" type="journal article" year="2002">2002</bibRefCitation>
), and makes further sense when considering that the Middle Miocene fauna deposited in a greenhouse interval (the Miocene Climatic Optimum;
<bibRefCitation id="EFCDEA82D073FFAFFEB49CBAFDB872AE" author="Steinthorsdottir, M. &amp; Coxall, H. &amp; de Boer, A. &amp; Huber, M. &amp; Barbolini, N. &amp; Bradshaw, C. D. &amp; Burls, N. J. &amp; Feakins, S. J. &amp; Gasson, E. &amp; Henderiks, J. &amp; Holbourn, A. E. &amp; Kiel, S. &amp; Kohn, M. J. &amp; Knorr, G. &amp; Kurschner, W. M. &amp; Lear, C. H. &amp; Liebrand, D. &amp; Lunt, D. J. &amp; Mors, T. &amp; Pearson, P. N. &amp; Pound, M. J. &amp; Stoll, H. &amp; Stromberg, C. A. E." box="[274,614,1160,1186]" pageId="39" pageNumber="40" pagination="2020" refId="ref37475" refString="Steinthorsdottir, M., Coxall, H., de Boer, A., Huber, M., Barbolini, N., Bradshaw, C. D., Burls, N. J., Feakins, S. J., Gasson, E., Henderiks, J., Holbourn, A. E., Kiel, S., Kohn, M. J., Knorr, G., Kurschner, W. M., Lear, C. H., Liebrand, D., Lunt, D. J., Mors, T., Pearson, P. N., Pound, M. J., Stoll, H., and Stromberg, C. A. E. 2021. The Miocene: the Future of the Past. Paleoceanography and Paleoclimatology, 36: e 2020 PA 004037. https: // doi. org / 10.1029 / 2020 PA 004037" type="journal article" year="2021">Steinthorsdottir et al. 2021</bibRefCitation>
), whereas the Late Pliocene one in a period of climate deterioration.
</paragraph>
<paragraph id="8BE39773D073FFAFFF5C9CDBFB85758E" blockId="39.[192,793,169,1890]" lastBlockId="39.[840,1441,168,1889]" pageId="39" pageNumber="40">
The faunal change between Hambach 6C on one side and Hambach 11 and 11C on the other is, thus, evident in several of its components, but it seems to affect the amphibians to a lower extent. Some amphibian taxa indeed disappeared between the Middle Miocene and the Late Pliocene (i.e., cryptobranchids, the
<taxonomicName id="4C5CECF0D073FFAFFDAA9D9BFD7473CE" authorityName="Macaluso, Villa &amp; Mörs" authorityYear="2021" box="[524,682,1449,1474]" class="Amphibia" family="Proteidae" genus="Euronecturus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D073FFAFFDAA9D9BFD7473CE" box="[524,682,1449,1474]" italics="true" pageId="39" pageNumber="40">Euronecturus</emphasis>
</taxonomicName>
lineage,
<taxonomicName id="4C5CECF0D073FFAFFF669DFAFE9F73ED" authorityName="Pomel" authorityYear="1853" box="[192,321,1480,1505]" class="Amphibia" family="Salamandridae" genus="Chelotriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D073FFAFFF669DFAFE9F73ED" box="[192,321,1480,1505]" italics="true" pageId="39" pageNumber="40">Chelotriton</emphasis>
</taxonomicName>
), but others persisted to even be among the last known representatives of their clades in northern/northwestern Europe (
<taxonomicName id="4C5CECF0D073FFAFFD1F9E3BFCCE702E" authorityName="von Meyer" authorityYear="1843" box="[697,784,1545,1570]" class="Amphibia" family="Alytidae" genus="Latonia" kingdom="Animalia" order="Anura" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D073FFAFFD1F9E3BFCCE702E" box="[697,784,1545,1570]" italics="true" pageId="39" pageNumber="40">Latonia</emphasis>
</taxonomicName>
, palaeobatrachids) or worldwide (
<taxonomicName id="4C5CECF0D073FFAFFDFA9E1BFCD6704E" baseAuthorityName="Vasilyan and Yanenko" baseAuthorityYear="2020" box="[604,776,1577,1602]" class="Amphibia" family="Batrachosauroididae" genus="Palaeoproteus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D073FFAFFDFA9E1BFCD6704E" box="[604,776,1577,1602]" italics="true" pageId="39" pageNumber="40">Palaeoproteus</emphasis>
</taxonomicName>
). The case of cryptobranchids is particularly interesting because their last reported occurrence in Europe is in Willershausen, an Upper Pliocene locality in
<collectingRegion id="49985991D073FFAFFEE59E9BFE2A70CE" box="[323,500,1704,1730]" country="Germany" name="Niedersachsen" pageId="39" pageNumber="40">Lower Saxony</collectingRegion>
, central
<collectingCountry id="F34BD7E3D073FFAFFDC29E9AFD0A70CE" box="[612,724,1704,1730]" name="Germany" pageId="39" pageNumber="40">Germany</collectingCountry>
. This makes it and Hambach 11 and 11C close or comparable in terms of both geographical position and age, leading to the question on why these urodeles are absent from the Upper Pliocene level in Hambach. The depositional environment in Willershausen was that of a large and deep lake in a sinkhole, surrounded by a hilly mesophytic woodland (
<bibRefCitation id="EFCDEA82D073FFAFFCE898DAFB9A770E" author="Kolibac, J. &amp; Adroit, B. &amp; Groning, E. &amp; Brauckmann, C. &amp; Wappler, T." box="[846,1092,232,258]" pageId="39" pageNumber="40" pagination="681 - 689" refId="ref33729" refString="Kolibac, J., Adroit, B., Groning, E., Brauckmann, C., and Wappler, T. 2016. First record of the family Trogossitidae (Insecta, Coleoptera) in the Late Pliocene deposits of Willershausen (Germany). PalZ, 90: 681 - 689. https: // doi. org / 10.1007 / s 12542 - 016 - 0316 - 6" type="journal article" year="2016">Kolibáč et al., 2016</bibRefCitation>
, and reference therein). The reconstructed palaeoclimate was cooler than what suggested for the Miocene Hambach, but with similar precipitations (
<bibRefCitation id="EFCDEA82D073FFAFFB85997BFB25776E" author="Thiel, C. &amp; Klotz, S. &amp; Uhl, D." box="[1059,1275,329,354]" pageId="39" pageNumber="40" pagination="263 - 287" refId="ref38369" refString="Thiel, C., Klotz, S., and Uhl, D. 2012. Palaeoclimate estimates for selected leaf floras from the Late Pliocene (Reuverian) of Central Europe based on different palaeobotanical techniques. Turkish Journal of Earth Sciences, 21: 263 - 287. https: // doi. org / 10.3906 / yer- 1007 - 41" type="journal article" year="2012">Thiel et al., 2012</bibRefCitation>
). Unlike other taxa present in Hambach 6C and absent in Hambach 11 and 11C, then, extirpation of cryptobranchids from there is likely not tied to a decrease in temperature, given their survival in the colder Willershausen. An alternative potential explanation may be the altitudinal distribution model proposed by
<bibRefCitation id="EFCDEA82D073FFAFFC6D9A1BFB0C744E" author="Bohme, M. &amp; Vasilyan, D. &amp; Winklhofer, M." box="[971,1234,553,578]" pageId="39" pageNumber="40" pagination="64 - 72" refId="ref30639" refString="Bohme, M., Vasilyan, D., and Winklhofer, M. 2012. Habitat tracking, range dynamics and palaeoclimatic significance of Eurasian giant salamanders (Cryptobranchidae) - indications for elevated Central Asian humidity during Cenozoic global warm periods. Palaeogeography, Palaeoclimatology, Palaeoecology, 342 - 343: 64 - 72. https: // doi. org / 10.1016 / j. palaeo. 2012.04.032" type="journal article" year="2012">Böhme et al. (2012)</bibRefCitation>
: cryptobranchids appear generally limited to higher elevations in drier periods with low basinal relief and colonize lowland environments in hyperhumid moments with high basinal relief. Giant salamanders may have disappeared from the Lower Rhine Embayment in one of such drier periods after the Middle Miocene, whereas the more elevated area near the Willershausen lake (i.e., the Harz Mountains) was able to sustain populations of these amphibians up to at least the late Neogene.
</paragraph>
<paragraph id="8BE39773D073FFA1FC249BBBFE31725E" blockId="39.[840,1441,168,1889]" lastBlockId="41.[192,793,1049,1906]" lastPageId="41" lastPageNumber="42" pageId="39" pageNumber="40">
Another locality that is worth comparing with Hambach is Tegelen, in
<collectingCountry id="F34BD7E3D073FFAFFBC39B9BFAF475CE" box="[1125,1322,937,962]" name="Netherlands" pageId="39" pageNumber="40">The Netherlands</collectingCountry>
(
<tableCitation id="C6DEA2C8D073FFAFFA919B9BFA4A75CE" box="[1335,1428,937,962]" captionStart="TABLE 2" captionStartId="40.[192,272,161,184]" captionTargetPageId="40" captionText="TABLE 2. Comparison between the palaeoherpetofaunas of Hambach and Tegelen. Pliocene sites in Hambach are considered together. Data for Hambach are from: Mörs et al., 2000; Mörs, 2002; Joyce, 2016; Čerňanský et al., 2017; Georgalis and Joyce, 2017; this work. Mauremys cf. sophiae (reported as Ocadia cf. sophiae by Mörs, 2002) is here considered as part of Mauremys following Hervet (2004). The Tegelen herpetofauna is mainly based on the Russel- Tiglia-Egypte pit assemblage (Villa et al., 2018c), but it also includes the Emys occurrences reported by Schreuder (1946) and Villa et al. (2018c). Abbreviation: Croc., Crocodylia." pageId="39" pageNumber="40">Table 2</tableCitation>
). Sediments at Tegelen were also deposited by the Rhine-Meuse river system, even though in a slightly younger, Early Pleistocene, time (
<bibRefCitation id="EFCDEA82D073FFAFFA9A9C3BFB9B724E" author="Van den Hoek Ostende, L. W." pageId="39" pageNumber="40" pagination="127 - 141" refId="ref38795" refString="Van den Hoek Ostende, L. W. 2004. The Tegelen clay-pits: a hundred year old classical locality. Scripta Geologica, Special Issue 4: 127 - 141." type="book chapter" year="2004">van den Hoek Ostende, 2004</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D073FFAFFBF49C1BFC23726E" author="Van den Hoek Ostende, L. W. &amp; de Vos, J." pageId="39" pageNumber="40" pagination="291 - 304" refId="ref38828" refString="Van den Hoek Ostende, L. W. and de Vos, J. 2006. A century of research on the classical locality of Tegelen (province of Limburg, The Netherlands). Courier Forschungsinstitut Senckenberg, 256: 291 - 304." type="journal article" year="2006">van den Hoek Ostende and de Vos, 2006</bibRefCitation>
). The Russel-Tiglia-Egypte pit at Tegelen is notable for being the
<typeStatus id="54E729D1D073FFAFFB749C5BFADA728E" box="[1234,1284,1129,1154]" pageId="39" pageNumber="40">type</typeStatus>
locality of
<emphasis id="B9284B61D073FFAFFA2C9C5BFC1F72AE" italics="true" pageId="39" pageNumber="40">
<taxonomicName id="4C5CECF0D073FFAFFA2C9C5BFC6372AE" class="Amphibia" family="Palaeobatrachidae" genus="Palaeobatrachus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="39" pageNumber="40" phylum="Chordata" rank="species" species="eurydices">P. eurydices</taxonomicName>
,
</emphasis>
the palaeobatrachid species found in the Upper Pliocene level at Hambach, and for having yielded a rather diverse assemblage of amphibians and reptiles (
<bibRefCitation id="EFCDEA82D073FFAFFBE79CDAFAF1730D" author="Villa, A. &amp; Rocek, Z. &amp; Tschopp, E. &amp; van den Hoek Ostende, L. W. &amp; Delfino, M." box="[1089,1327,1256,1281]" pageId="39" pageNumber="40" pagination="1211664" refId="ref39607" refString="Villa, A., Rocek, Z., Tschopp, E., van den Hoek Ostende, L. W., and Delfino, M. 2016. Palaeobatrachus eurydices, sp. nov. (Amphibia, Anura), the last western European palaeobatrachid. Journal of Vertebrate Paleontology, 36: e 1211664. https: // doi. org / 10.1080 / 02724634.2016.1211664" type="journal article" year="2016">Villa et al., 2016</bibRefCitation>
,
<bibRefCitation id="EFCDEA82D073FFAFFAE19CDAFA49730D" author="Villa, A. &amp; Blain, H. - A. &amp; van den Hoek Ostende, L. W. &amp; Delfino, M." box="[1351,1431,1256,1281]" pageId="39" pageNumber="40" pagination="203 - 219" refId="ref39874" refString="Villa, A., Blain, H. - A., van den Hoek Ostende, L. W., and Delfino, M. 2018 c. Fossil amphibians and reptiles from Tegelen (Province of Limburg) and the early Pleistocene palaeoclimate of The Netherlands. Quaternary Science Reviews, 187: 203 - 219. https: // doi. org / 10.1016 / j. quascirev. 2018.03.020" type="journal article" year="2018">2018c</bibRefCitation>
). Urodeles are much more diverse in Hambach, especially in the Miocene layers. Tegelen only yielded salamandrids (
<taxonomicName id="4C5CECF0D073FFAFFBCD9D7AFB1D736D" authorityName="Rafinesque" authorityYear="1815" box="[1131,1219,1352,1377]" class="Amphibia" family="Salamandridae" genus="Triturus" kingdom="Animalia" order="Caudata" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D073FFAFFBCD9D7AFB1D736D" box="[1131,1219,1352,1377]" italics="true" pageId="39" pageNumber="40">Triturus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C5CECF0D073FFAFFAB09D7AFA51736D" authorityName="Bell" authorityYear="1839" box="[1302,1423,1352,1377]" class="Amphibia" family="Salamandridae" genus="Lissotriton" kingdom="Animalia" order="Caudata" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D073FFAFFAB09D7AFA51736D" box="[1302,1423,1352,1377]" italics="true" pageId="39" pageNumber="40">Lissotriton</emphasis>
</taxonomicName>
), whereas Hambach has salamandrids, proteids, batrachosauroidids (both Miocene and Pliocene) and cryptobranchids (only Miocene). Among salamandrids, the presence of
<taxonomicName id="4C5CECF0D073FFAFFB379DFAFACC73ED" authorityName="Pomel" authorityYear="1853" box="[1169,1298,1480,1505]" class="Amphibia" family="Salamandridae" genus="Chelotriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D073FFAFFB379DFAFACC73ED" box="[1169,1298,1480,1505]" italics="true" pageId="39" pageNumber="40">Chelotriton</emphasis>
</taxonomicName>
in the Miocene level of Hambach expands the ecomorphological adaptations represented by a member of this clade in this assemblage compared to Tegelen. As already mentioned,
<taxonomicName id="4C5CECF0D073FFAFFB439E7AFAE3706D" authorityName="Rafinesque" authorityYear="1815" box="[1253,1341,1608,1633]" class="Amphibia" family="Salamandridae" genus="Triturus" kingdom="Animalia" order="Caudata" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D073FFAFFB439E7AFAE3706D" box="[1253,1341,1608,1633]" italics="true" pageId="39" pageNumber="40">Triturus</emphasis>
</taxonomicName>
is missing in the Pliocene of Hambach, but it is present in the Pleistocene of Tegelen. Given the scarcity of the remains, it is not clear whether the absence of this newt may just be artifactual or not, though. For anurans, significant is the persistence of palaeobatrachids in all levels at Hambach and then in the Russel-Tiglia-Egypte pit at Tegelen, and especially the same species being shared between the Upper Pliocene of the former and the Lower Pleistocene of the latter. In general, the overall diversity of frog and toads appears comparable, but there are strong differences in composition of the assemblages.
<taxonomicName id="4C5CECF0D07CFFA0FE829F3BFEA5712E" authorityName="von Meyer" authorityYear="1843" box="[292,379,1801,1826]" class="Amphibia" family="Alytidae" genus="Latonia" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07CFFA0FE829F3BFEA5712E" box="[292,379,1801,1826]" italics="true" pageId="40" pageNumber="41">Latonia</emphasis>
</taxonomicName>
is the most abundant anuran in Hambach, but it is absent in Tegelen. Pelobatids are possibly represented by
<taxonomicName id="4C5CECF0D07CFFA0FDAA9F7BFD7E716E" authorityName="Parker" authorityYear="1929" box="[524,672,1865,1890]" class="Amphibia" family="Pelobatidae" genus="Eopelobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07CFFA0FDAA9F7BFD7E716E" box="[524,672,1865,1890]" italics="true" pageId="40" pageNumber="41">Eopelobates</emphasis>
</taxonomicName>
at least in the Pliocene of Hambach, whereas
<taxonomicName id="4C5CECF0D07CFFA0FAAF9EBBFA5E70AE" authorityName="Wagler" authorityYear="1830" box="[1289,1408,1673,1698]" class="Amphibia" family="Pelobatidae" genus="Pelobates" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07CFFA0FAAF9EBBFA5E70AE" box="[1289,1408,1673,1698]" italics="true" pageId="40" pageNumber="41">Pelobates</emphasis>
</taxonomicName>
is identified at Tegelen. On the other hand, the Hambach assemblages lack
<taxonomicName id="4C5CECF0D07CFFA0FBC19EFBFAB770EE" authority="Oken, 1816" authorityName="Oken" authorityYear="1816" box="[1127,1385,1737,1763]" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07CFFA0FBC19EFBFB0E70EE" box="[1127,1232,1737,1762]" italics="true" pageId="40" pageNumber="41">Bombina</emphasis>
<bibRefCitation id="EFCDEA82D07CFFA0FB7F9EFBFAB770EE" author="Oken, L." box="[1241,1385,1737,1763]" pageId="40" pageNumber="41" refId="ref35201" refString="Oken, L. 1816. Lehrbuch der Naturgeschichte, vol. 3. Zoologie. Abtheilung 2. Atlas. C. H. Reclam, Leipzig." type="book" year="1816">Oken, 1816</bibRefCitation>
</taxonomicName>
, and
<taxonomicName id="4C5CECF0D07CFFA0FCEE9EDBFB52710E" authority="Bonaparte, 1838" authorityName="Bonaparte" authorityYear="1838" box="[840,1164,1769,1794]" class="Amphibia" family="Pelodytidae" genus="Pelodytes" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07CFFA0FCEE9EDBFC63710E" box="[840,957,1769,1794]" italics="true" pageId="40" pageNumber="41">Pelodytes</emphasis>
<bibRefCitation id="EFCDEA82D07CFFA0FC629EDBFB52710E" author="Bonaparte, C. L. J. L." box="[964,1164,1769,1794]" pageId="40" pageNumber="41" refId="ref30748" refString="Bonaparte, C. L. J. L. 1838. Iconographia della fauna italica per le quattro classi degli animali vertebrati. Tomo II. Amphibi. Fascicolo 23. Salviucci, Roma." type="book" year="1838">Bonaparte, 1838</bibRefCitation>
</taxonomicName>
, taxa otherwise present at Tegelen.
<taxonomicName id="4C5CECF0D07CFFA0FC759F3BFBD6712E" authorityName="Laurenti" authorityYear="1768" box="[979,1032,1801,1826]" class="Amphibia" family="Hylidae" genus="Hyla" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07CFFA0FC759F3BFBD6712E" box="[979,1032,1801,1826]" italics="true" pageId="40" pageNumber="41">Hyla</emphasis>
</taxonomicName>
, the common toad, and the ranids are shared by both the German and Dutch localities, even though less common in Hambach and restricted to specific levels as far as the toad and the ranids are concerned.
</paragraph>
<caption id="DF23C7FBD07CFFA0FF669893FC847746" pageId="40" pageNumber="41" startId="40.[192,272,161,184]" targetBox="[332,1317,353,1597]" targetIsTable="true" targetPageId="40" targetType="table">
<paragraph id="8BE39773D07CFFA0FF669893FC847746" blockId="40.[192,1446,161,331]" pageId="40" pageNumber="41">
<emphasis id="B9284B61D07CFFA0FF669893FEF276B4" bold="true" box="[192,300,161,184]" pageId="40" pageNumber="41">TABLE 2.</emphasis>
Comparison between the palaeoherpetofaunas of Hambach and Tegelen. Pliocene sites in Hambach are considered together. Data for Hambach are from:
<bibRefCitation id="EFCDEA82D07CFFA0FD72988CFC5276D9" author="Mors, T &amp; von der Hocht, F. &amp; Wutzler, B." box="[724,908,190,213]" pageId="40" pageNumber="41" pagination="145 - 170" refId="ref35109" refString="Mors, T, von der Hocht, F., and Wutzler, B. 2000. Die erste Wirbeltierfauna aus der miozanen Braunkohle der Niederrheinischen Bucht (Ville-Schichten, Tagebau Hambach). Palaontologische Zeitschrift, 74: 145 - 170. https: // doi. org / 10.1007 / BF 02987958" type="journal article" year="2000">Mörs et al., 2000</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D07CFFA0FC30988CFBCC76D9" author="Mors, T." box="[918,1042,190,213]" pageId="40" pageNumber="41" pagination="177 - 183" refId="ref34899" refString="Mors, T. 2002. Biostratigraphy and paleoecology of continental Tertiary vertebrate faunas in the Lower Rhine Embayment (NW-Germany). Netherlands Journal of Geosciences - Geologie en Mijnbouw, 81: 177 - 183. https: // doi. org / 10.1017 / S 0016774600022411" type="journal article" year="2002">Mörs, 2002</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D07CFFA0FBBB988CFB7C76D9" author="Joyce, W. G." box="[1053,1186,190,213]" pageId="40" pageNumber="41" pagination="21 - 56" refId="ref33516" refString="Joyce, W. G. 2016. A review of the fossil record of turtles of the clade Pan-Chelydridae. Bulletin of the Peabody Museum of Natural History, 57: 21 - 56. https: // doi. org / 10.3374 / 014.057.0103" type="journal article" year="2016">Joyce, 2016</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D07CFFA0FB0B9888FA7C76D9" author="Cernansky, A. &amp; Szyndlar, Z. &amp; Mors, T." box="[1197,1442,186,213]" pageId="40" pageNumber="41" pagination="329 - 354" refId="ref31064" refString="Cernansky, A., Szyndlar, Z., and Mors, T. 2017. Fossil squamate faunas from the Neogene of Hambach (northwestern Germany). Palaeobiodiversity and Palaeoenvironments, 97: 329 - 354. https: // doi. org / 10.1007 / s 12549 - 016 - 0252 - 1" type="journal article" year="2017">Čerňanský et al., 2017</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D07CFFA0FF6698E9FE3B76FF" author="Georgalis, G. L. &amp; Joyce, W. G." box="[192,485,219,243]" pageId="40" pageNumber="41" pagination="115 - 208" refId="ref32268" refString="Georgalis, G. L. and Joyce, W. G. 2017. A review of the fossil record of Old World turtles of the clade Pan-Trionychidae. Bulletin of the Peabody Museum of Natural History, 58: 115 - 208. https: // doi. org / 10.3374 / 014.058.0106" type="journal article" year="2017">Georgalis and Joyce, 2017</bibRefCitation>
; this work.
<taxonomicName id="4C5CECF0D07CFFA0FDF998EEFC9376FF" baseAuthorityName="Ammon" baseAuthorityYear="1911" box="[607,845,220,243]" class="Testudines" family="Geoemydidae" genus="Mauremys" isUncertain="true" kingdom="Animalia" order="Cryptodira" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="sophiae">
<emphasis id="B9284B61D07CFFA0FDF998EEFD1176FF" box="[607,719,220,243]" italics="true" pageId="40" pageNumber="41">Mauremys</emphasis>
cf.
<emphasis id="B9284B61D07CFFA0FD5F98EEFC9376FF" box="[761,845,220,243]" italics="true" pageId="40" pageNumber="41">sophiae</emphasis>
</taxonomicName>
(reported as
<taxonomicName id="4C5CECF0D07CFFA0FC4698E9FB7776FF" baseAuthorityName="Ammon" baseAuthorityYear="1911" box="[992,1193,219,243]" class="Testudines" family="Geoemydidae" genus="Ocadia" isUncertain="true" kingdom="Animalia" order="Cryptodira" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="sophiae">
<emphasis id="B9284B61D07CFFA0FC4698E9FBF276FE" box="[992,1068,219,242]" italics="true" pageId="40" pageNumber="41">Ocadia</emphasis>
cf.
<emphasis id="B9284B61D07CFFA0FBF398EEFB7776FF" box="[1109,1193,220,243]" italics="true" pageId="40" pageNumber="41">sophiae</emphasis>
</taxonomicName>
by
<bibRefCitation id="EFCDEA82D07CFFA0FB7498EEFA9076FF" author="Mors, T." box="[1234,1358,220,243]" pageId="40" pageNumber="41" pagination="177 - 183" refId="ref34899" refString="Mors, T. 2002. Biostratigraphy and paleoecology of continental Tertiary vertebrate faunas in the Lower Rhine Embayment (NW-Germany). Netherlands Journal of Geosciences - Geologie en Mijnbouw, 81: 177 - 183. https: // doi. org / 10.1017 / S 0016774600022411" type="journal article" year="2002">Mörs, 2002</bibRefCitation>
) is here considered as part of
<taxonomicName id="4C5CECF0D07CFFA0FE0898CBFDC0771C" authorityName="Gray" authorityYear="1869" box="[430,542,249,272]" class="Testudines" family="Geoemydidae" genus="Mauremys" kingdom="Animalia" order="Cryptodira" pageId="40" pageNumber="41" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07CFFA0FE0898CBFDC0771C" box="[430,542,249,272]" italics="true" pageId="40" pageNumber="41">Mauremys</emphasis>
</taxonomicName>
following
<bibRefCitation id="EFCDEA82D07CFFA0FD2B98CBFCFB771C" author="Hervet, S." box="[653,805,249,272]" pageId="40" pageNumber="41" pagination="13 - 78" refId="ref33017" refString="Hervet, S. 2004. Systematique du groupe &quot; Palaeochelys sensu lato - Mauremys &quot; (Chelonii, Testudinoidea) du Tertiaire d'Europe occidentale: principaux resultats. Annales de Paleontologie, 90: 13 - 78. https: // doi. org / 10.1016 / j. annpal. 2003.12.002" type="journal article" year="2004">Hervet (2004)</bibRefCitation>
. The Tegelen herpetofauna is mainly based on the Russel-Tiglia-Egypte pit assemblage (
<bibRefCitation id="EFCDEA82D07CFFA0FDA19924FD137721" author="Villa, A. &amp; Blain, H. - A. &amp; van den Hoek Ostende, L. W. &amp; Delfino, M." box="[519,717,278,301]" pageId="40" pageNumber="41" pagination="203 - 219" refId="ref39874" refString="Villa, A., Blain, H. - A., van den Hoek Ostende, L. W., and Delfino, M. 2018 c. Fossil amphibians and reptiles from Tegelen (Province of Limburg) and the early Pleistocene palaeoclimate of The Netherlands. Quaternary Science Reviews, 187: 203 - 219. https: // doi. org / 10.1016 / j. quascirev. 2018.03.020" type="journal article" year="2018">Villa et al., 2018c</bibRefCitation>
), but it also includes the
<taxonomicName id="4C5CECF0D07CFFA0FC469924FBC27721" authorityName="Dumeril" authorityYear="1806" box="[992,1052,278,301]" class="Testudines" family="Emydidae" genus="Emys" kingdom="Animalia" order="Cryptodira" pageId="40" pageNumber="41" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07CFFA0FC469924FBC27721" box="[992,1052,278,301]" italics="true" pageId="40" pageNumber="41">Emys</emphasis>
</taxonomicName>
occurrences reported by
<bibRefCitation id="EFCDEA82D07CFFA0FA909924FED87747" author="Schreuder, A." pageId="40" pageNumber="41" pagination="72 - 75" refId="ref37016" refString="Schreuder, A. 1946. De moerasschildpad, Emys orbicularis (L.), fossiel en levend in Nederland. Natuurhistorisch maandblad, 25: 72 - 75." type="journal article" year="1946">Schreuder (1946)</bibRefCitation>
and
<bibRefCitation id="EFCDEA82D07CFFA0FE9D9906FE207747" author="Villa, A. &amp; Blain, H. - A. &amp; van den Hoek Ostende, L. W. &amp; Delfino, M." box="[315,510,308,331]" pageId="40" pageNumber="41" pagination="203 - 219" refId="ref39874" refString="Villa, A., Blain, H. - A., van den Hoek Ostende, L. W., and Delfino, M. 2018 c. Fossil amphibians and reptiles from Tegelen (Province of Limburg) and the early Pleistocene palaeoclimate of The Netherlands. Quaternary Science Reviews, 187: 203 - 219. https: // doi. org / 10.1016 / j. quascirev. 2018.03.020" type="journal article" year="2018">Villa et al. (2018c)</bibRefCitation>
. Abbreviation: Croc.,
<taxonomicName id="4C5CECF0D07CFFA0FD479901FC887746" box="[737,854,307,330]" class="Crocodylia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="40" pageNumber="41" phylum="Chordata" rank="class">Crocodylia</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="8BE39773D07CFFA0FD799953FCFB7031" pageId="40" pageNumber="41">
<table id="F95C65D3D07C0077FEEA9953FAFB7031" box="[332,1317,353,1597]" colsContinueIn="41.[340,1217,233,963]" gridcols="5" gridrows="33" pageId="40" pageNumber="41">
<tr id="356C9531D07C0077FEEA9953FAFB779C" box="[332,1317,353,400]" gridrow="0" pageId="40" pageNumber="41" rowspan-0="1" rowspan-1="1">
<th id="76BDFC4DD07C0077FD6F9953FCA8779C" box="[713,886,353,400]" gridcol="2" gridrow="0" pageId="40" pageNumber="41">
<emphasis id="B9284B61D07CFFA0FD799953FCA8779C" bold="true" pageId="40" pageNumber="41">Hambach 6C (Middle Miocene)</emphasis>
</th>
<th id="76BDFC4DD07C0077FC2C9953FB99779C" box="[906,1095,353,400]" gridcol="3" gridrow="0" pageId="40" pageNumber="41">
<emphasis id="B9284B61D07CFFA0FC2C9953FBE3779C" bold="true" pageId="40" pageNumber="41">Hambach 11 &amp; 11C (Upper Pliocene)</emphasis>
</th>
<th id="76BDFC4DD07C0077FBFD9953FAFB779C" box="[1115,1317,353,400]" gridcol="4" gridrow="0" pageId="40" pageNumber="41">
<emphasis id="B9284B61D07CFFA0FB3E9953FAFB779C" bold="true" pageId="40" pageNumber="41">Tegelen (Lower Pleistocene)</emphasis>
</th>
</tr>
<tr id="356C9531D07C0077FEEA9996FAFB77B4" box="[332,1317,420,440]" gridrow="1" pageId="40" pageNumber="41" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE379996FD6277B4" box="[401,700,420,440]" gridcol="1" gridrow="1" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379996FDE777B4" authorityName="Fitzinger" authorityYear="1826" box="[401,569,420,440]" class="Amphibia" family="Cryptobranchidae" kingdom="Animalia" order="Caudata" pageId="40" pageNumber="41" phylum="Chordata" rank="family">Cryptobranchidae</taxonomicName>
indet.
</td>
<td id="76BDFC4DD07C0077FD6F9996FCA877B4" box="[713,886,420,440]" gridcol="2" gridrow="1" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA99FBFAFB77D1" box="[332,1317,457,477]" gridrow="2" pageId="40" pageNumber="41" rowspan-0="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE3799FBFD6277D1" box="[401,700,457,477]" gridcol="1" gridrow="2" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE3799FBFD7677D1" baseAuthorityName="Vasilyan and Yanenko" baseAuthorityYear="2020" box="[401,680,457,477]" class="Amphibia" family="Batrachosauroididae" genus="Palaeoproteus" higherTaxonomySource="GBIF" isUncertain="true" kingdom="Animalia" order="Caudata" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="miocenicus">
<emphasis id="B9284B61D07CFFA0FE3799FBFDC477D1" box="[401,538,457,477]" italics="true" pageId="40" pageNumber="41">Palaeoproteus</emphasis>
cf.
<emphasis id="B9284B61D07CFFA0FD9B99FBFD7677D1" box="[573,680,457,477]" italics="true" pageId="40" pageNumber="41">miocenicus</emphasis>
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FD6F99FBFCA877D1" box="[713,886,457,477]" gridcol="2" gridrow="2" pageId="40" pageNumber="41">x</td>
<td id="76BDFC4DD07C0077FC2C99FBFB9977D1" box="[906,1095,457,477]" gridcol="3" gridrow="2" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA99DDFAFB740F" box="[332,1317,495,515]" gridrow="3" pageId="40" pageNumber="41" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE3799DDFD62740F" box="[401,700,495,515]" gridcol="1" gridrow="3" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE3799DDFD93740F" authorityName="Macaluso, Villa, and Mors" authorityYear="2022" box="[401,589,495,515]" class="Amphibia" family="Proteidae" genus="Euronecturus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="grogu">
<emphasis id="B9284B61D07CFFA0FE3799DDFD93740F" box="[401,589,495,515]" italics="true" pageId="40" pageNumber="41">Euronecturus grogu</emphasis>
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FD6F99DDFCA8740F" box="[713,886,495,515]" gridcol="2" gridrow="3" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9A39FAFB7424" box="[332,1317,523,552]" gridrow="4" pageId="40" pageNumber="41" rowspan-2="1" rowspan-4="1">
<th id="76BDFC4DD07C0077FEEA9A39FE5E745A" box="[332,384,523,598]" gridcol="0" gridrow="4" pageId="40" pageNumber="41" rowspan="2" rowspanBelow="1">Urodela</th>
<td id="76BDFC4DD07C0077FE379A39FD627424" box="[401,700,523,552]" gridcol="1" gridrow="4" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379A26FD8D7424" box="[401,595,532,552]" class="Amphibia" family="Proteidae" genus="Mioproteus" higherTaxonomySource="GBIF" isUncertain="true" kingdom="Animalia" order="Caudata" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="wezei">
<emphasis id="B9284B61D07CFFA0FE379A26FE247424" box="[401,506,532,552]" italics="true" pageId="40" pageNumber="41">Mioproteus</emphasis>
cf.
<emphasis id="B9284B61D07CFFA0FDBB9A26FD8D7424" box="[541,595,532,552]" italics="true" pageId="40" pageNumber="41">wezei</emphasis>
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FC2C9A39FB997424" box="[906,1095,523,552]" gridcol="3" gridrow="4" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9A0BFAFB7441" box="[332,1317,569,589]" gridrow="5" pageId="40" pageNumber="41" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE379A0BFD62745A" box="[401,700,569,598]" gridcol="1" gridrow="5" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379A0BFDC27441" box="[401,540,569,589]" class="Amphibia" family="Salamandridae" genus="Chelotriton" kingdom="Animalia" order="Caudata" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FE379A0BFE267441" box="[401,504,569,589]" italics="true" pageId="40" pageNumber="41">Chelotriton</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FD6F9A0BFCA8745A" box="[713,886,569,598]" gridcol="2" gridrow="5" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9A6DFAFB747F" box="[332,1317,607,627]" gridrow="6" pageId="40" pageNumber="41" rowspan-0="1">
<td id="76BDFC4DD07C0077FE379A6DFD62747F" box="[401,700,607,627]" gridcol="1" gridrow="6" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379A6DFDC8747F" box="[401,534,607,627]" class="Amphibia" family="Salamandridae" genus="Lissotriton" kingdom="Animalia" order="Caudata" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FE379A6DFE2C747F" box="[401,498,607,627]" italics="true" pageId="40" pageNumber="41">Lissotriton</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FD6F9A6DFCA8747F" box="[713,886,607,627]" gridcol="2" gridrow="6" pageId="40" pageNumber="41">x</td>
<td id="76BDFC4DD07C0077FC2C9A6DFB99747F" box="[906,1095,607,627]" gridcol="3" gridrow="6" pageId="40" pageNumber="41">x</td>
<td id="76BDFC4DD07C0077FBFD9A6DFAFB747F" box="[1115,1317,607,627]" gridcol="4" gridrow="6" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9AB6FAFB7494" box="[332,1317,644,664]" gridrow="7" pageId="40" pageNumber="41" rowspan-0="1" rowspan-2="1" rowspan-3="1">
<td id="76BDFC4DD07C0077FE379AB6FD627494" box="[401,700,644,664]" gridcol="1" gridrow="7" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379AB6FE067494" authorityName="Rafinesque" authorityYear="1815" box="[401,472,644,664]" class="Amphibia" family="Salamandridae" genus="Triturus" kingdom="Animalia" order="Caudata" pageId="40" pageNumber="41" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07CFFA0FE379AB6FE067494" box="[401,472,644,664]" italics="true" pageId="40" pageNumber="41">Triturus</emphasis>
</taxonomicName>
gr.
<taxonomicName id="4C5CECF0D07CFFA0FE5D9AB6FD947494" baseAuthorityName="Laurenti" baseAuthorityYear="1768" box="[507,586,644,664]" class="Amphibia" family="Salamandridae" genus="Triturus" kingdom="Animalia" order="Caudata" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="cristatus">
<emphasis id="B9284B61D07CFFA0FE5D9AB6FD947494" box="[507,586,644,664]" italics="true" pageId="40" pageNumber="41">cristatus</emphasis>
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FBFD9AB6FAFB7494" box="[1115,1317,644,664]" gridcol="4" gridrow="7" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9A9BFAFB74B1" box="[332,1317,681,701]" gridrow="8" pageId="40" pageNumber="41" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE379A9BFD6274B1" box="[401,700,681,701]" gridcol="1" gridrow="8" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379A9BFE2274B1" box="[401,508,681,701]" class="Amphibia" family="Salamandridae" genus="Triturus" kingdom="Animalia" order="Caudata" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FE379A9BFE0674B1" box="[401,472,681,701]" italics="true" pageId="40" pageNumber="41">Triturus</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FD6F9A9BFCA874B1" box="[713,886,681,701]" gridcol="2" gridrow="8" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9AFDFAFB74EF" box="[332,1317,719,739]" gridrow="9" pageId="40" pageNumber="41" rowspan-0="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE379AFDFD6274EF" box="[401,700,719,739]" gridcol="1" gridrow="9" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379AFDFE2574EF" box="[401,507,719,739]" class="Amphibia" family="Alytidae" genus="Latonia" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FE379AFDFE0974EF" box="[401,471,719,739]" italics="true" pageId="40" pageNumber="41">Latonia</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FD6F9AFDFCA874EF" box="[713,886,719,739]" gridcol="2" gridrow="9" pageId="40" pageNumber="41">x</td>
<td id="76BDFC4DD07C0077FC2C9AFDFB9974EF" box="[906,1095,719,739]" gridcol="3" gridrow="9" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9AC6FAFB7504" box="[332,1317,756,776]" gridrow="10" pageId="40" pageNumber="41" rowspan-0="1" rowspan-2="1">
<td id="76BDFC4DD07C0077FE379AC6FD627504" box="[401,700,756,776]" gridcol="1" gridrow="10" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379AC6FD4C7504" box="[401,658,756,776]" class="Amphibia" family="Palaeobatrachidae" genus="Palaeobatrachus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="eurydices">
<emphasis id="B9284B61D07CFFA0FE379AC6FD4C7504" box="[401,658,756,776]" italics="true" pageId="40" pageNumber="41">Palaeobatrachus eurydices</emphasis>
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FC2C9AC6FB997504" box="[906,1095,756,776]" gridcol="3" gridrow="10" pageId="40" pageNumber="41">x</td>
<td id="76BDFC4DD07C0077FBFD9AC6FAFB7504" box="[1115,1317,756,776]" gridcol="4" gridrow="10" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9B2BFAFB7521" box="[332,1317,793,813]" gridrow="11" pageId="40" pageNumber="41" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE379B2BFD627521" box="[401,700,793,813]" gridcol="1" gridrow="11" pageId="40" pageNumber="41">
?
<taxonomicName id="4C5CECF0D07CFFA0FE3D9B2BFD917521" authorityName="Cope" authorityYear="1865" box="[411,591,793,813]" class="Amphibia" family="Palaeobatrachidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="family">Palaeobatrachidae</taxonomicName>
indet.
</td>
<td id="76BDFC4DD07C0077FD6F9B2BFCA87521" box="[713,886,793,813]" gridcol="2" gridrow="11" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9B0CFAFB755F" box="[332,1317,830,851]" gridrow="12" pageId="40" pageNumber="41" rowspan-0="1" rowspan-2="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE379B0CFD62755F" box="[401,700,830,851]" gridcol="1" gridrow="12" pageId="40" pageNumber="41">
cf.
<taxonomicName id="4C5CECF0D07CFFA0FE089B0DFD96755F" box="[430,584,831,851]" class="Amphibia" family="Pelobatidae" genus="Eopelobates" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FE089B0DFDFB755F" box="[430,549,831,851]" italics="true" pageId="40" pageNumber="41">Eopelobates</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FC2C9B0CFB99755F" box="[906,1095,830,851]" gridcol="3" gridrow="12" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9B56FAFB7574" box="[332,1317,868,888]" gridrow="13" pageId="40" pageNumber="41" rowspan-0="1" rowspan-2="1" rowspan-3="1">
<td id="76BDFC4DD07C0077FE379B56FD627574" box="[401,700,868,888]" gridcol="1" gridrow="13" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379B56FDEA7574" baseAuthorityName="Laurenti" baseAuthorityYear="1768" box="[401,564,868,888]" class="Amphibia" family="Pelobatidae" genus="Pelobates" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9284B61D07CFFA0FE379B56FDEA7574" box="[401,564,868,888]" italics="true" pageId="40" pageNumber="41">Pelobates fuscus</emphasis>
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FBFD9B56FAFB7574" box="[1115,1317,868,888]" gridcol="4" gridrow="13" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9BBBFAFB7591" box="[332,1317,905,925]" gridrow="14" pageId="40" pageNumber="41" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE379BBBFD627591" box="[401,700,905,925]" gridcol="1" gridrow="14" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379BBBFDDC7591" authorityName="Bonaparte" authorityYear="1850" box="[401,514,905,925]" class="Amphibia" family="Pelobatidae" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="family">Pelobatidae</taxonomicName>
indet.
</td>
<td id="76BDFC4DD07C0077FD6F9BBBFCA87591" box="[713,886,905,925]" gridcol="2" gridrow="14" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9B9CFAFB75CF" box="[332,1317,942,963]" gridrow="15" pageId="40" pageNumber="41" rowspan-2="1" rowspan-3="1">
<th id="76BDFC4DD07C0077FEEA9B9CFE5E75E4" box="[332,384,942,1000]" gridcol="0" gridrow="15" pageId="40" pageNumber="41" rowspan="2" rowspanBelow="1">
<taxonomicName id="4C5CECF0D07CFFA0FEFE9B9CFEAE75EB" authorityName="Fischer" authorityYear="1813" box="[344,368,942,999]" class="Amphibia" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="order">Anura</taxonomicName>
</th>
<td id="76BDFC4DD07C0077FE379B9CFD6275CF" box="[401,700,942,963]" gridcol="1" gridrow="15" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379B9DFDCC75CF" box="[401,530,943,963]" class="Amphibia" family="Pelodytidae" genus="Pelodytes" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FE379B9DFE3075CF" box="[401,494,943,963]" italics="true" pageId="40" pageNumber="41">Pelodytes</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FBFD9B9CFAFB75CF" box="[1115,1317,942,963]" gridcol="4" gridrow="15" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9BE6FAFB75E4" box="[332,1317,980,1000]" gridrow="16" pageId="40" pageNumber="41" rowspan-2="1" rowspan-3="1">
<td id="76BDFC4DD07C0077FE379BE6FD6275E4" box="[401,700,980,1000]" gridcol="1" gridrow="16" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379BE6FE6275E4" authorityName="Laurenti" authorityYear="1768" box="[401,444,980,1000]" class="Amphibia" family="Hylidae" genus="Hyla" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07CFFA0FE379BE6FE6275E4" box="[401,444,980,1000]" italics="true" pageId="40" pageNumber="41">Hyla</emphasis>
</taxonomicName>
gr.
<taxonomicName id="4C5CECF0D07CFFA0FE799BE6FDF675E4" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[479,552,980,1000]" class="Amphibia" family="Hylidae" genus="Hyla" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="arborea">
<emphasis id="B9284B61D07CFFA0FE799BE6FDF675E4" box="[479,552,980,1000]" italics="true" pageId="40" pageNumber="41">arborea</emphasis>
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FBFD9BE6FAFB75E4" box="[1115,1317,980,1000]" gridcol="4" gridrow="16" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9BCBFAFB7201" box="[332,1317,1017,1037]" gridrow="17" pageId="40" pageNumber="41" rowspan-0="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE379BCBFD627201" box="[401,700,1017,1037]" gridcol="1" gridrow="17" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379BCBFE3E7201" box="[401,480,1017,1037]" class="Amphibia" family="Hylidae" genus="Hyla" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FE379BCBFE627201" box="[401,444,1017,1037]" italics="true" pageId="40" pageNumber="41">Hyla</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FD6F9BCBFCA87201" box="[713,886,1017,1037]" gridcol="2" gridrow="17" pageId="40" pageNumber="41">x</td>
<td id="76BDFC4DD07C0077FC2C9BCBFB997201" box="[906,1095,1017,1037]" gridcol="3" gridrow="17" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9C2CFAFB723F" box="[332,1317,1054,1075]" gridrow="18" pageId="40" pageNumber="41" rowspan-0="1" rowspan-2="1">
<td id="76BDFC4DD07C0077FE379C2CFD62723F" box="[401,700,1054,1075]" gridcol="1" gridrow="18" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379C2CFE63723E" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[401,445,1054,1074]" class="Amphibia" family="Bufonidae" genus="Bufo" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07CFFA0FE379C2CFE63723E" box="[401,445,1054,1074]" italics="true" pageId="40" pageNumber="41">Bufo</emphasis>
</taxonomicName>
gr.
<emphasis id="B9284B61D07CFFA0FE479C2CFDD4723E" box="[481,522,1054,1074]" italics="true" pageId="40" pageNumber="41">bufo</emphasis>
</td>
<td id="76BDFC4DD07C0077FC2C9C2CFB99723F" box="[906,1095,1054,1075]" gridcol="3" gridrow="18" pageId="40" pageNumber="41">x</td>
<td id="76BDFC4DD07C0077FBFD9C2CFAFB723F" box="[1115,1317,1054,1075]" gridcol="4" gridrow="18" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9C76FAFB7254" box="[332,1317,1092,1112]" gridrow="19" pageId="40" pageNumber="41" rowspan-0="1" rowspan-2="1" rowspan-3="1">
<td id="76BDFC4DD07C0077FE379C76FD627254" box="[401,700,1092,1112]" gridcol="1" gridrow="19" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379C76FDD77254" box="[401,521,1092,1112]" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FE379C76FE3B7254" box="[401,485,1092,1112]" italics="true" pageId="40" pageNumber="41">Bombina</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FBFD9C76FAFB7254" box="[1115,1317,1092,1112]" gridcol="4" gridrow="19" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9C5BFAFB7271" box="[332,1317,1129,1149]" gridrow="20" pageId="40" pageNumber="41" rowspan-0="1" rowspan-3="1">
<td id="76BDFC4DD07C0077FE379C5BFD627271" box="[401,700,1129,1149]" gridcol="1" gridrow="20" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379C5BFDC37271" box="[401,541,1129,1149]" class="Amphibia" family="Ranidae" genus="Pelophylax" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FE379C5BFE277271" box="[401,505,1129,1149]" italics="true" pageId="40" pageNumber="41">Pelophylax</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FD6F9C5BFCA87271" box="[713,886,1129,1149]" gridcol="2" gridrow="20" pageId="40" pageNumber="41">x</td>
<td id="76BDFC4DD07C0077FBFD9C5BFAFB7271" box="[1115,1317,1129,1149]" gridcol="4" gridrow="20" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9CBDFAFB72AF" box="[332,1317,1167,1187]" gridrow="21" pageId="40" pageNumber="41" rowspan-0="1" rowspan-3="1">
<td id="76BDFC4DD07C0077FE379CBDFD6272AF" box="[401,700,1167,1187]" gridcol="1" gridrow="21" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379CBDFE3672AF" box="[401,488,1167,1187]" class="Amphibia" family="Ranidae" genus="Rana" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FE379CBDFE1A72AF" box="[401,452,1167,1187]" italics="true" pageId="40" pageNumber="41">Rana</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FD6F9CBDFCA872AF" box="[713,886,1167,1187]" gridcol="2" gridrow="21" pageId="40" pageNumber="41">x</td>
<td id="76BDFC4DD07C0077FBFD9CBDFAFB72AF" box="[1115,1317,1167,1187]" gridcol="4" gridrow="21" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9C86FAFB72C4" box="[332,1317,1204,1224]" gridrow="22" pageId="40" pageNumber="41" rowspan-0="1">
<td id="76BDFC4DD07C0077FE379C86FD6272C4" box="[401,700,1204,1224]" gridcol="1" gridrow="22" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379C86FE3F72C4" authorityName="Batsch" authorityYear="1796" box="[401,481,1204,1224]" class="Amphibia" family="Ranidae" kingdom="Animalia" order="Anura" pageId="40" pageNumber="41" phylum="Chordata" rank="family">Ranidae</taxonomicName>
indet.
</td>
<td id="76BDFC4DD07C0077FD6F9C86FCA872C4" box="[713,886,1204,1224]" gridcol="2" gridrow="22" pageId="40" pageNumber="41">x</td>
<td id="76BDFC4DD07C0077FC2C9C86FB9972C4" box="[906,1095,1204,1224]" gridcol="3" gridrow="22" pageId="40" pageNumber="41">x</td>
<td id="76BDFC4DD07C0077FBFD9C86FAFB72C4" box="[1115,1317,1204,1224]" gridcol="4" gridrow="22" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9CEBFAFB72E1" box="[332,1317,1241,1261]" gridrow="23" pageId="40" pageNumber="41" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE379CEBFD6272E1" box="[401,700,1241,1261]" gridcol="1" gridrow="23" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379CEBFDA772E1" baseAuthorityName="Bell" baseAuthorityYear="1836" box="[401,633,1241,1261]" class="Testudines" family="Chelydridae" genus="Chelydropsis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="murchisoni">
<emphasis id="B9284B61D07CFFA0FE379CEBFDA772E1" box="[401,633,1241,1261]" italics="true" pageId="40" pageNumber="41">Chelydropsis murchisoni</emphasis>
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FD6F9CEBFCA872E1" box="[713,886,1241,1261]" gridcol="2" gridrow="23" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9CCCFAFB731F" box="[332,1317,1278,1299]" gridrow="24" pageId="40" pageNumber="41" rowspan-0="1" rowspan-2="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE379CCCFD62731F" box="[401,700,1278,1299]" gridcol="1" gridrow="24" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379CCCFDEE731F" box="[401,560,1278,1299]" class="Testudines" family="Chelydridae" genus="Chelydropsis" kingdom="Animalia" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FE379CCCFDD2731E" box="[401,524,1278,1298]" italics="true" pageId="40" pageNumber="41">Chelydropsis</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FC2C9CCCFB99731F" box="[906,1095,1278,1299]" gridcol="3" gridrow="24" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9D16FAFB7334" box="[332,1317,1316,1336]" gridrow="25" pageId="40" pageNumber="41" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE379D16FD627334" box="[401,700,1316,1336]" gridcol="1" gridrow="25" pageId="40" pageNumber="41">Trionychinae indet.</td>
<td id="76BDFC4DD07C0077FD6F9D16FCA87334" box="[713,886,1316,1336]" gridcol="2" gridrow="25" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9D77FAFB7351" box="[332,1317,1349,1373]" gridrow="26" pageId="40" pageNumber="41" rowspan-2="1" rowspan-3="1">
<th id="76BDFC4DD07C0077FEEA9D77FE5E73A7" box="[332,384,1349,1451]" gridcol="0" gridrow="26" pageId="40" pageNumber="41" rowspan="3" rowspanBelow="2">
<taxonomicName id="4C5CECF0D07CFFA0FEFE9D77FEAE73A7" box="[344,368,1349,1451]" class="Testudines" kingdom="Animalia" pageId="40" pageNumber="41" phylum="Chordata" rank="class">Testudines</taxonomicName>
</th>
<td id="76BDFC4DD07C0077FE379D77FD627351" box="[401,700,1349,1373]" gridcol="1" gridrow="26" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379D7BFDF37351" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[401,557,1353,1373]" class="Testudines" family="Emydidae" genus="Emys" kingdom="Animalia" order="Cryptodira" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="orbicularis">
<emphasis id="B9284B61D07CFFA0FE379D7BFDF37351" box="[401,557,1353,1373]" italics="true" pageId="40" pageNumber="41">Emys orbicularis</emphasis>
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FBFD9D77FAFB7351" box="[1115,1317,1349,1373]" gridcol="4" gridrow="26" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9D5DFAFB738F" box="[332,1317,1391,1411]" gridrow="27" pageId="40" pageNumber="41" rowspan-2="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE379D5DFD62738F" box="[401,700,1391,1411]" gridcol="1" gridrow="27" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379D5DFE34738F" box="[401,490,1391,1411]" class="Testudines" family="Emydidae" genus="Emys" kingdom="Animalia" order="Cryptodira" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FE379D5DFE18738F" box="[401,454,1391,1411]" italics="true" pageId="40" pageNumber="41">Emys</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FC2C9D5DFB99738F" box="[906,1095,1391,1411]" gridcol="3" gridrow="27" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9DA6FAFB73A7" box="[332,1317,1428,1451]" gridrow="28" pageId="40" pageNumber="41" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE379DA6FD6273A7" box="[401,700,1428,1451]" gridcol="1" gridrow="28" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379DA6FD9E73A4" box="[401,576,1428,1448]" class="Testudines" family="Emydidae" genus="Clemmydopsis" kingdom="Animalia" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FE379DA6FDC373A4" box="[401,541,1428,1448]" italics="true" pageId="40" pageNumber="41">Clemmydopsis</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FD6F9DA6FCA873A7" box="[713,886,1428,1451]" gridcol="2" gridrow="28" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9D8BFAFB73C1" box="[332,1317,1465,1485]" gridrow="29" pageId="40" pageNumber="41" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE379D8BFD6273C1" box="[401,700,1465,1485]" gridcol="1" gridrow="29" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379D8BFDBC73C1" baseAuthorityName="Ammon" baseAuthorityYear="1911" box="[401,610,1465,1485]" class="Testudines" family="Geoemydidae" genus="Mauremys" isUncertain="true" kingdom="Animalia" order="Cryptodira" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="sophiae">
<emphasis id="B9284B61D07CFFA0FE379D8BFE2B73C1" box="[401,501,1465,1485]" italics="true" pageId="40" pageNumber="41">Mauremys</emphasis>
cf.
<emphasis id="B9284B61D07CFFA0FDB19D8BFDBC73C1" box="[535,610,1465,1485]" italics="true" pageId="40" pageNumber="41">sophiae</emphasis>
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FD6F9D8BFCA873C1" box="[713,886,1465,1485]" gridcol="2" gridrow="29" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9DEDFAFB73FF" box="[332,1317,1503,1523]" gridrow="30" pageId="40" pageNumber="41" rowspan-0="1" rowspan-2="1" rowspan-3="1">
<td id="76BDFC4DD07C0077FE379DEDFD6273FF" box="[401,700,1503,1523]" gridcol="1" gridrow="30" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379DEDFDC673FF" box="[401,536,1503,1523]" class="Testudines" family="Geoemydidae" genus="Mauremys" kingdom="Animalia" order="Cryptodira" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FE379DEDFE2B73FF" box="[401,501,1503,1523]" italics="true" pageId="40" pageNumber="41">Mauremys</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FBFD9DEDFAFB73FF" box="[1115,1317,1503,1523]" gridcol="4" gridrow="30" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9E36FAFB7014" box="[332,1317,1540,1560]" gridrow="31" pageId="40" pageNumber="41" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07C0077FE379E36FD627014" box="[401,700,1540,1560]" gridcol="1" gridrow="31" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379E36FDDE7014" box="[401,512,1540,1560]" class="Testudines" family="Testudinidae" genus="Testudo" kingdom="Animalia" order="Cryptodira" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FE379E36FE027014" box="[401,476,1540,1560]" italics="true" pageId="40" pageNumber="41">Testudo</emphasis>
sp.
</taxonomicName>
vel
<taxonomicName id="4C5CECF0D07CFFA0FD809E36FD627014" box="[550,700,1540,1560]" class="Testudines" family="Testudinidae" genus="Geochelone" kingdom="Animalia" order="Cryptodira" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FD809E36FD477014" box="[550,665,1540,1560]" italics="true" pageId="40" pageNumber="41">Geochelone</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FD6F9E36FCA87014" box="[713,886,1540,1560]" gridcol="2" gridrow="31" pageId="40" pageNumber="41">x</td>
</tr>
<tr id="356C9531D07C0077FEEA9E1BFAFB7031" box="[332,1317,1577,1597]" gridrow="32" pageId="40" pageNumber="41" rowspan-3="1" rowspan-4="1">
<th id="76BDFC4DD07C0077FEEA9E1BFE5E7031" box="[332,384,1577,1597]" gridcol="0" gridrow="32" pageId="40" pageNumber="41">Croc.</th>
<td id="76BDFC4DD07C0077FE379E1BFD627031" box="[401,700,1577,1597]" gridcol="1" gridrow="32" pageId="40" pageNumber="41">
<taxonomicName id="4C5CECF0D07CFFA0FE379E1BFDE87031" box="[401,566,1577,1597]" class="Crocodylia" family="Alligatoridae" genus="Diplocynodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07CFFA0FE379E1BFDCC7031" box="[401,530,1577,1597]" italics="true" pageId="40" pageNumber="41">Diplocynodon</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07C0077FD6F9E1BFCA87031" box="[713,886,1577,1597]" gridcol="2" gridrow="32" pageId="40" pageNumber="41">x</td>
</tr>
</table>
</paragraph>
<caption id="DF23C7FBD07DFFA1FF18989AFE7876B3" box="[190,422,168,191]" pageId="41" pageNumber="42" startId="41.[190,268,168,191]" targetType="table">
<paragraph id="8BE39773D07DFFA1FF18989AFE7876B3" blockId="41.[190,422,168,191]" box="[190,422,168,191]" pageId="41" pageNumber="42">
<emphasis id="B9284B61D07DFFA1FF18989AFEFE76B3" bold="true" box="[190,288,168,191]" pageId="41" pageNumber="42">TABLE 2</emphasis>
(continued).
</paragraph>
</caption>
<paragraph id="8BE39773D07DFFA1FE2B98DBFB1F75CF" pageId="41" pageNumber="42">
<table id="F95C65D3D07D0077FEF298DBFB1F75CF" box="[340,1217,233,963]" colsContinueFrom="40.[332,1317,353,1597]" gridcols="5" gridrows="17" pageId="41" pageNumber="42">
<tr id="356C9531D07D0077FEF298DBFB1F76F1" box="[340,1217,233,253]" gridrow="0" pageId="41" pageNumber="42" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<th id="76BDFC4DD07D0077FE2B98DBFD6676F1" box="[397,696,233,253]" gridcol="1" gridrow="0" pageId="41" pageNumber="42">
<taxonomicName id="4C5CECF0D07DFFA1FE2B98DBFDA176F1" authorityName="Cernansky" authorityYear="2010" box="[397,639,233,253]" class="Squamata" family="Chamaeleonidae" genus="Chamaeleo" isUncertain="true" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="andrusovi">
<emphasis id="B9284B61D07DFFA1FE2B98DBFE2476F1" box="[397,506,233,253]" italics="true" pageId="41" pageNumber="42">Chamaeleo</emphasis>
aff.
<emphasis id="B9284B61D07DFFA1FD8598DBFDA176F1" box="[547,639,233,253]" italics="true" pageId="41" pageNumber="42">andrusovi</emphasis>
</taxonomicName>
</th>
<th id="76BDFC4DD07D0077FCB098DBFCFF76F1" box="[790,801,233,253]" gridcol="2" gridrow="0" pageId="41" pageNumber="42">x</th>
</tr>
<tr id="356C9531D07D0077FEF2993DFB1F772F" box="[340,1217,271,291]" gridrow="1" pageId="41" pageNumber="42" rowspan-0="1" rowspan-2="1" rowspan-3="1">
<td id="76BDFC4DD07D0077FE2B993DFD66772F" box="[397,696,271,291]" gridcol="1" gridrow="1" pageId="41" pageNumber="42">
<taxonomicName id="4C5CECF0D07DFFA1FE2B993DFE26772F" box="[397,504,271,291]" class="Amphibia" family="Salamandridae" genus="Lacerta" kingdom="Animalia" order="Caudata" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07DFFA1FE2B993DFE0A772F" box="[397,468,271,291]" italics="true" pageId="41" pageNumber="42">Lacerta</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07D0077FB10993DFB1F772F" box="[1206,1217,271,291]" gridcol="4" gridrow="1" pageId="41" pageNumber="42">x</td>
</tr>
<tr id="356C9531D07D0077FEF29906FB1F7744" box="[340,1217,308,328]" gridrow="2" pageId="41" pageNumber="42" rowspan-0="1" rowspan-3="1">
<td id="76BDFC4DD07D0077FE2B9906FD667744" box="[397,696,308,328]" gridcol="1" gridrow="2" pageId="41" pageNumber="42">
<taxonomicName id="4C5CECF0D07DFFA1FE2B9906FE2F7744" box="[397,497,308,328]" class="Squamata" family="Lacertidae" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="family">Lacertidae</taxonomicName>
indet.
</td>
<td id="76BDFC4DD07D0077FCB09906FCFF7744" box="[790,801,308,328]" gridcol="2" gridrow="2" pageId="41" pageNumber="42">x</td>
<td id="76BDFC4DD07D0077FB109906FB1F7744" box="[1206,1217,308,328]" gridcol="4" gridrow="2" pageId="41" pageNumber="42">x</td>
</tr>
<tr id="356C9531D07D0077FEF2996BFB1F7761" box="[340,1217,345,365]" gridrow="3" pageId="41" pageNumber="42" rowspan-0="1" rowspan-2="1" rowspan-3="1">
<td id="76BDFC4DD07D0077FE2B996BFD667761" box="[397,696,345,365]" gridcol="1" gridrow="3" pageId="41" pageNumber="42">
<taxonomicName id="4C5CECF0D07DFFA1FE2B996BFE107761" authorityName="Linnaeus" authorityYear="1758" box="[397,462,345,365]" class="Squamata" family="Anguidae" genus="Anguis" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FE2B996BFE107761" box="[397,462,345,365]" italics="true" pageId="41" pageNumber="42">Anguis</emphasis>
</taxonomicName>
gr.
<emphasis id="B9284B61D07DFFA1FE54996BFDF17761" box="[498,559,345,365]" italics="true" pageId="41" pageNumber="42">fragilis</emphasis>
</td>
<td id="76BDFC4DD07D0077FB10996BFB1F7761" box="[1206,1217,345,365]" gridcol="4" gridrow="3" pageId="41" pageNumber="42">x</td>
</tr>
<tr id="356C9531D07D0077FEF2994DFB1F779F" box="[340,1217,383,403]" gridrow="4" pageId="41" pageNumber="42" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07D0077FE2B994DFD66779F" box="[397,696,383,403]" gridcol="1" gridrow="4" pageId="41" pageNumber="42">
<taxonomicName id="4C5CECF0D07DFFA1FE2B994DFD44779F" authorityName="Klembara" authorityYear="2012" box="[397,666,383,403]" class="Squamata" family="Anguidae" genus="Pseudopus" isUncertain="true" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="ahnikoviensis">
<emphasis id="B9284B61D07DFFA1FE2B994DFE26779F" box="[397,504,383,403]" italics="true" pageId="41" pageNumber="42">Pseudopus</emphasis>
cf.
<emphasis id="B9284B61D07DFFA1FDBC994DFD44779F" box="[538,666,383,403]" italics="true" pageId="41" pageNumber="42">ahnikoviensis</emphasis>
</taxonomicName>
</td>
<td id="76BDFC4DD07D0077FCB0994DFCFF779F" box="[790,801,383,403]" gridcol="2" gridrow="4" pageId="41" pageNumber="42">x</td>
</tr>
<tr id="356C9531D07D0077FEF29996FB1F77B4" box="[340,1217,420,440]" gridrow="5" pageId="41" pageNumber="42" rowspan-0="1" rowspan-2="1" rowspan-4="1">
<td id="76BDFC4DD07D0077FE2B9996FD6677B4" box="[397,696,420,440]" gridcol="1" gridrow="5" pageId="41" pageNumber="42">
<taxonomicName id="4C5CECF0D07DFFA1FE2B9996FD5977B4" baseAuthorityName="Kormos" baseAuthorityYear="1911" box="[397,647,420,440]" class="Squamata" family="Anguidae" genus="Pseudopus" isUncertain="true" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="pannonicus">
<emphasis id="B9284B61D07DFFA1FE2B9996FE2677B4" box="[397,504,420,440]" italics="true" pageId="41" pageNumber="42">Pseudopus</emphasis>
cf.
<emphasis id="B9284B61D07DFFA1FDBC9996FD5977B4" box="[538,647,420,440]" italics="true" pageId="41" pageNumber="42">pannonicus</emphasis>
</taxonomicName>
</td>
<td id="76BDFC4DD07D0077FC799996FC3477B4" box="[991,1002,420,440]" gridcol="3" gridrow="5" pageId="41" pageNumber="42">x</td>
</tr>
<tr id="356C9531D07D0077FEF299FBFB1F77D1" box="[340,1217,457,477]" gridrow="6" pageId="41" pageNumber="42" rowspan-0="1" rowspan-2="1" rowspan-3="1">
<td id="76BDFC4DD07D0077FE2B99FBFD6677D1" box="[397,696,457,477]" gridcol="1" gridrow="6" pageId="41" pageNumber="42">
cf.
<taxonomicName id="4C5CECF0D07DFFA1FE0C99FBFDE677D1" box="[426,568,457,477]" class="Squamata" family="Anguidae" genus="Pseudopus" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07DFFA1FE0C99FBFDCB77D1" box="[426,533,457,477]" italics="true" pageId="41" pageNumber="42">Pseudopus</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07D0077FB1099FBFB1F77D1" box="[1206,1217,457,477]" gridcol="4" gridrow="6" pageId="41" pageNumber="42">x</td>
</tr>
<tr id="356C9531D07D0077FEF299DDFB1F740F" box="[340,1217,495,515]" gridrow="7" pageId="41" pageNumber="42" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07D0077FE2B99DDFD66740F" box="[397,696,495,515]" gridcol="1" gridrow="7" pageId="41" pageNumber="42">
<taxonomicName id="4C5CECF0D07DFFA1FE2B99DDFDD9740F" box="[397,519,495,515]" class="Squamata" family="Aniliidae" genus="Eoanilius" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07DFFA1FE2B99DDFE3A740F" box="[397,484,495,515]" italics="true" pageId="41" pageNumber="42">Eoanilius</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07D0077FCB099DDFCFF740F" box="[790,801,495,515]" gridcol="2" gridrow="7" pageId="41" pageNumber="42">x</td>
</tr>
<tr id="356C9531D07D0077FEF29A26FB1F7494" box="[340,1217,532,664]" gridrow="8" pageId="41" pageNumber="42" rowspan-3="1" rowspan-4="1">
<th id="76BDFC4DD07D0077FEF29A26FEB27494" box="[340,364,532,664]" gridcol="0" gridrow="8" pageId="41" pageNumber="42">
<taxonomicName id="4C5CECF0D07DFFA1FEF29A17FEB2748A" box="[340,364,549,646]" class="Squamata" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="class">Squamata</taxonomicName>
</th>
<td id="76BDFC4DD07D0077FE2B9A26FD667494" box="[397,696,532,664]" gridcol="1" gridrow="8" pageId="41" pageNumber="42">
<taxonomicName id="4C5CECF0D07DFFA1FE2B9A26FDC37424" box="[397,541,532,552]" class="Squamata" family="Boidae" genus="Bavarioboa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07DFFA1FE2B9A26FE277424" box="[397,505,532,552]" italics="true" pageId="41" pageNumber="42">Bavarioboa</emphasis>
sp.
</taxonomicName>
cf.
<taxonomicName id="4C5CECF0D07DFFA1FE0C9A0BFE257441" authorityName="Szyndlar and Rage" authorityYear="2003" box="[426,507,569,589]" class="Squamata" family="Tropidophiidae" genus="Falseryx" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FE0C9A0BFE257441" box="[426,507,569,589]" italics="true" pageId="41" pageNumber="42">Falseryx</emphasis>
</taxonomicName>
sp. “
<taxonomicName id="4C5CECF0D07DFFA1FE329A6DFE01747F" authorityName="Linnaeus" authorityYear="1758" box="[404,479,607,627]" class="Squamata" family="Colubridae" genus="Coluber" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FE329A6DFE01747F" box="[404,479,607,627]" italics="true" pageId="41" pageNumber="42">Coluber</emphasis>
</taxonomicName>
” sp.
<taxonomicName id="4C5CECF0D07DFFA1FE2B9AB6FDC27494" box="[397,540,644,664]" class="Squamata" family="Colubridae" genus="Texasophis" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07DFFA1FE2B9AB6FE267494" box="[397,504,644,664]" italics="true" pageId="41" pageNumber="42">Texasophis</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07D0077FCB09A26FCFF7494" box="[790,801,532,664]" gridcol="2" gridrow="8" pageId="41" pageNumber="42">x x x x</td>
</tr>
<tr id="356C9531D07D0077FEF29A98FB1F74B2" box="[340,1217,682,702]" gridrow="9" pageId="41" pageNumber="42" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07D0077FE2B9A98FD6674B2" box="[397,696,682,702]" gridcol="1" gridrow="9" pageId="41" pageNumber="42">
<taxonomicName id="4C5CECF0D07DFFA1FE2B9A98FDC274B2" box="[397,540,682,702]" class="Squamata" family="Colubridae" genus="Telescopus" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07DFFA1FE2B9A98FE2674B2" box="[397,504,682,702]" italics="true" pageId="41" pageNumber="42">Telescopus</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07D0077FCB09A98FCFF74B2" box="[790,801,682,702]" gridcol="2" gridrow="9" pageId="41" pageNumber="42">x</td>
</tr>
<tr id="356C9531D07D0077FEF29AFDFB1F74EF" box="[340,1217,719,739]" gridrow="10" pageId="41" pageNumber="42" rowspan-0="1" rowspan-2="1">
<td id="76BDFC4DD07D0077FE2B9AFDFD6674EF" box="[397,696,719,739]" gridcol="1" gridrow="10" pageId="41" pageNumber="42">“Colubrinae”/”Colubrines” indet.</td>
<td id="76BDFC4DD07D0077FC799AFDFC3474EF" box="[991,1002,719,739]" gridcol="3" gridrow="10" pageId="41" pageNumber="42">x</td>
<td id="76BDFC4DD07D0077FB109AFDFB1F74EF" box="[1206,1217,719,739]" gridcol="4" gridrow="10" pageId="41" pageNumber="42">x</td>
</tr>
<tr id="356C9531D07D0077FEF29AC6FB1F7504" box="[340,1217,756,776]" gridrow="11" pageId="41" pageNumber="42" rowspan-0="1" rowspan-2="1" rowspan-3="1">
<td id="76BDFC4DD07D0077FE2B9AC6FD667504" box="[397,696,756,776]" gridcol="1" gridrow="11" pageId="41" pageNumber="42">
<taxonomicName id="4C5CECF0D07DFFA1FE2B9AC6FE217504" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[397,511,756,776]" class="Squamata" family="Colubridae" genus="Natrix" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="natrix">
<emphasis id="B9284B61D07DFFA1FE2B9AC6FE217504" box="[397,511,756,776]" italics="true" pageId="41" pageNumber="42">Natrix natrix</emphasis>
</taxonomicName>
</td>
<td id="76BDFC4DD07D0077FB109AC6FB1F7504" box="[1206,1217,756,776]" gridcol="4" gridrow="11" pageId="41" pageNumber="42">x</td>
</tr>
<tr id="356C9531D07D0077FEF29B28FB1F7522" box="[340,1217,794,814]" gridrow="12" pageId="41" pageNumber="42" rowspan-0="1" rowspan-4="1">
<td id="76BDFC4DD07D0077FE2B9B28FD667522" box="[397,696,794,814]" gridcol="1" gridrow="12" pageId="41" pageNumber="42">
<taxonomicName id="4C5CECF0D07DFFA1FE2B9B28FE367522" box="[397,488,794,814]" class="Squamata" family="Colubridae" genus="Natrix" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07DFFA1FE2B9B28FE1B7522" box="[397,453,794,814]" italics="true" pageId="41" pageNumber="42">Natrix</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07D0077FCB09B28FCFF7522" box="[790,801,794,814]" gridcol="2" gridrow="12" pageId="41" pageNumber="42">x</td>
<td id="76BDFC4DD07D0077FC799B28FC347522" box="[991,1002,794,814]" gridcol="3" gridrow="12" pageId="41" pageNumber="42">x</td>
</tr>
<tr id="356C9531D07D0077FEF29B0DFB1F755F" box="[340,1217,831,851]" gridrow="13" pageId="41" pageNumber="42" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07D0077FE2B9B0DFD66755F" box="[397,696,831,851]" gridcol="1" gridrow="13" pageId="41" pageNumber="42">
cf.
<taxonomicName id="4C5CECF0D07DFFA1FE0C9B0DFE27755F" box="[426,505,831,851]" class="Squamata" family="Elapidae" genus="Naja" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07DFFA1FE0C9B0DFE08755F" box="[426,470,831,851]" italics="true" pageId="41" pageNumber="42">Naja</emphasis>
sp.
</taxonomicName>
</td>
<td id="76BDFC4DD07D0077FCB09B0DFCFF755F" box="[790,801,831,851]" gridcol="2" gridrow="13" pageId="41" pageNumber="42">x</td>
</tr>
<tr id="356C9531D07D0077FEF29B56FB1F7574" box="[340,1217,868,888]" gridrow="14" pageId="41" pageNumber="42" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07D0077FE2B9B56FD667574" box="[397,696,868,888]" gridcol="1" gridrow="14" pageId="41" pageNumber="42">
<taxonomicName id="4C5CECF0D07DFFA1FE2B9B56FE307574" box="[397,494,868,888]" class="Squamata" family="Viperidae" genus="Vipera" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07DFFA1FE2B9B56FE147574" box="[397,458,868,888]" italics="true" pageId="41" pageNumber="42">Vipera</emphasis>
sp.
</taxonomicName>
(“Oriental viper”)
</td>
<td id="76BDFC4DD07D0077FCB09B56FCFF7574" box="[790,801,868,888]" gridcol="2" gridrow="14" pageId="41" pageNumber="42">x</td>
</tr>
<tr id="356C9531D07D0077FEF29BB8FB1F7592" box="[340,1217,906,926]" gridrow="15" pageId="41" pageNumber="42" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<td id="76BDFC4DD07D0077FE2B9BB8FD667592" box="[397,696,906,926]" gridcol="1" gridrow="15" pageId="41" pageNumber="42">
<taxonomicName id="4C5CECF0D07DFFA1FE2B9BB8FE307592" box="[397,494,906,926]" class="Squamata" family="Viperidae" genus="Vipera" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07DFFA1FE2B9BB8FE157592" box="[397,459,906,926]" italics="true" pageId="41" pageNumber="42">Vipera</emphasis>
sp.
</taxonomicName>
(“
<emphasis id="B9284B61D07DFFA1FDA79BB8FDED7592" box="[513,563,906,926]" italics="true" pageId="41" pageNumber="42">aspis</emphasis>
complex”)
</td>
<td id="76BDFC4DD07D0077FCB09BB8FCFF7592" box="[790,801,906,926]" gridcol="2" gridrow="15" pageId="41" pageNumber="42">x</td>
</tr>
<tr id="356C9531D07D0077FEF29B9DFB1F75CF" box="[340,1217,943,963]" gridrow="16" pageId="41" pageNumber="42" rowspan-0="1" rowspan-2="1" rowspan-3="1">
<td id="76BDFC4DD07D0077FE2B9B9DFD6675CF" box="[397,696,943,963]" gridcol="1" gridrow="16" pageId="41" pageNumber="42">
<taxonomicName id="4C5CECF0D07DFFA1FE2B9B9DFE3075CF" box="[397,494,943,963]" class="Squamata" family="Viperidae" genus="Vipera" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B9284B61D07DFFA1FE2B9B9DFE1575CF" box="[397,459,943,963]" italics="true" pageId="41" pageNumber="42">Vipera</emphasis>
sp.
</taxonomicName>
(small viper)
</td>
<td id="76BDFC4DD07D0077FB109B9DFB1F75CF" box="[1206,1217,943,963]" gridcol="4" gridrow="16" pageId="41" pageNumber="42">x</td>
</tr>
</table>
</paragraph>
<paragraph id="8BE39773D07DFFA1FF5C9C6BFCA4717E" blockId="41.[192,793,1049,1906]" lastBlockId="41.[840,1441,1049,1906]" pageId="41" pageNumber="42">
When looking at reptiles, crocodylians are absent in both the Pliocene of Hambach and at Tegelen, whereas several cranial and postcranial remains attributed to
<taxonomicName id="4C5CECF0D07DFFA1FE1F9C8BFD8472DE" authorityName="Pomel" authorityYear="1847" box="[441,602,1209,1234]" class="Crocodylia" family="Alligatoridae" genus="Diplocynodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FE1F9C8BFD8472DE" box="[441,602,1209,1234]" italics="true" pageId="41" pageNumber="42">Diplocynodon</emphasis>
</taxonomicName>
were recovered in the Miocene level of the former (
<bibRefCitation id="EFCDEA82D07DFFA1FD239CEBFF21731E" author="Mors, T &amp; von der Hocht, F. &amp; Wutzler, B." pageId="41" pageNumber="42" pagination="145 - 170" refId="ref35109" refString="Mors, T, von der Hocht, F., and Wutzler, B. 2000. Die erste Wirbeltierfauna aus der miozanen Braunkohle der Niederrheinischen Bucht (Ville-Schichten, Tagebau Hambach). Palaontologische Zeitschrift, 74: 145 - 170. https: // doi. org / 10.1007 / BF 02987958" type="journal article" year="2000">Mörs et al., 2000</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D07DFFA1FEB09CCBFE70731E" author="Mors, T." box="[278,430,1273,1298]" pageId="41" pageNumber="42" pagination="177 - 183" refId="ref34899" refString="Mors, T. 2002. Biostratigraphy and paleoecology of continental Tertiary vertebrate faunas in the Lower Rhine Embayment (NW-Germany). Netherlands Journal of Geosciences - Geologie en Mijnbouw, 81: 177 - 183. https: // doi. org / 10.1017 / S 0016774600022411" type="journal article" year="2002">Mörs, 2002</bibRefCitation>
). Turtles from Tegelen only include
<taxonomicName id="4C5CECF0D07DFFA1FE8C9D2BFDE4733E" authority="Dumeril, 1806" authorityName="Dumeril" authorityYear="1806" box="[298,570,1304,1330]" class="Testudines" family="Emydidae" genus="Emys" kingdom="Animalia" order="Cryptodira" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FE8C9D2BFEB3733E" box="[298,365,1305,1330]" italics="true" pageId="41" pageNumber="42">Emys</emphasis>
<bibRefCitation id="EFCDEA82D07DFFA1FE249D2AFDE4733E" author="Dumeril, A. M. C." box="[386,570,1304,1330]" pageId="41" pageNumber="42" refId="ref31625" refString="Dumeril, A. M. C. 1806. Zoologie Analytique, ou Methode Naturelle de Classification des Animaux, Rendue plus Facile a l'Aide de Tableaux Synoptiques. Allais, Paris. https: // doi. org / 10.5962 / bhl. title. 44835" type="book" year="1806">Duméril, 1806</bibRefCitation>
</taxonomicName>
, and
<taxonomicName id="4C5CECF0D07DFFA1FD329D2BFCCE733E" authorityName="Gray" authorityYear="1869" box="[660,784,1305,1330]" class="Testudines" family="Geoemydidae" genus="Mauremys" kingdom="Animalia" order="Cryptodira" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FD329D2BFCCE733E" box="[660,784,1305,1330]" italics="true" pageId="41" pageNumber="42">Mauremys</emphasis>
</taxonomicName>
, moreover never found together in the same pit (
<bibRefCitation id="EFCDEA82D07DFFA1FF609D6AFE4D737E" author="Schreuder, A." box="[198,403,1368,1394]" pageId="41" pageNumber="42" pagination="72 - 75" refId="ref37016" refString="Schreuder, A. 1946. De moerasschildpad, Emys orbicularis (L.), fossiel en levend in Nederland. Natuurhistorisch maandblad, 25: 72 - 75." type="journal article" year="1946">Schreuder, 1946</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D07DFFA1FE079D6BFEE4739E" author="Van den Hoek Ostende, L. W. &amp; de Vos, J." pageId="41" pageNumber="42" pagination="291 - 304" refId="ref38828" refString="Van den Hoek Ostende, L. W. and de Vos, J. 2006. A century of research on the classical locality of Tegelen (province of Limburg, The Netherlands). Courier Forschungsinstitut Senckenberg, 256: 291 - 304." type="journal article" year="2006">van den Hoek Ostende and de Vos, 2006</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D07DFFA1FEE39D4BFDC9739E" author="Villa, A. &amp; Blain, H. - A. &amp; van den Hoek Ostende, L. W. &amp; Delfino, M." box="[325,535,1401,1426]" pageId="41" pageNumber="42" pagination="203 - 219" refId="ref39874" refString="Villa, A., Blain, H. - A., van den Hoek Ostende, L. W., and Delfino, M. 2018 c. Fossil amphibians and reptiles from Tegelen (Province of Limburg) and the early Pleistocene palaeoclimate of The Netherlands. Quaternary Science Reviews, 187: 203 - 219. https: // doi. org / 10.1016 / j. quascirev. 2018.03.020" type="journal article" year="2018">Villa et al., 2018c</bibRefCitation>
). Hambach displays a higher diversity in the Miocene, with several distinct lineages and both aquatic/semiaquatic and terrestrial animals, but only two taxa in the Pliocene (
<bibRefCitation id="EFCDEA82D07DFFA1FEAF9DCBFE48701E" author="Mors, T." box="[265,406,1529,1554]" pageId="41" pageNumber="42" pagination="177 - 183" refId="ref34899" refString="Mors, T. 2002. Biostratigraphy and paleoecology of continental Tertiary vertebrate faunas in the Lower Rhine Embayment (NW-Germany). Netherlands Journal of Geosciences - Geologie en Mijnbouw, 81: 177 - 183. https: // doi. org / 10.1017 / S 0016774600022411" type="journal article" year="2002">Mörs, 2002</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D07DFFA1FE049DCBFD76701E" author="Klein, N. &amp; Mors, T." box="[418,680,1529,1554]" pageId="41" pageNumber="42" pagination="1 - 48" refId="ref33674" refString="Klein, N. and Mors, T. 2003. Die Schildkroten (Reptilia: Testudines) aus dem Mittel-Miozan von Hambach (Niederrheinische Bucht, NW-Deutschland). Palaeontographica A, 268: 1 - 48. https: // doi. org / 10.1127 / pala / 268 / 2003 / 1" type="journal article" year="2003">Klein and Mörs, 2003</bibRefCitation>
).
<taxonomicName id="4C5CECF0D07DFFA1FD1C9DCBFD23701E" authorityName="Dumeril" authorityYear="1806" box="[698,765,1529,1554]" class="Testudines" family="Emydidae" genus="Emys" kingdom="Animalia" order="Cryptodira" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FD1C9DCBFD23701E" box="[698,765,1529,1554]" italics="true" pageId="41" pageNumber="42">Emys</emphasis>
</taxonomicName>
is shared between the Pliocene of Hambach (even though this occurrence still lacks a detailed publication, with description and figures of the remains, and should be treated with caution) and the Pleistocene of Tegelen, but Hambach misses
<taxonomicName id="4C5CECF0D07DFFA1FD629EABFF2F70DE" authorityName="Gray" authorityYear="1869" class="Testudines" family="Geoemydidae" genus="Mauremys" kingdom="Animalia" order="Cryptodira" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FD629EABFF2F70DE" italics="true" pageId="41" pageNumber="42">Mauremys</emphasis>
</taxonomicName>
and Tegelen misses
<taxonomicName id="4C5CECF0D07DFFA1FE4B9E8AFD5970DE" authorityName="Peters" authorityYear="1868" box="[493,647,1720,1746]" class="Testudines" family="Chelydridae" genus="Chelydropsis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FE4B9E8AFD5970DE" box="[493,647,1720,1746]" italics="true" pageId="41" pageNumber="42">Chelydropsis</emphasis>
</taxonomicName>
(again, if its presence in Pliocene Hambach is confirmed). Squamate faunas from Hambach 6C and Tegelen are very different, with the former being far more diverse (especially in the snake component) and including several Palaeogene/Neogene lineages missing in Tegelen (i.e., chameleons,
<taxonomicName id="4C5CECF0D07DFFA1FA929C2BFC02725E" authority="Rage, 1974" authorityName="Rage" authorityYear="1974" class="Squamata" family="Aniliidae" genus="Eoanilius" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FA929C2BFA7E723E" box="[1332,1440,1049,1074]" italics="true" pageId="41" pageNumber="42">Eoanilius</emphasis>
<bibRefCitation id="EFCDEA82D07DFFA1FCEE9C0BFC02725E" author="Rage, J. - C." box="[840,988,1081,1106]" pageId="41" pageNumber="42" pagination="274 - 303" refId="ref35652" refString="Rage, J. - C. 1974. Les serpents des Phosphorites du Quercy. Palaeovertebrata, 6: 274 - 303." type="journal article" year="1974">Rage, 1974</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C5CECF0D07DFFA1FC489C0BFC59727E" authority="Szyndlar and Schleich, 1993" authorityName="Szyndlar and Schleich" authorityYear="1993" class="Squamata" family="Boidae" genus="Bavarioboa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FC489C0BFBAB725E" box="[1006,1141,1081,1106]" italics="true" pageId="41" pageNumber="42">Bavarioboa</emphasis>
<bibRefCitation id="EFCDEA82D07DFFA1FB259C0AFC59727E" author="Szyndlar, Z. &amp; Schleich, H. H." pageId="41" pageNumber="42" pagination="1 - 47" refId="ref38233" refString="Szyndlar, Z. and Schleich, H. H. 1993. Description of Miocene snakes from Petersbuch 2 with comments on the lower and middle Miocene ophidian faunas of Southern Germany. Stuttgarter Beitrage zur Naturkunde, Serie B (Geologie und Palaontologie), 192: 1 - 47." type="journal article" year="1993">Szyndlar and Schleich, 1993</bibRefCitation>
</taxonomicName>
, cf.
<taxonomicName id="4C5CECF0D07DFFA1FC679C6BFAAD727E" authority="Szyndlar and Rage, 2003" authorityName="Szyndlar and Rage" authorityYear="2003" box="[961,1395,1112,1138]" class="Squamata" family="Tropidophiidae" genus="Falseryx" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FC679C6BFBF8727E" box="[961,1062,1113,1138]" italics="true" pageId="41" pageNumber="42">Falseryx</emphasis>
<bibRefCitation id="EFCDEA82D07DFFA1FB949C6AFAAD727E" author="Szyndlar, Z. &amp; Rage, J. - C." box="[1074,1395,1112,1138]" pageId="41" pageNumber="42" refId="ref38194" refString="Szyndlar, Z. and Rage, J. - C. 2003. Non-erycine Booidea from the Oligocene and Miocene of Europe. Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Krakow." type="book" year="2003">Szyndlar and Rage, 2003</bibRefCitation>
</taxonomicName>
, cf.
<taxonomicName id="4C5CECF0D07DFFA1FCEE9C4BFB96729E" authority="Laurenti, 1768" authorityName="Laurenti" authorityYear="1768" box="[840,1096,1145,1170]" class="Squamata" family="Elapidae" genus="Naja" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FCEE9C4BFCA1729E" box="[840,895,1145,1170]" italics="true" pageId="41" pageNumber="42">Naja</emphasis>
<bibRefCitation id="EFCDEA82D07DFFA1FC369C4BFB96729E" author="Laurenti, J. N." box="[912,1096,1145,1170]" pageId="41" pageNumber="42" refId="ref34062" refString="Laurenti, J. N. 1768. Austriaci Viennensis specimen medicum, exhibens synopsin reptilium emendatam cum experimentis circa Venena et antidota reptilium Austriacorum quod authoritate et consensu. Joannis Thomae de Trattnern, Caes. Reg. Maj. Aulae Typographi et Bibliopolae, Wien. https: // doi. org / 10.5962 / bhl. title. 5108" type="book" year="1768">Laurenti, 1768</bibRefCitation>
</taxonomicName>
, and the “Oriental vipers”;
<bibRefCitation id="EFCDEA82D07DFFA1FCEE9CA6FB8272BE" author="Cernansky, A. &amp; Szyndlar, Z. &amp; Mors, T." box="[840,1116,1172,1202]" pageId="41" pageNumber="42" pagination="329 - 354" refId="ref31064" refString="Cernansky, A., Szyndlar, Z., and Mors, T. 2017. Fossil squamate faunas from the Neogene of Hambach (northwestern Germany). Palaeobiodiversity and Palaeoenvironments, 97: 329 - 354. https: // doi. org / 10.1007 / s 12549 - 016 - 0252 - 1" type="journal article" year="2017">Čerňanský et al., 2017</bibRefCitation>
). A small viper is present in both, even though it is not clear if they may represent the same viper group. “Colubrine” snakes and
<taxonomicName id="4C5CECF0D07DFFA1FCEE9CCBFC50731E" authorityName="Laurenti" authorityYear="1768" box="[840,910,1273,1298]" class="Squamata" family="Colubridae" genus="Natrix" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FCEE9CCBFC50731E" box="[840,910,1273,1298]" italics="true" pageId="41" pageNumber="42">Natrix</emphasis>
</taxonomicName>
are also shared between Tegelen and both Miocene and Pliocene levels in Hambach. However, Pleistocene and Pliocene “colubrines” remain unidentified at lower taxonomic ranks, whereas the Miocene ones include at least three different taxa (“
<taxonomicName id="4C5CECF0D07DFFA1FCFC9DAAFC6973BD" authority="Linnaeus, 1758" authorityName="Linnaeus" authorityYear="1758" box="[858,951,1432,1457]" class="Squamata" family="Colubridae" genus="Coluber" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FCFC9DAAFC6973BD" box="[858,951,1432,1457]" italics="true" pageId="41" pageNumber="42">Coluber</emphasis>
</taxonomicName>
<bibRefCitation id="EFCDEA82D07DFFA1FC769DABFB4D73BE" author="Linnaeus, C." box="[976,1171,1433,1458]" pageId="41" pageNumber="42" refId="ref34124" refString="Linnaeus, C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Tomus 1, Pars 1. Laurentii Salvii, Stockholm. https: // doi. org / 10.5962 / bhl. title. 542" type="book" year="1758">Linnaeus, 1758</bibRefCitation>
,
<taxonomicName id="4C5CECF0D07DFFA1FB019DABFC5973DE" authority="Holman, 1977" authorityName="Holman" authorityYear="1977" class="Squamata" family="Colubridae" genus="Texasophis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FB019DABFAF373BE" box="[1191,1325,1433,1458]" italics="true" pageId="41" pageNumber="42">Texasophis</emphasis>
<bibRefCitation id="EFCDEA82D07DFFA1FA9B9DABFC5973DE" author="Holman, J. A." pageId="41" pageNumber="42" pagination="391 - 403" refId="ref33217" refString="Holman, J. A. 1977. Amphibians and reptiles from the Gulf Coast Miocene of Texas. Herpetologica, 33: 391 - 403." type="journal article" year="1977">Holman, 1977</bibRefCitation>
</taxonomicName>
, and
<taxonomicName id="4C5CECF0D07DFFA1FC619D8BFB2973DE" authority="Wagler, 1830" authorityName="Wagler" authorityYear="1830" box="[967,1271,1465,1490]" class="Squamata" family="Colubridae" genus="Telescopus" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FC619D8BFB9373DE" box="[967,1101,1465,1490]" italics="true" pageId="41" pageNumber="42">Telescopus</emphasis>
<bibRefCitation id="EFCDEA82D07DFFA1FBF29D8BFB2973DE" author="Wagler, J." box="[1108,1271,1465,1490]" pageId="41" pageNumber="42" refId="ref40047" refString="Wagler, J. 1830. Naturliches System der Amphibien, mit vorangehender Classification der Saugthiere und Vogel. Ein Beitrag zur vergleichenden Zoologie. J. G. Cotta, Munchen, Stuttgart and Tubingen." type="book" year="1830">Wagler, 1830</bibRefCitation>
</taxonomicName>
). Not a lot can be said about lacertids, except for them being recovered both in the Middle Miocene of Hambach (
<bibRefCitation id="EFCDEA82D07DFFA1FCE89E26FBAF703E" author="Cernansky, A. &amp; Szyndlar, Z. &amp; Mors, T." box="[846,1137,1556,1586]" pageId="41" pageNumber="42" pagination="329 - 354" refId="ref31064" refString="Cernansky, A., Szyndlar, Z., and Mors, T. 2017. Fossil squamate faunas from the Neogene of Hambach (northwestern Germany). Palaeobiodiversity and Palaeoenvironments, 97: 329 - 354. https: // doi. org / 10.1007 / s 12549 - 016 - 0252 - 1" type="journal article" year="2017">Čerňanský et al., 2017</bibRefCitation>
) and the Pleistocene of Tegelen (
<bibRefCitation id="EFCDEA82D07DFFA1FC129E0BFB55705E" author="Villa, A. &amp; Blain, H. - A. &amp; van den Hoek Ostende, L. W. &amp; Delfino, M." box="[948,1163,1593,1618]" pageId="41" pageNumber="42" pagination="203 - 219" refId="ref39874" refString="Villa, A., Blain, H. - A., van den Hoek Ostende, L. W., and Delfino, M. 2018 c. Fossil amphibians and reptiles from Tegelen (Province of Limburg) and the early Pleistocene palaeoclimate of The Netherlands. Quaternary Science Reviews, 187: 203 - 219. https: // doi. org / 10.1016 / j. quascirev. 2018.03.020" type="journal article" year="2018">Villa et al., 2018c</bibRefCitation>
). Diversity seems to be comparable (two taxa), but there seems to be no evidence in Hambach of a possible green lizard as in Tegelen.
<taxonomicName id="4C5CECF0D07DFFA1FC719EABFB8270BE" authorityName="Merrem" authorityYear="1820" box="[983,1116,1689,1714]" class="Squamata" family="Anguidae" genus="Pseudopus" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="B9284B61D07DFFA1FC719EABFB8270BE" box="[983,1116,1689,1714]" italics="true" pageId="41" pageNumber="42">Pseudopus</emphasis>
</taxonomicName>
is present in all Hambach levels (
<bibRefCitation id="EFCDEA82D07DFFA1FC3A9E86FB6C70DE" author="Cernansky, A. &amp; Szyndlar, Z. &amp; Mors, T." box="[924,1202,1716,1746]" pageId="41" pageNumber="42" pagination="329 - 354" refId="ref31064" refString="Cernansky, A., Szyndlar, Z., and Mors, T. 2017. Fossil squamate faunas from the Neogene of Hambach (northwestern Germany). Palaeobiodiversity and Palaeoenvironments, 97: 329 - 354. https: // doi. org / 10.1007 / s 12549 - 016 - 0252 - 1" type="journal article" year="2017">Čerňanský et al., 2017</bibRefCitation>
; this work) and tentatively even in Tegelen. However, Miocene and Pliocene assemblages in Hambach bear two different
<taxonomicName id="4C5CECF0D07DFFA1FCDF9F2BFBBB713E" box="[889,1125,1817,1842]" pageId="41" pageNumber="42">
<emphasis id="B9284B61D07DFFA1FCDF9F2BFC20713E" box="[889,1022,1817,1842]" italics="true" pageId="41" pageNumber="42">Pseudopus</emphasis>
species
</taxonomicName>
, and it is not clear if the Tegelen one may be related to either one of the two.
</paragraph>
<paragraph id="8BE39773D07EFFA2FF5C9894FB957573" blockId="42.[192,793,166,1887]" lastBlockId="42.[840,1441,166,895]" pageId="42" pageNumber="43">
It seems, thus, that a major change in the reptilian palaeofauna happened already at some point in the Middle Miocene-Late Pliocene interval, with less significant differences between the Upper Pliocene assemblage of Hambach 11 and 11C and the Lower Pleistocene ones of the Tegelen pits (and in particular the Russel-Tiglia-Egypte pit). On the other hand, differences in the batrachofauna are more marked between Hambach 11 and 11C and Tegelen compared to the two levels in Hambach. Whilst the faunal change affecting reptiles agrees with the well-recognized pattern of extirpation of thermophilic taxa from northern Europe characterizing the late Neogene and Quaternary (e.g.,
<bibRefCitation id="EFCDEA82D07EFFA2FF669A54FE7D7473" author="Delfino, M. &amp; Sala, B." box="[192,419,614,639]" pageId="42" pageNumber="43" pagination="716 - 719" refId="ref31426" refString="Delfino, M. and Sala, B. 2007. Late Pliocene Albanerpetontidae (Lissamphibia) from Italy. Journal of Vertebrate Paleontology, 27: 716 - 719. https: // doi. org / 10.1671 / 0272 - 4634 (2007) 27 [716: LPALFI] 2.0. CO; 2" type="journal article" year="2007">Delfino et al., 2007</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D07EFFA2FE099A54FDE37473" author="Rage, J. - C." box="[431,573,614,639]" pageId="42" pageNumber="43" pagination="517 - 534" refId="ref35676" refString="Rage, J. - C. 2013. Mesozoic and Cenozoic squamates of Europe. Palaeobiodiversity and Palaeoenvironments, 93: 517 - 534. https: // doi. org / 10.1007 / s 12549 - 013 - 0124 - x" type="journal article" year="2013">Rage, 2013</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D07EFFA2FDEF9A54FCCA7473" author="Blain, H. - A. &amp; Bailon, S. &amp; Agusti, J." box="[585,788,614,639]" pageId="42" pageNumber="43" pagination="731 - 744" refId="ref30239" refString="Blain, H. - A., Bailon, S., and Agusti, J. 2016. The geographical and chronological pattern of herpetofaunal Pleistocene extinctions on the Iberian Peninsula. Comptes Rendus Palevol, 15: 731 - 744. https: // doi. org / 10.1016 / j. crpv. 2015.05.008" type="journal article" year="2016">Blain et al., 2016</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D07EFFA2FF669AB4FE027493" author="Villa, A. &amp; Delfino, M." box="[192,476,646,671]" pageId="42" pageNumber="43" pagination="177 - 211" refId="ref39467" refString="Villa, A. and Delfino, M. 2019. Fossil lizards and worm lizards (Reptilia, Squamata) from the Neogene and Quaternary of Europe: an overview. Swiss Journal of Palaeontology, 138: 177 - 211. https: // doi. org / 10.1007 / s 13358 - 018 - 0172 - y" type="journal article" year="2019">Villa and Delfino, 2019</bibRefCitation>
), amphibians here appear to be less affected by this trend. The differences between Hambach and Tegelen in the amphibian palaeocommunity may be simply explained by different palaeoenvironments (swamp vs floodplain, respectively), but the peculiarity of the batrachofauna in Hambach 11 and 11C (with several taxa otherwise unknown from northwestern Europe in contemporary times) and its similarities with the Hambach 6C one suggest that other factors may be also at play. Reconstructed palaeoclimate at Tegelen based on the herpetofaunistic association recovered from the Russel-Tiglia-Egypte pit (
<bibRefCitation id="EFCDEA82D07EFFA2FD459C34FE887233" author="Villa, A. &amp; Blain, H. - A. &amp; van den Hoek Ostende, L. W. &amp; Delfino, M." pageId="42" pageNumber="43" pagination="203 - 219" refId="ref39874" refString="Villa, A., Blain, H. - A., van den Hoek Ostende, L. W., and Delfino, M. 2018 c. Fossil amphibians and reptiles from Tegelen (Province of Limburg) and the early Pleistocene palaeoclimate of The Netherlands. Quaternary Science Reviews, 187: 203 - 219. https: // doi. org / 10.1016 / j. quascirev. 2018.03.020" type="journal article" year="2018">Villa et al., 2018c</bibRefCitation>
) indicate a humid subtropical climate, but with MAT and MAP significantly lower than those at Hambach during the Middle Miocene (quantitative climate reconstructions are not available for the Upper Pliocene Hambach sites). A light degree of aridity was also suggested by the same reconstruction. No evidence of a similar dryness is available for the Upper Pliocene level in Hambach. Increased aridization was proposed as a possible cause for the disappearance of some amphibians (i.e., palaeobatrachids;
<bibRefCitation id="EFCDEA82D07EFFA2FE799D54FD0E7373" author="Wuttke, M. &amp; Prikryl, T. &amp; Ratnikov, V. Yu. &amp; Dvorak, Z. &amp; Rocek, Z." box="[479,720,1382,1407]" pageId="42" pageNumber="43" pagination="367 - 395" refId="ref40324" refString="Wuttke, M., Prikryl, T., Ratnikov, V. Yu., Dvorak, Z., and Rocek, Z. 2012. Generic diversity and distributional dynamics of the Palaeobatrachidae (Amphibia: Anura). Palaeobiodiversity and Palaeoenvironments, 92: 367 - 395. https: // doi. org / 10.1007 / s 12549 - 012 - 0071 - y" type="journal article" year="2012">Wuttke et al., 2012</bibRefCitation>
) from Western Europe in the late Neogene, but it is evident that areas suitable for these animals persisted at least in the northwestern part of the continent (i.e., in the Lower Rhine Embayment and the Rhine-Meuse delta system) up to the Late Pliocene and, maybe to a lower extent, the Early Pleistocene. High humidity may have allowed the Lower Rhine Embayment to act as a refuge for amphibians during this time, while they were disappearing from other parts of Western Europe. Later, a combination of increasing aridity and potentially the onset of the Quaternary glaciation may have led to the ultimate loss of the refugial conditions and to a faunistic change towards the early Quaternary and subsequent modern batrachofauna of the area. As a matter of fact, various evidence points out to different trends shown by temperatures and precipitations in the Lower Rhine Embayment during the late Neogene, with MATs showing a clear decreasing trend after the early Middle Miocene and MAPs maintaining high values (&gt;
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) well into the Pliocene (
<bibRefCitation id="EFCDEA82D07EFFA2FC669954FB177773" author="Utescher, T. &amp; Mosbrugger, V. &amp; Ashraf, A. R." box="[960,1225,358,383]" pageId="42" pageNumber="43" pagination="430 - 449" refId="ref38493" refString="Utescher, T., Mosbrugger, V., and Ashraf, A. R. 2000. Terrestrial climate evolution in northwest Germany over the last 25 million years. Palaios, 15: 430 - 449. https: // doi. org / 10.2307 / 3515514" type="journal article" year="2000">Utescher et al., 2000</bibRefCitation>
,
<bibRefCitation id="EFCDEA82D07EFFA2FB7E9954FAC97773" author="Utescher, T. &amp; Mosbrugger, V. &amp; Ivanov, D. &amp; Dilcher, D. L." box="[1240,1303,358,383]" pageId="42" pageNumber="43" pagination="544 - 552" refId="ref38605" refString="Utescher, T., Mosbrugger, V., Ivanov, D., and Dilcher, D. L. 2009. Present-day climatic equivalents of European Cenozoic climates. Earth and Planetary Science Letters, 284: 544 - 552. https: // doi. org / 10.1016 / j. epsl. 2009.05.021" type="journal article" year="2009">2009</bibRefCitation>
,
<bibRefCitation id="EFCDEA82D07EFFA2FA819954FAB87773" author="Utescher, T. &amp; Ashraf, A. R. &amp; Dreist, A. &amp; Dybkjaer, K. &amp; Mosbrugger, V." box="[1319,1382,358,383]" pageId="42" pageNumber="43" pagination="289 - 314" refId="ref38663" refString="Utescher, T., Ashraf, A. R., Dreist, A., Dybkjaer, K., and Mosbrugger, V. 2012. Variability of Neogene continental climates in northwest Europe - A detailed study based on microfloras. Turkish Journal of Earth Sciences, 21: 289 - 314. https: // doi. org / 10.3906 / yer- 1005 - 3" type="journal article" year="2012">2012</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D07EFFA2FAD39954FC087793" author="van Dam, J. A." pageId="42" pageNumber="43" pagination="190 - 218" refId="ref38733" refString="van Dam, J. A. 2006. Geographic and temporal patterns in the late Neogene (12 - 3 Ma) aridification of Europe: The use of small mammals as paleoprecipitation proxies. Palaeogeography, Palaeoclimatology, Palaeoecology, 238: 190 - 218. https: // doi. org (10.1016 / j. palaeo. 2006.03.025" type="journal article" year="2006">van Dam, 2006</bibRefCitation>
;
<bibRefCitation id="EFCDEA82D07EFFA2FC4C99B4FA8B7793" author="Crampton-Flood, E. D. &amp; Peterse, F. &amp; Munsterman, D. &amp; Sinninghe Damste, J. D." box="[1002,1365,390,416]" pageId="42" pageNumber="43" pagination="193 - 205" refId="ref31294" refString="Crampton-Flood, E. D., Peterse, F., Munsterman, D., and Sinninghe Damste, J. D. 2018. Using tetraether lipids archived in North Sea Basin sediments to extract North Western European Pliocene continental air temperatures. Earth and Planetary Science Letters, 490: 193 - 205. https: // doi. org / 10.1016 / j. epsl. 2018.03.030" type="journal article" year="2018">Crampton-Flood et al., 2018</bibRefCitation>
).
<bibRefCitation id="EFCDEA82D07EFFA2FAC899B4FB9E77B3" author="Utescher, T. &amp; Ashraf, A. R. &amp; Dreist, A. &amp; Dybkjaer, K. &amp; Mosbrugger, V." pageId="42" pageNumber="43" pagination="289 - 314" refId="ref38663" refString="Utescher, T., Ashraf, A. R., Dreist, A., Dybkjaer, K., and Mosbrugger, V. 2012. Variability of Neogene continental climates in northwest Europe - A detailed study based on microfloras. Turkish Journal of Earth Sciences, 21: 289 - 314. https: // doi. org / 10.3906 / yer- 1005 - 3" type="journal article" year="2012">Utescher et al. (2012)</bibRefCitation>
further observed that cool events in the Zanclean of northwestern Europe are related to, or start with, wetter conditions, in contrast with the correlation of warmer and wetter periods in the Miocene. This may have favoured the survival of amphibians in this area. In a somehow similar way, a persistent humid climate allowed the Italian Peninsula to act as a preferential refuge for amphibians during the Quaternary glacial cycles, versus the role of reptile refuges that was played by the more arid Balkan and Iberian peninsulae (
<bibRefCitation id="EFCDEA82D07EFFA2FCE89B34FB887513" author="Macaluso, L. &amp; Villa, A. &amp; Carnevale, G. &amp; Delfino, M." box="[846,1110,774,799]" pageId="42" pageNumber="43" pagination="22139" refId="ref34264" refString="Macaluso, L., Villa, A., Carnevale, G., and Delfino, M. 2021. Past, present, and future climate space of the only endemic vertebrate genus of the Italian peninsula. Scientific Reports, 11: 22139. https: // doi. org / 10.1038 / s 41598 - 021 - 01492 - z" type="journal article" year="2021">Macaluso et al., 2021</bibRefCitation>
,
<bibRefCitation id="EFCDEA82D07EFFA2FBC59B34FB6D7513" author="Macaluso, L. &amp; Bertini, A. &amp; Carnevale, G. &amp; Eronen, J. T. &amp; Martinetto, E. &amp; Saarinen, J. &amp; Villa, A. &amp; Capasso, F. &amp; Delfino, M." box="[1123,1203,774,799]" pageId="42" pageNumber="43" pagination="111699" refId="ref34475" refString="Macaluso, L., Bertini, A., Carnevale, G. Eronen, J. T., Martinetto, E., Saarinen, J., Villa, A., Capasso, F., and Delfino, M. 2023 a. A combined palaeomodelling approach reveals the selective role as refugia of the Mediterranean peninsulas. Palaeogeography, Palaeoclimatology, Palaeoecology, 625: 111699. https: // doi. org / 10.1016 / j. palaeo. 2023.111699" type="journal article" year="2023">2023a</bibRefCitation>
). A comparable role may be advocated for the Lower Rhine Embayment as well during the late Neogene and maybe the Early Pleistocene.
</paragraph>
</subSection>
</treatment>
</document>