identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
E86287BFFFEEFFF9BA71A76EFB06FC04.text	E86287BFFFEEFFF9BA71A76EFB06FC04.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Urodela Dumeril 1805	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Urodela Duméril, 1805</p>
            <p>Salamandroidea Goldfuss, 1820</p>
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	https://treatment.plazi.org/id/E86287BFFFEEFFF9BA71A76EFB06FC04	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Georgalis, Georgios L.;Villa, Andrea;Ivanov, Martin;Delfino, Massimo	Georgalis, Georgios L., Villa, Andrea, Ivanov, Martin, Delfino, Massimo (2024): New diverse amphibian and reptile assemblages from the late Neogene of northern Greece provide novel insights into the emergence of extant herpetofaunas of the southern Balkans. Swiss Journal of Palaeontology (34) 143 (1): 1-91, DOI: 10.1186/s13358-024-00332-7, URL: https://doi.org/10.1186/s13358-024-00332-7
E86287BFFFEBFFFCBA71A30EFBD1F8E4.text	E86287BFFFEBFFFCBA71A30EFBD1F8E4.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Anura Dumeril 1805	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Anura Duméril, 1805</p>
            <p>Mediogyrinia Lataste, 1878</p>
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	https://treatment.plazi.org/id/E86287BFFFEBFFFCBA71A30EFBD1F8E4	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Georgalis, Georgios L.;Villa, Andrea;Ivanov, Martin;Delfino, Massimo	Georgalis, Georgios L., Villa, Andrea, Ivanov, Martin, Delfino, Massimo (2024): New diverse amphibian and reptile assemblages from the late Neogene of northern Greece provide novel insights into the emergence of extant herpetofaunas of the southern Balkans. Swiss Journal of Palaeontology (34) 143 (1): 1-91, DOI: 10.1186/s13358-024-00332-7, URL: https://doi.org/10.1186/s13358-024-00332-7
E86287BFFFE5FFF4B99EA1CFFAC1FB24.text	E86287BFFFE5FFF4B99EA1CFFAC1FB24.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Latonia ragei Hossini 1993	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Latonia cf. ragei</p>
            <p>Figures 7, 8, 9, 10, 11, 12</p>
            <p>Material. Spilia 1: one frontoparietal (UU SP1 1022), four maxillae (UU SP1 1018, UU SP1 1024, UU SP1 1044, and UU SP1 1046), two atlantes (UU SP1 1028), three trunk vertebrae [UU SP1 1021 (two elements) and UU SP1 1027], one urostyle (UU SP1 1134), and five ilia (UU SP1 1017, UU SP1 1026, UU SP1 1110, UU SP1 1133, and UU SP1 1141); Spilia 3: one frontoparietal (UU SP3 624), two maxillae (UU SP3 639 and UU SP3 672), four angulars (UU SP3 629– UU SP3 631, and UU SP3 680), one trunk vertebra (UU SP3 628), one sacral vertebra (UU SP3 637), one scapula (UU SP3 632), two humeri (UU SP3 636 and UU SP3 641), and seven ilia (UU SP3</p>
            <p>621– UU SP3 623, UU SP3 625– UU SP3 627, and UU SP3 638); Spilia 4: five maxillae (UU SP4 523, UU SP3 524, UU SP4 693, and UU SP4 737), four angulars (UU SP4 715 and UU SP4 728), one atlas (UU SP4 525), one trunk vertebra (UU SP4 712), four sacral vertebrae [UU SP4 529 and UU SP4 530 (three elements)], two urostyles (UU SP4 520 and UU SP4 521), two ribs (UU SP4 714), four humeri [UU SP4 517, UU SP4 518, and UU SP4 519 (two elements)], and seven ilia (UU SP4 513, UU SP4 526, UU SP4 528, UU SP4 531, UU SP4 532, and UU SP4 716).</p>
            <p>Description.</p>
            <p>UU SP1 1022 is a rather large element that most likely represents the anteromedial part of the frontoparietal</p>
            <p>(Fig. 7a, b). Te ventral surface is smooth, whereas the dorsal one is almost completely covered by a dense dermal sculpturing composed by tubercles, which fuses to form longitudinal ridges anteriorly. Te bone sections exposed along the broken margins clearly show that the ornamentation was secondarily connected to the frontoparietal table and not directly ossified on it. UU SP3 624 is a fragment of a moderately large frontoparietal, of which only the left anterolateral portion is preserved</p>
            <p>(Fig. 9a, b). Te ventral surface shows part of the incrassatio frontoparietalis, which is long and reaches the anterior end. Te lateral margin of the incrassatio is marked by a low and wavy ridge. Lateral to the incrassatio, there is a narrow pars contacta, the lateral portion of which bends ventrally. Anteriorly, the pars contacta develops an anterior horn, provided with a strongly striated ventral surface for the contact with the sphenethmoid. Te dorsal surface of the fragment of frontoparietal bears a well-developed dermal sculpturing. Te only unsculptured part is the anterior horn. Te sculpturing is made up by a moderately dense accumulation of small tubercles. Anteriorly, the tubercles are joined together to form longitudinal ridges. Laterally, the tectum supraorbitale is present, but poorly developed, at least in the preserved portion of the bone.</p>
            <p>Several fragments of maxillae are represented in the Spilia localities (Figs. 7c–f, 9c–e, 11). Te large UU SP3 639 (Fig. 9c–e) and the moderately large UU SP1 1018</p>
            <p>(Fig. 7c–e), UU SP1 1024, and UU SP4 523 (Fig. 11a, b) are the largest ones, whereas the others are slightly smaller. Medially, the maxillae display a mediolaterally short lamina horizontalis, which is rather high in medial view. Ventrally to the lamina, a high number of closely-spaced tooth positions is present. When preserved, the teeth are clearly pleurodont and cylindrical. When the posterior end is preserved [e.g., UU SP4 523 (Fig. 11a, b) and UU SP4 693 (Fig. 11d)], the tooth row extends slightly posteriorly to the lamina horizontalis. A well-developed processus pterygoideus is preserved only in few specimens (e.g., UU SP1 1044, UU SP3 639, and UU SP4 693), but it was probably present at least in other specimens where the posterior end is not missing also judging from the visible medially-directed curvature of the posterior end of the lamina horizontalis. Te posterior depression is present, but not marked by ridges. Te margo orbitalis is strongly concave. Te lateral surface of the maxillae is smooth. Only UU SP4 523 (Fig. 11b) displays very few and small rugosities towards the posterodorsal corner (i.e., the processus zygomaticomaxillaris). It has to be noted that the other largest specimens, UU SP1 1018, UU SP1 1024, and UU SP3 639, show no sculpturing.</p>
            <p>Te angulars from Spilia 3 (Fig. 9f, g) and Spilia 4 are medium to moderately large sized. None of them is completely preserved. Tey carry both a processus coronoideus and a processus paracoronoideus. Te former is distinctly dorsally directed and, when completely preserved, not extended posteriorly. Te lateral surface is characterized by a depressed area marked ventrally by a sharp crista mandibulae externa.</p>
            <p>Te atlantes from the Spilia localities are not complete and they preserve different portions of this element. UU SP4 525 preserves only the dorsal portion of the neural arch (Fig. 12a, b). It is moderately large and displays a long and robust posterior point. Te dorsal surface is smooth, but the remnants of a poorly preserved, low longitudinal ridge are recognisable in the middle. Te other atlantes preserved only the centrum and are smaller. Tey bear two anterior cotyles, which are somehow reniform in anterior view, dorsally inclined and distinctly separated medially by a spatium interglenoidale. Posteriorly, a subcircular cotyle is present. A distinct, variably developed longitudinal ridge stands out on the ventral surface of the centrum.</p>
            <p>Te trunk vertebrae from the Spilia localities are of medium size (Fig. 10j). Only the opisthocoelous centrum is preserved. It is subcylindrical, with circular anterior condyle and posterior cotyle. Te anterior condyle is followed by a poorly distinct neck. In lateral view, the centrum is distinctly ventrally concave.</p>
            <p>Te sacral vertebrae from Spilia are also medium-sized and preserving the centrum alone (Figs. 10a, b, 12c). Tey have an anterior condyle, which is elliptical, and two posterior condyles, which are elliptical as well and well separated. Te ventral surface of the sacrals is not distinctly concave in lateral view as it is in the trunk vertebrae.</p>
            <p>Te urostyles from Spilia are medium to moderately large sized and rather well preserved (Figs. 8a, 12d, e). Anteriorly, they bear two wide and roughly suboval cotyles, which are slightly mediolaterally extended and clearly separated medially. Posteriorly, the urostyle has a moderately high crista dorsalis, which is not closed dorsally for its whole length in UU SP4 521 and closed only by the anterior margin in UU SP4 520 (Fig. 12d, e). Two transverse processes are present by the anterior end of the crista. Teir distal ends are missing, but a narrowing is clearly visible. Only the right process in UU SP4 521 and the left one in UU SP4 520 (Fig. 12d, e) extend slightly posteriorly to form a short lamina, but this ends not far from the process itself. Te canalis coccygeus is wide and ogival in anterior view.</p>
            <p>Te ribs from Spilia are medium sized and moderately robust. Tey are constricted at mid-length and present a distinct process on the dorsal surface.</p>
            <p>Te single scapula (UU SP3 632) from Spilia 3 is large and fragmentary, preserving the pars suprascapularis alone (Fig. 10c). Te latter is very wide. A very well-developed crista anterior is present.</p>
            <p>Te humeri from Spilia (Fig. 12f–h) reach a rather large size (but a very small one is also present: UU SP3 636) and preserve only their distal half. Te shaft is moderately robust and straight. Te spherical eminentia capitata is shifted laterally. Te fossa cubitalis ventralis is present, but moderately shallow (somehow slightly deeper in the very small UU SP3 636). Te epicondylus ulnaris is large, whereas the radialis one is small. Cristae medialis and lateralis follow the same proportions, with the former distinctly more developed than the other. Te olecranon scar is well distinct and elongated.</p>
            <p>Te ilia from Spilia (Figs. 8b–e, 10d–i) are medium-sized, with only UU SP3 622 (Fig. 10f), UU SP3 623, UU SP4 513, UU SP4 531, and UU SP4 716 being smaller and UU SP1 1017 (Fig. 8b, c), UU SP1 1133, and UU SP3 638</p>
            <p>(Fig. 10h, i) being larger. None of them is completely preserved. Tey have a dorsal crest, which is well preserved only in UU SP3 638 (Fig. 10h, i) and UU SP4 528. In these specimens, the crest is visibly straight (i.e., not medially bending). In the smallest specimens, the posterior end of the dorsal crest merges with a laminar dorsal prominence, not clearly presenting a developed dorsal protuberance in this position. Larger specimens, on the other hand, display a slightly more evident protuberance. Te latter is elongated and presents a gently-curving dorsal margin. UU SP3 638 (Fig. 10h, i) and UU SP4 528 clearly shows that the dorsal prominence is not exceeded by the crista dorsalis in height. A distinct tubercular fossa (fossula tuberis superioris sensu Roček, 1994) is not clearly visible in any of the specimens, but few small foramina are present where it should be located. Te acetabulum is large and very deep, with a very strongly developed anterior margin. Anterior to the acetabulum, a distinct preacetabular fossa is present in UU SP3 622 (Fig. 10f), UU SP3 623, UU SP3 625 (Fig. 10g), UU SP3 627, UU SP4 513, UU SP4 526, and UU SP4 531, but clearly not in UU SP1 1141 (Fig. 8e) and UU SP4 528. A shallow supraacetabular fossa is visible dorsally in UU SP1 1141 (Fig. 8e), UU SP3 622, and UU SP4 513 and very poorly in UU SP3 638 (Fig. 10h, i), UU SP4 531, and UU SP4 532, but the same area is not preserved in the other specimens. Te base of the dorsal acetabular expansion is preserved only in UU SP3 627, UU SP3 638 (Fig. 10h, i), UU SP4 513, UU SP4 526, and UU SP4 531, which show apparently that the angle between the latter and the dorsal prominence is very wide (&gt; 90°). Te ventral acetabular expansion is regularly missing. On the medial side, a wide and deep interiliac groove (sensu Bailon, 1999) is present, but due to breakage an interiliac tubercle is clearly visible only in UU SP1 1141 (Fig. 8e) and UU SP3 627.</p>
            <p> Remarks.  Latonia is well represented in the Spilia localities. Among the diagnostic features reported for this genus by e.g., Roček (1994, 2013) and Syromyatnikova et. al. (2019b), we can mention the double processes on the angular and the secondary sculpturing on dermal bones as particularly significant. Compared to the five currently recognized extant and extinct species of the genus (Biton et al., 2013, 2016; Roček, 1994, 2013; Syromyatnikova &amp; Roček, 2019; Syromyatnikova et al., 2019b), the Spilia  Latonia differs from  Latonia nigriventer (Mendelssohn &amp; Steinitz, 1943) because of the highly sculptured frontoparietal, from  Latonia seyfriedi Meyer, 1843 (including  Latonia gigantea [Lartet, 1851]; Syromyatnikova et al., 2019b) in the smooth maxilla, from  Latonia vertaizoni (Friant, 1944) in the well-developed coronoid process, and from  Latonia caucasica Syromyatnikova &amp; Roček, 2019 , in the tooth row slightly extending posteriorly. A certain similarity in the significant features reported by Syromyatnikova and Roček (2019) is present between the  Latonia from Spilia and  Latonia ragei , at least as far as the elements known for both are concerned. Tus, we tentatively refer these fossils to the latter species. Te light rugosities present on the lateral surface of a single maxilla, UU SP4 523, which is otherwise undistinguishable from the other maxillae, are a peculiar feature, worth being highlighted here. However, the taxonomic significance of these rugosities, which do not represent clearly a dermal sculpturing, cannot be clearly evaluated here in the context of  Latonia species with smooth maxillae. </p>
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	https://treatment.plazi.org/id/E86287BFFFE5FFF4B99EA1CFFAC1FB24	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Georgalis, Georgios L.;Villa, Andrea;Ivanov, Martin;Delfino, Massimo	Georgalis, Georgios L., Villa, Andrea, Ivanov, Martin, Delfino, Massimo (2024): New diverse amphibian and reptile assemblages from the late Neogene of northern Greece provide novel insights into the emergence of extant herpetofaunas of the southern Balkans. Swiss Journal of Palaeontology (34) 143 (1): 1-91, DOI: 10.1186/s13358-024-00332-7, URL: https://doi.org/10.1186/s13358-024-00332-7
E86287BFFFFFFFECBA71A10EFEB3FD84.text	E86287BFFFFFFFECBA71A10EFEB3FD84.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Pelobates praefuscus Khosatzky 1985	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Pelobates praefuscus Khosatzky, 1985 Pelobates aff. praefuscus</p>
            <p>Figures 16, 17</p>
            <p>Material. Spilia 1: two humeri (UU SP1 1144); Spilia 3: four maxillae (UU SP3 649– UU SP3 652), one frontoparietal (UU SP3 658), three sacro-urostylar complexes (UU SP3 653– UU SP3 655), six humeri (UU SP3 618, UU SP3 633, and UU SP3 642– UU SP3 645), and three ilia (UU SP3 646– UU SP3 648); Spilia 4: eight maxillae (UU SP4 669– UU SP4 674 and UU SP4 736), one nasal (UU SP4 675), two frontoparietals (UU SP4 668 and UU SP4 691), one sacro-urostylar complex (UU SP4 676), and one humerus (UU SP4 711).</p>
            <p>Description.</p>
            <p>Tese small maxillae from Spilia are only partially preserved (Fig. 16a–f). Most of them are represented by a fragment of the mid-posterior portion of the bone, but UU SP3 651, UU SP3 652, and UU SP4 670 (Fig. 16e, f) preserve the area of the processus palatinus and UU SP3 649 (Fig. 16a, b) preserves most of the bone except for the anterior end and the tooth row. Te maxillae display a rather narrow (higher anteriorly) lamina horizontalis, which does not originate a processus pterygoideus posteriorly. Te tooth row carries closely-spaced and narrow tooth positions. When the teeth are preserved, these are pleurodont, pedicellated and bicuspid. Te tooth row ends in correspondence with the end of the lamina horizontalis in UU SP4 672, but it extends posterior to it in UU SP3 650 (Fig. 16c) and UU SP4 671. Te processus palatinus of UU SP3 649 (Fig. 16a, b), UU SP3 651, UU SP3 652, and UU SP4 670 (Fig. 16e, f), though highly damaged, is visibly narrow, anterodorsally projected and deeply grooved dorsally. Te margo orbitalis is gently concave. A distinct dermal sculpturing is present on the lateral surface. Te sculpturing is rather sparse [very slightly denser in UU SP4 670 (Fig. 16f) and UU SP4 671] and made up by small tubercles and short ridges.</p>
            <p>Te nasal (UU SP4 675) from Spilia 4 is of medium size and wide (Fig. 16g, h). Te dorsal surface bears a rather sparse dermal sculpturing made up by tubercles and short ridges.</p>
            <p>UU SP3 658 (Fig. 16i, j) and UU SP4 668 (Fig. 16k, l) are fragments of frontoparietalis, preserving approximately the left and right posterolateral quarter of the bone respectively. Te bone they represent is rather robustly built. On the ventral surface, the posterolateral corner of the incrassatio frontoparietalis is visible: it has a straight lateral margin (slightly sigmoid in the anterior portion in UU SP3 658), whereas the posterior margin has an anteromedial course. Lateral to the incrassatio, there is the rather shallow canalis arteriae orbitonasalis, starting from a wide foramen that is visible in ventral view. Te posterior portion of the canalis is open ventrally. A short, wide and distally rounded processus paraoccipitalis is present at the posterolateral corner of the bone. A low and dorsally flat ridge runs longitudinally on the dorsal surface of the latter process. Medial to the base of the same process, the rather wide foramen arteriae occipitalis is not exposed in dorsal view, being visible only in posterior view. UU SP4 691 is also a fragment of frontoparietal, but it preserves the middle portion of the left side of the bone. Te canalis arteriae occipitalis is visible, as well as the lateral margin of the incrassatio frontoparietalis, which appears slightly sigmoid in this portion of the bone. Te canalis opens ventrally in its posterior part and is moderately deep. Te dorsal surface of all specimens is largely covered by a sparse dermal sculpturing with tubercles and short ridges.</p>
            <p>Te sacrourostylar complexes from Spilia 3 (Fig. 17a, b) and Spilia 4 (Fig. 17c, d) are poorly preserved and medium to moderately small sized. Most of the urostylar component is missing, but the fusion of the sacral vertebra with the urostyle is still recognizable. Te centrum is cylindrical, with an anterior cotyle. Te neural canal is dorsoventrally compressed, reniform in anterior view. Te neural arch is dorsally flattened, devoid of any developed structure on its dorsal surface. Te lateral processes are very anteroposteriorly enlarged, but they are almost completely missing.</p>
            <p>Te humeri from Spilia preserve only the distal half of the bone (Fig. 17e, f). Te shaft is mostly missing and therefore it is not possible to state whether it was straight or curved. Te eminentia capitata is slightly laterally shifted. Te fossa cubitalis ventralis is moderately deep and opened on the lateral side. Te epicondylus ulnaris is almost double in size than the epicondylus radialis. Te cristae medialis and lateralis are not developed. Te olecranon scar is rather large, but somehow elongated.</p>
            <p>Te ilia from Spilia are medium sized (Fig. 17g –j). Tey have no dorsal crest and no dorsal tubercle. Te preacetabular and supraacetabular fossae are also absent. Te shaft is mostly missing in all specimens. Te acetabulum is rather wide and semicircular. Te spiral groove is deep. Te posteromedial surface of the bone displays a low degree of striation.</p>
            <p> Remarks. All these remains show combinations of features typical for  Pelobates (Bailon, 1999; Roček, 2013). Maxillae, nasal, and frontoparietals have a similar dermal sculpturing, which is coossified with the bone in contrast with that of  Latonia (Roček, 2013; Syromyatnikova et al., 2019b); in the latter genus, the sculpturing is a true exostosis and arises separately from dermal bones in outer layers of the dermis, fusing only secondarily with the dermal bones (see Georgalis et al., 2023c). Te humeri have a shifted eminentia capitata and a laterally-open fossa cubitalis ventralis. Te sacrourostylar complexes display an anterior cotyle and strongly-enlarged lateral processes. Tis sacral morphology is present in  Pelodytes Bonaparte, 1838b , as well, but fusion with the urostyle is absent in this taxon (Bailon, 1999). Ilia have no dorsal crest, no dorsal tubercle, a deep spiral groove (Böhme, 2010), and a striated posteromedial surface. Te Spilia  Pelobates differs from all extant species of the genus in having a ventrally open posterior portion of the canalis arteriae orbitonasalis (Syromyatnikova, 2019). Considering extinct occurrences, it closely recalls  Pelobates aff. praefuscus from the Late Miocene (MN 13) of Solnechnodolsk, Russia, in respect to the frontoparietal characters deemed significant by Syromyatnikova (2019). Similar to the taxon from Solnechnodolsk, the one from Spilia has a pustular sculpturing, a shallow canalis arteriae orbitonasalis (even though this may be slightly deeper at least in UU SP4 691), and the foramen arteriae occipitalis that is both located medial to the processus paraoccipitalis and not visible in dorsal view (even though both these latter features are variable in  Pelobates aff. praefuscus from Solnechnodolsk; Syromyatnikova, 2019). It differs from all other Miocene species in the pustular (i.e., not pit-and-ridges) sculpturing of the cranial bones.  Pelobates praefuscus has an extremely deep canalis artieriae orbitonasalis (Syromyatnikova, 2019), in contrast with the Spilia taxon.  Pelobates sp. from the Early Pliocene (MN 14) of Nizhniy Vodyanoi, Russia (Syromyatnikova, 2019) also shares affinities with the Spilia taxon, but only a few of the frontoparietal features are preserved in elements from this Russian locality. Following Syromyatnikova’s (2019) criteria, it appears thus reasonable to identify this Greek pelobatid as  Pelobates aff. praefuscus , in the same way as the Solnechnodolsk occurrence. </p>
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	https://treatment.plazi.org/id/E86287BFFFFFFFECBA71A10EFEB3FD84	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Georgalis, Georgios L.;Villa, Andrea;Ivanov, Martin;Delfino, Massimo	Georgalis, Georgios L., Villa, Andrea, Ivanov, Martin, Delfino, Massimo (2024): New diverse amphibian and reptile assemblages from the late Neogene of northern Greece provide novel insights into the emergence of extant herpetofaunas of the southern Balkans. Swiss Journal of Palaeontology (34) 143 (1): 1-91, DOI: 10.1186/s13358-024-00332-7, URL: https://doi.org/10.1186/s13358-024-00332-7
E86287BFFFFAFFE2B99EA0E9FCACF822.text	E86287BFFFFAFFE2B99EA0E9FCACF822.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rana dalmatina Fitzinger	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Rana dalmatina Fitzinger in Bonaparte, 1838b  Rana cf. dalmatina</p>
            <p>Figure 20</p>
            <p>Material. Spilia 4: one ilium (UU SP4 683).</p>
            <p>Description.</p>
            <p>Te moderately small ilium (UU SP4 683) from Spilia 4 is rather well preserved, even though most of the shaft is missing (Fig. 20). It has a well-developed dorsal crest and a distinct dorsal tubercle. Te tubercle is elongated and shows some low rugosities on its lateral surface. Its dorsalmost point does not exceed the dorsal crest in height. Te dorsal acetabular expansion is moderately long and creates an angle wider than 90° with the tubercle. A small supraacetabular fossa is visible between the tubercle and the dorsal expansion. Te ventral acetabular expansion is rather well developed. Tere is no preacetabular fossa. Te acetabulum is moderately wide. Te medial surface of the body of the ilium does not display any interiliac groove or tubercle. Te bone has a narrow and moderately high ilioischiatic junction.</p>
            <p> he did not mention this directly in the text. Similarities between the tubercle of  R. dalmatina and the irregular one of  R. arvalis are indeed reported, however (Gleed-Owen, 1998, p. 215), maybe involving the morphology of the lateral surface as well. Our personal observations confirm that rugosities are sometimes present on the tubercle of  R. dalmatina (AV, pers. obs.), even though this feature appears to be variable. Variation in the roughness/smoothness also occurs in  R. temporaria (Gleed-Owen, 1998; AV, pers. obs.). Pending a more complete study of the ilial comparative morphology of brown frogs, we here refer UU SP4 683 to  Rana cf. dalmatina , as this species is the one sharing the most similarities with it among the species for which osteological data are available. However, we acknowledge that this identification should only be considered as tentative, awaiting for more information on other species, such as the southern Balkan endemic  Rana graeca Boulenger, 1891 . </p>
            <p> Remarks. Tis ilium from Spilia 4 is assigned to the genus  Rana because of the following features: well-developed dorsal crest; angle between dorsal tubercle and dorsal acetabular expansion higher than 90°; and high and narrow ilioischiatic junction (Bailon, 1999; Gleed-Owen, 1998; Ratnikov, 2001). More than 100 species of  Rana are currently recognized (Dubois et al., 2021) and our knowledge of their comparative osteology is extremely limited. Tus, a real identification of this fossil at specific level is not possible, even though some features are useful to discuss. UU SP4 683 differs from  Rana temporaria Linnaeus, 1758 , in the well-developed ventral acetabular expansion (Bailon, 1999). A dorsal crest that is higher than the dorsal tubercle is a feature found in  Rana dalmatina and  Rana iberica Boulenger, 1879 , but not in  Rana arvalis Nilsson, 1842 ,  Rana pyrenaica Serra-Cobo, 1993 , and  Rana temporaria (Bailon, 1999; Blain &amp; Arribas, 2017). An irregular lateral surface of the dorsal tubercle is reported for  R. arvalis and  R. temporaria by Gleed-Owen (1998). Te irregularities seem to be present also in Gleed-Owen’s drawing of the ilium of  R. dalmatina (Gleed-Owen, 1998, fig. 5.46a), even though </p>
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	https://treatment.plazi.org/id/E86287BFFFFAFFE2B99EA0E9FCACF822	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Georgalis, Georgios L.;Villa, Andrea;Ivanov, Martin;Delfino, Massimo	Georgalis, Georgios L., Villa, Andrea, Ivanov, Martin, Delfino, Massimo (2024): New diverse amphibian and reptile assemblages from the late Neogene of northern Greece provide novel insights into the emergence of extant herpetofaunas of the southern Balkans. Swiss Journal of Palaeontology (34) 143 (1): 1-91, DOI: 10.1186/s13358-024-00332-7, URL: https://doi.org/10.1186/s13358-024-00332-7
E86287BFFFF6FFE6B99EA7AAFEB0FCC4.text	E86287BFFFF6FFE6B99EA7AAFEB0FCC4.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hyla Laurenti 1768	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Hyla gr.  arborea</p>
            <p>Figure 24</p>
            <p>Material. Spilia 1: one humerus (UU SP1 1100) and one ilium (UU SP1 1043); Spilia 3: one humerus (UU SP3 659) and one ilium (UU SP3 660); Spilia 4: one scapula (UU SP4 734) and one humerus (UU SP4 685).</p>
            <p>Description.</p>
            <p>Te poorly preserved scapula (UU SP4 734) from Spilia 4 shows a very slender and elongated body and a cavitas glenoidalis that is easily visible in ventral view.</p>
            <p>Te processus glenoidalis is broken off, but the processus acromialis is rather slender and straight.</p>
            <p>Te humeri from the Spilia localities are very small sized and slenderly built (Fig. 24a–c). Te diaphysis is straight and devoid of crista paraventralis. Te eminentia capitata is clearly laterally displaced compared to the main axis of the diaphysis. Te fossa cubitalis ventralis is very deep. Te cristae lateralis and medialis are moderately developed. On the dorsal surface, the olecranon scar is poorly visible.</p>
            <p>Te small ilia from Spilia are slenderly built (Fig. 24d– g). Tey have no dorsal crest, but a well-developed, globular and laterally slanted dorsal tubercle is present. Te acetabulum is semicircular. Te dorsal acetabular expansion is short. Te ventral acetabular expansion is strongly anteroventrally expanded. Te lateral surface of the body of the bone is flat.</p>
            <p> Remarks. Tese remains are here assigned to  Hyla based on the following combination of characters reported as diagnostic by Bailon (1999): slender and elongated scapula with a ventrally-exposed cavitas glenoidalis and a straight processus acromialis; straight humeri devoid of crista paraventralis and provided with a laterally-displaced eminentia capitata, a deep fossa cubitalis ventralis, and a moderately-developed crista radialis; ilia with a globular and laterally-bending dorsal tubercle, an expanded ventral acetabular expansion, and without dorsal crest. All these characters fit well with the European members of the  H. arborea group (Birbele et al., 2023). </p>
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	https://treatment.plazi.org/id/E86287BFFFF6FFE6B99EA7AAFEB0FCC4	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Georgalis, Georgios L.;Villa, Andrea;Ivanov, Martin;Delfino, Massimo	Georgalis, Georgios L., Villa, Andrea, Ivanov, Martin, Delfino, Massimo (2024): New diverse amphibian and reptile assemblages from the late Neogene of northern Greece provide novel insights into the emergence of extant herpetofaunas of the southern Balkans. Swiss Journal of Palaeontology (34) 143 (1): 1-91, DOI: 10.1186/s13358-024-00332-7, URL: https://doi.org/10.1186/s13358-024-00332-7
E86287BFFFF1FFDBBA71A56EFEE6F884.text	E86287BFFFF1FFDBBA71A56EFEE6F884.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Bufotes Rafinesque 1815	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Bufotes gr.  viridis</p>
            <p>Figure 25</p>
            <p>Material. Spilia 1: one femur (UU SP1 1057); Spilia 3: two ilia (UU SP3 669 and UU SP3 670); Spilia 4: two sacral vertebrae (UU SP4 679 and UU SP4 680).</p>
            <p>Description.</p>
            <p>UU SP4 679 is the best-preserved sacral vertebra from Spilia (Fig. 25a–e). It has a single anterior cotyle, which is elliptical and mediolaterally elongated, and two posterior condyles, subelliptical as well but less elongated. Te walls of the neural arch are robustly built. Dorsally, the neural arch bears a sort of pseudozygosphenal structure anteriorly and a distinct neural crest. Te transverse processes are broken, but a laterally open fossa is present by their bases. A single prezygapophysis is preserved: it is wide and subcircular. UU SP4 680 is less preserved, but it shares the same morphology. Only the pseudozygosphenal structure is not evidently developed in this second specimen.</p>
            <p>Te ilia from Spilia 3 are moderately small (Fig. 25f, i). Tey preserve a straight shaft devoid of dorsal crest and a small portion of the body. Te dorsal tubercle is better preserved in UU SP3 670 (Fig. 25h, i), displaying a rounded shape with small lobes dorsally. A deep preacetabular fossa is present, being more visible in UU SP3 669 (Fig. 25f, g).</p>
            <p>Te femur (UU SP1 1057) from Spilia 1 is rather small and slender (Fig. 25j). It displays a well-developed and sharp crista femoris.</p>
            <p> Remarks. Green toads (  Bufotes viridis complex) are present in the Spilia assemblage, identified through the criteria of Bailon (1999) and Ratnikov (2001): sacral vertebrae with an anterior cotyle, two posterior condyles, a distinct neural crest, and a laterally-open fossa close to each transverse process; ilia without dorsal crest, but provided with a multilobed dorsal tubercle and a deep preacetabular fossa; a femur with an undivided crista femoris. A pseudozygosphene is not commonly reported in green toad sacral vertebrae, but it was observed in at least few fossils from the Quaternary of southern Italy (Pirro Nord; M.D., pers. obs.) referred to this clade. Te taxonomic significance of this structure in these anurans may be worth further scrutiny. However, the structure observed in UU SP4 679 appears asymmetrical at close inspection, and so it may be interpreted as some sort of anomaly. </p>
            <p> Bufonidae indet. </p>
            <p>Figures 26, 27</p>
            <p>Material. Chalicorrema: one sacral vertebra (UU RA 501); Spilia 1: one humerus (UU SP1 1041); Spilia 3: three trunk vertebrae (UU SP3 661– UU SP3 663) and one humerus (UU SP3 667); Spilia 4: one humerus (UU SP4 681).</p>
            <p>Description.</p>
            <p>Te moderately large trunk vertebrae from Spilia 3 are robust (Fig. 26a–e). Te centrum is procoelous, well individualized and dorsoventrally compressed. Te neural arch has robust walls and a flat dorsal surface. A low carina neuralis is present. Te transverse processes are located posterior to the prezygapophyses.</p>
            <p>UU RA 501 from Chalicorrema is the only bufonid element from that locality, representing a small fragment of a sacral vertebra (Fig. 27). In spite of the small size, the vertebra was rather robustly built. Only the left half of the centrum and a small part of the left transverse process are preserved. Te centrum has an anterior cotyle. Posteriorly, there is an elliptical condyle. It can be assumed that another, similar condyle was present also on the missing right half of the centrum. Te two condyles were not coalescent. Te transverse process is too poorly preserved to give any significant morphological information. Te fragment has a total centrum length of 2.2 mm.</p>
            <p>Te humeri from Spilia are medium sized (Fig. 26f–h). Tey have a straight shaft with no crista paraventralis. Te eminentia capitata is slightly shifted laterally. A moderately shallow fossa cubitalis ventralis is present. Te cristae lateralis and medialis are moderately developed. Te distal epiphysis is poorly ossified distally. Te olecranon scar is moderately wide.</p>
            <p> Remarks. Tese fossils from Spilia share the diagnostic combination of features of bufonid anurans as reported by Bailon (1999). It is not possible, however, to identify them at lower taxonomic level, even though they could pertain to green toads as these are the only bufonids currently known from Spilia. Te straight humeral shaft, which is present in males of  Bufo bufo (Linnaeus, 1758) and  Bufotes viridis and absent in  Epidalea calamita (Laurenti, 1768) and females of the preceding taxa according to Bailon (1999), may support this hypothesis. </p>
            <p>Similarly, in spite of its very fragmentary status, the sacral vertebra UU RA 501 from Chalicorrema can still be assigned to a bufonid anuran, based on the overall robustness, the anterior cotyle, and the well-separated posterior condyles (Bailon, 1999). Further identification is not possible, but the size hints towards either a small species or a juvenile of a larger one.</p>
            <p> Anura indet. </p>
            <p>Figures 28, 29, 30</p>
            <p> Material. Chalicorrema: one trunk vertebra (UU RA 508), one humerus (UU RA 520), one radioulna (UU RA 506), one femur (UU RA 502), and two tibiofibulae (UU R 503 and UU RA 507);  Rema Marmara : eight bone fragments (UU RMA 401– UU RMA 405); Spilia 1: one premaxilla (UU SP1 1085), 14 maxillae (UU SP1 1045, UU SP1 1051, UU SP1 1052, UU SP1 1055, UU SP1 1056, UU SP1 1060, UU SP1 1062– UU SP1 1064, UU SP1 1111, and UU SP1 1138), six angulars (UU SP1 1106, UU SP1 1117, UU SP1 1139, and UU SP1 1149), one sculptured bone fragment (UU SP1 1020), two atlantes (UU SP1 1039 and UU SP1 1142), 10 trunk vertebrae (UU SP1 1054, UU SP1 1061, and UU SP1 1151), seven fragments of vertebra (UU SP1 1102, UU SP1 1104, UU SP1 1114, UU SP1 1116, UU SP1 1121, and UU SP1 1128), one sacral vertebra (UU SP1 1040), two urostyles (UU SP1 1137 and UU SP1 1148), one scapula (UU SP1 1124), three coracoids (UU SP1 1025, UU SP1 1118, and UU SP1 1153), 11 humeri (UU SP1 1019, UU SP1 1059, UU SP1 1105, UU SP1 1131, UU SP1 1132, UU SP1 1140, UU SP1 1146, and UU SP1 1147), 16 radioulnae (UU SP1 1050, UU SP1 1099, UU SP1 1103, UU SP1 1113, UU SP1 1122, UU SP1 1130, and UU SP1 1135), 13 ilia (UU SP1 1029, UU SP1 1030, UU SP1 1032, UU SP1 1036, UU SP1 1037, UU SP1 1042, and UU SP1 1123), eight femora (UU SP1 1058, UU SP1 1120, UU SP1 1129, UU SP1 1136, and UU SP1 1152), eight tibiofibulae (UU SP1 1101, UU SP1 1107, UU SP1 1119, UU SP1 1125, and UU SP1 1150), nine phalanges (UU SP1 1047– UU SP1 1049, UU SP1 1053, UU SP1 1112, UU SP1 1126, and UU SP1 1127), and 252 indeterminate elements (UU SP1 1023, UU SP1 1084, UU SP1 1086, UU SP1 1091, and UU SP1 1095); Spilia 2b: one radioulna (UU SP2b 503) and one indeterminate element (UU SP2b 502); Spilia 2d: one maxilla (UU SP2d 501) and one indeterminate element (UU SP2d 502); Spilia 3: one maxilla (UU SP3 671), seven trunk vertebrae (UU SP3 616, UU SP3 656, UU SP3 666, UU SP3 684, and UU SP3 687), three fragments of vertebra (UU SP3 682), one sacral vertebra (UU SP3 619), five urostyles (UU SP3 640, UU SP3 664, UU SP3 665, and UU SP3 686), one scapula (UU SP3 657), two coracoids (UU SP3 690), three humeri (UU SP3 634, UU SP3 635, and UU SP3 689), 25 radioulnae (UU SP3 620 and UU SP3 673), two ilia (UU SP3 688), one ischium (UU SP3 685), one tibiofibula (UU SP3 683), and six indeterminate elements (UU SP3 681 and UU SP3 691); Spilia 4: four premaxillae (UU SP4 738), 22 maxillae (UU SP4 512, UU SP4 689, UU SP4 690, UU SP4 696, and UU SP4 735), eight sculptured fragments (UU SP4 739), four atlantes (UU SP4 698 and UU SP4 731), 33 trunk vertebrae (UU SP4 624 and UU SP4 724), 13 fragments of vertebra (UU SP4 514), two sacral vertebrae (UU SP4 697 and UU SP4 713), four urostyles (UU SP4 </p>
            <p>522 and UU SP4 725), one scapula (UU SP4 700), five coracoids (UU SP4 677, UU SP4 678, and UU SP4 730), four humeri (UU SP4 699 and UU SP4 729), 10 radioulnae (UU SP4 614, UU SP4 694, and UU SP4 695), two ilia (UU SP4 527 and UU SP4 726), three tibiofibulae (UU SP4 727), two phalanges (UU SP4 732 and UU SP4 733), and 95 indeterminate elements (UU SP4 510 and UU SP4 511); Vevi: one premaxilla (UU VE 548), one sacral vertebra (UU VE 570), one humerus (UU VE 569), three radioulnae (UU VE 552), two tibiofibulae (UU VE 539 and UU VE 571), one indeterminate autopodial element (UU VE 572), one phalanx (UU VE 551), and one distal phalanx (UU VE 550).</p>
            <p>Description and remarks.</p>
            <p>  Among the remains from Spilia belonging to indeterminate anurans (Fig. 28), the largest and more robust likely pertain to either  Latonia or a bufonid. A brief description of the different morphological features encountered in the phalanges from Spilia can be provided. On the whole, the phalanges from Spilia are medium-sized, elongated and rather slender (Fig. 28c, d). Tey narrow distinctly toward the distal tip. Te bulb at the tip is either smooth [eight specimens: UU SP1 1047, UU SP1 1048, UU SP1 1049, UU SP1 1126 (Fig. 28d), UU SP1 1127, and UU SP4 732] or rough [two specimens: UU SP1 1053 (Fig. 28c) and UU SP4 733]. Two lobes are slightly recognizable on the tip of UU SP1 1048, UU SP1 1126, and UU SP1 1127. Specimen UU SP1 1112 is too poorly preserved to recognize the shape of its tip. It is currently not clear, however, to what degree these morphologies have a taxonomical significance  . </p>
            <p>Similarly, the indeterminate anuran remains from Vevi</p>
            <p> (Fig. 29) pertain perhaps to one of the taxa identified above from the same locality. Te distal phalanx (UU VE 550), in particular, is small and slender (Fig. 29c). It has an elongated triangular shape, strongly narrowing distally and ending with a globular tip with no evident lobes. Tis is similar to phalanxes of  Latonia figured by Vasilyan (2020, fig. 6), but clear diagnostic features of anuran phalanxes even at supraspecific levels are not known at the moment. We thus refrain from referring this phalanx to the Vevi  Latonia . </p>
            <p> Te remains from Chalicorrema are even more fragmentary (Fig. 30). Tese remains are too poorly preserved to give significant morphological information, allowing precise taxonomic allocation. Similarly, the even fewer remains from  Rema Marmara can be assigned to indeterminate anurans, but they cannot be identified more precisely due to either poor preservation or scarce diagnostic value. </p>
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	https://treatment.plazi.org/id/E86287BFFFF1FFDBBA71A56EFEE6F884	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Georgalis, Georgios L.;Villa, Andrea;Ivanov, Martin;Delfino, Massimo	Georgalis, Georgios L., Villa, Andrea, Ivanov, Martin, Delfino, Massimo (2024): New diverse amphibian and reptile assemblages from the late Neogene of northern Greece provide novel insights into the emergence of extant herpetofaunas of the southern Balkans. Swiss Journal of Palaeontology (34) 143 (1): 1-91, DOI: 10.1186/s13358-024-00332-7, URL: https://doi.org/10.1186/s13358-024-00332-7
E86287BFFFCCFFD9BA71A4EEFE68FE83.text	E86287BFFFCCFFD9BA71A4EEFE68FE83.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cryptodira Cope 1868	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Cryptodira Cope, 1868 (sensu Joyce et al., 2020b) </p>
            <p>Testudinoidea Fitzinger, 1826 (sensu Joyce et al., 2021)</p>
            <p> Emydidae Gray, 1825 (sensu Joyce et al., 2021) </p>
            <p> Emys Duméril, 1805</p>
            <p> Emys orbicularis (Linnaeus, 1758)</p>
            <p> Emys gr.  orbicularis</p>
            <p>Figure 31</p>
            <p>Material. Vevi: a left epiplastron fragment (UU VE 559), a right hyoplastron fragment (UU VE 567), a left hypoplastron fragment (UU VE 560), and a right xiphiplastron fragment (UU VE 561).</p>
            <p>Description.</p>
            <p>Te fragmentary left epiplastron UU VE 559 preserves on the external surface only part of the gular-humeral sulcus that does not correspond to a notch at the lateral edge of the bone (Fig. 31a–d). On the visceral surface, the preserved portion of the gular pad is low and with a width clearly tapering in medial direction.</p>
            <p>Even if only the posterolateral area of the right hyoplastron UU VE 567 is preserved, the axillary buttress is laterally delimited by a relatively smooth area indicating the presence of a ligamental hinge (Fig. 31e–g). Te buttress is ventrally marked by the lateral sulcus of the pectoral shield, which is also present at the posterior margin of the element, very close (about 1 mm) to the hyo-hypoplastral contact, represented by a hinge with few large bony interdigitations.</p>
            <p>UU VE 560 is a small posterolateral portion of a left hypoplastron (Fig. 31h–j), posterior to the abdominalfemoral sulcus that corresponds to the anterior edge of the fragment (only on the dorsal surface of the element there is a very little segment of the sulcus preserved); the dorsal overlap of the femoral shield is wide and slightly broadening posteriorly.</p>
            <p>Te right xiphiplastron UU VE 561 preserves only a small portion of the posterolateral region, not including the medial suture but clearly indicating a rather rounded lateral edge of the xiphiplastron and the absence of a deep anal notch (Fig. 31k–m). Te area covered by the dorsal overlap of the anal shield is rather tall in the anteriormost-preserved portion and tapers gently in posterior direction.</p>
            <p> Remarks. Te most diagnostic element is the fragmentary right hyoplastron UU VE 567 because it shows evidence of a ligamental hinge between the plastron and the carapace as well as between the hyoplastron and the hypoplastron. Among the Neogene European turtles, this morphology univocally characterizes the members of the genus  Emys but does not allow discriminating among the few extinct and extant species so far described (see among others: Fritz et al., 2005, 2011; Hervet, 2000; Młynarski, 1956; Ottonello et al., 2021). Te other elements from Vevi are referred to same taxon based on morphological congruence. </p>
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	https://treatment.plazi.org/id/E86287BFFFCCFFD9BA71A4EEFE68FE83	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Georgalis, Georgios L.;Villa, Andrea;Ivanov, Martin;Delfino, Massimo	Georgalis, Georgios L., Villa, Andrea, Ivanov, Martin, Delfino, Massimo (2024): New diverse amphibian and reptile assemblages from the late Neogene of northern Greece provide novel insights into the emergence of extant herpetofaunas of the southern Balkans. Swiss Journal of Palaeontology (34) 143 (1): 1-91, DOI: 10.1186/s13358-024-00332-7, URL: https://doi.org/10.1186/s13358-024-00332-7
E86287BFFFC8FFDFBA71A54EFBE4FE04.text	E86287BFFFC8FFDFBA71A54EFBE4FE04.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Squamata Oppel 1811	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Squamata Oppel, 1811b</p>
            <p> Scincoidea Oppel, 1811b (sensu Pyron et al., 2013)  Scincidae Oppel, 1811b</p>
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	https://treatment.plazi.org/id/E86287BFFFC8FFDFBA71A54EFBE4FE04	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Georgalis, Georgios L.;Villa, Andrea;Ivanov, Martin;Delfino, Massimo	Georgalis, Georgios L., Villa, Andrea, Ivanov, Martin, Delfino, Massimo (2024): New diverse amphibian and reptile assemblages from the late Neogene of northern Greece provide novel insights into the emergence of extant herpetofaunas of the southern Balkans. Swiss Journal of Palaeontology (34) 143 (1): 1-91, DOI: 10.1186/s13358-024-00332-7, URL: https://doi.org/10.1186/s13358-024-00332-7
E86287BFFFC1FFD5BA71A5EEFDEBFC24.text	E86287BFFFC1FFD5BA71A5EEFDEBFC24.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Agaminae Spix 1825	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Agaminae Spix, 1825</p>
            <p> Agaminae indet. </p>
            <p>Figure 41</p>
            <p>Material. Chalicorrema: one right maxilla (UU RA 403), three dentaries (UU RA 427– UU RA 429), two fragments of indeterminate tooth bearing bones (UU RA 401 and UU RA 402), and five caudal vertebrae (UU RA 404 and UU RA 430– UU RA 433).</p>
            <p>Description.</p>
            <p>UU RA 403 is a small bone fragment, representing the anterior end of the anterior premaxillary process of a right maxilla (Fig. 41a, b). In dorsal view, it has a straight anterior margin. Tere is no anterolateral process. An anteromedial process could, on the other hand, have been present in origin, but an evident breakage in the related area of the specimen hinders a clear evaluation of its possible development. Te dorsal surface of the anterior premaxillary process is strongly concave. Te specimen bears a single, well-preserved first tooth, plus fragments of a second one posteriorly. Te tooth is subpleurodont, straight and pointed. It has no striation and an unswollen base. In spite of being very poorly preserved, the second tooth also has a subpleurodont implantation. Te complete specimen is 2.4 mm long.</p>
            <p>Tree anterior portions of dentaries are present in our material from Chalicorrema (Fig. 41c–i). UU RA 428</p>
            <p> (Fig. 41f, g)  only preserves the anteriormost tip, where the subpleurodont teeth are located, whereas the other two specimens display also part of the acrodont part of the tooth row. In medial view, all specimens present a high subdental table. Te table is not distinctly developed medially in dorsal view, and bears no subdental ridge. Te Meckelian fossa is narrow and opens ventromedially. Anteriorly, the mandibular symphysis is wide. Te ventral margin of the dentaries is straight. Te lateral surface is smooth, with few anteroposteriorly-aligned mental foramina. In both most well-preserved dentaries, at least five foramina can be counted; these increase in width towards the posterior. Te dentition is heterodont, with two subpleurodont teeth followed by acrodont ones. UU RA 427 (Fig. 41c–e) and UU RA 429 (Fig. 41h, i) preserve five and three acrodont tooth positions, respectively. Only the first subpleurodont teeth in UU RA 428 (Fig. 41f, g) and UU RA 429 (Fig. 41h, i) are complete, whereas the second ones in these specimens and both ones in UU RA 427</p>
            <p>(Fig. 41c–e) preserve only their bases. Te most preserved subpleurodont teeth are straight, conical, non-striated, and with a rounded tip. A slightly-swollen base is visible in the first tooth of UU RA 428 (Fig. 41f, g). On both UU RA 427 (Fig. 41c–e) and UU RA 428 (Fig. 41f, g), the first subpleurodont tooth is clearly larger than the second one, whereas it is the opposite in UU RA 429 (Fig. 41h, i). Te acrodont teeth are triangular and closely packed. Ventrally, they extend to cover the underlying bone. Only in UU RA 427, distinct interdental grooves are present between each pair of acrodont teeth on the lateral surface of the bone</p>
            <p>(Fig. 41d). Te most preserved dentary, UU RA 427, is slightly more than 10 mm long (Fig. 41c–e).</p>
            <p>Te two fragments of indeterminate tooth bearing bones from Chalicorrema are too poorly preserved to be assigned to either maxillae or dentaries (Fig. 41j). Tey are rather small. UU RA 402 carries one subpleurodont tooth and one acrodont tooth, whereas UU RA 401 carries two acrodont teeth and a small fragment of a third one. Te subpleurodont tooth is pointed, not swollen, and slightly posteromedially curved by the tip. It has no striae. Te acrodont ones extend on the lingual margin of the related tooth bearing bone. Tey are less preserved in UU RA 401 than in UU RA 402. In the latter specimen, the acrodont teeth are small and triangular, with no distinct accessory cusps. Teeth of UU RA 401 are larger, representing either a more posterior portion of the tooth row or an older individual. Te total length of UU RA 401 is 3.0 mm, whereas it is 2.0 mm in UU RA 402.</p>
            <p>Te caudal vertebrae from Chalicorrema are slenderly built and elongated (Fig. 41k, l). Tey have wide and subelliptical anterior cotyle and posterior condyle. In anterior view, the neural canal is small, being roughly half as high as the cotyle. When preserved, the zygapophyses have subovoid and subvertical facets. Te ventral surface of the centrum is smooth, with no traces of haemapophyses or pedicles for the chevron bone. Tere is no autotomy plane. Te bases of laminar transverse processes are preserved in at least some vertebrae, being located roughly at mid-length. Te longest and most-preserved vertebra is slightly less than 10 mm long.</p>
            <p> Remarks. An agamid is represented in the Chalicorrema assemblage by a few fragmentary fossils. Te main character supporting this identification for the tooth bearing bones is the presence of both (sub)pleurodont and acrodont teeth in the dentition (Augé, 2005; Blain et al., 2014; Delfino et al., 2008; Georgalis et al., 2023a; Moody, 1980; Smith, 2011; Smith et al., 2011; Villa &amp; Delfino, 2019a). Another character that is commonly used to discriminate agamids from chamaeleonids is the extension of the acrodont teeth onto the lingual surface of the tooth bearing bone (Evans et al., 2002; Georgalis et al., 2023a; Villa &amp; Delfino, 2019a). Te validity of the latter distinction was doubted by Rage and Bailon (2011), but there is no evidence indicating the presence of a chamaeleonid in Chalicorrema and thus we here refer the only tooth bearing bone fragment not presenting subpleurodont teeth to the same taxon as the other specimens due to comparable size and overall morphology of the acrodont teeth. Te presence of two subpleurodont teeth is indicative of an agamine identity of this taxon (Maul et al., 2011). Te caudal vertebrae are also attributed to  Agaminae due to shared morphology with comparative specimens at our disposal (in i.e., overall elongated shape, subelliptical anterior cotyle, subvertical prezygapophyses). Te poor preservation of the remains does not allow a more precise identification. Smith et. al. (2016) reported sexual dimorphism in the size of subpleurodont teeth in the agamid  Stellagama stellio (Linnaeus, 1758) , with the second tooth being larger than the first one in males. Even though identification of the Chalicorrema material is limited to the suprageneric rank, a similar variation in tooth size is observed in the three dentaries available, possibly suggesting a comparable dimorphism. </p>
            <p> Agamidae indet. </p>
            <p>Figure 42</p>
            <p>Material. Vevi: one fragment of tooth bearing bone (UU VE 501).</p>
            <p>Description. Te fragment of tooth bearing bone (UU VE 501) from Vevi cannot be clearly assigned to a specific element, likely being either part of a maxilla or a dentary. It bears two acrodont teeth, which are triangular and provided with an incipient accessory cusp on both mesial and distal sides (Fig. 42). A low degree of wearing possibly due to tooth abrasion is visible, thus giving a rounded shape to the tip of both teeth. No distinct striae are visible on the labial and lingual sides. Te medial side of the tooth base expands ventrally to cover the alveolar surface of the tooth bearing bone. Te two teeth are separated by a distinct space. On the lateral side of the tooth bearing bone there are two vertical, deep and wide interdental grooves. Te fragment is 3.8 mm long.</p>
            <p> Remarks. Te bone fragment from Vevi is here referred to an agamid lizard because of the widely-separated acrodont teeth, which expand ventrally on the medial side of the bone (Delfino et al., 2008; Evans et al., 2002; Georgalis et al., 2023a; Villa &amp; Delfino, 2019a; but note possible variability, regarding features also present in chamaeleonids, reported by Rage &amp; Bailon, 2011 and Georgalis et al., 2023a). A more precise determination of this Vevi fragment within  Agamidae is not possible. It could be likely that the Vevi specimen could pertain to  Agaminae , as is the more complete material from Chalicorrema described above. </p>
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	https://treatment.plazi.org/id/E86287BFFFC1FFD5BA71A5EEFDEBFC24	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Georgalis, Georgios L.;Villa, Andrea;Ivanov, Martin;Delfino, Massimo	Georgalis, Georgios L., Villa, Andrea, Ivanov, Martin, Delfino, Massimo (2024): New diverse amphibian and reptile assemblages from the late Neogene of northern Greece provide novel insights into the emergence of extant herpetofaunas of the southern Balkans. Swiss Journal of Palaeontology (34) 143 (1): 1-91, DOI: 10.1186/s13358-024-00332-7, URL: https://doi.org/10.1186/s13358-024-00332-7
E86287BFFFDFFFCEB99EA094FB79FB84.text	E86287BFFFDFFFCEB99EA094FB79FB84.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Varanidae Gray 1827	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Varanidae Gray, 1827 (sensu Estes et al., 1988) </p>
            <p> cf.  Varanidae indet. </p>
            <p>Figure 46</p>
            <p>Material. Spilia 4: one isolated tooth (UU SP4 506).</p>
            <p>Description. Tis isolated tooth from Spilia 4 misses its proximal part (Fig. 46). Te preserved portion is 3.1 mm in length. Te tooth is strongly labiolingually compressed, slightly curved posteriorly, canine-like and pointed. It is smooth, without longitudinal grooves. Te mesial and distal cutting edges are slightly serrated, with the distal one being more serrated than the mesial one.</p>
            <p> Remarks. Te single and partial tooth from Spilia 4 is tentatively attributed to an indeterminate monitor lizard because of the overall similarity between it and the general morphology of varanid teeth: labiolingual compression that encompasses the pulp cavity, presence of serrated carinae, posterior bending (Bhullar &amp; Smith, 2008; Bullet, 1942; Georgalis et al., 2023b; Ivanov et al., 2018; Peyer, 1929; Villa et al., 2018). Nevertheless, the attribution cannot be confidently confirmed because of the missing tooth base, the absence of which hinders the recognition of the main apomorphy of varanoid teeth (namely, the presence of plicidentine; Estes et al., 1988; Georgalis &amp; Scheyer, 2019; Kearney &amp; Rieppel, 2006). Remarkable is also the absence of grooves in the preserved portion of the basal area of the tooth; such grooves are generally associated with the presence of the plicidentine (see for example Villa et al., 2021, fig. 17a–d). Nevertheless, the labiolingual tooth crown compression, with mesial and distal carinae is considered a synapomorphy of the varanoid genera  Lanthanotus Steindachner, 1878 , and  Varanus , with the latter genus having this compression extending to the tooth base and being visible in the pulp cavity (Bhullar &amp; Smith, 2008; Georgalis et al., 2023b). Most of the tooth base is not preserved in the Spilia tooth, but nevertheless it seems that the pulp cavity is compressed, a feature typical of  Varanus (Georgalis et al., 2023b) . </p>
            <p> (non-snake)  Squamata indet. </p>
            <p>Figure 47</p>
            <p>Material. Chalicorrema: one fragment of indeterminate tooth bearing bone (UU RA 405); Spilia 3: one cervical vertebra (UU SP3 615); Spilia 4: three quadrates (UU SP4 608, UU SP4 612, and UU SP4 613), one indeterminate cranial element (UU SP4 692), one axis (UU SP4 626), three trunk vertebrae (UU SP4 642, UU SP4 644, and UU SP4 710), two sacral vertebrae (UU SP4 703,</p>
            <p>708), three caudal vertebrae (UU SP4 621, UU SP4 628, and UU SP4 656), five ribs (UU SP4 505), two humeri (UU SP4 607 and UU SP4 610), and five femora (UU SP4 609, UU SP4 651, UU SP4 652, and UU SP4 702).</p>
            <p>Description and remarks.</p>
            <p>Te fossil remains from Spilia 3 and Spilia 4 pertain to lizards, but they cannot be confidently identified because of either an overall poor preservation or the current lack of data on the comparative postcranial osteology of these reptiles.</p>
            <p> Te three quadrate fragments from Spilia 4 (UU SP4 608, UU SP4 612, and UU SP4 613) only preserve the pillar, which is narrow and straight and expands dorsally and ventrally (Fig. 47c–k). In all three specimens, a low medial lamina is present but the lateral lamina is broken off . </p>
            <p>Te humerus (UU SP4 610) from Spilia 4 (Fig. 47m), as well as most femora, are very small, but well ossified. Tey could thus pertain to the small-sized lacertid identified based on the above-described coronoid UU SP4 704. Te other humerus (UU SP4 607) is also very small, but its epiphyses are not ossified, thus suggesting it belonged to a juvenile. At least one rather large femur (part of UU SP4 702) is also present, possibly attributable to the medium-sized lacertid.</p>
            <p>Te trunk vertebrae from Spilia 4 are small, with a centrum length ranging from 1.8 to 1.9 mm. Tey are poorly elongated and have a not compressed centrum. Tis morphology is more similar to the one of lacertid vertebrae, among non-snakes squamates from Spilia. Pending a real detailed comparative analysis of the vertebrae of these squamates, however, this possible identification should just remain tentative.</p>
            <p>It is also worth noticing the specimen UU SP4 656</p>
            <p> (Fig. 47l). Tis is a medium-sized caudal vertebra, with a not compressed, procoelous centrum. Te centrum length is about 3.5 mm and both cotyle and condyle are circular. Tere is no precondylar constriction. Te ventral surface of the centrum is distinctly keeled and posteriorly it displays two small articular surfaces for the chevron bone located immediately anterior to the posterior condyle. Te left transverse process is better preserved than the right one, even though the distal half is still missing. Te autotomy plane is not visible. Te neural arch is poorly preserved. A laminar neural spine is present, but broken. Te suboval left prezygapophysis is preserved, with a slight dorsal inclination in anterior view. On the lateral surface of the neural arch, a very low but distinct postzygoprezygapophyseal lamina is visible. Tis morphology fits well with caudals of lacertids, thus suggesting that this vertebra could pertain to the medium-sized one (cf.  Lacerta sp. ) identified in Spilia based on cranial bones. UU SP4 621 and UU SP4 628 are smaller (centrum length around 3 mm and 2.5 mm respectively) and less preserved, but the morphology is more or less the same. Tey could pertain to indeterminate lacertids as well. </p>
            <p>As for the single element from Chalicorrema, this specimen (UU RA 405) is a fragment of tooth bearing bone</p>
            <p>(Fig. 47a, b). It is either part of a maxilla or a dentary, but this cannot be clearly stated. It carries seven tooth positions, two of which are still occupied by moderately preserved teeth. Te teeth are pleurodont, cylindrical, slender and closely spaced. Te tooth base is not swollen. Both teeth have a poorly preserved crown, thus hindering a clear recognition of the morphology. Te crown of the (probably) most anterior tooth appears rounded, but this is due to breakage and wearing. Te other tooth seems to display a small anterior cusp mesially, but, at a closer inspection, this comes out as an artifact due to breakage. Te fragment is 4.0 mm long. According to this description, it is obvious that a third lizard taxon, a small lizard with a pleurodont dentition, is also present in the Chalicorrema fossil assemblage, but its taxonomy cannot be confidently stated based on this single and very poorly preserved specimen. Considering what is preserved of its tooth morphology and the fossil record of non-snake squamates in the Neogene of Europe, teeth of this element appear more similar to those of lacertid and scincoid lizards, and maybe to a lesser degree, gekkotans. However, a clear attribution is not possible for the moment, and we only refer this fossil to an indeterminate lizard, different from the above-described agamine and anguine from that locality.</p>
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	https://treatment.plazi.org/id/E86287BFFFDFFFCEB99EA094FB79FB84	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Georgalis, Georgios L.;Villa, Andrea;Ivanov, Martin;Delfino, Massimo	Georgalis, Georgios L., Villa, Andrea, Ivanov, Martin, Delfino, Massimo (2024): New diverse amphibian and reptile assemblages from the late Neogene of northern Greece provide novel insights into the emergence of extant herpetofaunas of the southern Balkans. Swiss Journal of Palaeontology (34) 143 (1): 1-91, DOI: 10.1186/s13358-024-00332-7, URL: https://doi.org/10.1186/s13358-024-00332-7
E86287BFFFDBFFC7BA71A4EEFA89FB64.text	E86287BFFFDBFFC7BA71A4EEFA89FB64.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Natrix rudabanyaensis Szyndlar 2005	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Natrix rudabanyaensis Szyndlar, 2005 Natrix aff. rudabanyaensis</p>
            <p>Figures 49, 50, 51, 52</p>
            <p>Material: Spilia 0: 12 trunk vertebrae (UU SP0 153– UU SP0 164) and one caudal vertebra (UU SP0 165); Spilia 1: one caudal vertebra (UU SP1 1012); Spilia 4: 19 trunk vertebrae [UU SP4 565, UU SP4 566, and UU SP4 568 (17 vertebrae)]; Vevi: Four anterior trunk vertebrae (UU VE 603– UU VE 606), 10 mid-trunk vertebrae (UU VE 607– UU VE 616), three posterior trunk vertebrae (UU VE 617– UU VE 619), and nine caudal vertebrae (UU VE 620– UU VE 628).</p>
            <p>Description.</p>
            <p>Te description of the trunk vertebrae is mainly based on the specimens from Vevi, which are more complete</p>
            <p> (Fig. 49).   Te vertebrae are lightly built. All specimens are at least partially fragmentary with the neural spine broken off close to its base (but see Fig. 51g for a perfectly preserved neural spine of a single specimen from Spilia 0).  In lateral view (Fig. 49a, f, k, n), the anterior margin of the neural spine of the best-preserved mid-trunk vertebra (UU VE 607) extends anteriorly to the level of the posterior margin of the zygosphenal facet.  Te zygosphenal facet is markedly large and rhomboid. Tere is a distinct blunt ridge developed at the posterior margin of the zygosphenal facet which extends from the base of the anterior neural spine base.  Te well-developed interzygapophyseal ridges are usually sharp.  Te lateral foramen is situated close below the interzygapophyseal ridge.  Te dorsally arched subcentral ridges are prominent.  Te diapophyses are well-separated from the equally sized parapophyses.  Te anteriorly directed parapophyseal processes are moderately long and they are well-separated from the parapophyseal facets.  Te only preserved complete hypapophysis of one specimen (UU VE 609) is sigmoid with blunt distal tip situated slightly anterior to the posterior margin of the condyle </p>
            <p>(Fig. 49k). Te condyle is developed on a rather short condylar neck. In dorsal view (Fig. 49b, g, o), the zygosphene has pointed lateral lobes and a rather wide medial lobe. Te prezygapophyseal articular facets are widely oval; however, prezygapophyseal accessory processes are broken off close to their base. Te epizygapophyseal spines are moderately developed but sometimes they are indistinct as a result of surface abrasion. Te posterior median notch of the neural arch is rather deep. Te diapophyses are directed posterolaterally. In ventral view (Fig. 49c, l, p), the hypapophysis extends anteriorly to form the triangular anterior keel with small subcotylar tubercles developed at the base of the cotylar rim. Teir distal tip is directed posterolaterally. Te converging subcentral ridges are straight and wide subcentral grooves occur between those ridges and narrow (almost sharp) hypapophysis. In posterior trunk vertebrae (UU VE 617; Fig. 49p), the sharp subcentral ridges are markedly developed and the subcentral grooves are rather deep. Subcentral foramina are very small. Te postzygapophyseal articular facets are irregularly shaped. In anterior view (Fig. 49d, i, m), the thin zygosphenal roof is straight with a ventrally bent medial lobe. Te zygosphenal facets are slightly uplifted above the zygosphenal roof. Te neural canal is sub-squared with rather small lateral sinuses. Paracotylar foramina are situated in depressions on either side of the rounded cotylar rims. In posterior view (Fig. 49e, j), the neural arch is moderately vaulted, with a vaulting ratio (sensu Georgalis et al., 2021b) ranging between 0.32 and 0.39. Te zygantrum is wide. Te number of small parazygantral foramina is variable from 1 to 3 on either side. Te small condyle is almost orbicular with its ventral margin slightly depressed.</p>
            <p> Trunk vertebrae from Spilia (Figs. 51, 52a–e) are in most cases more fragmentary than those from Vevi. Nevertheless, they still afford some anatomical observations. In one specimen (UU SP4 565; Fig. 52a–c) from Spilia 4, the hypapophysis is complete, projecting ventrally and its termination does not extend posteriorly from the condyle. Some variation exists among the shape and direction of the neural spine: in one specimen (UU SP4 566; Fig. 52d, e) from Spilia 4, the posterior margin of the neural spine strongly overhangs posteriorly, whereas in a vertebra from Spilia 0 (UU SP0 154; Fig. 51g), the neural spine has its anterior margin straight to slightly anteriorly inclined and its posterior margin posteriorly inclined, and in another vertebra from the latter locality (UU SP0 158; Fig. 51a) it seems that both anterior and posterior margins of the neural spine are almost straight. Te neural arch of the Spilia trunk vertebrae is moderately depressed, with a vaulting ratio (sensu Georgalis et al., 2021b) ranging between 0.34 and 0.39 . </p>
            <p>Te available caudal vertebrae from Vevi (Fig. 50), Spilia 0, and Spilia 1 (Fig. 52f) are fragmentary with broken off pleurapophyses and haemapophyses. In lateral view (Fig. 50d, i), the neural spine is about twice longer than high in more anteriorly located caudal vertebrae whereas in posterior caudal vertebrae the neural spine is about three to four times longer than high. In the single caudal vertebra (UU SP1 1012) from Spilia 1, the neural spine has its anterodorsal margin strongly inclined anteriorly (Fig. 52f). In dorsal view (Fig. 50b, e), the zygosphene has distinct lateral lobes and a wide medial lobe. Prezygapophyseal articular facets are oval with their axis elongated anterolaterally. Prezygapophyseal accessory processes are pointed distally and about half of the length of the prezygapophyseal articular facets or they are shorter. In ventral view (Fig. 50c, f), pleurapophyses of anterior caudal vertebrae have narrow base and the preserved portions of pleurapophyses indicate that they were laterally rather than antero-laterally directed. In posterior caudal vertebrae the pleurapophyses with wide base are directed anteriorly. Tere is a distinct, posteriorly directed spur developed on the posterior margin of the pleurapophysis base.</p>
            <p> Remarks. Te hypapophysis-bearing trunk vertebrae with triangular anterior keel, elongated centrum and well-developed epizygapophyseal spines correspond to a typical natricid morphology (Ivanov, 2002; Szyndlar, 1984, 1991b, 2005; Szyndlar &amp; Schleich, 1993). Te referral to the genus  Natrix is based on the following combination of features in mid-trunk vertebrae: 1, the neural spine is high; 2, the vertebrae with elongated centrum are cylindrical in shape; 3, the prezygapophyseal accessory processes are well developed; 4, the parapophyseal processes are moderately long and directed anteriorly rather than anteroventrally; 5, the hypapophysis is sigmoid with rounded distal tip. Tere are four extinct valid species of  Natrix in the European Neogene:  Natrix merkurensis Ivanov, 2002 (MN 3a–?MN 4; Ivanov, 2002; Rage &amp; Bailon, 2005),  Natrix sansaniensis (Lartet, 1851) (MN 3a–MN 4 and MN 6; Augé &amp; Rage, 2000; Ivanov, 2002; Szyndlar &amp; Schleich, 1993),  Natrix rudabanyaensis (MN 9; Szyndlar, 2005), and  Natrix longivertebrata Szyndlar, 1984 (widespread from the Early Miocene up to the Late Pliocene; Ivanov, 2022; Rage &amp; Szyndlar, 1986; Szyndlar, 1984, 1991b; Vasilyan et al., 2022). A further species,  Neonatrix natricoides Augé &amp; Rage, 2000 , from the Early and Middle Miocene (MN 4 and MN 5) of France (Augé &amp; Rage, 2000; Rage &amp; Bailon, 2005), was considered a member of the genus  Natrix by Szyndlar (2005, 2012); however, its rather short hypapophysis is unusual for  Natrix , and therefore, even generic allocation of that species is uncertain (Ivanov, 2022). Te  Natrix vertebrae from Vevi and Spilia particularly resemble those of  N. rudabanyaensis by the following combination of features (see Szyndlar, 2005): 1, relatively small dimensions; 2, moderately elongated centrum; 3, elongated prezygapophyseal articular facets; 4, prezygapophyseal accessory processes long and moderately flattened dorsoventrally. However, the parapophyseal processes of  N. aff. rudabanyaensis from Vevi and Spilia are shorter and the epizygapophyseal spines are less distinct, compared to those of  N. rudabanyaensis from the Late Miocene of Rudabanya (Szyndlar, 2005). Admittedly, several vertebral features that are used to differentiate Neogene  Natrix spp. are subjected to intracolumnar and intraspecific variation or are anyway widespread among natricids. A proper taxonomic referal of this Greek form can only be made once the taxonomy and diagnosis, and intracolumnar variation of the Neogene European species of  Natrix is better assessed and revised, something that is well beyond the scope of the present study. For these reasons, we only tentatively refer the Greek material as  Natrix aff. rudabanyaensis , as is the case with the material from Maramena that was recently described by Georgalis et. al. (2019b). </p>
            <p> ?  Natricidae indet. </p>
            <p>Figures 53, 54, 55</p>
            <p>Material. Chalicorrema: three trunk vertebrae (UU RA 408– UU RA 410); Spilia 0: two trunk vertebrae (UU SP0 151 and UU SP0 152); Spilia 2b: one trunk vertebra (UU SP2b 501); Vevi: one anterior trunk vertebra (UU VE 601).</p>
            <p>Description. All hypapophysis-bearing trunk vertebrae from Chalicorrema are rather fragmentary (Fig. 53). In the best-preserved anterior trunk vertebra (UU RA 408; Fig. 53a–c), only the centrum with broken off distal tip of the hypapophysis and incomplete neural arch is present. In lateral view (Fig. 53a), the short interzygapophyseal ridge is well developed. A small lateral foramen is situated in a shallow depression close below the interzygapophyseal ridge. Te subcentral ridges are arched dorsally and extend from the base of a non-preserved parapophysis up to the vicinity of the base of the short condylar neck. In dorsal view</p>
            <p> (Fig. 53d),  the only preserved left prezygapophyseal articular facet of one fragment (UU RA 409) is irregularly oval to subtriangular in outline. Te prezygapophyseal accessory processes, broken off at their wide base, were most probably well developed in life. In ventral view (Fig. 53b, e), the subcentral grooves are narrow. Subcentral ridges extend parallel along their entire length. Te hypapophysis extends to the close vicinity of the cotylar rim but moderately developed triangular expansion of its base occurs anterior to minute subcentral foramina. Small subcotylar tubercles are developed close behind the base of the cotylar rim. Te subcentral foramina are situated in the middle of the centrum length, the left one is doubled in UU RA 408 (Fig. 53b) but multiple foramina (three on the right side plus two on the left side) occur in UU RA 409 (Fig. 53e). Te condyle is relatively small. In anterior view (Fig. 53c, f), the neural canal is rounded with wide lateral sinuses. Te paracotylar foramina, situated in narrow depressions on either side of the circular cotylar rim, are either doubled with upper foramen larger than the lower one (UU RA 408; Fig. 53c). However, only one large single left paracotylar foramen is present in one specimen (UU RA 409; Fig. 53f). Te ventral border of the cotylar rim is flat in this specimen (Fig. 53f) . </p>
            <p>Te few vertebrae from Spilia 0 and Spilia 2 are rather fragmentary (Fig. 54). Tey all possess hypapophyses, which are, however, much incomplete in all specimens. Neural spines, paradiapophyses, and parts of the prezygapophyses are much damaged in all specimens.</p>
            <p>Te single specimen from Vevi (UU VE 601) represents an anterior trunk vertebra (Fig. 55). Its right side is damaged with broken off right postzygapophysis, incomplete prezygapophyses, and the right paradiapophysis eroded. In lateral view (Fig. 55a), the neural spine rises at the level of the posterior margin of the zygosphenal facets. Te neural spine is high with its basal anterior margin inclined slightly posteriorly whereas its posterior margin is inclined anteriorly. Te distal termination of the neural a spine is not preserved. Te long axis of the strongly elongated zygosphenal facet is directed anteriorly rather than anterodorsally. Te anteroventrally directed lamina, situated below the interzygapophyseal ridge, forms the dorsal limitation of the wide depression ventrally limited by a straight subcentral ridge. A large lateral foramen is situated close to the dorsal margin of this depression. Te diapophysis is well separated from the parapophysis. Te parapophyseal process is short but its anterior termination is slightly eroded. Te condyle is separated from the centrum by a wide groove. Te anterior keel of the hypapophysis is only moderately inclined in ventral direction but this inclination is well developed in the middle of the centrum length indicating most probably posteroventral inclination of the distal tip of the hypapophysis. In anterior view (Fig. 55d), the neural arch bears no epizygapophyseal spines. Te neural canal is rounded with shallow lateral sinuses. Te zygosphenal lip is slightly arched dorsally with straight medial part. Te prezygapophyses are horizontal with broken off prezygapophyseal accessory processes. Paracotylar foramina are situated in depressions on either side of the cotyle. Te cotyle is rounded with slightly depressed ventral margin. Te only complete right subcotylar tubercle occurs at the base of the cotyle. In posterior view (Fig. 55e), the neural arch is strongly vaulted, with a vaulting ratio (sensu Georgalis et al., 2021b) equal to 0.52. A very large parazygantral foramen is situated near the edge of the postzygapophysis.</p>
            <p> Remarks. All poorly preserved fragments of trunk vertebrae from Chalicorrema possess hypapophyses. Te well-developed subcentral grooves and ridges in the best-preserved vertebra indicate that this vertebra did not belong to the anterior trunk region. Although all hypapophyses are broken off close to their base, the anterior keel of the hypapophysis is triangular (UU RA 408, UU RA 409) with small subcotylar tubercles which frequently occurs in  Natricidae (e.g., Szyndlar &amp; Schleich, 1993; Szyndlar, 1984, 1991b, 2005). Te only preserved left prezygapophysis of UU RA 409 is horizontal in anterior view. Although the anterior keel of the hypapophysis is reminiscent of that of  Natrix aff. rudabanyaensis described above from Spilia and Vevi (see above), even an assignation of the Chalicorrema snakes to natricids is not safe and it should be eventually confirmed solely by better preserved material. </p>
            <p> As for the few specimens from Spilia 0 and Spilia 2, all preserved vertebrae are too fragmentary with broken off hypapophyses, neural spines and paradiapophyses (Fig. 54). Te elongated centrum, the presence of hypapophysis with distinct triangular anterior keel, as well as the presence of short epizygapophyseal ridges enable identification of these fragmentary vertebrae as possibly belonging to natricids (Head, 2005; Ivanov, 2002; Szyndlar, 1984, 1991b). Although an assignation of this material to the genus  Natrix , already identified in Spilia 0 on the basis of several more complete specimens, seems probable, the absence of important diagnostic structures hinders even an exact family-level identification. </p>
            <p> As for the Vevi specimen, this anterior trunk vertebra (UU VE 601) possesses a triangular anterior keel with distinct subcotylar tubercles and almost straight subcentral ridges in lateral view (Fig. 55). Te first two features typically occur in large specimens of  Natrix -like snakes. Tus, we tentatively assign UU VE 601 to?  Natricidae indet. </p>
            <p>Colubroidea indet.</p>
            <p>Figures 56, 57, 58 Material. Spilia 0: Two anterior trunk vertebrae (UU SP0 166 and UU SP0 167); Spilia 4: an anterior trunk vertebra (UU SP4 507) and one trunk vertebra (UU SP4 618); Vevi: a mid-trunk vertebra (UU VE 602).</p>
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	https://treatment.plazi.org/id/E86287BFFFDBFFC7BA71A4EEFA89FB64	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Georgalis, Georgios L.;Villa, Andrea;Ivanov, Martin;Delfino, Massimo	Georgalis, Georgios L., Villa, Andrea, Ivanov, Martin, Delfino, Massimo (2024): New diverse amphibian and reptile assemblages from the late Neogene of northern Greece provide novel insights into the emergence of extant herpetofaunas of the southern Balkans. Swiss Journal of Palaeontology (34) 143 (1): 1-91, DOI: 10.1186/s13358-024-00332-7, URL: https://doi.org/10.1186/s13358-024-00332-7
E86287BFFFADFFBBBA71A5AEFD94FBC4.text	E86287BFFFADFFBBBA71A5AEFD94FBC4.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Elapidae Boie 1827	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Elapidae Boié, 1827</p>
            <p> Elapidae gen. et sp. indet. (small form) </p>
            <p>Figure 59</p>
            <p>Material. Spilia 4: one trunk vertebra (UU SP4 537).</p>
            <p>Description.</p>
            <p>Te vertebra (UU SP4 537) is small and incomplete, missing most of the right prezygapophysis and postzygapophysis and parts of the neural spine, the zygosphene, and the left prezygapophyseal accessory process (Fig. 59). In anterior view (Fig. 59a), the zygosphene is thin and moderately arched. Te neural canal is large. Te prezygapophyses are only slightly dorsally inclined. Te cotyle is large; its precise outline cannot be assessed as it is damaged. A paracotylar foramen is visible next to the right side of the cotyle. In posterior view (Fig. 59b), the neural arch is depressed, with a vaulting ratio (sensu Georgalis et al., 2021b) equal to 0.25. Te condyle is elliptical and slightly laterally compressed. In lateral view (Fig. 59c), the neural spine commences to rise in height slightly posterior to the zygosphenal facets. Te neural spine is not well preserved and its original height cannot be precisely assessed; nevertheless, the thin base of the neural spine in dorsal view indicates that the neural spine was originally most probably very low in lateral view. Te zygosphenal facets are narrowly elliptical. A large and deep lateral foramen is situated below the relatively straight interzygapophyseal ridge. Te subcentral ridges are convex. Te hypapophysis runs below most of the centrum but its height (slightly) augments only towards its posterior termination; its distal tip extends to the condylar base. Even at this maximum height, the hypapophysis is still rather short, still projecting more posteriorly than ventrally. In dorsal view (Fig. 59d), the neural spine is thin and runs throughout about 3/4 of the midline of the neural arch. Te (sole preserved) left prezygapophysis is projecting anterolaterally. Te interzygapophyseal constriction is deep. Te posterior median notch of the neural arch is relatively deep. In ventral view (Fig. 59e), the centrum is elongated. Te subcentral grooves are deep. Te hypapophysis is thin, commencing anteriorly right below the ventral level of the cotyle and terminating posteriorly at the level of the condyle. Te postzygapophyses extend posterolaterally. Te postzygapophyseal articular facets are oval and elongated. A precondylar constriction is present, with the condyle developed at a distinct condylar neck.</p>
            <p> Remarks. Te trunk vertebra (UU SP4 537) can be attributed to a small-sized elapid (coral snakes) on the basis of its relatively small dimensions (centrum length), its elongated vertebral centrum, and the dorsoventrally short hypapophysis that is directed more posteriorly than ventrally (Ivanov, 2002; Zaher et al., 2019). It appears that there are some differences between the Spilia coral snake and  Micrurus Wagler, 1824 , which is also known from the Neogene of Europe (see Discussion below), as the latter is characterized by a longer hypapophysis and a low but well developed neural spine, compared to the Greek form (see figures in Camolez &amp; Zaher, 2010; Escobar et al., 2024; Ivanov &amp; Böhme, 2011; Onary et al., 2018; Rage &amp; Holman, 1984; Zaher et al., 2019). In fact, the Spilia specimen appears to be overall similar to the small indeterminate  Elapidae that has been described from Merkur-North, Czechia (Ivanov, 2002). </p>
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	https://treatment.plazi.org/id/E86287BFFFADFFBBBA71A5AEFD94FBC4	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Georgalis, Georgios L.;Villa, Andrea;Ivanov, Martin;Delfino, Massimo	Georgalis, Georgios L., Villa, Andrea, Ivanov, Martin, Delfino, Massimo (2024): New diverse amphibian and reptile assemblages from the late Neogene of northern Greece provide novel insights into the emergence of extant herpetofaunas of the southern Balkans. Swiss Journal of Palaeontology (34) 143 (1): 1-91, DOI: 10.1186/s13358-024-00332-7, URL: https://doi.org/10.1186/s13358-024-00332-7
E86287BFFFACFFBFB99EA0CEFA37FCA3.text	E86287BFFFACFFBFB99EA0CEFA37FCA3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Viperinae Oppel 1811	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Viperinae Oppel, 1811b</p>
            <p> Viperinae (“Oriental Vipers”) indet. </p>
            <p>Figure 60</p>
            <p>Material. Spilia 3: one fang (UU SP3 601), two trunk vertebrae (UU SP3 674 and UU SP3 675), and six fragments of trunk vertebrae (UU SP3 676).</p>
            <p>Description.</p>
            <p>Te isolated fang (UU SP3 601) from Spilia 3 is relatively complete (Fig. 60a–e). Te specimen is relatively large. Te apical termination is slightly curved, with a wide pulpal cavity and the venom canal situated anteriorly in central position. In dorsal view (Fig. 59e), the base of the entrance orifice, which is situated in the anteriormost proximal part of the element, is indicated by the separation of the dentine folds which form the anterior closure of the venom canal. In anterior view (Fig. 60c, d), there is a distinct suture close to the distal termination of the element. Tis suture turns proximally into a narrow groove, which diminishes in front of the entrance orifice base where the fang surface is completely smooth. Te discharge orifice is preserved. A wide groove occurs on both lateral sides of the fang along its entire length.</p>
            <p>Te few available vertebrae from Spilia 3 are rather fragmentary, missing in fact, most parts of the vertebral body (Fig. 60f, g). Despite their fragmentary nature, they are characterized by a large overall size, massive cotyle and condyle, and the presence of hypapophysis.</p>
            <p> Remarks. Te fang can be referred to  Viperidae based on the presence of a closed venom canal (Edmund, 1969; Kardong, 1979; Zahradnicek et al., 2008). Te large size of the fang suggests a probable referal to the “Oriental Vipers” complex. In fact, the fang looks very similar with the fang of “Oriental vipers” described by Georgalis et. al. (2019b, fig. 38.6–10) from the nearby (and slightly older; MN 13/14) locality of Maramena. Te fragmentary vertebrae can be referred to  Viperidae based on their massive nature, the relatively short centrum, the presence of hypapophysis, and the dorsal inclination of the prezygapophyses in anterior view (Georgalis et al., 2016a; Szyndlar, 1984, 1991b; Zaher et al., 2019). Unfortunately, important viperid features, such as the anteroventral inclination of the parapophyses, the shape and inclination of the hypapophysis, the height of the neural spine, and the strongly depressed neural arch, are not preserved in the available vertebrae from Spilia, precluding thus a more precise identification. Nevertheless, as in the case of the fang above, the large size of the vertebrae suggests a probable referral to the “Oriental Vipers” complex. </p>
            <p>Colubriformes incertae sedis</p>
            <p> Periergophis Georgalis, Villa, Ivanov, Vasilyan &amp; Delfino, 2019b</p>
            <p> Periergophis micros Georgalis, Villa, Ivanov, Vasilyan &amp; Delfino, 2019b</p>
            <p>Figures 61, 62</p>
            <p>Material. Spilia 1: one posterior mid- or posterior trunk vertebra (UU SP1 1011); Spilia 4: one anterior trunk vertebra (UU SP4 550), three trunk vertebrae (UU SP4 539, UU SP4 619, and UU SP4 623), and eight caudal vertebrae (UU SP4 542– UU SP4 545, UU SP4 547, UU SP4 548, UU SP4 620, and UU SP4 658).</p>
            <p>Description.</p>
            <p>A few trunk vertebrae were available in our sample from Spilia 1 and Spilia 4 (Fig. 61). UU SP4 550 is an anterior trunk vertebra, judging from the presence of a hypapophysis (Fig. 61a–c). Te specimen has small dimensions, blunt subcentral ridges, and the anterior margin of the neural spine is inclined posteriorly. Te zygosphene in this specimen is distinctly trilobed and the posterior median notch of the neural arch is rather deep. Te hypapophysis is not fully preserved, but it seems that it was somehow posteroventrally inclined in lateral view; in ventral view, this structure crosses the whole midline of the ventral surface of the centrum.</p>
            <p> Te remaining trunk vertebrae from Spilia 4 (UU SP4 539, UU SP4 619, and UU SP4 623) and the single vertebra from Spilia 1 (UU SP1 1011) are all posterior mid- or posterior trunk vertebrae, judging from the wide haemal keel and the deep subcentral grooves (Fig. 61d–m). However, they all lack the characteristic haemal keel tubercles that are known for the posterior trunk vertebrae of this species from the type locality of Maramena. In these small vertebrae, the haemal keel is considerably wide, running throughout the midline of the centrum. Te haemal keel possesses a distinctive constriction, situated posteriorly from the level of the synapophyses; the degree of this constriction varies, being apparently dependent on the vertebral position on the column, and is particularly prominent in UU SP4 619 (Fig. 61k). Te neural spine is dorsoventrally short; when fully preserved, its anterior margin is straight to posteriorly inclined. Te neural arch is much depressed, with a vaulting ratio sensu Georgalis et. al, (2021b) ranging between 0.22 and 0.28. Te zygosphene is trilobed, with the two lateral lobes being considerably prominent. Te prezygapophyseal accessory processes are relatively long and acute. Te synapophyses are clearly separated into diapophysis and parapophysis. Cotyles and condyles are small and almost circular . </p>
            <p>All available caudal vertebrae from Spilia 4 are incomplete, with their haemapophyses being mostly damaged</p>
            <p> (Fig. 62).  Because of this incompleteness, in most specimens it is not possible to assess whether or not the, presumably autapomorphic, haemapophyseal tubercles were indeed present in these vertebrae, as in the case of most caudal vertebrae from the type locality of Maramena. Te only two caudal vertebrae that do possess more or less complete haemapophyses (Fig. 62h, n) do not show any evidence of haemapophyseal tubercles, but as it was already suggested by Georgalis et. al. (2019b), this presence/absence across the column might be subjected to intracolumnar variation. Te caudal vertebrae possess large subcentral foramina and unusual “lateral wings”, situated close to the base of the broken off pleurapophyses. Tese distinctive “lateral wings” are actually present in all available caudal vertebrae. Te pleurapophyses are elongated, pointed, and face anteroventrally. Te neural spine is dorsoventrally rather short; its posterior margin is either almost vertical or inclined anteriorly (Fig. 62h, k, m, n, r). Large lateral foramina are present below the interzygapophyseal ridges. Cotyles and condyles are small and almost circular. Paracotylar foramina are present. Te more elongated vertebra UU SP4 543 represents a more posterior caudal vertebra than the other specimens (Fig. 62f–i) . </p>
            <p> Remarks. Te material from Spilia 1 and Spilia 4 resembles  Periergophis micros , otherwise known from the nearby Maramena, in regards to the following features: trunk vertebrae with anterior margin of the neural spine being posteriorly inclined, rather wide haemal keel with a constriction present posteriorly to the synapophyses, zygosphene with distinct lateral lobes, a depressed neural arch, large parapophyses, and a small size with an overall light structure (see Georgalis et al., 2019b). However, the most prominent diagnostic feature (the presumed autapomorphy) of this species is not evident in the Spilia sample: the distinct paired anteroventrally directed tubercles or short processes developed on the wide haemal keel of posterior mid- and posterior trunk vertebrae and the haemapophyseal tubercles separated from the remaining part of haemapophyses in caudal vertebrae, at least in a portion of the caudal series (Georgalis et al., 2019b). We here interprete this absence of this feature as: </p>
            <p>i) intracolumnar variation: the few available Spilia trunk vertebrae pertain to a part of the posterior mid- or anterior posterior trunk portion of the vertebral column that was genuinely not characterized by the presence of haemal keel tubercles. Similarly, the available caudal vertebrae from Spilia originate from a portion of the caudal vertebral column that was genuinely not characterized by the presence of haemapophyseal tubercles. As also suggested by Georgalis et. al. (2019b, p. 45), “an alternative explanation would be that in some portion of the caudal series, the haemapophyseal tubercles are substituted by strongly ventrally exposed anterior thickenings of the haemapophyses”.</p>
            <p> ii) chronospecific variation: Spilia is younger than Maramena and eventually this structure disappeared from this lineage. In this case, perhaps the Spilia sample would pertain to a different species within the genus  Periergophis . </p>
            <p>iii) taphonomy: these structures were originally present in (certain of) the Spilia specimens but have been eroded/faded out due to preservational/taphonomical issues.</p>
            <p> Worth mentioning is that in our sample there are more available caudal vertebrae compared to trunk ones; Smith (2013) had proposed that in fossil localities, such high percentages of caudal versus trunk vertebrae could be indicative of snakes with long tails. However, based on such limited sample from Spilia, it is premature to assume anything about the tail proportions of the Spilia  Periergophis . Moreover, the original material of  Periergophis micros from the type locality of Maramena consisted of much more trunk vertebrae compared to caudal ones. </p>
            <p> Paraxenophis Georgalis, Villa, Ivanov, Vasilyan &amp; </p>
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	https://treatment.plazi.org/id/E86287BFFFACFFBFB99EA0CEFA37FCA3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Georgalis, Georgios L.;Villa, Andrea;Ivanov, Martin;Delfino, Massimo	Georgalis, Georgios L., Villa, Andrea, Ivanov, Martin, Delfino, Massimo (2024): New diverse amphibian and reptile assemblages from the late Neogene of northern Greece provide novel insights into the emergence of extant herpetofaunas of the southern Balkans. Swiss Journal of Palaeontology (34) 143 (1): 1-91, DOI: 10.1186/s13358-024-00332-7, URL: https://doi.org/10.1186/s13358-024-00332-7
