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Testudinidae indet. (Fig. 4) LOCALITIES. — MWQ4/2018. STUDIED MATERIAL. — Czech Republic. South Moravia Region, Mokrá-Quarry, shell and postcranial remains (Fig. 4 A-M): Pal. 1363, shell fragment; Pal. 1364, scapula (i.e., anterodorsal process fragment); Pal. 1365, right fibula. DESCRIPTION Only a shell fragment formed by two portions of plates is known (Fig.4 A-D).Pal.1363 corresponds most likely to a carapace portion, but it is poorly preserved and is not possible to assess this confidently. The length and maximum width of the preserved plate fragment is 6 cm. The external part is completely smooth and is not crossed by any sulcus (Fig. 4C, D). A suture is recognized on top of plate, which is concave and approximately 2 cm wide. A partial bone, belonging to the shoulder girdle, has been identified (Pal.1364: Fig.4 E-H). Only the anterodorsal process fragment is preserved.Pal.1364 is subcylindrical in cross-section and distally rounded.The distal surface ends in a rough rounded area to join with the visceral part of the carapace (Fig. 4H). The hind limb skeleton is restricted to one partial fibula (Fig. 4 I-M) that is elliptical in cross-section. Its distal articular surface is slightly small, oval and convex (Fig.4M).Both postcranial bones are poorly preserved and no significant details can be discerned. REMARKS Fossil remains of giant tortoises are not very common in Miocene assemblages of Central Europe; their record being limited to few localities from Austria, Germany, Hungary and Switzerland (Alba et al. 2010, 2011; Carmona et al. 2011; Luján 2015). Loveridge & Williams (1957) proposed that all European giant tortoises should be transferred into the extant genus Geochelone. This proposal was adopted for some time, and consequently, large tortoise remains in Europe are still frequently referred to in the literature as Geochelone sp. (e.g., Auffenberg 1974; Młynarski 1976). However, current phylogenies do not support a close relationship between Mio- Pleistocene large tortoises and Geochelone. More recently, Bourgat & Bour (1983) referred all giant fossil tortoises to the genus Cheirogaster. Most subsequent works accepted this genus attribution (e.g., Luján et al. 2010, 2014), until recently when Pérez-García & Vlachos (2014) proposed that European Neogene giant tortoises constitute a clade that is more derived than the type species of Cheirogaster.To allocate these taxa, Pérez-García & Vlachos (2014) erected the genus Titanochelon, with Ti. bolivari (Hernández- Pacheco, 1917) as its type species. This genus is characterized by a shell reaching over 100 cm and the fusion of marginal scutes 12 (i.e., constituting a supracaudal scute). However, the evolution of gigantism amongst fossil tortoises is clearly a homoplastic phenomenon, mainly related to insular conditions, or adaptation to either global or local environmental changes (Kear 2010; Luján et al. 2010, 2017b; Itescu et al. 2014). Similarly, the fusion of marginal scutes 12 occurs in many extant and extinct genera and cannot be considered autapomorphic for the genus Titanochelon. In summary, the taxonomy of the Miocene giant tortoises of Europe is still a subject of debate and will require improvement of existing data matrix (e.g., including more skull characters) in order to decipher the phylogenetic relationships of Titanochelon (Luján et al. 2017b).

Cadurcogekko cf. piveteaui (Figs 2-4) REFERRED SPECIMENS. — A frontal (NHMW 2019/0052/0002); a left maxilla (NHMW 2019/0052/0001). DESCRIPTION Frontal NHMW 2019/0052/0002 (Fig. 2) The left and right frontals are fused in this specimen, forming a single element, with a length of 8.1 mm (Fig. 2). It is almost completely preserved and only its anterior portion is damaged. It is rectangular, with a slight mid-constriction. Thus, the lateral margins are rather concave in dorsal aspect and fluently continue into well laterally expanded posterolateral processes. These processes form narrow triangles with pointed lateral terminations. The anterolateral margins of both processes are stepped due to a presence of short and narrow facets for the postfrontal. An additional step is present on both sides in the anterior region of the lateral margin of the frontal. There, the large facet for prefrontal is located (Fig. 2B, C). It is mainly exposed in ventral aspect as a rough surface. It is wedge-shaped posteriorly, where its peak reaches posterior to the level of the step. The facet for prefrontal and postfrontal are not in contact, thus the frontal is not fully excluded from the orbit. In ventral view, the frontal cranial crests run medially from the posteroventral processes and merge together further anteriorly (Fig. 2B). They are completely fused to form a tunnel-like structure. The overall dorsal surface of the frontal, although slightly bulged in the central region, is slightly depressed, with dorsally inclined lateral portions. This external surface is sculptured, having a wrinkled appearance. The sculpturing pattern consists of ridges and pits (Fig. 2A). They are mainly developed in the mid-section and run to the periphery of the surface. A deep depression runs along the entire dorsal surface of both posterolateral processes. The posterior margin of the frontal is almost straight. Maxilla NHMW 2019/0052/0001 (Figs 3; 4) NHMW 2019/0052/0001 represents an almost complete left maxilla (Figs 3; 4). This element is elongate and lightly built, measuring 14.2 mm in length. It is straight with a small medial anterior curvature. A complete tooth row is preserved, bearing 37 tooth positions, of which six complete teeth are still attached. The premaxillary process is bifurcated. The external ramus of this process is short and blunt, whereas the internal ramus is more distinct and broader. The internal ramus is more medially oriented and more dorsally located relative to the external one. Between them, an oval and wide premaxillary notch is present. Dorsal to it, an anterior opening for the superior alveolar canal is located. The supradental shelf is thin. Its portion located in the anterior two thirds is slightly bent dorsally (convex) and expanded medially. In dorsal view, the medial margin of the supradental shelf is rounded, reaching the maximum of expansion at the level around of 18th tooth position (counted from anterior). This forms a palatine process. Slightly posteriorly to it, the rounded but flat superior alveolar foramen is located at the dorsal portion, at the level of the 21 st tooth position (counted from anterior). The posterior 1/3 of the supradental shelf is more or less straight in medial aspect. In posterior direction, it is gradually less expanded medially. The anteroposterior length of the nasal process is greater than its dorsal height. The dorsal margin of the nasal process appears to be slightly worn, but in the case that it represents more or less the original shape, it shows a lack of a pronounced dorsal process. This configuration of the nasal process contrasts sharply with the prominent, often pointed process typical of many gekkotans and resembles that seen in Euleptes Fitzinger, 1843 (e.g., Bauer et al. 1997; Čerňanský et al. 2018; Villa et al. 2018). The nasal process is trapezoidal in shape, with the posterior margin being more ventrally slopped relative to the anterior one. The anterior margin possesses a free terminus. It forms a triangular tip (this feature can be seen in extant and extinct species of the genus Euleptes but also in the genera Hemidactylus Goldfuss, 1820, and Tarentola Gray, 1825; see e.g., Čerňanský et al. 2018; Villa et al. 2018), well delimited from the dental portion of the maxilla. In the anterior region of the nasal process, on the medial side there is a fine ridge that runs posterodorsally. This ridge originates from the supradental shelf at the level of the fifth tooth position. The internal side of the nasal process posterior to this ridge is excavated, forming a cavity. In dorsal aspect, the process has a rounded, laterally convex appearance. The posteroventral process of maxilla is moderately long. It possesses a longitudinal deep groove, which is roofed by a bent, dorsally depressed bony flange. This flange continues here from the dorsolateral side of the posteroventral process. The lateral surface of maxilla is pierced by nine labial foramina located in the ventral series, where the posteriormost one is located at the level of the 27th tooth position (Figs 3A; 4A). Posteriorly from this foramen, a well-developed groove runs along the entire portion of the posteroventral process. Dorsal to this series, the surface is pieced by additional five, irregularly arranged foramina. The rest of the lateral surface is rather rugose and irregularly pitted. REMARKS The material can be referred to Gekkota on the basis of the following characters: 1) absence of osteoderms fused to the skull bones; 2) unpaired frontals; 3) fused subolfactory processes of frontal in the mid-line to form a tunnel like structure; and 4) the high number of conical, unicuspid pleurodont teeth (Estes 1983; Daza et al. 2014). The maxilla can be referred to the genus Cadurcogekko on the basis of the presence of rugose surface of the nasal process, an elongated groove associated with the most posterior neurovascular foramen, and a postnarial anterodorsal depression of the element (Augé 2005). The genus Cadurcogekko is known exclusively from localities within the Phosphorites du Quercy.Two species of this genus are currently regarded as valid: the type species, Cadurcogekko piveteaui from the old collections of the Phosphorites du Quercy, and Cadurcogekko verus Bolet, Daza, Augé & Bauer, 2015, from the late Eocene (MP 17) of Les Pradigues, Quercy (see Bolet et al. 2015). Notably, the holotype dentary of the type species, C. piveteaui, was originally described as a frog by Piveteau (1927), until it was eventually demonstrated by Hoffstetter (1946) that it pertained instead to gekkotans. The two valid species of Cadurcogekko can be distinguished by: 1) different size of the available elements, with Cadurcogekko verus being smaller than C. piveteaui; 2) lower maxillary tooth number – according to Bolet et al. (2015), C. verus possesses 30 vs. 50 in C. piveteaui. The tooth row in the referred maxilla MNHN.F.QU17734 of Cadurcogekko piveteaui from the late Eocene (MP 17) of Perrière, Quercy, is incomplete, lacking its posterior portion; it has 38 tooth positions (Augé 2005: fig. 59). Daza et al. (2014) estimated the tooth number in a complete tooth row to be between 40 and 44, contrasting with only 27 tooth positions in C. verus; and 3) a coarser sculpture on the external surface of the maxilla and possibly the frontal in C. verus. Note that a third named species of the genus, Cadurcogekko rugosus Augé, 2005, has recently been reidentified as a scincid and placed accordingly in its own genus, Gekkomimus Bolet, Daza, Augé & Bauer, 2015, by Bolet et al. (2015). Nevertheless, the figure of the type material of Cadurcogekko verus in Bolet et al. (2015: fig. 1) deserves a comment at this point. Bolet et al. (2015) established the species Cadurcogekko verus and they supposedly figured its holotype, the right maxilla UM PRA 9, in their fig. 1. That specimen, UM PRA 9, originated from the late Eocene (MP 17) of Les Pradigues, Quercy and was previously allocated to C. rugosus by Augé (2005). However, despite their figure caption which stated that UM PRA 9 was figured there, Bolet et al. (2015) did not figure at all this holotype, but instead their fig. 1 represents another specimen, the right maxilla MNHN.F.QU17734 of C. piveteaui originating from the late Eocene (MP 17) of Perrière, Quercy (see Augé 2005: fig. 59a-b [for MNHN.F.QU17734] vs. 64a-b [for UM PRA 9] and Daza et al. 2014: fig. 5a-b [for MNHN.F.QU17734]; this paper: Fig. 5 [for UM PRA 9]). In addition, Daza et al. (2014) mentioned in their text the maxilla UM PRA 9 (following the previously suggested referral by Augé [2005] to C. rugosus) and stated that they figured this specimen in their figure 5j-k. However, the photographs of the maxilla “ UM PRA 9” figured by Daza et al. (2014: fig. 5j-k) represent a different specimen from the drawings of the maxilla “ UM PRA 9” figured by Augé (2005: fig. 64a-b). Judging from newly furnished photographs of the specimen provided to us by UM, we here confirm that UM PRA 9 is the specimen figured by Augé (2005: fig. 64a-b) and not the one figured by Daza et al. (2014: fig. 5j-k). All these being said, despite the erroneous figure of Bolet et al. (2015), UM PRA 9 indeed represents the holotype of Cadurcogekko verus, as the diagnosis and description of this species is based on this specimen. We here provide photographs of this specimen, UM PRA 9, for the first time (Fig. 5). The left maxilla NHMW 2019/0052/0001 described herein exhibits a complete tooth row, possessing 37 tooth positions. This seems to be an intermediate condition between Cadurcogekko verus and that what has been previously estimated for Cadurcogekko piveteaui. This brings several questions about this character state in these two currently valid species. The frontal NHMW 2019/0052/0002 is almost identical with specimen MNHN.F.QU17165, which has been referred to C. piveteaui (see Augé 2005: fig. 60; Daza et al. 2014: fig. 5h-i). However, the sculpturing pattern present on the dorsal surface of NHMW 2019/0052/0002 appears to be slightly more strongly developed relative to that of MNHN.F.QU17165. The frontal NHMW 2019/0052/0002, despite the fact that its anterior portion is damaged, currently represents the best preserved frontal of Cadurcogekko. In any case, the overall morphologies of the specimens NHMW 2019/0052/0001 and NHMW 2019/0052/0002 fit better to the diagnosis previously stated for C. piveteaui, although with some doubts. Therefore, we have decided to refer this gekkotan material to as Cadurcogekko cf. piveteaui. LATERATA Vidal & Hedges, 2005 LACERTIFORMATA Vidal & Hedges, 2005 Family LACERTIDAE Oppel, 1811 Subfamily GALLOTIINAE Cano, Baez, López-Jurado & Ortega, 1984, Published as part of Georgalis, Georgios L., Čerňanský, Andrej & Klembara, Jozef, 2021, Osteological atlas of new lizards from the Phosphorites du Quercy (France), based on historical, forgotten, fossil material, pp. 219-293 in Geodiversitas 43 (9) on pages 223-227, DOI: 10.5252/geodiversitas2021v43a9, http://zenodo.org/record/4720776, {"references":["FITZINGER L. J. F. J. 1843. - Systema reptilium. Fasciculus primus. Amblyglossae. Braumuller et Seidel Bibliopolas, Vindobonae (Vienna), 106 p. https: // doi. org / 10.5962 / bhl. title. 4694","BAUER A. M., GOOD D. A. & BRANCH W. R. 1997. - The taxonomy of the Southern African leaf-toed geckos (Squamata: Gekkonidae) with a review of old World \" Phyllodactylus \" and the description of five new genera. Proceedings of the California Academy of Sciences 49: 447 - 497. https: // www. biodiversitylibrary. org / page / 15777227","CERNANSKY A., DAZA J. D. & BAUER A. M. 2018. - Geckos from the middle Miocene of Devinska Nova Ves (Slovakia): new material and a review of the previous record. Swiss Journal of Geosciences 111: 183 - 190. https: // doi. org / 10.1007 / s 00015 - 017 - 0292 - 1","VILLA A., DAZA J. D., BAUER A. M. & DELFINO M. 2018. - Comparative cranial osteology of European gekkotans (Reptilia, Squamata). Zoological Journal of the Linnean Society 184: 857 - 895. https: // doi. org / 10.1093 / zoolinnean / zlx 104","GOLDFUSS G. A. 1820. - Reptilia, in SCHUBERT G. H. (ed.), Handbuch der Naturgeschichte zum Gebrauch bei Vorlesungen. Vol. 3. Handbuch der Zoologie. J. L. Schrag, Nurnberg: 121 - 181.","GRAY J. E. 1825. - A synopsis of the genera of Reptiles and Amphibia, with a description of some new species. Annals of Philosophy, Series 2, 10: 193 - 217. https: // www. biodiversitylibrary. org / page / 2531387","ESTES R. 1983. - Sauria Terrestria, Amphisbaenia, in WELL- NHOFER P. (ed.), Encyclopedia of Paleoherpetology. Part 10 a. Gustav Fischer, Stuttgart, New York, 249 p.","DAZA J. D., BAUER A. M. & SNIVELY E. D. 2014. - On the fossil record of the Gekkota. The Anatomical Record 297: 433 - 462. https: // doi. org / 10.1002 / ar. 22856","AUGE M. 2005. - Evolution des lezards du Paleogene en Europe. Museum national d'Histoire naturelle, Paris: 1 - 369 (Memoires du Museum national d'Histoire naturelle; 192).","BOLET A., DAZA J. D., AUGE M., & BAUER A. M. 2015. - New genus and species names for the Eocene lizard Cadurcogekko rugosus Auge, 2005. Zootaxa 3985: 265 - 274. https: // doi. org / 10.11646 / zootaxa. 3985.2.5","PIVETEAU J. 1927. - Etudes sur quelques amphibiens et reptiles fossiles. Annales de Paleontologie 16: 59 - 99.","HOFFSTETTER R. 1946. - Sur les Gekkonidae fossiles. Bulletin du Museum national d'Histoire naturelle, 2 eme serie, 18: 195 - 203. https: // www. biodiversitylibrary. org / page / 53794135","VIDAL N. & HEDGES B. S. 2005. - The phylogeny of squamate reptiles (lizards, snakes, and amphisbaenians) inferred from nine nuclear protein-coding genes. Comptes Rendus Biologies 328: 1000 - 1008. https: // doi. org / 10.1016 / j. crvi. 2005.10.001","OPPEL M. 1811. - Die Ordnungen, Familien und Gattungen der Reptilien als Prodrom einer Naturgeschichte derselben. Joseph Lindauer, Munich, 87 p. https: // doi. org / 10.5962 / bhl. title. 4911"]}

Pseudeumeces sp. (Figs 15-17) REFERRED SPECIMENS. — A left maxilla (NHMW 2019/0051/0004); a left dentary (NHMW 2019/0051/0003); a right dentary (NHMW 2019/0051/0005). DESCRIPTION Maxilla NHMW 2019/0051/0004 (Fig. 15) Only one such element is available in our collection, the left maxilla NHMW 2019/0051/0004 (Fig. 15). This specimen is almost completely preserved. In medial view, the supradental shelf is well medially expanded, having rounded (dorsally convex) course (Fig. 15B). The maxilla bears 11 tooth positions (10 teeth are still attached). However, the premaxillary process is broken off and only its posterior root portion is preserved. Thus, it can be estimated that the tooth number in a complete tooth row was around 12. The superior alveolar foramen is located at the level between the 3rd and 4th tooth positions (counted from posterior). Posterior to this, the maxilla protrudes into the posteroventral process, having a facet for jugal on its dorsal internal side. A facet for the palatine is positioned medial to the superior alveolar foramen. The posteroventral process of the maxilla slightly narrows posteriorly, although its termination is not pointed, but rather stepped. This posteriormost portion does not bear dentition. In the anterior half of the bone, the nasal process is well dorsally elevated, being high. Its dorsal end slightly bents medially. However, the posterodorsal tip, which forms the contact with the fron- tal, is broken off. On the medial side of the nasal process, the carina maxillaris starts to rise dorsally at the level of the 3rd preserved tooth (counted from anterior). Further, it is inclined posteriorly and thus it does not reach a high level dorsally. In the dorsal portion of the process, a facet for the prefrontal is present. In lateral view, the ventral region of the maxilla is pierced by six labial foramina of various sizes (Fig. 15A). The posteriormost one is located at the level of the 4th tooth position (counted from posterior). The dorsally located nasal process is completely covered by three osteoderms. These are clearly demarked by sulci. The sulci meet all together at the level of the 4th tooth position (counted from ante- rior), forming a Y-shaped structure. The anteroventral osteoderm is the smallest one, whereas the largest is the posterior osteoderm. All three osteoderms are sculptured. The sculpture consists of densely arranged pits and ridges running to the periphery. The posteroventral process, posterior to the level of osteoderm, has bulged dorsal margin. Dentaries NHMW 2019/0051/0003 and NHMW 2019/0051/0005 (Figs 16; 17) Specimen NHMW 2019/0051/0005 is small in size and slightly damaged, whereas NHMW 2019/0051/0003 represents only fragment of the posterior portion of the dentary. NHMW 2019/0051/0005 possesses 15 tooth positions, with seven teeth being still attached (Fig. 16). The dorsal crest is high and the teeth only slightly exceed it dorsally. Meckel`s groove is fully open, but the ventral portion of the dentary is broken off. In any case, the dentary is narrow rather than robust. Its lateral surface is pierced by five labial foramina. In the posterodorsal region of the dentary, the wedge shaped, well defined facet for the anterolateral process of coronoid is present. It reaches the level of the penultimate tooth position. NHMW 2019/0051/0003 possesses only six tooth positions, with five teeth preserved (Fig. 17); its further anterior region is broken off and missing. The facet for the anteromedial process of the coronoid reaches the level of the last posterior tooth. On lateral side, the facet for the anterolateral process of coronoid is well defined, reaching the level between the last and penultimate tooth position. Dentition The dentition is pleurodont and amblyodont. The teeth are closely spaced. The tooth crowns bear delicate striations on both maxillary and dentary teeth. The maxillary tooth length varies, resulting in a sinuous occlusal surface. Here, the teeth in the posterior section are more robust except for the last two. The teeth in the anterior portion of the tooth row are slightly pointed and curved posterolingually. On some of those maxillary teeth, there is a very small indication of an indistinct, incipient mesial cusp. REMARKS The maxilla NHMW 2019/0051/0004 bears 12 tooth positions, whereas 15 are present in that of Pseudeumeces cadurcensis (see Augé 2005; Augé & Hervet 2009). For this reason, the maxilla NHMW 2019/0051/0004 potentially pertains to the above described species Pseudeumeces kyrillomethodicus n. sp. However, as there is a lack of a strong support for such association based on the available material, we decided to allocate this maxilla only as Pseudeumeces sp. Small differences in the anterior maxillary teeth of NHMW 2019/0051/0004 and the dentary teeth of Pseudeumeces kyrillomethodicus n. sp. can be explained by an ontogenetic change. Judging from the smaller size of the maxilla NHMW 2019/0051/0004 relative to dentaries, the former specimen most likely represents a late juvenile (or subadult) individual. Similar changes have been observed in both extant and extinct lacertids. For example, in the early Miocene Janosikia ulmensis (Gerhardt, 1903), vestiges of mesial cusps are present on some anterior maxillary teeth in a juvenile specimen (see Čerňanský et al. 2016b). Additionally, the ontogenetic change in the tooth morphology is sometimes observed in the extant Gallotia stehlini (Schenkel, 1901) as well, where the juvenile tricuspid teeth are replaced by multicuspid ones in adult individuals (Barahona et al. 2000). The maxilla NHMW 2019/0051/0004 further differs from that of Dracaenosaurus in the following features: 1) maxillary tooth number is ~12 rather than 7; 2) the posteroventral process of the maxilla is not markedly high as it is in D. croizeti; and 3) the presence of three well developed osteoderms attached to the nasal process of maxilla. Two dentaries are also referred to Pseudeumeces sp. The specimen NHMW 2019/0051/0005 represents the smallest lacertid dentary in our sample. It is very likely that it represents a juvenile (or subadult) ontogenetic stage, that could potentially pertain to the above described Pseudeumeces kyrillomethodicus n. sp. Nevertheless, in comparison with that taxon, NHMW 2019/0051/0005 does not appear to be so robust and the facet for the anterolateral process of coronoid reaches at the level of the penultimate tooth position.In the other specimen, the fragment of left dentary (NHMW 2019/0051/0003), this facet reaches the level between the last and penultimate tooth. Moreover, only the last posterior tooth was reduced (it is absent, but its size can be estimated based on its tooth loci). These characters are in a sharp contrast with the type material of Pseudeumeces kyrillomethodicus n. sp. Therefore, we cannot exclude that this material does not pertain to Pseudeumeces cadurcensis, which also occurs in the Oligocene of the Phosphorites du Quercy and shares these features (e.g., Augé & Hervet 2009). The proper taxonomic allocation of fragmentary material needs always to be met with caution. This is especially true for similar forms such as those discussed herein., Published as part of Georgalis, Georgios L., Čerňanský, Andrej & Klembara, Jozef, 2021, Osteological atlas of new lizards from the Phosphorites du Quercy (France), based on historical, forgotten, fossil material, pp. 219-293 in Geodiversitas 43 (9) on pages 236-240, DOI: 10.5252/geodiversitas2021v43a9, http://zenodo.org/record/4720776, {"references":["AUGE M. 2005. - Evolution des lezards du Paleogene en Europe. Museum national d'Histoire naturelle, Paris: 1 - 369 (Memoires du Museum national d'Histoire naturelle; 192).","AUGE M. & HERVET S. 2009. - Fossil lizards from the locality of Gannat (late Oligocene-early Miocene, France) and a revision of the genus Pseudeumeces (Squamata, Lacertidae). Palaeobiodiversity and Palaeoenvironments 89: 191 - 201. https: // doi. org / 10.1007 / s 12549 - 009 - 0009 - 1","GERHARDT K. 1903. - Ophisaurus ulmensis n. sp. aus dem Untermiocan von Ulm a. D. Jahreshefte des Vereins fur vaterlandische Naturkunde in Wurttemberg 59: 67 - 71.","CERNANSKY A., KLEMBARA J. & SMITH K. T. 2016 b. - Fossil lizard from central Europe resolves the origin of large body size and herbivory in giant Canary Island lacertids. Zoological Journal of the Linnean Society 176: 861 - 877. https: // doi. org / 10.1111 / zoj. 12340","SCHENKEL E. 1901. - Achter Nachtrag zum Katalog der herpetologischen Sammlung des Basler Museums. Verhandlungen der Naturforschenden Gesellschaft Basel 13: 142 - 199. https: // www. biodiversitylibrary. org / page / 32329988","BARAHONA F., EVANS S. E., MATEO J. A., GARCIA- MARQUEZ M., & LOPEZ- JURADO L. F. 2000. - Endemism, gigantism and extinction in island lizards: the genus Gallotia on the Canary Islands. Journal of Zoology 250: 373 - 388. https: // doi. org / 10.1111 / j. 1469 - 7998.2000. tb 00781. x"]}

Anguimorpha indet. (Figs 61; 62) REFERRED SPECIMENS. — Five presacral vertebrae (NHMW 2019/0046/0003- NHMW 2019/0046/0007); a partial pectoral girdle (NHMW 2019/0095/0001). DESCRIPTION AND REMARKS Presacral vertebrae (Fig. 61) These vertebrae are relatively large (Fig. 61), with centrum lengths ranging between 6.9 and 9.3 mm (see Appendix 1). The vertebrae demonstrate a mix of several features present in the above described specimens of Placosaurus, Anguinae indet., and Palaeovaranus. They have high neural spines, depressed cotyle and condyle, while the ventral surface of their centra is crossed by a wide surface or groove that is unlike the conditions seen above for the other taxa (Fig. 61). Considering the high intracolumnar variation observed in the vertebrae of extant lizards (e.g., Pseudopus), we are reluctant in assigning these specimens in a more precise taxonomic rank and we cannot even exclude that they (or part of them) pertain to some of the above described taxa. Pectoral girdle NHMW 2019/0095/0001 (Fig. 62) This specimen is incomplete, though preserving in relatively good state the right scapulocoracoid. The glenoid fossa is visible, well demarking the point of attachment with the humerus. Anteriorly to the glenoid fossa, lies the coracoid foramen. Dorsally to the foramen, the scapulocoracoid is of rectangular shape and is dorsoventrally elongated. The ventral portion of the element is anteroposteriorly elongated. It is readily obvious that this specimen apparently pertains to a rather large-sized lizard. Considering our currently inadequate state of knowledge of the appendicular skeleton of Paleogene European lizards, it is impossible to associate it with any of the above described glyptosaurines, palaeovaranids, and varanids, all of which could attain a considerably large size. Squamata indet. REFERRED SPECIMEN. — A?sacral vertebra (NHMW 2019/0095/0002). DESCRIPTION AND REMARKS. This vertebra is incomplete and not informative. Squamata indet. REFERRED SPECIMEN. — A?sacral vertebra (NHMW 2019/0095/0002). DESCRIPTION AND REMARKS. This vertebra is incomplete and not informative.

Mediolacerta sp. (Fig. 18) REFERRED SPECIMEN. — A left dentary (NHMW 2019/0050/0001). DESCRIPTION The only available specimen, the left dentary NHMW 2019/0050/0001, is almost completely preserved, with only the half posterior ventral portion being broken off and missing (Fig. 18). It is an anteroposteriorly long and massive element, with a slight medial curvature at its anterior end. The tooth row is completely preserved and the alveolar crest supports 23 tooth positions (17 teeth still attached). Meckel’s groove is fully open along its entire length, although it is narrow in the anterior region (Fig. 18B). The alveolar foramen is located at the level of the 6th tooth position (counted from posterior). The intramandibular septum is fused to the bone, being almost horizontal in this section. Meckel’s groove is roofed by a more or less straight subdental shelf (only its posterior portion is arched). The shelf is somewhat broad only in the anterior section, but it narrows posteriorly and thus is rather thin. This is mainly caused by the presence of the facet for the splenial, situated on its ventral margin. This facet reaches the level of the 8th tooth position (counted from anterior). Anteriorly, the shelf continues to the small rectangular symphysis. Posteriorly, the dentary protrudes into a short and low coronoid process, which bears a facet for the coronoid. The otherwise smooth lateral surface of the bone is pierced by six labial foramina (Fig. 18A). They are arranged in a single row, located in the dorsal half of the dentary. The posteriormost foramen is located at the level of the 6th tooth position (counted from posterior). A facet for the coronoid is present on the dorsolateral surface of the bone. Dentition The dentition is pleurodont and heterodont (Fig. 18). The tooth size gradually increases posteriorly (although it should be noted that the last posterior tooth is slightly smaller than the penultimate one). The teeth in the anterior section of the tooth row are small, slender, and somewhat pointed, with the tooth crowns slightly curved posterolingually. Posteriorly located teeth (from the 13th one if counted from the anterior) are markedly robust and blunt in comparison to those from the anterior region. The tooth apices of several teeth are worn (or weathered), but those which are complete show bicuspidity, with an incipient small mesial cusp being present. REMARKS The dentary NHMW 2019/0050/0001 described herein represents the largest lacertid from this collection. The anterior teeth are pointed, whereas those located further posteriorly are robust and blunt, but with some of them bearing mesial cusp - this character fits to the diagnosis of Mediolacerta and its so far sole named species, Mediolacerta roceki, also from the Phosphorites du Quercy (stratigraphic occurrence MP 23- MP 30; see Augé 2005). As such, in respect of these features, NHMW 2019/0050/0001 can be differentiated from both Pseudeumeces and Dracaenosaurus. Besides the dentition, NHMW 2019/0050/0001 also shares several other features with Mediolacerta: 1) rather thin subdental shelf; 2) small rectangular symphysis; and 3) the position of the alveolar foramen at the level of the 6th tooth position, counted from posterior. Nevertheless, there appear also to be differences among NHMW 2019/0050/0001 and other known specimens of Mediolacerta. Most principally, the tooth number of the holo- type dentary (MNHN.F. PFR11006) of Mediolacerta roceki is 19, whereas this number is 23 in NHMW 2019/0050/0001. Dentaries of the same taxon have been also described from the late Oligocene of Germany (Mediolacerta cf. roceki of Čerňanský et al. 2016a); the complete tooth row of the German material bears 18 tooth positions. However, it should be noted that tooth number in lacertids, like virtually most lizards, might be variable and also likely to be size related (the anteroposterior length of the dentary NHMW 2019/0050/0001 is 23.7 mm, thus this dentary is larger than the holotype of Mediolacerta roceki and the German material. These numbers should not be interpreted as absolutes. However, because the differences in the overall size and the dentary tooth count objectively exist, we decided to allocate NHMW 2019/0050/0001 only to the genus level, as Mediolacerta sp. ANGUIMORPHA Fürbringer, 1900 Family ANGUIDAE Gray, 1825 Subfamily GLYPTOSAURINAE Marsh, 1872, Published as part of Georgalis, Georgios L., Čerňanský, Andrej & Klembara, Jozef, 2021, Osteological atlas of new lizards from the Phosphorites du Quercy (France), based on historical, forgotten, fossil material, pp. 219-293 in Geodiversitas 43 (9) on pages 240-242, DOI: 10.5252/geodiversitas2021v43a9, http://zenodo.org/record/4720776, {"references":["AUGE M. 2005. - Evolution des lezards du Paleogene en Europe. Museum national d'Histoire naturelle, Paris: 1 - 369 (Memoires du Museum national d'Histoire naturelle; 192).","CERNANSKY A., KLEMBARA J. & MULLER J. 2016 a. - The new rare record of the late Oligocene lizards and amphisbaenians from Germany and its impact on our knowledge of the European terminal Palaeogene. Palaeobiodiversity and Palaeoenvironments 96: 559 - 587. https: // doi. org / 10.1007 / s 12549 - 015 - 0226 - 8","FURBRINGER M. 1900. - Zur vergleichenden anatomie des Brustschulterapparates und der Schultermuskeln. Jenaische Zeitschrift 34: 215 - 718. https: // doi. org / 10.5962 / bhl. title. 52377","GRAY J. E. 1825. - A synopsis of the genera of Reptiles and Amphibia, with a description of some new species. Annals of Philosophy, Series 2, 10: 193 - 217. https: // www. biodiversitylibrary. org / page / 2531387","MARSH O. C. 1872. - Preliminary description of new Tertiary reptiles. Parts I and II. American Journal of Science 4: 298 - 309. https: // doi. org / 10.2475 / ajs. s 3 - 4.22.298"]}

Squamata indet. REFERRED SPECIMEN. — A?sacral vertebra (NHMW 2019/0095/0002). DESCRIPTION AND REMARKS. This vertebra is incomplete and not informative., Published as part of Georgalis, Georgios L., Čerňanský, Andrej & Klembara, Jozef, 2021, Osteological atlas of new lizards from the Phosphorites du Quercy (France), based on historical, forgotten, fossil material, pp. 219-293 in Geodiversitas 43 (9) on page 271, DOI: 10.5252/geodiversitas2021v43a9, http://zenodo.org/record/4720776

Genus Placosaurus Gervais, 1848 -1852 TYPE SPECIES. — Placosaurus rugosus Gervais, 1848 -1852 (type species by original designation; Gervais 1848 -1852)., Published as part of Georgalis, Georgios L., Čerňanský, Andrej & Klembara, Jozef, 2021, Osteological atlas of new lizards from the Phosphorites du Quercy (France), based on historical, forgotten, fossil material, pp. 219-293 in Geodiversitas 43 (9) on page 242, DOI: 10.5252/geodiversitas2021v43a9, http://zenodo.org/record/4720776, {"references":["GERVAIS P. 1848 - 1852. - Zoologie et Paleontologie francaises (animaux vertebres): ou nouvelles recherches sur les animaux vivants et fossiles de la France. Arthus Bertrand, Paris, 271 p. https: // doi. org / 10.5962 / bhl. title. 39473"]}

Palaeovaranus lismonimenos n. sp. (Figs 37B, D; 38-40; 41B; 42; 43) urn:lsid:zoobank.org:act: 3C906259-DAB5-4B97-A1BB-7860695A76B8 HOLOTYPE. — An almost complete parietal (NHMW 2019 /0047/0001). ETYMOLOGY. — The new name originates from the Greek word “λησμονημένος” (“lismonimenos”) meaning “forgotten”, alluding to the fact that the holotype specimen was forgotten and unnoticed inside a museum drawer for more than a century. TYPE LOCALITY. — Imprecisely known locality, Phosphorites du Quercy, Department of Lot or Tarn-et-Garonne, Occitanie, southern France; probably late Eocene, around MP 17 (see Distribution below). DIAGNOSIS. — The parietal of Palaeovaranus lismonimenos n. sp. differs from that of Palaeovaranus cayluxi by the following distinguished features and the combination of features: 1) the dorsolateral crests are rather distinct, extending posterolaterally and dorsally, and their margins are distinctly crenulated; 2) the median crest is short (shorter than the length of the median triangular field measured in the median plane) and its posterior tip fits between the anterolateral processes of the triangular median field; 3) the anterior ends of the dorsolateral crests extend to the tips of the anterolateral processes; 4) the ornamentation of the parietal consists of small, more or less densely arranged mounds having more or less distinctly developed crest; and 5) the anterior margin of the parietal fossa lies anterior to the level of the junctions of the anterolateral margins of the supratemporal processes with the parietal plate. REFERRED SPECIMENS. — A complete parietal (MNHN.F.QU17177) and a partial parietal (UM BFI 1873), both from juvenile individuals. Tentatively also, two frontals (MNHN.F.QU17175 and UM PRA 8). DISTRIBUTION. — The holotype parietal NHMW 2019/0047/0001, the referred parietal MNHN.F.QU17177, and the tentatively referred frontal MNHN.F.QU17175, all originate from impre- cisely known localities in the Department of Lot or that of Tarnet-Garonne, within the Phosphorites du Quercy. The referred parietal UM BFI 1873 originates from the late Eocene (MP 17) of Bouffie (= La Bouffie), Quercy (Department of Lot), while the tentatively referred frontal UM PRA 8 originates from the coeval, late Eocene (MP 17) locality of Les Pradigues, also in Quercy (but in the Department of Tarn-et-Garonne). Accordingly, we here sug- gest that the specimens with imprecise locality date (including the holotype), originate also from late Eocene locality(ies), potentially also around the MP 17 stage. DESCRIPTION Holotype NHMW 2019 /0047/0001 (Figs 37B, D; 38-40; 41B) The holotype NHMW 2019 /0047/0001 has a length of its parietal table 14.5 mm (measured in mid-line). The parietal table is anteroposteriorly elongate (Figs 37B; 38B; 39A). The anterolateral process is slender. The dorsolateral crest is distinctly developed. It extends posterolaterally and dorsally. Its margin is distinctly crenulated. The crenulation consists of several more or less shallow and long notches. The anterior end of the dorsolateral crest reaches the tip of the anterolateral process. The dorsolateral crests and the anterior margin of the parietal limit a triangular field. Its deepest portion is pierced by the parietal foramen. The foramen lies in about the midlength of the anterior half of the parietal table. The surface of the triangular field is covered by small, but distinct mounds, most of them bearing the longitudinal crests on their surfaces. The median crest is short and its posterior pointed portion fits between the two anteriorly pointed anterolateral processes of the triangular median field (Fig. 39A). The supratemporal fossa is large and is inclined ventrolaterally. The ventral surface is smooth (Figs 37D; 38A; 39B). The anterior margin of the parietal fossa lies anterior to the level of the junctions of the anterolateral margins of the supratemporal processes with the parietal plate. The ventral cranial crest is rather low and runs closely medially to the lateral margin of the parietal table. The postfoveal crest passes along the medial margin of the root portion of the supratemporal process (Fig. 38A). REMARKS The two smaller specimens MNHN.F.QU17177 and UM BFI 1873 (Fig. 42), which we herein assign to Palaeovaranus lismonimenos n. sp. were originally described by Augé (2005) who also provided drawings of both (his Figs 194 and 195 respectively) and referred them to as “ Necrosaurus eucarinatus ” (see Discussion below about the status of this name). MNHN.F.QU17177 has a length of the parietal table 11.5 mm (measured in the mid-line) (Fig. 42A, B); thus, it is smaller than the holotype parietal NHMW 2019 /0047/0001 (14.5 mm). The right supratemporal process is completely preserved in MNHN.F.QU17177; it extends posterolaterally (Fig. 42A, B). The dorsolateral crests of the specimen MNHN.F.QU17177 bear slightly developed crenulations and do not meet in the mid-line, as is also the case in the still smaller specimen UM BFI 1873 (Fig. 42C, D). In this smallest specimen UM BFI 1873, the ornamentation is weakly developed and the dorsolateral crests are still more distantly placed one to another than in MNHN.F.QU17177. We may interpret this here by the hypothesis that during ontogenetic growth, the dorsolateral crests move one to another and finally fuse together in about their posterior portions to form a median crest. As a consequence, large anterior and small posterior median triangular fields are produced (Fig. 42). If so, the anatomy of three different size stages presented and discussed herein represent the first evidence of the medial movement of the dorsolateral crests to their final fusion in the median plane in adult specimens. We suppose the same process of the origin of the similar morphology of the dorsal surface of parietal in Palaeovaranus cayluxi. Although there is no palaeovaranid frontal material in the NHMW collection, there have been previously described such elements from the Phosphorites du Quercy (Fig. 43). Augé (2005) described the frontal MNHN.F.QU17175 as belonging to “ Necrosaurus cayluxi ” and that of the specimen UM PRA 8 as “ Necrosaurus eucarinatus ”. In contrast to Palaeovaranus cayluxi, however, both these frontals have the same type of mounds as those present in the dorsal surface of the parietals of Palaeovaranus lismonimenos n. sp. (Fig. 43). To the contrary, the dorsal surfaces of all three known parietals of Palaeovaranus cayluxi have no such type of ornamentation, as that exhibited in Palaeovaranus lismonimenos n. sp. Thus, it is highly probable that these two frontals belong to Palaeovaranus lismonimenos n. sp. and we accordingly, tentatively assign them to our new species.

Pseudeumeces kyrillomethodicus n. sp. (Figs 6-14) urn:lsid:zoobank.org:act: FE001956-70BD-4493-8FD4-2F15E9A4778D HOLOTYPE. — A left dentary (NHMW 2019 /0051/0001). PARATYPE. — A right dentary (NHMW 2019/0051/0002). ETYMOLOGY. — The new species epithet “ kyrillomethodicus ” honours Kyrillos (or Cyril; Greek: Κύριλλος) and Methodios (or Methodius; Greek: Μεθόδιος), the two Byzantine brothers from Thessaloniki that were sent by the Byzantine Emperor to the area of Bratislava (city of Dovina [currently Devin]) in 863 AD, where they created the first scripture for the people of Great Moravia. The name alludes to the fact that one of us (GLG) came from Thessaloniki to Bratislava (during which project also this paper was written) and generally to our nice collaboration among the three of us, Greek and Slovak scientists. Gender is masculine. TYPE LOCALITY. — Imprecisely known locality, Phosphorites du Quercy, Department of Lot orTarn-et-Garonne, Occitanie, southern France; probably Oligocene. DIAGNOSIS. — A medium sized lacertid, morphologically similar to Pseudeumeces cadurcensis, but differing from it in the following combination of features: 1) the last posterior three or four dentary teeth are reduced (instead of only the last one or two posterior teeth being reduced); 2) the dentary is short and massive rather than anteroposteriorly long relative to its dorsoventral height (the overall morphology of dentary in Pseudeumeces cadurcensis has narrower appearance relative to that in Pseudeumeces kyrillomethodicus n. sp.); 3) the facet for the anterolateral process of the coronoid reaches around the level of the 4 th tooth position (counted from posterior) rather than reaching the level of the posteriormost tooth position (in some cases, the level between the last and penultimate tooth positions); 4) the facet for anteromedial process of the coronoid reaches the level of the 3rd tooth position (counted from posterior) rather than only the level of the posteriormost tooth position; and 5) the dorsal elevation of the posterior section of the dentary is more pronounced. DISTRIBUTION. — Oligocene, Phosphorites du Quercy, southern France. DESCRIPTION Holotype NHMW 2019 /0051/0001 (Figs 6-9) The holotype left dentary NHMW 2019/0051/0001 is the most complete specimen (Figs 6-9). The specimen measures a total length of 16 mm. It is short, massively built, and ventrally deep. In cross-section, the dentary is C-shaped. In medial aspect, the dentary has a dorsally concave appearance and it gradually widens posteriorly. The CT scan reveals that the holotype dentary bears 15 tooth positions (see Fig. 9). On the medial side, the deep Meckel’s groove is fully open and distinctly broad posteriorly. It gradually narrows anteriorly, but it is still relatively wide even in this region. It disappears posterior to the large symphyseal region. CT images show that it continues inside to the bone further anteriorly only as a small internal canal. The alveolar foramen is small, being located at the level between the 4 th and 5 th tooth positions (counting from the posterior tooth; in other words, at the level of 11 th and 12 th tooth position counted from anterior). Further anteriorly, the alveolar canal is separated from the Meckel’s groove by the intramandibular septum, which is completely fused to the bone. Meckel’s groove is roofed dorsally by the subdental shelf, which bears a facet for the splenial on its ventral side. This facet reaches the level of around the 5th tooth position (counted from anterior) and its anterior end is well stepped. Dorsally, the subdental shelf bears the sulcus dentalis. The whole shelf gradually narrows posteriorly in medial aspect. The posterodorsal portion of the dentary is inclined dorsally, being well elevated. This inclination starts at the level of the 4th tooth position (counted from posterior). Posterior to the tooth row, there is a long and robust coronoid process. It reaches highly above the level of the largest tooth crowns. On its dorsomedial side, it bears a facet for the anteromedial process of the coronoid. It reaches the level of the 3 rd tooth position (counted from posterior). The posteroventral portion of the dentary is preserved. This region is slightly curved medially, forming a short angular process. Nevertheless, this region is also slightly weathered in this specimen. The otherwise smooth lateral surface of the dentary is pierced by five labial foramina (Figs 6A; 7A). The posteriormost one is located at the level of around the 5th tooth position (counted from posterior). In the posterodorsal region of the dentary, the wedge shaped facet for the anterolateral process of coronoid is present, being, however, very shallow. It appears to reach the level of the 4 th tooth position (counted from posterior). Paratype NHMW 2019/0051/0002 (Figs 10-14) Besides the holotype described above, only a single other dentary, the paratype NHMW 2019/0051/0002, can be confidently referred to Pseudeumeces kyrillomethodicus n. sp. This is rather similar to the holotype. The paratype NHMW 2019/0051/0002 is larger and more robust than the holotype NHMW 2019/0051/0001 (Figs 10-14). The posteroventral portion of the dentary that is preserved in the holotype is not preserved here and only 12 tooth positions can be therefore observed in the paratype NHMW 2019/0051/0002 (see Fig. 14). The facet for coronoid on the lateral side is more developed in the paratype dentary if compared to the holotype. When the specimen NHMW 2019/0051/0002 was originally complete and in the case it possessed 15 tooth positions as the holotype, then the facet for the anterolateral process of the coronoid reached the level of the 5th tooth position (counted from posterior). If the tooth count was only 14, the coronoid reached the level of the 4th tooth position, the condition being identical to the holotype. However, the difference between this specimen and the holotype is the position of the alveolar foramen, being located at the level of the 9th tooth position here (counted from anterior). Dentition The dentition is pleurodont and strongly heterodont. The teeth are closely spaced. While the tooth size increases posteriorly (except for the last three or four teeth that are again small, reduced), the teeth in the anterior portion of the tooth row are small and slender. The teeth in the posterior region are robust, forming blunt cylinders (Figs 8; 12; 13). The teeth (especially those in the posterior half of the tooth row) are anteroposteriorly compressed. The tooth crowns bear delicate striations. REMARKS Pseudeumeces cadurcensis was originally established by Filhol (1877a) as a scincid of the extant genus Plestiodon Duméril & Bibron, 1839. This taxonomic opinion was subsequently followed by others (e.g., Nopcsa 1908; Kuhn 1939), although affinities with the glyptosaurine Placosaurus also appeared in the literature (Lydekker 1888b; Leenhardt 1926). Hoffstetter (1944) was the first to realize its lacertid affinities and placed it into its own genus, Pseudeumeces. So far, Pseudeumeces cadurcensis represented the only currently recognized species of this genus, as other two species that have in the past been referred to Pseudeumeces are now known to pertain to other genera or represent indeterminate lizards (i.e., Glyptosaurus walbeckensis Kuhn, 1940, which was recombined into Pseudeumeces by Estes [1983], and Pseudeumeces pouiti Augé, 1993, which was subsequently recombined to its own genus Ligerosaurus Augé, Bailon & Malfay, 2003, as Ligerosaurus pouiti by Augé et al. [2003]). We consider that the dentition and overall morphology of our new species Pseudeumeces kyrillomethodicus n. sp. appears to bear a resemblance with Pseudeumeces cadurcensis. Similarly to the case of our new taxon, the type material of Pseudeumeces cadurcensis also originates from an imprecise locality within the Phosphorites du Quercy (Filhol 1877a; for this taxon see also Augé 2005; Augé & Hervet 2009; Čerňanský & Augé 2012; Čerňanský et al. 2016a; Bolet et al. 2017). Nevertheless, the material described herein clearly exhibits several obvious differences relative to Pseudeumeces cadurcensis (see Diagnosis above). Besides the features stated in the diagnosis, there is one additional difference – the dentary tooth number in Pseudeumeces cadurcensis is usually 17 (note that the tooth number in that taxon can range from 15-17; the holotype of the species [the left dentary AMNH FARB 241A] is incomplete but the preserved portion bears seven teeth and at least five other empty tooth positions), whereas the dentary tooth number of Pseudeumeces kyrillomethodicus n. sp., based on material described here, is around 14-15. Although such small differences in tooth counts can be informative in some cases, it should be noted that the tooth number in lacertids (see e.g., Čerňanský & Syromyatnikova 2019), like virtually all lizards, should not be interpreted as absolute due to its variability. So, whether it seems that the tooth number of Pseudeumeces cadurcensis appears to be slightly higher than the new species, we cannot fully demonstrate it and we refrain from formally considering this feature as a diagnostic character. Besides the resemblance with Pseudeumeces cadurcensis discussed above, it should be noted also that the dentaries of Pseudeumeces kyrillomethodicus n. sp. slightly resemble those of Dracaenosaurus Pomel, 1846, in the following features (see Müller 2004; Augé 2005; Čerňanský et al. 2016 a, 2017): 1) dentary is a rather short, massive, and deep element; 2) the presence of a dorsally elevated posterior portion of the dentary; 3) the presence of the amblyodont dentition, where the posterior robust teeth are low and form blunt cylinders (this is more pronounced in Dracaenosaurus); and 4) the presence of striations on the tooth crown (note that the last two features are not unique to these two forms). However, there are some important differences between these two forms, where Pseudeumeces kyrillomethodicus n. sp. can be differentiated from Dracaenosaurus croizeti Gervais, 1848 -1852, by the following combination of features (for Dracaenosaurus, see Müller 2004; Augé 2005; Čerňanský et al. 2016 a, 2017): 1) the dentary tooth number is around 14-15 rather than seven or eight; 2) the tooth size increases posteriorly, however, the largest tooth is the 4th or 5th one (counted from posterior), whereas the further posterior teeth decrease in size (to the contrary, the largest tooth in D. croizeti is usually the posteriormost one or sometimes the penultimate one); 3) the alveolar foramen, although its position can vary, is located further anteriorly (at the level between the 4th and 5th tooth positions in the holotype; counted from posterior) rather than at the level of the posteriormost tooth (or between last and penultimate tooth positions); 4) the facet for the anterolateral process of coronoid reaches around the level of the 4 th tooth position (counted from posterior) rather than terminating posterior to the tooth row; and 5) teeth (especially those in the posterior half of the tooth row) are anteroposteriorly compressed rather than mediolateraly compressed. Note that for Dracaenosaurus we follow recent workers and treat Dracaenosaurus sauvagei (Filhol, 1882) as a junior synonym of the type species Dracaenosaurus croizeti (see e.g., Augé 2005). In any case, the holotype dentary of D. sauvagei is different from that of Pseudeumeces kyrillomethodicus n. sp. and its tooth count is within the range of D. croizeti. In regards to our material, the specimen MNHN.F.QU17169 (see Augé & Hervet 2009: fig. 1) deserves a comment. This specimen, which has been allocated to Pseudeumeces cadurcensis by Augé (2005) and Augé & Hervet (2009), has only 12 tooth positions instead of usual 16-17. Moreover, the dentary of this specimen appears to be robust rather than narrow. This would point to a huge level of variability. However, in MNHN.F.QU17169, only the last posterior tooth is reduced and the coronoid reaches the level of this last posterior tooth position on both sides as it is typical, indeed, for Pseudeumeces cadurcensis (in contrast to our material described herein). In general, it can be expected stratigraphically that not identical but slightly similar forms of a lineage (the exact age of our material is unfortunately unknown, as is also that of the type material of Pseudeumeces cadurcensis) would exhibit a higher degree of morphological disparity reflecting the evolution through time than specimens collected from a single stratigraphic level. In those cases, it is of course difficult to add an exact border between such forms to distinguish taxa as units for science. However, we are convinced that all the above mentioned differences allow to erect a new taxon based on our type material. Even in extant herpetofaunas, lacertids include several morphologically cryptic species for which determination based on morphology can be even more difficult than the situation discussed here. Therefore, we consider the obvious differences in our material relative to the previously described forms as sufficient. Due to a high level of similarities of the currently known limited material with Pseudeumeces cadurcensis, we decided to allocate this new taxon Pseudeumeces kyrillomethodicus n. sp. to the same genus instead of erecting a new one. It should be further noted that the extinct genera Pseudeumeces, Dracaenosaurus, and Janosikia Čerňanský, Klembara, & Smith, 2016, have been recovered as sister taxa to the extant Gallotia Boulenger, 1916, from the Canary Islands, and all these taxa together with Psammodromus Fitzinger, 1826, form the clade Gallotiinae. This was firstly observed by Čerňanský et al. (2016b, 2017), who also applied this revelation on the principles of the island rules, and later supported by Garcia-Porta et al. (2019) by their analyses based on a supermatrix relying on novel phylogenomic datasets. Therefore, we allocate Pseudeumeces kyrillomethodicus n. sp., as a member of Pseudeumeces, to the Gallotiinae as well. Recently described fossil material from the early Eocene (MP 8-9) French locality of Mutigny (Paris Basin) indicates that not only stem but also morphologically mod- ern-like (potentially crown or close to crown) lacertids were present on the European continent already in the early Eocene (Čerňanský et al. 2020). And later, lacertids were a rather diverse group during the Paleogene. The new taxon described herein, Pseudeumeces kyrillomethodicus n. sp., fully supports this high diversity and abundance of European Paleogene lacertids., Published as part of Georgalis, Georgios L., Čerňanský, Andrej & Klembara, Jozef, 2021, Osteological atlas of new lizards from the Phosphorites du Quercy (France), based on historical, forgotten, fossil material, pp. 219-293 in Geodiversitas 43 (9) on pages 227-236, DOI: 10.5252/geodiversitas2021v43a9, http://zenodo.org/record/4720776, {"references":["FILHOL H. 1877 a. - Recherches sur les Phosphorites du Quercy. Etude des fossiles qu'on y rencontre et specialement des mammiferes. Pt. II. Annales des Sciences geologiques 8: 1 - 340. https: // gallica. bnf. fr / ark: / 12148 / bpt 6 k 432584 w / f 5. item","DUMERIL A. M. C. & BIBRON G. 1839. - Erpetologie generale ou histoire naturelle complete des reptiles. Tome cinquieme. Contenant l'histoire de quatre-vingt-trois genres et de deux cent sept especes des trois dernieres familles de l'ordre des sauriens, savoir: les lacertiens, les chalcidiens et les scincoIdiens. Librairie encyclopedique de Roret, Paris, 854 p. https: // doi. org / 10.5962 / bhl. title. 45973","NOPCSA F. 1908. - Zur Kenntnis der fossilen Eidechsen. Beitrage zur Palaontologie und Geologie Osterreich-ungarns und des Orients 21: 33 - 62. https: // www. biodiversitylibrary. org / page / 15264653","KUHN O. 1939. - Squamata: Lacertilia et Ophidia. Fossilium Catalogus. I: Animalia. Pars 86. Verlag Gustav Feller, Neubrandenburg, 89 p. [Lacertilia] + 3 pp [Ophidia].","LYDEKKER R. 1888 b. - Notes on Tertiary Lacertilia and Ophidia. Geological Magazine 5: 110 - 113. https: // doi. org / 10.1017 / S 0016756800173480","LEENHARDT H. 1926. - Sur quelques sauriens de l'Eocene superieur de la France. Bulletin de la Societe geologique de France, serie 4, 26: 371 - 374.","HOFFSTETTER R. 1944. - Sur les Scincidae fossiles. I. Formes europeennes et nord-americaines. Bulletin du Museum national d'Histoire naturelle, 2 eme serie, 16: 547 - 553. https: // www. biodiversitylibrary. org / page / 54154543","ESTES R. 1983. - Sauria Terrestria, Amphisbaenia, in WELL- NHOFER P. (ed.), Encyclopedia of Paleoherpetology. Part 10 a. Gustav Fischer, Stuttgart, New York, 249 p.","AUGE M. 1993. - Une nouvelle espece de Lacertide (Reptilia, Lacertilia) des Faluns miocenes de l'Anjou-Touraine. Bulletin de la Societe de Sciences naturelles de l'Ouest de la France 15: 69 - 74.","AUGE M., BAILON S. & MALFAY J. - P. 2003. - Un nouveau genre de Lacertidae (Reptilia, Lacertilia) dans les faluns miocenes de l'Anjou- Touraine (Maine-et-Loire, France). Geodiversitas 25: 289 - 295.","AUGE M. 2005. - Evolution des lezards du Paleogene en Europe. Museum national d'Histoire naturelle, Paris: 1 - 369 (Memoires du Museum national d'Histoire naturelle; 192).","AUGE M. & HERVET S. 2009. - Fossil lizards from the locality of Gannat (late Oligocene-early Miocene, France) and a revision of the genus Pseudeumeces (Squamata, Lacertidae). Palaeobiodiversity and Palaeoenvironments 89: 191 - 201. https: // doi. org / 10.1007 / s 12549 - 009 - 0009 - 1","CERNANSKY A. & AUGE M. 2012. - Additions to the lizard fauna (Squamata: Lacertilia) of the Upper Oligocene (MP 28) of Herrlingen 8, Southern Germany. Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen 264: 11 - 19.","CERNANSKY A., KLEMBARA J. & MULLER J. 2016 a. - The new rare record of the late Oligocene lizards and amphisbaenians from Germany and its impact on our knowledge of the European terminal Palaeogene. Palaeobiodiversity and Palaeoenvironments 96: 559 - 587. https: // doi. org / 10.1007 / s 12549 - 015 - 0226 - 8","BOLET A., RAGE J. - C. & CONRAD J. L. 2017. - Rediscovery of the long-lost holotype of the lacertid lizard Pseudeumeces cadurcensis (Filhol, 1877). Journal of Vertebrate Paleontology 37: e 1315669. https: // doi. org / 10.1080 / 02724634.2017.1315669","CERNANSKY A. & SYROMYATNIKOVA E. V. 2019. - The first Miocene fossils of Lacerta cf. trilineata (Squamata, Lacertidae) with a comparative study of the main cranial osteological differences in green lizards and their relatives. PLoS ONE 14: e 0216191. https: // doi. org / 10.1371 / journal. pone. 0216191","POMEL A. 1846. - Memoire pour servir a la geologie paleontologique des terrains tertiaires du departement de l'Allier. Bulletin de la Societe geologique de France, 2 eme serie, 3: 353 - 373. https: // www. biodiversitylibrary. org / page / 54525673","MULLER J. 2004. - Cranial osteology of Dracaenosaurus croizeti, a lacertid lizard from the Oligocene of France (Reptilia, Squamata). Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen 232: 253 - 266. https: // doi. org / 10.1127 / njgpa / 232 / 2004 / 253","CERNANSKY A., BOLET A., MULLER J., RAGE J. - C., AUGE M. & HERREL A. 2017. - A new exceptionally preserved specimen of Dracaenosaurus (Squamata, Lacertidae) from the Oligocene of France as revealed by micro-computed tomography. Journal of Vertebrate Paleontology 37: e 1384738. https: // doi. org / 10.1 080 / 02724634.2017.1384738","GERVAIS P. 1848 - 1852. - Zoologie et Paleontologie francaises (animaux vertebres): ou nouvelles recherches sur les animaux vivants et fossiles de la France. Arthus Bertrand, Paris, 271 p. https: // doi. org / 10.5962 / bhl. title. 39473","BOULENGER G. A. 1916. - On the lizards allied to Lacerta muralis with an account of Lacerta agilis and L. parva. Transactions of the Zoological Society of London 21: 1 - 104. https: // doi. org / 10.1111 / j. 1096 - 3642.1916. tb 00480. x","FITZINGER L. J. F. J. 1826. - Neue Classification der Reptilien nach ihren Naturlichen Verwandtschaften. Nebst einer Verwandtschafts- Tafel und einem Verzeichnisse der Reptilien-Sammlung des k. k. zoologischen Museums zu Wien. J. G. Huebner, Wien, 66 p. https: // doi. org / 10.5962 / bhl. title. 4683","CERNANSKY A., KLEMBARA J. & SMITH K. T. 2016 b. - Fossil lizard from central Europe resolves the origin of large body size and herbivory in giant Canary Island lacertids. Zoological Journal of the Linnean Society 176: 861 - 877. https: // doi. org / 10.1111 / zoj. 12340","CERNANSKY A., AUGE M. & PHELIZON A. 2020. - Dawn of lacertids (Squamata, Lacertidae): new finds from the Upper Paleocene and the Lower Eocene. Journal of Vertebrate Paleontology 40: e 1768539. https: // doi. org / 10.1080 / 02724634. 2020.1768539"]}

Saniwa sp. (Figs 59-61) REFERRED SPECIMENS. — Three presacral vertebrae (NHMW 2019/0065/0001-NHMW 2019/0065/0003). DESCRIPTION Presacral vertebrae (Figs 59-61) The three vertebrae are large (Figs 59-61), with centrum lengths ranging from about 8.8 to 9.6 mm (seeAppendix 1). The centrum is almost triangular in ventral view and widens anteriorly (though not to that extent as in melanosaurine vertebrae described above). The subcentral ridges are straight in ventral view. The prezygapophyses are either much dorsally tilted (NHMW 2019/0065/0003) or only slightly so (in the other two specimens). The prezygapophyseal articular facets are massive and broad in dorsal view. The postzygapophyseal articular facets are also massive. The neural spine develops in height mostly in the posterior half of the neural arch. The neural arch is vaulted in posterior view. There are slight signs of “pseudozygosphene” and “pseudozygantrum” (sensu Hoffstetter 1969). The cotyle and the condyle are strongly depressed. The centrum appear more convex in lateral view is NHMW 2019/0065/0001 and NHMW 2019/0065/0002, while it is more straight in NHMW 2019/0065/0003. In all specimens though, the dorsal level of the cotyle can be clearly visible in ventral view of the specimen. Precondylar constriction can be observed (even slightly though) in NHMW 2019/0065/0001, as the respective portion of the centrum is eroded in the other two specimens. Anocotylar foramina are present and are most prominent in the largest vertebra NHMW 2019/0065/0001 (Fig. 59A). REMARKS These three vertebrae can be referred to Saniwa on the basis of their triangular centrum that widens anteriorly and the slight presence of “pseudozygosphene” and “pseudozygantrum” (Gilmore 1922; Rage & Augé 2003; Augé 2005). See Discussion below for further information on European material of Saniwa. Anguimorpha indet. (Figs 61; 62) REFERRED SPECIMENS. — Five presacral vertebrae (NHMW 2019/0046/0003- NHMW 2019/0046/0007); a partial pectoral girdle (NHMW 2019/0095/0001). DESCRIPTION AND REMARKS Presacral vertebrae (Fig. 61) These vertebrae are relatively large (Fig. 61), with centrum lengths ranging between 6.9 and 9.3 mm (see Appendix 1). The vertebrae demonstrate a mix of several features present in the above described specimens of Placosaurus, Anguinae indet., and Palaeovaranus. They have high neural spines, depressed cotyle and condyle, while the ventral surface of their centra is crossed by a wide surface or groove that is unlike the conditions seen above for the other taxa (Fig. 61). Considering the high intracolumnar variation observed in the vertebrae of extant lizards (e.g., Pseudopus), we are reluctant in assigning these specimens in a more precise taxonomic rank and we cannot even exclude that they (or part of them) pertain to some of the above described taxa. Pectoral girdle NHMW 2019/0095/0001 (Fig. 62) This specimen is incomplete, though preserving in relatively good state the right scapulocoracoid. The glenoid fossa is visible, well demarking the point of attachment with the humerus. Anteriorly to the glenoid fossa, lies the coracoid foramen. Dorsally to the foramen, the scapulocoracoid is of rectangular shape and is dorsoventrally elongated. The ventral portion of the element is anteroposteriorly elongated. It is readily obvious that this specimen apparently pertains to a rather large-sized lizard. Considering our currently inadequate state of knowledge of the appendicular skeleton of Paleogene European lizards, it is impossible to associate it with any of the above described glyptosaurines, palaeovaranids, and varanids, all of which could attain a considerably large size. Squamata indet. REFERRED SPECIMEN. — A?sacral vertebra (NHMW 2019/0095/0002). DESCRIPTION AND REMARKS. This vertebra is incomplete and not informative. Anguimorpha indet. (Figs 61; 62) REFERRED SPECIMENS. — Five presacral vertebrae (NHMW 2019/0046/0003- NHMW 2019/0046/0007); a partial pectoral girdle (NHMW 2019/0095/0001). DESCRIPTION AND REMARKS Presacral vertebrae (Fig. 61) These vertebrae are relatively large (Fig. 61), with centrum lengths ranging between 6.9 and 9.3 mm (see Appendix 1). The vertebrae demonstrate a mix of several features present in the above described specimens of Placosaurus, Anguinae indet., and Palaeovaranus. They have high neural spines, depressed cotyle and condyle, while the ventral surface of their centra is crossed by a wide surface or groove that is unlike the conditions seen above for the other taxa (Fig. 61). Considering the high intracolumnar variation observed in the vertebrae of extant lizards (e.g., Pseudopus), we are reluctant in assigning these specimens in a more precise taxonomic rank and we cannot even exclude that they (or part of them) pertain to some of the above described taxa. Pectoral girdle NHMW 2019/0095/0001 (Fig. 62) This specimen is incomplete, though preserving in relatively good state the right scapulocoracoid. The glenoid fossa is visible, well demarking the point of attachment with the humerus. Anteriorly to the glenoid fossa, lies the coracoid foramen. Dorsally to the foramen, the scapulocoracoid is of rectangular shape and is dorsoventrally elongated. The ventral portion of the element is anteroposteriorly elongated. It is readily obvious that this specimen apparently pertains to a rather large-sized lizard. Considering our currently inadequate state of knowledge of the appendicular skeleton of Paleogene European lizards, it is impossible to associate it with any of the above described glyptosaurines, palaeovaranids, and varanids, all of which could attain a considerably large size. Squamata indet. REFERRED SPECIMEN. — A?sacral vertebra (NHMW 2019/0095/0002). DESCRIPTION AND REMARKS. This vertebra is incomplete and not informative. Squamata indet. REFERRED SPECIMEN. — A?sacral vertebra (NHMW 2019/0095/0002). DESCRIPTION AND REMARKS. This vertebra is incomplete and not informative.

Genus Mediolacerta Augé, 2005 TYPE SPECIES. — Mediolacerta roceki Augé, 2005 (type species by original designation; Augé 2005)., Published as part of Georgalis, Georgios L., Čerňanský, Andrej & Klembara, Jozef, 2021, Osteological atlas of new lizards from the Phosphorites du Quercy (France), based on historical, forgotten, fossil material, pp. 219-293 in Geodiversitas 43 (9) on page 240, DOI: 10.5252/geodiversitas2021v43a9, http://zenodo.org/record/4720776, {"references":["AUGE M. 2005. - Evolution des lezards du Paleogene en Europe. Museum national d'Histoire naturelle, Paris: 1 - 369 (Memoires du Museum national d'Histoire naturelle; 192)."]}

Genus Palaeovaranus Zittel, 1887 -1890 TYPE SPECIES. — Palaeovaranus cayluxi Zittel, 1887 -1890 (type species by subsequent designation; Georgalis 2017). EMENDED GENERIC DIAGNOSIS. — 1) Presence of a distinctly developed nasal crest on the dorsomedial surface of the nasal process of maxilla; and 2) the dorsolateral crests on the dorsal surface of the parietal meeting in the median plane to form a median crest in adult specimens., Published as part of Georgalis, Georgios L., Čerňanský, Andrej & Klembara, Jozef, 2021, Osteological atlas of new lizards from the Phosphorites du Quercy (France), based on historical, forgotten, fossil material, pp. 219-293 in Geodiversitas 43 (9) on page 251, DOI: 10.5252/geodiversitas2021v43a9, http://zenodo.org/record/4720776, {"references":["ZITTEL K. A. 1887 - 1890. - Handbuch der Palaontologie. Palaeozoologie. III. Pisces, Amphibia, Reptilia, Aves. Druck und Verlag von R. Oldenbourg, Munchen, Leipzig, 900 p. https: // www. biodiversitylibrary. org / page / 40393265"]}

Paraplacosauriops quercyi (Filhol, 1882) (Figs 23-29) REFERRED SPECIMENS. — An incomplete left maxilla (NHMW 2019/0049/0003); a right dentary (NHMW 2019/0049/0001); a left dentary (NHMW 2019/0049/0002). DESCRIPTION Maxilla (Figs 23; 24) About the anterior half of the maxilla is preserved, however, its anteriormost portion is missing (Figs 23; 24). The external wall of the maxilla consists of two complete and three incomplete ornamented shields divided by more or less distinct grooves (Figs 23A; 24A). One of the shields lying between the two anterior and two posterior shields is the largest and is of pentagonal shape. The ornamentation consists of small tubercles (Fig. 23E). The ornamented surface is ventrally delimited by a shallow and narrow, anteroposteriorly running groove. Immediately ventral to the groove, the surface of the maxilla is smooth and four labial foramina are present there (Figs 23A; 24A). The smooth medial surface bears two depressions; a large one anteriorly (lacrimal recess) and a smaller one posteriorly (Figs 23B; 24B). The supradental shelf is straight and the bone immediately laterally to the posterior portion of the shelf is burrowed by the superior alveolar canal. The teeth are mesiodistally narrow and the apex is pointed. Dentaries (Figs 25-29) The right dentary NHMW 2019/0049/0001 is almost completely preserved, with only several pieces of the posteriormost portion are missing (Figs 25-28), while the left dentary NHMW 2019/0049/0002 is much more incomplete, missing both anterior and posterior portions (Fig. 29). Accordingly, we base our description on the almost complete right dentary NHMW 2019/0049/0001. The dentary is massively built. Its external surface is smooth. A distinct groove runs immediately anteroventrally to the root portion of the coronoid process (Figs 25-28). From there, a shallow sulcus runs anteriorly along the dorsal crest of the dentary. Ventrally to this sulcus, five labial foramina are present. The coronoid process is small and short. The posteriormost portion of the dentary has an almost perpendicular margin and represents a surangular process (Figs 26C, D; 28A). The most distinctive feature of the dentary is a huge alveolar canal (Figs 25-28). The intramandibular septum runs in dorsolateral-ventromedial direction and, in the place of the alveolar foramen, it is distinctly embayed anteriorly. The ventral margin of the intramandibular septum is divided from the wall of the dentary by a distinct groove (Figs 25-28). The posteroventral margin of the intramandibular septum extends to a distinct posterior spine (Figs 25-28). The dental crest extends medioventrally. The Meckel’s groove opens medioventrally. There are 22 tooth positions, while at least 10 teeth are rather well preserved (Figs 25-27). The teeth are dorsoventrally straight and they increase in size mesiodistally (Figs 25-28). The most robust are the 3rd and 4th teeth from posterior. The lingual walls of the teeth are medially bulged. The apices become mesiodistally gradually more robust and bear distinct mesiodistally straight cutting edges (Figs 25-28). The lingual and labial surfaces of the apices are distinctly striated. REMARKS The almost complete dentary NHMW 2019/0049/0001 bears strong resemblance with the neotype dentary of Para- placosauriops quercyi (MNHN.F.QU16569). Most characteristically, both specimens share the strongly heterodont dentition, with the anterior teeth being much slender and the posterior ones robust, a developed intramandibular septum, with its ventral margin fused to the floor of the Meckel’s groove (Augé & Sullivan 2006). The fragmentary dentary NHMW 2019/0049/0002, although much incomplete, seems to possess also this dental morphology. The general morphology of the preserved portion of the maxilla and the type of ornamentation correspond to those described by Augé & Sullivan (2006). The specimen NHMW 2019/0049/0001 represents the most complete dentary assigned to Paraplacosauriops quercyi and thus, enhances our understanding of this element in this melanosaurine taxon. We agree with recent authors that the species Diploglossus cadurcensis De Stefano, 1903, and Placosaurus leenhardti Leenhardt, 1926 (both from the old collections of the Phosphorites du Quercy) are junior synonyms of Paraplacosauriops quercyi (e.g., Augé 2005), although we have to highlight that the completeness of the holotype of Placosaurus leenhardti is remarkable and this taxon is certainly worth of a comprehensive redescription. Melanosaurini indet. (Figs 30-32) REFERRED SPECIMENS. — Four presacral vertebrae (NHMW 2019/0094/0001- NHMW 2019/0094/0003 and NHMW 2019/0094/0005); one sacral vertebra (NHMW 2019/0094/0004). DESCRIPTION Presacral vertebrae (Figs 30; 31) In all specimens, the centrum is significantly anteriorly widened (Figs 30; 31). Their size varies, with centrum lengths ranging between 6 and 9.4 mm (Appendix 1). There is a distinct and rather wide groove in the ventral surface of the centrum, originating anteriorly almost at the level of the cotyle and terminating posteriorly at around the level of the condyle, being almost uniform in wideness across its length. The prezygapophyses are strongly dorsally inclined in anterior view. Both cotyle and condyle are strongly dorsoventrally compressed. The neural spine, when preserved, develops mostly at the posterior half of the neural arch, however, its base extends anteriorly in the shape of a narrow longitudinal ridge until the anterior most edge of the neural arch. The height of the neural spine varies, being either high (NHMW 2019/0094/0002) or rather short (NHMW 2019/0094/0001).In one specimen (NHMW 2019/0094/0001), the posterior edge of the neural spine (as seen in dorsal view) is bifurcated. The postzygapophyses are large and extend much laterally in dorsal view. The neural canal is relatively large and triangular in shape. The shape of the neural arch in posterior view varies, apparently depending on the exact position of the vertebra in the column; as such it can be either depressed (e.g., NHMW 2019/0094/0005 and NHMW 2019/0094/0003) or relatively vaulted (e.g., NHMW 2019/0094/0002). Sacral vertebra NHMW 2019/0094/0004 (Fig. 32) The sacral vertebra NHMW 2019/0094/0004 is rather similar to the above described presacral ones, especially at the degree of the anterior widening of its centrum, the dorsally inclined prezygapophyses, and the much dorsoventrally compressed cotyle and condyle (Fig. 32). The prezygapophyses are robust. The postzygapophyses are short and do not extend significantly laterally. The neural spine is high, and is mostly developed and augmenting in height in the posterior half of the neural arch. The neural arch is moderately vaulted in posterior view. Subcentral foramina are present.Two distinct foramina, each situated between each prezygapophysis, are present above the cotyle, a structure herein defined as “anocotylar” foramina (see Remarks below). Interestingly also, this specimen is pierced by distinct foramina in the dorsal surface of its neural arch. REMARKS The referral of this vertebral material to Melanosaurini is made primarily on the basis of the much anteriorly widened centrum than in other glyptosaurines, similar to that observed for the North American Melanosaurus maximus Gilmore, 1928, and, to a lesser degree, Paraplacosauriops from the Eocene of Europe (see figures in Gilmore 1928 and Augé 2003, 2005). One other important difference between NHMW 2019/0094/0001 (but not the other melanosaurine vertebrae from our collection) and the above ones referred to Placosaurus is that the former possess much more massive postzygapophyses that extend more prominently laterally in dorsal view. A plausible taxonomic scenario could be that these specimens pertain to Paraplacosauriops quercyi described above from cranial material, however, on the absence of articulated specimens and the imprecisely known locality Meckel’s groove data (including the fact that the material was probably collected from different localities), we refrain from referring them to the same taxon. The presence of two distinct foramina above the cotyle of the sacral vertebra, a feature also prominent in several presacral vertebrae of Palaeovaranus (see below), is interesting. We acknowledge the presence of these structures in large-sized vertebrae of extant specimens of the anguid Pseudopus Merrem, 1820, and the varanid Varanus Merrem, 1820. We consider that their presence is widespread in large-sized anguimorphs and is apparently correlated with large size; we define these structures as “anocotylar foramina”, from the Greek words “ἄνω” (“ano”), meaning “above” and “κότυλος” (“cotylos”), meaning “cup”, in a similar trend of the term “paracotylar foramina”, which applies in snake vertebrae terminology. The potential taxonomic utility of anocotylar foramina needs to be further investigated in the light of detailed quantitative analyses on extant forms, as well as articulated fossil specimens. Melanosaurini indet. (Figs 30-32) REFERRED SPECIMENS. — Four presacral vertebrae (NHMW 2019/0094/0001- NHMW 2019/0094/0003 and NHMW 2019/0094/0005); one sacral vertebra (NHMW 2019/0094/0004). DESCRIPTION Presacral vertebrae (Figs 30; 31) In all specimens, the centrum is significantly anteriorly widened (Figs 30; 31). Their size varies, with centrum lengths ranging between 6 and 9.4 mm (Appendix 1). There is a distinct and rather wide groove in the ventral surface of the centrum, originating anteriorly almost at the level of the cotyle and terminating posteriorly at around the level of the condyle, being almost uniform in wideness across its length. The prezygapophyses are strongly dorsally inclined in anterior view. Both cotyle and condyle are strongly dorsoventrally compressed. The neural spine, when preserved, develops mostly at the posterior half of the neural arch, however, its base extends anteriorly in the shape of a narrow longitudinal ridge until the anterior most edge of the neural arch. The height of the neural spine varies, being either high (NHMW 2019/0094/0002) or rather short (NHMW 2019/0094/0001).In one specimen (NHMW 2019/0094/0001), the posterior edge of the neural spine (as seen in dorsal view) is bifurcated. The postzygapophyses are large and extend much laterally in dorsal view. The neural canal is relatively large and triangular in shape. The shape of the neural arch in posterior view varies, apparently depending on the exact position of the vertebra in the column; as such it can be either depressed (e.g., NHMW 2019/0094/0005 and NHMW 2019/0094/0003) or relatively vaulted (e.g., NHMW 2019/0094/0002). Sacral vertebra NHMW 2019/0094/0004 (Fig. 32) The sacral vertebra NHMW 2019/0094/0004 is rather similar to the above described presacral ones, especially at the degree of the anterior widening of its centrum, the dorsally inclined prezygapophyses, and the much dorsoventrally compressed cotyle and condyle (Fig. 32). The prezygapophyses are robust. The postzygapophyses are short and do not extend significantly laterally. The neural spine is high, and is mostly developed and augmenting in height in the posterior half of the neural arch. The neural arch is moderately vaulted in posterior view. Subcentral foramina are present.Two distinct foramina, each situated between each prezygapophysis, are present above the cotyle, a structure herein defined as “anocotylar” foramina (see Remarks below). Interestingly also, this specimen is pierced by distinct foramina in the dorsal surface of its neural arch. REMARKS The referral of this vertebral material to Melanosaurini is made primarily on the basis of the much anteriorly widened centrum than in other glyptosaurines, similar to that observed for the North American Melanosaurus maximus Gilmore, 1928, and, to a lesser degree, Paraplacosauriops from the Eocene of Europe (see figures in Gilmore 1928 and Augé 2003, 2005). One other important difference between NHMW 2019/0094/0001 (but not the other melanosaurine vertebrae from our collection) and the above ones referred to Placosaurus is that the former possess much more massive postzygapophyses that extend more prominently laterally in dorsal view. A plausible taxonomic scenario could be that these specimens pertain to Paraplacosauriops quercyi described above from cranial material, however, on the absence of articulated specimens and the imprecisely known locality Meckel’s groove data (including the fact that the material was probably collected from different localities), we refrain from referring them to the same taxon. The presence of two distinct foramina above the cotyle of the sacral vertebra, a feature also prominent in several presacral vertebrae of Palaeovaranus (see below), is interesting. We acknowledge the presence of these structures in large-sized vertebrae of extant specimens of the anguid Pseudopus Merrem, 1820, and the varanid Varanus Merrem, 1820. We consider that their presence is widespread in large-sized anguimorphs and is apparently correlated with large size; we define these structures as “anocotylar foramina”, from the Greek words “ἄνω” (“ano”), meaning “above” and “κότυλος” (“cotylos”), meaning “cup”, in a similar trend of the term “paracotylar foramina”, which applies in snake vertebrae terminology. The potential taxonomic utility of anocotylar foramina needs to be further investigated in the light of detailed quantitative analyses on extant forms, as well as articulated fossil specimens., Published as part of Georgalis, Georgios L., Čerňanský, Andrej & Klembara, Jozef, 2021, Osteological atlas of new lizards from the Phosphorites du Quercy (France), based on historical, forgotten, fossil material, pp. 219-293 in Geodiversitas 43 (9) on pages 246-250, DOI: 10.5252/geodiversitas2021v43a9, http://zenodo.org/record/4720776, {"references":["SULLIVAN R. M. & AUGE M. 2006. - Redescription of the holotype of Placosaurus rugosus Gervais 1848 - 1852 (Squamata, Anguidae, Glyptosaurinae) from the Eocene of France. Journal of Vertebrate Paleontology 26: 127 - 132. https: // doi. org / cqfr 8 h","DE STEFANO G. 1903. - I sauri del Quercy appartenenti alla collezione Rossignol. Atti della Societa Italiana di Scienze Naturali e del Museo Civili di Storia Naturale, Milan 42: 382 - 418.","LEENHARDT H. 1926. - Sur quelques sauriens de l'Eocene superieur de la France. Bulletin de la Societe geologique de France, serie 4, 26: 371 - 374.","AUGE M. 2005. - Evolution des lezards du Paleogene en Europe. Museum national d'Histoire naturelle, Paris: 1 - 369 (Memoires du Museum national d'Histoire naturelle; 192).","GILMORE C. W. 1928. - The fossil lizards of North America. Memoirs of the National Academy of Sciences 11: 1 - 197.","MERREM B. 1820. - Versuch eines systems der Amphibien, Vol. 8. J. C. Krieger, Marburg, 191 p. https: // doi. org / 10.5962 / bhl. title. 5037"]}

Anguinae indet. (Fig. 33) REFERRED SPECIMENS. — Two presacral vertebrae (NHMW 2019/0093/0001 and NHMW 2019/0093/0002). DESCRIPTION The presacral vertebrae NHMW 2019/0093/0001 and NHMW 2019/0093/0002 are almost totally complete (Fig. 33). They are relatively large, both having a centrum length of 8.4 mm. In anterior view (Fig. 33A, G), the prezygapophyses are dorsolaterally inclined. The neural canal is triangular in shape. The cotyle is exceedingly depressed. In posterior view (Fig. 33B, H), the neural arch is moderately vaulted. The condyle is rather depressed, with its ventral level being flattened. In dorsal view (Fig. 33D, G), the neural spine extends across the whole midline of the neural arch. The neural spine is relatively thickened in its posterior portion, while it is much thinner throughout its middle and anterior portions, where it takes the shape of a sharp, longitudinal ridge. The prezygapophyseal articular facets are enlarged. In ventral view (Fig. 33E, K), the centrum is widened anteriorly; its surface is flattened, with only a slight median ridge running throughout its midline. The subcentral ridges are straight; they are not parallel. Two prominent subcentral foramina pierce the centrum of NHMW 2019/0093/0001, while in the other specimen (NHMW 2019/0093/0002) these are smaller. In lateral view (Fig. 33C, F, I), the neural spine is rather short. It augments in height gradually towards the posterior portion of the neural arch, reaching its maximum height at its posteriormost portion. Its dorsal surface is straight, with its posterodorsal edge being slightly inclined posteriorly. The synapophyses are large and elongated. REMARKS These two vertebrae are strongly resembling to the ones of the genus Pseudopus on the basis of their wide centrum, being wider anteriorly, straight subcentral ridges in ventral view, and their neural spine slightly inclined posteriorly (Klembara 1979, 1981; Klembara & Rummel 2018; Čerňanský et al. 2019). Such resemblance is also supported by a biogeographic and stratigraphic rationale, as material assigned (or tentatively assigned) to Pseudopus is known in the Oligocene of Western Europe (Boettger 1875). However, it is known that at least other three non-glyptosaurine anguid genera were present in the Paleogene of Western and Central Europe, i.e., Helvetisaurus Augé, 2005, Ophisauromimus Čerňanský, Klembara & Müller, 2016, and Ophisauriscus Kuhn, 1940 (Augé 2005; Čerňanský et al. 2016a). Considering that the vertebral morphology of Ophisauromimus is currently unknown, we refrain from further assigning these two NHMW vertebrae to Pseudopus, although their overall large size, may suggest that such taxonomic referral may be most plausible., Published as part of Georgalis, Georgios L., Čerňanský, Andrej & Klembara, Jozef, 2021, Osteological atlas of new lizards from the Phosphorites du Quercy (France), based on historical, forgotten, fossil material, pp. 219-293 in Geodiversitas 43 (9) on page 251, DOI: 10.5252/geodiversitas2021v43a9, http://zenodo.org/record/4720776, {"references":["KLEMBARA J. 1979. - Neue funde der gattungen Ophisaurus und Anguis (Squamata, Reptilia) aus dem Untermiozan Westbohmens (CSSR). VestnIk UstrednIho ustavu geologickeho 54: 163 - 169.","KLEMBARA J. 1981. - Beitrag zur Kenntnis der Subfamilie Anguinae (Reptilia, Anguidae). Acta Universitatis Carolinae, Geologica 2: 121 - 168.","KLEMBARA J. & RUMMEL M. 2018. - New material of Ophisaurus, Anguis and Pseudopus (Squamata, Anguidae, Anguinae) from the Miocene of the Czech Republic and Germany and systematic revision and palaeobiogeography of the Cenozoic Anguinae. Geological Magazine 155: 20 - 44. https: // doi. org / 10.1017 / S 0016756816000753","BOETTGER O. 1875. - Ueber die Gliederung der Cyrenenmergelgruppe im Mainzer Becken. Bericht uber die Senckenbergische Naturforschende Geselschaft 1873 - 1874: 50 - 102. https: // www. biodiversitylibrary. org / page / 9351642","AUGE M. 2005. - Evolution des lezards du Paleogene en Europe. Museum national d'Histoire naturelle, Paris: 1 - 369 (Memoires du Museum national d'Histoire naturelle; 192).","CERNANSKY A., KLEMBARA J. & MULLER J. 2016 a. - The new rare record of the late Oligocene lizards and amphisbaenians from Germany and its impact on our knowledge of the European terminal Palaeogene. Palaeobiodiversity and Palaeoenvironments 96: 559 - 587. https: // doi. org / 10.1007 / s 12549 - 015 - 0226 - 8"]}

Genus Cadurcogekko Hoffstetter, 1946 TYPE SPECIES. — Cadurcogekko piveteaui Hoffstetter, 1946 (type species by original designation; Hoffstetter 1946)., Published as part of Georgalis, Georgios L., Čerňanský, Andrej & Klembara, Jozef, 2021, Osteological atlas of new lizards from the Phosphorites du Quercy (France), based on historical, forgotten, fossil material, pp. 219-293 in Geodiversitas 43 (9) on page 223, DOI: 10.5252/geodiversitas2021v43a9, http://zenodo.org/record/4720776, {"references":["HOFFSTETTER R. 1946. - Sur les Gekkonidae fossiles. Bulletin du Museum national d'Histoire naturelle, 2 eme serie, 18: 195 - 203. https: // www. biodiversitylibrary. org / page / 53794135"]}

Palaeovaranus cayluxi Zittel, 1887 -1890 (Figs 34-36; 37A, C; 41A) EMENDED DIFFERENTIAL DIAGNOSIS. — The parietal of Palaeovaranus cayluxi differs from that of the sole other recognized species of the genus, Palaeovaranus lismonimenos n. sp., described below, in the following distinguished characters and the combination of features: 1) presence of a long median crest (longer than the length of the median triangular field measured in the median plane); 2) dorsolateral crests are low and without crenulations; 3) anterior end of the dorsolateral crests disappears on the dorsal surface of the root of the anterolateral process; 4) the ornamentation is weakly developed consisting of only several low ridges of various lengths running medially to the medial margins of the dorsolateral crests, as well as small mounds; and 5) the anterior margin of the parietal fossa lies at or posterior to the level of the junctions of the anterolateral margins of the supratemporal processes with the parietal plate. REFERRED SPECIMENS. — Two almost complete parietals (NHMW 2019/0048/0001 and MNHN.F.QU17176). DESCRIPTION The parietal plate is rectangular; only the basis of the supratemporal processes is preserved in both parietals (Figs 34-36). The anterolateral process is slender. The parietal foramen lies in about the mid-length of the anterior half of the parietal plate. The most distinctive feature of the dorsal surface of the parietal are two dorsolateral crests. The crests run in anterolateral-posteromedial direction. The anterior end of each crest gradually diminishes and terminates on the dorsal surface of the anterolateral process. The posterior ends of the dorsolateral crests meet in the median plane. The crests, together with the anterior margin of the parietal, limit a triangular field containing the parietal foramen. The surface bears several low mounds and more or less long ridges running along the medial margins of the dorsolateral crests. From the junction of the dorsolateral crests, a median crest extends posteriorly. The length of the crest increases with the size of the parietal and it seems that this increase in length comes to the negative expense of the midline length of the anterior triangular surface, which gradually throughout ontogeny becomes proportionally shorter.This can be demonstrated by comparing the largest known parietal of this species (the one figured by Rage 1978) relative to the two ones described in our paper. From the posterior end of the median crest, a median triangular field is located. The triangular field is a space between the posteriormost portions of the dorsolateral crests. The triangular field achieves its largest width posteriorly; its posterior end is the posteromedian margin of the parietal table (Figs 34-36). The supratemporal fossa is mediolaterally broad indicating a strongly developed adductor musculature. The ventral surface of the parietal is smooth (Figs 34B; 36B; 37C). The anterior margin of the parietal fossa lies at the level (or posterior to the level in large specimens) of the junctions of the anterolateral margins of the supratemporal processes with the parietal table. The ventral cranial crest is low and runs immediately medially to the lateral margin of the parietal. Its posterior end is turned posteromedially. The length of the juxtaotic and postfoveal crests is about the same. The posterior portion of the postfoveal crest runs immedi- ately laterally to the medial margin of the basal portion of the supratemporal process. REMARKS Although we acknowledge that the holotype of Palaeovaranus cayluxi is a maxilla (see Georgalis 2017 for details), we assign these parietals to the same species on the basis of the referral of a parietal by Rage (1978) to the same species (see Discussion below for details). The so far three known parietals of this species (the two ones documented herein plus the one described by Rage [1978]) enhance our understanding of the parietal morphology and variation in this species and allow a confident distinguishment from its new congeneric species described below., Published as part of Georgalis, Georgios L., Čerňanský, Andrej & Klembara, Jozef, 2021, Osteological atlas of new lizards from the Phosphorites du Quercy (France), based on historical, forgotten, fossil material, pp. 219-293 in Geodiversitas 43 (9) on pages 252-253, DOI: 10.5252/geodiversitas2021v43a9, http://zenodo.org/record/4720776, {"references":["ZITTEL K. A. 1887 - 1890. - Handbuch der Palaontologie. Palaeozoologie. III. Pisces, Amphibia, Reptilia, Aves. Druck und Verlag von R. Oldenbourg, Munchen, Leipzig, 900 p. https: // www. biodiversitylibrary. org / page / 40393265","RAGE J. - C. 1978. - Squamates, in GEZE B., RAGE J. - C., VERGNAUD- GRAZZINI F., DE BROIN F., BUFFETAUT E., MOURI- ER- CHAUVIRE C., CROCHET J. - Y., SIGE B., SUDRE J., REMY A., LANGEBADRE L., BONIS L. DE, HARTENBERGER J. L. & VIANEY- LI- AUD M. (eds), La poche a Phosphate de Ste-Neboule (Lot) et sa faune de vertebres du Ludien superior. Palaeovertebrata 8: 201 - 215."]}

Genus Pseudeumeces Hoffstetter, 1944 TYPE SPECIES. — Plestiodon cadurcense Filhol, 1877 (type species by original designation; Hoffstetter 1944)., Published as part of Georgalis, Georgios L., Čerňanský, Andrej & Klembara, Jozef, 2021, Osteological atlas of new lizards from the Phosphorites du Quercy (France), based on historical, forgotten, fossil material, pp. 219-293 in Geodiversitas 43 (9) on page 227, DOI: 10.5252/geodiversitas2021v43a9, http://zenodo.org/record/4720776, {"references":["HOFFSTETTER R. 1944. - Sur les Scincidae fossiles. I. Formes europeennes et nord-americaines. Bulletin du Museum national d'Histoire naturelle, 2 eme serie, 16: 547 - 553. https: // www. biodiversitylibrary. org / page / 54154543"]}

Genus Saniwa Leidy, 1870 TYPE SPECIES. — Saniwa ensidens Leidy, 1870 (type species by original designation; Leidy 1870)., Published as part of Georgalis, Georgios L., Čerňanský, Andrej & Klembara, Jozef, 2021, Osteological atlas of new lizards from the Phosphorites du Quercy (France), based on historical, forgotten, fossil material, pp. 219-293 in Geodiversitas 43 (9) on page 269, DOI: 10.5252/geodiversitas2021v43a9, http://zenodo.org/record/4720776, {"references":["LEIDY J. 1870. - Descriptions of Emys jeanesi, E. haydeni, Baena arenosa, and Saniwa ensidens. Proceedings of the Academy of Natural Sciences, Philadelphia 22: 123 - 124. https: // www. biodiversitylibrary. org / page / 26300494"]}

Bavarioboa cf. hermi Szyndlar & Schleich, 1993 (Fig. 7) Bavarioboa cf. hermi – Ivanov & Musil 2004: 228, 229, fig. 3C, D. — Ivanov et al. 2006: 229, table 2. MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 1/2001 Turtle Joint: One posterior trunk vertebra (Pal. 1448). DESCRIPTION Trunk vertebra The only preserved trunk vertebra is almost complete with partial damage to the large paradiapophyses, the cranial margin of the zygosphene and the caudal margin of the condyle. In lateral view, the neural spine rises at about the level of the zygosphenal base. The cranial margin of the neural spine is inclined caudally with a rounded anterodorsal margin. The articular surfaces of the zygosphene are widely oval. The short interzygapophyseal ridges are rather sharp. The small lateral foramina, situated close below the interzygapophyseal ridges, do not sit within depressions. The short and dorsally bent subcentral ridges are well-developed, especially in the anterior half of the centrum. In dorsal view, the partially damaged prezygapophyseal articular facets were originally broadly subtriangular to oval in outline. The prezygapophyseal processes are rather short (about a quarter of the prezygapophyseal facets length). The zygosphene was almost straight with small lateral lobes; however, the medial part of the zygosphenal lip is rather damaged in the preserved specimen. The dorsal margin of the neural spine becomes thick towards its caudal margin. The caudal margin of the neural arch forms a relatively shallow notch. In ventral view, the haemal keel is laterally wide with a rounded ventral surface. The subcentral foramina are rather small and developed on either side of the haemal keel base. The subcentral grooves are wide and deep. The postzygapophyseal articular facets are subrectangular in shape. In cranial view, the prezygapophyses are tilted slightly dorsally. Prezygapophyseal articular facets are situated high above the neural canal base roughly at the level of the dorsal margin of the lateral sinuses. The cranial margin of the zygosphene is slightly concave with strongly build lateral sinuses and a rather thin central part. The neural arch is vaulted. The neural canal is rounded with rather wide and shallow lateral sinuses. Depressions are developed on either side of the slightly dorsoventrally depressed cotyle. The paracotylar foramina are absent. In caudal view, the zygantrum is wide. The condyle is slightly depressed dorsoventrally. The vertebral dimensions are as follows (Pal. 1448): cl: = 5.19 mm; naw = 6.73 mm; cl/naw = 0.77. REMARKS The massive structure of vertebra, the cl/naw ratio Bavarioboa by the almost straight cranial margin of the zygosphenal lip in dorsal view, as well as the dorsally thickened neural spine that is typical for the posterior trunk vertebrae of Bavarioboa hermi (see Szyndlar & Rage 2003). B. ultima Szyndlar & Rage, 2003 from the German late early Miocene Rothenstein 13 locality (late Burdigalian, MN 5), originally assigned to B. hermi by Szyndlar & Schleich (1993), differs mainly by the lower neural spine as well as longer prezygapophyseal processes (Szyndlar & Rage 2003)., Published as part of Ivanov, Martin, Čerňanský, Andrej, Bonilla-Salomón, Isaac & Luján, Àngel Hernández, 2020, Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications, pp. 343-376 in Geodiversitas 42 (20) on pages 356-357, DOI: 10.5252/geodiversitas2020v42a20, http://zenodo.org/record/4447563, {"references":["SZYNDLAR Z. & SCHLEICH H. H. 1993. - Description of Miocene snakes from Petersbuch 2 with comments on the Lower and Middle Miocene ophidian faunas of Southern Germany. Stuttgarter Beitrage zur Naturkunde Serie B (Geologie und Palaontologie) 192: 1 - 47.","IVANOV M. & MUSIL R. 2004. - Preliminary results of investigation of Neogene vertebrates from the Mokra-Quarry site. Acta Musei Moraviae, Scientiae geologicae 89: 223 - 236 (in Czech).","IVANOV M., MUSIL R. & BRZOBOHATY R. 2006. - Terrestrial and Marine Faunas from the Miocene Deposits of the Mokra Plateau (Drahany Upland, Czech Republic) - Impact on Palaeogeography. Beitrage zur Palaontologie 30: 223 - 239.","SZYNDLAR Z. & RAGE J. - C. 2003. - Non-erycine Booidea from the Oligocene and Miocene of Europe. Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Krakow, 109 p."]}

‘ BOOIDEA’ indet. MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 1/2001 Turtle Joint: three trunk vertebrae (Pal. 1451-1453); 2/2003 Reptile Joint: five trunk vertebrae (Pal. 1951-1955). DESCRIPTION Trunk vertebrae The preserved trunk vertebrae are too fragmentary. Their massive structure, combined with the wide haemal keel as well as the absence of paracotylar foramina enables assignment only to the indeterminate ‘Booidea’., Published as part of Ivanov, Martin, Čerňanský, Andrej, Bonilla-Salomón, Isaac & Luján, Àngel Hernández, 2020, Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications, pp. 343-376 in Geodiversitas 42 (20) on page 358, DOI: 10.5252/geodiversitas2020v42a20, http://zenodo.org/record/4447563

VIPERINAE (‘ Oriental vipers’ group) (Fig. 14 A-N; Fig. 15 A-J) Vipera sp. 1 (‘ Oriental vipers’) – Ivanov & Musil 2004: 230. Vipera sp. (‘ Oriental vipers’ group) – Ivanov et al. 2006: 229, table 2. MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 1/2001 Turtle Joint: one left maxilla (Pal. 1501), isolated fang (Pal. 1502), 30 trunk vertebrae (Pal. 1503-1532). 2/2003 Reptile Joint: 28 trunk vertebrae (Pal. 1994-2021). DESCRIPTION Maxilla Only a left maxilla is preserved (Fig. 14A, B). In rostral view, the body of the bone is high, especially along its medial margin. The ascending process is long, and it is inclined medially. A small process for the prefrontal connection occurs medially at its distal termination. A ridge extends from the base of the ascending process as far as its distal termination. A large, slightly oval, foramen occurs opposite to the medially directed maxillary-prefrontal process. In caudal view, a wide groove for connection with the ectopterygoid occurs above the base of the fangs. This groove is deep, and it is restricted dorsally by a transverse ridge. This ridge extends from the medial margin of the bone (where it forms two small processes) as far as the middle of the ascending process width. In medial view an oval foramen occurs within the deep, wide orifice. This orifice is restricted dorsally by a ridge. Dentition A single fang of about 1 cm length is preserved (Fig. 14C). It is long and slender, and is curved slightly caudally. A canal extends within the tooth and has a strongly elongated/slit-like orifice situated mesially, close to the pointed tip of the fang. The slightly widened basal portion of the fang is damaged and does not preserve the orifice of the venom canal. Trunk vertebrae Numerous fragmentary trunk vertebrae (Fig. 14 D-N; Fig. 15 A-J) are preserved, mostly with broken-off neural spines and hypapophyses. In lateral view, the neural spine was about as high as long in anterior precaudal vertebrae. The most complete neural spine lacks its cranial margin, but it seems possible that it was vertical or slightly anteriorly inclined. The interzygapophyseal ridges are welldeveloped and sometimes rather sharp. Lateral foramina are large and are situated in shallow depressions. The parapophyses are well-separated from the diapophyses and the parapophyseal processes are strongly built and directed antero-ventrally. The subcentral ridges are conspicuous. The condyle is developed on a very short neck. The rarely preserved hypapophyses of anterior trunk vertebrae are long, straight, and directed posteroventrally. The pointed distal termination of the hypapophysis of middle and posterior trunk vertebrae is directed caudally. The hypapophysis of the posteriormost trunk vertebrae have indication of bifurcation on its distal tip. In dorsal view, the vertebrae are markedly short and wide. The cranial margin of the zygosphenal lip is concave, straight, or with a small medial lobe. The medial lobe is better developed in posterior trunk vertebrae. The prezygapophyseal articular facets are widely oval to subtriangular (although partial damage cannot be excluded). The prezygapophyseal processes are broken-off close to their bases. Epizygapophyseal spines are absent. In ventral view, the subcentral grooves are shallow, and the large subcentral foramina are situated on both sides of the base of the wide hypapophysis. The ventromedial margin of parapophyseal processes is always medially enlarged and it is usually fused with the ventrolateral extensions of the cotylar rim (subcotylar tubercles). The blood vessels of the circulatory system passed through the canals on both sides of the hypapophyseal base. The postzygapophyseal articular facets have an irregularly triangular outline. In cranial view, the neural arch is strongly flattened dorsoventrally. The cranial margin of the zygosphene is straight. The prezygapophyses are tilted up dorsally. The paracotylar foramina are situated on both sides of the rounded cotyle. The subcotylar tubercles are usually fused with bases of parapophyseal processes. The vertebral dimensions of the largest vertebrae from 1/2001 Turtle Joint are as follows (n = 7): cl: or = 6.98-7.76 mm; naw: or = 6.31-7.03 mm; cl/ naw: or = 1.02-1.15, mean 1.09 ± 0.05. REMARKS The maxilla partially resembles that of the ‘ xanthina ’ clade of Montivipera in the shape of the medial margin of the body of maxilla. The maxilla of Viperinae (‘Oriental vipers’ group) differs from that of extinct Macrovipera gedulyi (Bolkay, 1913) (for current generic allocation see Cordea et al. 2017) in the medially strongly inclined ascending process. However, the presence of a sharp ridge situated on the rostromedial margin of the process, as well as the single orifice of the dental canal being located close to the distal termination of the process typically occur in this extinct species (Szyndlar & Rage 2002; Cordea et al. 2017). A study at the late Miocene (MN 13) Polgárdi site in Hungary shows that maxilla morphology is highly variable in M. gedulyi (Szyndlar & Rage 2002: 421, fig. 4). Findings of venomous fangs are rarely discussed in palaeoherpetological literature because isolated teeth do not allow identification even at the subfamily level (Szyndlar & Rage 2002). However, the venom fang is typified by its large dimensions, and the maxilla as well as almost all the viperid vertebrae at MWQ belonged to ‘Oriental vipers’. Therefore, it is almost certain that the isolated venom fang belonged to a large ‘Oriental viper’. The large massive vertebrae with a low cl/naw ratio as well as long,straight hypapophyses and strongly dorsoventrally depressed neural arches enable identification of vertebrae as belonging to ‘ Oriental vipers’. Distinct subcotylar tubercles are not usually observed in recent ‘ Oriental vipers’ (Szyndlar & Rage 1999; pers. observation) but in fossil representatives, at least small subcotylar tubercles frequently occur (Szyndlar 1988; Zerova 1992; Szyndlar & Rage 1999). However, the fusion of the ventromedial margin of the parapophyseal processes with strongly developed subcotylar tubercles has not been observed in either extant or extinct ‘ Oriental vipers’. A medial elongation of the medial margin of the parapophyseal processes was reported in posteriormost trunk vertebrae just anterior to the cloacal region, e.g. in Macrovipera ukrainica (Zerova, 1992) (Zerova 1992: fig. 10). The strange development of the parapophyseal region in trunk vertebrae of ‘ Oriental vipers’ from MWQ is probably not due to intraspecific variation or abnormal (pathological) development because the same vertebrae have been reported from the coeval (MN 4), still unpublished, locality 3/ 2005 in Mokrá-Central Quarry. Although ‘ Oriental viper’ from MWQ is not identified below the subfamily level, it is probable that this true viper could represent the genus Macrovipera. If it is true, the first occurrence of the genus Macrovipera could be placed to the late early Miocene., Published as part of Ivanov, Martin, Čerňanský, Andrej, Bonilla-Salomón, Isaac & Luján, Àngel Hernández, 2020, Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications, pp. 343-376 in Geodiversitas 42 (20) on pages 365-369, DOI: 10.5252/geodiversitas2020v42a20, http://zenodo.org/record/4447563, {"references":["IVANOV M. & MUSIL R. 2004. - Preliminary results of investigation of Neogene vertebrates from the Mokra-Quarry site. Acta Musei Moraviae, Scientiae geologicae 89: 223 - 236 (in Czech).","IVANOV M., MUSIL R. & BRZOBOHATY R. 2006. - Terrestrial and Marine Faunas from the Miocene Deposits of the Mokra Plateau (Drahany Upland, Czech Republic) - Impact on Palaeogeography. Beitrage zur Palaontologie 30: 223 - 239.","BOLKAY S. J. 1913. - Additions to the fossil herpetology of Hungary from the Pannonian and Praeglacial periode. Mitteilungen aus dem Jahrbuch der koniglichen ungarischen Geologischen Reichsanstalt 21: 217 - 230.","CORDEA V., VENCZEL M., URSACHI L. & RATOI B. 2017. - A large viper from the early Vallesian (MN 9) of Moldova (E-Romania) with notes on the palaeobiogeography of late Miocene ' Oriental vipers'. Geobios 50: 401 - 411. https: // doi. org / 10.1016 / j. geobios. 2017.07.001","SZYNDLAR Z. & RAGE J. - C. 2002. - Fossil record of the true viper, in SCHUETT G. W., HOGGREN M., DOUGLAS M. E. & GREENE H. W. (eds), Biology of the Vipers. Eagle Mountain Publishing, Eagle Mountain: 419 - 444.","SZYNDLAR Z. & RAGE J. - C. 1999. - Oldest fossil vipers (Serpentes: Viperidae) from the Old World. Kaupia 8: 9 - 20.","SZYNDLAR Z. 1988. - Two new extinct species of the genera Malpolon and Vipera (Reptilia, Serpentes) from the Uppermost Miocene of Algora (Spain). Acta Zoologica Cracoviensia 31 (11): 687 - 706.","ZEROVA G. A. 1992. - Vipera (Daboia) ukrainica - a new viper (Serpentes; Viperidae) from the Middle Sarmatian (Upper Miocene). Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen 184: 235 - 249."]}

Ophisaurus sp. (Fig. 6 A-E) MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 1/2001 Turtle Joint: one trunk vertebra (Pal. 1407). DESCRIPTION Trunk vertebra The vertebra is robustly built. The neural spine starts to rise dorsally from the area of anterior region of the neural arch. However, the dorsal portion of the neural spine is damaged. In lateral aspect, the neural spine reaches posteriorly only to the level of the anterior end of the condyle. The oval, tunnel-like neural canal is medium-sized. Pre- and postzygapophyses are well expanded laterally,but the interzygapophyseal constriction is relatively shallow, notas deep as it is inthe Pseudopus vertebrae described above.This gives Pal.1407 a broad appearance in dorsal aspect. The prezygapophyses are dorsally inclined at an angle of around 33°.The articular surfaces are large, elliptical and oblong laterally. In anterior aspect, a pair of foramina is located ventral to the prezygapophyses, approximately at the level of the dorsal margin of the cotyle.In lateral aspect,the postzygapophyses reach posteriorly approximately to the level of the mid-region of the condyle.Both condyle and cotyle are depressed.The synapophyses are well-developed, large, and located close to the anterior edge of the neural arch pedicel. They are slightly posterolaterally oriented. The ventral region of the centrum is flat, although it has two visible edges which run from the level of the cotyle and gradually weaken posteriorly. A pair of subcentral foramina is located close to the cotyle. The subcentral ridges are concave. REMARKS The trunk vertebra described here can be allocated to Ophisaurus based on the following features (Čerňanský et al. 2019): 1) the lateral margins of the centrum (subcentral ridges) are concave; and 2) the height of the neural canal is greater than the height of the cotyle., Published as part of Ivanov, Martin, Čerňanský, Andrej, Bonilla-Salomón, Isaac & Luján, Àngel Hernández, 2020, Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications, pp. 343-376 in Geodiversitas 42 (20) on page 354, DOI: 10.5252/geodiversitas2020v42a20, http://zenodo.org/record/4447563, {"references":["CERNANSKY A., YARYHIN O., CICEKOVA J., WERNEBURG I., HAIN M. & KLEMBARA J. 2019. - Vertebral Comparative Anatomy and Morphological Differencesin Anguine Lizards with a Special Reference to Pseudopus apodus. The Anatomical Record 302 (2): 232 - 257. https: // doi. org / 10.1002 / ar. 23944"]}

LACERTOIDEA indet. (Fig. 4A, B) Lacerta sp., small form – Ivanov et al. 2006: 229, table 2 (in part). MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 2/2003 Reptile Joint: two left dentaries (Pal. 1571, 1572). DESCRIPTION Dentary The preserved fragments represent the anterior and mid-portion of the two left dentaries. Meckel’s groove is fully open, although narrow in this region. It is roofed by a subdental shelf. The shelf gradually narrows posteriorly, caused by the presence of the facet for the splenial. This facet is situated on the ventral margin of the shelf. This facet reaches anteriorly to the level of the tenth tooth position (counted from anterior).The shelf is only slightly concave, so the small symphyseal region is only weakly elevated dorsally if compared to the posteriorly located shelf. A sulcus dentalis is present. The preserved portion of the dentary bears sixteen and half tooth positions (eight teeth are attached). The dental crest, which supports the teeth, is high(higherthan the ventrally located Meckel’s groove).Except for four labial foramina,which pierce the lateral side of the bone in its mid-line,the external surface is smooth. Dentition The implantation is pleurodont.Teeth are closely spaced with small interdental gaps. All teeth are badly preserved, with heavily weathered tooth crowns. REMARKS Unfortunately, the poor preservation does not allow a more precise determination of this specimen. The heavily weathered tooth crown might be the result of predation and digestion, especially by birds of prey. This dentary very likely represents a lacertid, but this cannot be fully demonstrated. If such an allocation is correct, then it resembles the material described here as Lacertidae indet. tooth morphotype 2 rather than 1. The main similarity is that the interdental gaps of teeth here are small and the dental crest is high., Published as part of Ivanov, Martin, Čerňanský, Andrej, Bonilla-Salomón, Isaac & Luján, Àngel Hernández, 2020, Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications, pp. 343-376 in Geodiversitas 42 (20) on pages 349-350, DOI: 10.5252/geodiversitas2020v42a20, http://zenodo.org/record/4447563, {"references":["IVANOV M., MUSIL R. & BRZOBOHATY R. 2006. - Terrestrial and Marine Faunas from the Miocene Deposits of the Mokra Plateau (Drahany Upland, Czech Republic) - Impact on Palaeogeography. Beitrage zur Palaontologie 30: 223 - 239."]}

“ NATRICINAE ” indet. (Fig. 12 F-I) cf. Neonatrix sp. – Ivanov et al. 2006: 229, table 2. MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 1/2001 Turtle Joint: One trunk vertebra (Pal. 1483). 2/2003 Reptile Joint: Four trunk vertebrae (Pal. 1970-1973). DESCRIPTION Trunk vertebrae The most complete vertebra (Pal. 1483) has strongly damaged prezygapophyses and the zygosphene, and the neural spine is broken-off close to its base.In lateral view,the interzygapophyseal ridges are moderately developed and lateral foramina are situated within deep depressions.The subcentral ridges are nearly straight. The distal termination of sigmoid hypapophysis does not reach posterior to the caudal border of condyle. In dorsal view, the vertebra is cylindrical, the damaged zygosphene was wide relative to the neural arch width. Epizygapophyseal ridges are absent. In ventral view, the subcentral grooves are very shallow and short and the subcentral foramina are rather small and situated just posterior to the base of the hypapophysis. The subcotylar tubercles are missing.The damaged postzygapophyseal articular facets have a subrectangular outline and are not elongated laterally. In cranial view, the neural arch is slightly vaulted, and the large neural canal has developed conspicuously large and wide lateral sinuses.The cranial margin of the zygosphene was arched dorsally.The small paracotylar foramina are situated within wide depressions on either side of the rounded cotyle. The vertebral dimensions of the best-preserved specimen (Pal. 1483) are: cl = 3.51 mm; naw = 2.24 mm; cl/naw = 1.57. REMARKS The sigmoid shape of the hypapophysis, with an anterior keel sloping towards the cotylar rim is reminiscent of precaudal vertebrae of the genus Palaeonatrix Szyndlar, 1982 (seeMłynarski et al. 1982) reported from the early Miocene (MN 4) of Dolnice, Czech Republic (Rage & Roček 1983; Szyndlar 1987), Petersbuch 2 and Langenau, Germany (Szyndlar & Schleich 1993), Oberdorf, Austria (Szyndlar 1998) and the middle Miocene of Sansan, France (MN 6; Augé & Rage 2000) and Opole, Poland (MN 7+8; Młynarski et al. 1982). However, a generic allocation is impossible because the relatively low neural spine which characterizes the genus Palaeonatrix (Szyndlar 1987, 1991b) is not preserved in studied specimens from MWQ. Although the very strongly developed subcentral ridges are typical for the genus Palaeonatrix (Szyndlar 1987) we cannot exclude the possibility that the subcentral ridges were shorter and less distinct in vertebrae situated anterior to the preserved vertebra. Because of the limited fossil material and lack of information on intracolumnar variability, a more precise assignment is not possible., Published as part of Ivanov, Martin, Čerňanský, Andrej, Bonilla-Salomón, Isaac & Luján, Àngel Hernández, 2020, Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications, pp. 343-376 in Geodiversitas 42 (20) on page 365, DOI: 10.5252/geodiversitas2020v42a20, http://zenodo.org/record/4447563, {"references":["IVANOV M., MUSIL R. & BRZOBOHATY R. 2006. - Terrestrial and Marine Faunas from the Miocene Deposits of the Mokra Plateau (Drahany Upland, Czech Republic) - Impact on Palaeogeography. Beitrage zur Palaontologie 30: 223 - 239.","RAGE J. - C. & ROCEK Z. 1983. - Dolniceophis lehmani (Serpentes, Colubridae), a new fossil snake from the Lower Miocene of Czechoslovakia. Casopis pro mineralalogii a geologii 28: 17 - 21.","SZYNDLAR Z. 1987. - Snakes from the Lower Miocene locality of Dolnice (Czechoslovakia). Journal of Vertebrate Paleontology 7 (1): 55 - 71. https: // doi. org / 10.1080 / 02724634.1987.10011637","SZYNDLAR Z. & SCHLEICH H. H. 1993. - Description of Miocene snakes from Petersbuch 2 with comments on the Lower and Middle Miocene ophidian faunas of Southern Germany. Stuttgarter Beitrage zur Naturkunde Serie B (Geologie und Palaontologie) 192: 1 - 47.","SZYNDLAR Z. 1998. - Vertebrates from the Early Miocene lignite deposits of the opencast mine Oberdorf (Western Styrian Basin, Austria). Annalen des Naturhistorischen Museums inWien A 99: 31 - 38.","AUGE M. & RAGE J. - C. 2000. - Les Squamates (Reptilia) du Miocene moyen de Sansan (Gers, France), in GINSBURG L. (ed.), La faune miocene de Sansan et son environnement. Museum national d'Histoire naturelle, Paris: 263 - 313 (Memoires du Museum national d'Histoire naturelle; 183).","MLYNARSKI M., SZYNDLAR Z., ESTES R. & SANCHIZ B. 1982. - Lower vertebrate fauna from the Miocene of Opole (Poland). Estudios geologicos 38 (1 - 2): 103 - 119.","SZYNDLAR Z. 1991 b. - A rewiew of Neogene and Quaternary snakes of Central and Eastern Europe. Part II: Natricinae, Elapidae, Viperidae. Estudios geologicos 47 (3 - 4): 237 - 266. https: // doi. org / 10.3989 / egeol. 91473 - 4422"]}

Natrix sp. (Fig. 12 A-E) Natrix sp. – Ivanov & Musil 2004: 229. Natrix sp., type I – Ivanov et al. 2006: 229, table 2 (in part). MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 1/2001 Turtle Joint:Two trunk vertebrae (Pal. 1481-1482); 2/2003 Reptile Joint: Two trunk vertebrae (Pal. 1968-1969). DESCRIPTION Trunk vertebrae The most complete specimen, Pal. 1968, is fragmentary with the hypapophysis and right prezygapophyseal process broken off, and strongly damaged paradiapophyses (Fig. 12 A-E). The vertebral centrum is cylindrical. In lateral view, the interzygapophyseal ridges are moderately developed. The completely preserved neural spine is about twice as long as high. Its cranial margin is inclined anteriorly whereas the caudal margin is inclined posteriorly. The small lateral foramina are situated in shallow depressions. The well-developed subcentral ridges are arched dorsally. In dorsal view, the prezygapophyseal articular facets are almost oval.The prezygapophyseal processes are about two thirds of the prezygapophyseal facets length. The cranial margin of the mostly damaged zygosphene has developed a wide medial lobe and small lateral lobes. The epizygapophyseal spines are absent. In ventral view, the hypapophysis expands laterally in a cranial direction to form a triangular anterior keel. The very small subcentral foramina are situated on both sides of the hypapophysis.In cranial view, the neural arch is slightly vaulted, and the neural canal is rounded with short lateral sinuses. The parapophyseal processes are separated from the rounded cotyle by deep furrows.The paracotylar foramina are situated on both sides of the cotyle. The left paracotylar foramen is doubled in Pal. 1968. The small paracotylar tubercles occur at the ventral margin of the cotyle. The vertebral dimensions of the figured specimen (Pal. 1968) are as follows: cl = 4.60 mm; naw = 2.72 mm; cl/naw = 1.69. The largest specimen (Pal.1481) measures as follows: cl = 4.93 mm; naw = 2.85 mm; cl/naw = 1.73. REMARKS Assignment to the “natricine” snakes is based on the presence of hypapophyses in precloacal vertebrae, the vaulted neural arch, the high neural spine, and the presence of paracotylar foramina on either side of the cotyle. The most complete vertebra with laterally directed prezygapophyses, an elongated centrum with a triangular anterior keel on the hypapophysis, and a neural spine inclined both anteriorly and posteriorly permit identification of the preserved vertebrae to the genus Natrix whose fossil remains are abundant in Central Europe as early as the early Miocene (Ivanov 2002a; Čerňanský et al. 2015). Natrix sp. differs from early Miocene N. merkurensis Ivanov, 2002 (MN 3a-?MN 4; Ivanov 2002a; Rage & Bailon 2005) in its smaller dimensions, elongated prezygapophyseal facets, shorter prezygapophyseal processes, and more distinct lateral lobes of the zygosphene. It differs from N. sansaniensis (Lartet, 1851), reported from the early and middle Miocene (MN 3-?MN 7+8; Augé & Rage 2000; Ivanov 2000, 2002a, b), in the less vaulted neural arch and flattened prezygapophyseal processes (Szyndlar 2005). N. longivertebrata Szyndlar, 1984 reported from the late Miocene (MN 9, MN 10/11; Ivanov 1997; Tempfer 2005) and Pliocene (MN 14-MN 16; Szyndlar 1984, 1991b; Venczel 2001), and doubtfully as early as the middle Miocene (?MN 6-?MN 7+8; Rage & Szyndlar 1986; Szyndlar 1991c), has more elongated vertebrae and a much lower neural spine. Natrix sp. closely resembles the extinct N. rudabanyaensis Szyndlar, 2005 reported from the early late Miocene of Rudabánya, Hungary (MN 9a; Szyndlar 2005) and perhaps the middle Miocene of Tauţ, Romania (MN 7+8; Venczel & Ştiucă 2008) in: 1) the same height and shape of the neural spine; 2) moderately developed subcentral ridges; 3) elongated prezygapophyseal facets; and 4) flattened prezygapophyseal processes (Szyndlar 2005). However, the absence of the hypapophysis and paradiapophyses prevents alpha taxonomy., Published as part of Ivanov, Martin, Čerňanský, Andrej, Bonilla-Salomón, Isaac & Luján, Àngel Hernández, 2020, Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications, pp. 343-376 in Geodiversitas 42 (20) on page 363, DOI: 10.5252/geodiversitas2020v42a20, http://zenodo.org/record/4447563, {"references":["IVANOV M. & MUSIL R. 2004. - Preliminary results of investigation of Neogene vertebrates from the Mokra-Quarry site. Acta Musei Moraviae, Scientiae geologicae 89: 223 - 236 (in Czech).","IVANOV M., MUSIL R. & BRZOBOHATY R. 2006. - Terrestrial and Marine Faunas from the Miocene Deposits of the Mokra Plateau (Drahany Upland, Czech Republic) - Impact on Palaeogeography. Beitrage zur Palaontologie 30: 223 - 239.","IVANOV M. 2002 a. - The oldest known Miocene snake fauna from Central Europe: Merkur-North locality, Czech Republic. Acta Palaeontologica Polonica 47 (3): 513 - 534.","CERNANSKY A., RAGE J. C. & KLEMBARA J. 2015. - The Early Miocene squamates of Amoneburg (Germany): the first stages of modern squamates in Europe. The Journal of Systematic Palaeontology 13 (2): 97 - 128. https: // doi. org / 10.1080 / 14772019.20 14.897266","RAGE J. - C. & BAILON S. 2005. - Amphibians and squamate reptiles from the late early Miocene (MN 4) of Beon 1 (Montrealdu-Gers, southwestern France). Geodiversitas 27 (3): 413 - 441.","AUGE M. & RAGE J. - C. 2000. - Les Squamates (Reptilia) du Miocene moyen de Sansan (Gers, France), in GINSBURG L. (ed.), La faune miocene de Sansan et son environnement. Museum national d'Histoire naturelle, Paris: 263 - 313 (Memoires du Museum national d'Histoire naturelle; 183).","SZYNDLAR Z. 2005. - Snake fauna from the Late Miocene of Rudabanya. Palaeontographia Italica 90 (2003): 31 - 52.","SZYNDLAR Z. 1984. - Fossil snakes from Poland. Acta Zoologica Cracoviensia 28 (1): 3 - 156.","IVANOV M. 1997. - Hadi evropskeho kenozoika. Unpublished PhD thesis, Masaryk University, Department of Geology and Palaeontology), Brno, 217 p. (in Czech).","TEMPFER P. M. 2005. - The Herpetofauna (Amphibia: Caudata, Anura; Reptilia: Scleroglossa) of the Upper Miocene Locality Kohfidisch (Burgerland, Austria). Beitrage zur Palaontologie 29: 145 - 253.","SZYNDLAR Z. 1991 b. - A rewiew of Neogene and Quaternary snakes of Central and Eastern Europe. Part II: Natricinae, Elapidae, Viperidae. Estudios geologicos 47 (3 - 4): 237 - 266. https: // doi. org / 10.3989 / egeol. 91473 - 4422","VENCZEL M. 2001. - Anurans and squamates from the Lower Pliocene (MN 14) Osztramos 1 locality (Northern Hungary). Fragmenta Palaeontologica Hungarica 19: 79 - 90.","RAGE J. - C. & SZYNDLAR Z. 1986. - Natrix longivertebrata from the European Neogene, a snake with one of the longest known stratigraphic ranges. Neues Jahrbuch fur Geologie und Palaontologie, Mh. 1: 56 - 64. https: // doi. org / 10.1127 / njgpm / 1986 / 1986 / 56","SZYNDLAR Z. 1991 c. - Ancestry of the Grass Snake (Natrix natrix): Paleontological evidence. Journal of Herpetology 25 (4): 412 - 418. https: // doi. org / 10.2307 / 1564762","VENCZEL M. & STIUCA E. 2008. - Late middle Miocene amphibians and squamate reptiles from Taut, Romania. Geodiversitas 30 (4): 731 - 763."]}

Vipera sp. (‘European vipers’ group) (Fig. 13 A-H) Vipera sp. 2 (‘European vipers’) – Ivanov & Musil 2004: 230. Vipera sp. (‘European vipers’ group) – Ivanov et al. 2006: 229, table 2. MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 2/2003 Reptile Joint: three trunk vertebrae (Pal. 2022-2024). DESCRIPTION Trunk vertebrae The vertebrae are rather fragmentary with the neural spines broken-off close to their bases, and loss of the paradiapophyses and hypapophysis. In lateral view, the interzygapophyseal ridge of the best-preserved vertebra (Pal. 2022) is short. The lateral foramina are situated just below the interzygapophyseal ridges. The subcentral ridges are clearly bent dorsally. In dorsal view, the cranial margin of the zygosphene has developed small lateral lobes and a wide medial lobe. The right prezygapophyseal articular facet is subtriangular to oval in outline. The prezygapophyseal process is rather short. The epizygapophyseal spines are moderately developed. In ventral view, the subcentral grooves are deep, possibly indicating a more posterior position of the vertebra within the precloacal region. The hypapophysis is strongly built, extending anteriorly into a short triangular anterior keel. In cranial view, the neural arch is depressed dorsoventrally, and the neural canal is rounded with prominent lateral sinuses. The cranial margin of the zygosphene is vaulted dorsally. Large paracotylar foramina occur on either side of the rounded cotyle. The vertebral dimensions of the largest vertebra (Pal. 2023) are as follows: cl = 3.75 mm; naw = 2.85 mm; cl/naw = 1.32. REMARKS The vertebrae are typically viperine with a dorsoventrally depressed neural arch and prezygapophyses tilted up dorsally. The vertebrae are assigned to the ‘European vipers’. Because of their small dimensions we conclude that the vertebrae could probably belong to the ‘ Vipera aspis ’ complex. A more precise identification is not possible because of poor preservation., Published as part of Ivanov, Martin, Čerňanský, Andrej, Bonilla-Salomón, Isaac & Luján, Àngel Hernández, 2020, Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications, pp. 343-376 in Geodiversitas 42 (20) on page 365, DOI: 10.5252/geodiversitas2020v42a20, http://zenodo.org/record/4447563, {"references":["IVANOV M. & MUSIL R. 2004. - Preliminary results of investigation of Neogene vertebrates from the Mokra-Quarry site. Acta Musei Moraviae, Scientiae geologicae 89: 223 - 236 (in Czech).","IVANOV M., MUSIL R. & BRZOBOHATY R. 2006. - Terrestrial and Marine Faunas from the Miocene Deposits of the Mokra Plateau (Drahany Upland, Czech Republic) - Impact on Palaeogeography. Beitrage zur Palaontologie 30: 223 - 239."]}

SCINCOIDEA indet. (Fig. 4C, D) MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 2/2003 Reptile Joint: one left maxilla (Pal. 1573). DESCRIPTION Maxilla Only a fragment of the left maxilla is preserved. It bears five tooth positions (only one and half teeth are attached). The supradental shelf is thin and straight in the preserved portion. The nasal process is partly preserved. On the medial side of the nasal process, there is a fine ridge (carina maxillaris sensu Müller 1996) that runs posterodorsally. The ridge originates from the subdental shelf. This gives an estimation that this fragment represents the anterior region of a maxilla. In the lower region of the external surface, three irregularly spaced labial foramina are located. Besides this, the rest of the surface is smooth. Dentition The implantation is pleurodont. Only one tooth is more-orless completely preserved. The lingual aspect of the crown is bordered by the culmen lateralis anterior and culmen lateralis posterior (terms after Richter 1994), between which the area is striated. This fine striation is formed by approximately six striae. The tooth crown termination is blunt. In medial aspect, it appears to be divided into rounded labial cusp, and a smaller, medially located lingual cusp. The tooth bases are pierced by small rounded resorption pits. REMARKS The tooth morphology resembles that of Scincidae or Cordylidae (see Kosma 2004). Both groups have been previously described from several lower Miocene sites of Central Europe (e.g., Roček 1984; Čerňanský 2012; Čerňanský 2016). Unfortunately, the character of preservation of this maxillary fragment from Mokrá does not allow a more specific allocation. Suborder ANGUIMORPHA Fürbringer, 1900 Family ANGUIDAE Gray, 1825 Subfamily ANGUINAE Gray, 1825, Published as part of Ivanov, Martin, Čerňanský, Andrej, Bonilla-Salomón, Isaac & Luján, Àngel Hernández, 2020, Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications, pp. 343-376 in Geodiversitas 42 (20) on page 350, DOI: 10.5252/geodiversitas2020v42a20, http://zenodo.org/record/4447563, {"references":["MULLER J. 1996. - Eine neue Art der Echten Eidechsen (Reptilia: Lacertilia: Lacertidae) aus dem Unteren Miozan. Mainzer Geowissenschaftliche Mitteilungen 25: 79 - 88.","RICHTER A. 1994. - Lacertilia aus der Unteren Kreide von Una und Galve (Spanien) und Anoual (Marokko). Berliner geowissenschaftliche Abhandlungen (E: Palaobiologie) 14: 1 - 147.","KOSMA R. 2004. - The Dentition of Recent and Fossil Scincomorphan Lizards (Lacertilia, Squamata) - Systematics, Functional Morphology, Palecology. Unpublished PhD thesis, University of Hannover, Hannover. 187 p.","ROCEK Z. 1984. - Lizards (Reptilia: Sauria) from the Lower Miocene locality Dolnice (Bohemia, Czechoslovakia). Rozpravy Ceskoslovenske Akademie Ved, Rada Matematickych a Prirodnich Ved 94: 1 - 69.","CERNANSKY A. 2012. - The oldest known European Neogene girdled lizard fauna (Squamata, Cordylidae), with comments on early Miocene immigration of African taxa. Geodiversitas 34 (4): 837 - 847. https: // doi. org / 10.5252 / g 2012 n 4 a 6"]}

AMPHISBAENIA indet. (Fig. 3) Blanus sp. – Ivanov et al. 2006: 229, table 2. MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 2/2003 Reptile Joint: One trunk vertebra (Pal. 1570). DESCRIPTION Trunk vertebra A single trunk vertebra is preserved. It is small in size. A neural spine is absent, and the dorsal portion of the neural arch forms a median edge. In lateral view, the synapophysis is simple and large. The posterior portion of the neural arch is fused with the postzygapophyses, forming the dorsal roof (or lamina) between the left and right postzygapophyses. The neural canal is subtriangular with distinct lateral sinuses. The interzygapophyseal constriction is distinct and it occurs in the anterior half of the anteroposterior vertebral length. The dorsally tilted prezygapophyseal articular facets have an elliptical shape. A zygosphene is absent. The ventral side of the depressed centrum is flat, pierced by a pair of large subcentral foramina in the anterior 1/3 of the anteroposterior length. The lateral margins (subcentral ridges) are roughly parallel in ventral aspect. No constriction is developed at the base of the damaged condyle. The postzygapophyseal articular facets are oval and slightly enlarged posteriorly. The cotyle is distinctly laterally enlarged. REMARKS The vertebra described here can be attributed to Amphisbaenia based on the following combination of features (see Estes 1983): 1) the depressed centrum, having a flat ventral surface; 2) roughly parallel lateral margins in ventral aspect; 3) massive synapophyses; 4) the absence of a zygosphene (enabling distinction of amphisbaenians from scolecophidian snakes (Estes 1983; Rage 1984); and 5) the sinusoidal neural arch lacking a neural spine. Family level allocation of an isolated vertebra is limited by a lack of clear diagnostic features for identification (Estes 1983; Augé 2005, 2012; Georgalis et al. 2016b).We can exclude rhineurids, which have a denticulate vertebral posterior margin. The same feature can be observed in trogonophiids as well (Kearney 2003; Augé 2012; Čerňanský et al. 2016a). Based on the geographical position of the locality and the age of the sediments, this vertebra most likely represents a blanid taxon. According to cranial elements, amphisbaenians reported from the Central European late Oligocene and Miocene localities are almost exclusively identified as belonging to the clade Blanidae (Roček 1984; Schleich 1988; Čerňanský & Venczel 2011; Čerňanský et al. 2016a). The morphology and dimensions of the vertebra described here are very similar to those of trunk vertebra of Blanus gracilis Roček, 1984 reported from the Czech early Miocene (MN 4b) Dolnice site (Roček 1984: 5, table 16).

Coluber (s.l.) sp. (Fig. 9) Coluber sp. 1 – Ivanov & Musil 2004: 229 (in part). Coluber sp., type I – Ivanov et al. 2006: 229, table 2 (in part). MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 1/2001 Turtle Joint: Two anterior trunk vertebrae (Pal. 1454-1455), 6 middle trunk vertebrae (Pal. 1456-1461), 2 caudal vertebrae (Pal. 1462, 1463). 2/2003 Reptile Joint:Two trunk vertebrae (Pal. 1956- 1957), 2 caudal vertebrae (Pal. 1958-1959). DESCRIPTION Anterior and middle trunk vertebrae The only two preserved anterior trunk vertebrae differ from those from the middle trunk portion by the presence of hypapophysis instead of haemal keel and higher neural spine (Pal. 1454; Fig. 9 A-E). All preserved middle trunk vertebrae are at least partially fragmentary with neural spines mostly broken-off close to their bases. In lateral view, the preserved neural spine base of the most complete specimen (Pal. 1456; Fig. 9 F-J) indicates that the neural spine was probably at least twice longer than high. The caudal margin of the neural spine was inclined posteriorly as documented by specimen Pal. 1956. The well-developed interzygapophyseal ridges are short. The lateral foramina occur just below these ridges. The prominent subcentral ridges are moderately dorsally arched. The large diapophyses are well-separated from somewhat smaller parapophyses. Although parapophyses are incomplete in all specimens, their ventral margin apparently extended below the cotylar rim. In dorsal view, the wide zygosphene has developed lateral lobes. The medial lobe is rather wide. The prezygapophyseal articular facets are oval with long axis directed antero-laterally. The prezygapophyseal processes are broken-off close to their bases. The epizygapophyseal spines are moderately developed. A deep notch occurs at the caudal margin of the neural arch. In ventral view, the straight subcentral ridges form the lateral margins of a cranio-caudally elongated centrum of narrowly triangular shape. The subcentral grooves are wide and shallow. The subcentral foramina are rather small. The haemal keel is narrow and reaches posteriorly almost to the cranial margin of the small rounded condyle. Small subcotylar tubercles occur at the ventral margin of the cotylar rim. The postzygapophyseal articular facets are subrectangular and slightly laterally elongated. In cranial view, the neural arch is vaulted, and the neural canal is wide and rounded with wide and shallow lateral sinuses. The cranial margin of the zygosphenal lip arches dorsally. The small paracotylar foramina occur within depressions on either side of the rounded cotyle. In caudal view, the zygantrum is wide. The vertebral dimensions of the largest specimen (Pal. 1456) are as follows: cl = 4.52 mm; naw = 3.21 mm; cl/naw = 1.41. Caudal vertebrae The rarely preserved caudal vertebrae are relatively short. In lateral view, the neural spine of the better-preserved specimen was twice as long as high. Its cranial margin rises in the middle of the zygosphene length. The zygosphenal facets are oval. The preserved base of the right pleurapophysis is directed antero-ventrally. Haemapophyses are broken-off close to their bases. In dorsal view, the zygosphene is almost straight. The prezygapophyseal articular facets are oval in outline. The preserved base of the right pleurapophysis in specimen Pal. 1462 (Fig. 9 K-O) is anterolaterally directed indicating anterior caudal position within the vertebral column. In cranial view, the zygosphene is arched dorsally. The small paracotylar foramina occur on either side of the circular cotylar rim. REMARKS The gracile structure of vertebrae with cl/naw ratio> 1, the presence of paracotylar foramina, as well as well-developed neural spine, the presence of prezygapophyseal processes, and the haemal keel developed in trunk vertebrae enable assignment to “ Colubrinae ”. The middle trunk vertebrae resemble those of the genus “ Coluber ” on the basis of the following combination of characters: 1) strongly craniocaudally elongated centrum of trunk vertebrae; 2) the vaulted neural arch; 3) the well-developed and narrow haemal keel; and 4) the prezygapophyseal processes which were probably long, based on the well-developed prezygapophyseal processes in anterior trunk vertebrae. “Colubrines” referred to the genus “ Coluber ” have frequently been reported from the European Neogene (e.g., Szyndlar 1991a, 2005, 2009, 2012; Szyndlar & Schleich 1993; Ivanov 2002a, b; Ivanov & Böhme 2011; Rage & Bailon 2005; Venczel 1994, 1998, 2001). There are four large “colubrine” species in the European Miocene: Coluber dolnicensis Szyndlar, 1987 (MN 3a-MN 4), C. caspioides Szyndlar & Schleich, 1993 (MN 3a-? MN 6), C. suevicus (Fraas, 1870) (MN 3a-MN 7+8) and C. pouchetii (de Rochebrune, 1880) (MN 4- MN 9). Coluber (s.l.) sp. differs from C. dolnicensis in the absence of a prominent step in the anterior part of the haemal keel (Szyndlar 1987; Ivanov 2002a). It differs from C. caspioides in the smaller dimensions and the wider zygosphene (Szyndlar & Schleich 1993; Ivanov 2002a). Coluber (s.l.) sp. differs from C. suevicus in the more vaulted neural arch and clearly smaller diameter of the prezygapophyseal articular facets (Szyndlar & Böhme 1993; Ivanov 2002a). It differs from C. pouchetii in the apparently much lower neural spine and a cervical hypapophysis inclined posteroventrally rather than ventrally (Augé & Rage 2000; Szyndlar 2009). Coluber (s.l.) sp., type 1 resembles C. hungaricus (Bolkay, 1913) reported from the early middle Miocene (MN 6) of Germany (Ivanov & Böhme 2011), the middle Miocene to early Pliocene (? MN 6- MN 9; MN 13- MN 14) of Hungary (Venczel 1994, 1998, 2001), and the middle Miocene of Kazakhstan (Ivanov et al. 2019) in: 1) the same dimensions; 2) most probably in the same height of the neural spine; 3) a similarly wide medial lobe of the zygosphene; and 4) a parapophysis of the same length as the diapophysis (Venczel 1994; Szyndlar 2005). C. hungaricus displays a high intraspecific variability and the largest specimens have a dorsally thickened neural spine, parapophyses larger than diapophyses, as well as well-developed subcotylar tubercles (Venczel 1998). Despite this variability, Coluber (s.l.) sp. cannot be attributed to C. hungaricus because of the presence of much more distinct, sharp, and straight subcentral ridges, a deeper notch in the caudal margin of the neural arch, as well as the smaller condyle. Because of the incomplete preservation of the rather scarce material we avoid a species level identification., Published as part of Ivanov, Martin, Čerňanský, Andrej, Bonilla-Salomón, Isaac & Luján, Àngel Hernández, 2020, Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications, pp. 343-376 in Geodiversitas 42 (20) on pages 358-360, DOI: 10.5252/geodiversitas2020v42a20, http://zenodo.org/record/4447563, {"references":["IVANOV M. & MUSIL R. 2004. - Preliminary results of investigation of Neogene vertebrates from the Mokra-Quarry site. Acta Musei Moraviae, Scientiae geologicae 89: 223 - 236 (in Czech).","IVANOV M., MUSIL R. & BRZOBOHATY R. 2006. - Terrestrial and Marine Faunas from the Miocene Deposits of the Mokra Plateau (Drahany Upland, Czech Republic) - Impact on Palaeogeography. Beitrage zur Palaontologie 30: 223 - 239.","SZYNDLAR Z. 1991 a. - A rewiew of Neogene and Quaternary snakes of Central and Eastern Europe. Part I: Scolecophidia, Boidae, Colubridae. Estudios geologicos 47 (1 - 2): 103 - 126. https: // doi. org / 10.3989 / egeol. 91471 - 2412","SZYNDLAR Z. 2005. - Snake fauna from the Late Miocene of Rudabanya. Palaeontographia Italica 90 (2003): 31 - 52.","SZYNDLAR Z. 2009. - Snake fauna (Reptilia: Serpentes) from the Early / Middle Miocene of Sandelzhausen and Rothenstein 13 (Germany). Palaontologische Zeitschrift 83 (1): 55 - 66. https: // doi. org / 10.1007 / s 12542 - 009 - 0009 - 5","SZYNDLAR Z. 2012. - Early Oligocene to Pliocene Colubridae of Europe: a review. Bulletin de la Societe geologique de France 183 (6): 661 - 681. https: // doi. org / 10.2113 / gssgfbull. 183.6.661","SZYNDLAR Z. & SCHLEICH H. H. 1993. - Description of Miocene snakes from Petersbuch 2 with comments on the Lower and Middle Miocene ophidian faunas of Southern Germany. Stuttgarter Beitrage zur Naturkunde Serie B (Geologie und Palaontologie) 192: 1 - 47.","IVANOV M. 2002 a. - The oldest known Miocene snake fauna from Central Europe: Merkur-North locality, Czech Republic. Acta Palaeontologica Polonica 47 (3): 513 - 534.","IVANOV M. & BOHME M. 2011. - Snakes from Griesbeckerzell (Langhian, Early Badenian), North Alpine Foreland Basin (Germany), with comments on the evolution of snake fauna in Central Europe during the Miocene Climatic Optimum. Geodiversitas 33 (3): 411 - 449. https: // doi. org / 10.5252 / g 2011 n 3 a 2","RAGE J. - C. & BAILON S. 2005. - Amphibians and squamate reptiles from the late early Miocene (MN 4) of Beon 1 (Montrealdu-Gers, southwestern France). Geodiversitas 27 (3): 413 - 441.","VENCZEL M. 1994. - Late Miocene snakes from Polgardi (Hungary). Acta Zoologica Cracoviensia 37 (1): 1 - 29.","VENCZEL M. 1998. - Late Miocene snakes (Reptilia: Serpentes) from Polgardi (Hungary): a second contribution. Acta Zoologica Cracoviensia 41 (1): 1 - 22.","VENCZEL M. 2001. - Anurans and squamates from the Lower Pliocene (MN 14) Osztramos 1 locality (Northern Hungary). Fragmenta Palaeontologica Hungarica 19: 79 - 90.","SZYNDLAR Z. 1987. - Snakes from the Lower Miocene locality of Dolnice (Czechoslovakia). Journal of Vertebrate Paleontology 7 (1): 55 - 71. https: // doi. org / 10.1080 / 02724634.1987.10011637","SZYNDLAR Z. & BOHME W. 1993. - Die fossilen Schlangen Deutschlands: Geschichte der Faunen und ihrer Erforschung. Mertensiella 3: 381 - 431.","AUGE M. & RAGE J. - C. 2000. - Les Squamates (Reptilia) du Miocene moyen de Sansan (Gers, France), in GINSBURG L. (ed.), La faune miocene de Sansan et son environnement. Museum national d'Histoire naturelle, Paris: 263 - 313 (Memoires du Museum national d'Histoire naturelle; 183).","BOLKAY S. J. 1913. - Additions to the fossil herpetology of Hungary from the Pannonian and Praeglacial periode. Mitteilungen aus dem Jahrbuch der koniglichen ungarischen Geologischen Reichsanstalt 21: 217 - 230.","IVANOV M., VASILYAN D., BOHME M. & ZAZHIGIN V. S. 2019. - Miocene snakes from northeastern Kazakhstan: new data on the evolution of snake assemblages in Siberia. Historical Biology 31 (10): 1284 - 1303. https: // doi. org / 10.1080 / 08912963.2018. 1446086"]}

Python sp. (Fig. 8) Boidae gen. et sp. indet. (large form) – Ivanov & Musil 2004: 228, 229, fig. 3A, B. Boidae gen. et sp. indet – Ivanov et al. 2006: 229, table 2. MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 1/2001 Turtle Joint: two trunk vertebrae (Pal. 1449, 1450). DESCRIPTION Trunk vertebrae The more complete middle trunk vertebra (Pal. 1449) has lost the left prezygapophyseal facet as well as the lateral extension of the left postzygapophysis. In lateral view, the vertebra is anteroposteriorly shorter than high. The strongly vaulted neural arch is caudally upswept above the zygantrum. The neural spine has a gently eroded dorsal margin and is slightly longer than high. Its cranial margin is vertical and rises in the middle of the zygosphene length. The caudal margin of the neural spine is inclined posteriorly behind the neural arch. The interzypapophyseal ridges are rather sharp. The zygosphenal surfaces are wide and irregularly oval. They are characterised by conspicuously large dimensions. The lateral foramina are large, and they sit in shallow depressions. The haemal keel is arched upwards. In dorsal view, the wide zygosphene possesses distinct lateral lobes; the damaged medial lobe was rather small. The neural spine is thick. The right prezygapophyseal articular facet is subtriangular. The prezygapophyseal process is rather short and it is hardly visible from the dorsal aspect. The interzygapophyseal constriction is shallow. The neural arch widens triangularly in a caudal direction. The median notch, developed at the caudal margin of the neural arch, reaches anteriorly as far as the cranial margin of the postzygapophysis. In ventral view, the paradiapophyses are massively developed. The large right postzygapophyseal articular facet has a subtriangular outline and is laterally elongated. In this view, the haemal keel is relatively wide with subcentral foramina situated at both sides of its base. In cranial view, the relatively thin prezygapophyses are tilted slightly dorsally with their base situated just above the floor of the neural canal. The lateral extension of the preserved right postzygapophysis is conspicuously thin. The neural arch is strongly vaulted, and the neural canal is rounded with short lateral sinuses. The straight cranial margin of the zygosphene is thick along its entire width. Deep depressions occur on either side of a large cotyle of almost circular outline. Paracotylar foramina are absent. In caudal view, the zygantrum is large and laterally wide. The postzygapophyses are tilted dorsally like the prezygapophyses. Several very small foramina are arranged in a line on caudal side of both postzygapophyses. The condyle is almost circular with a depressed ventral margin. The vertebral dimensions are as follows: larger specimen (Pal. 1449): cl = 7.32 mm; naw = 10.39 mm; cl /naw = 0.70. Smaller specimen (Pal. 1450): cl = 5.45 mm; naw = 8.15 mm; cl/naw = 0.67. REMARKS The following combination of features indicate assignment of the better preserved vertebra to the extant genus Python (see Szyndlar & Rage 2003): 1) the large vertebra is massively built with cl/naw ratio Python sp. from MWQ particularly resembles the only known extinct Python species, Python europaeus Szyndlar & Rage, 2003, reported from French early Miocene Béon 1 (MN 4) and Vieux-Collonges (MN 5) localities (Szyndlar & Rage 2003; Rage & Bailon 2005), and from the middle Miocene (MN 6, base) of Griesbeckerzell 1a, Germany (Ivanov & Böhme 2011). However, Python sp. from MWQ differs from P. europaeus by the following features: 1) the zygosphene of Python sp. is not straight – conspicuous lateral lobes are present; 2) in cranial view, the zygosphenal lip is slightly less massive compared to that of P. europaeus; and 3) lateral extensions of postzygapophyses are more slender than the same structures in P. europaeus. The trunk vertebrae of Python sp. from MWQ are smaller than middle trunk vertebrae of P. europaeus (Szyndlar & Rage 2003). As the morphology and build of the zygosphene strongly depends on the ontogenetic stage, it is possible that vertebrae of Python sp. belonged to subadult specimens., Published as part of Ivanov, Martin, Čerňanský, Andrej, Bonilla-Salomón, Isaac & Luján, Àngel Hernández, 2020, Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications, pp. 343-376 in Geodiversitas 42 (20) on pages 357-358, DOI: 10.5252/geodiversitas2020v42a20, http://zenodo.org/record/4447563, {"references":["IVANOV M. & MUSIL R. 2004. - Preliminary results of investigation of Neogene vertebrates from the Mokra-Quarry site. Acta Musei Moraviae, Scientiae geologicae 89: 223 - 236 (in Czech).","IVANOV M., MUSIL R. & BRZOBOHATY R. 2006. - Terrestrial and Marine Faunas from the Miocene Deposits of the Mokra Plateau (Drahany Upland, Czech Republic) - Impact on Palaeogeography. Beitrage zur Palaontologie 30: 223 - 239.","SZYNDLAR Z. & RAGE J. - C. 2003. - Non-erycine Booidea from the Oligocene and Miocene of Europe. Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Krakow, 109 p.","RAGE J. - C. & BAILON S. 2005. - Amphibians and squamate reptiles from the late early Miocene (MN 4) of Beon 1 (Montrealdu-Gers, southwestern France). Geodiversitas 27 (3): 413 - 441.","IVANOV M. & BOHME M. 2011. - Snakes from Griesbeckerzell (Langhian, Early Badenian), North Alpine Foreland Basin (Germany), with comments on the evolution of snake fauna in Central Europe during the Miocene Climatic Optimum. Geodiversitas 33 (3): 411 - 449. https: // doi. org / 10.5252 / g 2011 n 3 a 2"]}

ELAPIDAE gen. et sp. indet. (Fig. 16) Elapidae gen. et sp. indet. – Ivanov & Musil 2004: 230. MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 1/2001 Turtle Joint: One trunk vertebra (Pal. 1500). DESCRIPTION Trunk vertebra The only preserved vertebra is fragmentary with loss of the left prezygapophysis and the right prezygapophyseal process. In lateral view, the neural spine is broken-off close to its base, but the caudal part of the neural spine indicates that it was originally rather low. The interzygapophyseal ridges are moderately developed. The epizygapophyseal ridges are missing. The subcentral ridges are arched slightly dorsally. The hypapophysis is straight and its base rises at one quarter of the centrum length. The distal termination of the hypapophysis is absent but apparently it was short and did not reach behind the caudal margin of the condyle. In ventral view, the subcentral grooves are shallow and subcentral foramina are rather small. The postzygapophyseal articular facets are subcircular to irregularly shaped. In cranial view, the neural arch is slightly vaulted, and the neural canal is rounded with a wide diameter and short lateral sinuses. The damaged cranial margin of the zygosphenal lip was arched dorsally with raised zygosphenal facets. The paracotylar foramina occur on either side of the rounded cotyle. The small laterally directed subcotylar tubercles occur at the ventral margin of the rounded cotylar rim. The vertebral dimensions are as follows: cl = 5.40 mm; naw = 3.57 mm; cl/naw = 1.51. REMARKS The single preserved vertebra was assigned to the family Elapidae on the basis of the presence of a short and straight hypapophysis that begins far from the ventral margin of the cotylar rim, the absence of epizygapophyseal ridges and the likely presence of a low neural spine. The morphology of Elapidae gen. et sp. indet. is identical with that of a vertebra of Elapidae indet. reported from the late Miocene (MN 9) of Rudabánya, Hungary (Szyndlar 2005). According toSzyndlar (2005), the only preserved Rudabánya specimen is similar to numerous small-sized elapids known from the European Miocene including Micrurus (M. gallicus Rage & Holman, 1984). However, the Mokrá specimen is larger. Both the morphology and dimensions of the largest vertebra resemble those of vertebral morphotype Elapidae B reported from the French early Miocene (MN 5) Vieux-Collonges site, with the exception of the rather short condylar neck in French specimens (Ivanov 2000: fig. 14). In accordance with Ivanov (2000), on the basis of relatively high cl/naw ratio, we conclude that this middle trunk vertebra of an indeterminate Elapidae from 1/2001 Turtle Joint can be attributed to large-sized elapids of possibly Asiatic origin (perhaps Naja)., Published as part of Ivanov, Martin, Čerňanský, Andrej, Bonilla-Salomón, Isaac & Luján, Àngel Hernández, 2020, Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications, pp. 343-376 in Geodiversitas 42 (20) on page 369, DOI: 10.5252/geodiversitas2020v42a20, http://zenodo.org/record/4447563, {"references":["IVANOV M. & MUSIL R. 2004. - Preliminary results of investigation of Neogene vertebrates from the Mokra-Quarry site. Acta Musei Moraviae, Scientiae geologicae 89: 223 - 236 (in Czech).","SZYNDLAR Z. 2005. - Snake fauna from the Late Miocene of Rudabanya. Palaeontographia Italica 90 (2003): 31 - 52.","RAGE J. - C. & HOLMAN J. A. 1984. - Des serpents (Reptilia, Squamata) de type Nord-Americain dans le Miocene francais. Evolution parallele ou dispersion? Geobios 17 (1): 89 - 104. https: // doi. org / 10.1016 / S 0016 - 6995 (84) 80007 - 8"]}

LACERTIDAE indet. (Fig. 2A, B) MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 2/2003 Reptile Joint: one left frontal (Pal. 1565). DESCRIPTION Frontal Only the anterior portion of the left frontal is preserved. The dorsal surface bears two osteodermal shields fused to the bone. The main region is occupied by the frontal shield, whereas the prefrontal shield is located in the anterolateral section. The dorsal regions of the shields are sculptured. The sculpture consists of irregularly distributed pits and several connected ridges. Both shields are separated one from another by a slightly rounded (medially convex) sulcus, which runs posterolaterally to the lateral margin of the bone. The anterior termination of the frontal is divided into larger lateral process and slightly smaller medial process. Between them, a wedge-shaped facet for the nasal is present, forming a bony septum. The facet for the posterodorsal termination of the maxillary nasal process is located lateral to the lateral process, however, this region is damaged. The medial margin, which forms a contact with the right frontal, is straight. On the internal surface, the frontal cranial ridge can be observed. This portion is unfortunately badly preserved – its section that would form an anteroventrally oriented subolfactory process is broken off. LACERTIDAE indet. tooth morphotype 1 (Fig. 2 C-G) MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 1/2001 Turtle Joint: One left dentary (Pal. 1400). 2/2003 Reptile Joint: one right maxilla (Pal. 1566), 3 dentaries, 1 left + 2 right (Pal. 1567-1569). DESCRIPTION Maxilla Only a small fragment of the right maxilla is preserved (Fig. 2C, D). This portion bears two teeth, which are bordered dorsally by the supradental shelf. The lateral surface is pierced by a large labial foramen. Dentary The description is based on two fragments – one represents a left dentary, whereas the second is a right dentary. The left dentary fragment (Fig. 2E, F) bears four tooth positions (two teeth are still attached). The right dentary (Fig. 2G) exhibits five and half tooth positions (four teeth are still attached, but the tooth crown of one tooth is broken off). The dental crest is low, and teeth exceed it by 1/2 of their height. The subdental shelf (sensu Rage & Augé 2010) is well developed, robust. However only its short portion is preserved. It gradually becomes thinner posteriorly (this can be observed mainly in the right dentary fragment), partly as a result of the presence of the facet for the splenial, situated on its ventral margin. The shelf forms the dorsal roof of the Meckel’s groove, which is open but narrow. The lateral surface of the bone is smooth. In the preserved section, it is pierced by two labial foramina located slightly above the mid-section of the bone. Dentition The implantation is pleurodont.Teeth are high. The interdental gaps are large – the size of the gap forms approximately the 1/2 of the mesiodistal length of the tooth neck. The tooth crowns are bicuspid with a dominant distal (central) cusp and a smaller mesial cusp. The distal cusp is pointed in most cases and slightly directed posterolingually. The lingual portion of the crowns bears vertical striations. The striae are almost parallel, and their number is around ten. The tooth necks are slightly expanded lingually and they appear lightly more swollen if compared to the tooth crowns. The central part of the tooth base is pierced by a small resorption pit. REMARKS The maxilla and dentary have identical tooth morphology and thus can be attributed to the single taxon. Several features in the material from Mokrá described here resemble Lacerta poncenatensis: 1) the presence of robust bicuspid teeth; 2) the wide interdental gaps; and 3) the low dental crest. This taxon was originally described by Müller (1996) from the French locality of Poncenat (early Miocene, MN 2a). Later, it was also recognized in Germany (Čerňanský et al. 2015; early Miocene, MN 2) and Austria (Čerňanský 2016; early Miocene, MN 4). However, the fragmentary nature of the Mokrá material does not allow confident alpha taxonomy. LACERTIDAE indet. tooth morphotype 2 (Fig. 2H) MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 1/2001 Turtle Joint: One left dentary (Pal. 1401). DESCRIPTION Dentary The description is based on the fragment of the anterior half of a left dentary. The element bears ten tooth positions (four teeth are still attached). The dental crest is high, and the teeth extend above it only in a quarter of their total height. The subdental shelf (sensu Rage & Augé, 2010) is robust, being only slightly concave in this section. Meckel’s groove is open, narrow in the preserved section, but gradually widening posteriorly. The lateral surface of the bone is smooth, pierced by several labial foramina. Dentition The implantation is pleurodont. Teeth are tall and robust. The interdental gaps are small – the size of the gap forms approximately only a 1/4 of the mesiodistal length of the tooth neck. The tooth crowns are bicuspid, with a dominant distal (central) cusp and a smaller mesial cusp. The lingual portion of the crowns bears vertical striations. The striae are almost parallel, and their number is around six. In medial aspect, the tooth necks are more or less as wide as the tooth crowns, in some cases gradually narrowing slightly ventrally. The necks appear slightly more swollen lingually if compared to the tooth crowns. The central part of the tooth base is pierced by a resorption pit. REMARKS Although the dentary described here possesses some similarities with the above described lacertid material, e.g. bicuspid teeth, several important differences can be observed: 1) large size; 2) the high dental crest; 3) small interdental gaps; 4) more robust teeth; and 5) low number of lingual striae on the tooth crowns. Because not all of those differences can be explained by ontogenetic changes, we suggest the presence of at least two lacertid taxa in MWQ. Unranked clade AMPHISBAENIA Gray, 1844 AMPHISBAENIA indet. (Fig. 3) Blanus sp. – Ivanov et al. 2006: 229, table 2. MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 2/2003 Reptile Joint: One trunk vertebra (Pal. 1570). DESCRIPTION Trunk vertebra A single trunk vertebra is preserved. It is small in size. A neural spine is absent, and the dorsal portion of the neural arch forms a median edge. In lateral view, the synapophysis is simple and large. The posterior portion of the neural arch is fused with the postzygapophyses, forming the dorsal roof (or lamina) between the left and right postzygapophyses. The neural canal is subtriangular with distinct lateral sinuses. The interzygapophyseal constriction is distinct and it occurs in the anterior half of the anteroposterior vertebral length. The dorsally tilted prezygapophyseal articular facets have an elliptical shape. A zygosphene is absent. The ventral side of the depressed centrum is flat, pierced by a pair of large subcentral foramina in the anterior 1/3 of the anteroposterior length. The lateral margins (subcentral ridges) are roughly parallel in ventral aspect. No constriction is developed at the base of the damaged condyle. The postzygapophyseal articular facets are oval and slightly enlarged posteriorly. The cotyle is distinctly laterally enlarged. REMARKS The vertebra described here can be attributed to Amphisbaenia based on the following combination of features (see Estes 1983): 1) the depressed centrum, having a flat ventral surface; 2) roughly parallel lateral margins in ventral aspect; 3) massive synapophyses; 4) the absence of a zygosphene (enabling distinction of amphisbaenians from scolecophidian snakes (Estes 1983; Rage 1984); and 5) the sinusoidal neural arch lacking a neural spine. Family level allocation of an isolated vertebra is limited by a lack of clear diagnostic features for identification (Estes 1983; Augé 2005, 2012; Georgalis et al. 2016b).We can exclude rhineurids, which have a denticulate vertebral posterior margin. The same feature can be observed in trogonophiids as well (Kearney 2003; Augé 2012; Čerňanský et al. 2016a). Based on the geographical position of the locality and the age of the sediments, this vertebra most likely represents a blanid taxon. According to cranial elements, amphisbaenians reported from the Central European late Oligocene and Miocene localities are almost exclusively identified as belonging to the clade Blanidae (Roček 1984; Schleich 1988; Čerňanský & Venczel 2011; Čerňanský et al. 2016a). The morphology and dimensions of the vertebra described here are very similar to those of trunk vertebra of Blanus gracilis Roček, 1984 reported from the Czech early Miocene (MN 4b) Dolnice site (Roček 1984: 5, table 16). LACERTIDAE indet. tooth morphotype 1 (Fig. 2 C-G) MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 1/2001 Turtle Joint: One left dentary (Pal. 1400). 2/2003 Reptile Joint: one right maxilla (Pal. 1566), 3 dentaries, 1 left + 2 right (Pal. 1567-1569). DESCRIPTION Maxilla Only a small fragment of the right maxilla is preserved (Fig. 2C, D). This portion bears two teeth, which are bordered dorsally by the supradental shelf. The lateral surface is pierced by a large labial foramen. Dentary The description is based on two fragments – one represents a left dentary, whereas the second is a right dentary. The left dentary fragment (Fig. 2E, F) bears four tooth positions (two teeth are still attached). The right dentary (Fig. 2G) exhibits five and half tooth positions (four teeth are still attached, but the tooth crown of one tooth is broken off). The dental crest is low, and teeth exceed it by 1/2 of their height. The subdental shelf (sensu Rage & Augé 2010) is well developed, robust. However only its short portion is preserved. It gradually becomes thinner posteriorly (this can be observed mainly in the right dentary fragment), partly as a result of the presence of the facet for the splenial, situated on its ventral margin. The shelf forms the dorsal roof of the Meckel’s groove, which is open but narrow. The lateral surface of the bone is smooth. In the preserved section, it is pierced by two labial foramina located slightly above the mid-section of the bone. Dentition The implantation is pleurodont.Teeth are high. The interdental gaps are large – the size of the gap forms approximately the 1/2 of the mesiodistal length of the tooth neck. The tooth crowns are bicuspid with a dominant distal (central) cusp and a smaller mesial cusp. The distal cusp is pointed in most cases and slightly directed posterolingually. The lingual portion of the crowns bears vertical striations. The striae are almost parallel, and their number is around ten. The tooth necks are slightly expanded lingually and they appear lightly more swollen if compared to the tooth crowns. The central part of the tooth base is pierced by a small resorption pit. REMARKS The maxilla and dentary have identical tooth morphology and thus can be attributed to the single taxon. Several features in the material from Mokrá described here resemble Lacerta poncenatensis: 1) the presence of robust bicuspid teeth; 2) the wide interdental gaps; and 3) the low dental crest. This taxon was originally described by Müller (1996) from the French locality of Poncenat (early Miocene, MN 2a). Later, it was also recognized in Germany (Čerňanský et al. 2015; early Miocene, MN 2) and Austria (Čerňanský 2016; early Miocene, MN 4). However, the fragmentary nature of the Mokrá material does not allow confident alpha taxonomy. LACERTIDAE indet. tooth morphotype 2 (Fig. 2H) MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 1/2001 Turtle Joint: One left dentary (Pal. 1401). DESCRIPTION Dentary The description is based on the fragment of the anterior half of a left dentary. The element bears ten tooth positions (four teeth are still attached). The dental crest is high, and the teeth extend above it only in a quarter of their total height. The subdental shelf (sensu Rage & Augé, 2010) is robust, being only slightly concave in this section. Meckel’s groove is open, narrow in the preserved section, but gradually widening posteriorly. The lateral surface of the bone is smooth, pierced by several labial foramina. Dentition The implantation is pleurodont. Teeth are tall and robust. The interdental gaps are small – the size of the gap forms approximately only a 1/4 of the mesiodistal length of the tooth neck. The tooth crowns are bicuspid, with a dominant distal (central) cusp and a smaller mesial cusp. The lingual portion of the crowns bears vertical striations. The striae are almost parallel, and their number is around six. In medial aspect, the tooth necks are more or less as wide as the tooth crowns, in some cases gradually narrowing slightly ventrally. The necks appear slightly more swollen lingually if compared to the tooth crowns. The central part of the tooth base is pierced by a resorption pit. REMARKS Although the dentary described here possesses some similarities with the above described lacertid material, e.g. bicuspid teeth, several important differences can be observed: 1) large size; 2) the high dental crest; 3) small interdental gaps; 4) more robust teeth; and 5) low number of lingual striae on the tooth crowns. Because not all of those differences can be explained by ontogenetic changes, we suggest the presence of at least two lacertid taxa in MWQ. Unranked clade AMPHISBAENIA Gray, 1844 AMPHISBAENIA indet. (Fig. 3) Blanus sp. – Ivanov et al. 2006: 229, table 2. MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 2/2003 Reptile Joint: One trunk vertebra (Pal. 1570). DESCRIPTION Trunk vertebra A single trunk vertebra is preserved. It is small in size. A neural spine is absent, and the dorsal portion of the neural arch forms a median edge. In lateral view, the synapophysis is simple and large. The posterior portion of the neural arch is fused with the postzygapophyses, forming the dorsal roof (or lamina) between the left and right postzygapophyses. The neural canal is subtriangular with distinct lateral sinuses. The interzygapophyseal constriction is distinct and it occurs in the anterior half of the anteroposterior vertebral length. The dorsally tilted prezygapophyseal articular facets have an elliptical shape. A zygosphene is absent. The ventral side of the depressed centrum is flat, pierced by a pair of large subcentral foramina in the anterior 1/3 of the anteroposterior length. The lateral margins (subcentral ridges) are roughly parallel in ventral aspect. No constriction is developed at the base of the damaged condyle. The postzygapophyseal articular facets are oval and slightly enlarged posteriorly. The cotyle is distinctly laterally enlarged. REMARKS The vertebra described here can be attributed to Amphisbaenia based on the following combination of features (see Estes 1983): 1) the depressed centrum, having a flat ventral surface; 2) roughly parallel lateral margins in ventral aspect; 3) massive synapophyses; 4) the absence of a zygosphene (enabling distinction of amphisbaenians from scolecophidian snakes (Estes 1983; Rage 1984); and 5) the sinusoidal neural arch lacking a neural spine. Family level allocation of an isolated vertebra is limited by a lack of clear diagnostic features for identification (Estes 1983; Augé 2005, 2012; Georgalis et al. 2016b).We can exclude rhineurids, which have a denticulate vertebral posterior margin. The same feature can be observed in trogonophiids as well (Kearney 2003; Augé 2012; Čerňanský et al. 2016a). Based on the geographical position of the locality and the age of the sediments, this vertebra most likely represents a blanid taxon. According to cranial elements, amphisbaenians reported from the Central European late Oligocene and Miocene localities are almost exclusively identified as belonging to the clade Blanidae (Roček 1984; Schleich 1988; Čerňanský & Venczel 2011; Čerňanský et al. 2016a). The morphology and dimensions of the vertebra described here are very similar to those of trunk vertebra of Blanus gracilis Roček, 1984 reported from the Czech early Miocene (MN 4b) Dolnice site (Roček 1984: 5, table 16). LACERTIDAE indet. tooth morphotype 2 (Fig. 2H) MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 1/2001 Turtle Joint: One left dentary (Pal. 1401). DESCRIPTION Dentary The description is based on the fragment of the anterior half of a left dentary. The element bears ten tooth positions (four teeth are still attached). The dental crest is high, and the teeth extend above it only in a quarter of their total height. The subdental shelf (sensu Rage & Augé, 2010) is robust, being only slightly concave in this section. Meckel’s groove is open, narrow in the preserved section, but gradually widening posteriorly. The lateral surface of the bone is smooth, pierced by several labial foramina. Dentition The implantation is pleurodont. Teeth are tall and robust. The interdental gaps are small – the size of the gap forms approximately only a 1/4 of the mesiodistal length of the tooth neck. The tooth crowns are bicuspid, with a dominant distal (central) cusp and a smaller mesial cusp. The lingual portion of the crowns bears vertical striations. The striae are almost parallel, and their number is around six. In medial aspect, the tooth necks are more or less as wide as the tooth crowns, in some cases gradually narrowing slightly ventrally. The necks appear slightly more swollen lingually if compared to the tooth crowns. The central part of the tooth base is pierced by a resorption pit. REMARKS Although the dentary described here possesses some similarities with the above described lacertid material, e.g. bicuspid teeth, several important differences can be observed: 1) large size; 2) the high dental crest; 3) small interdental gaps; 4) more robust teeth; and 5) low number of lingual striae on the tooth crowns. Because not all of those differences can be explained by ontogenetic changes, we suggest the presence of at least two lacertid taxa in MWQ. Unranked clade AMPHISBAENIA Gray, 1844 AMPHISBAENIA indet. (Fig. 3) Blanus sp. – Ivanov et al. 2006: 229, table 2. MATERIAL. — MWQ, early Miocene, Burdigalian, Orleanian, MN 4: 2/2003 Reptile Joint: One trunk vertebra (Pal. 1570). DESCRIPTION Trunk vertebra A single trunk vertebra is preserved. It is small in size. A neural spine is absent, and the dorsal portion of the neural arch forms a median edge. In la, Published as part of Ivanov, Martin, Čerňanský, Andrej, Bonilla-Salomón, Isaac & Luján, Àngel Hernández, 2020, Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications, pp. 343-376 in Geodiversitas 42 (20) on pages 346-349, DOI: 10.5252/geodiversitas2020v42a20, http://zenodo.org/record/4447563, {"references":["RAGE J. - C. & AUGE M. L. 2010. - Squamate reptiles from the middle Eocene of Lissieu (France). A landmark in the middle Eocene of Europe. Geobios 43 (2): 253 - 268. https: // doi. org / 10.1016 / j. geobios. 2009.08.002","MULLER J. 1996. - Eine neue Art der Echten Eidechsen (Reptilia: Lacertilia: Lacertidae) aus dem Unteren Miozan. Mainzer Geowissenschaftliche Mitteilungen 25: 79 - 88.","CERNANSKY A., RAGE J. C. & KLEMBARA J. 2015. - The Early Miocene squamates of Amoneburg (Germany): the first stages of modern squamates in Europe. 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