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A fragment of the posterior right portion of parietal Pb 02024 of the green lizard Lacerta group L. viridis: A-dorsal view; B-ventral view. Scale 2 mm.

A fragment of the posterior right portion of parietal Pb 02024 of the green lizard Lacerta group L. viridis: A-dorsal view; B-ventral view. Scale 2 mm.

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The earliest world record of the green lizards, Lacerta viridis group, is described from the lower Miocene of Central Europe. The fossils come from greenish, calcareous marls and limnic clayey silts of the Ottnangian zone MN 4 of the Dolnice locality near Cheb in the Czech Republic. Sediments are interpreted as marginal, riparian facies. The materi...

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Context 1
... development, whereas Pb 02054 undoubtedly represents an adult on- togenetic stage. Parietal. The parietal is a large azygous element con- sisting of the parietal plate with an ornamented surface, and two posterolaterally diverging supratemporal pro- cesses. Description is based on a fragment Pb 02024 of the right posterior portion of the parietal (Fig. 2). The ornamentation of the parietal plate is identical to that of the frontals described above (Fig. 2A). The pre- served portion of parietal plate is divided by a distinct, medially arched groove. It demarcates the border be- tween the parietal and interparietal osteodermal plate. The occipital plate is not preserved. The supratempo- ...
Context 2
... is a large azygous element con- sisting of the parietal plate with an ornamented surface, and two posterolaterally diverging supratemporal pro- cesses. Description is based on a fragment Pb 02024 of the right posterior portion of the parietal (Fig. 2). The ornamentation of the parietal plate is identical to that of the frontals described above (Fig. 2A). The pre- served portion of parietal plate is divided by a distinct, medially arched groove. It demarcates the border be- tween the parietal and interparietal osteodermal plate. The occipital plate is not preserved. The supratempo- ral process continues posterolaterally. However, it is in- complete. The distal tip is broken and ...
Context 3
... of the parietal, the crista cranii parie- talis runs along the lateral margin of the supratemporal process. The crista cranii continues at the level of the parietal plate and becomes gradually less well devel- oped. The rest is broken and missing. The crista cranii forms the border between a medial depression and a lateral facies semilunaris (Fig. ...

Citations

... For example, the influence of the African-Eurasian terrestrial contact, which occurred during the Early Miocene, remains poorly understood. In regards to the later stages of the Early Miocene (MN 3-4), the lizard faunas are well known from Central and other parts of Europe (Roček, 1984;Schleich, 1988;Fejfar and Schleich, 1994;Klembara, 2008Klembara, , 2015Č erň anský , 2010a, 2010bČ erň anský and Bauer, 2010;Č erň anský et al., 2015aČ erň anský et al., , Georgalis et al., 2016aVasilyan et al., 2022), but they are more-or-less only occasionally documented from the Western part of the continent (Rage and Bailon, 2005). The data indicates that the main event occurred during the Early Burdigalian (Early Orleanian in terms of continental stratigraphy), and more precisely in zones MN 3 and MN 4, when a large wave of immigrants entered Europe (Ivanov 2001(Ivanov , 2002Szyndlar and Rage, 2003;Č erň anský , 2010aČ erň anský et al., 2015;Rage, 2013). ...
... Besides these, two other species of the genus were described from the Early Miocene: Ophisaurus fejfari Klembara, 1979 and Ophisaurus robustus (Klembara, 1979), e.g., from the locality Dolnice (MN 4; see Klembara and Rummel, 2018). Lacertids are also described from these localities (Č erň anský and Joniak, 2009;Č erň anský , 2010b, Č erň anský et al., 2015a. Interestingly, chameleonids and cordylids are not documented in the Crémat material. ...
Article
Squamate faunas from the MN 1–3 intervals (earliest Miocene) are scarcely documented from the European continent. I here describe squamate faunas from two French localities – Montaigu-le-Blin (MN 2) and Crémat (MN 3), the latter being the youngest locality of the Phosphorites du Quercy. The palaeobiodiversity of squamates from these sites is low relative to the faunas described from coeval localities of Amöneburg (MN 2) in Germany and Merkur-North (MN 3) in the Czech Republic. The beginning of the Miocene represents the temporary return to a paratropical humid climate after the relatively cool and dry Oligocene, and the Montaigu and Crémat materials provide previously undocumented components of herpetofaunas and their changes during this crucial time interval in France. The importance of fossil squamates from the area of the Phosphorites du Quercy is therefore highlighted for the beginning of the Miocene as well as the better known Eocene-Oligocene sites. The lizard material of Montaigu includes lacertids and blanids, the latter being represented by Blanus cf. gracilis, one of the oldest records of the genus. Moreover, it shows a higher spatial and temporal distribution of this amphisbaenian species during the Early Miocene than previously known. The material from Crémat consists of few elements which can be allocated to anguids and potentially to lacertids. Detailed figures of the specimens are provided through the means of both photography and micro-CT scanning.
... In older, Eocene and Oligocene localities in the neighbouring north-western Czech Republic and south-eastern Germany (Saxony and southeastern Saxony-Anhalt), rich assemblages of terrestrial-aquatic tetrapod fauna have been documented, comprising frogs, salamanders, choristoderans, crocodiles (also crocodile coprolites, see Kasi nski et al. (2015), and literature cited therein), turtles, lizards, and snakes (Table 1). On the other hand, in the lower Miocene clays and sands of North Bohemian Brown Coal Basin, a very rich fauna assemblage was reported (Klembara, 1979(Klembara, , 1981Roček, 1984;Szyndlar, 1991aSzyndlar, , 1991bSzyndlar & Schleich, 1993;Szyndlar & Rage, 2003;Čerňanský, 2010a, 2010bDvořák et al., 2010). The latter mentioned and illustrated numerous vertebrates represented by osteichthyan fish, amphibians, reptiles, birds, and mammals. ...
... The chemical composition, morphology of ferruginous masses matrix and microstructures topography were investigated using the desktop scanning electron microscopy (SEM) Phenom XL, Phenom World (Thermo Fisher Scientific, Eindhoven, Netherlands) equipped with a fully integrated energy-dispersive X-ray spectroscopy (EDS) detector and secondary electron detector (SED) located in the Faculty of Natural Sciences at the Table 2 Reptiles recorded in the lower Miocene deposits of North Bohemia, Czech Republic (taken from Klembara, 1981Klembara, , 2008Klembara, , 2012Klembara, , 2015Młynarski & Roček, 1985;Ivanov, 2002;Čerňanský & Joniak, 2009;Čerňanský, 2010a, 2010bČerňanský & Bauer, 2010;Čerňanský, Rage & Klembara, 2015;Dvořák et al., 2010;Joyce, 2016;Georgalis & Joyce, 2017;Klembara & Rummel, 2018;Chroust et al., 2021). University of Silesia in Katowice (Bankowa, Katowice, Poland) (Fig. 5). ...
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Excrement-shaped ferruginous masses have been recovered from the Miocene of Turów mine in south-western Poland. These siderite masses have been the subject of much controversy, having been interpreted either as being coprolites, cololithes or pseudofossils created by mechanical deformation of plastic sediment. Here we present the results of mineralogical, geochemical, petrographic and microtomographical analyses. Our data indicate that these masses consist of siderite and iron oxide rather than phosphate, and rarely contain recognizable food residues, which may suggest abiotic origins of these structures. On the other hand, evidence in support of a fecal origin include: (i) the presence of two distinct morphotypes differing in size and shape, (ii) the presence of rare hair-like structures or coalified inclusions and (iii) the presence of rare fine striations on the surface. Importantly, comparative actualistic study of recent vertebrate feces shows overall resemblance of the first morphotype (sausage-shaped with rare coalified debris) to excrements of testudinoid turtles (Testudinoidea), whose shell fragment was found in the investigated locality. The second morphotype (rounded to oval-shaped with hair-like structures), in turn, is similar to the feces of some snakes (Serpentes), the remains of which were noted in the Miocene of the neighborhood areas. Other potential producers (such as lizards and crocodiles) and even abiotic origins cannot be fully excluded but are less likely
... SZYNDLAR and SCHLEICH 1993;SZYNDLAR and BÖHME 1993;SZYNDLAR and RAGE 2003;SZYNDLAR 2009;IVANOV and BÖHME 2011) and several localities in Czech Republic (e.g. ROČEK 1984;SZYND LAR 1987;ČERŇANSKÝ 2010aČERŇANSKÝ , 2010bIVANOV et al. 2020) and Austria (SZYNDLAR 1998, ČERŇANSKÝ 2016. Although Aquitanian and early Burdigalian (MN 1 -MN 3) squa mates are still poorly known in Europe (GEORGALIS and SCHEYER 2021), at least four Central European localities provided rather diversified communities including the German localities Amöneburg (MN 2;ČERŇANSKÝ et al. 2015), Ulm-Westtangente (MN 2; ČERŇAN-SKÝ et al. 2016;KLEMBARA et al. 2019) and Wintershof-West (MN 3;IVANOV 2015;PACLÍK et al. 2018) and the Czech locality of Merkur-North (MN 3a;e.g. ...
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Squamate assemblages of the earliest Miocene (Aquitanian) are extremely rare in Europe due to the lack of well-documented (or dated) localities. We describe a small collection of squamates from the early Miocene (MN 1–2?) locality of Weißenburg 6, Germany. The community consists of four squamate taxa including Lacertidae indet., cf. Eoanilius sp., ‘Colubridae’ gen. et sp. indet. and Viperidae gen. et sp. indet. The presence of Eoanilius indicates that thermophilic ‘anilioid’, whose extant relatives occur under the tropical humid climate, might have survived locally the global decrease in temperatures around the Oligocene/Miocene transition (23.03 Ma). The ‘colubrid’ snake from Weißenburg 6 differs from known late Palaeogene and earliest Miocene Colubroidea (Colubridae + Natricidae). Although colubroids were apparently common in Europe since the earliest Miocene (Aquitanian), the first significant post-Oligocene dispersal of Colubroidea from Asia and/or North Africa into Europe, accompanied by a quick dispersal of Viperidae (both ‘European’ and ‘Oriental’ vipers) and small Elapoidea, can be dated to the early Budrigalian (MN 3). The indeterminate viperid from Weißenburg 6 is among the oldest known fossil record of Viperidae.
... Several major clades can be recognized: Gekkota, Chamaeleonidae, Anguidae and Scincidae. Overall, the Echzell lizard assemblage is similar to other European localities from MN4, e.g., Dolnice and Mokrá Western-Quarry in the Czech Republic (see Roček 1984;Čerňanský 2010b, 2010cIvanov et al. 2020) and Oberdorf in Austria (Čerňanský 2016) (see Table 2). Interestingly, amphisbaenians, varanids, cordylids and the genus Pseudopus were not recognized in the material available to us. ...
... Interestingly, amphisbaenians, varanids, cordylids and the genus Pseudopus were not recognized in the material available to us. Lacertids are very limited in Echzell based on the material available to us, although they are very abundant in other localities of this age (Čerňanský 2010cIvanov et al. 2020). The missing group represents large-sized animals, which might point to potential taphonomic and/or sampling biases. ...
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The present study describes a rich amphibian and reptilian assemblage from the early Miocene locality Echzell, Germany. It consists of one allocaudate, five salamander, five frog, one gecko, chamaeleonids, anguine lizards, one lacertid, one skink and five snake taxa. The entire herpetofauna of Echzell is represented by genera and/or families very broadly known from the early Miocene of Europe. Contrary to other early Miocene herpetofaunas, the Echzell assemblage includes surprisingly only one form of crocodile-newts (Chelotriton). The Echzell Palaeobatrachus robustus represents the youngest record of the species and extends its stratigraphic range to the late early Miocene. Regarding chameleons, the frontal is partly preserved, but represents the first described frontal of the extinct species Chamaeleo andrusovi. The only anguine lizard that can be identified in the assemblage is represented by a new genus and species Smithosaurus echzellensis. Our phylogenetic analyses consistently recovered it as the sister taxon to either [Ophisauriscus quadrupes + Ophisaurus holeci] + [Anguis + Ophisaurus] (in the first analysis) or [Anguis + Ophisaurus] (in the second analysis). However, the results are based on limited fossil material-the parietal-and the support for the clade is very low. Thus, the interpretation of the Smithosaurus relationship among anguines needs to be taken with caution and has to be tested in further studies. Among snakes, Natrix longivertebrata represents the oldest record of the species and extends the stratigraphic range of this fossil snake back to the early Miocene. In addition, we provide here a broader comparison of the Echzell amphibian and reptilian assemblage with their European records for the MN3 and MN4 biostratigraphical units. Besides that, the entire herpetofauna of Echzell includes very broadly known early Miocene European forms. Remains of other groups of the same period such as Bufonidae, Hylidae, Pelodytidae, Amphisbaenia, Varanidae, Cordylidae, Pseudopus, are not found in the material available to us. We also conclude that the amphibian and reptilian fossil record across MN3-MN4 is significantly biased by taphonomic and/or environmental conditions. The amphibian and reptilian assemblage of Echzell is rich in forms living in humid and warm environments with forested areas, permanent water bodies and also some open habitats. The following climatic parameters can be reconstructed based on the herpetofauna: a mean annual temperature of 17.4-28.8 °C, minimal warm month temperature 18-28.3 °C, minimal cold month temperature 8-22.2 °C, and mean annual precipitation with a value of 791±254 mm.
... The emergence of the Canary Island of El Hierro representing the divergence of Gallotia caesaris caesaris and Gallotia caesaris gomerae (1.0 Mya [40]) was used selecting a normal prior distribution (sigma = 0.02). The split between Lacerta and Timon was dated based using as a prior a gamma distribution based on a minimum age of 17.5 Mya [41] with shape and scale set to 1.0. The same prior distribution was used to calibrate the divergence between L. viridis and L. bilineata 8.7 Mya [42]. ...
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South-western Europe has a rich diversity of lacertid lizards. In this study, we evaluated the occupancy patterns and niche segregation of five species of lacertids, focusing on large-bodied species (i.e., adults having ˃75 mm snout-vent length) that occur in south-western Europe (Italian to the Iberian Peninsula). We characterized the niches occupied by these species based on climate and vegetation cover properties. We expected some commonality among phylogenetically related species, but also patterns of habitat segregation mitigating competition between ecologically equivalent species. We used multivariate ordination and probabilistic methods to describe the occupancy patterns and evaluated niche evolution through phylogenetic analyses. Our results showed climate niche partitioning, but with a wide overlap in transitional zones, where segregation is maintained by species-specific responses to the vegetation cover. The analyses also showed that phylogenetically related species tend to share large parts of their habitat niches. The occurrence of independent evolutionary lineages contributed to the regional species richness favored by a long history of niche divergence.
... 7) The Paleogene Plesiolacerta lydekkeri 29 from France, scored for 19 characters for dentary, maxilla, frontal, and parietal. 8) "Lacerta viridis group" from the early Miocene of Dolnice near Cheb, Czech Republic, scored for 6 characters for the frontal (see also 30 ). 9) Pseudeumeces cardurcensis 31 from the early Oligocene of France 32 . ...
... Ma) is based on the oldest-known reliable fossil of the Lacerta agilis lineage, which comes from the Pliocene of Ivanovce, Slovakia 40 . We decided not to use the fossil lacertids originally described as members of the Lacerta viridis group from the Lower Miocene of the Czech Republic for calibration 30 , although they had been previously used for molecular dating 41 , because of their uncertain position in our phylogenetic analysis. Also in the original description, the systematic placement of these fossils was not tested phylogenetically. ...
... This is not present in other Solnechnodolsk specimens, where this facet is reduced. The facet for the maxilla in GIN 1145/277 forms a similar condition to that in adult individuals of extant species L. pamphylica studied herein and to the frontals described from the lower Miocene of the Czech Republic [18]. The frontals of extant European species of Lacerta usually have either a reduced or absent articulation facet for the maxilla (in dorsal Parietal. ...
... The anterior region in front of the sulcus interfacialis is long, forming more-or-less 2/3 of the anteroposterior length of the frontal. In Timon lepidus, this anterior region of the frontal is short and the ratio of the anteroposterior length of the frontal shield vs. frontoparietal shield on frontal is approximatelly 1:1 (see [18]; or Fig 34A here). (2) Premaxilla bears nine teeth. ...
... Lacertids are often reported from the fossil record of Europe [18,54,[74][75][76][77] and from areas adjacent to Solnechnodolsk (the Black Sea area), including lacertid material previously described from the early Pliocene of Turkey. However due to the fragmentary nature of the elements, they were allocated only to cf. ...
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We here describe the first fossil remains of a green lizard of the Lacerta group from the late Miocene (MN 13) of the Solnechnodolsk locality in southern European Russia. This region of Europe is crucial for our understanding of the paleobiogeography and evolution of these middle-sized lizards. Although this clade has a broad geographical distribution across the continent today, its presence in the fossil record has only rarely been reported. In contrast to that, the material described here is abundant, consists of a premaxilla, maxillae, frontals, parietals, jugals, quadrate, pterygoids, dentaries and vertebrae. The comparison of these elements to all extant green lizard species shows that these fossils are indistinguishable from Lacerta trilineata. Thus, they form the first potential evidence of the occurrence of this species in the Miocene. This may be also used as a potential calibration point for further studies. Together with other lizard fossils, Solnechnodolsk shows an interesting combination of survivors and the dawn of modern species. This locality provides important evidence for the transition of an archaic Miocene world to the modern diversity of lizards in Europe. In addition, this article represents a contribution to the knowledge of the comparative osteological anatomy of the selected cranial elements in lacertids. This study gives special emphasis to the green lizards, but new data are also presented for related taxa, e.g., Timon lepidus, Podarcis muralis or Zootoca vivipara. Although the green lizards include several cryptic species for which determination based on isolated osteological material would be expected to be difficult, our comparisons show several important morphological differences, although a high degree of variability is present.
... frontoparietal shield on frontal is approximatelly 1:1 (see [18]; or Fig. 25K here). ...
... doi: bioRxiv preprint first posted online Apr. 17, 2019; Lacertids are often reported from the fossil record of Europe [18,54,[72][73][74][75] and from areas adjacent to Solnechnodolsk (the Black Sea area), including lacertid material previously described from the early Pliocene of Turkey. However due to the fragmentary nature of the elements, they were allocated only to cf. ...
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We here describe the first fossil remains of a green lizardof the Lacerta group from the late Miocene (MN 13) of the Solnechnodolsk locality in southern European Russia. This region of Europe is crucial for our understanding of the paleobiogeography and evolution of these middle-sized lizards. Although this clade has a broad geographical distribution across the continent today, its presence in the fossil record has only rarely been reported. In contrast to that, the material described here is abundant, consists of a premaxilla, maxillae, frontals, parietals, jugals, quadrate, pterygoids, dentaries and vertebrae. The comparison of these elements to all extant green lizard species shows that these fossils are indistinguishable from Lacerta trilineata. Thus, they form the first potential evidence of the occurrence of this species in the Miocene. This may be also used as a potential calibration point for further studies. Together with other lizard fossils, Solnechnodolsk shows an interesting combination of survivors and the dawn of modern species. This locality provides important evidence for the transition of an archaic Miocene world to the modern diversity of lizards in Europe. In addition, this article represents a contribution to the knowledge of the comparative osteological anatomy of the selected cranial elements in lacertid. This study gives special emphasis to the green lizards, but new data are also presented for related taxa, e.g., Timon lepidus, Podarcis muralis or Zootoca vivipara. Although the green lizards include several cryptic species for which determination based on isolated osteological material would be expected to be difficult, our comparisons show several important morphological differences.
... They remained an important component of the European fauna since then, as is testified by the high number of fossils found on the continent documenting a wide variety of different morphologies and taxa (see e.g., Rage 2013 and reference therein). The modern European lizard fauna started to set up in the Neogene, with the first representatives of taxa closely related to modern ones appearing in the early Miocene (Č erňanský 2010b;Rage 2013;Č erňanský et al. 2015). Neogene and Quaternary fossils are, therefore, pivotal to understand the origin and evolution of the current lizard assemblage in Europe. ...
Article
Lizards were and still are an important component of the European herpetofauna. The modern European lizard fauna started to set up in the Miocene and a rich fossil record is known from Neogene and Quaternary sites. At least 12 lizard and worm lizard families are represented in the European fossil record of the last 23 Ma. The record comprises more than 3000 occurrences from more than 800 localities, mainly of Miocene and Pleistocene age. By the beginning of the Neogene, a marked faunistic change is detectable compared to the lizard fossil record of Palaeogene Europe. This change is reflected by other squamates as well and might be related to an environmental deterioration occurring roughly at the Oligocene/Miocene boundary. Nevertheless, the diversity was still rather high in the Neogene and started to decrease with the onset of the Quaternary glacial cycles. This led to the current impoverished lizard fauna, with the southward range shrinking of the most thermophilic taxa (e.g., agamids, amphisbaenians) and the local disappearance of other groups (e.g., varanids). Our overview of the known fossil record of European Neogene and Quaternary lizards and worm lizards highlighted a substantial number of either unpublished or poorly known occurrences often referred to wastebasket taxa. A proper study of these and other remains, as well as a better sampling of poorly explored time ranges (e.g., Pliocene, Holocene), is needed and would be of utmost importance to better understand the evolutionary history of these reptiles in Europe.
... According to Vidal and Hedges (2009), Lacertidae diverged from their sister lineage before the Mesozoic/Cenozoic boundary, although their fossil record is unknown in the Mesozoic and doubtful in the Paleocene ( Rage, 2013). On the other hand, fossils of this clade are well documented in the post-Paleocene Cenozoic of Europe (e.g., Ro cek, 1984;Aug e, 2005;Cer nansk y, 2010;Cer nansk y et al., 2015). Most finds are, however, represented by disarticulated elements. ...
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The best-preserved material of Dracaenosaurus croizeti, an almost complete and previously unpublished skull with a few associated postcranial bones (stylopodium, zeugopodium, and cervical vertebra), is described. The material comes from the locality of Cournon, a late Oligocene site in south-central France. Micro-computed tomography applied to this specimen revealed previously unknown internal osteological characters. Among lacertids, this taxon represents a notable phenomenon: it is an extreme durophagous specialist. Many of the newly observed cranial character states reflect the lifestyle of this lizard, because animals with a hard-shelled diet display a specialized cranial morphology associated with more massive cranial muscles. One unique character for Lacertidae is observed: the parietal-supraoccipital contact is formed by a ventrally deep parietal crest that fits into a bifurcate ascending process of the supraoccipital. In fact, such a connection represents the opposite to the connection in modern members of Lacertidae. Phylogenetic analysis recovered Dracaenosaurus inside Gallotiinae, a clade that would consist of the mainly Oligocene genera Pseudeumeces and Dracaenosaurus, the Miocene genus Janosikia, and the extant Psammodromus and Gallotia. Our study supports previous phylogenetic results and provides an example of the achievement of large size in mainland members of the stem of Gallotia, previously exemplified by Janosikia and Pseudeumeces. The extreme amblyodonty of Dracaenosaurus also confirms the view that herbivory in Gallotia is derived and may be the result of insularity. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP Citation for this article: Čerňanský, A., A. Bolet, J. Müller, J.-C. Rage, M. Augé, and A. Herrel. 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. DOI: 10.1080/02724634.2017.1384738.