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.We here describe the first fossil remains of a green lizard of the  Lacerta. Molecular and morphological phylogenetic analyses show that Lacerta is the sister taxon to Timon [Although the fossil record of squamates is well documented in western and central Europe, many aspects of eastern Europe remain unknown. However, this area can be crucial for our understanding of the biogeography and evolution of this currently dominant group of non-avian reptiles –3. We heto Timon –17, 6.Lacerta cf.viridis have been described on the basis of several isolated jaw elements from the late Miocene (MN 11) localities of Kohfidish in Austria [L. agilis, L. schreiberi , L. strigata (the Caucasus green lizard), L. trilineata  and L. viridis. Based on morphological studies [L. trilineata was recently split into three species, L. trilineata, L. media (Levant green lizard) and L. pamphylica (Turkisch pamphylic green lizard). Within Lacerta viridis clade, two main lineages were recognized—Western European lineage of L. bilineata and Eastern European L. viridis [Lacerta is a comparative study of extant taxa for the purpose of discovering characters useful in species level identification.The oldest finds of green lizards are documented from the lower Miocene (MN 4) locality Dolnice near Cheb in the Czech Republic . The old Austria . and Pol Austria . Nowaday Austria . However Austria  for whic Austria . Even th Austria  already  studies , L. tril viridis . The tax viridis , 25–27.  viridis , is almo viridis , 27–28. Pelobates, amphisbaenians and anguimorphs have recently been described [The Solnechnodolsk locality  is situated in the Northern Caucasus, 40 km NW of the city of Stavropol. In paleogeographical terms, it is located at the southern shore of the Pontian marine basin. The vertebrate fossils occurred as disassociated bones coming from fluviatile and lacustrine beds incised in middle Sarmatian (Bessarabian) limestones. Mostly preliminary accounts of the fauna have been so far published –31. The escribed . In addiThe aims of this paper are as follows: (1) to describe the fossil lacertid material from the late Miocene of Solnechnodolsk in detail; (2) to compare it with other lacertids, with the main focus on extant green lizards; (3) to do a comparative anatomy of selected cranial elements useful for diagnostic purposes in extant lizards. This will serve as a reference for other comparative anatomical studies of any other lacertids extant, or extinct .The lizard specimens described here are housed in the Geological Institute of the Russian Academy of Sciences, Moscow, Russia, prefixed under individual GIN numbers. Since the site discovery in 2009, the material was sampled in 2009, 2010, 2014, and 2017 by expeditions from of the Russian Academy of Sciences. Fossils were extracted from sediments by screen washing with the mesh size 0.5, 0.7, and 1 mm. The excavated vertebrate material includes fishes, amphibians, reptiles, birds, small and large mammals.Lacerta viridis , L. schreiberi , L. bilineata , L. pamphylica , L. strigata , L. trilineata , L. media , L. agilis , Timon lepidus (NHMV 10921–1), Podarcis muralis , Zootoca vivipara (NHMV 32438–1), Takydromus sexlineatus (DE 134), Meroles ctenodactylus (NHMV 31376); Psammodromus algirus (NHMV36038-2) and Gallotia stehlini (NHMV 11031–1).The following specimens of extant lizard species were used for comparison: Institutional abbreviations: DE, Department of Ecology, Comenius University in Bratislava, Slovakia; GIN, Geological Institute of the Russian Academy of Sciences, Moscow, Russia; NHMV, Museum of Natural History, Vienna; UF, University of Florida, Florida Museum of Natural History (USA); UMJGP, Universalmuseum Joanneum, Graz, Austria; ZSM, The Bavarian State Collection of Zoology, Munich, Germany.Lacerta viridis (DE 51) were photographed under a Leica M125 binocular microscope with axially mounted DFC 500 camera LAS software (Leica Application Suite) version 4.1.0 (build 1264).The specimens of extant lacertids see  were scaThe standard anatomical orientation system is used throughout this paper. The image processing program ImageJ  was usedSquamata Oppel, 1811 Lacertoidea Estes, Queiroz and Gauthier, 1988 Lacertidae Oppel, 1811 Lacerta Linnaeus, 1758 [us, 1758 Lacerta cf. trilineata Bedriaga, 1886 [ga, 1886 Figs 7Material––One premaxilla GIN 1145/270, six right maxillae 1145/271-276, five right frontals 1145/277-281, one left frontal 1145/282, four parietals 1145/283-285, 304, one left prefrontal 1145/286, two right jugals 1145/287-288, one left jugal 1145/289, two right postfrontals 1145/290-291, one right quadrate 1145/292, one right pterygoid 1145/293, two left dentaries 1145/294-295, four right dentaries 1145/296-299.Locality and horizon––Solnechnodolsk ; upper Miocene, MN 13.The premaxilla is a triradiate element . The nasMaxilla. The description is based on the specimen GIN 1145/271 , which The lateral face of the maxilla is pierced by 6 larger labial foramina arranged in a single line and several smaller, scattered in the anterior region. The posteriormost one of the larger series is located at the level between the 13th and 14th tooth position (counted from anterior) in 1145/271, or at the level of 7th tooth position (counted from posterior) in 1145/272. The nasal process bears attached osteodermal shields. At least two well separated osteoderms fused to lateral side of the ventral portion of the nasal process of maxilla can be identified here. They are seaparated by a sulcus. This sulcus has a posteroventral course and virtually points to the 4th labial foramen (counted from anterior). The sculpture of these osteoderms consists of densely spaced grooves, pits and ridges. The anteriorly located osteoderm is large. The length of its ventral margin forms 1/3 of the entire length of the sculptured region. The posteroventral process of maxilla does not gradually narrow to its end, but it is stepped. The notch located between the dorsal and ventral portions of the posteroventral process is shallow, weakly developed . The posPrefrontal. The left prefrontal is preserved  individual. This is also supported by the apparent presence of somewhat stronger degree of mid-constriction . The facet for the maxilla is well-developed here and reaches even further posteriorly than the facet for the nasal. 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 [Lacerta usually have either a reduced or absent articulation facet for the maxilla . However in some cases, this facet can have a similar morphology to that of GIN 1145/277, e.g. the left frontal of L. viridis NHMV 40137. For this reason, we regard this difference between GIN 1145/277 and other Solnechnodolsk fossil frontals as a result of individual and/or ontogenetic variation.Republic . The froParietal. All parietals are incompletely preserved (Timon). Laterally located parietal shields appear to be large, however they are not completely preserved. Only the right one shows the lateral margin, which appears to be slightly concave medially. The frontoparietal shields are only partly preserved on the left side in 1145/283. However, this region is better preserved in 1145/285, showing that the interparietal shield was entirely restricted to the parietal bone ; upper Miocene, MN 13.Three dorsal vertebrae are preserved, of which the specimen GIN 1145/300 is almost completely preserved . In lateRemarks: The dorsal vertebrae of green lizards have tendency to uniform morphology, which encourages caution in assigning isolated vertebrae to particular species. The dorsal vertebrae from Solnechnodolsk show a high similarity to that of L. trilineata, rather than to e.g. L. strigata. The posterior centrosynapophyseal lamina is absent in L. strigata and the synapophysis here is small . There All fossil bones described here can be allocated to Lacertidae without doubt. They exhibit the following synapomorphies of Lacertidae, e.g., sulcus dentalis, lateral overlap of the posterodorsal margin of the dentary by the coronoid, maxilla-frontal contact, closure of the supratemporal fenestra, interparietal shield is restricted on parietal bone, and parietal cranial crests meet medially , 45. ThuLacerta agilis is traditionally called the sand lizard, it has been shown that this taxon branches within the green lizards [L. agilis, L. viridis, L. bilineata, L. pamphylica, L. strigata, L. schreiberi, L. trilineata and L. media  and, moreover, to record the distribution of some character states, we also used P. muralis  on the basis of dentition, sculpture of osteodermal shields and great similarity with the morphology presented by the extant green lizards (see below). Moreover they are comparable in size and come from a single site. Green lizards form a uniform group of lacertids. Their fossil record is rather poor see , what is lizards , 26. Forsee Figs –33. Timoson Figs  and 35.  muralis , Zootocavivipara , Takydrolineatus , Merolesdactylus , Psammod algirus  and Gallstehlini .Solnechnodolsk lacertid material can be allocated to green lizards without doubts. This can be supported by the combination of several features, e.g.: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 [(1) The frontal bone. 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 1:1  Premaxilla bears nine teeth. This is true for all green lizards . In T. lepidus, the parietal fossa has a somewhat peculiar morphology, with an additional posterior constriction of the parietal fossa. This gives a lens shape to this structure . However, it never reaches such a strong degree as in T. lepidus . Very unique morphology can be seen in the Oligocene durophagous lacertid specialist Dracaenosaurus croizeti, where the fossa is absent. In contrast, a parietal flange is present here [(3) Parietal fossa. This structure in the posterior ventral section of the parietal forms an area where the ascending process of the supraoccipital fits into a groove . In soml shield , 54. Not 35860–2  and L. sent here .T. lepidus or G. stehlini, but absent in Po. muralis, Z. vivipara, T. sexlineatus and M. ctenodactylus.(4) Medial expansion of the frontal process of the postfrontal, forming a broad area with a contact to frontal, is present. This is present in all green lizards, but is not unique to them . It is aLacerta, Timon, Anatololacerta, Gallotia, and Psammodromus algirus [Po. muralis and Z. vivipara. Among green lizards, the pterygoid teeth show several arrangements. The teeth on the Solnechnodolsk pterygoid are arranged strictly in a single line. This is present in several specimens of green lizards studied here. In others, teeth are arranged in the elliptical area, where the central region is wider; here at least two teeth can be found next to each other . Therefore, these two conditions seem to be a matter of individual (or ontogenetic) variations rather than being taxonomically important among green lizards. The second condition is also present in the specimens of T. lepidus and G. stehlini available to us. In Ps. algirus, teeth on pterygoid are arranged in a wide area and not in a single line. Because the clade Gallotiinae is sister to Lacertinae [(5) Pterygoid dentition. Teeth on the pterygoid are often found in species of  algirus , 55. Amocertinae , 56, we certinae , 57.In the following section, osteological differences which cannot be observed in the Solnechnodolsk fossils, but are present among green lizards, are discussed:Lacerta viridis and L. trilineata  have distictly laterally expanded cheeks relative to other taxa  Lateral expansion of cheeks in adults. In dorsal view, see Figs  and 26.  lepidus .L. agilis , L. pamphylica  and L. media  in L. trilineata seems to be the case of individual variation. The presence of both character states could reflect the topology of L. trilineata within green lizards (see (2) Osteodems covering cheeks laterally see Figs  and 10. geny see . Althoug6 mm see . However6 mm see . One posards see . But allL. schreiberi, fragmented discontinuous osteoderms in cheeks are present in NHMV 10809  evolved independently. Its absence in L. agilis, L. pamphylica and L. media is regarded as reversal.The cheeks are strongly covered by osteoderms in idus see , in contvivipara . In Gall algirus , but thestehlini . However goliath . The chaL. agilis  and L. media. In T. lepidus (G. stehlini (33A), the supraoccipital is completely covered by the parietal in dorsal view. In contrast to that, it is highly exposed in Po. muralis and Z. vivipara. Regarding the posterior region of the parietal, the Z. vivipara + Ta. sexlineatus clade can be characterized by the presence of a smooth area of the parietal table posterior to the ornamented region and by the presence of a short bilobed, posteriorly located process in the posterior mid-region.(3) In dorsal view, the supraoccipital in adults is highly exposed  The width of the supratemporal processes of the parietal Figs  and 20:  lepidus . Thus thstehlini .L. viridis  In lateral view, the skull is slightly depressed in the preorbital region, so the premaxillary region is slightly stepped Figs  and 10 imphylica , L. stri. agilis , as wellilineata . Howevereveloped . The pre lepidus . The dep algirus  and weakstehlini .L. trilineata, L. media, L. pamphylica  The position of the anterior mylohyoid foramen on the splenial relative to the anterior inferior alveolar foramen. The anterior mylohyoid foramen is located ventrally to the anterior inferior alveolar foramen, but its position varies Figs  and 25: L. agilis, L. viridis and L. strigata. It is located at the level of the dorsal process of coronoid or only slightly posterior to it in L. bilineata, L. pamphylica, L. schreiberi, L. trilineata and L. media. This second character state is also present in T. lepidus, whereas it is posteriorly located in Po. muralis and Z. vivipara, but anterior to the dorsal process of coronoid in G. stehlini.(7) The position of the posterior mylohyoid foramen on the angular relative to the top of the dorsal process of coronoid Figs  and 25: L. media in dorsal aspect  Among green lizards, the dentary of l aspect  is markel aspect . Althougped here .Lacerta agilis, L. strigata and L. media; whereas L. viridis is distributed in northern, western and southern regions  of the the Black Sea [L. trilineata can be found in southern and western areas around the Black Sea [L. trilineata among extant species. However, caution is needed here, because the sample size here does not fully allow to discuss the patterns of ontogenetic and sexual variation that must be present in these lizards and which would introduce more variation. For this reason, we decided to allocate this material as L. cf. trilineata. This combination of features is as follows:Today, the area to the east of the Black Sea (where Solnechnodolsk is located) is occupied by lack Sea , 59, 60.lack Sea . The lacL. trilineata. The process is very broad in Lacerta agilis  is similar to that of L. trilineata. However in L. schreiberi , the base of the process is already slightly expanded laterally, so the lateral margins of the mid-region of the process are slightly concave  The nasal process of premaxilla is moderately wide, gradually expanding laterally from its base in a posterodorsal direction. This is identical to that of a agilis , whereassee Figs  and 33K.rrow see . A distispanicus ) represeL. bilineata  The vomerine processes  are moderately long. They are small, not posteriorly expanded in the specimen NHMV 18599–1 of juvenile .L. trilineata . However, the notch appears to be fully absent in L. strigata (only weakly developed in NHMV 10917–1), and L. schreiberi . A strong notch is developed in L. pamphylica. The notch is usually shallow in L. media  The posteroventral process of the maxilla is stepped as it is in all green lizards studied here Figs  and 14. see also ). This c0137 see , but is  18599–1 , but not 18599–1 . This shT. lepidus  The posteroventral process rises anteriorly, gradually continuing to the nasal process. Where the two processes contact, only a small dorsal curvature is present . In L. trilineata, but not in L. media, L. schreiberi, L. pamphylica, L. bilineata. The separation of the anterior osteoderm by a sulcus is present in L. strigata and L. viridis as well. However, the anterior osteoderm in these species is distictly smaller in comparison to the dominant posterior osteoderm. In L. agilis, there is usually an absence of fused osteoderms  The presence of well separated osteoderms fused to the lateral side of the ventral portion of the nasal process of maxilla and the ratio of the anteroposterior length of the anterior osteoderm relative to the posteriorly located sculptured region. The anteriorly located osteoderm is large. The sulcus which separates the anterior osteoderm from the posteriorly located sculptured region, has a posteroventral course virtually pointed to the 4th labial foramen (counted from anterior). The anteroposterior length of its ventral margin forms 1/3 of the entire ventral margin of the sculptured region. Such a condition is present in erms see . In T. lis small .L. trilineata, L. viridis and L. bilineata. It is located at the level of around the 4th–5th tooth position in adults of L. strigata, 6th in L. agilis  The last labial foramen is located at the level of the 7th tooth position (counted from posterior). This is present in adults of L. trilineata, L. schreiberi, L. media and L. agilis. The opposite condition can be found in L. bilineata and L. strigata. These two rami are almost equally developed in L. viridis and L. pamphylica. In T. lepidus, the septomaxillary ramus of the premaxillary process is more robust relative to the external one.(7) The septomaxillary  ramus of the premaxillary process of the maxilla is larger and more robustly developed than the external one. This condition is present in L. trilineata. It is around 75° in L. bilineata. 70° can be found in L. media and L. viridis, whereas it is 67° in L. strigata and L. agilis, 65° in L. pamphylica and 60° in L. schreiberi. It is 60° in T. lepidus. It should be noted that the morphology of the Solnechnodolsk jugal slightly resembles that in the specimen NHMV 18599–1 of L. bilineata . In T. lepidus, the posteroventral process is broad and well developed (see (8) The angle between the central line of the posteroventral and postorbital process of jugal is around 78° in both the fossil taxon and ilineata , but notilineata . In L. blunt see  and thislunt see . A condihreiberi , but heroped see .L. trilineata, L. viridis, L. bilineata, but is somewhat narrow in L. schreiberi, L. strigata, L media, L. pamphylica. In L. agilis, the postorbital process is moderately narrow if compared to others. In T. lepidus, the postorbital process is wide.(9) Width of postorbital process relative to posteroventral process: The postorbital process is wide in the fossil material studied here. The wide postorbital process is present in L. trilineata. The relative length of these two anterior processes varies among species: a) the anterolateral processes are long whereas the anteromedial ones are indistinct in adults of L. viridis, L. bilineata and L. pamphylica. L. bilineata deserves a comment here—both processes are weakly developed and reach more-or-less the same level anteriorly in NHMV 18599–1  The length of the anterolateral process relative to the anteromedial process. Although only the base of the anteromedial process is preserved in the fossil material, it can be estimated that it did not reach the level of the anterior end of the anterolateral process, thus was shorter in comparison to the anterolateral process. This is identical to that of 99–1 see . However99–1 see , whereascies see ; b) the lis Figs  and 28E ata Figs  and 32A.L. trilineata. The most pronounced mid-constriction of the frontals is present in L. strigata and a distinct constriction can be also found in L. media. The widest frontals relative to their antero-posterior length is present in L. schreiberi and L. bilineata. In Timon lepidus and Zootoca vivipara, frontals are wide, however, they are constricted in Podarcis muralis. The marked constriction can be seen in Meroles ctenodactylus and a similarly strong constriction appears to be common in members of the tribe Eremiadini  The lateral mid-constriction of the frontal. The position and level of the constriction is identical to that of rus, see ).L. trilineata . In all other green lizards , or more or less in the mid-region . One exception is the specimen NHMV 35860–2 of L. bilineata, where the parietal foramen is located anteriorly to the mid-length of the interparietal shield. In T. lepidus, the interparietal shield is pierced in its anterior region  On the dorsal surface of the parietal, the interparietal shield is pierced by the parietal foramen in the anterior half of its anteroposterior length. This is identical to ulmensis ). The inL. trilineata, L. pamphylica, L. schreiberi and L. viridis, the interparietal shield is entirely restricted to the parietal by the frontoparietal shields that meet to form a medial suture that interpolates between the anterior margin of the parietal shield and the suture with the frontal bone. In L. agilis and L. bilineata the frontoparietals are in point contact, but this can vary. In L. bilineata, the interparietal reaches the frontoparietal suture in NHMV 35860–1, whereas it is fully restricted to the parietal table in NHMV 35860–2 . In the NHMV 39765 of L. strigata, a small anterior portion of interparietal shield continues on the posterior mid-region of frontals. However, in two other specimens, NHMV 10917–1 and 10917–2, the interparietal shield is completely restricted to the parietal table.(13) In the Solnechnodolsk fossils, as well as in 60–2 see . In L. mL. trilineata is moderately large, based on the specimen NHMV 27665  The occipital shield of the parietal in MV 27665 , occupinMV 27665 . The occMV 27665 . This casee Figs . The larsee Figs  and 32E.see also ) and extL. trilineata, L. media, L. pamphylica and L. viridis. In L. trilineata and L. media, the small mid-constriction is present and the posterior portion is equally wide as the anterior portion. Such constriction is also present in L. viridis and L. pamphylica, but here the posterior portion is slightly wider than the anterior one. In L. agilis (slightly concave margins) and L. bilineata (slightly convex margins or more or less straight margins), the whole parietal narrows anteriorly. In L. schreiberi, the lateral margins are more-or-less straight. In L. strigata, the lateral margins diverge laterally from each other anteriorly, thus the anterior portion appears to be usually slighty wider than the posterior one. In T. lepidus, the parietal table is anteroposteriorly long and slightly narrows anteriorly. The lateral margins of the parietal table are slightly concave in this taxon.(15) The lateral margins of the parietal are not completelly preserved and can be only estimated. However, they appear to have a lateral concave course similar to L. trilineata, L. viridis and L. bilineata and slightly similar condition can be observed in L. media. The diffence here between L. trilineata and L. viridis is in the course of the parietal crest. In L. viridis, the crest on each side continues anterolaterally in a more or less straight course. Thus, the virtual line extending anteriorly from the posterior section of the branch would meet the anterolateral corner of the parietal , the parietal crest on each side is angled approximately at the level of the anterior margin of the parietal foramen. Here, the parietal crest turns more anteromedial relative to its posterior section. Thus, the virtual line extending anteriorly from the posterior section  would not meet the corner of the parietal table, but would finish on its anterolateral side. This is identical to the condition present in the Solnechnodolsk parietal GIN 1145/283  On the ventral surface of the parietal in the Solnechnodolsk fossil lacertid, the parietal cranial crests converge strongly posteromedially, forming a distinct median crest slightly posterior to the parietal foramen (for this character see ). This isee also ). In L. /283 see . The conilineata  might sllica see . The ten 18599–1 . Among lsee Figs  and 28G.L. trilineata, L. viridis, L. bilineata, L. pamphylica, L. strigata and L. media, but not in L. agilis  The ventrolateral ridge in the internal region gradually disappears anteromedially. This character state can be observed in see also ) and in L. media here. This taxon is characterized by very narrow wedge shaped fossa, with an additional process located in the anterior end of the fossa  Parietal fossa—in regard to the Solnechnodolsk parietals, we can exlude L. viridis; see L. agilis, partial fusion of these two elements has occured, with traces of the suture being easily recognized. However in L. schreiberi, the fusion is present in a stronger degree, where traces of the suture can be recognized only in the posterior region of the postorbitofrontal element  In Solnechnodolsk fossils, the postfrontal and postorbital are separated individual elements. This is present in all specimens of green lizards  The tympanic crest, which forms the anterolateral margin of the quadrate, is angled approximatelly in the mid-region. This is present in eri Figs  and 23. L. trilineata, L. viridis and L. media. However in the latter taxon, the quadrate is more robustly built if compared to other taxa. This difference between the width of the dorsal and ventral portion is not so pronounced in other taxa. In T. lepidus, the whole quadrate is robust.(21) In lateral aspects, the ventral half of quadrate  gradually markedly narrows ventrally. Thus this portion is much narrower than the dorsal portion. This condition is identical to that of L. viridis and L. trilineata. In contrast, unicuspidity is restricted only to the first two anterior teeth in L. agilis [(22) Number of unicuspid teeth. In the Solnechnodolsk material, bicuspidity sometimes starts with the 7th tooth, but this can be variable even in extant taxa. Kosma  stated t. agilis .L. trilineata and L. schreiberi, whereas it is present at the level of the 6th tooth position in L. bilineata, L. strigata, L. agilis, L. viridis. The alveolar foramen is located further anteriorly, at the level of the 8th tooth position, in L. pamphylica and at the level between 9th-10th tooth positions in L. media. Although the location of the alveolar foramen can be informative, it should be noted that the positions of the alveolar foramen in lacertids, like virtually all lizards, should not be interpreted as absolute due to its variations.(23) The position of alveolar foramen in dentary at the level of the 7th tooth position (counted from posterior). Such condition is present in Lacerta cf. trilineata—it shows that the Solnechnodolsk lacertid material shares the complete combinations of character states  only with L. trilineata. According to Ahmadzadeh et al. [L. trilineata seems to have evolved and diverged in western Anatolia during the Pliocene or Early Pleistocene. Sagonas et al. [L. viridis and L. bilineata is also dated to the late Miocene (6.78 Mya). This is in a strong contradiction with the recently published results of Kornilios et al. [trilineata+pamphylica, occurred in parallel in the late Pliocene. All three studies used the same calibration points and the same data (mtDNA) for their time estimates, but differed in their inference methods. The Solnechnodolsk material shows support for the hypothesis of Sagonas et al. [L. trilineata already in the late Miocene.The detailed comparison above strongly supports an allocation of the Solnechnodolsk lizard fossil to h et al. , L. trils et al.  suggestes et al. . These as et al.  and formVaranus sp. in Solnechnodolsk suggests a mean annual temperature not less than around 15 °C [Lacerta trilineata occupies Mediterranean-type shrubby vegetation, sandy shores, arable land, pastureland, plantations, and rural gardens. The diet of this taxon consists mainly of insects [Lacerta together with previously-described lizard fossils [Green lizards form a dominant component of the squamate paleofauna from the Solnechnodolsk locality , sh insects , 73. In Lacerta sp. [Lacerta sp.Lacertids are often reported from the fossil record of Europe , 74–77 aerta sp. . A localerta sp. . It shouerta sp. . The lacPlesiolacerta lydekkeri is regarded as in or close to the crown Lacertidae [Timon lepidus [carina maxillaris) starts from the supradental shelf at the level of the 4th tooth position; (7) the course of the maxillary crest—its anterior section runs posteriorly almost in a horizontal level (this gives the appereance to this section as being depressed). It starts to rise posterodorsally at the level of the 6th tooth position; (8) similar ratio of the anteroposterior length of the frontal and frontoparietal shields; (9) fragmentation of the interparietal shield  the size of the elements, indicating animals with a similar large body size.In this section, previously described lacertid fossils from the European Cenozoic, which can may have strong affinities with modern lineages, are discussed. The Eocene taxon certidae . As was certidae , this ta lepidus . These sosition;  similar see e.g. ); (10) lGallotia, e.g. G. stehlini). It seems to be more plausible that such similarities might reflect possible relationship rather than represent just homoplasies. Plesiolacerta is also known from late Oligocene deposits of Germany, where the species P. eratosthenesi is described [Lacerta and Timon dates back to 18.6 Mya [Plesiolacerta has a close relationship to Timon, it would show a much longer evolutionary history of lineage. In comparison with other lizard lineages, this would be not so unusual- one example is the presence of the modern genus Ophisaurus  in the late Eocene [Timon lineage were recognized in deposits close to the Oligocene/Miocene boundary.These are not just shallow similarities, because some features e.g., additional posterior constriction of the parietal fossa, are present only in these two taxa and cannot be only a matter of a large size  from the late middle Miocene of Gratkorn in Austria, allocated to Lacertidae Vasilyan  deservesope, see ). TherefS1 DatasetThe CT data links for Figshare and Morphosource.(DOCX)Click here for additional data file.


Reich, D. Guilligay, A. Pflug, H. Malet, I. Berger, et al.  made an impact on her by providing new grounds to study the influenza virus polymerase and its role in virus biology and evolution.Mathilde Richard works in the field of virology, more specifically on the evolution and pathogenesis of influenza viruses. In this mSphere of Influence article, she reflects on how the two articles “Structure of Influenza A Polymerase Bound to the Viral RNA Promoter” by A. Pflug, D. Guilligay, S. Reich, and S. Cusack  and “Structural Insight into Cap-Snatching and RNA Synthesis by Influenza Polymerase” by S. Reich, D. Guilligay, A. Pflug, H. Malet, I. Berger, et al.  made an impact on her by providing new grounds to study the influenza virus polymerase and its role in virus biology and evolution.Mathilde Richard works in the field of virology, more specifically on the evolution and pathogenesis of influenza viruses. In this mSphere of Influence article, she reflects on how the two articles “Structure of Influenza A Polymerase Bound to the Viral RNA Promoter” by A. Pflug, D. Guilligay, S. Reich, and S. Cusack (Nature 516:355–360, 2014,  In 2014, the Cusack research group described the high-resolution structures of the influenza A virus and influenza B virus polymerases in complex with the viral promoter in two articles published back-to-back in Nature  using the ComplexLINK and MultiBac technologies . The genThis work has influenced me in two ways. First, it taught me that perseverance, resilience, and hard work are immense qualities for a researcher. I remember attending a lecture by Cusack in the late 2000s, early in my Ph.D. trajectory. He presented the structures of fragments of the influenza virus polymerase and reported that his group had been working on resolving the structure of the full complex for years but had been unsuccessful because of the technical difficulties of working with such large protein complexes. When this work was published, years after that lecture, I remember being very impressed and thinking, “that’s it, they’ve done it!” Second, both the technological and theoretical advances from this work opened avenues for my own research. The improvement in the biochemistry of the influenza virus RdRp, i.e., the possibility of purifying and producing it in large and pure quantities, allows the development of new assays to study the RdRp in its full context, in the absence or in the presence of the nucleoprotein. Novel techniques that have been successfully used with other purified recombinant RdRps, such as single-molecule assays , 5, can In a broader perspective, these two articles are a game changer for the influenza virus field. The resolution of influenza virus RdRp structures provides an extraordinary basis for future work, both fundamental and applied. In fundamental studies, similar systems have since been used to resolve the structures of RdRps of other influenza viruses: that of the apo (unbound to promoter) influenza C virus RdRp , the inf


Rhipicephalus evertsi evertsi, 19.4% with Amblyomma hebraeum and 15.6% with Rhipicephalus decoloratus. Rhipicephalus evertsi evertsi was recovered from horses in all nine provinces of South Africa and R. decoloratus in eight provinces. Donkeys were infested with eight tick species, and 81.6% were infested with R. evertsi evertsi, 23.7% with A. hebraeum and 10.5% with R. decoloratus. Several tick species collected from the horses and donkeys are the vectors of economically important diseases of livestock. Rhipicephalus evertsi evertsi is the vector of Theileria equi, the causative organism of equine piroplasmosis. It also transmits Anaplasma marginale, the causative organism of anaplasmosis in cattle. Amblyomma hebraeum is the vector of Ehrlichia ruminantium, the causative organism of heartwater in cattle, sheep and goats, whereas R. decoloratus transmits Babesia bigemina, the causative organism of babesiosis in cattle.The aim of the study was to determine the species spectrum of ixodid ticks that infest horses and donkeys in South Africa and to identify those species that act as vectors of disease to domestic livestock. Ticks were collected opportunistically from 391 horses countrywide by their owners or grooms, or by veterinary students and staff at the Faculty of Veterinary Science, University of Pretoria. Ticks were also collected from 76 donkeys in Limpopo Province, 2 in Gauteng Province and 1 in North West province. All the ticks were identified by means of a stereoscopic microscope. Horses were infested with 17 tick species, 72.1% with  Despite the long association between horses, donkeys and humans in South Africa, it is strange that so little attention has been paid to the ixodid ticks with which they are infested. With the exception of surveys on the ticks that infest donkeys in Botswana  and a tick belonging to the Haemapysalis leachi group were identified. Norval and his co-workers in Zimbabwe examined collections from 39 horses and 11 donkeys and identified Amblyomma hebraeum, H. rufipes, Haemapysalis truncatum, R. appendiculatus, R. decoloratus, R. evertsi evertsi, Rhipicephalus kochi, Rhipicephalus sp. , R. simus, Rhipicephalus turanicus and Rhipicephalus zambeziensis in the collections from horses, and A. hebraeum, H. rufipes, R. appendiculatus, R. decoloratus, R. evertsi evertsi and R. simus in those from donkeys  spp., Rhipicephalus muhsamae and a tick they referred to as R. sanguineus from a total of 450 donkeys in a survey conducted in two districts in central Oromia Regional State, Ethiopia. In the same region of Oromia, Kumsa et al. , Cape mountain zebras (Equus zebra zebra) and Hartmann’s mountain zebras (Equus zebra hartmannae) than from domesticated horses or donkeys in South Africa or Namibia  collected ticks from horses presented for treatment at the university’s Equine Clinic. Ticks previously collected from two horses in the Mountain Zebra National Park  and the collection of a male tick from a donkey is further proof of its presence there. R. zambeziensis is present in the Limpopo Valley and adjoining areas (Norval et al. R. appendiculatus. It is also a vector of East Coast fever (Lawrence, Norval & Uilenberg r et al. . Its prer et al.  as R. spR. evertsi evertsi was the most widespread. The prevalence of this tick is potentially important because it is the vector of the causative organisms of equine piroplasmosis. Various other tick species collected from the horses are important vectors of diseases in domestic cattle. Donkeys in Limpopo Province were infested with eight tick species of which R. evertsi evertsi was the most prevalent.Horses examined countrywide were found to be infested with a large variety of tick species of which


Scientific Reports 10.1038/s41598-018-34226-9, published online 25 October 2018Correction to: The original version of this Article contained a typographical error in the spelling of the author Emeline Julie Ribot, which was incorrectly given as Emeline Ribot. This has now been corrected in the PDF and HTML versions of the Article, and in the accompanying Supplementary Information file.The Author Contributions section now reads:O.K. and E.J.R. conceived the idea, carried out the experiments, analyzed the data and wrote the manuscript with assistance from all the authors. J.C.F. participated to the experiments and revised the manuscript. R.D. and S.M. participated to the experiments, contributed to discussions of results, coordinated and supervised the overall project. All authors reviewed and approved the final manuscript.


Mosquito-borne pathogens are important causes of diseases in the Kingdom of Saudi Arabia. Knowledge of the mosquito fauna is needed for the appropriate control of the vectors that transmit the pathogens and prevent the diseases they cause. An important first step is to have an up-to-date list of the species known to be present in the country. Original occurrence records were obtained from published literature and critically scrutinized to compile a list of the mosquito species that occur within the borders of the Kingdom.Fifty-one species have been recorded in the Kingdom; however, the occurrence of two of these species is unlikely. Thus, the mosquito fauna of the Kingdom comprises 49 species that include 18 anophelines and 31 culicines. Published records are provided for each species. Problematic records based on misidentifications and inappropriate sources are discussed and annotated for clarity.Integrated morphological and molecular methods of identification are needed to refine the list of species and accurately document their distributions in the Kingdom. The KSA occupies the major portion of the peninsula (c.1.97 m km2), and includes 13 provinces. Mosquito-borne pathogens cause several diseases in the KSA. Malaria has been considered an endemic disease in the country from as early as 1900 ..OchlerotAe. (Ochlerotatus) detritus 4. Ochlerotatus detritus).Records: , 7 (as OAe. (Ochlerotatus) caspius 5. Ochlerotatus caspius).Records: , 28–40; SubgenusStegomyiaTheobald, 1901Ae. (Stegomyia) aegypti 6. Stegomyia aegypti).Records: , 41–49; Ae. (Stegomyia) unilineatus 7. Records: , 24, 50.GenusAnophelesMeigen, 1818SubgenusAnophelesMeigen, 1818An. (Anopheles) coustani Laveran, 19008. Records: , 51–54.An. (Anopheles) tenebrosus Dönitz, 19029. An. coustani var. tenebrosus).Records: , 31, 35;SubgenusCelliaTheobald, 1902An. (Cellia) arabiensis Patton, 190510. An. gambiae); ; ;  (requirexiguus); , 28; [30xiguus);   nebulosus Theobald, 190141. Records: , 24, 37.SubgenusMaillotiaTheobald, 1907Cx. (Maillotia) arbieeni Salem, 193842. Records: , 30, 36.Cx. (Maillotia) salisburiensis Theobald, 190143. Records: , 21, 24.SubgenusOculeomyiaTheobald, 1907Cx. (Oculeomyia) bitaeniorhynchus Giles, 190144. Records: , 26, 63.GenusCulisetaFelt, 1894SubgenusAllotheobaldiaBrolemann, 1919Cs.  longiareolata 45. Records: , 39–41.SubgenusCulisetaFelt, 1894Cs. (Culiseta) subochrea 46. Cs. annulata and Cs. subochrea); [ochracea annulata).Records:  (as Cs. ochrea); , 36, 40;ochrea);   tigripes 47. Cx. tigripes); [Records: , 22, 30 gripes); , 26, 36.GenusCoquillettidiaDyar, 1905Cq. (Coquillettidia) richiardii ?48. Mansonia sp.).Record:   unguiculata pefflyi Stone, 196149. Ur. unguiculata); [Records: , 31, 35 culata); .The ecological diversity of the KSA allows multiple opportunities for the evolution of variation in both mosquitoes and their pathogens. Large-scale movements of people, including the annual pilgrimage to Mecca, large-scale use of foreign workers and new developmental projects engender environmental heterogeneity and complex epidemiology. The wide variation in ecological settings may complicate the understanding of spatial/temporal characteristics and population dynamics of mosquito vectors of pathogens that cause diseases in humans.Anopheles species complexes in the country, including the An. gambiae, An. fluviatilis, An. culicifacies and An. subpictus complexes. The Afrotropical An. gambiae complex consists of eight species, which includes malaria vectors  and non-vectors (An. quadriannulatus and An. amharicus) [Anopheles arabiensis is the only member of the complex with a range that extends outside of Africa into the southwestern and western areas of the Arabian Peninsula, occurring in the KSA and Yemen, where it is the major malaria vector [An. arabiensis (often as An. gambiae) in different life stages [An. arabiensis (GenBank: KM068071) is the only member of An. gambiae complex present in the KSA [Investigators may be reluctant to conduct ecological studies in the KSA because field work is labour-intensive and time-consuming. Moreover, due to habitat heterogeneity, which in the KSA is often extreme, there are species of only seven mosquito genera in the country. Here we present the first complete list (see above) of the mosquito species that definitely are known to occur in the KSA. In some cases, ecological and morphological studies have shown that some species are members of species complexes. There are members of four haricus) , 71. Anoa vector –75. Moste stages , 30, 51. the KSA .An. fluviatilis complex is a group of mosquitoes that is widely distributed in southern and southwestern Asia and includes important malaria vectors [Anopheles fluviatilis S is mainly anthropophilic, endophilic and a very competent malaria vector [Anopheles fluviatilis (s.l.) was recorded in the eastern and western regions of the KSA by Mattingly & Knight [The  vectors . The com vectors –81. Anopa vector . Speciesa vector , 82. Ano& Knight  and Al-G& Knight . It is nAn. culicifacies complex consists of five sibling species, informally named species A, B, C, D and E, based on cytotaxonomic and molecular evidence [The evidence , 76, 83.evidence , 84, 85.evidence . Speciesevidence . The othevidence , 88. Avaevidence , but onlevidence . FurtherAn. subpictus complex consists of four species in India, informally denoted as species A, B, C and D, based on morphological, chromosomal and molecular differences [The ferences , 92. As ferences , it is nAnopheles stephensi is the main malaria vector in the eastern region of the KSA, but members of the An. fluviatilis and An. culicifacies complexes, which are important malaria vectors in other parts of Asia [An. stephensi, An. culicifacies (s.l.) and An. fluviatilis (s.l.) consist of different ecoforms that are restricted to rural or urban areas with different degrees of involvement in malaria transmission [Anopheles stephensi is a major vector in Asia and the Middle East, including, in addition to the eastern KSA, areas of Iran, Pakistan, Afghanistan, India and southern China [mysorensis, typical and intermediate forms, which are based and distinguished on egg morphology [mysorensis forms are mainly found in rural environments [mysorensis form is mainly zoophilic and a poor- or non-vector [mysorensis form is an important malaria vector in India [cox1) did not reveal any differences between An. stephensi  in two distant cities of the KSA (Al-Ahsaa and Dammam) where different phenotypes might be present [ of Asia , 55, 74,smission , 76, 85.rn China . It inclrphology , 93–96. ronments . The blon-vector . In contin India . These d present .Anopheles azaniae was first encountered in Saudi Arabia during the collection of mosquitoes for Rift Valley virus isolation in the Jizan Region [An. dthali or An. rhodesiensis rupicolus. Anopheles azaniae was previously known to occur in neighbouring Yemen [An. azaniae [n Region , and subn Region  and Khatn Region . The recn Region , which in Region  is doubtng Yemen , where t azaniae . A singlCoquillettidia collected in the Eastern Province of the KSA was identified as a species of Mansonia in the study of Wills et al. [Cq. (Coquillettidia) buxtoni in Israel and Syria and Cq. (Coq.) richiardii in Iran, Syria and Turkey. With that in mind, it seems likely that the generic identification of Wills et al. [Coquillettidia as a subgenus of Mansonia. In view of the proximity of eastern areas of the KSA to Iran, we are confident that the mosquitoes identified as a species of Mansonia by Wills et al. [Cq. richiardii.A species of the genus s et al. . Accordis et al. , only tws et al.  was bases et al. , which ts et al.  were mosCx. pipiens and Cx. quinquefasciatus in the KSA are problematic. Culex pipiens generally occurs north of a line transecting the KSA between Jeddah in the west and Al-Hasa in the east. Culex quinquefasciatus occurs in the more southerly region of the Arabian Peninsula, with isolated populations of Cx. pipiens occurring in the southwestern mountains [Cx. pipiens or Cx. quinquefasciatus. Larvae and adults collected north of the hybrid zone that have been identified as Cx. quinquefasciatus are misidentified specimens of Cx. pipiens, and those collected south of the zone that have been identified as Cx. pipiens, with the possible exception of those collected in the southwestern mountains, are misidentified specimens of Cx. quinquefasciatus. Future studies based on dissections of male genitalia or DNA sequence analysis of larvae and adults are needed to substantiate species identifications and collection records. Culex quinquefasciatus is an important vector of the microfilariae that cause lymphatic filariasis.Many of the occurrence records for ountains . Introgrountains ). Both sountains , 103. ItCulex torrentium is listed as a species of the KSA in the online catalogue of the Walter Reed Biosystematics Unit [Cx. torrentium in the KSA by these authors is based on unreliable characters of larvae and adult habitus principally using the keys of Harbach [Cx. pipiens complex with certainty based on features of the male genitalia. Additionally, Cx. torrentium is a Palaearctic species whose distribution only extends south into Turkey and possibly into northern Iran [Cx. torrentium in Iraq are doubtful, and the species is not now listed as present in that country [Cx. torrentium is excluded from the list of mosquito species known to occur in the KSA.ics Unit  based onics Unit , 19, Al ics Unit , 28, Kheics Unit , Alahmedics Unit  and Hassics Unit . However Harbach , 5. Thisern Iran , 105. As country , 107. FoCulex of the genus Culex occur in the southwestern Afrotropical Region of the Arabian Peninsula, i.e. Cx. decens, Cx. duttoni, Cx. simpsoni and Cx. univittatus. These species were only known to occur in Yemen [Cx. simpsoni and Cx. univittatus, require confirmation because their identification was based on larvae and adults. They are only reliably distinguished from Cx. sinaiticus and Cx. perexiguus, respectively, based on the morphology of the male genitalia. Records of the two species in more northerly areas of the country are more doubtful.Four species of the subgenus in Yemen , 108 untin Yemen , Jizan  and Harbach (Cx. nebulosus and Cx. perexiguus). Use of the badly outdated keys of Hopkins [Cx. wigglesworthi, is problematic, nonetheless because it is a sub-Sharan species with closest records in southern Sudan, South Sudan and Kenya. Consequently, as it seems likely that the two larvae were misidentified specimens of either Cx. perexiguus or Cx. univittatus, Cx. wigglesworthi is excluded from the list of mosquito species known to occur in the KSA.Since nce 1953 , there ince 1953 , 67. On  Hopkins , Matting Hopkins  and Harb Hopkins , 5. The  Hopkins  for AfriUranotaenia  unguiculata in eastern and southwestern areas of the KSA. The authors, however, were apparently unaware of the earlier description of Ur. unguiculata pefflyi by Stone [Ur. unguiculata unguiculata, is present in the country, only the subspecies described and named by Stone [Alahmed , Alahmedby Stone , and theby Stone . Consequby Stone  is incluCulex, have not confirmed identifications with correlated diagnostic features of the male genitalia. Future studies in the KSA must overcome these shortcomings by individually rearing larvae to obtain adults of both sexes for integrated morphological and molecular study, and use the resultant unequivocal molecular markers for the identification of larvae and adults collected in subsequent studies. Based on currently available information, it is concluded that 49 species of Culicidae, 18 anophelines and 31 culicines, are present in various regions of the KSA.It is obvious from this study that researchers working in the KSA have generally been unaware of the limitations of available keys, have failed to critically evaluate published records and, at least in the case of


Cynomorium songaricum Rupr. is a well-known and widespread plant in China. It has very high medicinal values in many aspects. The study aimed at discriminating and predicting C. songaricum from major growing areas in China. An electronic tongue was used to analyze C. songaricum based on flavor. Discrimination was achieved by principal component analysis and linear discriminant analysis. Moreover, a prediction model was established, and C. songaricum was classified by geographical origins with 100% degree of accuracy. Therefore, the identification method presented will be helpful for further study of C. songaricum. Cynomorium songaricum Rupr. of the family Cynomoriaceae is a desert, holoparasitic perennial plant found in China, Mongolia, Iran, and Afghanistan [C. songaricum grows in Xinjiang, Inner Mongolia, Ningxia, Qinghai, and Gansu [C. songaricum, called Suo Yang in China, is a known food, nutrient, and a tonic herb for improving kidney and immunity function and treating constipation [C. songaricum to date [hanistan , 2. In Cnd Gansu   and linear discriminant analysis (LDA), were used for data analysis in this research. In addition, this study provided a simple approach for identifying the geographical origins of C. songaricum, and the acquired information can be used for evaluating the quality of C. songaricum growing in China.In this work, we first developed a method to discriminate and predict the geographical origin of C. songaricum samples were collected from different areas in China  (ongolia) . All of The electronic tongue system  consists of a reference electrode and seven liquid sensors  with a cross-selection function, a fully automated sample injector, and a personal computer with a software for sample injection, data acquisition, and chemometric analysis.Pieces of each sample (10 g) were placed in a beaker, soaked with 200 mL of pure water for 30 min, and then decocted for 30 min. The solution was filtered immediately. The residue was processed according to the abovementioned method twice. Afterward, all filtrates were combined. The obtained solution was placed into the special beaker of the electronic tongue and detected at room temperature.Each sensor collected data from each sample for 120 s and was cleaned for 10 s. Then, data were recorded by the data acquisition system. All assays were carried out in triplicate.C. songaricum originating from different places.In this paper, PCA and LDA were used to differentiate PCA is a multivariate statistical method that reduces the dimensionality of data while retaining most of the variation in the data . This apLDA is another commonly used technique for data discrimination and dimensionality reduction. LDA is strongly linked to regression analysis and analysis of variance (ANOVA), which also aims at expressing one dependent variable as a combination of other measurements or features . The metC. songaricum groups where possible. The accumulated explained variance was 89.6%, which was distributed in 79.5% (PC1) and 10.1% (PC2). C. songaricum samples can be classified in general. Moreover, C. songaricum samples from Xinjiang are discriminated clearly between samples from other provinces. Similar samples appear in the same location of the graph. Thus, C. songaricum samples from Gansu, Ningxia, Qinghai, and Inner Mongolia are similar. The chemical constituents of these samples may be similar as well.The first discrimination model was established using PCA to visualize the different C. songaricum samples by the LDA model. Compared with the PCA model, the LDA model shows a clearer discrimination among the eight types of C. songaricum. The explained variances by each discriminant function (DF) were 96.0% (DF1) and 2.9% (DF2). Each group of C. songaricum samples can be distinctly classified with others. As a result, the LDA model is a superior method to discriminate C. songaricum from different growing areas in China.C. songaricum samples. As shown in C. songaricum by geographical origins with 100% degree of accuracy.Given the good discrimination feature of the model, we used the LDA model to predict the geographical origin of unknown C. songaricum samples from different places of origin. PCA and LDA were used for discrimination. The LDA model shows a clearer discrimination than the PCA model. The eight groups of C. songaricum samples can be classified precisely using the LDA model. On this basis, we used the LDA model to predict the geographical origin of several unknown C. songaricum samples. A prediction model with 100% degree of accuracy was achieved. We present for the first time a method for the discrimination and geographical origin prediction of C. songaricum from different growing areas in China according to their flavor by an electronic tongue. The operations of data acquisition and processing are simpler and more convenient than the traditional chemical methods. The acquired information can be used for evaluating the quality of C. songaricum growing in China according to our previous work. Moreover, the identification and quality analysis method presented by us will be helpful for further study of C. songaricum. Further efforts should be focused on investigating the connection between the flavor and chemical constituents of C. songaricum samples and correlating electronic tongue signals with human perceptions of taste.We applied an electronic tongue to classify and predict


Liolaemus conspersus and L. hieroglyphicus, one syntype of Callopistes maculatus (here designated as the lectotype) and two syntypes of L. lineatus (one of which is herein designated as the lectotype). Reexamination of these specimens indicates that previous synonymies proposed for L. conspersus and two syntypes of L. hieroglyphicus are problematic; furthermore, more complex taxonomic work is needed to resolve this issue. Two rediscovered syntypes of L. lineatus differ in several scalation traits and are possibly not conspecific. The type specimens of several other species of lizards from Gravenhorst’s collection  were not found and are probably lost.Johann Ludwig Christian Gravenhorst’s herpetological collection at the Museum of Natural History, University of Wrocław included numerous important specimens of amphibians and reptiles. The majority, if not the entirety, of this collection has long been thought to be lost. However, we were able to rediscover some type specimens of lizards. The rediscovered specimens include the holotypes of  Liolaemus lemniscatusGerman naturalist Johann Ludwig Christian Gravenhorst (1777–1857) was the founder and first director of the Zoological Museum at the University of Wrocław . His main interests lay in entomology, particularly beetles and, later, ichneumonid wasps . HoweverLiolaemus, including all those named by Gravenhorst (except L. lemniscatus) were synonymised by L. oxycephalus  in an attempt to extract DNA. Unfortunately, these attempts were unsuccessful, so all descriptions and taxonomic assessments were made entirely based on morphology.A small amount of muscle tissue was taken from putative holotypes of http://zoobank.org/. The LSID for this publication is: urn:lsid:zoobank.org:pub:3CDA8E5A-30C8-4847-B4A4-C644595361FC. The online version of this work is archived and available from the following digital repositories: PeerJ, PubMed Central and CLOCKSS.The electronic version of this article in portable document format will represent a published work according to the Reptilia Laurenti, 1768Squamata Oppel, 1811Teiidae Gray, 1827Callopistes maculatusGravenhorst, 1838Original name.Callopistes maculatusType specimens. Two syntypes  and Phymaturus flagellifer . They attempted to synonymise Callopistes maculatus with Callopistes palluma and designated a neotype (MNHNCL 2909) for the latter name. However, these proposals were later rejected . Posterior to the mental scale, there are four pairs of large chinshields. They are separated from the infralabials by one (anteriorly) or two (posteriorly) rows of sublabials. The first pair of chinshields contact at the midline, but the second and farther pairs are separated by small, oval gular scales (it is impossible to tell the exact number). The whole throat is covered by small, oval or roughly pentagonal, juxtaposed gulars, all about the same size. The interangular sulcus is absent and the intertympanic sulcus is present. Between the throat and the posterior surface of the arms, the scales are also juxtaposed, but larger; some of them are oval, elongated, pentagonal or hexagonal. The whole venter (behind the posterior end of the arms) is covered by large, rectangular, juxtaposed scales. Similar scales, though slightly smaller and imbricate, also cover the entire tail, forming numerous annuli. The dorsal surface of the forelimbs is covered by roughly round, juxtaposed or slightly imbricate scales. The ventral forelimb scales are much smaller, round and juxtaposed. The dorsal hindlimb scales are relatively large, quadrangular, juxtaposed or slightly imbricate. The posterior surface of the thighs is covered by much smaller, round or quadrangular, juxtaposed scales. The ventral thigh scales are roughly rectangular, juxtaposed or minimally imbricate. The ventral shank scales are much larger, rounded and juxtaposed . The subdigital lamellae on both the fore- and hindlimbs are impossible to count precisely because of the distortion of the specimen  collected by F.S. Scholtz in ‘Cauquenes’ . This specimen still exists in Wrocław.Present name.Liolaemus conspersusRemarks. This species was described by Ptychodeira Fitzingerii (Liolaemus fitzingerii) by L. nigromaculatus  is 13. The interparietal scale is large , subtriangular, with slightly concave right and slightly convex left margins; it contacts six scales. The frontal scale is single, approximately dumbbell-shaped, with a straight anterior margin and a posterior margin forming an obtuse angle. Five scales separate the rostral from the frontal, and seven scales separate it from the interparietal. The rostral contacts the nasal at a point. Nostrils are directed laterally on the right side and dorsolaterally on the left, probably due to distortion of the specimen. The nasal is separated from the canthal by one scale, and contacts eight scales. There are eight scales between the external nares. There are five flat and long supralabials; the fourth one is located below the eye and has an oblique posterior border. There are five loreals, one of which contacts the subocular. There is one row of seven lorilabials between the supralabials and loreals, four of which contact the subocular. Five infralabials are present; the second one contacts four scales, including two other infralabials . There are five enlarged postmental scales (chinshields) on the right side; on the left, there are only four such scales, but the second one clearly originated by the fusion of two scales. The second postmentals are separated from each other by two gular scales. The anteriormost gulars are slightly elongated and juxtaposed, while the others are wider than they are long and imbricate. The auricular scale is not differentiated. Temporal scales have varying shapes: some are subtriangular, some quadrangular, some pentagonal and some hexagonal; all are juxtaposed and unkeeled. There are approximately 73 dorsal scales between the occiput and the anterior surface of the thighs. There are 115 ventral scales between the mental scale and the vent and 77 scales around the midbody. Most of the ventrals are quadrangular and juxtaposed or subimbricate, but some are pentagonal. The dorsal neck scales are rounded or lanceolate, strongly keeled, while the lateral neck scales are small and bead-like. The dorsal scales are larger than the dorsal neck scales but otherwise very similar . InterstThe dorsal colouration is light grey, with the tail being slightly more yellowish. There is no difference in background colour between the vertebral and paravertebral fields. Vertebral, dorso- and ventrolateral stripes are absent. Numerous dark spots or short stripes are faintly visible on almost the entire dorsum and limbs; they are much more visible on the dorsal and lateral aspects of the head. The antehumeral spot cannot be unambiguously identified. Several darker longitudinal stripes are faintly visible on the throat. Apart from that, the ventral colouration is uniformly grey-yellowish.Liolaemus conspersus does not have a patch of enlarged scales on the posterior surface of the thighs and its fourth supralabial scale is upturned, so its affiliation to the subgenus Eulaemus, and thus synonymy with L. fitzingerii, can be excluded  than any of the above-named lizards  than to L. flavipiceus but the converse is true for the number of midbody scales (L. buergeri has more than both L. conspersus and L. flavipiceus). It is also worth noting that L. flavipiceus is one of the few species in which males do not have precloacal pores . At present, we regard it as a species inquirenda, that is, a species of doubtful identity, needing further investigation  differ d traits , so theil scales . The numvipiceus . Howeveral pores . If the tigation .Liolaemus hieroglyphicusGravenhorst, 1838Original name.Liolaemus hieroglyphicusType specimen. A single specimen  collected by F.S. Scholtz before 1838 in ‘Cauquenes’ . This specimen still exists in Wrocław.Present name.Liolaemus hieroglyphicusRemarks.Ptychodeira signifera (Liolaemus signifer) by L. olivaceus ; (3) the individual is a similar size and colour to those reported originally by ignifera   from the rostral. The frontal scale is hexagonal but with a nearly straight anterior margin, and is separated from the rostral by five scales. The rostral and nasal are in broad contact. The nasal is separated from the canthal by one scale. There are two pairs of internasals, the first one is in medial contact, the second pair is separated by one scale. Posteriorly, they contact two large, hexagonal frontonasals, also separated medially by one scale. There are five supralabials on both the left and right sides, the fourth one is located below the eye and has an upturned posterior margin. One row of lorilabials separates the supralabials from the loreals. The subocular contacts four lorilabials on the right side but only two on the left side. The loreal region is concave, and contains four loreals on the left and four on the right. The temporal scales are polygonal, some with a rounded posterior margin, juxtaposed or slightly imbricate, not keeled . The auricular scale is present, and its ventral border does not reach the auditory meatus midline. There are three enlarged supraoculars, with the first one being the largest. Seven supraciliaries are present. Four infralabials are present on the right side and five on the left side; the second one contacts four scales, including two other infralabials. There are four pairs of enlarged postmental scales (chinshields); the first pair is in contact, and the second pair is separated by two scales. Most gulars are oval, slightly imbricate.The nuchal and dorsal scales are imbricate, keeled and lanceolate with mucrons  brown stripes extending from the occiput to the preserved part of the tail, resembling a dotted line running along the spine. Similar stripes are also present in the temporal region. Numerous small brown spots occur on almost the entire pileus , the supralabials and the infralabials. The paravertebral fields are covered by many brown spots, usually one to two scales long and two to four scales wide. Whitish dorso- and ventrolateral lines are present. Between them, the lateral fields are also covered by brown-black spots, which are less numerous but larger than the paravertebral spots. The whole ventral body part is uniformly whitish.Liolaemus hieroglyphicus does not have a patch of enlarged scales on the posterior surface of the thighs and its fourth supralabial scale is upturned, so its affiliation to the subgenus Eulaemus, and thus synonymy with L. signifer, can be excluded  by its larger size , more loreal scales , fewer subdigital lamellae on the fourth toe  and in the shape of the dorsal scales, which are more rounded and do not have a mucron in L. septentrionalis , but more subdigital lamellae on the fourth toe (24–28). The shape of the ventral scales is very variable in this species—they can be triangular, subtriangular or rounded, as in L. hieroglyphicus. The lateral neck scales are markedly keeled in L. schroederi . Howevey keeled . It furte latter , while ihroederi , while in Region .Liolaemus hieroglyphicus is not synonymous with L. signifer, L. chiliensis or L. lemniscatus. It is similar to L. cyanogaster and L. schroederi (particularly the latter), but at present cannot be unambiguously synonymised with any of these. We regard it as a species inquirenda, that is, a species of doubtful identity, needing further investigation  collected by F.S. Scholtz in ‘Valparaíso’, Chile.Present name.Liolaemus lemniscatusRemarks. The holotype of this species survived the war and was recently redescribed  of undetermined sex, collected by F.S. Scholtz in ‘Valparaíso’, Chile. Two specimens  still exist in Wrocław, and the two others are probably lost.Present name.Liolaemus nitidus . L. nitidus. Liolaemus lineatusLiolaemus lineatusLiolaemus nigroviridisAll specimens of livaceus   is a relatively large and robust lizard with a stout appearance  and 11. The dorsal scales are large, imbricate, strongly keeled and mucronate. There are 33 dorsal scales, 69 ventral scales and 46 scales around the midbody. The tail scales form numerous annuli, which are particularly visible from the ventral side. The ventral tail scales have smaller mucrons. The ventral scales (including the gulars) are also relatively large, lanceolate or rounded (particularly the gulars), imbricate and not keeled. There are 29 gular scales between auditory meatus. The posterior parts of the thighs are covered by much smaller, oval, juxtaposed scales. No precloacal pores are visible. The numbers of subdigital lamellae on the left forelimb are as follows: I-8, II-15, III-17, IV-21, V-12; on the right forelimb, they are: I-9, II-15, III-19, IV-20, V-11; on the left hindlimb, they are: I-13, II-17, III-21, IV-25, V-14; on the right hindlimb, they are: I-10, II-15, III-18, IV-25, V-14.The colour of the head and dorsum is blue-grey. Numerous darker spots are visible on the lateral side of the head; they cover the background colour on the pileus. The ventral body part is almost uniformly white-yellow, with the throat covered by several faintly visible darker longitudinal stripes.The type specimen of the third variety MNHUW 1323b) is much smaller than MNHUW 1323a  and 13. 23b is muThe dorsal and ventral scalation is very similar to the second variety—dorsal scales are imbricate, lanceolate , keeled and mucronate , while the ventrals are large, flat, rounded  or lanceolate, imbricate and unkeeled. There are 33 dorsal scales, 64 ventral scales and 43 scales around the midbody. A total of 30 gulars are present between the auditory meatus. No precloacal pores can be observed. The number of scales between the posterior border of the auditory meatus and the shoulder is 16 on the left side and 18 on the right side. The numbers of subdigital lamellae on the left forelimb are as follows: I-9, II-15, III-18, IV-16, V-12; on the right forelimb, they are: I-9, II-14, III-18, IV-16, V-10; on the left hindlimb, they are: I-10, II-16, III-20, IV-26, V-14; on the right hindlimb, they are: I-11, II-15, III-21, IV-25, V-14.The colouration is very similar to the second variety. The dark stripes on the throat are much more visible in this specimen than in MNHUW 1323a. On the other hand, the spots on the lateral part of the head are better preserved in the type specimen of the second variety; however, this may be an artefact of preservation.Liolaemus nitidus have long been difficult to establish  and the contact between the nasal and rostral scales. In all 16 specimens of L. nitidus sampled by Troncoso-Palacios, the nasal and rostral scales are in contact; this trait is also apparently fixed in L. chiliensis . The synonymy of the third variety with L. nitidus is more problematic. It is more similar to L. chiliensis, a species with which it was synonymised by L. nitidus by the presence of contact between the rostral and nasal scales , Chile. We did not find this specimen and consider it to be lost.Present name.Liolaemus nitidus . This specimen is probably lost.Present name.Liolaemus nitidus  is consistent with L. unicolor but no unambiguous identification can be made. Therefore, this name must be considered a nomen dubium.Reptilia Laurenti, 1768Squamata Oppel, 1811Leiocephalidae Frost & Etheridge, 1989Leiocephalus schreibersii  by natural history dealer Ludwig Parreys (1796–1879) from Vienna. We did not find this specimen and consider it to be lost.Present name.Leiocephalus schreibersii  by a natural history dealer, Bescke; there are several naturalists with this name, so it is unclear whether one of them collected lizards for Gravenhorst. We were not able to locate any of these specimens and consider them to be lost.Present name.Chalcides viridanus , preferably also with molecular data, especially mitochondrial DNA, because there are several mitochondrial clades within this species.r islets . Populatr islets . The popr islets . Howeverr islets . Recentlr islets  suggesteLiolaemus, with an often controversial taxonomy  tend to have significantly fewer precloacal pores than do members of Eulaemus  or a few (L. lineatus) specimens and no new individuals have been reported for 180 years. However, there are several Liolaemus species that are currently considered valid and are known only from the type locality or only from the type specimen  of the Liolaemus nomina coined by nomina oblita (To the best of our knowledge, none (except a oblita .Callopistes maculatus and liolaemids Liolaemus conspersus, L. hieroglyphicus and L. lineatus. Reexamination of the morphology of liolaemids revealed several taxonomically informative differences between these specimens and their presumed senior synonyms. While the synonymy of L. lineatus with L. nitidus is supported, the same cannot be done for L. conspersus and L. hieroglyphicus and their supposed senior synonyms, so we regard them as species inquirendae. Unfortunately, our attempts at molecular analyses were unsuccessful, so resolving the status of these taxa requires further, more complex studies. Nonetheless, the rediscovery of these important specimens underscores the importance of natural history collections, and their proper management and protection, a point that was recently further strengthened by the tragic fire at the National Museum of Brazil in Rio de Janeiro in September 2018.We rediscovered several important specimens from the Gravenhorst’s long thought to be lost herpetological collection at the University of Wrocław: type specimens of the teiid lizard