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The lacertid Latastia ornata was known to date only by its holotype collected in 1938 in Bafatá, central Guinea-Bissau. We report new specimens and localities from Guinea-Conakry, a new country record and major range extension of 700 km SE of the type-locality. We provide an updated diagnosis of the species, including the first genetic and osteological data, and confirm that Latastia ornata is closely related to, but distinct from, L. longicaudata based on external morphology, cranial osteology, DNA data and zoogeography.

Urban habitats receive an increasing number of species due to anthropogenic activities, mainly transportations. Here, we report a new addition to the herpetofauna of Athens (Greece): a small population of the Pelasgian wall lizard (Anatololacerta pelasgiana) was found in a suburb of the Athenian metropolitan area. The species normally occurs in southwestern Anatolia and southeastern Aegean islands and this is the first record in the Greek mainland. Allochthonous species that successfully colonize cities raise new challenges to urban ecology.

We present a comprehensive catalogue of the reptiles of Equatorial Guinea, consisting of 118 species belonging to 67 genera and 22 families. There are two species of Crocodylia, ten of Testudines and 106 of Squamata; this last taxon is represented by 62 species of snakes, two amphisbaenians and 42 lizards. Of these 118 species, seven are present only in Annobon, seven only in Bioko, 47 only in Río Muni, 53 occur both in Bioko and Río Muni (or Bioko, Río Muni and other islands), and four are sea turtles. Despite its high diversity, the level of endemism of Bioko is relatively low, with only four endemic species out of the 60 species reported for the island. In contrast, despite its low diversity, Annobon has the highest endemicity level, with five endemic species (and two introduced). No endemic species are known for the rest of the regions of Equatorial Guinea, which contain 100 species. We reveal several new country and species records, and point to some pending taxonomic questions to be addressed. Among the new species records, we highlight the presence of Cyclanorbis elegans, which represents the most meridional known population of the genus. Additional species are expected to be found in Equatorial Guinea as further field and taxonomic work is developed.

Automated image classification is a thriving field of machine learning, and various successful applications dealing with biological images have recently emerged. In this work, we address the ability of these methods to identify species that are difficult to tell apart by humans due to their morphological similarity. We focus on distinguishing species of wall lizards, namely those belonging to the Podarcis hispanicus species complex, which constitutes a well-known example of cryptic morphological variation. We address two classification experiments: (1) assignment of images of the morphologically relatively distinct P. bocagei and P. lusitanicus; and (2) distinction between the overall more cryptic nine taxa that compose this complex. We used four datasets (two image perspectives and individuals of the two sexes) and three deep-learning models to address each problem. Our results suggest a high ability of the models to identify the correct species, especially when combining predictions from different perspectives and models (accuracy of 95.9% and 97.1% for females and males, respectively, in the two-class case; and of 91.2% to 93.5% for females and males, respectively, in the nine-class case). Overall, these results establish deep-learning models as an important tool for field identification and monitoring of cryptic species complexes, alleviating the burden of expert or genetic identification.

Podarcis wall lizards mainly feed on coleopterans, orthopterans, arachnids, and other small invertebrates. However, Aegean wall lizards, Podarcis erhardii, are widely distributed across Aegean islands and are increasingly observed eating non-traditional food items ranging from plant material to conspecific eggs and body parts. Here, we report the first documented case of P. erhardii consuming a large centipede, Scolopendra cingulata. The predator-prey relationship between these species has appeared to go both ways and may intensify on islands.

The genus Heliobolus comprises four recognized species, all endemic to sub-Saharan Africa. Of these, only Heliobolus lugubris occurs in southern Africa, its distribution extending from Angola in the west to Mozambique in the east and reaching as far south as parts of northern South Africa. Like many of the reptile species that occur in southern Africa, Heliobolus lugubris is poorly studied, and preliminary investigation suggested that it may contain cryptic diversity. The present work focusses on the Angolan population of H. lugubris and uses an integrative taxonomic approach based on morphological, coloration and DNA sequence data. The results indicate that some of the current and historical specimens of H. lugubris from Angola do not correspond to the nominotypical form, and that differences between specimens suggest the presence of two additional species, described here as Heliobolus bivarisp. nov. from the southernmost xeric/desertic regions and plateau of Namibe Province, southwestern Angola and H. crawfordisp. nov. from the Serra da Neve inselberg north through the sub-desert coastal regions of northern Namibe, Benguela, and Kwanza Sul provinces. Nominotypical Heliobolus lugubris is confirmed to occur in Cuando Cubango Province, southeastern Angola.

Elmberg, Johan (2023): Amphibians and reptiles in North Sweden: distribution, habitat affinities, and abundance (Classes: Amphibia and Reptilia). Zootaxa 5301 (3): 301-335, DOI: 10.11646/zootaxa.5301.3.1, URL: http://dx.doi.org/10.11646/zootaxa.5301.3.1

Viviparous Lizard Zootoca vivipara (Jacquin 1787) Distribution (Figure 7). Included records from Artportalen (N=775): all reports have been included, as there are not any confusion species. Widespread and common in the Southern, Middle, and Northern Boreal. Widespread but scarce in the Subalpine zone. Locally occurring above treeline in the Low-Alpine zone in favorable microclimates. As expected, the highest known occurrences in the Scandic Mountain range are gradually lower towards the north: 1000–1050 m altitude in Härjedalen (Sånfjället and Flatruet), 740 m in Pite lappmark (west of Vuoggatjålme), and 690 m in Lule lappmark (Vastenjaure). There is just one record from a truly far offshore island (Stora Fjäderägg, Västerbotten; Figure 7; Elmberg 1995). Although common in seashore habitats on the mainland along the entire Baltic coast of North Sweden, there are surprisingly few records even from nearshore islands. An exception may be the archipelago in southern Norrbotten, where the species occurs on some outer islands (e.g., Stor-Räbben and Vargön, green offshore area in Figure 7; Stefan Andersson, personal communication). For North Sweden as a whole, this indicates a limited dispersal capacity over brackish water. There are no indications of changes in distribution over the last 50 years. Habitat and movements. Found in almost any habitat offering a combination of basking sites and protective low vegetation. Favored natural habitats are forest edges and clearings, stony slopes, rock outcrops, sandy areas, and shores of lakes, rivers and the sea (Figures 12, 14). It often occurs among Juniperus communis, Calluna vulgaris, Empetrum nigrum and other plants typical of dry sun-exposed conditions. Closed forest, tall grass, and wet habitats are avoided. Anthropogenic habitats are widely used, for example clearings under powerlines, clear-cuts, edges of fields and meadows, stone walls, cairns, and roadsides (Figures 13, 19). There have not been any dedicated studies of this species in North Sweden. As far as known, it spends the entire annual activity period in the habitats mentioned above. Daily and annual movements are not known but appear very limited. Subterranean hibernation sites are found in or close to the summer habitat, usually in south-facing situations. Abundance estimates and trends. There are not any published abundance data, but estimates based on extensive field work in the Umeå area (Västerbotten) 1975–1994 run in the neighborhood of> 500 adults /km 2 in representative landscapes near the coast (Elmberg, unpublished). There are no indications of changes in abundance over the last 50 years., Published as part of Elmberg, Johan, 2023, Amphibians and reptiles in North Sweden: distribution, habitat affinities, and abundance (Classes: Amphibia and Reptilia), pp. 301-335 in Zootaxa 5301 (3) on page 316, DOI: 10.11646/zootaxa.5301.3.1, http://zenodo.org/record/8030434, {"references":["Elmberg, J. (1995) Groddjurens och kraldjurens utbredning i Norrland. Natur i Norr, 15 (2), 57 - 82. [in Swedish]"]}

Redescription of external morphology Based on a re-examination of the holotype and on the examination of the recently collected specimens RBINS 20301–20302, shown on Figures 3–6. Raw measurements and scale counts of the three specimens are provided in Table 2. Snout-vent length to 76 mm; total length> 229 mm (holotype). Tail 2.5 times SVL (based on RBINS 20302, the only specimen with a complete, original tail; the tail tip of RBINS 20301 is missing and is not healed). Body moderately depressed. Head distinct from neck, narrow (HL/HW ratio 1.8–2.1), long (HL/SVL ratio 0.25–0.30, proportionally longest in the subadult), depressed (HH/HL ratio 0.41–0.49, least depressed in the subadult). Head covered with symmetric plates. Rostral well visible in dorsal view. Frontonasal rounded anteriorly, slightly wider than long. A line of small tubercles along the posterior borders of the internasals, frontonasal and prefrontals (also along the lateral borders of the prefrontals in the subadult), “en forme de perles” (i.e., pearl-shaped) as described by Monard (1940). Suture between internasals subequal to suture between prefrontals (respectively 0.8 and 0.9 mm in RBINS 16301). Pupil round. Lower eyelid scaly. Canthus rostralis rounded. Lores near-vertical. Nostril opening in contact with 1 st supralabial, the anterior nasal and the two posterior, superposed, nasals. Four supraorbital scales, the anterior- and posteriormost small, separated from the supraciliaries by a continuous row of granular scales. In the holotype the anterior- and posteriormost supraoculars are entire, while in RBINS 20301 the anterior supraoculars are divided into two (left) or three (right) fragments and the posterior ones into three (left) or two (right) fragments. In RBINS 20302 the anterior supraoculars are divided into two fragments on each side, and the posterior supraoculars are unfragmented. Tympanic opening large, rounded, surrounded by smooth scales (i.e., no auricular denticulation), including a crescentic scale bordering the antero-dorsal limit of the tympanum. Seven or eight supralabials; one of them much enlarged and bordering the orbit, separating the five anterior supralabials (sometimes four, as on the left side of the holotype) from the two posterior ones. Frontal hexagonal, surrounded anteriorly by the two prefrontals, laterally by three supraoculars on each side, and posteriorly by the two frontoparietals. Anterior border of frontal pointing forward. Frontal narrow posteriorly. Length of frontal slightly smaller than distance between frontal and snout tip (respectively 4.1 and 5.1 mm in RBINS 16301). Frontoparietals pentagonal. Parietals large, nearly as long as frontal, separated by the interparietal scale and a small occipital. Pineal eye visible through the interparietal scale. Scales on the upper surface of the head smooth, except the pearl-shaped tubercles. Temporal area covered by an elongate, thin temporal plate along the parietal, and by smaller to granular scales. Six (in a single case seven) infralabials. Mental followed by four pairs of sublinguals, the first three in contact on the midline, the fourth pair separated from each other. Sublinguals progressively increasing in size posteriorly. Gular collar present and distinct, ventrally including five distinctly enlarged scales. The right profile and ventral view of the head, which had never been illustrated for the holotype so far, are shown on Figure 3. Mediodorsal scale rows not widened. Dorsal scales granular, in 67–70 longitudinal rows at midbody, those on the lower flanks nearly smooth, those on upper flanks and dorsum with a median, single, longitudinal keel. About 40 dorsal scales between legs. Six longitudinal rows of parallelepipedal or trapezoidal, widened, smooth, ventral scales; the two medioventral rows narrower than the lateral ones. Between the gular collar and the line of porebearing scales, 27 or 28 transversal rows of ventrals. A distinctly enlarged preanal plate, bordered laterally by a row of a few small preanals on each side. Femoral pores in a continuous row of 17–20. The left and right series of pores of RBINS 20301 and RBINS 20302 are separated by respectively three and two poreless scales (two in the holotype according to the Figure 3 in the original description, but actually three according to our observations). Subdigital lamellae of fingers and toes with two keels each. First finger shortest. Second finger longer than 5 th. Third and 4 th fingers longest, of subequal length. RBINS 20301 and RBINS 20302 both show 16/16 subdigital lamellae under the 4 th finger. When the leg is stretched alongside the body it extends anteriorly beyond the gular collar. Length of the feet comparable to head length. Toes without lateral denticulate fringes. First toe shortest. Second and 5 th toe of subequal length, shorter than the 3 rd and much shorter than the 4th. Subdigital lamellae under 4 th toe 23 to 26 (23 in the holotype, 25 or 26 in the two other specimens). Supracaudals much larger than dorsal scales. All supracaudal scales rectangular, presenting a strong median keel, each keel bearing a small tubercle at its posterior extremity. Subcaudals at the base of the tail smooth and rounded posteriorly, quickly becoming rectangular with a medial strong keel, but without a terminal tubercle on the keel. The tail of the holotype is broken. Monard (1940) mentioned 87 subcaudals, but the tail tip shows a uniform color contrasting with the anterior part of the tail, indicating that it is regenerated. The tail of the subadult RBINS 20302 is original, and shows 143 subcaudals. Coloration in life. Based on original description, RBINS 20301–20302 and Figures 3–7. The dorsal surface of the head is uniformly dark brown. The ventral surface of the head, the body and of the base of the tail is uniformly white. The background color of the sides of the head above the mouth line is dark brown, progressively darkening posteriorly to become black as the background color of the flanks and the first half of the dorsum. There is an alignment of white spots on the upper and lower lips and another on the temporal area, in continuity with four similar, irregular alignments of white spots along the whole length of the flanks. Four continuous, parallel white dorsal stripes extend from just behind the parietals till the posterior part of the dorsum where they fade and disappear (Figures 4–5 and 7). In the subadult these white stripes are irregular and discontinuous (Figure 6). From half-length of the trunk, the dorsal background color turns to reddish-brown (similar to the color of laterite), continuing to the tail. The upper surfaces of the proximal parts of the arms are black with white spots, turning to reddish-brown with irregular lighter spots on the distal parts of the arms. The upper surfaces of the legs show a reddish-brown background color with irregular lighter spots and markings. The lower surface of the tail progressively reaches posteriorly the uniform reddish-brown color uniformly covering the dorsal and lateral surfaces of the tail until its tip. The lower surfaces of the arms and legs are grayish-white, the palms are reddish-brown. Cranial osteology. Based on subadult male RBINS 20302. Snout and palatomaxillary bones The premaxilla is a single, dentigerous bone (Figure 8A–D). The alveolar shelf carries probably seven teeth (counting the sockets) and juts out caudad as two triangular processes touching the maxilla. There is a narrow, tapering, dorsocaudally directed nasal process that is almost as long as the nasals themselves and wedges the tapering tip between the nasals. The nasals are paired, almost flat elements that form a straight suture between themselves and an interdigitating suture with the frontal (Figure 8A, D). The anterior tips are pointed and diverge from each other to make room for the intercalating premaxillary nasal process. The nasals are the widest at about the mid-length where they project out laterally into a shark fin-shaped, anteriorly embayed protuberance slightly overlapping the maxillary facial process. The maxilla has a high facial process with a triangular posterodorsal process reaching the frontal and another small posterolateral process below it touching the prefrontal (Figure 8A–B). There is a semilunar embayment between these two processes. The alveolar border bears 15 pleurodont teeth. Right above the teeth, a palatal shelf medially expands (Figure 8C). The premaxillary process is short and slightly upturned and bifurcated into an anterolateral and an anteromedial process, as is common in many lacertids (Villa & Delfino 2019). The palatine is an edentulous, ventrally concave bone that overlaps the palatal shelf of maxilla laterally with a maxillary process (Figure 8C). This process also establishes contact with the prefrontal dorsolaterally. The squarish vomerine process overlaps the palatine process of the vomer. Between these two processes, namely the maxillary and the palatine, there is an anterior embayment and a ventral concavity corresponding to choana. Posteriorly the palatine ends in three little triangular protuberances of which the medial two overlap the pterygoid, and the lateral one laterally articulates to that bone. The palatines do not contact each other medially. The pterygoid is an edentulous, triradiate bone (Figure 8C). The anteromedial process of the pterygoid, which articulates with the palatine, is longer than the anterolateral process of the same that articulates with the ectopterygoid. The quadrate process of the pterygoid is slender, lateromedially compressed and is directed posterolateral. The dorsolateral surface of the quadrate process bears a small facet for the epipterygoid. The medial surface of this process bears a longitudinal groove for the attachment of the pterygomandibularis muscle (Daza et al. 2011; Das & Pramanick 2019). The ectopterygoid articulates medially to the ectopterygoid articular facet on the pterygoid anterolateral process (Figure 8C). Anteriorly it overlaps the palatal shelf of the maxilla and just contacts the palatine. The vomer forms the casing of the vomeronasal organ together with the septomaxilla (Figure 8C). Vomers contact each other medially except from their posterior medial margins. Anteriorly, they contact the maxilla. Posteriorly vomers touch the palatine. Anterolaterally vomers are narrow to create an opening for the vomeronasal fenestra. Behind this, vomers expand laterally, only to taper laterally again to for choana. The septomaxillae form the dorsal encasing of the vomeronasal organ (Figure 8A, D). The septomaxillae are concave ventrally. Their medial edges are turned dorsad. Anteroventrally the septomaxilla ends with two small, pointed processes. Except a small protuberance, there is no clear posteromedial process as in some Palaearctic lacertids (Villa & Delfino 2019). Chondrocranial braincase bones The braincase bones show some degree of fusion, although sutures are detectable, at least partially, between the otic capsule elements and the ventral braincase elements, namely the parabasisphenoid and the basioccipital. Prootic is a prominent element housing cochlea, anterior and horizontal (partly) semicircular canals and their ampullae (Figure 8B–C). The anterior semicircular canal forms a prominent bulge on the anterolateral surface, just behind the crista alaris. In this species, the crista alaris is a narrow semilunar projection immediately rostrad and somewhat medial to the anterior semicircular canal bulge (Figure 8B). Immediately below this bulge, begins a rather weakly developed crista prootica that runs caudad from this point. Ventrad to the crista prootica, the anterior margin of the prootic is embayed by the incisura prootica (Figure 8B). Ventrad to the incisura prootica notch, prootic projects rostrad into an obtusely triangular anterior inferior process. The bulge along the horizontal semicircular canal continues posteriorly as a prootic process to reach the anterior surface of the paroccipital process. On the medial surface of the prootic, there are two auditory nerve foramina. The prootic articulates (in this specimen, fused) with the otoccipital along the former’s posterior lateral margin. The paired otoccipital forms the occipital condyle with the basioccipital (Figure 8A, E). The otoccipitals are composite (of opisthotic and exoccipital), hollow, bulbous bones that encase the internal ear. Dorsally the otoccipitals project out into a prominent, posterolaterally directed, axe head-shaped (in posterior view) paroccipital process that is approximately one-third the length of the quadrate. The supratemporal attaches to the anterior surface of the lateral end of this process. Ventrad to the supratemporal facet, there is a facet for the quadrate. Dorsad to the basal tubera, the otoccipital has a very prominent, deep embayment of the recessus scalae tympani which is bordered anterodorsally by the crista interfenestralis. Posterodorsal to the lateral opening of the recessus scalae tympani, there is a vagus foramen. The position of the hypoglossal foramina could not be detected clearly in the scan. One of the two single elements of the ventral braincase is the parabasisphenoid (Figure 8C). The bone has an elongated, narrow parasphenoid rostrum. Behind this parasphenoid element, the basisphenoid begins to expand. On both sides of the base of the parasphenoid rostrum, a truncated looking trabecula cranii is present. Between the trabeculae, on the dorsal surface is situated the sella turcica.Two very small internal carotid foramina open within the sella turcica. Caudad to the sella turcica, the crista sellaris is present transversely. Two short, anteroventrally directed basipterygoid processes project out from the ventrolateral base of the basisphenoid (Figure 8C). These processes expand at their end. Dorsally and medially, the anterior vidian foramen pierces the basipterygoid process. The basioccipital is the ventral element of the braincase participating in the formation of the occipital condyle (Figure 8A, C, E). The bone is hexagonal, dorsally concave and in this specimen, partly fused with the otoccipital, the prootic and the parabasisphenoid. Ventrad to the lateral opening of the recessus scalae tympani, the basioccipital has a small protuberance, the basal tubera. The basioccipital forms the floor of the recessus scalae tympani. The supraoccipital consists of a dorsomedian roof for the foramen magnum and two expanded and hollowed lateroventral wings partly encasing the inner ear (Figure 8A). In this species the processus ascendens is a very small, anteriorly truncated protuberance which does not reach the parietal (Figure 8E). This bone articulates with the prootic and the otoccipital. Dermal skull roofing bones The frontal bone is a single (fused in this specimen, condition in hatchlings is not known), elongated, skull roof element, being almost twice as long as the parietal in this species (Figure 8A, D). The frontal is wide at the rostral end and very wide at the caudal end and relatively narrower in the middle. A few digitiform processes from the frontals form interdigitating sutures with the nasals and the maxillae. The frontal articulates with the prefrontal along the anterior one-third of the former’s lateral margin. On the ventral surface of the frontal, along its outer margin, there is a crest, the crista cranii. The caudal end of the frontal expands markedly into two posterolateral processes with a squarish end and articulates with the parietal and the postfrontal. This particular specimen does not show any rugosity on the dorsal surface of the frontal. The parietal is a squarish, short, wide skull roof bone (Figure 8A–B). There are no anterolateral processes. However, there is a narrow, tapering posterolateral process on each side that articulates with the supratemporal and is just separated from the squamosal. These processes do not reach the paroccipital process of the otoccipital. There is a pineal foramen (Ledesma & Scarpetta 2018) piercing the parietal. There is a fossa parietalis (Oelrich 1956) at the midpoint of the embayed posterior margin of the parietal, though the processus ascendens from the supraoccipital is very weakly developed in this species and does not reach the fossa. Circumorbital bones The prefrontals are cavernous (in anteromedial view) bones that meet the frontals laterally, the maxillary facial process anteriorly and anterodorsally, the palatines ventrally and the lacrimals ventrolaterally (Figure 8A–B). The ventrolateral border of the orbitonasal flange of the parietal has a deep lacrimal notch. The orbitonasal flange projects ventrally into a posteroventral process medial to the lacrimal notch. Dorsally, the flange projects into a caudally directed process along the crista cranii. There is rather weak palpebral crest. The jugals are paired bones consisting of an anterior and a posterodorsal process, the former being more robust, especially close to the junction between the two, than the latter (Figure 8A–C). The anterior process tapers anteriorly and meets the lacrimal.This process articulates with the frontal anteroventrally and the ectopterygoid ventromedially. There is a faintly developed, caudally directed protuberance, the quadratojugal process, at the junction between the anterior and the posterodorsal processes. The slender posterodorsal process touches the postorbital. The lacrimal is a small sliver of a bone overlying the junction of the prefrontal, jugal and the maxilla (the prefrontal is just separated from the jugal), immediately caudad to the prefrontal lacrimal notch (Figure 8B). The postfrontal is a triradiate bone with a very slender anterolateral process articulating to the posterior lateral margin of the frontal, a weakly developed, triangular protuberance articulating to the anterodorsal margin of the postorbital and a robust, dagger-like posterior process wedged between the frontal and the parietal medially and the postorbital laterally (Figure 8A–B). The postorbital is situated below the postfrontal (Figure 8A–B). It consists of a short, triangular anteroventral process that contacts the posterodorsal process of the jugal and a longer posterior process touching the postfrontal and the squamosal, but the resolution of the scan did not permit determination of the exact end point of this process. In this specimen, the postfrontal and the postorbital do not extend more than one-third the length of the squamosal and thus, leave a prominent upper temporal fenestra open. There are prominent supraorbital ossifications above and scleral ring within the orbit (Figure 8A–B). Suspensorial and palatoquadrate derived bones The squamosals are a pair of J-shaped suspensorial bones (Figure 8A–B). Rostrally the squamosal is tapered and articulates medially to the postorbital. Posteriorly the bone curves ventrad and contacts the supratemporal dorsomedially and the quadrate ventrally. The ventrally curved caudal end of this bone is squarish. The supratemporals are small, curved bones with a tapered rostral and widened caudoventral end (Figure 8A–B). The caudoventral end bears an articulatory facet for the quadrate. Anterolaterally and anteromedially the supratemporal articulates with the squamosal and the posterolateral process of the parietal, respectively. The epipterygoid is a rather nondescript rod-like bone that articulates to the pterygoid right behind t, Published as part of Pauwels, Olivier S. G., Das, Sunandan, Camara, Lewei Boyo, Chirio, Laurent, Doumbia, Joseph, D'Acoz, Cédric D'Udekem, Dufour, Sylvain, Margraf, Nicolas & Sonet, Gontran, 2023, Rediscovery, range extension, phylogenetic relationships and updated diagnosis of the Ornate Long-tailed Lizard Latastia ornata Monard, 1940 (Squamata: Lacertidae), pp. 501-524 in Zootaxa 5296 (4) on pages 507-517, DOI: 10.11646/zootaxa.5296.4.1, http://zenodo.org/record/7984314, {"references":["Monard, A. (1940) Resultats de la mission scientifique du Dr. Monard en Guinee portugaise, 1937 - 1938. VIII. Reptiles. Arquivos do Museu Bocage, 11, 147 - 182, pl.","Villa, A. & Delfino, M. (2019) A comparative atlas of the skull osteology of European lizards (Reptilia: Squamata). Zoological Journal of the Linnean Society, 187 (3), 829 - 928. https: // doi. org / 10.1093 / zoolinnean / zlz 035","Daza, J. D., Diogo, R., Johnston, P. & Abdala, V. (2011) Jaw adductor muscles across lepidosaurs: a reappraisal. The Anatomical Record, 294 (10), 1765 - 1782. https: // doi. org / 10.1002 / ar. 21467","Das, S. & Pramanick, K. (2019) Comparative anatomy and homology of jaw adductor muscles of some South Asian colubroid snakes (Serpentes: Colubroidea). Vertebrate Zoology, 69 (1), 93 - 102. https: // doi. org / 10.26049 / VZ 69 - 1 - 2019 - 04","Ledesma, D. T. & Scarpetta, S. G. (2018) The skull of the gerrhonotine lizard Elgaria panamintina (Squamata: Anguidae). PloS ONE, 13 (6), e 0199584. https: // doi. org / 10.1371 / journal. pone. 0199584","Oelrich, T. M. (1956) The anatomy of the head of Ctenosaura pectinata (Iguanidae). University of Michigan Museum of Zoology Miscellaneous Publications, 94, 1 - 122.","Good, D. A. (1987) A phylogenetic analysis of cranial osteology in the gerrhonotine lizards. Journal of Herpetology, 21, 285 - 297. https: // doi. org / 10.2307 / 1563970"]}

Eremias scripta Strauch, 1867 — Erroneous. Sand Racerunner Singapore records. No published records. Remarks. A specimen of Eremias scripta (MCZ R-460) said to be from Singapore was collected by Putnam on 30 September 1859. The specimen could not have originated from Singapore as E. scripta is native to western Asia (Ananjeva et al. 2006). Unfortunately, we were unable to examine the specimen as it is no longer at MCZ, having been sent to Cope in 1891. The specimen must have either been introduced into Singapore or the locality written in the catalogue is incorrect. LKCNHM & NHMUK Museum specimens. No specimens. Additional Singapore museum specimens. Singapore (no locality): MCZ. Family Leiolepididae Fitzinger, 1843 (2 species) Genus Leiolepis Cuvier, 1829 (2 species), Published as part of Figueroa, Alex, Low, Martyn E. Y. & Lim, Kelvin K. P., 2023, Singapore's herpetofauna: updated and annotated checklist, history, conservation, and distribution, pp. 1-378 in Zootaxa 5287 (1) on pages 262-263, DOI: 10.11646/zootaxa.5287.1.1, http://zenodo.org/record/7960319, {"references":["Ananjeva, N. B., Orlov, N. L., Khalikov, R. G., Darevsky, I. S., Ryabov, S. A. & Barabanov, A. V. (2006) The Reptiles of Northern Eurasia: Taxonomic Diversity, Distribution, Conservation Status. Pensoft Publishers, Sofia, Bulgaria, 245 pp."]}

Figueroa, Alex, Low, Martyn E.Y., Lim, Kelvin K.P. (2023): Singapore's herpetofauna: updated and annotated checklist, history, conservation, and distribution. Zootaxa 5287 (1): 1-378, DOI: https://doi.org/10.11646/zootaxa.5287.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5287.1.1

Takydromus sexlineatus Daudin, 1802 — Erroneous. Six-lined Long-tailed Grass Lizard Singapore records. Takydromus sexlineatus —Chan-ard et al., 2015: 104.—Hawkeswood & B. Sommung, 2017a: 1.—Wiradana et al., 2021: S143. Remarks. Chan-ard et al. (2015) listed Singapore as part of the distribution for T. sexlineatus, but we are unsure where they acquired the record from since there are no publications reporting it from Singapore. Takydromus sexlineatus ranges from India east to southern China, south to northern Peninsular Malaysia, and Borneo (Grismer 2011b). LKCNHM & NHMUK Museum specimens. No specimens. Additional Singapore museum specimens. No specimens. Family Scincidae Oppel, 1811 (3 species) Genus Lygosoma Hardwicke & Gray, 1827 (1 species), Published as part of Figueroa, Alex, Low, Martyn E. Y. & Lim, Kelvin K. P., 2023, Singapore's herpetofauna: updated and annotated checklist, history, conservation, and distribution, pp. 1-378 in Zootaxa 5287 (1) on page 284, DOI: 10.11646/zootaxa.5287.1.1, http://zenodo.org/record/7960319, {"references":["Grismer, L. L. (2011 b) Lizards of Peninsular Malaysia, Singapore and Their Adjacent Archipelagos. Edition Chimaira, Frankfurt am Main, 728 pp.","Hardwicke, T. & Gray, J. E. (1827) A synopsis of the species of saurian reptiles, collected in India by Hardwicke. Zoological Journal, 3 (10), 213 - 229."]}

Atlantolacerta andreanskyi (Werner, 1929) is an endemic lizard from the High Atlas Mountains region of Morocco. A previous molecular assessment of this species using mitochondrial and nuclear DNA markers uncovered extensive genetic diversity with seven lineages indicative of a species complex. A morphological assessment of six of these lineages did not establish simple diagnostic features, and proposed these should be considered as a cryptic species, while highlighting the need for greater sampling across the range. In this study, we sampled 8 individuals from 5 previously unsampled localities and carried out genetic analyses to compare these populations to the known variation. Phylogenetic reconstruction based on mitochondrial DNA markers (12S rRNA and ND4) corroborates the previously described lineages and identified a new one. Interestingly, the two samples that account for this newly identified lineage have been collected from distinct localities – M'goun and Toumliline – that form a sister taxon to the population of Jbel Azourki.

DISCRIMINATION BETWEEN PODARCIS BOCAGEI AND P. LUSITANICUS The overall performance of the three methods for image classification of P. bocagei and P. lusitanicus in the four different datasets is shown in Table 2. Detailed results, including training, validation and test-set evaluation for all cross-validation sets, are shown in the Supporting Information, Tables S1–S 4. Accuracy is generally high, ranging from 87.3% in the case of InceptionV 3 in female dorsal images to 94.8% in male dorsal images when applying InceptionResNetV2. AUC ranges from 0.931 using InceptionV 3 in female dorsal images to 0.984 using Inception-ResNetV 2 in male dorsal and head lateral images. F1-scores show that, typically, P. lusitanicus is more frequently misclassified than P. bocagei, for both types of images and for both sexes. All three methods perform similarly in all datasets considering the three performance metrics. Identification of males is generally more accurate than that of females. Considering all five cross-validation replicates of the three models, the identification accuracy of males is significantly higher than that of females only when considering dorsal images (P = 0.048, Mann–Whitney–Wilcoxon test). The same result is obtained, but even more pronounced, using other metrics (P = 0.009 and P = 0.030 for AUC and F1-scores, respectively). With respect to head lateral images, the difference in identification accuracy between sexes also exists but it is significant only for differences in AUC (P = 0.046, Mann–Whitney– Wilcoxon test). There is no difference in performance using different image perspectives, neither in the case of males nor in that of females. As an extension to this basic approach, we tested whether model ensembles (calculated by averaging predictions of different models) would increase classification success. Model ensembles within each of the four datasets do not always improve classification success compared to the best single model (see results in Table 2). For instance, in the case of head lateral images, prediction performance is worse with the model ensemble than when using the best-performing model alone. In the case of dorsal images, the improvement is slight for males and more substantial for females. By contrast, combining the predictions from different views results in a much higher classification success in all cases, where accuracy reaches as high as 97.1% for males and 95.9% for females. These results are presented in Table 3 and the corresponding confusion matrices in Figure 2. Grad-CAM heatmaps were produced only for the model showing the highest accuracy in each case (Inception-ResNet V 2 in the case of male dorsal and head lateral images, ResNet 50 in the case of female dorsal images and Inception V 3 in the case of female head lateral images). Visualization of the heatmaps confirms that the models are indeed considering the lizard images for classification and not external features (like human fingers, writings, shadows and other non-lizard elements that appear in some images). Examples of heatmaps used to discriminate the two classes are shown in Figure 3. In dorsal images, the model often uses the middle area of the trunk to discriminate the two classes. Still, the head region is also used (and both regions combined). In female dorsal images, the head is not as frequently used as the trunk, but the portion of the trunk used for discrimination is generally more anterior than in males. In both male and female head lateral images, the area around the ear is the one most frequently used for classification, although this region could be more or less shifted towards the throat in both sexes. *AUC refers to the area under the ROC curve. Podarcis bocagei was the positive class. F1, harmonic mean of precision and recall; Pboc, Podarcis bocagei; Plus, Podarcis lusitanicus. AUC was calculated assuming P. bocagei as the positive case; F1, precision and recall were macro-averaged. DISCRIMINATION BETWEEN THE NINE GROUPS Overall, the performance of the different models for classification of the nine classes is worse than in the two-class case. Unlike the experiment involving only P. bocagei and P. lusitanicus, in all analyses considering nine classes there is some evidence of overfitting (see the Supporting Information, Tables S5–S 9 for detailed training, validation and testing evaluation scores), which could not be completely overcome by varying the hyperparameters. A summary of the performance of each model is presented in Table 4. In general, accuracy ranges from 76.3% for ResNet 50 in female head perspectives to 85.3% for InceptionResNetV 2 in male dorsal views. A striking result is the highly significant difference between male and female image identification accuracy, with consistently higher accuracies in male datasets, which holds for both types of images (P P = 0.0325 for both accuracy and F1-score, Wilcoxon signed rank test). Unlike the two-class case, in which the utility of ensemble models is mostly restricted to the combination of predictions from different perspectives, without important improvements in the within-dataset case, in the nine-class experiment ensemble models combining predictions from the three architectures for each image perspective greatly improve classification accuracy when compared to the best single model (see Table 4). Using estimates from different views by averaging across the six model predictions improves classification success even further. These results are shown in Table 5 and the respective confusion matrices shown in Figure 4. In this case, prediction accuracy reaches as high as 93.5% for males and 91.2% for females. The distribution of classification metrics according to the species is shown in Table 5. Taking a deeper look into these classification scores, it appears that several species are fairly well recognizable, with F1 scores above 0.90: this is the case for P. bocagei, P. carbonelli, P. lusitanicus, P. Ʋaucheri s.l., P. Ʋaucheri s.s. and P. Ʋirescens in males, and for the same species except P. lusitanicus in females. The most problematic species is, in both sexes, P. liolepis. Considering confusion matrices, it is noticeable that individuals of this species are often misclassified as P. Ʋirescens (more so in the case of females than males). Noteworthy is that the misclassification between the cryptic P. guadarramae and P. lusitanicus is minimal (7.9% of P. lusitanicus females and 4.6% of males are classified as P. guadarramae and 0 and 1.6% of P. guadarramae females and males are classified as P. lusitanicus; see Fig. 4). As for the two-class problem, Grad-CAM analyses show that, typically, the models use lizard – and not other – features for classification. However, even with the visualization tool available, it is not straightforward to understand what the model considers for discrimination. More precisely, the same regions seem to be used to classify distinct species, and it is not evident how differences in these regions are used. The most common patterns for each species are summarized in Tables 6 and 7 (for males and females, respectively)., Published as part of Pinho, Catarina, Kaliontzopoulou, Antigoni, Ferreira, Carlos A & Gama, João, 2023, Identification of morphologically cryptic species with computer vision models: wall lizards (Squamata: Lacertidae: Podarcis) as a case study, pp. 184-201 in Zoological Journal of the Linnean Society 198 (1) on pages 188-190, DOI: 10.1093/zoolinnean/zlac087, http://zenodo.org/record/7926780

In the summer of 2021 we identified three new distribution localities of Darevskia praticola north of the Mureș River, and one locality south of the river. The habitats populated by D. praticola (broad-leaved forest with wet areas) and the altitude (175–245 m) of the new records are typical for this species. Nevertheless, D. praticola had not been recorded in 12 other localities with similar conditions from an area previously considered suitable for this species. Thus, D. praticola may be slowly expanding from a bridgehead north of the Mureș River, occupying new favorable habitats. Probably, D. praticola recently crossed the Mureș River, possibly on a bridge, or with the timber trucks which exploit the woods from both sides of the river.

Understanding the genomic basis of adaptation to different abiotic environments is important in the context of climate change and resulting short-term environmental fluctuations. Using functional and comparative genomics approaches, we here investigated whether signatures of genomic adaptation to a set of environmental parameters are concentrated in specific subsets of genes and functions in lacertid lizards and other vertebrates. We first identify 200 genes with signatures of positive diversifying selection from transcriptomes of 24 species of lacertid lizards and demonstrate their involvement in physiological and morphological adaptations to climate. To understand how functionally similar these genes are to previously predicted candidate functions for climate adaptation and to compare them with other vertebrate species, we then performed a meta-analysis of 1, 100 genes under selection obtained from -omics studies in vertebrate species adapted to different abiotic factors. We found that the vertebrate gene set formed a tightly connected interactome, which was to 23% enriched in previously predicted functions of adaptation to climate, and to a large part (18%) involved in organismal stress response. We found a much higher degree of identical genes being repeatedly selected among different animal groups (43.6%), and of functional similarity and post-translational modifications than expected by chance, and no clear functional division between genes used for ectotherm and endotherm physiological strategies. In total, 171 out of 200 genes of Lacertidae were part of this network. These results highlight an important role of a comparatively small set of genes and their functions in environmental adaptation and narrow the set of candidate pathways and markers to be used in future research on adaptation and stress response related to climate change.

Darevskia mirabilis Arribas, Ilgaz, Kumlutaş, Durmuş, Avcı & Üzüm, 2013. stat. nov. (Fig. 12b). Type Locality: Ovit Pass, Kaçkar Mountains, Rize, Turkey. Distribution: It is known from the southern parts of Rize and Trabzon, especially around Kaçkar region. Comments: Distinctiveness of it already mentioned in other previous genetic studies (Rato et al. 2021; Candan et al. 2021), and whose isolated presence in the Kaçkar mountains, without contact with other forms, has made its classification oscillate between rudis and cf. valentini, and that has been genetically revealed in another distant locality (Sarıkamış, Kars, Turkey), a question that will be more deeply studied. Darevskia rudis rudis seems to be distinct from other forms that have been assigned to rudis s. lat., and that shall be considered now as nominally belonging to another taxon different from D. rudis: D. obscura stat. nov. (see below). Darevskia rudis would have as subspecies Darevskia rudis lantzicyreni (Darevsky & Eiselt, 1967) comb. nov. (Fig. 12d) and D. r. bolkardaghica (Fig. 12c). Darevskia bithynica, together with Darevskia b. tristis, perhaps paraphyletic and harboring more than one taxon within, or perhaps the results (paraphyly) are due to an ancient introgression that obscures its homogeneity. Darevskia valentini (s. str.) (Fig. 12f), monotypical, without any of its former subspecies (latzicyreni or spitzenbergerae) that belong to other species or are taxa on its own. Darevskia obscura (Lantz & Cyrén, 1936) stat. nov., including D. obscura bischoffi comb. nov. and D. obscura macromaculata comb. nov.. The latter seems to be identical in the different analyses done and could be synonymous with obscura s. str. (almost the Turkish populations). Must be mentioned that D. obscura has been postulated as a species on its own by other authors (Gabelaia et al. 2018 – by geometric morphometrics; Tarkhnishvili et al. 2020b - by head shape morphometrics-; Gabelaia 2019). It remains to clarify the status of the two forms of the Greater Caucasus (D. r. chechenica, and D. r. svanetica): independent from the others or probably closer to D. obscura, but not to the true D. rudis. This point has to be confirmed, however. DISCUSSION Phylogenetic reconstruction The complex structure of the studied group, D. valentini, D. rudis, and their relatives, has been recognized from the first studies to the present (Lantz & Cyren 1936; Darevsky & Eiselt 1967; Darevsky 1967; 1972; Darevsky & Lukina 1977; Eiselt et al. 1992; Arribas et al. 2013; Rato et al. 2021; Candan et al. 2021). This complexity has always been attractive to researchers who apply both kinds of markers, morphology and/or more recently genetics trying to solve it. Elaborated recent assessments using genetic markers point out that there are more lineages within the D. valentini / D. rudis complexes than the previously suspected (Candan et al. 2021; Rato et al. 2021). In this study, we aimed to increase the knowledge of the status of the currently recognized genetic lineages by creating the largest datasets, including a remote subspecies not studied so far – D. v. spitzenbergerae – for the first time, to use in both morphological and molecular analyses to clarify the problem. Our phylogenetic results show the presence of several monophyletic clades that reveal themselves as different species (Fig. 9). Of these distinct clades, some had been well-documented for the first time in a recently published study (Candan et al. 2021), and the authors have accepted that D. valentini s. lat. has more genetic lineages than previously suspected, two of which have been presented there as they should have to be described and named. However, two important shortcomings that we have tried to eliminate here prevented them from their taxonomic description: the lack of morphological study for diagnoses and the absence of samples of one of the up to now two unique subspecies of D. valentini (D. v. spitzenbergerae), whose study was unavoidable to make taxonomic decisions. As seen from the tree topology obtained here, genetically divergent lineages, clades A and B, were detected as monophyletic (Fig. 9). The occurrence of these two highly divergent monophyletic lineages is not only confirmed by the tree topology but also the species delimitation analyses revealed both clades as different species, which is one of the most important factors that paved the way for the here proposed taxonomic revision. Network analyses based on both genetic markers also supported this distinction. In Cyt-b, all clades were placed into their unique positions and they did not share any haplotypes (Fig. 10A). In MC1R, which is a nuclear marker and has slower substitution rates, an agreement relatively with a more complex structure was showed. Clade A is represented by a single haplotype (Hap16), while clade B appears to have two haplotypes (Hap16 and Hap26) (Fig. 10B). Although these results were suggested by Candan et al. (2021), a definite conclusion could not be made due to the absence of subspecies D. v. spitzenbergerae, a problem now solved. Considering sampling data used in our phylogenetic construction, clade A consists of both D. s. spitzenbergerae from Mergan Plateau (type locality of this relevant and geographically extreme subspecies) and D. s. wernermayeri ssp. nov. from Narlıca Valley as sister taxa (Fig. 9). Although the population located in Narlıca Valley (Van, Turkey) is morphologically included in D. v. lantzicyreni (Eiselt et al. 1992), it is genetically more closely related to D. s. spitzenbergerae than to the former. In addition to this, the populations, which were assimilated to D. v. lantzicyreni according to morphology (Eiselt et al. 1992), represent a completely different lineage (clade B) according to genetics. Such discordant patterns called cryptic speciation are often shown in the lizards (Ahmadzadeh et al. 2013; Kornilios et al. 2018; Karakasi et al. 2021; Arribas et al. 2022). Another major point is the status of D. r. mirabilis (clade C). This subspecies was first described by Arribas et al. (2013) from Ovit Pass, a very isolated geographic region in Kaçkar Mountains. Its phylogenetic position is obvious here and reveals that it should be a species as different as clades A and B (Fig. 9). The genetic difference of this taxon was demonstrated by two independent studies. Firstly, Rato et al. (2021) suggested that a clade, called Trabzon-Rize in their study, is genetically distinct and that it should be considered one of the four main lineages of D. rudis. Since they did not distinguish any subspecies, they could not determine that this clade belongs to D. r. mirabilis. The fact that the D. rudis specimens used in their study share the same branch with a specimen we know for certain to be D. r. mirabilis, undoubtedly proves that this clade is a new taxon and the corresponding samples of Rato et al. (2021) belong to it. Secondly, Candan et al. (2021) has also mentioned that it has isolated genetic structure and that its distribution area may be wider than expected because a datum retrieved from GenBank (Tarkhnishvili et al. 2013), which is located around Sarıkamış (Kars, Turkey), clustered with D. r. mirabilis in the same branch. Similar to Candan et al. (2021), one of the interesting results obtained within D. valentini / D. rudis complexes is that the specimens belonging to D. r. rudis, D. r. bolkardaghica and D. v. lantzicyreni, cluster together with overlapping. This unexpected pattern makes it difficult to engage the complexity of the group, which unables to apply the current nomenclature and difficulties understanding the main processes underlying genetic variation. Considering the genetic (Fig. 9) and morphological (see results section) evidence together, the most possible scenario seems to accept that D. v. lantzicyreni is really a subspecies of D. rudis, not from D. valentini. Thus, nominal form of D. valentini is only limited to northeast Anatolia (with areas of Georgia, Armenia and Azerbaijan), while the distribution of D. rudis sensu novo, extends from the northeastern Black Sea region to the inner Anatolia and from there to the south up to the Bolkar Mountains. Finally, the status of some former subspecies of D. rudis also inevitably needs revision. The claim that a member of this group, D. r. obscura, is different has been put forward in a previous study including phenotypic comparison (Gabelaia et al. 2018; Tarkhnishvili et al. 2020b). The phylogenetic results strongly support these morphological findings (clade G, Fig. 9). Above all, D. r. obscura has a phylogenetic position quite closely related to other two former D. rudis subspecies: D. r. bischoffi and D. r. macromaculata. Considering all these results, it seems that accepting the first described form, D. saxicola obscura Lantz & Cyren 1936, as a species: D. obscura will contribute positively to the clarification of this group. Morphology derived structure Considering the studied complex group it seems that there are three large groups, which obviously coincide with the current taxonomy based on morphology (we still use here the old nomenclature to refer them). The most different includes D. bithynica s. str. and D. b. tristis, which had longer heads both concerning its width, and also about their body length, but not in their pilei because other species (especially of the former rudis complex) had smaller (in size and length) but very wide heads. Similarly, the scales that cover the upper part of the crus are small and barely keeled. Also, they had comparatively longer hindlimbs (are the more climbing, based on this characteristic). Osteologically, they have very rarely any B-Type pre-autotomic vertebrae. The sternal fontanelle is frequently reduced or absent in D. b. bithynica. Postorbital and postfrontal are subequal or the postorbital is a bit smaller (different to D. valentini, clade B and D. v. spitzenbergerae). Squamosal and postorbital overlap commonly in half of the second’s length (as also in D. r. bolkardaghica), more usually than in other forms of the group. Darevskia b. bithynica and D. b. tristis are identical in ANOSIM. This species is also recovered by genetics. Genetics indicates the possibility that tristis is paraphyletic as presently understood. The former valentini complex has a broad overlap among the different forms in CDA. These valentini complex samples had comparatively longer limbs, comparatively smaller heads, a greater number of scales in the crus (which in this case corresponds also to smaller scale size, and are no or almost-none keeled), and less markedly, a greater number of ventral and dorsal scales. Anal index, a bit greater (scale comparatively wider) in D. valentini than in D. bithynica or the former rudis complex. One of their supposed taxa, D. v. lantzicyreni, perhaps due to its wide dispersal and the presence of isolated populations, appears somewhat heterogeneous. Darevskia v. lantzicyreni overlaps a few with D. r. bolkardaghica (in males, and even more in females, which would be in agreement with the genetic results and the taxonomic changes proposed above). In turn, D. v. lantzicyreni has the higher dorsalia among the former valentini complex and is the closer of this complex to D. rudis s. str., which would also agree with the genetic analysis (see above) and its relation as conspecific. Darevskia r. bolkardaghica is characterized by low lamellae (in males and females), and osteologically is characterized because not infrequently shows a weakly ossified rib associated to the third vertebra (an extremely rare character, probably atavistic, associated to small and isolated populations), and the sternal fontanelle adopt singular shapes in sand-clock, irregular cordiform or trilobate in its forepart.Also its squamosal and postorbital bones overlap commonly in half of the second’s length (as in D. bithynica). Genetically, it is related to D. v. lantzicyreni and D. r. rudis (all three are proposed here as subspecies of D. rudis) Darevskia v. spitzenbergerae, clade A (here treated as the nominal ssp. of D. spitzenbergerae), and clade B (here described as a new species) (in males) and in general, as all the former valentini complex (in females) have higher ventralia counts than in D. bithynica or the former rudis complex taxa. Osteologically, this singular form (spitzenbergerae) has the interclavicle lateral branches inclined forwards (with only this model in typical D. s. spitzenbergerae) and in “clade A” (coexisting with some branches backward). Postorbital and postfrontal are subequal or the postorbital is a bit greater (as in D. valentini s. str. or clade B). Nominal taxa spitzenbergerae +clade A, and clade B are recovered as two different species by genetics. Darevskia spitzenbergerae and clade A are primitive forms, among the closely related to D. rudis (sensu novo) and their subspecies (lantzicyreni and bolkardaghica, especially this latter). Darevskia valentini s. str. seems to be a different taxon (genetics) without its formerly assigned subspecies (is nominotypical). It has Temporalia2 a bit lower than in related taxa. In D. valentini s. str. not infrequently appear some B-Type autotomic vertebrae. Postorbital and postfrontal are subequal or the postorbital is a bit greater (as in clade B or D. spitzenbergerae). Darevskia rudis complex is characterized by smaller dorsalia in a great part of the rudis complex (except in D. r. rudis –yet indicated in Arribas et al. 2013, that also is recovered as a different species in genetic analyses), than in the former valentini complex and D. bithynica ssp. (except in D. v. valentini that has lower scores similar to the main former rudis complex). Hindlimb relative length is also comparatively smaller when compared with the former valentini complex and even more with D. bithynica. These differences shall be considered as characteristic of D. obscura and their newly assigned subspecies, which are very few differentiated. Darevskia r. obscura and D. r. macromaculata are near the same by morphology, as suggested in Arribas et al. (2013). They are so similar in all analyses (including non-significant differences in ANOSIM) that they appear to be the same (increased pigmentation in typical Georgian macromaculata, but perhaps not in Turkish specimens, a question to be studied in future). Temporalia1 is somewhat smaller in D. r. macromaculata and D. r. obscura (M, F), and SVL (size) is greater in D. r. bischoffi (M, F). Paradoxically, D. rudis s. str. is morphologically extreme and a differentiated form within “its” former complex, and is distinguished from the other former rudis complex taxa (now D. obscura sspp.) by its greater values of dorsalia. Also, it is basal to the group in UPGMA. Osteologically, in D. rudis s. str. postorbital and postfrontal are subequal or the first is smaller than the second (as in D. obscura and D. bithynica). The two extreme populations displaced towards the south of both classical complexes (D. rudis bolkardaghica and D. valentini spitzenbergerae) result in the ones that connect morphologically the former rudis and valentini complexes.This may be because they are the most primitive in both groups, or because of an ecoclimatic convergence in their scalation. Both live on calcareous substrates (siliceous, even volcanic in the other forms), so they have a lighter background color than other forms (darker). Concordance of genetic and morphological results a) The classic morphological groupings/species (rudis and valentini complexes) seem to be no longer valid, due to newly discovered “intermediate” taxa, recent speciation, and multiple past and present introgression. The situation is fairly more complex than previously expected. b) The above-mentioned morphological characteristics of D. bithynica are valid for this species. c) The above-mentioned morphological characteristics of D. rudis complex are valid for D. obscura (and its sspp. macromaculata and bischoffi). d) As stated above D. rudis s. str. is distinguished from the other former rudis complex taxa (hereinafter D. obscura sspp.) by its greater values of dorsalia. Also, it is basal to the group in UPGMA. e) Darevskia r. lantzicyreni comb. nov. and D. r. bolkardaghica are subspecies of D. rudis. f) Darevskia spitzenbergerae is a different taxon. The fourth axis of females analysis (at the limit of significance) discriminates specially clade A (D. s. wernermayeri nov. ssp.) and in a lesser degree D. v. spitzenbergerae (that genetically cluster together and are very similar in ANOVA), characterized by higher values of lamellae, preanalia and femoralia. Inside this spitzenbergerae genetic clade, preanalia is higher in clade A (D. s. wernermayeri nov. ssp.) and strongly characterizes it (M, F) concerning near all the taxa studied here (and if the Tukey-Kramer multiple comparison test is used instead of the much stricter of Scheffe, is significantly different to all the taxa, including the nominal D. s. spitzenbergerae) g) Darevskia mirabilis is another taxon, genetically singular, and only moderately differentiated in its morphology within the former rudis complex (morphologically seems more related to D. o. obscura or D. r. bolkardaghica –Anatolian diagonal effect? -), and longtime approached to D. valentini by its pattern, but well isolated genetically. In ANOVA, very low supraciliar granula (M, F) (especially distinctive of this taxon) and gularia (M), and higher circumanalia (M) counts are the most diagnostic characters. h) The number (tibialia), size and keeling of the crus scales was formerly used to distinguish between the forms assigned to “ rudis ” and “ valentini ” (sensu auctt.) and is distinctive with higher counts (and smaller scales size and keeling) in the former valentini complex (and D. b. bithynica), appearing, to the contrary, the lower counts (with big size and strong keeling) in part of the former rudis, Published as part of Arribas, Oscar, Candan, Kamil, Kornilios, Panagiotis, Ayaz, Dinçer, Kumlutaş, Yusuf, Gül, Serkan, Yilmaz, Can, Caynak, Elif Yildirim & Ilgaz, Çetin, 2022, Revising the taxonomy of Darevskia valentini (Boettger, 1892) and Darevskia rudis (Bedriaga, 1886) (Squamata, Lacertidae): a Morpho-Phylogenetic integrated study in a complex Anatolian scenario, pp. 1-68 in Zootaxa 5224 (1) on pages 25-29, DOI: 10.11646/zootaxa.5224.1.1, http://zenodo.org/record/7517907, {"references":["Arribas, O., Ilgaz, C., Kumlutas, Y., Durmus, S. H., Avci, A. & Uzum, N. (2013) External morphology and osteology of Darevskia rudis (Bedriaga, 1886), with a taxonomic revision of the Pontic and Small-Caucasus populations (Squamata: Lacertidae). Zootaxa, 3626 (4), 401 - 428. https: // doi. org / 10.11646 / zootaxa. 3626.4.1","Rato, C., Stratakis, M., Sousa-Guedes, D., Sillero, N., Corti, C., Freitas, S., Harris, D. J. & Carretero, M. A. (2021) The more you search, the more you find: Cryptic diversity and admixture within theAnatolian rock lizards (Squamata, Darevskia). Zoologica Scripta, 50 (2), 193 - 209. https: // doi. org / 10.1111 / zsc. 12462","Candan, K., Kornilios, P., Ayaz, D., Kumlutas, Y., Gul, S., Yildirim-Caynak, E. & Ilgaz, C. (2021) Cryptic genetic structure within Valentin's Lizard, Darevskia valentini (Boettger, 1892) (Squamata, Lacertidae), with implications for systematics and origins of parthenogenesis. Systematics and Biodiversity, 19 (7), 665 - 681. https: // doi. org / 10.1080 / 14772000.2021.1909171","Darevsky, I. S. & Eiselt, J. (1967) Ein neuer Name fur Lacerta saxicola mehelyi Lantz & Cyren 1936. Annalen des Naturhistorischen Museums in Wien, 70, 107.","Lantz, L. A. & Cyren, O. (1936) Description of Darevskia bithynica tristis. In: Contribution a la connaissance de Lacerta saxicola Eversmann. Bulletin de la Societe Zoologique de France, Paris, 61, pp. 159 - 181.","Gabelaia, M., Tarkhnishvili, D. & Adriaens, D. (2018) Use of three-dimensional geometric morphometrics for the identification of closely related species of Caucasian rock lizards (Lacertidae: Darevskia). Biological Journal of the Linnean Society, 125, 709 - 717. https: // doi. org / 10.1093 / biolinnean / bly 143","Tarkhnishvili, D., Gabelaia, M. & Adriaens, D. (2020 b) Phenotypic divergence, convergence and evolution of Caucasian rock lizards (Darevskia). Biological Journal of the Linnean Society, 130, 142 - 155. https: // doi. org / 10.1093 / biolinnean / blaa 021","Gabelaia, M. (2019) Phylogeny and morphological variation in the rock lizards of the genus Darevskia. Thesis, Ilia State University and Ghent University, Tbilisi, 121 pp.","Darevsky, I. S. & Lukina, G. P. (1977) Rock lizards of the Lacerta saxicola Eversmann group (Sauria, Lacertidae) collected in Turkey by Richard and Erica Clark. Proceedings of the Zoological Institute of the Academy of Sciences, U. S. S. R., 1977, 60 - 63.","Eiselt, J., Darevsky, I. S. & Schmidtler, J. F. (1992) Untersuchungen an Felseneidechsen (Lacerta saxicola komplex) in der ostlichen Turkei, I. Lacerta valentini Boettger. Annalen des Naturhistorischen Museums in Wien, 93 (B), 1 - 18.","Ahmadzadeh, F., Flecks, M., Carretero, M. A., Mozaffari, O., Bohme, W., Harris, D. J., Freitas, S. & Rodder, D. (2013) Cryptic speciation patterns in Iranian Rock Lizards uncovered by integrative taxonomy. Plos One, 8 (12), 1 - 17. https: // doi. org / 10.1371 / journal. pone. 0080563","Kornilios, P., Kumlutas, Y., Lymberakis, P. & Ilgaz, C. (2018) Cryptic diversity and molecular systematics of the Aegean Ophiomorus skinks (Reptilia: Squamata), with the description of a new species. Journal of Zoological Systematics and Evolutionary Research, 56 (3), 364 - 381. https: // doi. org / 10.1111 / jzs. 12205","Karakasi, D., Ilgaz, C., Kumlutas, Y., Candan, K., Guclu, O., Kankilic, T., Beser, N., Sindaco, R., Lymberakis, P. & Poulakakis, N. (2021) More evidence of cryptic diversity in Anatololacerta species complex Arnold, Arribas and Carranza, 2007 (Squamata: Lacertidae) and re-evaluation of its current taxonomy. Amphibia-Reptilia, 42 (2), 201 - 216. https: // doi. org / 10.1163 / 15685381 - bja 10045","Arribas, O., Candan, K., Kurnaz, M., Kumlutas, Y., Yildirim-Caynak, E. & Ilgaz, C. (2022) A new cryptic species of the Darevskia parvula group from NE Anatolia (Squamata, Lacertidae). Organisms Diversity & Evolution, 22, 475 - 490. https: // doi. org / 10.1007 / s 13127 - 022 - 00540 - 4","Tarkhnishvili, D., Murtskhvaladze, M. & Gavashelishvili, A. (2013) Speciation in Caucasian lizards: Climatic dissimilarity of the habitats is more important than isolation time. Biological Journal of the Linnean Society, 109 (4), 876 - 892. https: // doi. org / 10.1111 / bij. 12092","Murphy, R. W., Fu, J., MacCulloch, R. Darevsky, I. S. & Kupriyanova, L. (2000) A fine line between sex and unisexuality: the phylogenetic constraints on parthenogenesis in lacertid lizards. Zoological Journal of the Linnean Society, 130, 527 - 549. https: // doi. org / 10.1111 / j. 1096 - 3642.2000. tb 02200. x"]}

Darevskia bithynica bithynica 16(M), 4(F) 1. ZDEU 15 /2009. (N=20), Kirazlı Plateau, Uludağ, Bursa, Turkey, 23.06.2009, Leg. Y. KUMLUTAŞ, Ç. ILGAZ [Map ID: 1]., Published as part of Arribas, Oscar, Candan, Kamil, Kornilios, Panagiotis, Ayaz, Dinçer, Kumlutaş, Yusuf, Gül, Serkan, Yilmaz, Can, Caynak, Elif Yildirim & Ilgaz, Çetin, 2022, Revising the taxonomy of Darevskia valentini (Boettger, 1892) and Darevskia rudis (Bedriaga, 1886) (Squamata, Lacertidae): a Morpho-Phylogenetic integrated study in a complex Anatolian scenario, pp. 1-68 in Zootaxa 5224 (1) on page 56, DOI: 10.11646/zootaxa.5224.1.1, http://zenodo.org/record/7517907

Darevskia spitzenbergerae (Eiselt, Darevsky & Schmidtler, 1992)stat. nov. (Fig. 12g). Type Locality: Mergan Plateau, Cilo Mountain, Hakkari, Turkey. Distribution: It is known from only two locations: Mergan Plateau, Hakkari, Turkey and Narlıca Valley, Van, Turkey (this study). Comments: It includes one of the subspecies of D. valentini previously described and a population from Narlıca Valley (called the New Clade A in Candan et al. 2021). Since they are morphologically distinct and D. s. spitzenbergerae s tat. et comb. nov. is so singular in pattern, we think it may be subspecifically different. Clade A is described as subspecies of Darevskia spitzenbergerae s tat. nov.

Darevskia bithynica tristis 60(M), 54(F) 1. ZDEU 12 /2009. (N=15), Güzeldere Village, Düzce, Turkey, 24.06.2009, Leg. Y. KUMLUTAŞ, Ç. ILGAZ [Map ID: 2]. 2. ZDEU 6 /2009. (N=23), Samandere Waterfall, Düzce, Turkey, 24.06.2009, Leg. Y. KUMLUTAŞ, Ç. ILGAZ [Map ID: 3]. 3. ZDEU 10 /2009. (N=6), Between Yığılca and Bolu 30.km., Bolu, Turkey, 27.06.2009, Leg. Y. KUMLUTAŞ, Ç. ILGAZ [Map ID: 4]. 4. ZDEU 14 /2009. (N=11), Between Yığılca and Alaplı 12.km., Bolu, Turkey, 27.06.2009, Leg. Y. KUMLUTAŞ, Ç. ILGAZ [Map ID: 5]. 5. ZDEU 16 /2009. (N=19), Between Zonguldak and Gökçebey 15.km., Zonguldak, Turkey, 28.06.2009, Leg. Y. KUMLUTAŞ, Ç. ILGAZ [Map ID: 6]. 6. ZDEU 13 /2009. (N=8), Yenice, Karabük, Turkey, 28.06.2009, Leg. Y. KUMLUTAŞ, Ç. ILGAZ [Map ID: 8]. 7. ZDEU 11 /2009. (N=8), Between Safranbolu and Bartın 14.km., Bartın, Turkey, 29.06.2009, Leg. Y. KUMLUTAŞ, Ç. ILGAZ [Map ID: 7]. 8. ZDEU 17 /2009. (N=5), Ulus, Bartın, Turkey, 29.06.2009, Leg. Y. KUMLUTAŞ, Ç. ILGAZ [Map ID: 9]. 9. ZDEU 7 /2009. (N=7), Amasra, Bartın, Turkey, 29.06.2009, Leg. Y. KUMLUTAŞ, Ç. ILGAZ [Map ID: 10]. 10. ZDEU 9 /2009. (N=12), Kapısuyu, Kurucaşile, Bartın, Turkey, 30.06.2009, Leg. Y. KUMLUTAŞ, Ç. ILGAZ [Map ID: 11]., Published as part of Arribas, Oscar, Candan, Kamil, Kornilios, Panagiotis, Ayaz, Dinçer, Kumlutaş, Yusuf, Gül, Serkan, Yilmaz, Can, Caynak, Elif Yildirim & Ilgaz, Çetin, 2022, Revising the taxonomy of Darevskia valentini (Boettger, 1892) and Darevskia rudis (Bedriaga, 1886) (Squamata, Lacertidae): a Morpho-Phylogenetic integrated study in a complex Anatolian scenario, pp. 1-68 in Zootaxa 5224 (1) on page 56, DOI: 10.11646/zootaxa.5224.1.1, http://zenodo.org/record/7517907

Darevskia spitzenbergerae spitzenbergerae stat. et comb. nov. 10(M), 6(F) 1. ZDEU 1 /2020. (N=16), Mergan Plateau, Hakkari, Turkey, 15.08.2020, Leg. C. YILMAZ [Map ID: 78]., Published as part of Arribas, Oscar, Candan, Kamil, Kornilios, Panagiotis, Ayaz, Dinçer, Kumlutaş, Yusuf, Gül, Serkan, Yilmaz, Can, Caynak, Elif Yildirim & Ilgaz, Çetin, 2022, Revising the taxonomy of Darevskia valentini (Boettger, 1892) and Darevskia rudis (Bedriaga, 1886) (Squamata, Lacertidae): a Morpho-Phylogenetic integrated study in a complex Anatolian scenario, pp. 1-68 in Zootaxa 5224 (1) on page 53, DOI: 10.11646/zootaxa.5224.1.1, http://zenodo.org/record/7517907

Darevskia obscura obscura stat. et comb. nov. 38(M), 35(F) 1. ZDEU 43 /2016. (N=7), Kutul Plateau, Ardahan, Turkey, 18.07.2016, Leg. Ç. ILGAZ, K. CANDAN [Map ID: 59]. 2. ZDEU 17 / 2010. (N=22), Kutul Plateau, Ardahan, Turkey 14.07.2010, Leg. Y. KUMLUTAŞ, Ç. ILGAZ, A. AVCI, N. ÜZÜM, B. ÜZÜM [Map ID: 59]. 3. ZDEU 156 /2001. (N=44), Between Geçitli Village and Bilbilen Plateau, Ardanuç, Artvin, Turkey, 06.07.2001, Leg. Y. KUMLUTAŞ, K. OLGUN, Ç. ILGAZ, A. AVCI, F. İRET [Map ID: 60].

Darevskia valentini (Boettger, 1892) - Darevskia josefschmidtleri sp. nov. - Darevskia spitzenbergerae (Eiselt, Darevsky & Schmidtler, 1992) stat. nov. - D. spitzenbergerae spitzenbergerae (Eiselt, Darevsky & Schmidtler, 1992) comb. nov. - D. spitzenbergerae wernermayeri ssp. nov. - Darevskia mirabilis Arribas, Ilgaz, Kumlutaş, Durmuş, Avcı & Üzüm, 2013 stat. nov. - Darevskia rudis (Bedriaga, 1886) - D. rudis rudis (Bedriaga, 1886) - D. rudis lantzicyreni (Darevsky & Eiselt, 1967) comb. nov. - D. rudis bolkardaghica Arribas, Ilgaz, Kumlutaş, Durmuş, Avcı & Üzüm, 2013 - Darevskia obscura (Lantz & Cyrén, 1936) stat. nov. - D. obscura obscura (Lantz & Cyrén, 1936) comb. nov. - D. obscura bischoffi (Böhme & Budak, 1977) comb. nov. - D. obscura macromaculata (Darevsky, 1977) comb. nov. - Darevskia bithynica (Méhely, 1909) - D. bithynica bithynica (Méhely, 1909) - D. bithynica tristis (Lantz & Cyrén, 1936) This classification can be completed in future studies with the analysis of more informative genetic markers than the single-copy nuclear markers used here, such as microsatellite DNA or genomic SNPs. ACKNOWLEDGEMENTS The data used in the morphology section were mostly obtained from Kamil Candan’s PhD thesis, supervised by Dr. Dinçer Ayaz, which was supported by Dokuz Eylül University with project number 2017.KB.FEN.039. We thank to Dr. Mehmet Kürşat Şahin and Nurettin Beşer for their helps in the field. Finally, we wish to thank both reviewers and the editor for their contributions to improve our manuscript. CONFLICT OF INTEREST We declare that we have no conflict of interest. ETHICS To realize this study, the ethical committee document numbered 11/04/2016 was received from the Dokuz Eylül University Faculty of Medicine Animal Experiments Local Ethics Committee at the meeting dated 23.02.2016 and decided. In addition, the necessary application for the realization of the study was approved by the General Directorate of Nature Conservation and National Parks within the Ministry of Agriculture and Forestry, Turkey on 05.04.2016., Published as part of Arribas, Oscar, Candan, Kamil, Kornilios, Panagiotis, Ayaz, Dinçer, Kumlutaş, Yusuf, Gül, Serkan, Yilmaz, Can, Caynak, Elif Yildirim & Ilgaz, Çetin, 2022, Revising the taxonomy of Darevskia valentini (Boettger, 1892) and Darevskia rudis (Bedriaga, 1886) (Squamata, Lacertidae): a Morpho-Phylogenetic integrated study in a complex Anatolian scenario, pp. 1-68 in Zootaxa 5224 (1) on page 29, DOI: 10.11646/zootaxa.5224.1.1, http://zenodo.org/record/7517907, {"references":["Eiselt, J., Darevsky, I. S. & Schmidtler, J. F. (1992) Untersuchungen an Felseneidechsen (Lacerta saxicola komplex) in der ostlichen Turkei, I. Lacerta valentini Boettger. Annalen des Naturhistorischen Museums in Wien, 93 (B), 1 - 18.","Arribas, O., Ilgaz, C., Kumlutas, Y., Durmus, S. H., Avci, A. & Uzum, N. (2013) External morphology and osteology of Darevskia rudis (Bedriaga, 1886), with a taxonomic revision of the Pontic and Small-Caucasus populations (Squamata: Lacertidae). Zootaxa, 3626 (4), 401 - 428. https: // doi. org / 10.11646 / zootaxa. 3626.4.1","Darevsky, I. S. & Eiselt, J. (1967) Ein neuer Name fur Lacerta saxicola mehelyi Lantz & Cyren 1936. Annalen des Naturhistorischen Museums in Wien, 70, 107.","Lantz, L. A. & Cyren, O. (1936) Description of Darevskia bithynica tristis. In: Contribution a la connaissance de Lacerta saxicola Eversmann. Bulletin de la Societe Zoologique de France, Paris, 61, pp. 159 - 181.","Bohme, W. & Budak, A. (1977) Uber die rudis-Gruppe des Lacerta saxicola - Komplexes in der Turkei, II (Reptilia: Sauria: Lacertidae). Salamandra, 13 (3 / 4), 141 - 149.","Darevsky, I. S. & Lukina, G. P. (1977) Rock lizards of the Lacerta saxicola Eversmann group (Sauria, Lacertidae) collected in Turkey by Richard and Erica Clark. Proceedings of the Zoological Institute of the Academy of Sciences, U. S. S. R., 1977, 60 - 63."]}

Darevskia rudis bolkardaghica 9(M), 9(F) 1. ZDEU 4 /2017. (N=7), Karagöl, Ulukışla, Niğde, Turkey, 13.05.2017, Leg. Ç. ILGAZ, K. CANDAN, E. YILDIRIM CAYNAK [Map ID: 12]. 2. ZDEU 36 /2009. (N=11), Karagöl, Ulukışla, Niğde, Turkey, 19.07.2009, Leg. Y. KUMLUTAŞ [Map ID: 12]., Published as part of Arribas, Oscar, Candan, Kamil, Kornilios, Panagiotis, Ayaz, Dinçer, Kumlutaş, Yusuf, Gül, Serkan, Yilmaz, Can, Caynak, Elif Yildirim & Ilgaz, Çetin, 2022, Revising the taxonomy of Darevskia valentini (Boettger, 1892) and Darevskia rudis (Bedriaga, 1886) (Squamata, Lacertidae): a Morpho-Phylogenetic integrated study in a complex Anatolian scenario, pp. 1-68 in Zootaxa 5224 (1) on page 55, DOI: 10.11646/zootaxa.5224.1.1, http://zenodo.org/record/7517907

Darevskia mirabilis stat. nov. 17(M), 16(F) 1. ZDEU 142 /2014. (N=11), Ovit Pass, Rize, Turkey, 06.08.2014, Leg. Y. KUMLUTAŞ, Ç. ILGAZ [Map ID: 42]. 2. ZDEU 145 /2002. (N=22), Ovit Pass, Rize, Turkey, 06.09.2002, Leg. İ. BARAN, Y. KUMLUTAŞ, Ç. ILGAZ, A. ÖZDEMİR [Map ID: 42]., Published as part of Arribas, Oscar, Candan, Kamil, Kornilios, Panagiotis, Ayaz, Dinçer, Kumlutaş, Yusuf, Gül, Serkan, Yilmaz, Can, Caynak, Elif Yildirim & Ilgaz, Çetin, 2022, Revising the taxonomy of Darevskia valentini (Boettger, 1892) and Darevskia rudis (Bedriaga, 1886) (Squamata, Lacertidae): a Morpho-Phylogenetic integrated study in a complex Anatolian scenario, pp. 1-68 in Zootaxa 5224 (1) on page 53, DOI: 10.11646/zootaxa.5224.1.1, http://zenodo.org/record/7517907

Darevskia mirabilis Arribas, Ilgaz, Kumlutaş, Durmuş, Avcı & Üzüm, 2013. stat. nov. (Fig. 12b). Type Locality: Ovit Pass, Kaçkar Mountains, Rize, Turkey. Distribution: It is known from the southern parts of Rize and Trabzon, especially around Kaçkar region. Comments: Distinctiveness of it already mentioned in other previous genetic studies (Rato et al. 2021; Candan et al. 2021), and whose isolated presence in the Kaçkar mountains, without contact with other forms, has made its classification oscillate between rudis and cf. valentini, and that has been genetically revealed in another distant locality (Sarıkamış, Kars, Turkey), a question that will be more deeply studied. Darevskia rudis rudis seems to be distinct from other forms that have been assigned to rudis s. lat., and that shall be considered now as nominally belonging to another taxon different from D. rudis: D. obscura stat. nov. (see below). Darevskia rudis would have as subspecies Darevskia rudis lantzicyreni (Darevsky & Eiselt, 1967) comb. nov. (Fig. 12d) and D. r. bolkardaghica (Fig. 12c). Darevskia bithynica, together with Darevskia b. tristis, perhaps paraphyletic and harboring more than one taxon within, or perhaps the results (paraphyly) are due to an ancient introgression that obscures its homogeneity. Darevskia valentini (s. str.) (Fig. 12f), monotypical, without any of its former subspecies (latzicyreni or spitzenbergerae) that belong to other species or are taxa on its own. Darevskia obscura (Lantz & Cyrén, 1936) stat. nov., including D. obscura bischoffi comb. nov. and D. obscura macromaculata comb. nov.. The latter seems to be identical in the different analyses done and could be synonymous with obscura s. str. (almost the Turkish populations). Must be mentioned that D. obscura has been postulated as a species on its own by other authors (Gabelaia et al. 2018 – by geometric morphometrics; Tarkhnishvili et al. 2020b - by head shape morphometrics-; Gabelaia 2019). It remains to clarify the status of the two forms of the Greater Caucasus (D. r. chechenica, and D. r. svanetica): independent from the others or probably closer to D. obscura, but not to the true D. rudis. This point has to be confirmed, however. DISCUSSION Phylogenetic reconstruction The complex structure of the studied group, D. valentini, D. rudis, and their relatives, has been recognized from the first studies to the present (Lantz & Cyren 1936; Darevsky & Eiselt 1967; Darevsky 1967; 1972; Darevsky & Lukina 1977; Eiselt et al. 1992; Arribas et al. 2013; Rato et al. 2021; Candan et al. 2021). This complexity has always been attractive to researchers who apply both kinds of markers, morphology and/or more recently genetics trying to solve it. Elaborated recent assessments using genetic markers point out that there are more lineages within the D. valentini / D. rudis complexes than the previously suspected (Candan et al. 2021; Rato et al. 2021). In this study, we aimed to increase the knowledge of the status of the currently recognized genetic lineages by creating the largest datasets, including a remote subspecies not studied so far – D. v. spitzenbergerae – for the first time, to use in both morphological and molecular analyses to clarify the problem. Our phylogenetic results show the presence of several monophyletic clades that reveal themselves as different species (Fig. 9). Of these distinct clades, some had been well-documented for the first time in a recently published study (Candan et al. 2021), and the authors have accepted that D. valentini s. lat. has more genetic lineages than previously suspected, two of which have been presented there as they should have to be described and named. However, two important shortcomings that we have tried to eliminate here prevented them from their taxonomic description: the lack of morphological study for diagnoses and the absence of samples of one of the up to now two unique subspecies of D. valentini (D. v. spitzenbergerae), whose study was unavoidable to make taxonomic decisions. As seen from the tree topology obtained here, genetically divergent lineages, clades A and B, were detected as monophyletic (Fig. 9). The occurrence of these two highly divergent monophyletic lineages is not only confirmed by the tree topology but also the species delimitation analyses revealed both clades as different species, which is one of the most important factors that paved the way for the here proposed taxonomic revision. Network analyses based on both genetic markers also supported this distinction. In Cyt-b, all clades were placed into their unique positions and they did not share any haplotypes (Fig. 10A). In MC1R, which is a nuclear marker and has slower substitution rates, an agreement relatively with a more complex structure was showed. Clade A is represented by a single haplotype (Hap16), while clade B appears to have two haplotypes (Hap16 and Hap26) (Fig. 10B). Although these results were suggested by Candan et al. (2021), a definite conclusion could not be made due to the absence of subspecies D. v. spitzenbergerae, a problem now solved. Considering sampling data used in our phylogenetic construction, clade A consists of both D. s. spitzenbergerae from Mergan Plateau (type locality of this relevant and geographically extreme subspecies) and D. s. wernermayeri ssp. nov. from Narlıca Valley as sister taxa (Fig. 9). Although the population located in Narlıca Valley (Van, Turkey) is morphologically included in D. v. lantzicyreni (Eiselt et al. 1992), it is genetically more closely related to D. s. spitzenbergerae than to the former. In addition to this, the populations, which were assimilated to D. v. lantzicyreni according to morphology (Eiselt et al. 1992), represent a completely different lineage (clade B) according to genetics. Such discordant patterns called cryptic speciation are often shown in the lizards (Ahmadzadeh et al. 2013; Kornilios et al. 2018; Karakasi et al. 2021; Arribas et al. 2022). Another major point is the status of D. r. mirabilis (clade C). This subspecies was first described by Arribas et al. (2013) from Ovit Pass, a very isolated geographic region in Kaçkar Mountains. Its phylogenetic position is obvious here and reveals that it should be a species as different as clades A and B (Fig. 9). The genetic difference of this taxon was demonstrated by two independent studies. Firstly, Rato et al. (2021) suggested that a clade, called Trabzon-Rize in their study, is genetically distinct and that it should be considered one of the four main lineages of D. rudis. Since they did not distinguish any subspecies, they could not determine that this clade belongs to D. r. mirabilis. The fact that the D. rudis specimens used in their study share the same branch with a specimen we know for certain to be D. r. mirabilis, undoubtedly proves that this clade is a new taxon and the corresponding samples of Rato et al. (2021) belong to it. Secondly, Candan et al. (2021) has also mentioned that it has isolated genetic structure and that its distribution area may be wider than expected because a datum retrieved from GenBank (Tarkhnishvili et al. 2013), which is located around Sarıkamış (Kars, Turkey), clustered with D. r. mirabilis in the same branch. Similar to Candan et al. (2021), one of the interesting results obtained within D. valentini / D. rudis complexes is that the specimens belonging to D. r. rudis, D. r. bolkardaghica and D. v. lantzicyreni, cluster together with overlapping. This unexpected pattern makes it difficult to engage the complexity of the group, which unables to apply the current nomenclature and difficulties understanding the main processes underlying genetic variation. Considering the genetic (Fig. 9) and morphological (see results section) evidence together, the most possible scenario seems to accept that D. v. lantzicyreni is really a subspecies of D. rudis, not from D. valentini. Thus, nominal form of D. valentini is only limited to northeast Anatolia (with areas of Georgia, Armenia and Azerbaijan), while the distribution of D. rudis sensu novo, extends from the northeastern Black Sea region to the inner Anatolia and from there to the south up to the Bolkar Mountains. Finally, the status of some former subspecies of D. rudis also inevitably needs revision. The claim that a member of this group, D. r. obscura, is different has been put forward in a previous study including phenotypic comparison (Gabelaia et al. 2018; Tarkhnishvili et al. 2020b). The phylogenetic results strongly support these morphological findings (clade G, Fig. 9). Above all, D. r. obscura has a phylogenetic position quite closely related to other two former D. rudis subspecies: D. r. bischoffi and D. r. macromaculata. Considering all these results, it seems that accepting the first described form, D. saxicola obscura Lantz & Cyren 1936, as a species: D. obscura will contribute positively to the clarification of this group. Morphology derived structure Considering the studied complex group it seems that there are three large groups, which obviously coincide with the current taxonomy based on morphology (we still use here the old nomenclature to refer them). The most different includes D. bithynica s. str. and D. b. tristis, which had longer heads both concerning its width, and also about their body length, but not in their pilei because other species (especially of the former rudis complex) had smaller (in size and length) but very wide heads. Similarly, the scales that cover the upper part of the crus are small and barely keeled. Also, they had comparatively longer hindlimbs (are the more climbing, based on this characteristic). Osteologically, they have very rarely any B-Type pre-autotomic vertebrae. The sternal fontanelle is frequently reduced or absent in D. b. bithynica. Postorbital and postfrontal are subequal or the postorbital is a bit smaller (different to D. valentini, clade B and D. v. spitzenbergerae). Squamosal and postorbital overlap commonly in half of the second’s length (as also in D. r. bolkardaghica), more usually than in other forms of the group. Darevskia b. bithynica and D. b. tristis are identical in ANOSIM. This species is also recovered by genetics. Genetics indicates the possibility that tristis is paraphyletic as presently understood. The former valentini complex has a broad overlap among the different forms in CDA. These valentini complex samples had comparatively longer limbs, comparatively smaller heads, a greater number of scales in the crus (which in this case corresponds also to smaller scale size, and are no or almost-none keeled), and less markedly, a greater number of ventral and dorsal scales. Anal index, a bit greater (scale comparatively wider) in D. valentini than in D. bithynica or the former rudis complex. One of their supposed taxa, D. v. lantzicyreni, perhaps due to its wide dispersal and the presence of isolated populations, appears somewhat heterogeneous. Darevskia v. lantzicyreni overlaps a few with D. r. bolkardaghica (in males, and even more in females, which would be in agreement with the genetic results and the taxonomic changes proposed above). In turn, D. v. lantzicyreni has the higher dorsalia among the former valentini complex and is the closer of this complex to D. rudis s. str., which would also agree with the genetic analysis (see above) and its relation as conspecific. Darevskia r. bolkardaghica is characterized by low lamellae (in males and females), and osteologically is characterized because not infrequently shows a weakly ossified rib associated to the third vertebra (an extremely rare character, probably atavistic, associated to small and isolated populations), and the sternal fontanelle adopt singular shapes in sand-clock, irregular cordiform or trilobate in its forepart.Also its squamosal and postorbital bones overlap commonly in half of the second’s length (as in D. bithynica). Genetically, it is related to D. v. lantzicyreni and D. r. rudis (all three are proposed here as subspecies of D. rudis) Darevskia v. spitzenbergerae, clade A (here treated as the nominal ssp. of D. spitzenbergerae), and clade B (here described as a new species) (in males) and in general, as all the former valentini complex (in females) have higher ventralia counts than in D. bithynica or the former rudis complex taxa. Osteologically, this singular form (spitzenbergerae) has the interclavicle lateral branches inclined forwards (with only this model in typical D. s. spitzenbergerae) and in “clade A” (coexisting with some branches backward). Postorbital and postfrontal are subequal or the postorbital is a bit greater (as in D. valentini s. str. or clade B). Nominal taxa spitzenbergerae +clade A, and clade B are recovered as two different species by genetics. Darevskia spitzenbergerae and clade A are primitive forms, among the closely related to D. rudis (sensu novo) and their subspecies (lantzicyreni and bolkardaghica, especially this latter). Darevskia valentini s. str. seems to be a different taxon (genetics) without its formerly assigned subspecies (is nominotypical). It has Temporalia2 a bit lower than in related taxa. In D. valentini s. str. not infrequently appear some B-Type autotomic vertebrae. Postorbital and postfrontal are subequal or the postorbital is a bit greater (as in clade B or D. spitzenbergerae). Darevskia rudis complex is characterized by smaller dorsalia in a great part of the rudis complex (except in D. r. rudis –yet indicated in Arribas et al. 2013, that also is recovered as a different species in genetic analyses), than in the former valentini complex and D. bithynica ssp. (except in D. v. valentini that has lower scores similar to the main former rudis complex). Hindlimb relative length is also comparatively smaller when compared with the former valentini complex and even more with D. bithynica. These differences shall be considered as characteristic of D. obscura and their newly assigned subspecies, which are very few differentiated. Darevskia r. obscura and D. r. macromaculata are near the same by morphology, as suggested in Arribas et al. (2013). They are so similar in all analyses (including non-significant differences in ANOSIM) that they appear to be the same (increased pigmentation in typical Georgian macromaculata, but perhaps not in Turkish specimens, a question to be studied in future). Temporalia1 is somewhat smaller in D. r. macromaculata and D. r. obscura (M, F), and SVL (size) is greater in D. r. bischoffi (M, F). Paradoxically, D. rudis s. str. is morphologically extreme and a differentiated form within “its” former complex, and is distinguished from the other former rudis complex taxa (now D. obscura sspp.) by its greater values of dorsalia. Also, it is basal to the group in UPGMA. Osteologically, in D. rudis s. str. postorbital and postfrontal are subequal or the first is smaller than the second (as in D. obscura and D. bithynica). The two extreme populations displaced towards the south of both classical complexes (D. rudis bolkardaghica and D. valentini spitzenbergerae) result in the ones that connect morphologically the former rudis and valentini complexes.This may be because they are the most primitive in both groups, or because of an ecoclimatic convergence in their scalation. Both live on calcareous substrates (siliceous, even volcanic in the other forms), so they have a lighter background color than other forms (darker). Concordance of genetic and morphological results a) The classic morphological groupings/species (rudis and valentini complexes) seem to be no longer valid, due to newly discovered “intermediate” taxa, recent speciation, and multiple past and present introgression. The situation is fairly more complex than previously expected. b) The above-mentioned morphological characteristics of D. bithynica are valid for this species. c) The above-mentioned morphological characteristics of D. rudis complex are valid for D. obscura (and its sspp. macromaculata and bischoffi). d) As stated above D. rudis s. str. is distinguished from the other former rudis complex taxa (hereinafter D. obscura sspp.) by its greater values of dorsalia. Also, it is basal to the group in UPGMA. e) Darevskia r. lantzicyreni comb. nov. and D. r. bolkardaghica are subspecies of D. rudis. f) Darevskia spitzenbergerae is a different taxon. The fourth axis of females analysis (at the limit of significance) discriminates specially clade A (D. s. wernermayeri nov. ssp.) and in a lesser degree D. v. spitzenbergerae (that genetically cluster together and are very similar in ANOVA), characterized by higher values of lamellae, preanalia and femoralia. Inside this spitzenbergerae genetic clade, preanalia is higher in clade A (D. s. wernermayeri nov. ssp.) and strongly characterizes it (M, F) concerning near all the taxa studied here (and if the Tukey-Kramer multiple comparison test is used instead of the much stricter of Scheffe, is significantly different to all the taxa, including the nominal D. s. spitzenbergerae) g) Darevskia mirabilis is another taxon, genetically singular, and only moderately differentiated in its morphology within the former rudis complex (morphologically seems more related to D. o. obscura or D. r. bolkardaghica –Anatolian diagonal effect? -), and longtime approached to D. valentini by its pattern, but well isolated genetically. In ANOVA, very low supraciliar granula (M, F) (especially distinctive of this taxon) and gularia (M), and higher circumanalia (M) counts are the most diagnostic characters. h) The number (tibialia), size and keeling of the crus scales was formerly used to distinguish between the forms assigned to “ rudis ” and “ valentini ” (sensu auctt.) and is distinctive with higher counts (and smaller scales size and keeling) in the former valentini complex (and D. b. bithynica), appearing, to the contrary, the lower counts (with big size and strong keeling) in part of the former rudis

Darevskia obscura obscura stat. et comb. nov. 38(M), 35(F) 1. ZDEU 43 /2016. (N=7), Kutul Plateau, Ardahan, Turkey, 18.07.2016, Leg. Ç. ILGAZ, K. CANDAN [Map ID: 59]. 2. ZDEU 17 / 2010. (N=22), Kutul Plateau, Ardahan, Turkey 14.07.2010, Leg. Y. KUMLUTAŞ, Ç. ILGAZ, A. AVCI, N. ÜZÜM, B. ÜZÜM [Map ID: 59]. 3. ZDEU 156 /2001. (N=44), Between Geçitli Village and Bilbilen Plateau, Ardanuç, Artvin, Turkey, 06.07.2001, Leg. Y. KUMLUTAŞ, K. OLGUN, Ç. ILGAZ, A. AVCI, F. İRET [Map ID: 60]., Published as part of Arribas, Oscar, Candan, Kamil, Kornilios, Panagiotis, Ayaz, Dinçer, Kumlutaş, Yusuf, Gül, Serkan, Yilmaz, Can, Caynak, Elif Yildirim & Ilgaz, Çetin, 2022, Revising the taxonomy of Darevskia valentini (Boettger, 1892) and Darevskia rudis (Bedriaga, 1886) (Squamata, Lacertidae): a Morpho-Phylogenetic integrated study in a complex Anatolian scenario, pp. 1-68 in Zootaxa 5224 (1) on page 55, DOI: 10.11646/zootaxa.5224.1.1, http://zenodo.org/record/7517907

Darevskia rudis rudis 89(M), 72(F) 1. ZDEU 200 /2014. (N=10), Karacaören Village, Başçiftlik, Tokat, Turkey, 18.06.2014, Leg. Y. KUMLUTAŞ [Map ID: 16]. 2. ZDEU 99 /2013. (N=2), Tekkeköy, Samsun, Turkey, 02.08.2013, Leg. Y. KUMLUTAŞ [Map ID: 15]. 3. 1 ZDEU 43 /2014. (N=7), Şalpazarı, Trabzon, Turkey, 04.08.2014, Leg. Y. KUMLUTAŞ, Ç. ILGAZ, K. CANDAN [Map ID: 35]. 4. ZDEU 133 /2014. (N=4), Zigana Pass, Gümüşhane, Turkey, 04.08.2014, Leg.Y. KUMLUTAŞ, Ç. ILGAZ, K. CANDAN [Map ID: 34]. 5. ZDEU 2 / 2010. (N=23), Between Sürmene and Köprübaşı 8.km., Trabzon, Turkey, 17.07.2010, Leg. Y. KUMLUTAŞ, Ç. ILGAZ, A. AVCI, N. ÜZÜM, B. ÜZÜM [Map ID: 41]. 6. ZDEU 40 / 2010. (N=23), Maçka, Trabzon, Turkey, 18.07.2010, Leg. Y. KUMLUTAŞ, Ç. ILGAZ, A. AVCI, N. ÜZÜM, B. ÜZÜM [Map ID: 40]. 7. ZDEU 60 / 2010. (N=27), Between Akçaabat and Düzköy 14.km., Trabzon, Turkey, 18.07.2010, Leg. Y. KUMLUTAŞ, Ç. ILGAZ, A. AVCI, N. ÜZÜM, B. ÜZÜM [Map ID: 38]. 8. ZDEU 50 /2003. (N=19), Zigana Pass, Trabzon, Turkey, 10.07.2003, Leg. İ. BARAN, Y. KUMLUTAŞ, Ç. ILGAZ, C.V. TOK, F. İRET [Map ID: 34]. 9. ZDEU 51 /2003. (N=20), Between Sümele and Maçka 10.km., Trabzon, Turkey, 10.07.2003, Leg. İ. BARAN, Y. KUMLUTAŞ, Ç. ILGAZ, C.V. TOK, F. İRET [Map ID: 39]. 10. ZDEU 53 /2003. (N=10), Between Beşikdüzü and Şalpazarı 7.km., Trabzon, Turkey, 11.07.2003, Leg. İ. BARAN, Y. KUMLUTAŞ, Ç. ILGAZ, C.V. TOK, F. İRET [Map ID: 36]. 11. ZDEU 54 /2003. (N=16), Between Tonya and Vakfıkebir 10-15.km., Trabzon, Turkey, 11.07.2003, Leg. İ. BARAN, Y. KUMLUTAŞ, Ç. ILGAZ, C.V. TOK, F. İRET [Map ID: 37]., Published as part of Arribas, Oscar, Candan, Kamil, Kornilios, Panagiotis, Ayaz, Dinçer, Kumlutaş, Yusuf, Gül, Serkan, Yilmaz, Can, Caynak, Elif Yildirim & Ilgaz, Çetin, 2022, Revising the taxonomy of Darevskia valentini (Boettger, 1892) and Darevskia rudis (Bedriaga, 1886) (Squamata, Lacertidae): a Morpho-Phylogenetic integrated study in a complex Anatolian scenario, pp. 1-68 in Zootaxa 5224 (1) on pages 54-55, DOI: 10.11646/zootaxa.5224.1.1, http://zenodo.org/record/7517907

Darevskia obscura macromaculata comb. nov. 24(M), 27(F) 1. ZDEU 35 /2010. (N=51), Sahara National Park, Ardahan, Turkey, 13.07.2010, Leg. Y. KUMLUTAŞ, Ç. ILGAZ, A. AVCI, N. ÜZÜM [Map ID: 58]., Published as part of Arribas, Oscar, Candan, Kamil, Kornilios, Panagiotis, Ayaz, Dinçer, Kumlutaş, Yusuf, Gül, Serkan, Yilmaz, Can, Caynak, Elif Yildirim & Ilgaz, Çetin, 2022, Revising the taxonomy of Darevskia valentini (Boettger, 1892) and Darevskia rudis (Bedriaga, 1886) (Squamata, Lacertidae): a Morpho-Phylogenetic integrated study in a complex Anatolian scenario, pp. 1-68 in Zootaxa 5224 (1) on page 55, DOI: 10.11646/zootaxa.5224.1.1, http://zenodo.org/record/7517907

Darevskia spitzenbergerae wernermayeri ssp. nov. (Appendix 5; Fig. 12a). Synonymy/Chresonymy: Lacerta valentini lantzicyreni; Eiselt, Darevsky & Schmidtler, 1992. (from “Yukarı Narlıca”-sic.!-). Darevskia valentini “Clade A”; Candan et al. (2021). (same locality as this study) ZooBank registration (http://zoobank.org): urn:lsid:zoobank.org:act: 927894FD-EEFD-450A-81D9-0A68188EDC3B. Holotype: ZDEU123 /2015 (n.3). ♁, Yukarınarlıca Village, Çatak, Van, Turkey. leg. Yusuf Kumlutaş, Çetin Ilgaz, 29.07.2015. Paratypes: 7 ♁♁, 10 ♀♀. Same locality, date and collectors as holotype. Derivatio nominis: The specific epithet refers to Dr. Werner Mayer (1943-2015), for his remarkable work on the knowledge of lacertid genera relationships and species taxonomy. Diagnosis: Darevskia spitzenbergerae wernermayeri ssp. nov. differs from nominate form (D. s. spitzenbergerae) in having a higher number of supraciliar granules (18-31 vs. 14-23), supratemporal (4-6 vs. 3-6), ventrals (29-31 vs. 26-30) (females), preanals (1-3 vs. 1-3), tibial scales (16-21 vs. 15-19), temporal scales 1 (5-7 vs. 2-6) and temporal scales 2 (4-6 vs. 2-6). Darevskia s. wernermayeri ssp. nov. has a relatively smaller head relative length (0.18- 0.21 vs. 0.19-0.24). It also differs by a characteristic color pattern of the body. The main osteological diagnostic characters that differ from the nominate form can be specified as follows: The higher number of maxillary (19-21 vs. 16-18) and dentary teeth (23-24 vs. 20-23). Postorbital greater or equal than postfrontal (greater, rarely equal in D. s. spitzenbergerae). Description of holotype: An adult male. Tail regenerated (see Appendix 5c). Fixed with ethanol and conserved in 96% ethanol. Scalation: Rostral not in contact with the nostril. Single postnasal on each side. Width of frontonasal (internasal) plate subequal to length, not in contact with rostral. Sutures between prefrontal plates and frontal plate straight. Parietal plates in contact with postorbital plates on each side. Supraciliar granules 13 and 12, interrupted series on left, continuous on right. Supraciliar plates 6 on each side. Supralabial and sublabial plates 4 and 6 on each side, respectively. Plates in supratemporal region 6 on left, 5 on right side. The first supratemporal plate large narrows towards the back, and ends bluntly. Masseteric large, in one piece on left and two pieces on right, separated from the first supratemporal plate by three longitudinal scales on each side. Tympanic obvious, separated from masseteric by three and two rows of small scales on left and right, respectively. Nine flat collaria. Gularia 33. Ventralia contains 6 longitudinal and 28 transverse rows of plates. Preanal scale singular, surrounded by 6 rows of plates. Femoral pores 20 on each side. Subdigital lamellae 28 on each side. Tibialia 20. Dorsalia 51. Biometry: SVL 63.57 mm. Pileus width 6.90 mm, pileus length 12.96 mm, head width 8.26 mm, head length 13.74 mm. Length of forelimb 22.02 mm, length of hindlimb 31.37 mm. Anal plate width and length are 4.12 mm and 1.84 mm, respectively. Coloration and pattern (in alcohol): Ground color of dorsum greenish-brown, with two irregular paravertebral rows of dark spots. Similar dark spotting is present on each side of the body, that show the reminiscent of a reticulate pattern reduced to only isolated irregular spots. Between these two areas of dark spots, located both in the middle of the dorsum and flank, a paler and spot-free double strip area extends from nape to base of the tail (see photos in Appendix 5). There are few pale spots on dark bandings on the flanks, and more on the forelimb. A few spots near the basis of the forelimb are bluish. The belly, along with the head and neck, is whitish (Appendix 5). The background color of head plates is brownish, with a few scattered and small black spots. The first longitudinal row of ventral plates has dark spots on each side. Variations of paratypes: Descriptive statistics and variation range of the morphometric and scalation characters are given in Table 8. Frontonasal rarely contacts with rostral. The suture between the prefrontal and frontal is usually slanted. The Parietal is in contact with the postorbital in general. Supraciliar granules sometimes are double rows. Tight scales are feeble-keeled. In four specimens, transverse dark spots on ground color are combined with spots on each side, do not form two separate rows. In seven samples, paler spots on the forelimb base are not blue. Belly, along with throat and neck, whitish in seven specimens. In three samples, no dotting on head plates. In fifteen samples, the first longitudinal row of ventral plates contains bluish spots on each side. Distribution: Around Narlıca Valley, Çatak (Van) in the south of Van Lake, Turkey. Habitat: Subalpine-like vegetation of Irano-Turanian Region, on rocky and stony areas, 2363 m. No other reptile species could be identified in the area during study., Published as part of Arribas, Oscar, Candan, Kamil, Kornilios, Panagiotis, Ayaz, Dinçer, Kumlutaş, Yusuf, Gül, Serkan, Yilmaz, Can, Caynak, Elif Yildirim & Ilgaz, Çetin, 2022, Revising the taxonomy of Darevskia valentini (Boettger, 1892) and Darevskia rudis (Bedriaga, 1886) (Squamata, Lacertidae): a Morpho-Phylogenetic integrated study in a complex Anatolian scenario, pp. 1-68 in Zootaxa 5224 (1) on pages 24-25, DOI: 10.11646/zootaxa.5224.1.1, http://zenodo.org/record/7517907, {"references":["Eiselt, J., Darevsky, I. S. & Schmidtler, J. F. (1992) Untersuchungen an Felseneidechsen (Lacerta saxicola komplex) in der ostlichen Turkei, I. Lacerta valentini Boettger. Annalen des Naturhistorischen Museums in Wien, 93 (B), 1 - 18.","Candan, K., Kornilios, P., Ayaz, D., Kumlutas, Y., Gul, S., Yildirim-Caynak, E. & Ilgaz, C. (2021) Cryptic genetic structure within Valentin's Lizard, Darevskia valentini (Boettger, 1892) (Squamata, Lacertidae), with implications for systematics and origins of parthenogenesis. Systematics and Biodiversity, 19 (7), 665 - 681. https: // doi. org / 10.1080 / 14772000.2021.1909171"]}

Darevskia josefschmidtleri sp. nov. (Appendix 5; Fig. 12e). Synonymy/Chresonymy: Lacerta valentini “Zwischenpopulation ”(intermediate population); Eiselt, Darevsky & Schmidtler, 1992. (from “Çaldıran”- sic.!-) Darevskia valentini “Clade B”; Candan et al. (2021) (same locality as this study) ZooBank registration (http://zoobank.org): urn:lsid:zoobank.org:act: 56CFE08E-164E-485B-8F4C-94EA76293128. Holotype: ZDEU220 /2016 (n.2). ♁, Başeğmez Village, Çaldıran, Van, Turkey. leg. Kamil Candan, Nurettin Beşer and Mehmet Kürşat Şahin, 24.06.2016. Conserved in ZDEU collection. Paratypes: 8 ♁♁, 8 ♀♀, 2 subadult ♀♀. Same locality, date and collectors as holotype. ZDEU221 /2016, 6 ♁♁, 12 ♀♀, 1 juvenile, Çirişli Village, Çat, Erzurum, Turkey. leg. Kamil Candan, Nurettin Beşer, Mehmet Kürşat Şahin, 22.06.2016. ZDEU222 /2016, 5 ♁♁, 8 ♀♀, 1 juvenile, Palandöken Mountain, Erzurum, Turkey. leg. Kamil Candan, Nurettin Beşer, Mehmet Kürşat Şahin, 01.07.2016. ZDEU119 /2015, 1 ♁♁, 3 ♀♀, 2 ♀♀ subadults, Balık Lake, Taşlıçay, Ağrı, Turkey. leg. Kamil Candan, Elif Yıldırım Caynak, 26.07.2015. ZDEU121 /2015, 2 ♁♁, 4 ♀♀, Güzeldere Village, Hınıs, Erzurum, Turkey, leg. Kamil Candan, Elif Yıldırım Caynak, 25.07.2015. Derivatio nominis: The specific epithet refers to Josef Friederich Schmidtler (born 1942), for his remarkable work on the knowledge of Turkish herpetofauna and its rich diversity. Comparative diagnosis (Morphology): Darevskia josefschmidtleri sp. nov. is a medium sized species (adults SVL: 53.25–67.95 mm). It is characterized by medium or small-sized scales with feeble keeling (even barely visible). Darevskia josefschmidtleri sp. nov. differs from D. valentini in that there is a higher number of lamellae (46–59 vs. 42–53) and dorsals (47–58 vs. 41–52); Darevskia josefschmidtleri sp. nov. males have a higher number of preanal than D. valentini (1–3 vs. 1) while females have different collar scores (8–12 vs. 7–10) and gulars (23–31 vs. 21–29). It also differs from D. valentini in having a greater head relative length (0.18–0.23 vs. 0.16.–0.21) for males. Darevskia josefschmidtleri sp. nov. differs from D. spitzenbergerae spitzenbergerae s tat. et comb. nov. (see below) in that there is higher number of tibials (16–24 vs. 15–19) and dorsals (48–58 vs. 44–53) for males. Also, it differs from D. spitzenbergerae spitzenbergerae in having a shorter head relative length (0.18–0.23 vs 0.19–0.24). Darevskia josefschmidtleri sp. nov. differs from “Clade A” from Candan et al. (2021) (described below as D. spitzenbergerae wernermayeri ssp. nov.) in that there is a lower number of ventrals (26–29 vs. 26–31), temp 2 (2–6 vs. 4–6) in males and preanals (1–3 vs. 2–3) in females. Darevskia josefschmidtleri sp. nov. differs from D. valentini in that there is a lower number of maxillary (15–18 vs. 16–20) and dentary teeth (20–22 vs. 18–25). Darevskia josefschmidtleri sp. nov. differs from Clade A (D. spitzenbergerae wernermayeri ssp. nov. see below) in that there is a lower number of maxillary (15–18 vs. 19–21) and dentary teeth (20–22 vs. 23–24). Diagnosis (Molecular): It can be distinguished from other former D. valentini populations by unique nucleotide combination located on the mitochondrial gene Cyt-b and nuclear loci MC1R. The consensus sequence (Cyt-b) for Darevskia josefschmidtleri sp. nov. is found in Appendix 6, together with the respective sequence for the D. valentini s. str. In this Table, the thirty simple nucleotide diagnostic characters between the consensus sequences are highlighted. Similarly, the molecular diagnostic characters for Darevskia josefschmidtleri sp. nov. regarding the nuclear loci are shown in Appendix 6. Description of holotype: An adult male. Tail in regeneration process. Fixed with ethanol and conserved in 96% ethanol. Scalation: Rostral not in contact with the nostril. Single postnasal on each side. Width of frontonasal (internasal) plate subequal to length, not contact with rostral. Sutures between prefrontal plates and frontal plate straight. Parietal plates in contact with postorbital plates on each side. Supraciliar granules 9 on each side, interrupted series on right, not on left side. Supraciliar plates 6 on each side. Supralabial and sublabial plates 4 and 6 on each side, respectively. Plates in supratemporal region 3 on left, 4 on right. The first supratemporal plate large, narrow towards the back, ends bluntly. Masseteric large, in one piece on each side, separated from the first supratemporal plate by a row of small scales. Tympanic obvious, in two pieces on the left and one piece on right, separated from masseteric by three and two rows of small scales on the left and right, respectively. Eight flat and smooth collaria. Gularia 27. Ventralia contains 6 longitudinal and 28 transverse rows of plates. Preanal scale in two pieces, surrounded by 6 rows of plates. Femoral pores 21 on left, 20 on right side. Subdigital lamellae 27 on left, 26 on right. Tibial scale 19. Dorsalia 53. Biometry: SVL 64.68 mm., pileus width 6.79 mm, pileus length 13.27 mm, head width 8.13 mm, head length 13.14 mm. Length of forelimb 18.39 mm, length of hindlimb 27.90 mm. Anal plate width 4.69 mm, length 1.71 mm. Coloration and pattern (in alcohol): The ground color of the dorsum is greenish brown. Dorsal tract with a wide vertebral band composed of fairly irregular transverse spots nost covering almost its complete width. Similar dark spotting is present on each side of the body (temporal or lateral band) that appears reticulated. Between these two dark spots, located both in the middle of the dorsum and flanks, a paler double line extends from the nape to the base of the tail (Appendix 5). There are few pale spots inside the reticulate on dark bands on flanks, with more in the forelimb. A few light bluish spots near the forelimb basis. Belly, along with head and neck, whitish (see photos in Appendix 5), with dark and blue spots in the outermost rows of ventrals. Background color of head plates brownish, with few scattered and small black spots on it. Variations of paratypes: Descriptive statistics and variation range of the morphometric and scalation characters are given in Table 7. Frontonasal (internasal) rarely is in contact with rostral. Sutures between the prefrontal and frontal are usually slanted. Parietal is rarely in contact with postorbital. Masseteric is sometimes divided into two pieces. Tight scales are feeble-keeled, smaller than dorsal ones. In one specimen, transverse dark spots on ground color combined with spots on each side, do not form two separate rows. In addition, transverse dark spots, longitudinally above ground color, are faint in ten specimens. In four samples, pale spots on the forelimb base were not blue (white). Belly, along with head and neck, yellowish in nine specimens. In fourteen samples, no spotting on head plates. Distribution: Confirmed localities draw an area around Çaldıran (Van) located in the east of Van Lake, around Balık Lake (Ağrı), Palandöken Mountain and around Çat and Hınıs (Erzurum) which is in the south of this massif, all in Turkey. Probably also in intermediate areas among these localities. Habitat: Subalpine-like vegetation from Irano-Turanian Region, in rocky and stony areas near 2000 m or higher: 2095 m. (Çaldıran), 2270 m. (Balık Lake), 2429 m. (Palandöken), 1946 m. (Çat), 2643 m. (Hınıs). Darevskia unisexualis was found sympatric in the Palandöken Mountain, while no other reptile species were detected in the same and other localities of the area. Comments: This new taxon seems to be the parental species that gave origin by hybridization to the parthenogenetic D. unisexualis, D. sapphirina and D. bendimahiensis according to Z-chromosome inheritance (Yanchukov et al. 2022).

Darevskia spitzenbergerae (Eiselt, Darevsky & Schmidtler, 1992)stat. nov. (Fig. 12g). Type Locality: Mergan Plateau, Cilo Mountain, Hakkari, Turkey. Distribution: It is known from only two locations: Mergan Plateau, Hakkari, Turkey and Narlıca Valley, Van, Turkey (this study). Comments: It includes one of the subspecies of D. valentini previously described and a population from Narlıca Valley (called the New Clade A in Candan et al. 2021). Since they are morphologically distinct and D. s. spitzenbergerae s tat. et comb. nov. is so singular in pattern, we think it may be subspecifically different. Clade A is described as subspecies of Darevskia spitzenbergerae s tat. nov., Published as part of Arribas, Oscar, Candan, Kamil, Kornilios, Panagiotis, Ayaz, Dinçer, Kumlutaş, Yusuf, Gül, Serkan, Yilmaz, Can, Caynak, Elif Yildirim & Ilgaz, Çetin, 2022, Revising the taxonomy of Darevskia valentini (Boettger, 1892) and Darevskia rudis (Bedriaga, 1886) (Squamata, Lacertidae): a Morpho-Phylogenetic integrated study in a complex Anatolian scenario, pp. 1-68 in Zootaxa 5224 (1) on page 23, DOI: 10.11646/zootaxa.5224.1.1, http://zenodo.org/record/7517907, {"references":["Eiselt, J., Darevsky, I. S. & Schmidtler, J. F. (1992) Untersuchungen an Felseneidechsen (Lacerta saxicola komplex) in der ostlichen Turkei, I. Lacerta valentini Boettger. Annalen des Naturhistorischen Museums in Wien, 93 (B), 1 - 18.","Candan, K., Kornilios, P., Ayaz, D., Kumlutas, Y., Gul, S., Yildirim-Caynak, E. & Ilgaz, C. (2021) Cryptic genetic structure within Valentin's Lizard, Darevskia valentini (Boettger, 1892) (Squamata, Lacertidae), with implications for systematics and origins of parthenogenesis. Systematics and Biodiversity, 19 (7), 665 - 681. https: // doi. org / 10.1080 / 14772000.2021.1909171"]}

Darevskia obscura bischoffi comb. nov. 82(M), 86(F) 1. ZDEU 38 /2015. (N=7), Cankurtaran Pass, Hopa, Artvin, Turkey, 23.07.2015, Leg. Y. KUMLUTAŞ, Ç. ILGAZ [Map ID: 51]. 2. ZDEU 201 /2014. (N=2), Maden village, Artvin, Turkey, 23.07.2014, Leg. Y. KUMLUTAŞ [Map ID: 57]. 3. ZDEU 7 / 2010. (N=15), Balcılar Village, Borçka, Artvin, Turkey, 15.07.2010, Leg. Y. KUMLUTAŞ, Ç. ILGAZ, A. AVCI, N. ÜZÜM, B. ÜZÜM [Map ID: 55]. 4. ZDEU 6 / 2010. (N=16), Between Borçka and Hopa 8.km., Artvin, Turkey, 15.07.2010, Leg. Y. KUMLUTAŞ, Ç. ILGAZ, A. AVCI, N. ÜZÜM, B. ÜZÜM [Map ID: 52]. 5. ZDEU 31 / 2010. (N=10), Between Arhavi and Güneşli Village, 2.km., Artvin, Turkey, 15.07.2010, Leg. Y. KUMLUTAŞ, Ç. ILGAZ, A. AVCI, N. ÜZÜM, B. ÜZÜM [Map ID: 47]. 6. ZDEU 4 / 2010.(N=11), Between Çamlıhemşin and Ayder Plateau 3.km., Rize, Turkey, 16.07.2010, Leg.Y. KUMLUTAŞ, Ç. ILGAZ, A. AVCI, N. ÜZÜM, B. ÜZÜM [Map ID: 46]. 7. ZDEU 25 / 2010. (N=19), Hemşin, Rize, Turkey, 16.07.2010, Leg. Y. KUMLUTAŞ, Ç. ILGAZ, A. AVCI, N. ÜZÜM, B. ÜZÜM [Map ID: 45]. 8. ZDEU 10 / 2010. (N=17), Between İkizdere and İspir, 19.km., Rize, Turkey, 17.07.2010, Leg. Y. KUMLUTAŞ, Ç. ILGAZ, A. AVCI, N. ÜZÜM, B. ÜZÜM [Map ID: 43]. 9. ZDEU 158 /2001. (N=9), Between Borçka and Camili 10-21.km., Artvin, Turkey, 07.07.2001, Leg. Y. KUMLUTAŞ, K. OLGUN, Ç. ILGAZ, A. AVCI, F. İRET [Map ID: 56]. 10. ZDEU 116 /2002. (N=16), Between Borçka and Balcılar, Artvin, Turkey, 12.07.2002, Leg. İ. BARAN, Y. KUMLUTAŞ, Ç. ILGAZ, A. ÖZDEMİR [Map ID: 54]. 11. ZDEU 130 /2002. (N=13), Between Rize and Küçükçayır 18.km., Rize, Turkey, 14.07.2002, Leg. İ. BARAN, Y. KUMLUTAŞ, Ç. ILGAZ, A. ÖZDEMİR [Map ID: 44]. 12. ZDEU 124 /2002. (N=16), Between Ortacalar and Dülgerli 16-24.km., Artvin, Turkey, 13.07.2002, Leg. İ. BARAN, Y. KUMLUTAŞ, Ç. ILGAZ, A. ÖZDEMİR [Map ID: 48]. 13. ZDEU 163 /2001. (N=3), Esenkıyı Village, Hopa, Artvin, Turkey, 07.07.2001, Leg. Y. KUMLUTAŞ, K. OLGUN, Ç. ILGAZ, A. AVCI, F. İRET [Map ID: 49]. 14. ZDEU 105 /2000. (N=6), Çamurköy, Sarp, Artvin, Turkey, 24.04.2000, Leg. K. OLGUN [Map ID: 50]. 15. ZDEU 102 /2002. (N=8), Between Artvin and Hatila Plateau 35.km., Artvin, Turkey, 09.07.2002, Leg. İ. BARAN, Y. KUMLUTAŞ, Ç. ILGAZ, A. ÖZDEMİR [Map ID: 53]., Published as part of Arribas, Oscar, Candan, Kamil, Kornilios, Panagiotis, Ayaz, Dinçer, Kumlutaş, Yusuf, Gül, Serkan, Yilmaz, Can, Caynak, Elif Yildirim & Ilgaz, Çetin, 2022, Revising the taxonomy of Darevskia valentini (Boettger, 1892) and Darevskia rudis (Bedriaga, 1886) (Squamata, Lacertidae): a Morpho-Phylogenetic integrated study in a complex Anatolian scenario, pp. 1-68 in Zootaxa 5224 (1) on pages 55-56, DOI: 10.11646/zootaxa.5224.1.1, http://zenodo.org/record/7517907

Darevskia rudis lantzicyreni comb. nov. 53(M), 42(F) 1. ZDEU 105 /2015. (N=3), Balıklı Village, Kelkit, Gümüşhane, Turkey, 05.05.2015, Leg. K. CANDAN, S. GÜL [Map ID: 27]. 2. ZDEU 115 /2015. (N=4), Firdevs Hatun Türbesi, Şiran, Gümüşhane, Turkey, 05.05.2015, Leg. K. CANDAN, S. GÜL [Map ID: 26]. 3. ZDEU 109 /2015. (N=3), Mahmatlı village, Kelkit, Gümüşhane, Turkey, 05.05.2015, Leg. K. CANDAN, S. GÜL [Map ID: 28]. 4. ZDEU 107 /2015. (N=8), Yukarı Kulaca Village, Şiran, Gümüşhane, Turkey, 06.05.2015, Leg. K. CANDAN, S. GÜL [Map ID: 25]. 5. ZDEU 103 /2015. (N=11), Kırkpınar Village, Bayburt, Turkey, 30.07.2015, Leg. K. CANDAN, E. YILDIRIM CAYNAK [Map ID: 32]. 6. ZDEU 110 /2015. (N=1), Bayburt, Turkey, 25.05.2015, Leg. K. CANDAN, E. YILDIRIM CAYNAK [Map ID: 33]. 7. ZDEU 109 /2011. (N=2), Akçainiş, Sivas, Turkey, 09.06.2011, Leg. Y. KUMLUTAŞ [Map ID: 14]. 8. ZDEU 108 /2011. (N=6), Yaylacık, İmranlı, Sivas, Turkey, 10.06.2011, Leg. Y. KUMLUTAŞ [Map ID: 22]. 9. ZDEU 223 /2016. (N=19), Erciyes Mountain, Kayser, Turkey, 20.06.2016, Leg. K. CANDAN, M.K. ŞAHİN, N. BEŞER [Map ID: 13]. 10. ZDEU 21 /2017. (N=6), Otlukbeli Lake, Erzincan, Turkey, 18.05.2017, Leg. K. CANDAN, S. GÜL [Map ID: 31]. 11. ZDEU 58 /2014. (N=1), Doğanşar, Sivas, Turkey, 19.06.2014, Leg. Y. KUMLUTAŞ [Map ID: 17]. 12. ZDEU 68 /2014. (N=1), Armutçayırı Village, Zara, Sivas, Turkey, 19.06.2014, Leg. Y. KUMLUTAŞ [Map ID: 18]. 13. ZDEU 3 /2011. (N=1), Çilhoroz Village, Çayırlı, Erzincan, Turkey, 28.06.2011, Leg. Y. KUMLUTAŞ [Map ID: 29]. 14. ZDEU 197 /2014. (N=10), Çamur Village, Kelkit, Gümüşhane, Turkey, 13.07.2014, Leg. Y. KUMLUTAŞ [Map ID: 30]. 15. ZDEU 9 /2017. (N=4), Kümbet Village, Zara, Sivas, Turkey, 18.07.2017, Leg. Y. KUMLUTAŞ [Map ID: 19]. 16. ZDEU 69 /2011. (N=1), Sucak, Zara, Sivas, Turkey, 10.06.2011, Leg. Y. KUMLUTAŞ [Map ID: 20]. 17. ZDEU 114 /2013. (N=2), Yukarıboğaz, İmranlı, Sivas, Turkey, 20.06.2013, Leg. Y. KUMLUTAŞ [Map ID: 23]. 18. ZDEU 78 /2014. (N=2), Karalar Village, Suşehri, Sivas, Turkey, 11.07.2014, Leg. Y. KUMLUTAŞ [Map ID: 21]. 19. ZDEU 68 /2016. (N=10), Gemecik Village, Refahiye, Erzincan, Turkey, 25.06.2016, Leg. K. CANDAN [Map ID: 24]., Published as part of Arribas, Oscar, Candan, Kamil, Kornilios, Panagiotis, Ayaz, Dinçer, Kumlutaş, Yusuf, Gül, Serkan, Yilmaz, Can, Caynak, Elif Yildirim & Ilgaz, Çetin, 2022, Revising the taxonomy of Darevskia valentini (Boettger, 1892) and Darevskia rudis (Bedriaga, 1886) (Squamata, Lacertidae): a Morpho-Phylogenetic integrated study in a complex Anatolian scenario, pp. 1-68 in Zootaxa 5224 (1) on page 54, DOI: 10.11646/zootaxa.5224.1.1, http://zenodo.org/record/7517907

Darevskia spitzenbergerae wernermayeri ssp. nov. (Appendix 5; Fig. 12a). Synonymy/Chresonymy: Lacerta valentini lantzicyreni; Eiselt, Darevsky & Schmidtler, 1992. (from “Yukarı Narlıca”-sic.!-). Darevskia valentini “Clade A”; Candan et al. (2021). (same locality as this study) ZooBank registration (http://zoobank.org): urn:lsid:zoobank.org:act: 927894FD-EEFD-450A-81D9-0A68188EDC3B. Holotype: ZDEU123 /2015 (n.3). ♁, Yukarınarlıca Village, Çatak, Van, Turkey. leg. Yusuf Kumlutaş, Çetin Ilgaz, 29.07.2015. Paratypes: 7 ♁♁, 10 ♀♀. Same locality, date and collectors as holotype. Derivatio nominis: The specific epithet refers to Dr. Werner Mayer (1943-2015), for his remarkable work on the knowledge of lacertid genera relationships and species taxonomy. Diagnosis: Darevskia spitzenbergerae wernermayeri ssp. nov. differs from nominate form (D. s. spitzenbergerae) in having a higher number of supraciliar granules (18-31 vs. 14-23), supratemporal (4-6 vs. 3-6), ventrals (29-31 vs. 26-30) (females), preanals (1-3 vs. 1-3), tibial scales (16-21 vs. 15-19), temporal scales 1 (5-7 vs. 2-6) and temporal scales 2 (4-6 vs. 2-6). Darevskia s. wernermayeri ssp. nov. has a relatively smaller head relative length (0.18- 0.21 vs. 0.19-0.24). It also differs by a characteristic color pattern of the body. The main osteological diagnostic characters that differ from the nominate form can be specified as follows: The higher number of maxillary (19-21 vs. 16-18) and dentary teeth (23-24 vs. 20-23). Postorbital greater or equal than postfrontal (greater, rarely equal in D. s. spitzenbergerae). Description of holotype: An adult male. Tail regenerated (see Appendix 5c). Fixed with ethanol and conserved in 96% ethanol. Scalation: Rostral not in contact with the nostril. Single postnasal on each side. Width of frontonasal (internasal) plate subequal to length, not in contact with rostral. Sutures between prefrontal plates and frontal plate straight. Parietal plates in contact with postorbital plates on each side. Supraciliar granules 13 and 12, interrupted series on left, continuous on right. Supraciliar plates 6 on each side. Supralabial and sublabial plates 4 and 6 on each side, respectively. Plates in supratemporal region 6 on left, 5 on right side. The first supratemporal plate large narrows towards the back, and ends bluntly. Masseteric large, in one piece on left and two pieces on right, separated from the first supratemporal plate by three longitudinal scales on each side. Tympanic obvious, separated from masseteric by three and two rows of small scales on left and right, respectively. Nine flat collaria. Gularia 33. Ventralia contains 6 longitudinal and 28 transverse rows of plates. Preanal scale singular, surrounded by 6 rows of plates. Femoral pores 20 on each side. Subdigital lamellae 28 on each side. Tibialia 20. Dorsalia 51. Biometry: SVL 63.57 mm. Pileus width 6.90 mm, pileus length 12.96 mm, head width 8.26 mm, head length 13.74 mm. Length of forelimb 22.02 mm, length of hindlimb 31.37 mm. Anal plate width and length are 4.12 mm and 1.84 mm, respectively. Coloration and pattern (in alcohol): Ground color of dorsum greenish-brown, with two irregular paravertebral rows of dark spots. Similar dark spotting is present on each side of the body, that show the reminiscent of a reticulate pattern reduced to only isolated irregular spots. Between these two areas of dark spots, located both in the middle of the dorsum and flank, a paler and spot-free double strip area extends from nape to base of the tail (see photos in Appendix 5). There are few pale spots on dark bandings on the flanks, and more on the forelimb. A few spots near the basis of the forelimb are bluish. The belly, along with the head and neck, is whitish (Appendix 5). The background color of head plates is brownish, with a few scattered and small black spots. The first longitudinal row of ventral plates has dark spots on each side. Variations of paratypes: Descriptive statistics and variation range of the morphometric and scalation characters are given in Table 8. Frontonasal rarely contacts with rostral. The suture between the prefrontal and frontal is usually slanted. The Parietal is in contact with the postorbital in general. Supraciliar granules sometimes are double rows. Tight scales are feeble-keeled. In four specimens, transverse dark spots on ground color are combined with spots on each side, do not form two separate rows. In seven samples, paler spots on the forelimb base are not blue. Belly, along with throat and neck, whitish in seven specimens. In three samples, no dotting on head plates. In fifteen samples, the first longitudinal row of ventral plates contains bluish spots on each side. Distribution: Around Narlıca Valley, Çatak (Van) in the south of Van Lake, Turkey. Habitat: Subalpine-like vegetation of Irano-Turanian Region, on rocky and stony areas, 2363 m. No other reptile species could be identified in the area during study.

Ecological pressure is the major driver of morphological adaptation. Different habitat preferences even among closely related species, often result in the evolution of different body shapes. In the present study, we employed geometric morphometric and principal component analyses (PCA) to compare body shape and head plate morphology among seven species in the genus Darevskia Arribas, 1999 from the Elburz Mountains, Iran that occur in either rocky or forested habitats. The geometric morphometric analysis and the PCA of meristic characters recovered a wide degree of overlap between the rock and forest dwelling species. The PCA of the morphometric characters showed wide separation among the rock and forest dwelling species as well as among some of the rock dwelling species. These results strongly suggest that body shape is correlated with the habitat type whereas head plate morphology and scale meristics are not. Furthermore, the results suggest that the rock dwelling species may be occupying and navigating their microhabitat in different ways. Ecological observations are needed to test this hypothesis.

Poromera fordii (Hallowell, 1857) Figure 15B Tachydromus fordii Hallowell, 1857: 48. Type locality: “Gaboon”. Gabon. Poromera fordii (Hallowell, 1857): Boulenger 1887: 6. Distribution. It occurs in Central Africa, from Cameroon to Central African Republic, southward to the Democratic Republic of the Congo. In Equatorial Guinea it has been recorded in Bioko at Bonyoma (at 450 m a.s.l.) and Parador (North of Musola) (Mertens 1968), and in Río Muni at Cabo San Juan (Boulenger 1905), Benito River (Boulenger 1900, 1921), and Monte Alén National Park (Lasso et al. 2002) (Map 17C). Comments. Photographic records by Twan Leenders from Los Altos de Nsork National Park are available in “https://calphotos.berkeley.edu/”, likely belonging to one of the specimens catalogued as “YPM HERR 014402” and “YPM HERR 014402” held in the Yale University Peabody Museum. Specimens examined. One specimen. Río Muni: Cabo San Juan, likely part of the material examined by Boulenger (1905) (MNCN 7983)., Published as part of Sánchez-Vialas, Alberto, Calvo-Revuelta, Marta & Riva, Ignacio De La, 2022, Synopsis of the terrestrial Reptiles of Equatorial Guinea, pp. 1-197 in Zootaxa 5202 (1) on page 64, DOI: 10.11646/zootaxa.5202.1.1, http://zenodo.org/record/7285600, {"references":["Hallowell, E. (1857) Notice of a collection of reptiles from the Gaboon country, West Africa, recently presented to the Academy of Natural Sciences of Philadelphia, by Dr. Henry A. Ford. Proceedings of the Academy of Natural Sciences of Philadelphia, 9 (3), 48 - 72.","Boulenger, G. A. (1887) Catalogue of the Lizards in the British Museum (Natural History), Vol. 3. The Trustees of the British Museum, London, 575 pp., 40 pls.","Mertens, R. (1968) Zur Kenntnis der Herpetofauna von Kamerun und Fernando Poo. Bonner Zoologische Beitrage, 19, 69 - 84.","Boulenger, G. A. (1905) Reptiles de la Guinee Espagnole. Memorias de la Sociedad Espanola de Historia Natural, 1 (8), 183 - 186.","Boulenger, G. A. (1900) A list of the batrachians and reptiles of the Gaboon (French Congo), with descriptions of new genera and species. Proceedings of the Zoological Society of London, 1900, 433 - 456.","Boulenger, G. A. (1921) Monograph of the Lacertidae. Vol. 2. Trustees of the British Museum, London, viii + 451 pp. https: // doi. org / 10.5962 / bhl. title. 54022","Lasso, C. A., Rial, A. I., Castroviejo, J. & De la Riva, I. (2002) Herpetofauna del Parque Nacional de Monte Alen (Rio Muni, Guinea Ecuatorial). Graellsia, 58, 21 - 34. https: // doi. org / 10.3989 / graellsia. 2002. v 58. i 2.276"]}

Gastropholis echinata (Cope, 1862) Figure 14C Lacerta (Zootoca) echinata Cope, 1862: 189. Type locality: “West Africa”. Very likely Liberia according to Loveridge (1941). Lacerta hirticauda Vaillant, 1884: 344. Type locality: “Kinjabo” Kinjaho, Assini, Ghana. Lacerta langi Schmidt, 1919: 492. Type locality: “Medje”, Democratic Republic of the Congo. Centromastix echinatus (Cope, 1862): Laurent 1958: 118. Gastropholis echinata (Cope, 1862): Arnold 1989: 543. Distribution. West and Central Africa, from Liberia to the Northeast of the Democratic Republic of the Congo. In Equatorial Guinea it has been recorded in Río Muni at Benito River (Boulenger 1921) and Bioko (Pérez del Val 2001) (but see comments) (Map 17A). Comments. Mertens (1964) did not include this species among the reptiles of Bioko. Later, Pérez del Val (2001) recorded a single specimen from the island (MNCN 6231). However, the collection locality of this specimen is tentative, and there are no recent confirmed records of this species in Bioko. In this sense, following Mertens (1964), we do not consider the presence of this species in Bioko. As this species inhabits the forest canopy, it is rarely observed and collected (Arnold 1989). Future prospections should be made to detect the presence (and confirm the Biokoan record) of this elusive species in Equatorial Guinea. Specimens examined. One specimen. Equatorial Guinea, without specific locality (Bioko?) (MNCN 6231)., Published as part of Sánchez-Vialas, Alberto, Calvo-Revuelta, Marta & Riva, Ignacio De La, 2022, Synopsis of the terrestrial Reptiles of Equatorial Guinea, pp. 1-197 in Zootaxa 5202 (1) on pages 61-64, DOI: 10.11646/zootaxa.5202.1.1, http://zenodo.org/record/7285600, {"references":["Cope, E. D. (1862) On Lacerta echinata and Tiliqua dura. Proceedings of the Academy of Natural Sciences of Philadelphia, 1862, 189 - 191.","Loveridge, A. (1941) Report on the Smithsonian-Firestone Expedition's collection of reptiles and amphibians from Liberia. Proceedings of the United States National Museum, 91 (3128), 113 - 140. https: // doi. org / 10.5479 / si. 00963801.91 - 3128.113","Vaillant, M. L. (1884) Catalogue raisonne des reptiles et batraciens d'Assinie donnes par M. Chaper au Museum d'Histoire Naturelle. Bulletin de la Societe Zoologique de France, 9, 343 - 354.","Schmidt, K. P. (1919) Contributions to the herpetology of the Belgian Congo based on the collection of the American Congo Expedition, 1909 - 1915. Part I. Turtles, crocodiles, lizards and chameleons; with field notes by Herbert Lang and James P. Chapin. Bulletin of the American Museum of Natural History, 39, 385 - 624.","Laurent, R. F. (1958) Notes herpetologiques africaines II. Revue de Zoologie et de Botanique Africaines, 58 (1 - 2), 115 - 128.","Arnold, E. N. (1989) Systematics and adaptive radiation of Equatorial African lizards assigned to the genera Adolfus, Bedriagaia, Gastropholis, Holaspis and Lacerta (Reptilia: Lacertidae). Journal of Natural History, London, 23, 525 - 555. https: // doi. org / 10.1080 / 00222938900770311","Boulenger, G. A. (1921) Monograph of the Lacertidae. Vol. 2. Trustees of the British Museum, London, viii + 451 pp. https: // doi. org / 10.5962 / bhl. title. 54022","Perez del Val, J. (2001) Catalogo de las Colecciones Zoologicas de Guinea Ecuatorial del Museo Nacional de Ciencias Naturales, II. Vertebrados. Manuales Tecnicos de Museologia 11, MNCN-CSIC, Madrid, 90 pp."]}

As the only viviparous reptile in China that has both temperature-dependent sex determination (TSD) and genetic-dependent sex determination (GSD) mechanisms, Eremias multiocellata is considered as an ideal species for studying the sex determination mechanism in viviparous lizards. However, studies on embryonic stage of viviparous lizards and morphological characteristics of each stage are limited. In the present study, the embryonic development process of E. multiocellata is divided into 15 stages (stages 28–42) according to the morphology of embryos. Embryos sizes are measured and continuous dynamic variation of some key features, including limbs, genitals, eyes, pigments, and brain scales are color imaged by a stereoscopic microscope. Furthermore, based on these morphological characteristics, we compare the similarities and differences in the embryonic development of E. multiocellata with other squamate species. Our results not only identified the staging table of E. multiocellata with continuous changes of external morphological characteristics but also developed a staging scheme for an important model species that provides a necessary foundation for study of sex determination in a viviparous lizard.

Abstract The Lindholm rock lizard, Darevskia lindholmi, is the only member of the genus Darevskia whose range is restricted solely to Europe, representing a local endemism found only in the Crimean Mountains. In our study, we investigated the cytochrome b gene (mtDNA) of 101 D. lindholmi sequences from 65 Crimean localities, representing its entire range. We found that D. lindholmi is highly genetically structured, and its range is divided into populations belonging to three mitochondrial lineages. The Lindholm rock lizard populations inhabiting the middle part of the Crimean Mountains (further referred to as the Central lineage) are sharply differentiated from the other two lineages (the Common and the Southwestern lineages), which are present in most of the species range. The genetic distance between the Central lineage and the other two taken together is 4.6%, according to our results, suggesting that the divergence occurred during the Early Pleistocene. The narrowly distributed Southwestern lineage and the widespread Common lineage, on the other hand, are differentiated by 1%. Field observations on the representatives of the main evolutionary groups show that their ecology is also different: the Central lineage is a mesophilic and cold-resistant form, while the other two closely related lineages are more xerophilic and thermophilic. Results of the potential ranges modeling and ecological niche analysis confirm that the genetic lineages occupy different niches of the Crimea. Furthermore, the area of inhabitation of the Central lineage splits the western and eastern parts of the Common lineage range, while the Southwestern lineage is restricted along the coast of the southwestern coast of the peninsula. The long-term co-existence of deeply divergent sister mitochondrial lineages in a relatively small (circa 7,000 km2) isolated mountain system serves as a mesocosm for understanding the speciation process. Our data suggest that the Central lineage warrants further taxonomic investigation.

Acanthodactylus boskianus is a widespread species in Northern Africa and Western Asia. In this study, we used morphological and genetic approaches to study populations of A. boskianus from the Zagros Mountains in western Iran, the easternmost limit of the species’ distribution. Our morphological and genetic data indicate that populations of A. boskianus in Iran are distinct from other populations of A. boskianus. Therefore, we describe the Iranian populations as Acanthodactylus zagrosicus sp. nov. The new species is the third endemic species of Acanthodactylus in Iran and the ninth Acanthodactylus species distributed in Iran overall. According to our surveys, this species is distributed widely in the Zagros Mountains including Kermanshah, Lorestan, Ilam, and Khuzestan Provinces.

Small islets in the Mediterranean Sea are often home to reptiles, typically representing an impoverished sample of the continental fauna, yet with high population densities and signs of rapid morphological and behavioral evolution. In this paper, we present the first herpetofaunal survey of several small islet clusters in close proximity to the Mediterranean coast of Israel, only recently geologically separated from the mainland. We performed surveys of five islets during March of 2017 – 2018 and recorded the presence of five different species of reptiles on four of the surveyed islets. Species richness varied between 1 and 4 species, and appeared to be correlated with island area, with a distinct nested structure. Reptile species may have colonized the islets by natural dispersal from nearby coastal populations, or by hitch-hiking on fishing boats and similar methods of human-assisted dispersal. Alternatively, the recorded reptiles may represent relictual populations from earlier geologic periods, when lower sea-levels supported continuous land-bridges between the islets and the mainland. These insular reptile populations require further study to establish the exact means of colonization and describe if and how they differ from mainland populations. We stress the importance of such small Mediterranean islets such as these as centers of unique biodiversity and encourage future study and conservation action aimed at them and similar islets.

Body size evolution on islands is widely studied and hotly debated. Gigantism and dwarfism are thought to evolve under strong natural selection, especially on small remote islands. We report a curious co-occurrence of both dwarf and giant lizards on the same small, remote island (Plakida): the largest Podarcis erhardii (Lacertidae) and smallest Mediodactylus kotschyi sensu lato; Gekkonidae — the two commonest insular reptiles in the Aegean Sea. The geckos of Plakida have a peculiar tail-waving behavior, documented here for the first time in this genus. We suspect that P. erhardii evolved large size to consume geckos and the geckos evolved a unique tail-waving behavior as a defensive mechanism.

The aim of this study is to detect areas of endemism of lizards in Iran. This is the first study of its kind focusing on this subject. Areas of endemism for two families of lizards (Lacertidae and Gekkonidae) that have the highest number of endemic species than other lizard families in Iran were identified by Parsimony Analysis of Endemicity (PAE). Distribution data were collected from previous studies on the Iranian lizards and also from the recent literature on the descriptions of new endemic species. A total of 81 species of lizards were available for analysis. The study area was divided into a 2° × 2° grid of 63 Operational Geographic Units (OGUs). PAE was applied to the data matrix to detect areas of endemism and detected eight areas of endemism in southwestern Iran and near the Persian Gulf. Southern Iran is the main region that most species from Arabia came into Iran and were stopped in their dispersal. Two grids in southern and northeastern Iran were recognized as the areas with the highest density of species in the studied families. Lacertidae and Gekkonidae did not have a shared endemic species in the region (cells 59 and 60) but the area of endemism identified by PAE in the Persian Gulf region suggests that exchange between Iranian and Arabian herpetofauna was very high during interglacial periods. The distribution pattern of the endemic species of these families is concentrated in the region of the Alborz and the Zagros Mountains, but the single area of endemism in southern Iran has an important role in the historical biogeography of the Iranian herpetofauna. During interglacial periods, the Persian Gulf acted as a corridor between the herpetofauna of the two sides and this suggests the importance of this area of endemism for the Gekkonidae family. Also, the OGUs with the highest density of species are located around the country and, the lowest density is in the Central Plateau. PAE detected eight areas of endemism in southwestern Iran, but according to the number of species per units, two OGUs can be identified as high density in northeastern and southern Iran.

Identification key to the Pakistani species of the genus Eremias (modified from Masroor et al. 2020a) 1. Subocular bordering mouth............................................................................. 2 - Subocular not bordering mouth................................................................ E. acutirostris 2. A complete row of lateral scales of the 4 th toe forming a distinct fringe or comb on its entire length..................... 3 - Lateral scales of 4 th toe not forming distinct fringe........................................................... 4 3. Row of femoral pores reaches well short of the knee; the median dark dorsal stripes interrupted and form reticulate pattern.............................................................................................. E. scripta - Row of femoral pores reaches to knee; dorsal stripes without any sign of vermiculation................... E. cholistanica 4. Back with 5–11 dark stripes, broader than interspaces, none of the stripes containing light ocelli or spots; stripes persistent in adults, but sometimes indistinct so that back appears almost uniform sandy; usually only single median collar scale distinctly larger than adjacent gulars.............................................................................. 5 - Dark stripes on the dorsum of juvenile breaking up in adults to form spots or broken lines; usually, several collar scales distinctly larger than adjacent gulars...................................................................... 6 5. 4 th toe with two complete rows of subdigital scales and a complete row of sharply pointed lateral scales, i.e., a total of 4 scales counted around penultimate phalanx.......................................................... Eremias fasciata - 4 th toe with one complete row of subdigital scales and a complete row of lateral scales, i.e., total of three scales counted around penultimate phalanx............................................................................. E. kakari 6. Adults with four more or less regular rows of disconnected dark spots on dorsum between dorsolateral broader dark stripes, the latter with white ocelli at the edges and within each stripe; infranasal in contact with the rostral........... E. rafiqi sp. nov. - Adults with seven light stripes on the neck, transforming into disconnected series of white ocelli edged with black; no dorsolateral dark stripes, an outer-most series of white and black ocelli starts behind the eyes on each side, onto tympanum and flanks above the forelimb and hindlimb insertion; infranasal not in contact with the rostral................... E. killasaifullahi sp. nov., Published as part of Masroor, Rafaqat, Khan, Muazzam Ali, Nadeem, Muhammad Sajid, Amir, Shabir Ali, Khisroon, Muhammad & Jablonski, Daniel, 2022, Appearances often deceive in racerunners: integrative approach reveals two new species of Eremias (Squamata: Lacertidae) from Pakistan, pp. 55-87 in Zootaxa 5175 (1) on page 78, DOI: 10.11646/zootaxa.5175.1.3, http://zenodo.org/record/7003222, {"references":["Masroor, R., Khisroon, M., Khan, M. A. & Jablonski, D. (2020 a) A new species of Microgecko Nikolsky, 1907 (Squamata: Gekkonidae) from Pakistan. Zootaxa, 4780 (1), 147 - 164. https: // doi. org / 10.11646 / zootaxa. 4780.1.7"]}

Eremias rafiqi sp. nov. (Table 2, Figs. 4, 5, 6) Suggested vernacular name: Rafiq’s Racerunner Pashto name: Holotype. PMNH 856 (cyt b: n/a; Rag1: n/a), an adult male, collected from Tanishpa village, Torghar Mountains, Killa Saifullah district, Balochistan (31.1869º N, 68.4126º E; Fig. 1), elevation 2,506 m a. s. l., May 25, 1997, leg. Khalid Javed Baig (Fig. 4). Paratypes. Males: PMNH 855 (cyt b: n/a; Rag 1: n/a), 857 (cyt b: n/a; Rag 1: n/a), PMNH 859 (cyt b: n/a; Rag 1: n/a), PMNH 861 (cyt b: n/a; Rag 1: n/a), PMNH 4056 (cyt b: n/a; Rag 1: MT 554487). Females : PMNH 3724 (cyt b: n/a; Rag 1: MT 554476), PMNH 3735 (cyt b: MT 554461; Rag 1: MT 554477), PMNH 4058 (cyt b: n/a; Rag 1: n/a). Juveniles : PMNH 837 (cyt b: n/a; Rag 1: n/a), PMNH 3723 (cyt b: MT 554470; Rag 1: MT 554496), PMNH 4053 (cyt b: MT 554457; Rag 1: MT 554485), PMNH 4054 (cyt b: MT 554454; Rag 1: MT 554480). PMNH 857, collected along with the holotype; PMNH 855 and 861, May 26, 1997, Ashewat, Qamar Din Karez, Zhob district (31.3448ºN, 68.6307ºE), leg. Khalid Javed Baig; PMNH 859, May 24,1997, Tanishpa village, Torghar, Killa Saifullah district, leg. Khalid Javed Baig; PMNH 3723–24, October 09, 2017, Khar, Nushki district, Balochistan (29.5879ºN, 65.6609ºE), leg. Muazzam Ali Khan; PMNH 3735, October 21, 2017, Khar, Nushki district, leg. Muazzam Ali Khan; PMNH 4053, September 05, 2018, Zamkai Nala, Tanishpa village, Torghar, Killa Saifullah district (31.1930ºN, 68.4111ºE), leg. Rafaqat Masroor; PMNH 4054, September 01, 2018, Ashewat, Qamar Din Karez, Zhob district, leg. Rafaqat Masroor; PMNH 4056 and 4058, August 30, 2018, Kunder, Torghar, Killa Saifullah district, leg. Ibad ur Rehman (Figs. 5 & 6). Morphological diagnosis. A large-sized lacertid lizard, maximum snout-vent length (SVL) = 99.3 mm, tail 1.67 to 1.89 times longer than body length (SVL), hindlimbs relatively long (HLL / SVL ratio 0.6–0.8); subocular scale reaching to the edge of the mouth, 5–7 (mainly 6, rarely 5) anterior to subocular; dorsals 56–67; ventrals in 14–17 oblique longitudinal series; frontal separated from supraoculars; the height of the first two to three transverse rows of ventral scales in the pectoral region more than its breadth; 17–21 femoral pores on each side, separated medially by 1–4 scales (mainly 3, rarely 1), the space between the femoral pores less than one–fourth length of each row; toes without fringe, encircled by three scales in a single series of 22–27 unicarinate and bicarinate scales underneath; the tip of the fourth toe reaches to the forelimb and extends to just behind the collar. The adult specimens are grayish in life with four series of longitudinal black ocelli on the dorsum originating from behind the parietals and extending onto the tail; on each lateral side, a broader dark stripe originates from behind the eye and continues onto the tail with disconnected white round ocelli at the margins as well as white ocelli inside the stripe. Molecular data. Eremias rafiqi sp. nov. represents the so-called Zabol clade sensu Rastegar-Pouyani et al. (2010) from eastern Iran and the clade E identified by Khan et al. (2021) from northeastern Balochistan, Pakistan. Whereas Rastegar-Pouyani et al. (2010) identified this clade solely on mtDNA (cyt b, 12S), Khan et al. (2021) used mitochondrial (16S, COI, cyt b) as well as nuclear data (Rag 1) that clearly showed deep differentiation of this new species from other available sequences of the genus. The distinction of Eremias rafiqi sp. nov. is supported by its phylogenetic position (monophyletic clade among other species of E. persica complex, sister to E. fahimii but with weak statistical support; Fig. 1), the differentiation on solely Rag1 dataset (Fig. 2), and the value of the uncorrected p distances reaching from 8.5% (E. fahimii) to 20.7% (E. strauchi) (Table 3). The intraclade genetic diversity (cyt b) is 2 % with six detected haplotypes found in Iran and Pakistan (Fig. 2, Table 3). Etymology. The species epithet “ rafiqi ” is taken from the first name of late Rafique Ahmed Rajput (1968– 2008) to whom the new species is dedicated. The deceased Rajput served in Sindh Wildlife Department from 1986 till his demise. With no formal education in wildlife research, conservation and management, his passion for the conservation of wildlife in Pakistan remain unparalleled. From collecting the first-ever data of Ursus arctos isabellinus in the high-altitude Deosai Plateau, Gilgit-Baltistan, to the faunistic studies in the Indus River Delta and desert areas, he was a symbol of hard work. He also has very sound techniques for the collection of lizards and snakes. Unfortunately, during one such endeavor for gathering faunistic data, he collected a juvenile venomous krait, Bungarus sp. (possibly B. persicus) from Jiwani town, Gwadar District, Balochistan and mistaken its identity with non-venomous Lycodon species. On the morning of October 13, 2008, when he was shifting the live snake from a container to permanently preserve it for research purposes, the snake bit him multiple times.After a couple of hours, Rajput felt severe pain, anxiety and dizziness. He was immediately taken to the hospital in Karachi, where the doctors told his relatives that anti-snake venom is not available and asked them to get it from the pharmacy market. By the time the anti-venom was arranged, Rajput breathed his last. Description of the holotype. SVL: 99.3, TL: 168.0, HL: 26.9, HW: 14.5, HH: 15.5, TrL: 42.5, HLL: 66.8, FLL: 37.3, FrL: 6.3, FrW: 3.4.An adult male preserved in formalin in a good state of preservation (Fig. 4); head and body moderately depressed; tail long, ca. 1.7 times longer than the body, cylindrical and depressed at the base. Head relativelylong (HL/SVL, 0.27) (Fig. 4), ca. 1.7 times longer than wide (HW/HL, 0.58), head height slightly less than head width (HH/HW, 0.89). Limbs strong, hindlimbs ca. 1.8 times longer than the length of forelimbs (FLL/HLL, 0.56), hindlimbs comprise 1.4 times of the body length (HLL/SVL, 0.67). Head slightly broader than the neck.Head shields including nasals,frontonasal,prefrontals,frontal,frontoparietals, interparietal and parietals are smooth and convex. Nasals moderately swollen, three nasals, the lower in contact with two supralabials on both right and left side and in contact with the rostral (Fig. 4D). Supranasals in contact with rostral but lack such contact with first supralabial, the suture between them is 3.7 times the length of frontonasal, whose breadth is slightly more than its length; length of prefrontals 1.6 times its width, forming a median suture; length of frontal ca. 1.8 times as long as broad, its length slightly less than its distance from the tip of the snout, narrow behind; parietals smooth, slightly longer than its width; interparietal smooth, more than half the length of frontoparietals, about equal to the suture of frontoparietal; no occipital. Two large supraoculars, about equal in size, the space anterior to supraoculars filled by few small and three to five larger granules; both supraoculars in contact with frontal of their sides while separated from supraciliaries by a series of granules (Fig. 4C); six supraciliaries, first longest, its length shorter than its distance from the first loreal. Rostral pentagonal, broader than high, narrower beneath than above; anterior loreal slightly higher than wide, shorter than the second loreal which is longer than high; supralabials 10 on the right side, 9 on the left side; subocular keeled just below the eye, bordering the mouth, wedged between sixth and seventh supralabials on the right side and fifth and sixth supralabials on the left side (Fig. 4D). Temporals smooth, a large scale above ear; auricular denticulation indistinct or three small scales forming slight denticulation anteriorly. Lower eyelid covered with numerous small semi-transparent scales. Six infralabials on the right side, seven on the left side, gradually increasing in size posteriorly. Five pairs of chin shields; anterior three completely in contact, the fourth pair separated by 10 to 11 smaller gulars in a straight line, fifth not in contact with infralabials, separated by a single row of scales. Collar curved, free, serrated and composed of ten plates larger than adjacent gulars, the middle one quite enlarged than others. Gular fold distinct, 32 gular scales in a straight line between the symphysis of the chin shields and the collar (Fig. 4B). Dorsal scales granular, smooth, 62 across the middle of the body. Ventral plates broader than long (except for outermost series), forming oblique longitudinal series of 16 plates across mid-belly and 29 transverse rows counted from behind collar to vent; first three rows of ventral scales in the pectoral region behind collar longer than broad, the first row is twice as long as broad. Precloacal region with a pair of the enlarged median plates just above the vent, surrounded by six large scales. Forelimb ca. 1.4 times longer than the head, the upper surface of the arm with rhombic, smooth scales. Scales on the upper surface of hindlimbs similar to dorsals, equal in size; ventral surface of hindlimbs covered by enlarged plates, the ventral surface of the tibia with one row of very large and one comparatively smaller plates; the tip of the fourth toe reaches to the forelimb and extends to just behind the collar; 21 femoral pores on the right side, most of the left side damaged, the two femoral pore series separated by two scales, length of the interfemoral space not greater than one–fourth length of each row. Toes slender, compressed, with no fringe; subdigital lamellae unicarinate, in a single row of 25 scales under the 4th toe, a total of three scales around the 4 th toe. Upper caudal scales oblique, truncate, strongly and diagonally keeled, 30 scales in the 9 th –10 th annulus behind the postcloacal granules. Coloration in life. The adult specimens are grayish in life with four more or less regular rows of black spots on the light dorsum, originating from behind the parietals, smaller on the nape, larger on the middle of the dorsum, disappearing on proximal one-fourth of the tail. The middle two rows of black spots have white spots along each black spot of the rows. On each lateral side, a broader dorsolateral dark stripe originates from behind the eye and continues onto the tail with disconnected white ocelli at both margins as well as white ocelli inside the stripe; next to the broader dark stripe, a lateral-most stripe is composed of disconnected black ocelli, originating from behind tympanum and reaching to the hindlimb. Upper parts of both hindlimbs and forelimbs with white and black ocelli. Head gray without any markings or spots; labials white with black markings. Belly and underside of tail creamy white, tail dorsum sandy grayish. The juveniles and subadults are nearly similar in coloration to the adults except for the following details; four longitudinal dark stripes on the body, the outermost originate from anterior parietals on the outer side and continue onto the tail, the innermost originate from the posterior of parietals and merge after running a while on the tail. A broader lateral stripe on each side originates from behind the eye and continues on the lateral side of the body and tail with interspersed white ocelli between the forelimb and hindlimb. The dorsal forelimb and hindlimb are dark gray with white ocelli. Variations in paratypes. The paratypes of E. rafiqi sp. nov. agree with the holotype with some differences given in Table 2 (Figs. 5, 6). Besides sex, the specimens differ in the arrangement of supralabials i.e. subocular wedged between 6 th and 7 th in all the type series except PMNH 861 (between 7 th and 8 th) and PMNH 3724 (between 5 th and 6 th). The arrangement of postmentals has a similar pattern in the paratypes except PMNH 855, 861, 837, 3723, 3735 and 4054, where the fifth chin shield is in contact with the infralabials. The contact of postmental shield with the supralabials varies in the paratypes; PMNH 855 and 3724, the fifth postmental shield is in contact with 7 th supralabial; PMNH 837 and 3735, the fifth postmental shield is in contact with sixth supralabial; PMNH 861, the fifth postmental shield is in contact with 7 th and 8 th supralabials. The infranasal scale in PMNH 837, 855, 859, 861, 3723, 3735, 4053–4054 and 4058 rests on first, second and third supralabials. The scale count of dorsals, ventrals, gulars, collars, caudals at 9 th –10 th annuli and lamellae under 4 th toe, however, show a unique value for every specimen within a certain range. Sexual and age dimorphism. Apparently, males attain larger sizes than females in E. rafiqi sp. nov.: male SVL to 99.3 mm, female SVL 82.1 mm. Moreover, males have generally longer hindlimbs and shorter trunks as compared to females. For a larger female having SVL of 98.1 mm (PMNH 4058), the hindlimb is 56.0 mm against a smaller-sized male (PMNH 855, SVL 92.2 mm) which has a hindlimb length of 57.1 mm. Similarly, the trunk length of a smaller female (PMNH 3724, SVL 75.1 mm) is 36.8 mm against a larger male (PMNH 857, SVL 78.5 mm) which has a trunk length of 34.5 mm. The dorsal body color and pattern, however, varies in juveniles and adults of both genders (Figs. 5, 6). Comparison. The new species Eremias rafiqi sp. nov. is strikingly different from species exhibiting striped and ocellate patterns in the subgenus Aspidorhinus (E. kopetdaghica Szczerbak, 1972, E. lalezharica Moravec, 1994, E. papenfussi Mozaffari et al., 2011, Eremias persica Blanford, 1874, E. regeli Bedriaga, 1905, E. fahimii Mozaffari et al., 2020, E. isfahanica Rastegar-Pouyani et al., 2016, E. montana Rastegar-Pouyani & RastegarPouyani, 2001, E. nikolskii Bedriaga, 1905, E. velox Pallas, 1771) and ocellate pattern (E. killasaifullahi sp. nov., E. afghanistanica Böhme & Szczerbak, 1991, E. roborowskii Bedriaga, 1912, E. strauchi Kessler, 1878, E. suphani Başoğlu & Hellmich, 1968). The new species E. rafiqi sp. nov. can also be differentiated from the geographically closely-distributed members of the subgenus Eremias having striped and ocellate pattern (E. aria Anderson & Leviton 1967) and ocellate pattern (E. nigrocellata Nikolsky 1896) by the arrangement of subocular scale which borders the mouth (Supplementary Tab. 1; see published data in Lantz 1928, Szczerbak 1974, Bischoff & Böhme 1980, Böhme & Szczerbak 1991, Anderson 1999). A brief of morphological differences is provided (the material used for a first-hand comparison is listed in parentheses at each species; see also Table 2 and S 1). Besides striped and ocellate body pattern (vs. ocellate), E. rafiqi sp. nov. can be distinguished from E. afghanistanica by its larger size (SVL up to 99.3 mm vs. 67.0 mm), higher count of dorsals (56–67 vs. 44–46), gulars (30–36 vs. 25–28), femoral pores (17–22 vs. 16–18), caudal scales in the 9 th –10 th annulus (24–33 vs. 22–25), 5–7 supralabials (mainly 6, rarely 5) located anterior to subocular (vs. 5) and lower number of ventral scales in a single row from the posterior edge of collar to the vent (29–33 vs. 37–38). From E. persica, that is partly close in dorsal coloration, pattern and size, E. rafiqi sp. nov. differs in the length of interparietal to the length of suture of parietals (longer vs. shorter), length of frontonasal to its width (longer vs. as long as wide), size of the second loreal scale to first loreal scale (more than three times vs. two times), supracaudals (strongly keeled vs. weakly keeled) and tail coloration in the juveniles (sandy grayish vs. bluish). Besides distant distribution, Eremias rafiqi sp. nov. differs from the recently described E. fahimii by its larger size (SVL up to 99.3 mm vs. 56.0 mm), more SDLT 4 th (22–27 vs. 20–21), the greater number of scales separating the femoral pores (1–4 vs. 1) and the dorsal color and pattern in adults (dorsal stripes broken into ocelli vs. dorsal stripes persistent throughout life). From E. isfahanica, E. rafiqi sp. nov. differs in the following morphological characters apart from its distant distribution: higher count of supralabials (8–10 vs. 6–8), 5–7 (mainly 6, rarely 5) of them located anterior to subocular (vs. 5), lower count of collars (8–12 vs. 12–15) and the dorsal color pattern in adults (dorsal stripes broken into ocelli vs. dorsal stripes persistent throughout life). Eremias rafiqi sp. nov. differs from E. kopetdaghica in having a higher count of dorsals (56–67 vs. 48–59), gulars (30–36 vs. 19–28), caudal scales in the 9 th –10 th annulus (24–33 vs. 20–26) and collars (8–12 vs. 7) and the dorsal color and pattern in adults (presence of a broader dark stripe on each lateral side above flanks with disconnected white ocelli at the margins as well as white ocelli inside the stripe vs. no such lateral broader stripes). Eremias rafiqi sp. nov. can be distinguished from E. lalezharica in having a higher count of dorsals (56–67 vs. 54–59), femoral pores (17–22 vs. 15–19), pair of chin shields/ submaxillary shields (5 vs. 4), lower number of collars (8–12 vs. 13–15), contact of gulars with second pair of submaxillary shields (none vs. 1-2 rows of gulars with the second pair of submaxillary shields) and dorsal color and pattern. Apart from its peculiar distribution in the remote valley in Torghar Mountains, a part of the Palearctic region, E. rafiqi sp. nov. can be differentiated from E. montana in the following set of characters: larger size (SVL up to 99.3 mm vs. 58.5 mm), higher count of ventral scales in a row across mid-belly in the widest part (14–17 vs. 13–14), number of ventral scales in a single row from the posterior edge of collar to the vent (29–33 vs. 27–28), gulars (30–36 vs. 23–25), infralabials (6–10 vs. 4–6), number of supralabials anterior to the subocular (5–7 vs. 4–5), generally more SDLT 4 th (22–27 vs. 18–25), generally higher count of scales separating the femoral pores (1–4 vs. 2) and dorsal color and pattern. From E. nigrocellata, E. rafiqi sp. nov. differs in dorsal body pattern (striped and ocellate vs. ocellate), higher count of dorsals (56–67 vs. 42–56), lower number of ventral scales in a row across mid-belly i, Published as part of Masroor, Rafaqat, Khan, Muazzam Ali, Nadeem, Muhammad Sajid, Amir, Shabir Ali, Khisroon, Muhammad & Jablonski, Daniel, 2022, Appearances often deceive in racerunners: integrative approach reveals two new species of Eremias (Squamata: Lacertidae) from Pakistan, pp. 55-87 in Zootaxa 5175 (1) on pages 70-75, DOI: 10.11646/zootaxa.5175.1.3, http://zenodo.org/record/7003222, {"references":["Rastegar-Pouyani, E., Rastegar-Pouyani, N., Kazemi-Noureini, S., Joger, U. & Wink, M. (2010) Molecular phylogeny of the Eremias persica complex of the Iranian plateau (Reptilia: Lacertidae), based on mtDNA sequences. Zoological Journal of the Linnaean Society, 158, 641 - 660. https: // doi. org / 10.1111 / j. 1096 - 3642.2009.00553. x","Khan, M. A., Jablonski, D., Nadeem, M. S., Masroor, R., Kehlmaier, C., Spitzweg, C. & Fritz, U. (2021) Molecular phylogeny of Eremias spp. from Pakistan contributes to a better understanding of the diversity of racerunners. Journal of Zoological Systematics and Evolutionary Research, 59, 466 - 483. https: // doi. org / 10.1111 / jzs. 12426","Szczerbak, N. N. (1972) New subspecies of Eremias strauchi - Eremias strauchi kopetdaghica ssp. nova (Sauria, Reptilia) from Turkmenia. Vestnik Zoologii, 6, 83 - 86 [in Russian]","Moravec, J. (1994) A new lizard from Iran, Eremias (Eremias) lalezharica sp. n. (Reptilia: Lacertilia: Lacertidae). Bonner zoologische Beitrage, Bonn, 45, 61 - 66.","Mozaffari, O., Ahmadzadeh, F. & Parham, J. F. (2011) Eremias papenfussi sp. nov., a new lacertid lizard (Sauria: Lacertidae) from Tehran Province, Iran. Zootaxa, 3114 (1), 57 - 62. https: // doi. org / 10.11646 / zootaxa. 3114.1.6","Bedriaga, J. V. (1905) Neue Saurier aus Russisch-Asien. Annu. In: Nikolsky, A. M. (Ed.), Herpetologia rossica. Memoires de l`Academie imperiale des sciences de St. Petersbourg, XVII (1), pp. 1 - 518.","Mozaffari, O., Ahmadzadeh, F. & Saberi-Pirooz, R. (2020) Fahimi's racerunner, a new species of the genus Eremias Fitzinger, 1834 (Sauria: Lacertidae) from Iran. Zootaxa, 4768 (4), 565 - 578. https: // doi. org / 10.11646 / zootaxa. 4768.4.7","Rastegar-Pouyani, E., Yousefkhani Hosseinian, S. S., Soolmaz, R., Kami, H. G., Mehdi, R. & Wink, M. (2016) A new species of the genus Eremias Fitzinger, 1834 (Squamata: Lacertidae) from Central Iran, supported by mtDNA sequences and morphology. Zootaxa, 4132 (2), 207 - 220. https: // doi. org / 10.11646 / zootaxa. 4132.2.2","Bedriaga, J. von (1912) Wissenschaftliche Resultate der von N. M. Przewalski nach Central-Asien unternommenen Reisen, Zoologischer Theil. Band III, Abtheilung 1. Amphibien und Reptilien. Kaiserliche Akademie der Wissenschaften, St. Petersburg, 769 pp. [in Russian and German]","Kessler, K. F. (1878) Transcaucasian voyage. Travaux de la Societe des Naturalistes de St. Petersbourg, 8 (Supplement), 1 - 200. [in Russian]","Basoglu, M. & Hellmich, W. (1968) Eine neue Eremias - Form aus Ost-Anatolien (Reptilia, Lacertidae). Ege Universitesi Fen Fakultesi Ylmi Raporlar Serisi, Bornova-Yzmir, 67, 3 - 7.","Anderson, S. C. & Leviton, A. E. (1967) A new species of Eremias (Reptilia: Lacertidae) from Afghanistan. Occasional Papers of the California Academy of Sciences, 64, 1 - 4.","Nikolsky, A. M. (1896) Diagnoses reptilium et amphibiorum novorum in Persia orientali a N. Zarudny collectorum. Annuaire Musee Zoologique de l'Academie Imperiale des Sciences de St. - Petersbourg, 1, 369 - 372.","Lantz, L. A. (1928) Les Eremias de l'Asie Occidentale. Bulletin du Museum de Georgie, 4, 1 - 136.","Szczerbak, N. N. (1974) Racerunners of the Palaearctic. Naukova Dumka Press, Kiev, Ukraine, 296 pp. [in Russian]","Bischoff, W. & Bohme, W. (1980) Der systematische Status der turkischen Wustenrenner des Subgenus Eremias (Sauria: Lacertidae). Zoologische Beitrage, 26, 297 - 306.","Anderson, S. C. (1999) The Lizards of Iran. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, 442 pp.","Masroor, R., Khisroon, M., Khan, M. A. & Jablonski, D. (2020 b) A new species of Eremias Fitzinger, 1834 (Squamata: Lacertidae) from the arid mountains of Pakistan. Zootaxa, 4786 (1), 101 - 121. https: // doi. org / 10.11646 / zootaxa. 4786.1.8"]}

Eremias killasaifullahi sp. nov. (Table 1, Figs. 3, 5, 6) Suggested vernacular name: Killa Saifullah’s Racerunner Pashto name: Holotype. PMNH 3613 (cyt b: MT 554460; Rag1: MT 554498), an adult male, collected from Kunder, Torghar Mountains, Killa Saifullah district, Balochistan (31.3247º N, 68.5452º E; Fig. 7D), elevation 1,920 m a. s. l., March 23, 2017, leg. Rafaqat Masroor (Fig. 3). Paratypes. Males: PMNH 3614–3616 (cyt b: MT554466, MT554456, n/a; Rag1: MT554478, MT554482, n/a). Females: PMNH 4046 (cyt b: MT 554453; Rag1: MT 554479), PMNH 4050 (cyt b: MT 554473; Rag1: MT 554483), PMNH 4055 (cyt b: MT 554455; Rag1: MT 554481). Juveniles: PMNH 3673 (cyt b: n/a; Rag1: n/a), PMNH 4045 (cyt b: MT 554467; Rag1: MT 554486), PMNH 4052 (cyt b: MT 554459; Rag1: MT 554497). PMNH 3614–16, 3673 collected along with the holotype; PMNH 4045, 4052, September 5, 2018, Zamkai Nala, Tanishpa, Killa Saifullah district, leg. Rafaqat Masroor; PMNH 4046, 4055 August 31, 2018, Ashewat, Qamar Din Karez, Zhob district, leg. Rafaqat Masroor; PMNH 4050, September 1, 2018, Zamkai Nala, Tanishpa, Killa Saifullah district, leg. Rafaqat Masroor (Figs. 5, 6). Morphological diagnosis. A medium-sized lacertid lizard, maximum snout-vent length (SVL) = 70.5 mm, tail 1.67 to 1.97 times longer than body length (SVL), hindlimbs relatively long (HLL / SVL ratio 0.6–0.8); subocular scale reaching to the edge of the mouth, 5–7 (mainly 6, rarely 5) anterior to subocular; dorsals 53–63; ventrals in 14–18 oblique longitudinal series; frontal separated from supraoculars; the height of the first two to three transverse rows of ventral scales in the pectoral region more than its breadth; 17–24 femoral pores on each side, separated medially by 1–5 scales (mainly 2–4, rarely 1), the space between the femoral pores less than one–fourth length of each row; toes without fringe, encircled by three scales in a single series of 21–25 unicarinate and bicarinate scales underneath; tip of the fourth toe reaches to the forelimb and extends to just behind the collar. The adult specimens are creamy beige in life with seven light stripes appearing on the neck which transforms into ocelli and vermiculation behind the neck. No dorsolateral broader dark stripes, an outer-most series of white and black ocelli starts behind the eyes on each side, onto the tympanum and flanks above the forelimb and hindlimb insertion. ......continued on the next page TABLE 1. (Continued) ......continued on the next page TABLE 2. (Continued) ......continued on the next page TABLE 2. (Continued) ......continued on the next page TABLE 2. (Continued) Molecular data. Eremias killasaifullahi sp. nov. represents a newly detected evolutionary lineage (Fig. 1) of the genus Eremias (Aspidorhinus) that was firstly detected by Khan et al. (2021) as the clade F (with subclades F1, F2) based on four studied genetic markers (16S, COI, cyt b, Rag1). This lineage was detected occurring in NE Balochistan in Pakistan and represents local microendemism (Fig. 2). The lineage deeply diverges and is sister to all other lineages of such called E. persica complex (see Khan et al. 2021 and Fig. 1 in this study) and well differentiated in the Rag1 dataset (Fig. 2). The lineage genetically (uncorrected p distances) differs from 14.5% (E. strauchi) to 21.6% (E. velox) (Table 3) among species of the subgenus Aspidorhinus. Its average intraclade genetic variability (cyt b) is 3% (Fig. 2). Despite a very small known range of distribution, the distances between F1 and F2 subclades sensu Khan et al. (2021) reached 4.6% and the haplotype network based on cyt b dataset showed six different haplotypes. High allele diversity was also detected by analyzing the Rag1 marker (Fig. 2). Etymology. We derived the name of the new species from Killa Saifullah (Pashto:; also Qilla Saifullah), a city and district in northwestern Balochistan province, Pakistan that represents the area, from where this newly discovered endemic species of Eremias (subgenus Aspidorhinus) is currently known. The region plays an important role for producing fruits, nuts and vegetables in Pakistan. The discovery of this species of lizards thus highlights the importance of this region from the biodiversity point of view. Description of the holotype. SVL: 65.3, TL: 109.7, HL: 17.4, HW: 10.2, HH: 8.1, TrL: 28.3, HLL: 44.8, FLL: 26.6, FrL: 4.6, FrW: 2.3.An adult male of E. killasaifullahi sp. nov. preserved in ethanol in a good state of preservation (Fig. 3); head and body moderately depressed; tail long, ca. 1.7 times longer than the body, cylindrical and depressed at the base. Head relativelylong (HL/SVL ratio 0.27) (Fig. 3), 1.7 times longer than wide (HW/HL ratio 0.59), head height less than head width (HH/HW, 0.79). Limbs strong, hindlimbs 1.6 times more than the length of forelimbs (FLL/HLL, 0.59), hindlimbs comprise 1.4 times the body length (HLL/SVL, 0.69). Head broader than the neck; nasals, frontonasal, prefrontals, frontal, frontoparietals, interparietal and parietals are smooth and convex. Nasals are moderately swollen, three nasals, the lower in contact with three supralabials on the right and left side, its contact with the rostral lacking (Fig. 3D). Supranasals in contact with rostral and first supralabial, the suture between them is four times the length of frontonasal, whose breadth is ca. 1.1 times its length; length of prefrontals 1.4 times its width, joined by a median suture; frontal two times as long as broad, its length slightly less than its distance from the tip of the snout, narrow behind; parietals smooth, slightly longer than wide; interparietal smooth, more than half of the length of frontoparietals; no occipital. Two large supraoculars, about equal in size, the space anterior to supraoculars filled by few small and three to five larger granules; both supraoculars in contact with frontal of their sides while separated from supraciliaries by a series of granules (Fig. 3C), behind the two large spraoculars a single, comparatively medium-sized, granule exist; six supraciliaries, first longest, its length shorter than its distance from the first loreal. Rostral pentagonal, broader than high, narrower beneath than above; anterior loreal slightly higher than wide, shorter than the second loreal which is longer than high; supralabials 8; subocular keeled just below the eye, bordering the mouth, wedged between fifth and sixth supralabials (Fig. 3D). Temporals smooth, a large scale above ear; auricular denticulation indistinct or three small scales forming slight denticulation anteriorly. Lower eyelid covered with numerous small semi-transparent scales. Six infralabials, gradually increasing in size posteriorly. Five pairs of chin shields; anterior three completely in contact, the fourth one separated by six smaller gulars, the fifth one is in contact with fifth and sixth infralabials on both sides. Collar curved, free, serrated and composed of 10 plates larger than adjacent gulars, the middle one slightly enlarged than others. Gular fold distinct, 20 gular scales in a straight line between the symphysis of the chin shields and the collar (Fig. 3B). Dorsal scales granular, smooth, 60 across the middle of the body. Ventral plates broader than long (except for outermost series), forming oblique longitudinal series of 16 plates across mid-belly and 25 transverse rows counted from behind collar to vent; first three rows of ventral scales in the pectoral region behind collar longer than broad, the first row is twice as long as broad. Precloacal region with an enlarged median plate just above the vent, surrounded by four large scales. Forelimb ca. 1.5 times longer than the head, upper surface of the arm with rhombic, smooth scales. Scales on the upper surface of hindlimbs similar to dorsals, varying in size; ventral surface of hindlimbs covered by enlarged plates, the lower surface of the tibia with one row of very large and one comparatively smaller plates, the tip of the fourth toe reaches to the forelimb and extends to just behind the collar; 17 femoral pores on the right side, the left side damaged, the two series separated by two scales, length of the interfemoral space not greater than one-fourth length of each row. Toes slender, compressed, with no fringe. Subdigital lamellae unicarinate, in a single row of 21 scales under the 4th toe, a total of three scales around the 4 th toe. Upper caudal scales oblique, truncate, strongly and diagonally keeled, 26 scales in the 9 th –10 th annulus behind the postcloacal granules. Coloration in life. The adult specimens (Fig. 7E) are creamy beige with ocellate body pattern. Seven light stripes appear on the neck which transforms into ocelli and vermiculation behind the neck. Of the seven, the lateralmost light stripe originates from behind the eye and runs on the outer edge of the parietal, transforming into a disconnected series of white ocelli edged with black, running up to anterior one-third of the tail. Next to the lateralmost, the paravertebral light stripe originates from behind the parietal and transforms into closely-connected white ocelli edged with black and runs on the tail short of lateral-most ocelli. Next to paravertebral light stripe, there exists a light nuchal stripe on each side and the light vertebral stripe, the three joins behind the neck and transform into white ocelli edged with black in the pattern of vermiculation. In addition to seven light stripes on the neck, an outermost series of white and black ocelli starts behind the eyes on each side, onto the tympanum and flanks above the forelimb and hindlimb insertion. The upper parts of both hindlimbs and forelimbs are provided with white and black ocelli. Head gray with black mottled markings or spots; supralabials white with black markings. Belly and underside of tail creamy white, tail dorsum grayish. The juveniles and subadults (Fig. 6) are nearly similar in coloration to the adults except for the following details; seven longitudinal light stripes on the neck, the lateral-most originate from behind the eye, running on the outer parietals and continuing onto the dorsum in the form of connected small white ocelli, terminating on the one-third of the tail, the paravertebral light stripe originates from the posterior of parietals, and merge short of the lateral-most stripe on the tail, the nuchal of each side and vertebral light stripe merge after the neck to form light vermiculation up to the base of the tail. An additional outer-most light stripe originates from behind the tympanum and is produced in the form of disconnected white ocelli above the insertion of forelimbs and hindlimbs. The upper parts of hindlimbs and forelimbs are provided with white and black ocelli. Head gray with black mottled markings or spots; supralabials white with black markings. Belly and underside of tail creamy white, tail dorsum creamy grayish. Variations in paratypes. Paratypes of E. killasaifullahi sp. nov. agree with the holotype with some differences given in Table 1 and Figs. 5, 6. Besides sex, the specimens differ in the arrangement of supralabials i.e. subocular wedged between 6 th and 7 th supralabials in all the type series except PMNH 4055 where it is wedged between 5 th and 6 th supralabials. The arrangement of postmentals has a similar pattern in the paratypes except PMNH 3673, where the fifth chins shield is not in contact with the infralabials. In all the type series including the holotype, the fifth chin shield is in contact with the infralabials. The scale count of dorsals, ventrals, gulars, collars, caudals at 9 th –10 th whorl and lamellae under 4 th toe, however, show a unique value for every specimen within a certain range. The infranasal is not in contact with the rostral in all type specimens including the holotype (Figs. 3, 5, 6). Sexual and age dimorphism. Apparently, males attain larger sizes than females in E. killasaifullahi sp. nov.: male SVL to 70.5 mm, female SVL 58.1 mm. Moreover, males have generally longer hindlimbs and shorter trunks as compared to females. For a larger female having SVL of 58.5 mm (PMNH 4050), the hindlimb is 37.9 mm against a same-sized male (PMNH 3616, SVL 59.4 mm) which has a hindlimb length of 43.1 mm. Similarly, the trunk length of a smaller female PMNH 4050 (SVL 58.5 mm) is 29.0 mm against a larger male (PMNH 3614, SVL 67.9 mm) which has a trunk length of 28.1 mm. The dorsal body color and pattern are, however, similar in juveniles and adults of both genders (Figs. 3, 5–7). Comparison. The new species Eremias killasaifullahi sp. nov. is strikingly different from species exhibiting striped and ocellate pattern (E. aria; E. kopetdaghica; E. lalezharica; E. papenfussi; Eremias persica; E. regeli; E. fahimii; E. isfahanica; E. montana; E. nikolskii; E. velox) and ocellate pattern (E. afghanistanica; E. nigrocellata; E. strauchi; E. suphani; Table 1 and S1). Eremias killasaifullahi sp. nov. can be distinguished from E. afghanistanica by a higher count of dorsals (53–63 vs. 44–46), caudal scales in the 9 th –10 th annulus (24–33 vs. 20–26) and a lower number of ventral scales in a single row from the posterior edge of collar to the vent (25–29 vs. 37–38). From E. persica, E. killasaifullahi sp. nov. differs by its smaller size (SVL up to 70.5 mm vs. 98.0 mm), size of the second loreal scale to first loreal scale (more than two times vs. two times), supracaudals (strongly keeled vs. weakly keeled), the dorsal color and pattern in adults (ocellate without broader lateralmost stripe vs. striped and ocellate with broader lateralmost stripe) and tail coloration in the juveniles (creamy grayish vs. bluish). Besides distant distribution, Eremias killasaifullahi sp. nov. differs from the recently described E. fahimii by its comparatively larger size (SVL up to 70.5 mm vs. 56.0 mm), more SDLT 4 th (21–25 vs. 20–21), lower count of caudal scales in the 9 th –10 th annulus (22–27 vs. 31), the greater number of scales separating the femoral pores (1–5 vs. 1) and the dorsal color and pattern in adults (dorsal stripes broken into ocelli without broader lateralmost stripe vs. dorsal stripes persistent throughout life with broader lateralmost stripe). From E. isfahanica, E. killasaifullahi sp. nov. differs in the following morphological characters apart from its distant distribution: higher count of supralabials (8–11 vs. 6–8), 5–7 of them (mainly 6, rarely 5) located anterior to subocular (vs. 5), lower count of collars (10–12 vs. 12–15), number of ventral scales in a single row from the posterior edge of collar to the vent (25–29 vs. 30–33) and the dorsal color pattern in adults (dorsal stripes broken into ocelli vs. dorsal stripes persistent throughout life). Eremias killasaifullahi sp. nov. differs from E. kopetdaghica in having comparatively higher count of dorsals (53–63 vs. 48–59), collars (10–12 vs. 7) and the dorsal color and pattern in adults. Eremias killasaifullahi sp. nov. can be distinguished from E. lalezharica in having a lower number of ventral scales in a single row from posterior edge of collar to the vent (25–29 vs. 30–33), gulars (20–33 vs. 33–40), collars (10–12 vs. 13–15), generally higher count of femoral pores (17–24 vs. 15–19), pair of chin shields/ submaxillary shields (5 vs. 4), contact of gulars with second pair of submaxillary shields (none vs. 1-2 rows) and dorsal color and pattern (ocellate vs. ocellated and striped). Apart from its peculiar distribution in the remote valley in Torghar Mountains, E. killasaifullahi sp. nov. can be differentiated from E. montana in the following set of characters: comparatively larger size (SVL up to 70.5 mm vs. 58.5 mm), lower count of dorsals (53–63 vs. 63–68), higher number of ventral scales in a row across mid-belly in the widest part (14–18 vs. 13–14), infralabials (6–10 vs. 4–6), number of supralabials anterior to the subocular (5–6 vs. 4–5), generally higher count of scales separating the femoral pores (1–5 vs. 2), three scales around the penultimate phalanx of 4 th toe (vs. 4) and dorsal color and pattern (ocellated vs. striped and ocellate). Besides having a subocular scale bordering mouth and ocellate dorsal pattern, E. killasaifullahi sp. nov. differs from E. nigrocellata by its smaller size (SVL up to 70.5 mm vs. 83.0 mm), higher count of dorsals (53–63 vs. 42–56) and the number of femoral pores on each side (17–24 vs. 11–13). E. killasaifullahi sp. nov. differs from E. nikolskii by having a higher count of ventral scales in a row across mid-belly in the widest part (14–18 vs. 14), lower number of ventral scales in a single row from the posterior edge of collar to the vent (25–29 vs. 28–32) and dorsal color and pattern (ocellate vs. striped and ocellate). Besides the dorsal color and pattern, our new species stands distinguished from E. papenfussi by having a lower number of ventral scales in a single row from the posterior edge of the collar to the vent (25–29 vs. 30–33), 5–6 (mainly 6) number of scales anterior to subocular (vs. 5), generally higher count of scales separating the femoral pores (1–5 vs. 1–2). From E. regeli, E. killasaifullahi sp. nov. differs in having three scales around the penultimate phalanx of 4 th toe (vs. four scales), higher count of gulars (20–33 vs. 14–24), ventral scales in a row across mid-belly in the widest part (14–18 vs. 13), generally higher count of caudal scales in the 9 th –10 th annulus (22–27 vs. 17–25) and dorsal color and pattern (ocellate vs. striped and ocellate). The new species E. killasaifullahi sp. nov. can be easily differentiated from E. strauchi by its distant distribution, lower number of ventral scales in a single row from the posterior edge of collar to the vent (25–29 vs. 28–33), and 5–6 (mainly 6) number of scales anterior to subocular (vs. 7). From E. suphani, E. killasaifullahi sp. nov. differs by its distant distribution, lower number of ventral scales in a single row from the posterior edge of collar to the vent (25–29 vs. 29–34) and arrangement of gulars (2 rows of gulars reaching to the second pair of chin

Eremias rafiqi sp. nov. (Table 2, Figs. 4, 5, 6) Suggested vernacular name: Rafiq’s Racerunner Pashto name: Holotype. PMNH 856 (cyt b: n/a; Rag1: n/a), an adult male, collected from Tanishpa village, Torghar Mountains, Killa Saifullah district, Balochistan (31.1869º N, 68.4126º E; Fig. 1), elevation 2,506 m a. s. l., May 25, 1997, leg. Khalid Javed Baig (Fig. 4). Paratypes. Males: PMNH 855 (cyt b: n/a; Rag 1: n/a), 857 (cyt b: n/a; Rag 1: n/a), PMNH 859 (cyt b: n/a; Rag 1: n/a), PMNH 861 (cyt b: n/a; Rag 1: n/a), PMNH 4056 (cyt b: n/a; Rag 1: MT 554487). Females : PMNH 3724 (cyt b: n/a; Rag 1: MT 554476), PMNH 3735 (cyt b: MT 554461; Rag 1: MT 554477), PMNH 4058 (cyt b: n/a; Rag 1: n/a). Juveniles : PMNH 837 (cyt b: n/a; Rag 1: n/a), PMNH 3723 (cyt b: MT 554470; Rag 1: MT 554496), PMNH 4053 (cyt b: MT 554457; Rag 1: MT 554485), PMNH 4054 (cyt b: MT 554454; Rag 1: MT 554480). PMNH 857, collected along with the holotype; PMNH 855 and 861, May 26, 1997, Ashewat, Qamar Din Karez, Zhob district (31.3448ºN, 68.6307ºE), leg. Khalid Javed Baig; PMNH 859, May 24,1997, Tanishpa village, Torghar, Killa Saifullah district, leg. Khalid Javed Baig; PMNH 3723–24, October 09, 2017, Khar, Nushki district, Balochistan (29.5879ºN, 65.6609ºE), leg. Muazzam Ali Khan; PMNH 3735, October 21, 2017, Khar, Nushki district, leg. Muazzam Ali Khan; PMNH 4053, September 05, 2018, Zamkai Nala, Tanishpa village, Torghar, Killa Saifullah district (31.1930ºN, 68.4111ºE), leg. Rafaqat Masroor; PMNH 4054, September 01, 2018, Ashewat, Qamar Din Karez, Zhob district, leg. Rafaqat Masroor; PMNH 4056 and 4058, August 30, 2018, Kunder, Torghar, Killa Saifullah district, leg. Ibad ur Rehman (Figs. 5 & 6). Morphological diagnosis. A large-sized lacertid lizard, maximum snout-vent length (SVL) = 99.3 mm, tail 1.67 to 1.89 times longer than body length (SVL), hindlimbs relatively long (HLL / SVL ratio 0.6–0.8); subocular scale reaching to the edge of the mouth, 5–7 (mainly 6, rarely 5) anterior to subocular; dorsals 56–67; ventrals in 14–17 oblique longitudinal series; frontal separated from supraoculars; the height of the first two to three transverse rows of ventral scales in the pectoral region more than its breadth; 17–21 femoral pores on each side, separated medially by 1–4 scales (mainly 3, rarely 1), the space between the femoral pores less than one–fourth length of each row; toes without fringe, encircled by three scales in a single series of 22–27 unicarinate and bicarinate scales underneath; the tip of the fourth toe reaches to the forelimb and extends to just behind the collar. The adult specimens are grayish in life with four series of longitudinal black ocelli on the dorsum originating from behind the parietals and extending onto the tail; on each lateral side, a broader dark stripe originates from behind the eye and continues onto the tail with disconnected white round ocelli at the margins as well as white ocelli inside the stripe. Molecular data. Eremias rafiqi sp. nov. represents the so-called Zabol clade sensu Rastegar-Pouyani et al. (2010) from eastern Iran and the clade E identified by Khan et al. (2021) from northeastern Balochistan, Pakistan. Whereas Rastegar-Pouyani et al. (2010) identified this clade solely on mtDNA (cyt b, 12S), Khan et al. (2021) used mitochondrial (16S, COI, cyt b) as well as nuclear data (Rag 1) that clearly showed deep differentiation of this new species from other available sequences of the genus. The distinction of Eremias rafiqi sp. nov. is supported by its phylogenetic position (monophyletic clade among other species of E. persica complex, sister to E. fahimii but with weak statistical support; Fig. 1), the differentiation on solely Rag1 dataset (Fig. 2), and the value of the uncorrected p distances reaching from 8.5% (E. fahimii) to 20.7% (E. strauchi) (Table 3). The intraclade genetic diversity (cyt b) is 2 % with six detected haplotypes found in Iran and Pakistan (Fig. 2, Table 3). Etymology. The species epithet “ rafiqi ” is taken from the first name of late Rafique Ahmed Rajput (1968– 2008) to whom the new species is dedicated. The deceased Rajput served in Sindh Wildlife Department from 1986 till his demise. With no formal education in wildlife research, conservation and management, his passion for the conservation of wildlife in Pakistan remain unparalleled. From collecting the first-ever data of Ursus arctos isabellinus in the high-altitude Deosai Plateau, Gilgit-Baltistan, to the faunistic studies in the Indus River Delta and desert areas, he was a symbol of hard work. He also has very sound techniques for the collection of lizards and snakes. Unfortunately, during one such endeavor for gathering faunistic data, he collected a juvenile venomous krait, Bungarus sp. (possibly B. persicus) from Jiwani town, Gwadar District, Balochistan and mistaken its identity with non-venomous Lycodon species. On the morning of October 13, 2008, when he was shifting the live snake from a container to permanently preserve it for research purposes, the snake bit him multiple times.After a couple of hours, Rajput felt severe pain, anxiety and dizziness. He was immediately taken to the hospital in Karachi, where the doctors told his relatives that anti-snake venom is not available and asked them to get it from the pharmacy market. By the time the anti-venom was arranged, Rajput breathed his last. Description of the holotype. SVL: 99.3, TL: 168.0, HL: 26.9, HW: 14.5, HH: 15.5, TrL: 42.5, HLL: 66.8, FLL: 37.3, FrL: 6.3, FrW: 3.4.An adult male preserved in formalin in a good state of preservation (Fig. 4); head and body moderately depressed; tail long, ca. 1.7 times longer than the body, cylindrical and depressed at the base. Head relativelylong (HL/SVL, 0.27) (Fig. 4), ca. 1.7 times longer than wide (HW/HL, 0.58), head height slightly less than head width (HH/HW, 0.89). Limbs strong, hindlimbs ca. 1.8 times longer than the length of forelimbs (FLL/HLL, 0.56), hindlimbs comprise 1.4 times of the body length (HLL/SVL, 0.67). Head slightly broader than the neck.Head shields including nasals,frontonasal,prefrontals,frontal,frontoparietals, interparietal and parietals are smooth and convex. Nasals moderately swollen, three nasals, the lower in contact with two supralabials on both right and left side and in contact with the rostral (Fig. 4D). Supranasals in contact with rostral but lack such contact with first supralabial, the suture between them is 3.7 times the length of frontonasal, whose breadth is slightly more than its length; length of prefrontals 1.6 times its width, forming a median suture; length of frontal ca. 1.8 times as long as broad, its length slightly less than its distance from the tip of the snout, narrow behind; parietals smooth, slightly longer than its width; interparietal smooth, more than half the length of frontoparietals, about equal to the suture of frontoparietal; no occipital. Two large supraoculars, about equal in size, the space anterior to supraoculars filled by few small and three to five larger granules; both supraoculars in contact with frontal of their sides while separated from supraciliaries by a series of granules (Fig. 4C); six supraciliaries, first longest, its length shorter than its distance from the first loreal. Rostral pentagonal, broader than high, narrower beneath than above; anterior loreal slightly higher than wide, shorter than the second loreal which is longer than high; supralabials 10 on the right side, 9 on the left side; subocular keeled just below the eye, bordering the mouth, wedged between sixth and seventh supralabials on the right side and fifth and sixth supralabials on the left side (Fig. 4D). Temporals smooth, a large scale above ear; auricular denticulation indistinct or three small scales forming slight denticulation anteriorly. Lower eyelid covered with numerous small semi-transparent scales. Six infralabials on the right side, seven on the left side, gradually increasing in size posteriorly. Five pairs of chin shields; anterior three completely in contact, the fourth pair separated by 10 to 11 smaller gulars in a straight line, fifth not in contact with infralabials, separated by a single row of scales. Collar curved, free, serrated and composed of ten plates larger than adjacent gulars, the middle one quite enlarged than others. Gular fold distinct, 32 gular scales in a straight line between the symphysis of the chin shields and the collar (Fig. 4B). Dorsal scales granular, smooth, 62 across the middle of the body. Ventral plates broader than long (except for outermost series), forming oblique longitudinal series of 16 plates across mid-belly and 29 transverse rows counted from behind collar to vent; first three rows of ventral scales in the pectoral region behind collar longer than broad, the first row is twice as long as broad. Precloacal region with a pair of the enlarged median plates just above the vent, surrounded by six large scales. Forelimb ca. 1.4 times longer than the head, the upper surface of the arm with rhombic, smooth scales. Scales on the upper surface of hindlimbs similar to dorsals, equal in size; ventral surface of hindlimbs covered by enlarged plates, the ventral surface of the tibia with one row of very large and one comparatively smaller plates; the tip of the fourth toe reaches to the forelimb and extends to just behind the collar; 21 femoral pores on the right side, most of the left side damaged, the two femoral pore series separated by two scales, length of the interfemoral space not greater than one–fourth length of each row. Toes slender, compressed, with no fringe; subdigital lamellae unicarinate, in a single row of 25 scales under the 4th toe, a total of three scales around the 4 th toe. Upper caudal scales oblique, truncate, strongly and diagonally keeled, 30 scales in the 9 th –10 th annulus behind the postcloacal granules. Coloration in life. The adult specimens are grayish in life with four more or less regular rows of black spots on the light dorsum, originating from behind the parietals, smaller on the nape, larger on the middle of the dorsum, disappearing on proximal one-fourth of the tail. The middle two rows of black spots have white spots along each black spot of the rows. On each lateral side, a broader dorsolateral dark stripe originates from behind the eye and continues onto the tail with disconnected white ocelli at both margins as well as white ocelli inside the stripe; next to the broader dark stripe, a lateral-most stripe is composed of disconnected black ocelli, originating from behind tympanum and reaching to the hindlimb. Upper parts of both hindlimbs and forelimbs with white and black ocelli. Head gray without any markings or spots; labials white with black markings. Belly and underside of tail creamy white, tail dorsum sandy grayish. The juveniles and subadults are nearly similar in coloration to the adults except for the following details; four longitudinal dark stripes on the body, the outermost originate from anterior parietals on the outer side and continue onto the tail, the innermost originate from the posterior of parietals and merge after running a while on the tail. A broader lateral stripe on each side originates from behind the eye and continues on the lateral side of the body and tail with interspersed white ocelli between the forelimb and hindlimb. The dorsal forelimb and hindlimb are dark gray with white ocelli. Variations in paratypes. The paratypes of E. rafiqi sp. nov. agree with the holotype with some differences given in Table 2 (Figs. 5, 6). Besides sex, the specimens differ in the arrangement of supralabials i.e. subocular wedged between 6 th and 7 th in all the type series except PMNH 861 (between 7 th and 8 th) and PMNH 3724 (between 5 th and 6 th). The arrangement of postmentals has a similar pattern in the paratypes except PMNH 855, 861, 837, 3723, 3735 and 4054, where the fifth chin shield is in contact with the infralabials. The contact of postmental shield with the supralabials varies in the paratypes; PMNH 855 and 3724, the fifth postmental shield is in contact with 7 th supralabial; PMNH 837 and 3735, the fifth postmental shield is in contact with sixth supralabial; PMNH 861, the fifth postmental shield is in contact with 7 th and 8 th supralabials. The infranasal scale in PMNH 837, 855, 859, 861, 3723, 3735, 4053–4054 and 4058 rests on first, second and third supralabials. The scale count of dorsals, ventrals, gulars, collars, caudals at 9 th –10 th annuli and lamellae under 4 th toe, however, show a unique value for every specimen within a certain range. Sexual and age dimorphism. Apparently, males attain larger sizes than females in E. rafiqi sp. nov.: male SVL to 99.3 mm, female SVL 82.1 mm. Moreover, males have generally longer hindlimbs and shorter trunks as compared to females. For a larger female having SVL of 98.1 mm (PMNH 4058), the hindlimb is 56.0 mm against a smaller-sized male (PMNH 855, SVL 92.2 mm) which has a hindlimb length of 57.1 mm. Similarly, the trunk length of a smaller female (PMNH 3724, SVL 75.1 mm) is 36.8 mm against a larger male (PMNH 857, SVL 78.5 mm) which has a trunk length of 34.5 mm. The dorsal body color and pattern, however, varies in juveniles and adults of both genders (Figs. 5, 6). Comparison. The new species Eremias rafiqi sp. nov. is strikingly different from species exhibiting striped and ocellate patterns in the subgenus Aspidorhinus (E. kopetdaghica Szczerbak, 1972, E. lalezharica Moravec, 1994, E. papenfussi Mozaffari et al., 2011, Eremias persica Blanford, 1874, E. regeli Bedriaga, 1905, E. fahimii Mozaffari et al., 2020, E. isfahanica Rastegar-Pouyani et al., 2016, E. montana Rastegar-Pouyani & RastegarPouyani, 2001, E. nikolskii Bedriaga, 1905, E. velox Pallas, 1771) and ocellate pattern (E. killasaifullahi sp. nov., E. afghanistanica Böhme & Szczerbak, 1991, E. roborowskii Bedriaga, 1912, E. strauchi Kessler, 1878, E. suphani Başoğlu & Hellmich, 1968). The new species E. rafiqi sp. nov. can also be differentiated from the geographically closely-distributed members of the subgenus Eremias having striped and ocellate pattern (E. aria Anderson & Leviton 1967) and ocellate pattern (E. nigrocellata Nikolsky 1896) by the arrangement of subocular scale which borders the mouth (Supplementary Tab. 1; see published data in Lantz 1928, Szczerbak 1974, Bischoff & Böhme 1980, Böhme & Szczerbak 1991, Anderson 1999). A brief of morphological differences is provided (the material used for a first-hand comparison is listed in parentheses at each species; see also Table 2 and S 1). Besides striped and ocellate body pattern (vs. ocellate), E. rafiqi sp. nov. can be distinguished from E. afghanistanica by its larger size (SVL up to 99.3 mm vs. 67.0 mm), higher count of dorsals (56–67 vs. 44–46), gulars (30–36 vs. 25–28), femoral pores (17–22 vs. 16–18), caudal scales in the 9 th –10 th annulus (24–33 vs. 22–25), 5–7 supralabials (mainly 6, rarely 5) located anterior to subocular (vs. 5) and lower number of ventral scales in a single row from the posterior edge of collar to the vent (29–33 vs. 37–38). From E. persica, that is partly close in dorsal coloration, pattern and size, E. rafiqi sp. nov. differs in the length of interparietal to the length of suture of parietals (longer vs. shorter), length of frontonasal to its width (longer vs. as long as wide), size of the second loreal scale to first loreal scale (more than three times vs. two times), supracaudals (strongly keeled vs. weakly keeled) and tail coloration in the juveniles (sandy grayish vs. bluish). Besides distant distribution, Eremias rafiqi sp. nov. differs from the recently described E. fahimii by its larger size (SVL up to 99.3 mm vs. 56.0 mm), more SDLT 4 th (22–27 vs. 20–21), the greater number of scales separating the femoral pores (1–4 vs. 1) and the dorsal color and pattern in adults (dorsal stripes broken into ocelli vs. dorsal stripes persistent throughout life). From E. isfahanica, E. rafiqi sp. nov. differs in the following morphological characters apart from its distant distribution: higher count of supralabials (8–10 vs. 6–8), 5–7 (mainly 6, rarely 5) of them located anterior to subocular (vs. 5), lower count of collars (8–12 vs. 12–15) and the dorsal color pattern in adults (dorsal stripes broken into ocelli vs. dorsal stripes persistent throughout life). Eremias rafiqi sp. nov. differs from E. kopetdaghica in having a higher count of dorsals (56–67 vs. 48–59), gulars (30–36 vs. 19–28), caudal scales in the 9 th –10 th annulus (24–33 vs. 20–26) and collars (8–12 vs. 7) and the dorsal color and pattern in adults (presence of a broader dark stripe on each lateral side above flanks with disconnected white ocelli at the margins as well as white ocelli inside the stripe vs. no such lateral broader stripes). Eremias rafiqi sp. nov. can be distinguished from E. lalezharica in having a higher count of dorsals (56–67 vs. 54–59), femoral pores (17–22 vs. 15–19), pair of chin shields/ submaxillary shields (5 vs. 4), lower number of collars (8–12 vs. 13–15), contact of gulars with second pair of submaxillary shields (none vs. 1-2 rows of gulars with the second pair of submaxillary shields) and dorsal color and pattern. Apart from its peculiar distribution in the remote valley in Torghar Mountains, a part of the Palearctic region, E. rafiqi sp. nov. can be differentiated from E. montana in the following set of characters: larger size (SVL up to 99.3 mm vs. 58.5 mm), higher count of ventral scales in a row across mid-belly in the widest part (14–17 vs. 13–14), number of ventral scales in a single row from the posterior edge of collar to the vent (29–33 vs. 27–28), gulars (30–36 vs. 23–25), infralabials (6–10 vs. 4–6), number of supralabials anterior to the subocular (5–7 vs. 4–5), generally more SDLT 4 th (22–27 vs. 18–25), generally higher count of scales separating the femoral pores (1–4 vs. 2) and dorsal color and pattern. From E. nigrocellata, E. rafiqi sp. nov. differs in dorsal body pattern (striped and ocellate vs. ocellate), higher count of dorsals (56–67 vs. 42–56), lower number of ventral scales in a row across mid-belly i

Eremias killasaifullahi sp. nov. (Table 1, Figs. 3, 5, 6) Suggested vernacular name: Killa Saifullah’s Racerunner Pashto name: Holotype. PMNH 3613 (cyt b: MT 554460; Rag1: MT 554498), an adult male, collected from Kunder, Torghar Mountains, Killa Saifullah district, Balochistan (31.3247º N, 68.5452º E; Fig. 7D), elevation 1,920 m a. s. l., March 23, 2017, leg. Rafaqat Masroor (Fig. 3). Paratypes. Males: PMNH 3614–3616 (cyt b: MT554466, MT554456, n/a; Rag1: MT554478, MT554482, n/a). Females: PMNH 4046 (cyt b: MT 554453; Rag1: MT 554479), PMNH 4050 (cyt b: MT 554473; Rag1: MT 554483), PMNH 4055 (cyt b: MT 554455; Rag1: MT 554481). Juveniles: PMNH 3673 (cyt b: n/a; Rag1: n/a), PMNH 4045 (cyt b: MT 554467; Rag1: MT 554486), PMNH 4052 (cyt b: MT 554459; Rag1: MT 554497). PMNH 3614–16, 3673 collected along with the holotype; PMNH 4045, 4052, September 5, 2018, Zamkai Nala, Tanishpa, Killa Saifullah district, leg. Rafaqat Masroor; PMNH 4046, 4055 August 31, 2018, Ashewat, Qamar Din Karez, Zhob district, leg. Rafaqat Masroor; PMNH 4050, September 1, 2018, Zamkai Nala, Tanishpa, Killa Saifullah district, leg. Rafaqat Masroor (Figs. 5, 6). Morphological diagnosis. A medium-sized lacertid lizard, maximum snout-vent length (SVL) = 70.5 mm, tail 1.67 to 1.97 times longer than body length (SVL), hindlimbs relatively long (HLL / SVL ratio 0.6–0.8); subocular scale reaching to the edge of the mouth, 5–7 (mainly 6, rarely 5) anterior to subocular; dorsals 53–63; ventrals in 14–18 oblique longitudinal series; frontal separated from supraoculars; the height of the first two to three transverse rows of ventral scales in the pectoral region more than its breadth; 17–24 femoral pores on each side, separated medially by 1–5 scales (mainly 2–4, rarely 1), the space between the femoral pores less than one–fourth length of each row; toes without fringe, encircled by three scales in a single series of 21–25 unicarinate and bicarinate scales underneath; tip of the fourth toe reaches to the forelimb and extends to just behind the collar. The adult specimens are creamy beige in life with seven light stripes appearing on the neck which transforms into ocelli and vermiculation behind the neck. No dorsolateral broader dark stripes, an outer-most series of white and black ocelli starts behind the eyes on each side, onto the tympanum and flanks above the forelimb and hindlimb insertion. ......continued on the next page TABLE 1. (Continued) ......continued on the next page TABLE 2. (Continued) ......continued on the next page TABLE 2. (Continued) ......continued on the next page TABLE 2. (Continued) Molecular data. Eremias killasaifullahi sp. nov. represents a newly detected evolutionary lineage (Fig. 1) of the genus Eremias (Aspidorhinus) that was firstly detected by Khan et al. (2021) as the clade F (with subclades F1, F2) based on four studied genetic markers (16S, COI, cyt b, Rag1). This lineage was detected occurring in NE Balochistan in Pakistan and represents local microendemism (Fig. 2). The lineage deeply diverges and is sister to all other lineages of such called E. persica complex (see Khan et al. 2021 and Fig. 1 in this study) and well differentiated in the Rag1 dataset (Fig. 2). The lineage genetically (uncorrected p distances) differs from 14.5% (E. strauchi) to 21.6% (E. velox) (Table 3) among species of the subgenus Aspidorhinus. Its average intraclade genetic variability (cyt b) is 3% (Fig. 2). Despite a very small known range of distribution, the distances between F1 and F2 subclades sensu Khan et al. (2021) reached 4.6% and the haplotype network based on cyt b dataset showed six different haplotypes. High allele diversity was also detected by analyzing the Rag1 marker (Fig. 2). Etymology. We derived the name of the new species from Killa Saifullah (Pashto:; also Qilla Saifullah), a city and district in northwestern Balochistan province, Pakistan that represents the area, from where this newly discovered endemic species of Eremias (subgenus Aspidorhinus) is currently known. The region plays an important role for producing fruits, nuts and vegetables in Pakistan. The discovery of this species of lizards thus highlights the importance of this region from the biodiversity point of view. Description of the holotype. SVL: 65.3, TL: 109.7, HL: 17.4, HW: 10.2, HH: 8.1, TrL: 28.3, HLL: 44.8, FLL: 26.6, FrL: 4.6, FrW: 2.3.An adult male of E. killasaifullahi sp. nov. preserved in ethanol in a good state of preservation (Fig. 3); head and body moderately depressed; tail long, ca. 1.7 times longer than the body, cylindrical and depressed at the base. Head relativelylong (HL/SVL ratio 0.27) (Fig. 3), 1.7 times longer than wide (HW/HL ratio 0.59), head height less than head width (HH/HW, 0.79). Limbs strong, hindlimbs 1.6 times more than the length of forelimbs (FLL/HLL, 0.59), hindlimbs comprise 1.4 times the body length (HLL/SVL, 0.69). Head broader than the neck; nasals, frontonasal, prefrontals, frontal, frontoparietals, interparietal and parietals are smooth and convex. Nasals are moderately swollen, three nasals, the lower in contact with three supralabials on the right and left side, its contact with the rostral lacking (Fig. 3D). Supranasals in contact with rostral and first supralabial, the suture between them is four times the length of frontonasal, whose breadth is ca. 1.1 times its length; length of prefrontals 1.4 times its width, joined by a median suture; frontal two times as long as broad, its length slightly less than its distance from the tip of the snout, narrow behind; parietals smooth, slightly longer than wide; interparietal smooth, more than half of the length of frontoparietals; no occipital. Two large supraoculars, about equal in size, the space anterior to supraoculars filled by few small and three to five larger granules; both supraoculars in contact with frontal of their sides while separated from supraciliaries by a series of granules (Fig. 3C), behind the two large spraoculars a single, comparatively medium-sized, granule exist; six supraciliaries, first longest, its length shorter than its distance from the first loreal. Rostral pentagonal, broader than high, narrower beneath than above; anterior loreal slightly higher than wide, shorter than the second loreal which is longer than high; supralabials 8; subocular keeled just below the eye, bordering the mouth, wedged between fifth and sixth supralabials (Fig. 3D). Temporals smooth, a large scale above ear; auricular denticulation indistinct or three small scales forming slight denticulation anteriorly. Lower eyelid covered with numerous small semi-transparent scales. Six infralabials, gradually increasing in size posteriorly. Five pairs of chin shields; anterior three completely in contact, the fourth one separated by six smaller gulars, the fifth one is in contact with fifth and sixth infralabials on both sides. Collar curved, free, serrated and composed of 10 plates larger than adjacent gulars, the middle one slightly enlarged than others. Gular fold distinct, 20 gular scales in a straight line between the symphysis of the chin shields and the collar (Fig. 3B). Dorsal scales granular, smooth, 60 across the middle of the body. Ventral plates broader than long (except for outermost series), forming oblique longitudinal series of 16 plates across mid-belly and 25 transverse rows counted from behind collar to vent; first three rows of ventral scales in the pectoral region behind collar longer than broad, the first row is twice as long as broad. Precloacal region with an enlarged median plate just above the vent, surrounded by four large scales. Forelimb ca. 1.5 times longer than the head, upper surface of the arm with rhombic, smooth scales. Scales on the upper surface of hindlimbs similar to dorsals, varying in size; ventral surface of hindlimbs covered by enlarged plates, the lower surface of the tibia with one row of very large and one comparatively smaller plates, the tip of the fourth toe reaches to the forelimb and extends to just behind the collar; 17 femoral pores on the right side, the left side damaged, the two series separated by two scales, length of the interfemoral space not greater than one-fourth length of each row. Toes slender, compressed, with no fringe. Subdigital lamellae unicarinate, in a single row of 21 scales under the 4th toe, a total of three scales around the 4 th toe. Upper caudal scales oblique, truncate, strongly and diagonally keeled, 26 scales in the 9 th –10 th annulus behind the postcloacal granules. Coloration in life. The adult specimens (Fig. 7E) are creamy beige with ocellate body pattern. Seven light stripes appear on the neck which transforms into ocelli and vermiculation behind the neck. Of the seven, the lateralmost light stripe originates from behind the eye and runs on the outer edge of the parietal, transforming into a disconnected series of white ocelli edged with black, running up to anterior one-third of the tail. Next to the lateralmost, the paravertebral light stripe originates from behind the parietal and transforms into closely-connected white ocelli edged with black and runs on the tail short of lateral-most ocelli. Next to paravertebral light stripe, there exists a light nuchal stripe on each side and the light vertebral stripe, the three joins behind the neck and transform into white ocelli edged with black in the pattern of vermiculation. In addition to seven light stripes on the neck, an outermost series of white and black ocelli starts behind the eyes on each side, onto the tympanum and flanks above the forelimb and hindlimb insertion. The upper parts of both hindlimbs and forelimbs are provided with white and black ocelli. Head gray with black mottled markings or spots; supralabials white with black markings. Belly and underside of tail creamy white, tail dorsum grayish. The juveniles and subadults (Fig. 6) are nearly similar in coloration to the adults except for the following details; seven longitudinal light stripes on the neck, the lateral-most originate from behind the eye, running on the outer parietals and continuing onto the dorsum in the form of connected small white ocelli, terminating on the one-third of the tail, the paravertebral light stripe originates from the posterior of parietals, and merge short of the lateral-most stripe on the tail, the nuchal of each side and vertebral light stripe merge after the neck to form light vermiculation up to the base of the tail. An additional outer-most light stripe originates from behind the tympanum and is produced in the form of disconnected white ocelli above the insertion of forelimbs and hindlimbs. The upper parts of hindlimbs and forelimbs are provided with white and black ocelli. Head gray with black mottled markings or spots; supralabials white with black markings. Belly and underside of tail creamy white, tail dorsum creamy grayish. Variations in paratypes. Paratypes of E. killasaifullahi sp. nov. agree with the holotype with some differences given in Table 1 and Figs. 5, 6. Besides sex, the specimens differ in the arrangement of supralabials i.e. subocular wedged between 6 th and 7 th supralabials in all the type series except PMNH 4055 where it is wedged between 5 th and 6 th supralabials. The arrangement of postmentals has a similar pattern in the paratypes except PMNH 3673, where the fifth chins shield is not in contact with the infralabials. In all the type series including the holotype, the fifth chin shield is in contact with the infralabials. The scale count of dorsals, ventrals, gulars, collars, caudals at 9 th –10 th whorl and lamellae under 4 th toe, however, show a unique value for every specimen within a certain range. The infranasal is not in contact with the rostral in all type specimens including the holotype (Figs. 3, 5, 6). Sexual and age dimorphism. Apparently, males attain larger sizes than females in E. killasaifullahi sp. nov.: male SVL to 70.5 mm, female SVL 58.1 mm. Moreover, males have generally longer hindlimbs and shorter trunks as compared to females. For a larger female having SVL of 58.5 mm (PMNH 4050), the hindlimb is 37.9 mm against a same-sized male (PMNH 3616, SVL 59.4 mm) which has a hindlimb length of 43.1 mm. Similarly, the trunk length of a smaller female PMNH 4050 (SVL 58.5 mm) is 29.0 mm against a larger male (PMNH 3614, SVL 67.9 mm) which has a trunk length of 28.1 mm. The dorsal body color and pattern are, however, similar in juveniles and adults of both genders (Figs. 3, 5–7). Comparison. The new species Eremias killasaifullahi sp. nov. is strikingly different from species exhibiting striped and ocellate pattern (E. aria; E. kopetdaghica; E. lalezharica; E. papenfussi; Eremias persica; E. regeli; E. fahimii; E. isfahanica; E. montana; E. nikolskii; E. velox) and ocellate pattern (E. afghanistanica; E. nigrocellata; E. strauchi; E. suphani; Table 1 and S1). Eremias killasaifullahi sp. nov. can be distinguished from E. afghanistanica by a higher count of dorsals (53–63 vs. 44–46), caudal scales in the 9 th –10 th annulus (24–33 vs. 20–26) and a lower number of ventral scales in a single row from the posterior edge of collar to the vent (25–29 vs. 37–38). From E. persica, E. killasaifullahi sp. nov. differs by its smaller size (SVL up to 70.5 mm vs. 98.0 mm), size of the second loreal scale to first loreal scale (more than two times vs. two times), supracaudals (strongly keeled vs. weakly keeled), the dorsal color and pattern in adults (ocellate without broader lateralmost stripe vs. striped and ocellate with broader lateralmost stripe) and tail coloration in the juveniles (creamy grayish vs. bluish). Besides distant distribution, Eremias killasaifullahi sp. nov. differs from the recently described E. fahimii by its comparatively larger size (SVL up to 70.5 mm vs. 56.0 mm), more SDLT 4 th (21–25 vs. 20–21), lower count of caudal scales in the 9 th –10 th annulus (22–27 vs. 31), the greater number of scales separating the femoral pores (1–5 vs. 1) and the dorsal color and pattern in adults (dorsal stripes broken into ocelli without broader lateralmost stripe vs. dorsal stripes persistent throughout life with broader lateralmost stripe). From E. isfahanica, E. killasaifullahi sp. nov. differs in the following morphological characters apart from its distant distribution: higher count of supralabials (8–11 vs. 6–8), 5–7 of them (mainly 6, rarely 5) located anterior to subocular (vs. 5), lower count of collars (10–12 vs. 12–15), number of ventral scales in a single row from the posterior edge of collar to the vent (25–29 vs. 30–33) and the dorsal color pattern in adults (dorsal stripes broken into ocelli vs. dorsal stripes persistent throughout life). Eremias killasaifullahi sp. nov. differs from E. kopetdaghica in having comparatively higher count of dorsals (53–63 vs. 48–59), collars (10–12 vs. 7) and the dorsal color and pattern in adults. Eremias killasaifullahi sp. nov. can be distinguished from E. lalezharica in having a lower number of ventral scales in a single row from posterior edge of collar to the vent (25–29 vs. 30–33), gulars (20–33 vs. 33–40), collars (10–12 vs. 13–15), generally higher count of femoral pores (17–24 vs. 15–19), pair of chin shields/ submaxillary shields (5 vs. 4), contact of gulars with second pair of submaxillary shields (none vs. 1-2 rows) and dorsal color and pattern (ocellate vs. ocellated and striped). Apart from its peculiar distribution in the remote valley in Torghar Mountains, E. killasaifullahi sp. nov. can be differentiated from E. montana in the following set of characters: comparatively larger size (SVL up to 70.5 mm vs. 58.5 mm), lower count of dorsals (53–63 vs. 63–68), higher number of ventral scales in a row across mid-belly in the widest part (14–18 vs. 13–14), infralabials (6–10 vs. 4–6), number of supralabials anterior to the subocular (5–6 vs. 4–5), generally higher count of scales separating the femoral pores (1–5 vs. 2), three scales around the penultimate phalanx of 4 th toe (vs. 4) and dorsal color and pattern (ocellated vs. striped and ocellate). Besides having a subocular scale bordering mouth and ocellate dorsal pattern, E. killasaifullahi sp. nov. differs from E. nigrocellata by its smaller size (SVL up to 70.5 mm vs. 83.0 mm), higher count of dorsals (53–63 vs. 42–56) and the number of femoral pores on each side (17–24 vs. 11–13). E. killasaifullahi sp. nov. differs from E. nikolskii by having a higher count of ventral scales in a row across mid-belly in the widest part (14–18 vs. 14), lower number of ventral scales in a single row from the posterior edge of collar to the vent (25–29 vs. 28–32) and dorsal color and pattern (ocellate vs. striped and ocellate). Besides the dorsal color and pattern, our new species stands distinguished from E. papenfussi by having a lower number of ventral scales in a single row from the posterior edge of the collar to the vent (25–29 vs. 30–33), 5–6 (mainly 6) number of scales anterior to subocular (vs. 5), generally higher count of scales separating the femoral pores (1–5 vs. 1–2). From E. regeli, E. killasaifullahi sp. nov. differs in having three scales around the penultimate phalanx of 4 th toe (vs. four scales), higher count of gulars (20–33 vs. 14–24), ventral scales in a row across mid-belly in the widest part (14–18 vs. 13), generally higher count of caudal scales in the 9 th –10 th annulus (22–27 vs. 17–25) and dorsal color and pattern (ocellate vs. striped and ocellate). The new species E. killasaifullahi sp. nov. can be easily differentiated from E. strauchi by its distant distribution, lower number of ventral scales in a single row from the posterior edge of collar to the vent (25–29 vs. 28–33), and 5–6 (mainly 6) number of scales anterior to subocular (vs. 7). From E. suphani, E. killasaifullahi sp. nov. differs by its distant distribution, lower number of ventral scales in a single row from the posterior edge of collar to the vent (25–29 vs. 29–34) and arrangement of gulars (2 rows of gulars reaching to the second pair of chin, Published as part of Masroor, Rafaqat, Khan, Muazzam Ali, Nadeem, Muhammad Sajid, Amir, Shabir Ali, Khisroon, Muhammad & Jablonski, Daniel, 2022, Appearances often deceive in racerunners: integrative approach reveals two new species of Eremias (Squamata: Lacertidae) from Pakistan, pp. 55-87 in Zootaxa 5175 (1) on pages 58-70, DOI: 10.11646/zootaxa.5175.1.3, http://zenodo.org/record/7003222, {"references":["Khan, M. A., Jablonski, D., Nadeem, M. S., Masroor, R., Kehlmaier, C., Spitzweg, C. & Fritz, U. (2021) Molecular phylogeny of Eremias spp. from Pakistan contributes to a better understanding of the diversity of racerunners. Journal of Zoological Systematics and Evolutionary Research, 59, 466 - 483. https: // doi. org / 10.1111 / jzs. 12426","Masroor, R., Khisroon, M., Khan, M. A. & Jablonski, D. (2020 b) A new species of Eremias Fitzinger, 1834 (Squamata: Lacertidae) from the arid mountains of Pakistan. Zootaxa, 4786 (1), 101 - 121. https: // doi. org / 10.11646 / zootaxa. 4786.1.8"]}

Subgenus Aspidorhinus Eichwald, 1841 Type species: Eremias velox (Pallas, 1771), Published as part of Masroor, Rafaqat, Khan, Muazzam Ali, Nadeem, Muhammad Sajid, Amir, Shabir Ali, Khisroon, Muhammad & Jablonski, Daniel, 2022, Appearances often deceive in racerunners: integrative approach reveals two new species of Eremias (Squamata: Lacertidae) from Pakistan, pp. 55-87 in Zootaxa 5175 (1) on page 58, DOI: 10.11646/zootaxa.5175.1.3, http://zenodo.org/record/7003222