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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 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 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 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

Lizardscomprise 7176 described species all over the world. Of these, 362 speciesbelong to the family Lacertidae. Herein, the cranial skeletal differences andsimilarities of Anatololacerta anatolica (Werner, 1900) and Anatololacertapelasgiana (Mertens 1959) species, which are distributed in the south andnorth of the Büyük Menderes River in Turkey and are included in the familyLacertidae, were investigated. Osteologicalcharacters are important for biological information. In the crrent study, thecranial osteology of A. anatolicaand A. pelasgiana are described based on cleared anddouble-stained specimens. We used 15 museum samples, which stored in the FaunaFlora Research and Applied Centre (Dokuz Eylül University), for each species inthe current study. Our result suggests that at least the skull roof, nasal andpalatal regions of skull of A. anatolica can be distinguished from thoseof specimens of A. pelasgiana. These differences are caused by the nasalprocess of the premaxilla, the posterior process of the nasal, the junction ofthe anterolateral and anteromedial process of the frontal, the posterolateralmargin of the frontal and the numbers of teeth that are on the premaxilla, maxilla,dentary, and pterygoid bones. Despite thelarge number of lacertid lizards, information about the detailed descriptionsof their cranial osteology is scarce. In this study, comparative detailedcranial osteology of these two allopatric species separated by the BüyükMenderes River as a geographic barrier was performed. In this way, it is aimedto provide important data for future comparative morphological studies.

LAUDAKIA VULGARIS (SONNINI & LATREILLE, 1802) STAT. NOV. Included taxa: Laudakia s. picea (Parker, 1935), L. s. brachydactyla (Haas, 1951), L. s. vulgaris (Sonnini & Latreille, 1802), L. s. salehi (Werner, 2006) and the Jordan phylogenetic subclade. Type specimen: Agama stellio vulgaris (Sonnini & Latreille, 1802), FMNH 153134 (neotype) Crochet et al. (2006) (Field Museum of Natural History, Chicago, USA). Type locality: El Amiria, Alexandria, Lower Egypt, by present neotype designation. Distribution: All localities from Egypt, Jordan, Israel and south-western and western Syria (east to Palmyra). Remarks: According to the principle of priority of the ICZN (International Code of Zoological Nomenclature), this distinct genetic lineage was named vulgaris as that is the oldest available name. It includes the morphological subspecies of L. s. picea, L. s. brachydactyla, L. s. vulgaris and L. s. salehi, as well as a new phylogenetic subclade that corresponds to the Jordan populations from the eastern side of the Wadi–Araba valley, previously reported as an unnamed subspecies. The subspecific description of the latter is under preparation, but we have to note that the subspecific status of L. s. vulgaris and L. s. salehi is ambiguous and requires further investigation.

LAUDAKIA STELLIO (LINNAEUS, 1758) S.S. Included taxa: Laudakia s. stellio (Linnaeus, 1758), L. s. daani (Beutler & Frör, 1980) and the subclades from the western Taurus, Symi–Kaş, Anatolia, Lebanon and Hatay, as shown in this study (Figs 2, 4). Type specimen: Laudakia stellio stellio (Linnaeus, 1758), ZFMK 2063 (neotype), Crochet et al. (2006) (Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany). Type locality: Delos Island, Cyclades, Greece. Distribution: All localities from Greece (Cyclades, Crete, Thessaloniki, Corfu and East Aegean islands) and Turkey, north-western Syria and northern Lebanon. Remarks: According to the principle of priority of the ICZN, this distinct genetic lineage should be named stellio because that is the oldest available name. It includes already described morphological subspecies and other cryptic lineages of Clade 2, revealed by this study: the nominal L. s. stellio and its sister Anatolia subclade, L. s. daani and its sister W Taurus–Symi–Kaş subclades, as well as the Lebanon and Hatay subclades. The subspecific status of L. s. stellio and L. s. daani is retained, while the elevation of the Lebanon and Hatay lineages to the subspecies level is under preparation., Published as part of Karameta, Emmanouela, Lymberakis, Petros, Grillitsch, Heinz, Ilgaz, Çetin, Avci, Aziz, Kumlutaş, Yusuf, Candan, Kamil, Wagner, Philipp, Sfenthourakis, Spyros, Pafilis, Panayiotis & Poulakakis, Nikos, 2022, The story of a rock-star: multilocus phylogeny and species delimitation in the starred or roughtail rock agama, Laudakia stellio (Reptilia: Agamidae), pp. 195-219 in Zoological Journal of the Linnean Society 195 (1) on page 212, DOI: 10.1093/zoolinnean/zlab107, http://zenodo.org/record/6530771, {"references":["Beutler A, Fror E. 1980. Die Amphibien und Reptilien der Nordkykladen (Griechenland). Mitteilungen der Zoologischen Gesellschaft Braunau 3: 255 - 290.","Crochet PA, Lymberakis P, Werner YL. 2006. The type specimens of Laudakia stellio (Linnaeus) (Reptilia: Agamidae) and its subspecies. Journal of Natural History 40: 461 - 471."]}

WOS:000787283000002 The dwarf lizard, Parvilacerta parva, is a characteristic member of the steppe biome in Irano-Anatolian biological hotspot. While this lizard has been included in local faunistic surveys and its morphological variation was addressed, no targeted study has been performed on the ecology and distribution of the species. Here we investigate the range dynamics of dwarf lizards during recent glacial and interglacial periods. We looked at the effects of climatic oscillations on species distribution at Present, Last Glacial Maximum (LGM), and Last Interglacial (LIG) periods using ecological niche modeling (ENM), based on our fieldwork and literature data. The model results suggest that the range of dwarf lizards contracted during the LIG and expanded during the LGM, opposite to the pattern observed in many other temperate reptiles. During the LIG, the distribution of the dwarf lizards had been restricted to the mountainous steppe habitats in Northeastern Anatolia, but during the LGM it expanded to the west by including the new steppe habitats in Sultan, Emir, and Murat mountains and adjacent areas. Climatic factors had a strong influence on shaping the spatiotemporal habitat. The Anatolian Biogeographic Region overlaps with Irano–anatolian biodiversity hotspot, reflecting remarkable species richness in this area. However, faunal elements of the hotspots are under threat due to not only global climate change, but also anthropogenic pressures, such as habitat loss and overgrazing. Our results suggest that the dwarf lizards have potential as indicators for tracking the local effects of global climate change as well as human-induced degradation of the steppe habitat.

Age structure and longevity in 47 museum specimens (31 females and 16 males) of Lacerta agilis from two different populations in Turkey were skeletochronologically studied. The mean SVL of males was not significantly different from those of females, although intersexual differences in body size are female-biased. The age structure ranged from 3 to 6 years in male individuals, and 3 to 8 years in females. Age at sexual maturity was 2 years for males, while females reached sexual maturity after the third year of life. The age did not significantly differ between the sexes. Additionally, age structure and SVL did not significantly differ between the two populations. A significant positive correlation between SVL and age structure was present for the Kars population, although there was a weak correlation for the Ardahan population. Considering sex, there was a significant difference in terms of SVL for the Ardahan population, while age structure significantly differed for the Kars population., Türkiye'deki iki farklı popülasyondan Lacerta agilis'in 47 müze örneğinde (31 dişi ve 16 erkek) yaş yapısı ve yaşam süresi iskelet kronolojisi kullanılarak incelenmiştir. Cinsiyete bağımlı vücut büyüklüğü dişi eğilimli olmasına rağmen, erkeklerin ortalama SVL’si dişilerden önemli ölçüde farklı değildir. Yaş dağılımı erkek bireylerde 3 ile 6 yıl arasında, dişilerde ise 3 ile 8 yıl arasında değişmektedir. Erkekler iki, dişiler ise yaşamın üçüncü yılından sonra cinsel olgunluğa ulaşır. Yaş, cinsiyetler arasında önemli ölçüde farklılık göstermez. Bununla birlikte, yaş yapısı ve SVL iki popülasyon arasında önemli ölçüde farklı değildir. SVL ile yaş arasında Ardahan popülasyonu için zayıf bir korelasyon olsa da Kars popülasyonu için anlamlı ölçüde bir pozitif korelasyon vardır. Cinsiyetler göz önüne alındığında, Ardahan popülasyonunda SVL için, Kars popülasyonda ise yaş dağılımı için anlamlı bir fark söz konusudur.