| Darevskia adjarica (DAREVSKY & EISELT, 1980) Darevskia alpina (DAREVSKY, 1967) Darevskia armeniaca (MÉHELY, 1909) Darevskia arribasi TUNIYEV, PETROVA & LOTIEV, 2023 Darevskia bendimahiensis (SCHMIDTLER et al., 1994) Darevskia bithynica (MÉHELY, 1909) Darevskia brauneri (MÉHELY, 1909) Darevskia caspica AHMADZADEH et al. 2013 Darevskia caucasica (MÉHELY, 1909) Darevskia chlorogaster (BOULENGER, 1908) Darevskia clarkorum (DAREVSKY & VEDMEDERJA, 1977) Darevskia daghestanica (DAREVSKY, 1967) Darevskia dahli (DAREVSKY, 1957) Darevskia defilippii (CAMERANO, 1877) Darevskia derjugini (NIKOLSKY, 1898) Darevskia josefschmidtleri ARRIBAS et al., 2022 Darevskia kamii AHMADZADEH et al. 2013 Darevskia kopetdaghica AHMADZADEH et al. 2013 Darevskia lindholmi (SZCZERBAK, 1962) Darevskia mirabilis ARRIBAS, ILGAZ, KUMLUTAS, DURMUS, AVCI & ÜZÜM, 2013 Darevskia mixta (MÉHELY, 1909) Darevskia nairensis (DAREVSKY, 1967) Darevskia parvula (LANTZ & CYRÉN, 1913) Darevskia pontica (LANTZ & CYRÉN, 1918) Darevskia portschinskii (KESSLER, 1878) Darevskia praticola (EVERSMANN, 1834) Darevskia raddei (BOETTGER, 1892) Darevskia rostombekowi (DAREVSKY, 1957) Darevskia rudis (BEDRIAGA, 1886) Darevskia salihae KURNAZ, ŞAHIN & EROĞLU, 2022 Darevskia sapphirina (SCHMIDTLER et al., 1994) Darevskia saxicola (EVERSMANN, 1834) Darevskia schaekeli AHMADZADEH et al. 2013 Darevskia spitzenbergerae (EISELT, DAREVSKY & SCHMIDTLER, 1992) Darevskia steineri (EISELT, 1995) Darevskia szczerbaki (LUKINA, 1963) Darevskia tuniyevi ARRIBAS et al., 2022 Darevskia unisexualis (DAREVSKY, 1966) Darevskia uzzelli (DAREVSKY & DANIELYAN, 1977) Darevskia valentini (BOETTGER, 1892) |
| Caucasilacerta Harris, Arnold & Thomas, 1998 has been considered as a nomen nudum during the last 18 years (Arnold et al., 2007). Recently a paper by Busack et al. (2016) suggest that the publication of Arribas (1997) in microfiche is not valid, and that Caucasilacerta Harris, Arnold and Thomas, 1998 is valid, and has priority over Darevskia Arribas, 1999. In fact Busack et al. (2016) were unaware that a case was submitted to the ICZN to validate the Arribas (1997) publication in microfiche, which was perfectly valid when published (according the 1985 version of the Code), however not according the current version (ICZN, 2000 and actualizations) that only allow publication in paper and optical disks, with retroactive effect. If a case is accepted for consideration, Art 82.1 of the Code protects the widespread used nomen, until the Comission reach a verdict. A paper has been submitted which will reply to Busack et al. (2016), explaining in detail why Caucasilacerta is really a nomen nudum (Arribas, 2016).
Osсar Arribas, Igor Doronin
The International Commission on Zoological Nomenclature has conserved the names Iberolacerta and Darevskia for two genera of lacertid lizards by confirming their availability from Arribas (1999). The microfiche publication (Arribas, 1997) is
confirmed as unavailable and placed on the Official Index of Rejected and Invalid Works in Zoological Nomenclature. The name Caucasilacerta Harris, Arnold & Thomas, 1998 is confirmed as a nomen nudum and placed on the Official Index of Rejected and Invalid Generic Names in Zoology. |
Uzzell, T. & Darevsky, I.S. (1975) - Biochemical evidence for the hybrid origin of the parthenogenetic species of Lacerta saxicola complex (Sauria, Lacertidae) with a discussion of some ecological and evolutionary implications. - Copeia, 1975 (2): 204-222.  × Five proteins (mannosephosphate isomerase, glucosephosphate isomerase, lactate dehydrogenase, creatine kinase, hemoglobin) were examined for six bisexual Transcaucasian taxa related to Lacerta saxicola (Lacerta valentini, L. portschinskii, L. raddei raddei, L. r. nairensis, L. mixta, L. parvula) and four unisexual taxa (Lacerta armeniaca, L. dahli, L. rostombekovi, L. unisexualis). Heterozygosity in the bisexual taxa was very limited for this sample of proteins. All individuals examined of each unisexual were heterozygous at loci specifying at least two of these proteins (L. armeniaca: mannosephosphate isomerase and creatine kinase; L. dahli, mannosephosphate isomerase and hemoglobin; L. rostombekovi, glucosephosphate isomerase, mannosephosphate isomerase and hemoglobin; L. unisexualis, glucosephosphate isomerase, mannosephosphate isomerase and creatine kinase). High levels of heterozygosity in unisexuals appears to result from the hybrid origin of the unisexuals. The exact combinations of alleles present in unisexuals would readily result from certain crosses among the bisexuals, and from no others. The first point supports the hypothesis that hybridization accounts for the observed heterozygosity, the second identifies, biochemically at least, the probable parents. On these biochemical grounds, L. armeniaca arose by hybridization of L. valentini and L. mixta; L. unisexualis from L. valentini and L. r. nairensis; L. rostombekovi from L. r. raddei and L. portschinskii; and L. dahli from L. portschinskii and L. mixta. With regard to altitudinal distribution, vegetational associations and geographic distribution, each unisexual species appears to be intermediate between its putative parental species. L. armeniaca, L. unisexualis and L. rostombekovi all live in drier situations than either parental species. L. dahli is an exception since it occupies slightly more moist habitats than L. portschinskii. Biogeographical considerations appear to place the age of L. unisexualis and L. rostombekovi at greater than 5000 years. It is possible that their occupancy of more extreme habitats than their parental species represents a relic ecology, reflecting the adaptations of the parental bisexual species when the unisexual species arose, rather than a weed habitat into which they moved to escape from competition with their parental species. The fixed heterozygosity of the unisexual species of Lacerta indicates that the restitution of somatic diploidy results either from a premeiotic endoduplication without cytokinesis, or from fusion of the female pronucleus with one of the second division meiotic products of the first polar nucleus. The first mechanism is inconsistent with the number of bivalents reported, the second is otherwise unknown in organisms in which the first polar nucleus becomes a polar body. The bisexual taxa are treated as several distinct species, four pairs of which are partly sympatric with little or no hybridization. Morphological and ecological differences between the other taxa are so great that there seems little question about specific distinctness. The degree of biochemical and morphological difference between these non-sympatric taxa is as great as that between those that do occur sympatrically without fusing. The formation of parthenogenetic species as a result of past hybridization between some pairs indicates a great selective disadvantage to these pairs of hybridizing, and is thus compelling evidence that the two members of each pair are not conspecific. Moritz, C. & Uzzell, T. & Spolsky, C. & Hotz, H. & Darevsky, I.S. & Kupriyanova, L. & Danielyan, F. (1992) - The maternal ancestry and approximate age of parthenogenetic species of Caucasian rock lizards (Lacerta: Lacertidae). - Genetica, The Hague, 87 (1): 53-62.  × Restriction enzymes were used to assay variation among mitochondrial DNAs from parthenogenetic and sexual species of Lacerta. This permitted identification of the sexual species that acted as the maternal parent of the various hybrid-parthenogenetic lineages. Lacerta mixta was the maternal parent for both L. dahli and L. armeniaca, L. valentini was the maternal parent for L. uzzelli, and L. raddei was the maternal parent of L. rostombekovi. The maternal ancestry of L. unisexualis is not as clear. The sample of L. nairensis was very similar to one from a population of L. raddei and either species could be the maternal parent of L. unisexualis. The parthenogenetic species all had very low nucleotide diversity in absolute terms and in comparison to their sexual relatives. The close similarity between mtDNAs from the parthenogenetic species and their respective sexual maternal ancestor species provides strong evidence for the recent origin of the parthenogens. The low diversity of the parthenogens indicates that few females were involved in their origins; the maternal parents of L. dahli and L. armeniaca could have come from a single population. The patterns of mtDNA variation in Lacerta are very similar to those in Cnemidophorus and Heteronotia, establishing recent and geographically restricted origins as a general feature of parthenogenetic lizards. Arribas, O.J. (1997) - Morfología, filogenia y biogeografía de las lagartijas de alta montaña de los Pirineos. - Ph.D. Thesis. Universidad Autónoma de Barcelona. 353 pp. (8 pp in paper and 353 pp in microfiche). Publications Universitat Autònoma Barcelona. Bischoff, W. (2003) - Die Eidechsenfauna Georgiens. Teil II. Die Gattung Darevskia. - Die Eidechse, Bonn, 14 (3): 65-93. × In Georgia, 16 species of rock lizards of the Genus Darevskia are occuring (D. alpina, D.
„armeniaca`, D. brauneri, D. caucasica, D. clarkorum, D. daghestanica, D. „dahli`, D.
derjugini, D. mixta, D. nairensis, D. parvula, D. portschinskii, D. praticola, D. rudis, D.
„unisexualis` and D. Valentini). Besides short presentations of the single species and hints on their distribution and habitats, also some systematic remarks are given. Arnold, E.N. & Arribas, O. & Carranza, S. (2007) - Systematics of the Palaearctic and Oriental lizard tribe Lacertini (Squamata: Lacertidae: Lacertinae), with descriptions of eight new genera. - Zootaxa, 1430: 1-86. × DNA sequence indicates the Lacertidae contain two subfamilies, Gallotiinae and Lacertinae, the latter comprising two
monophyletic tribes, the Eremiadini of Africa and arid southwest and central Asia, and the Lacertini of Europe, northwest
Africa and southwest and east Asia. Relationships within the 108 species of Lacertini are explored using mtDNA
(291 bp cytochrome b; 329 bp 12S rRNA for 59 nominal species, and reanalysis of the data of Harris et al. 1998, and Fu
2000). The morphology of the tribe is reviewed and 64 of its characters (equivalent to 83 binary ones) also used to assess
relationships. The Lacertini are assigned to 19 monophyletic units of 1 to 27 species, recognised here as the following
genera (contents are indicated in brackets): Algyroides, Anatololacerta gen. nov. (L. danfordi group), Apathya (L. cappadocica
group), Archaeolacerta (L. bedriagae), Dalmatolacerta gen. nov. (L. oxycephala), Darevskia (L. saxicola group),
Dinarolacerta gen. nov. (L. mosorensis), Hellenolacerta gen. nov. (L. graeca), Iberolacerta (L. monticola group), Iranolacerta
gen. nov. (L. brandtii and L. zagrosica), Lacerta s. str. (sand and green lizards, L. agilis group), Parvilacerta gen.
nov. (L. parva and L. fraasii), Phoenicolacerta gen. nov. (L. laevis group), Podarcis (wall lizards), Scelarcis (L. perspicillata),
Takydromus (Asian grass lizards), Teira (L. dugesii), Timon (ocellated lizards, L. lepida group) and Zootoca (L.
vivipara). Both mtDNA and morphology indicate that Lacerta and Timon are sister taxa, and DNA suggests further possible
relationships among genera (Fig. 1, p. 6). Neither DNA nor morphology indicates that the archaeolacertas (sometimes
formalised as Archaeolacerta sens. lat.) form a clade. Instead, they are representatives of an ecomorph associated
with living on rock exposures and using the narrow crevices that these contain.
The Lacertidae probably arose in the European area, with the Gallotiinae later reaching Northwest Africa and the
Canary Islands, and the ancestor of the Eremiadini invading Africa in the mid-Miocene. The Lacertini spread through
much of their present European range and diversified, perhaps largely by repeated vicariance, around 12–16 My ago,
producing the ancestors of the present mainly small-bodied genera, which then underwent often modest speciation. Three
units spread more widely: the Lacerta-Timon clade of large-bodied lizards probably dispersed earliest, followed by Algyroides
and then Podarcis. Overall, European Lacertidae show a pattern of repeated spread, often accompanied by restriction
of previous groups. Expansion of Lacertini may have displaced earlier lacertid lineages from all or much of Europe;
while spread of Podarcis may have restricted many other genera of Lacertini. The earlier expansion of the Lacerta-Timon
clade probably did not have this effect, as difference in adult body size restricted competitive interaction with other
forms. Several invasions of more distant areas also occurred: of East Asia by Takydromus over 10 My ago, and more
recently of northwest Africa by Podarcis, Scelarcis and Timon, and Madeira by Teira.
Relationships within the Eremiadini estimated from both mtDNA, and nDNA differ considerably from those based
on morphology. They indicate relatively mesic forms may have diversified widely across Africa and given rise to at least
three independent invasions of arid habitats. MtDNA also indicates that Lacerta andreanskyi belongs in the Eremiadini
and may occupy a basal position there. It is assigned to a further new genus, Atlantolacerta gen. nov. Ahmadzadeh, F. & Flecks, M. & Carretero, M.A. & Mozaffari, O. & Böhme, W. & Harris, D.J. & Freitas, S. & Rödder, D. (2013) - Cryptic speciation patterns in Iranian Rock Lizards uncovered by integrative taxonomy. - PLoS ONE 8 (12): e80563. × While traditionally species recognition has been based solely on morphological differences either typological or quantitative, several newly developed methods can be used for a more objective and integrative approach on species delimitation. This may be especially relevant when dealing with cryptic species or species complexes, where high overallresemblance between species is coupled with comparatively high morphological variation within populations. Rock lizards, genus Darevskia, are such an example, as many of its members offer few diagnostic morphological features. Herein, we use a combination of genetic, morphological and ecological criteria to delimit cryptic species within two species complexes, D. chlorogaster and D. defilippii, both distributed in northern Iran. Our analyses are based on molecular information from two nuclear and two mitochondrial genes, morphological data (15 morphometric, 16 meristic and four categorical characters) and eleven newly calculated spatial environmental predictors. The phylogeny inferred for Darevskia confirmed monophyly of each species complex, with each of them comprising several highly divergent clades, especially when compared to other congeners. We identified seven candidate species within each complex, of which three and four species were supported by Bayesian species delimitation within D. chlorogaster and D. defilippii, respectively. Trained with genetically determined clades, Ecological Niche Modeling provided additional support for these cryptic species. Especially those within the D. defilippii-complex exhibit well-differentiated niches. Due to overall morphological resemblance, in a first approach PCA with mixed variables only showed the separation between the two complexes. However, MANCOVA and subsequent Discriminant Analysis performed separately for both complexes allowed for distinction of the species when sample size was large enough, namely within the D. chlorogaster-complex. In conclusion, the results support four new species, which are described herein. Arribas, O. & Candan, K. & Kornilios, P. & Ayaz, D. & Kumlutas, Y. & Gül, S. & Yilmaz, C. & Yildirim Caynak, E. & Ilgaz, C. (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. - Zootaxa, 5224 (1): 1-68. × Revealing biodiversity allows the accurate determination of the underlying causes of many biological processes such as speciation and hybridization. These processes contain many complex patterns, especially in areas with high species diversity. As two of the prominent zoogeographic areas, Anatolia and Caucasus are also home to the genus Darevskia, which has a complex morphological structure and parthenogenetic speciation. Darevskia valentini and D. rudis are two largely distributed taxa of this genus, both of which have a controversial taxonomic delimitation. Here we performed both a highly detailed morphological comparison and a molecular evaluation for the populations in both species groups. The most comprehensive taxonomic revision of this complex was carried out to determine the cases where the data obtained were compatible or not with each approach. As a result of the obtained outputs, it seems that D. spitzenbergerae stat. nov., D. mirabilis stat. nov. and D. obscura stat. nov. should be accepted as the species level, this later with subspecies D. o. bischoffi comb. nov. and D. o. macromaculata comb. nov.. Also, we propose two new taxa: D. josefschmidtleri sp. nov. and D. spitzenbergerae wernermayeri ssp. nov.. It has also been shown that “lantzicyreni” subspecies belong to D. rudis instead of D. valentini. The extensive revision has contributed to subsequent studies to more accurately understand the past histories of species in the genus Darevskia.
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