Darevskia

Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Cladus: Craniata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Lepidosauromorpha
Superordo: Lepidosauria
Ordo: Squamata
Cladus: Unidentata Episquamata

Subordo: Lacertoidea
Infraordo: Lacertibaenia

Familia: Lacertidae
Subfamilia: Lacertinae
Genus: Darevskia
Species (33): D. aghasyani – D. alpina – D. armeniaca – D. bendimahiensis – D. bithynica – D. brauneri – D. caspica – D. caucasica – D. chlorogaster – D. clarkorum – D. daghestanica – D. dahli – D. defilippii – D. derjugini – D. dryada – D. kamii – D. kopetdaghica – D. lindholmi – D. mixta – D. parvula – D. portschinskii – D. praticola – D. raddei – D. rostombekovi – D. rudis – D. sapphirina – D. saxicola – D. schaekeli – D. steineri – D. szczerbaki – D. unisexualis – D. uzzelli – D. valentini
Name

Darevskia Arribas, 1999: 17 [conserved name]

Type species: Lacerta saxicola Eversmann, 1834, by original designation.
Gender: feminine.
Etymology: named after Ilya Sergeevich Darevsky.
Placed on the Official List of Generic Names in Zoology by Opinion 2461 (2020: 135).

Synonymy

Darevskia Arribas, 1997: 305 [unavaliable name by Opinion 2461 (2020: 135)]
Type species: Lacerta saxicola Eversmann, 1834, by original designation.
Caucasilacerta Harris, Arnold & Thomas, 1998: 1947 [nomen nudum fide Arribas (2016: 306) and Opinion 2461 (2020: 135)]
Type species: Lacerta saxicola Eversmann, 1834, by original designation.
Placed on the Official Index of Rejected and Invalid Generic Names in Zoology by Opinion 2461 (2020: 135).

References
Primary references

Arribas, O.J. 1997. Morfología, filogenia y biogeografía de las lagartijas de alta montaña de los Pirineos. Publications Universitat Autònoma Barcelona: Barcelona, Spain. 353 pp. Reference page.
Harris, D.J., Arnold, E.N. & Thomas, R.H. 1998. Relationships of lacertid lizards (Reptilia: Lacertidae) estimated from mitochondrial DNA sequences and morphology. Proceedings of the Royal Society B: Biological Sciences 265(1409): 1939–1948. DOI: 10.1098/rspb.1998.0524 Paywall Reference page.
Arribas, O.J. 1999. Phylogeny and relationships of the mountain lizards of Europe and Near East (Archaeolacerta Mertens, 1921, sensu lato) and their relationships among the Eurasian lacertid radiation. Russian Journal of Herpetology 6(1): 1–22. DOI: 10.30906/1026-2296-1999-6-1-1-22 [nonfunctional] Broken access Reference page.
International Commission on Zoological Nomenclature. 2020. Opinion 2461. (Case 3711) – Iberolacerta Arribas, 1999 and Darevskia Arribas, 1999 (Chordata, Squamata, Lacertidae): names confirmed as available. Bulletin of Zoological Nomenclature 77: 135–137. DOI: 10.21805/bzn.v77.a043 Paywall Reference page.

Additional references

Fu, J.-Z., Murphy, R.W. & Darevsky, I.S. 1997. Toward the phylogeny of Caucasian rock lizards: implications from mitochondrial DNA gene sequences (Reptilia: Lacertidae). Zoological Journal of the Linnean Society 120(4): 463–477. DOI: 10.1111/j.1096-3642.1997.tb01283.x Open access Reference page.
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. Abstract & excerpt. Reference page.
Kosushkin, S.A. & Grechko, V.V. 2013. Molecular genetic relationships and some issues of systematics of rock lizards of the genus Darevskia (Squamata: Lacertidae) based on locus analysis of SINE-type repeats (Squam1). Russian Journal of Genetics 49(8): 857–869. DOI: 10.7868/S0016675813070084 Paywall Reference page.
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. DOI: 10.1371/journal.pone.0080563 Reference page.
Arribas, O.J. 2016. Why Caucasilacerta Harris, Arnold et Thomas, 1998 is a nomen nudum? Russian Journal of Herpetology 23(4): 305–306. DOI: 10.30906/1026-2296-2016-23-4-305-306 [nonfunctional] Broken access Reference page.
Busack, S.D., Salvador, A., Bauer, A.M. & Kaiser, H. 2016. Darevskia and Iberolacerta (Reptilia, Lacertidae): Arribas, 1997 or 1999? The correct dating of two nomenclatural acts affecting Palearctic lizards, and validation of the name Caucasilacerta Harris, Arnold & Thomas, 1998. Bionomina 10: 61–73. DOI: 10.11646/bionomina.10.1.4 Paywall Reference page.
Arribas, O.J., Ananjeva, N.B., Carranza, S., Doronin, I.V., Harris, D.J., Orlov, N.L. & Orlova, V.F. 2017. The pernicious effect of retroactive changes in the Code: Darevskia and nomenclatorial stability, a reply to Busack et al. (2016). Basic and Applied Herpetology 31: 125-129. DOI: 10.11160/bah.75 Open access Reference page.
Arribas, O.J., Ananjeva, N.B., Carranza, S., Doronin, I.V., Orlov, N.L. & Orlova, V.F. 2018. Case 3711 – Iberolacerta Arribas and Darevskia Arribas (Chordata, Squamata, Lacertidae): proposals to deem these names available either from Arribas (1997) or from Arribas (1999). Bulletin of Zoological Nomenclature 75(1): 122–129. DOI: 10.21805/bzn.v75.a026 Paywall Reference page.
Tuniyev, B.S. & Petrova, T.V. 2019. A new lizard species of the genus Darevskia Arribas, 1997 from Southern Armenia. Proceedings of the Zoological Institute RAS 323(2): 136–148. DOI: 10.31610/trudyzin/2019.323.2.136 Open access Reference page.

Links

Uetz, P. & Hallermann, J. 2022. Darevskia . The Reptile Database. Accessed on 10 August 2021.

Darevskia is a genus of wall lizards of the family Lacertidae living in the Caucasus, Iran and Turkey in forest and grassy habitats with numerous rock outcrops. Among rock lizards, 7 parthenogenetic species are known.

Description

These are small lizards with a body length of 50–85 mm and with about two times longer tail. The body is usually flattened, the head is pointed in shape and in most species flattened in a vertical plane, which allows lizards to hide in narrow crevices between stones and rocks. Rock lizards have relatively long legs with special calluses on the inner surfaces of the paws and sharp claws, thanks to which they quickly move along the vertical rough surfaces of rocks and stones.

The color of rock lizards varies from different tones of green to sand. Females are usually colored paler than males. On the dorsal side of the body, rock lizards have an occipital stripe composed of a set of black or brown spots and a wide line of the lizard's main color, and dark patterns on the sides of the body. In some species, blue or violet spots with white circles in the center and / or monotonous blue-violet spots at the junction of the abdominal scutes with trunk scales are located on the anterior third of the body. Most types of rock lizards are characterized by a diverse color of the abdominal side of the body, ranging from various shades of pink, red and orange to yellow and green.
Etymology

The generic name, Darevskia, is in honor of Russian herpetologist Ilya Sergeyevich Darevsky.[2]

In 1830, Professor of Kazan University E. A. Eversman (1794-1860) made an expedition to the North Caucasus, as a result of which he described two new species: a meadow lizard (Lacerta praticola (Eversman 1834)) and a rock lizard (Lacerta saxicola (Eversman 1834)). At that time, European scientists did not recognize the independence of the species L. saxicola, considering it as part of the European L. muralis (Laurenti 1768). But at the beginning of the XX century between the two zoologists Mecheli (1862–1953) and Boulanger (1858–1937) there was a lengthy discussion about the taxonomic position of L. saxicola, which was resolved in favor of the former, and L. saxicola and subspecies were further considered separately from L. muralis. For several decades, scientists from different countries studied the monophyletic group of rock lizards independently, highlighting new subspecies, simplifying some taxa and describing new species. It was on this group of lizards that the phenomenon of parthenogenesis in amniotic vertebrates was first discovered by the Russian zoologist I. S. Darevsky (1924-2009), who also made a significant contribution to understanding the ecology, systematics, and morphology of rock lizards. And in 1997, the Spanish scientist Arribas named the genus of rock lizards Darevskia and identified the type species D. saxicola.
Taxonomy

According to Arribas (1997), the genus Darevskia includes four groups (clades), combining species by origin and kinship: “raddei”, “rudis”, “saxicola” and “caucasica”. Subsequently, three more clades were identified: “praticola”, “chlorogaster”, and “defilippii”. In total, the genus includes 33 species, 7 of which breed parthenogenetically, and 22 subspecies.

Raddei	Rudis	Saxicola	Caucasica
Darevskia raddei (Boettger, 1892)	Darevskia rudis (Bedriaga, 1886)	Darevskia saxicola (Eversmann, 1834)	Darevskia caucasica (Mehely, 1909)
Darevskia nairensis (Darevsky, 1967)	Darevskia parvula (Lantz & Cyren, 1913)	Darevskia brauneri (Mehely, 1909)	Darevskia alpina (Darevsky, 1967)
Darevskia bithynica (Mehely, 1909)	Darevskia valentini (Boettger, 1892)	Darevskia sczerbaki (Lukina, 1963)	Darevskia daghestanica (Darevsky, 1967)
Darevskia clarkorum (Darevsky & Vedmederja, 1977)	Darevskia portschinskii (Kessler, 1878)	Darevskia lindholmi (Lantz & Cyren, 1936)	Darevskia derjugini (Nikolsky, 1898)
Darevskia mixta (Mehely, 1909)
Darevskia dryada (Darevsky & Tuniyev, 1997)


Praticola	Chlorogaster	Defilippii	Parthenogenetic species
Darevskia praticola (Eversmann, 1834)	Darevskia chlorogaster (Boulenger, 1908)	Darevskia defilippii (Camerano, 1877)	Darevskia armeniaca (Mehely, 1909)
	Darevskia kamii Ahmadzadeh, Flecks, Carretero, Mozaffari, Bohme, Harris, Freitas & Rodder, 2013	Darevskia kopetdaghica Ahmadzadeh, Flecks, Carretero, Mozaffari, Bohme, Harris, Freitas, Rodder, 2013	Darevskia bendimahiensis (Schmidtler, Eiselt & Darevsky, 1994)
	Darevskia caspica Ahmadzadeh, Flecks, Carretero, Mozaffari, Bohme, Harris, Freitas & Rodder, 2013	Darevskia schaekeli Ahmadzadeh, Flecks, Carretero, Mozaffari, Bohme, Harris, Freitas & Rodder, 2013	Darevskia dahli(Darevsky, 1957)
		Darevskia steineri (Eiselt, 1995)	Darevskia rostombekowi (Darevsky, 1957)
			Darevskia unisexualis (Darevsky, 1966)
			Darevskia uzzelli (Darevsky & Danielyan, 1977)
			Darevskia sapphirina (Schmidtler, Eiselt & Darevsky, 1994)

Species

The following species are recognized as being valid.[3]

Darevskia aghasyani Tuniyev & Petrova, 2019
Darevskia alpina (Darevsky, 1967)
Darevskia armeniaca (Méhelÿ, 1909) - Armenian lizard
Darevskia bendimahiensis (Schmidtler, Eiselt & Darevsky, 1994)
Darevskia bithynica (Méhelÿ, 1909)
Darevskia brauneri (Méhelÿ, 1909) - Brauner's rock lizard
Darevskia caspica Ahmadzadeh et al., 2013
Darevskia caucasica (Méhelÿ, 1909)
Darevskia chlorogaster (Boulenger, 1908) - greenbelly lizard
Darevskia clarkorum (Darevsky & Vedmederja, 1977)
Darevskia daghestanica (Darevsky, 1967)
Darevskia dahli (Darevsky, 1957)
Darevskia defilippii (Camerano, 1877) - Elburs lizard
Darevskia derjugini (Nikolsky, 1898)
Darevskia dryada (Darevsky & Tuniyev, 1997) - Charnali lizard
Darevskia kamii Ahmadzadeh et al., 2013
Darevskia kopetdaghica Ahmadzadeh et al., 2013
Darevskia lindholmi (Szczerbak, 1962)
Darevskia mixta (Méhelÿ, 1909)
Darevskia parvula (Lantz & Cyrén, 1913) - red-bellied lizard
Darevskia portschinskii (Kessler, 1878)
Darevskia praticola (Eversmann, 1834) - meadow lizard
Darevskia raddei (Boettger, 1892) - Azerbaijan lizard
Darevskia rostombekowi (Darevsky, 1957)
Darevskia rudis (Bedriaga, 1886)
Darevskia sapphirina (Schmidtler, Eiselt & Darevsky, 1994)
Darevskia saxicola (Eversmann, 1834)
Darevskia schaekeli Ahmadzadeh et al., 2013
Darevskia steineri (Eiselt, 1995) - Steiner's lizard
Darevskia szczerbaki (Lukina, 1963) - Szczerbak's lizard
Darevskia unisexualis (Darevsky, 1966)
Darevskia uzzelli (Darevsky & Danielyan, 1977)
Darevskia valentini (Boettger, 1892) - Caucasian rock lizard, Valentin's lizard

Nota bene: A binomial authority in parentheses indicates that the species was originally described in a genus other than Darevskia.
Distribution

Rock lizards are common in Abkhazia, Azerbaijan, Armenia, Georgia, Iran, Nagorno-Karabakh, Russia (Adygea, Dagestan, Ingushetia, Kabardino-Balkaria, Karachay-Cherkessia, Krasnodar Territory, Republic of Crimea, North Ossetia-Alania, Stavropol Territory and Chechnya), in Turkey and South Ossetia. It is worth noting that the boundaries of the range of some species are not reliably known, but the expected areas of their encounters coincide with the already indicated distribution sites for the whole genus
Habitat

Rock lizards are found in various high-altitude zones from 0 to 3000 meters above sea level and occupy a variety of landscapes: mountain-steppe, forest-steppe, mountain meadow, mountain forest, anthropogenic and coastal. By confinement to one or another habitat, they can be conditionally divided into several groups:

1) Lizards living in the forest, according to the occupied microreliefs, are divided into: adhering to rock outcrops (D. raddei, D. brauneri) and independent of them, able to live in habitats in which there are no rocks, using rodent burrows as shelters, deciduous litter, cavities in trees and bark (e.g. D. chlorogaster, D. armeniaca).

2) Inhabitants in areas of bedrock outcrops and clayey cliffs in alpine and subalpine meadows. As shelters, they often use rodent burrows, cavities between stones and cracks in the rocks. These habitats are adhered to by D. alpine and D. mixta, D. armeniaca and D. valentini.

3) Rock lizards of dry and moderately dry landscapes (alpine steppes) of rocks and their feet on slopes with dry-loving shrubbery and grassy vegetation, road slopes. Such habitats have a large number of crevices and voids serving as shelters for lizards. Such species are found in such landscapes as: D. rudis, D. portschinski, D. daghestanica, D. raddei, D. saxicola.

4) Occupying anthropogenic habitats: an abandoned building, walls in cities, abandoned temples, monasteries, etc., where their number often exceeds that in natural habitats. For example, D. armeniaca, D. lindholmi, D. dahli.

Rock lizards are found at heights of 0 – 3000 m above sea level. Zonal and geographical distribution is determined by the amount of precipitation, average annual temperature, duration of the adverse season, and exposure of the slope. For example, D. daghestanica on the southern slope of the Greater Caucasus Mountain Range (South Ossetia) is distributed at altitudes of 1500–1800 meters above sea level, and on the northern slopes (Dagestan) - 50 – 2100 meters above sea level.
Nutrition

Rock lizards feed on various invertebrates with a body size from a few millimeters to 4 cm: spiders, dipterous, lepidoptera, hymenoptera, cockroaches, orthoptera, semi-rigid-winged, coleoptera, wood lice, worms, slugs, marine and freshwater amphipods, caddis flies, chironomids and sometimes parts of plants.. Also, particular cases of cannibalism were recorded when adults ate juvenile individuals.

Despite the diversity of the food supply, rock lizards can develop preferences for feeding invertebrates of a certain group (for example, flying forms of ants), which is caused by seasonal changes in the availability or abundance of this type of prey. Even after a significant reduction in the density of invertebrates of this group, lizards continue to hunt for some time in the presence of more accessible food objects.
Population density and spatial structure of the population

Rock lizards are extremely rare on their own, usually forming settlements. The population density of parthenogenetic rock lizards can vary in a wider range than that of bisexual species: up to 200 individuals per 1 km of the route in unisexual species and up to 80 individuals in bisexual species, which is explained by the fact that parthenogenetic species are less aggressive and have a high population growth rate.

Rock lizards are characterized by complex and diverse social systems, which, in particular, are characterized by stable long-term friendly relations between the male and the female and territorial or hierarchical relations between individuals of the same sex.

The basis of the settlements of bisexual rock lizards are sedentary males and females with individual sites, often overlapping. In a number of species, some males possess territories protected from other males. The territories of males never overlap, but their centers of activity, primarily associated with basking, coincide with the centers of activity of females that live within their territories.

The study of the social behavior and spatial structure of rock lizard populations has been the subject of a number of scientific papers published based on the results of many years of research
Activity

The seasonal activity of rock lizards is determined by temperature indicators and, therefore, species living at different heights differ in terms of exit from wintering shelters, mating period, laying of eggs, hatching of young individuals and time of leaving for wintering. Around the end of February until the end of May, exit from wintering shelters occurs, and the active period is from 6 – 7 months (in the mountains) and up to 9 – 10 months (in the valleys and on the seashore). During this period, mature individuals mate, and females lay eggs. Hibernation takes place from late September to mid November.

The beginning and end of the daily activity of the lizard is determined by the lighting conditions within the individual area, and in some individuals it can start early in the morning, while individuals living on the slopes of the northern exposure or in deep forest valleys are active for several hours in the middle of the day [4]. After heating (basking), the body temperature of the lizard reaches about 30 - 34 °C, and it begins a routine activity aimed at supporting the body . Late in the evening, when the heat subsides, the animals return to the basking places and stay there for some time, after which they go to their night shelters.
Reproduction

Rock lizards reach maturity in the second (females) and third (males) year of life with a total life expectancy of up to 13 years. Mating of some types of rock lizards occurs after the first molt, approximately 3–5 weeks after leaving wintering (D. brauneri), in others, immediately after leaving wintering shelter (D. valentini). Egg laying begins in the second half of June and lasts until early August. In clutch usually 2 to 7 eggs. The incubation period lasts approximately 55 – 65 days. Young animals with a body length of about 25 mm are born in late summer.
The evolution and origin of parthenogenetic species of Darevskia

Separation of the genus Darevskia from the subfamily Lacertinae occurred approximately 12-16 million years ago, in the middle of the Miocene. Presumably, the ancestral form of the genus Darevskia penetrated the territory of the Western Caucasus in the Middle Miocene or in the Middle Pliocene, when there was a land connection between Asia Minor and the Balkans.

The genus of rock lizards has 7 parthenogenetic species: D. armeniaca, D. bendimahiensis, D. dahli, D. rostombekovi, D. sapphirina, D. unisexualis and D. uzzelli, resulting from the hybridization of males and females of different bisexual species. Moreover, D. valentini and D. portschinskii always enter as paternal species, and D. mixta, D. nairensis, D. raddei always enter as maternal species. From the point of view of the classical concept of a species, homosexual taxa cannot be assigned a species status due to the lack of exchange of genetic material between individuals within the same population. However, on the basis of morphological and cytogenetic characters, they are assigned a species rank.

The appearance of parthenogenetic species is usually associated with the last, Wurm glaciation. Parthenogenetic rock lizards appeared about 10 thousand years ago when, due to the formation of mountain glaciers, the habitats of the parent bisexual species were disturbed, which led to the overlap of their ranges and the hybridogenic formation of parthenogenetic individuals better adapted to the conditions of short summers and long winters. Due to the doubled reproduction rate and successful resettlement, same-sex populations later began to exist independently of the parent species.

Hybridization occurs in places where the ranges of parthenogenetic and bisexual species overlap, resulting in the appearance of triploid sterile females and a small number of males, with changes in the reproductive system (malfunctions in the formation of reproductive products, hermaphroditism). Nevertheless, presumably, these males are able to give offspring, which leads to the appearance of tetraploid hybrids. Sometimes males emerge from unfertilized eggs laid by parthenogenetic females. Like males of hybrid origin, they have reproductive disorders, which may not interfere with their offspring. A small amount of parthenogenetic female males emerging from clutches is explained by frequently occurring mutations incompatible with life.

Despite the fact that as a result of parthenogenesis, individuals are born that receive hereditary material only from the mother's body, a small intraspecific genetic diversity has been revealed due to mutational processes, genetic instability and the appearance of same-sex species as a result of repeated and independent crossing of parental species among themselves.
See also

Cnemidophorus, another lizard genus containing several parthenogenic species.

References

Ananjeva, Natalia B.; Orlov, Nikolai L.; Khalikov, Roman G.; Darevsky, Ilya S.; Ryabov, Sergei A.; Barabanov, Andrei V. (2006), The Reptiles of Northern Eurasia: Taxonomic Diversity, Distribution, Conservation Status, Faunastica, vol. 47, Sofia, Bulgaria: Pensoft Publishers, p. 79, ISBN 954-642-269-X.
Beolens, Bo; Watkins, Michael; Grayson, Michael (2011). The Eponym Dictionary of Reptiles. Baltimore: Johns Hopkins University Press. xiii + 296 pp. ISBN 978-1-4214-0135-5. (Darevskia, p. 65).

"Darevskia ". The Reptile Database. https://www.reptile-database.org.