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Literature- and poster projects
of the real lizards, family Lacertidae
Darevskia ARRIBAS, 1999
Arakelyan, M. & Spangenberg, V. & Petrosyan, V. & Ryskov, A. & Kolomiets, O. & Galoyan, E. (2022) -
Despite numerous works devoted to hybrid origin of parthenogenesis in reptiles, the causes of hybridization between different species, resulting in the origin of parthenogenetic forms, remain uncertain. Recent studies demonstrate that sexual species considered parental to parthenogenetic rock lizards (Darevskia spp.) avoid interspecific mating in the secondary overlap areas. A specific combination of environmental factors during last glaciation period was critical for ectotherms, which led to a change in their distribution and sex ratio. Biased population structure (e.g., male bias) and limited available distributional range favored the deviation of reproductive behavior when species switched to interspecific mates. To date, at least 7 diploid parthenogenetic species of rock lizards (Darevskia, Lacertidae) originated through interspecific hybridization in the past. The cytogenetic specifics of meiosis, in particular the weak checkpoints of prophase I, may have allowed the formation of hybrid karyotypes in rock lizards. Hybridization and polyploidization are 2 important evolutionary forces in the genus Darevskia. At present, throughout backcrossing between parthenogenetic and parental species, the triploid and tetraploid hybrid individuals appear annually, but no triploid species found among Darevskia spp. on current stage of evolution. The speciation by hybridi- zation with the long-term stage of diploid parthenogenetic species, non-distorted meiosis, together with the high ecological plasticity of Caucasian rock lizards provide us with a new model for considering the pathways and persistence of the evolution of parthenogenesis in vertebrates.
Abrahamyan, M.R. & Petrosyan, R.K. & Galoyan, E.A. & Danielyan, F.D. & Arakelyan, M.S. (2014) -
The patterns of seasonal and daily activities of two parthenogenetic species of lizards (Darevskia armeniaca, D. unisexualis) sharing the same territories in the zone of hybridization in central region of Armenia demonstrate ecologically similar potential niches.
Abramjan, A. (2008) -
Parthenogenesis in reptiles was fort he first time discovered in the Caucasian rock lizards of the genus Darevskia (family Lacertidae). There re recognized 7 all-female diploid species occupying rocky habitats of the Armenian Highland. All oft hem (as well as other parthenogenetoc reptiles) originated from hybridisation between bisexual species and display high levels of heterozygosity. The hybridisations most probably occurred after the last glaciations period. It was found out that maternal, respectivily paternal species are phylogenetically related and that most probably the origian of parthenogenesis is Phylogenetically constrained in these lizards. The role of sex chromosomes, determination of sex and other possible factors determining the origin of parthenogenesis are also discussed. The paerthenogenetic Darevskia and other unisexual reptiles have in common relatively young age and very little genetic variability. However, some hyperunstable microsatellite loci were also detected. The evolutionary perspective of parthenogens doesn´t seem tob e long-termed, but there have been observed also peculiarities concerning adaptability of these species. Some questions like the meiotic mechanisms, maintenance of high heterozygosity or effects of epigenetic factors on clonal species aren´t yet fully resolved.
Abramjan, A. (2011) -
Parthenogenetic lizards of the genus Darevskia as an evolutionary model.
Several parthenogenetic lineages occur within the lizards of the genus Darevskia (Sauria: Lacertidae) which are endemic to southern Transcaucasus. High level of heterozygosity, cause by thein hybrid origin, is one of the crucial aspects of thein evolutionary potential, as well as the asexual reproduction. Heterosis on one side is in the opposition to the outbreeding depression and genetic uniformity of the clones on the other side. Aim of this work is to evaluace if these aspects influence viability of parthenogenetic species and differ them from the sexual ones. We chose the amount of asymmetries as a measure of developmental instability, which we studied on three meristic characters. We also evaluated the pattern of asymmetries in lateral blue spots, which are of signaling importace in lacertid lizards. Our results suggest that there isn`t significant difference between parthenogenetic and sexual species in developmental stability, but the sexual ones are more sensitive to population changes. Absence of males may have perhaps the greatest influence on coloration, resulting in loss of symmetry in the blue spots.
Abramjan, A. & Frýdlová, P. & Juncúchová-Lásková, J. & Suchomelová, P. & Landová, E. & Yavruyan, E. (2019) -
Parthenogenetic species are usually considered to be short‐lived due to the accumulation of adverse mutations, lack of genetic variability, and inability to adapt to changing environment. If so, one may expect that the phenotype of clonal organisms may reflect such genetic and/or environmental stress. To test this hypothesis, we compared the developmental stability of bisexual and parthenogenetic lizards of the genus Darevskia. We assessed asymmetries in three meristic traits: ventral, preanal, and supratemporal scales. Our results suggest that the amount of ventral and preanal asymmetries is significantly higher in clones compared with their maternal, but not paternal, progenitor species. However, it is questionable, whether this is a consequence of clonality, as it may be considered a mild form of outbreeding depression as well. Moreover, most ventral asymmetries were found in the bisexual species Darevskia valentini. We suggest that greater differences in asymmetry levels among bisexuals may be, for instance, a consequence of the population size: the smaller the population, the higher the inbreeding and the developmental instability. On the basis of the traits examined in this study, the parthenogens do not seem to be of significantly poorer quality.
Afsar, M. & Sahin, M.K. & Afsar, B. & Cicek, K. & Tok, C.V. (2018) -
Interactions between the environment and internal regulation drive the biophysiological dynamics of lizards. Although diurnal lizards are usually heliothermic, they can sometimes be active in the absence of sunlight. Here, we report, for the first time, a case of nocturnal behavior (aided by the artificial light) in the spiny-tailed lizard - Darevskia rudis (Bedriaga, 1886) – a species that normally exhibits diurnal activity.
Agasyan, A. & Ananjeva, N. (2009) -
Agasyan, A. & Avci, A. & Tuniyev, B. & Crnobrnja-Isailovic, J. & Lymberakis, P. & Andrén, C. & Cognbalniceanu, D. & Wilkinson, J. & Ananjeva, N. & Üzüm, N. & Orlov, N. & Podloucky, R. & Tuniyev, S. & (2009) -
Ahmadzadeh, F. & Flecks, M. & Carretero, M.A. & Mozaffari, O. & Böhme, W. & Harris, D.J. & Freitas, S. & Rödder, D. (2013) -
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.
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 overall resemblance 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 (two nuclear and two mitochondrial loci), morphological (15 morphometric, 16 meristic and four categorical characters) and ecological (eleven newly calculated spatial environmental predictors) criteria to delimit cryptic species within two species complexes, D. chlorogaster and D. defilippii, both distributed in northern Iran. Phylogenetic analyses of the molecular data confirmed the monophyly of D. chlorogaster, while D. defilippii is paraphyletic in respect to D. steineri. However, each of the complexes comprises several highly divergent clades, especially when compared to other congeners. We identified seven candidate species within each complex, of which three and four species are supported by Bayesian species delimitation within D. chlorogaster and D. defilippii (including D. steineri), respectively. Although the species within one complex lack clear diagnostic features, they can be well separated based on morphological variables when sample size is appropriate. Ecological Niche Modelling provided additional support for the identified species and niche overlaps between them are generally low, especially in the D. defilippii complex.
Ahmadzadeh, F. & Kheyrandish, A. (2006) -
The Northwest of Iran, covering the three provinces of Ardabil, East Azerbaijan and West Azerbaijan, is a special area that, due to its particular geographical and climatic conditions has a rich biodiversity. The present study was conducted with the aim of studying the habitat, substrate type, feeding, conservation and distribution of lizards in this area from 2003 to now. Our results have revealed that there are 20 lizard species belonging to 13 genera and 5 families in this area. Lacertid lizards have the highest species diversity with 12 species. We also found that high ranges of habitats with a different substrate types are inhabited by lizard species. The number of species in pine regions compared with other types of area is considerably greater. The conservation status of lizards is unknown and requires further study.
Ahmadzadeh, F. & Kiabi, B.H. & Kami, H.G. & Hojjati, V. (2008) -
Northwestern Iran has unique geographical and climatic conditions that support a rich flora and fauna. In
view of the lack of in-depth studies on the lizards of the region, an investigation was started in the northern part of
Ardabil Province for an inventory of this component of the fauna and their habitats. Collections were made from
October 2003 to June 2005 and 165 specimens were collected and identified. Five families, 12 genera and 15 species
are represented, including Agamidae: Laudakia caucasia, Phrynocephalus persicus, Trapelus ruderatus; Lacertidae:
Lacerta media media, Lacerta strigata, Lacerta brandtii, Darevskia raddei raddei, Eremias strauchi strauchi,
Eremias arguta, Ophisops elegans; Scincidae: Mabuya aurata transcaucasica, Eumeces schneiderii princeps,
Abelepharus bivittatus; Anguidae: Pseudopus apodus and Gekkonidae: Cyrtopodion caspium caspium. Comparing
this list to the data provided by Anderson (1999), it seems that most of the lizards are being reported for the Province
for the first time. The families Gekkonidae and Anguidae are newly recorded, and the gecko Cyrtopodion caspium is
first recorded from the west and northwest of Iran. With seven species represented in the area, lacertids have the highest
species diversity among the lizard families and need further study. Habitat features also have been given for all
Akarsu, F. & Tuniyev, B. & Ananjeva, N. & Agasyan, A. & Orlov, N. & Tuniyev, S. (2009) -
Akman, B. & Yildiz, M.Z. & Igci, N. & Tel, A.Z. & Azidel, Ö. & Bulum, E. & Göcmen, B. (2016) -
Akman, B. & Yildizt, M.Z. & Özcan, A.F. & Bozkurt, M.A. & Igci, N. & Göcmen, B. (2018) -
Anhand ihrer Feld- und Literaturstudien identifizierten die Autoren 36 Amphibien- und Reptilienarten, die nachweislich in der turkischen Provinz Bitlis vorkommen, wobei die Fundorte und die an ihnen angetroffenen Habitattypen angegeben werden. die durch Beobachtung festgestellten Hauptgefahrdungsursachen werden benannt. insgesamt kennt man aus dem Untersuchungsgebiet Vorkommen von vier Froschlurch-, zwei Schwanzlurch- und Schildkrötenarten sowie 15 Echsen- und 13 Schlangenarten. Hyla savignyi Audouin, 1827, Heremites vittatus (Olivier, 1804), Timon kurdistanicus (Suchow, 1936) and Eirenis thospitis Schmidtler & Lanza, 1990, stellen Erstnachweise fur die Provinz Bitlis dar.
Alekperov A.M. (1978) -
Алекперов, А.M. (1978) -
Altunisik, A. & Eksilmez, H. (2018) -
Knowing the age structure of endangered species is important in order to contribute to future conservation studies for such species. In this context, we investigated age structure, age at sexual maturity, potential reproductive lifespan and longevity in a population of the Charnali lizard, Darevskia dryada, an endangered species from Turkey. The results show that the Charnali lizard has a longer life span than other lizards of the genus Darevskia that live in the same region. We estimated that these lizards attain their sexual maturity at the age of one or two years and the potential reproductive life span for males and females is six and five years, respectively. As in many other lizards, the Charnali lizard exhibited a low-level male-biased sexual dimorphism in terms of increased size. We believe that this study, in which prior information related to the Charnali lizard is shared, will contribute to future conservation activities for this critically endangered species.
Altunisik, A. & Eksiolmez, H. (2020) -
We investigated age structure, age at sexual maturity, lifespan, growth and survival rate and adult life expectancy (as demographic life-history traits) as well as body size of two Darevskia derjugini (Artvin lizard) populations from different altitudes, using skeletochronology. Our findings indicated that age upon attaining sexual maturity was two or three years in the low-altitude population (Fındıklı) while it was three years in the high-altitude population (Murgul) for both sexes. The maximum longevity was seven years in the high-elevation site while it was six years in the low-elevation site. As reported for many lizards, we found a significant positive relationship between age and body size within each sex of Artvin lizard at both altitudes. High- and low-altitude populations did not differ in age structure, survival rates, adult life expectancy and body size. Rather than the effect of altitude, which is hard to compare without replication of other low and high altitude populations, the fact that these two populations have similar growth rates and the similarity of local conditions (food availability and predator density) may indicate similarity between the two regions.
Ananjeva, N. & Borkin, L. & Darevsky, I. & Orlov, N. (1988) -
Ananjeva, N.B. (1972) -
This paper reports the results of newest investigations of lizards` demonstration behaviour. A special attention was paid to lizards` aposematic and epigamious reactions which are of great importance when forming couples and defending individual territories.
The species differences represent in quite a number of cases the most essential obsacle for the formation of couples of the individuals of different specis, even if the morphological differences of these species are hardly perceptible. Being the determining feature in intra- and inter- species contacts, the demonstration behaviour ensures the integrity of a species.
Ананьева, Н.Б. (1972) -
Ananjeva, N.B. & Borkin, L.Y. & Darevsky, I.S. & Orlov, N.L. (1998) -
Ananjeva, N.B. & Darevsky, I.S. (2004) -
Ананьева Н.Б. & Даревский И.С. (2004) -
Ananjeva, N.B. & Orlov, N.L. & Khalikov, R.G. & Darevsky, I.S. & Ryabov, S.A. & Barabanov, A.V. (2006) -
TERRA TYPICA. Settlement Terskol on the Baksan River, Kabardino-
DISTRIBUTION. Endemic of the high-mountainous Caucasus. It
occurs in the high-altitude part of the western half of the Great
Caucasus Range – from Elbrus in the east to the mountains Fisht
and Osten in the west. The eastern border of the distribution
range runs in the upper reaches of Baksan River on the south-eastern
slopes of Elbrus and mountain Cheget in Kabardino-Balkaria.
Further to the west it is widely distributed in the upper reaches of
the rivers Kuban’, Teberda, Zelenchuck, Laba, Belaya and others
flowing down to the north. It inhabits the southern slopes of the
Caucasus: in the upper reaches of the Bzyb River and its tributaries
in Abkhazia and the upper reaches of Mzymta River in the southeast
of the Krasnodar Territory.
CONSERVATION STATUS. The number is rather stable everywhere
within of its distribution range. The species is included into the
Red Data Book of IUCN with the status DD.
TERRA TYPICA. Settlement Yelenovka (= Sevan) on the shore of
the Sevan Lake, Armenia.
DISTRIBUTION. It is widely distributed in the mountains of northwestern
and northern Armenia and adjacent regions of southern
Georgia and western Azerbaijan. Outside of the Caucasus it is
also known from the north-western Turkey. In 1967 D. armeniaca
was introduced into the Ukraine. At present this parthenogenetic
species formes a stable extensive population in the rock canyon
of Teterev River of the Zhitomir region.
CONSERVATION STATUS. The number is high and stable everywhere
within the whole distribution range.
TERRA TYPICA. Settlement Krasnaya Polyana in the south of the
Krasnodar Territory, Russia.
DISTRIBUTION. Endemic of the montane Caucasus. It is widely
distributed in the western part of the Great Caucasus Range, on
northern, western and south-western slopes of the Black Sea
chain of the Great Range in the Krasnodar Territory, Abkhazia and
north-western and south-western Georgia, to the canyons of the
rivers Kodory and Ingury in lower and upper Svanetia in the east.
The northern border of its distribution passes in the area of the
middle current of numerous left tributaries of the Kuban River. The
most northern habitats of the species are known in the vicinity of
Goryachyi Klyuch on Psekupsa River and the vicinity of the town
Maikop. The southern border of the distribution range stretches
on the southern slopes of the Great Range and, gradually lowing
down, on the western spurs of the Gagra range. It comes out
to the sea in the south of the Krasnodar Territory and western
Abkhazia, where it is sporadically recorded on the coast from the
ALPINE LIZARD DAREVSKIA ALPINA (DAREVSKY, 1967)
ARMENIAN LIZARD DAREVSKIA ARMENIACA (MEHELY, 1909)
BRAUNER’S ROCK LIZARD DAREVSKIA BRAUNERI (MEHELY, 1909)
L A C E R T I D A E
town Anapa in the north up to the canyons of the rivers Khoby
and Riony in the south-west of Georgia.
The nominative subspecies D. b. brauneri occurs in the south
of the Krasnodar Territory, in Abkhazia and adjacent regions of
Georgia. D. b. darevskii (Szczerbak, 1962) inhabits foothills of
the western part of the Great Caucasus range, mainly within the
Krasnodar Territory. A comparatively small range of the subspecies
D. b. szczerbaki (Lukina, 1963) occupies a narrow strip of
coastal rocky steeps, sporadically occurring along the coast of the
Black Sea from the town Anapa in the north and for more than
100 km further to the south-east towards Novorossiisk.
CONSERVATION STATUS. On the most part of the distribution range
the number of the species is rather high and stable.
TERRA TYPICA. Kazbek Mountain, Georgia.
DISTRIBUTION. Endemic of the Great Caucasus. It is widely distributed
on the northern and southern slopes of the Great Caucasus
Range from the northern foothills of Elbrus in Kabardino-Balkaria
in the west up to the south-eastern extreme point of its Caspian
chain within Azerbaijan in the east. South-western border of the
distribution range passes on the southern foothills of Megrel’sky,
Svanetsky and Lekhchumsky mountain ranges in lower Svanetia,
and then coming down within South Ossetia approximately up to
the latitude of the health-resort Dzhava. Along the southern spurs
of Kharul’sky, Kartalinsky and Kakhetinsky mountain ranges it comes
out into the slopes of the Water Dividing (Watershed) Range in the
Alazan River valley of Georgia. In Azerbaijan this border continues
in the upper reaches of the rivers beginning from the Great Range,
and further in the east – from the slopes of the mountains Baba-
Dag and Dyubrar, in particular of the rivers Pirsagat and Sumgait.
This lizard is also widely distributed on the northern slopes of the
Great Caucasus, from the Baksan ravine in Kabardino-Balkaria in
the west to Samur range in the South Dagestan in the east.
The north-western border of the distribution area passes
here in the Baksan ravine approximately from the town Tegenekly
in the south, continuing then to the east on the northern foothills
of the Skalisty and Bokovoi ranges. There the most northern
habitats are known from the ravine Adyr-su, near the waterfall
in the Chegemskaya gorge, from the vicinity of the Blue Lakes in
Kabardino-Balkaria and the vicinity of Alagir and the settlement
Balta in the lower part of the Dar’yal ravine of North Ossetia. In
the foothills and on the northern slopes of the Andiysky mountain
range in the south-eastern Chechnya and possibly in the adjacent
part of Ingooshetia occurs a narrow-ranged subspecies D. c. vedenica
(Darevsky et Roitberg, 1999).
CONSERVATION STATUS. The number of the species everywhere
within its distribution range is high and stable.
TERRA TYPICA. Town Enzely, Iran, southern coast of the Caspian
DISTRIBUTION. It inhabits mainly forested northern and northwestern
foothills of the Elburz range in northern Iran and Lenkoran
lowland in the south-eastern foothills of the Talysh range in the
south-east of Azerbaijan. There the most northern habitats are
known from the valley of Vilyazhchai River.
CONSERVATION STATUS. On the most part of the distribution range
the number of the species is high and stable.
TERRA TYPICA. 20 km to the west from the settlement Borchka,
vilajet Artvin, north-eastern Turkey.
DISTRIBUTION. The main distribution range of the species occupies
the northern spurs of the East-Pontic mountains in the
north-east of Turkey, at least up to the vilajet Giresun in the west.
A small isolated area of the distribution range is known on the
western slopes of the mountain Mtirala in the vicinity of the town
Batumi in the south-west of Ajaria (Georgia).
CONSERVATION STATUS. As a rare species the species was included
into the Red Data Book of the USSR (1984) – category 3 and the
Red Data Book of IUCN with the status ENC2a.
TERRA TYPICA. Settlement Levashi in the region with the same
name of central Dagestan.
DISTRIBUTION. It occurs in the montane Dagestan, southern
Chechnya, Ingooshetia and south-east of north Ossetia, from the
lower part of the Dar’yal ravine in the west up to Gimrinsky range
in the east. The northern border of the distribution area passes in
the foothills of the Skalisty range in the upper current of numerous
right tributaries of Sunja River. Further on the extreme north-eastern
foothills of the frontal ranges of Dagestan it attains the nearest
vicinity of Makhachkala city in the east. An isolated population
lives also on the southern slope of the Great Caucasus range in
the canyon of the river Bol’shaya Liakhvy on the territory of South
Ossetia in Georgia.
CONSERVATION STATUS. On the most part of the distribution range
the number is stable.
TERRA TYPICA. Settlement Shagaly, Armenia.
DISTRIBUTION. It is comparatively widely distributed in the north
Armenia and southern Georgia up to the foothills area in the valley
of Kura River in the north. The recent distribution range is divided
into some isolated populations.
CONSERVATION STATUS. On the most part of the distribution range
the number is stable.
TERRA TYPICA. Valley of Lar, to the north-west of Teheran, northern
DISTRIBUTION. The distribution range of the species covers practically
all Alborz range in the north of Iran and adjacent east Kopet
Dagh range both in its Iranian part and in the south of the neighbouring
Turkmenistan. In Turkmenistan the species is very rare and
once it was considered to be extinct. In central Kopet Dagh it is
known here from a few points in the ravines Sushanka and Bol’shie
CONSERVATION STATUS. In Iran the number of the species is high
and stable. In Turkmenistan the number is catastrophically declining.
The species is included into the Red Data Book of Turkmenistan
(1999): category 3 – a rare species on the periphery of the distribution
TERRA TYPICA. Vicinity of the town Artvin, north-eastern Turkey.
DISTRIBUTION. South-west of the Krasnodar Territory, western
and north-eastern Georgia and the extreme north-west of
Azerbaijan. Outside of the Caucasus it occurs on the extreme
north-east of Turkey. Of 6 described subspecies 5 are distributed
in the Caucasus. D. d. silvatica (Bartenjev et Reznikova, 1931) inhabits
submontane part of the Krasnodar Territory mainly in the
Caucasian Biosphere State Reserve and the adjacent spurs of
the Great Caucasus. D. d. boehmei (Bischoff, 1982) lives in the
western spurs of the Great Caucasus in Krasnodar Territory and
adjoining regions of Abkhazia up to the valley of the river Kodor in
the south-east. D. d. abchasica (Bischoff, 1982) inhabits the coast
of the Black Sea of Abkhazia and the adjacent regions of western
Georgia, D. d. barani (Bischoff, 1982) is known from coastal mountain
regions of Ajaria and of neighbouring part of north-eastern
Turkey. D. d. orlowae (Bischoff, 1984) inhabits southern slopes
and foothills of the Great Caucasus in Georgia and north-western
CONSERVATION STATUS. The number is rather high and stable.
TERRA TYPICA. Ravine of small Charnali River in the lower reaches
of the river Chorokh in the extreme south-west of Ajaria (southwestern
Georgia) and the adjacent part of Turkey.
DISTRIBUTION. It occurs on a limited territory in the north-eastern
foothills of the Pontic range, within extreme south-west of Ajaria in
Georgia and adjoining regions of north-eastern Turkey. The distribution
is not sufficiently studied.
CONSERVATION STATUS. The number is rather high and stable.
TERRA TYPICA. The Crimean Peninsula.
DISTRIBUTION. The distribution range of the species is limited by
the mountain Crimea from Sevastopol in the west up to the cape
Kiik-Atlam near the town Feodosia in the east. The northern border
of the distribution range runs on the northern slopes of the
Crimean Mountains through the towns Bakhchisarai, Simferopol
and the vicinity of the settlement Vishennoe in the Belgorod region.
A small isolated population lives on the rocks Adalary near
CONSERVATION STATUS. The number is rather high and stable.
TERRA TYPICA. Town Abastumani in south-western Georgia.
DISTRIBUTION. Endemic of Georgia. The recent distribution range
forms some more or less extensive isolates mainly situated on the
slopes of the Meskhetsky range within Ajaria and western Georgia
up to Borzhomi ravine in the valley of Kura River and north-western
slopes of Trialetsky range in the east. Isolated populations live
on the southern slopes of Lechkhumsky and Rachinsky ranges in
the foothills of Great Caucasus, where this lizard is known from the
vicinity of the towns Kutaisi, Tkvibuli and the health-resort Labarde.
Its distribution in the foothills of the Great Caucasus Range is still
not sufficiently studied.
CONSERVATION STATUS. The number in the most of known populations
is rather high and stable.
TERRA TYPICA. Town Artvin, north-eastern Turkey.
DISTRIBUTION. The distribution range of the species covers
north-eastern Turkey (approximately from Erzerum in the west and
from the valley of the upper current of Aras River in the south) and
northern and southern foothills and spurs of the Meskhetsky range
within Ajaria and neighbouring regions of Georgia from the coast
of the Black Sea in the west up to the middle current of Kura River
in the east. There are local isolated populations in the ravines of
several lateral tributaries of the river Kura (Akhaldaba, Baniskhevi,
Azkuri and others). In Georgia and in the ravine of the lower current
of river Chorokh in Turkey occurs subspecies D. p. adjarica
(Darevsky et Eiselt, 1980). The remaining part of the distribution
range is occupied by the nominative subspecies D. p. parvula.
CONSERVATION STATUS. On the most part of the distribution range
the number of the species is high and stable.
TERRA TYPICA. City Tiflis (= Tbilisi), Georgia.
DISTRIBUTION. The main distribution range of the species occupies
right-bank section of the valley of middle current of Kura
River and the ravine of its right tributaries within Georgia, northern
Armenia and north-western Azerbaijan. It covers the area from
Gori in the west up to northern foothills of the ranges of the Lesser
Caucasus in the lower and middle current of the rivers flowing
into Kura River: Algeti, Khrami, Debeta, Akstafa-chai, Gyandzhachai
and others in the south-east. Big population, isolated now
from the main distribution range, is known in the valley of middle
current of the left tributary of Kura River – river Iori on the southern
slopes of the Zivgamborsky range. A great zoogeographical
interest represents an isolated population of this species discovered
in south-western Azerbaijan, in the ravine of upper current
of Akar River, belonging to the basin of Aras River. Presence of D.
portschinskii in the southern foothills of Lesser Caucasus can be
connected with a radical reconstruction of river systems which
was taking place on the territory of Transcaucasia in the postglacial
period. The nominative subspecies D. p. portschinskii occupies
the most part of the distribution range. In the ravines of right tributaries
of Kura River – the rivers Mashavera in Georgia and Debeta
in Armenia – occurs D. p. nigrita (Bakradze, 1976).
CONSERVATION STATUS. On the most part of the distribution range
the number is high and stable.
TERRA TYPICA. Settlement Narzan, North Caucasus.
DISTRIBUTION. It occurs in the north-west of the Balkan Peninsula
from one side and within the Caucasus and adjacent part of Iran
from the other side. It is sporadically distributed on the both sides
of the Great Caucasus Range within the limits of the Krasnodar
and Stavropol Territories, in Kabardino-Balkaria, North Ossetia,
Chechnya, Ingooshetia and Dagestan, and in the Transcaucasia
– on the coast of the Black Sea of Abkhazia, in north Georgia,
northern Azerbaijan. Isolated part of range lies in south-east
Azerbaijan and in the north of Armenia. D. p. praticola inhabits
the most part of the distribution range. D. d. pontica (Lantz et
Cyren, 1919) inhabits the coast of the Black Sea and north-western
CONSERVATION STATUS. On the most part of the distribution range
the number is high and stable.
TERRA TYPICA. Vicinity of the settlement Ijevan, Armenia.
DISTRIBUTION. It has a comparatively limited distribution range in
the northern Armenia and adjacent regions of Azerbaijan. The recent
distribution range is divided into several more or less isolated
populations. One of them, now extinct, existed on the north-eastern
shore of the Lake Sevan in Armenia.
CONSERVATION STATUS. The number is stable.
TERRA TYPICA. Town Trapezund (Trabzon), north-eastern
DISTRIBUTION. The species is widely distributed, mainly in
the western Transcaucasia, and in the north – in the montane
Ingooshetia, Chechnya, neighbouring Dagestan and in northern
Ossetiya. Outside of the Caucasus it is widely distributed in the
northern half of Turkey. Of eight described subspecies the five
occur in the Caucasus. D. r. bischoffi (Boehme et Budak, 1977)
occurs in Ajaria. D. r. obscura (Lantz et Cyren, 1936) inhabits the
valley and adjoining slopes of mountains of the upper current of
Kura river within Georgia. D. r. macromaculata (Darevsky, 1967)
is recorded in the ravine of the right tributary of Kura River, river
Akhalkalakichai in southern Georgia. D. r. svanetica (Darevsky et
Eiselt, 1980) occurs in the ravine of the upper current of the river
Inguri in Svanetia (western Georgia). D. r. chechenica (Eiselt et
Darevsky, 1991) is widely distributed on the southern slopes of
the Great Caucasus from lower Svanetia in the west up to northwestern
Azerbaijan in the east, and also isolatedly on the northern
slopes within the limits of the montane Chechnya and Dagestan.
CONSERVATION STATUS. The number of the species is high and
TERRA TYPICA. Town Kislovodsk, Stavropol Territory.
DISTRIBUTION. The main distribution range of the species
covers the northern slopes of the Rocky range within Karachaevo-
Cherkesskaya Autonomous Region, south of the Stavropol Territory
and Kabardino-Balkaria, from the valley of the upper current of
Kuban’ River and its tributaries in the west up to the ravine of the
middle current of the river Chegem in the east. The most northern
isolated population (which was considered as extinct) lives in the southern foothills of the Stavropol Upland, in the limits of
the Alexandrovsky region of the Stavropol Territory. The southern
border of the distribution range passes on the north-eastern
spurs of the mountain Elbrus in the ravine of the river Kuban, and,
crossing the Rocky Range (known from the Mariinsky pass), continues
further on its watershed in the upper reaches of the rivers
Kuma, Podkumok, Alikanovka, Beryozovaya, Zolka and others. On
the ravine of the river Baksan it comes up to the town Tyrny-Auza
and goes from the Chegem waterfalls up along the Chegem River
CONSERVATION STATUS. The number is high and stable.
TERRA TYPICA. Settlement Akhta, ravine of the river Razdan,
DISTRIBUTION. The distribution range of the species is divided
into several isolated populations within the northern and central
high-mountain Armenia and the adjacent regions of eastern
Turkey, from where along the valley of Kura River this lizard penetrates
into southern Georgia. Outside of the Caucasus it occurs in
the valley of the river Aras in eastern Turkey.
CONSERVATION STATUS. The number is high and stable.
TERRA TYPICA. Town Bazarchai (Basarkent), Sisiansky region,
DISTRIBUTION. The recent distribution range of the species is divided
into several of more or less extensive isolates, attributed to
the steppe high-mountain regions of the mountain-steppe zone
of Armenia, Azerbaijan, southern Georgia and eastern Turkey. The
most extensive areas of the distribution range are occupied by
the high-mountain zone of the Gegamsky range up to the shores
of Sevan Lake in the north, a ring-like high-mountain area of the
Aragaz mount and a high-mountain plateau in the foothills of the
Childyrsky and Javakhetsky ranges in the extreme north-west of
Armenia and adjoining regions of southern Georgia. Besides, it is
encountered on the Ardenissky range confining the basin of Sevan
Lake from the south. Further to the east it inhabits all high-mountain
part of the Karabakh Upland within the limits of Nagorny Karabakh.
Outside of the Caucasus, covering the mountains, surrounding the
Lake Van, the species is widely distributed in north-eastern, eastern
and south-eastern Turkey, at least from the range Palandeken
in the vilajet Erzerum in the west. Probably it penetrates into the
high-mountain region of extreme west on the territory of the
Three subspecies are described. The nominative subspecies
D. v. valentini is distributed in the whole Transcaucasia. D. v. lantzicyreni
(Darevsky et Eiselt, 1967) occurs on the most part of Turkey.
The third subspecies D. v. spitzenbergerae (Eiselt, Darevsky et
Schmidtler, 1992) is known only from the type locality in the vilajet
Khakyari in the extreme south-east of Turkey.
CONSERVATION STATUS. On the most part of the distribution area
the number of the species is high and stable.
TERRA TYPICA. Settlement Nyuvadi in the ravine of the Aras River,
DISTRIBUTION. It is widely distributed in the mountains of Lesser
Caucasus, in the most part of the southern and northern Armenia,
southern and southern-east Azerbaijan and the ravine of the upper
current of Kura River in western Georgia. In the Transcaucasia
the distribution range consists of many isolated populations of different
size associated with the ravines of small rivers flowing down
to the river Aras in the south, and tributaries of the river Kura in
the north. Outside the Caucasus it occurs in eastern Turkey (to
the east of the Lake Van) and in the adjacent north-western Iran.
AZERBAIJAN LIZARD DAREVSKIA RADDEI (BOETTGER, 1892)
MEADOW LIZARD DAREVSKIA PRATICOLA (EVERSMANN, 1834)
RI V ER KURA LIZARD DAREVSKIA PORTSCHINSKI I (KESSLER, 1678)
88 88L A
The southern border of its distribution range is not exactly determined.
Intraspecific taxonomy is still not clear. Three subspecies
are described. The nominative subspecies D. r. raddei inhabits
the most part of Armenia and Azerbaijan, in the basin of the Lake
Sevan in Armenia (mainly in the western part of the republic). In
the ravine of the upper current of the river Kura in Georgia occurs
D. r. nairensis, considered by some taxonomists as a distinct species.
D. r. vanensis (Eiselt, Schmidtler et Darevsky, 1993) occurs
in the eastern part of the basin of Lake Van in Turkey and in the
CONSERVATION STATUS. In the most part of the distribution range
the number of the species is high and stable.
Anderson, S. & Sharifi, M. & Papenfuss, T. & Bafti, S.S. (2009) -
Anderson, S.C. (1968) -
Anderson, S.C. (1974) -
Acanthodactylus cantoris blanfordi, Acanthodactylus cantoris schmidti, Acanthodactylus fraseri, Acanthodactylus micropholis, Apathya cappadocica urmiana, Eremias arguta, Eremias brevirostris, Eremias fasciata, Eremias grammica, Eremias guttulata, Eremias intermedia, Eremias lineolata, Eremias nigrocellata, Eremias persica, Eremias pleskei, Eremias scripta, Eremias strauchi, Eremias velox velox, Lacerta brandtii, Lacerta chlorogaster, Lacerta princeps princeps, Lacerta princeps kurdistanica, Lacerta saxicola defilippii. Lacerta saxicola raddei, Lacerta strigata, Lacerta trilineata media, Ophisops elegans.
Anderson, S.C. (1999) -
Acanthodactylus blanfordi, Acanthodactylus boskianus, Acanthodactylus grandis, Acanthodactylus micropholis, Acanthodactylus nilsoni, Acanthodactylus opheodurus, Acanthodactylus schmidti, Eremias acutirostris, Eremias andersoni, Eremias arguta, Eremias fasciata, Eremias grammica, Eremias intermedia, Eremias lalezharica, Eremias lineolata, Eremias nigrocellata, Eremias nigrolateralis, Eremias persica, Eremias pleskei, Eremias scripta, Eremias strauchi strauchi, Eremias strauchi kopetdaghica, Eremias velox velox, Lacerta brandtii, Lacerta cappadocica urmiana, Lacerta chlorogaster, Lacerta defilippii, Lacerta media media, Lacerta mostoufi, Lacerta praticola praticola, Lacerta princeps princeps, Lacerta princeps kurdistanica. Lacerta raddei raddei, Lacerta raddei vanensis, Lacerta steineri, Lacerta strigata, Lacerta valentini valentini, Lacerta zagrosica, Mesalina brevirostris brevirostris, Mesalina brevirostris fieldi, Mesalina watsonana, Ophisops elegans.
Anderson, S.C. (2000) -
Relatively little is known about the ecology of lizards in Iran. This paper serves as a brief introduction to five representative biotopes and their associated lizard species living syntopically, based largely on my own observations. The selected localities are found in 1) Masjed Soleyman, foothills of the Zagros Mountains, Khuzestan; 2) Kupal dunes, on the Mesopotamian Plain, Khuzestan; 3) Shah ‘Abbas Caravanserai, Dasht-e Kavir, on the Central Plateau; 4) an abandoned village in Sistan, ruined buildings and loose loess substrate; 5) Hyrcanian Forest, northern slopes of lower Alborz Mountains and the Gorgan region. The biotopes are characterized briefly and the concept of syntopy is discussed.
Andrei, M.-D. (2002) -
The author updates the knowledge on the herpetofauna of southern Dobruja according to his
own studies within the area, during 1993-1998, when he identified 18 taxa. They represent 37.5% of
the total amphibian and reptile species from Romania. Also the author makes some remarks on
respective ecosystems and on the urgent protection need of some species.
Andronikov, V.B. Danielyan, F.D. & Passinkova, R.A. (1994) -
The extent of resistance of animal cells and tissues to injurious heat effect is a cytophysiological criterion of species. In poikilotherms it can be determined both by phylogenetic relations and ecological conditions of a species` formation and existence. To analyze the relative role of these factors in formation of the tissue heat resistance level as characteristic of a species, a study was made on 9 species of lizards of the `Lacerta saxicola` complex. A characteristic feature of the complex is that some of the species belonging it are of hybrid origin, which implies that genetically they are closely related to other, parental forms. Heat resistance level of muscle tissue in a hybrid species may differ from the parental forms if the conditions of its existence are different. On other hand, genetically unrelated species living under similar conditions may have the same level of heat resistance of muscle tissue. Interspecific differences in heat resistance of muscles cannot be explained by phylogenetic relations only. Of major importance in formation of the cellular heat resistance level in the process of speciation are ecological, and primarily, thermal conditions of the environment.
Arakelyan, M. (2001) -
Arakelyan, M. (2002) -
Skeletochronological study of rock lizards hybrids Darevskia valentini × D. unisexualis, D. valentini × D. armeniaca, D. nairensis × D. unisexualis, and their parental species, collected from three mixed populations (near village of Kuchak, Lchashen, Lchap — mountainous regions of central Armenia) proved their similar pattern of growth. However, the hybrids are characterized by faster rate of growth and predominantly bigger size of body in each age group contrary to their parental species. The longevity of studied hybrids is same as in parental species and reaches 6 – 7 years.
Arakelyan, M. (2004) -
Arakelyan, M. (2007) -
Arakelyan, M. & Danielyan, F. & Corti, C. & Sindaco, R. & Leviton, A. (2011) -
Arakelyan, M. & Harutyunyan, T. & Aghayan, A. & Carretero, M.A. (2019) -
Sexual organisms should be better suited than asexual ones in a context of continuous evolution in response to opposite organisms in changing environments (“Red Queen” hypothesis of sex). However, sex also carries costs associated with the maintenance of males and mating (sex cost hypothesis). Here, both non-mutually excluding hypotheses are tested by analysing the infestation by haemogregarines of mixed communities of Darevskia rock lizards composed of parthenogens generated by hybridisation and their bisexual relatives. Prevalence and intensity were recorded from 339 adult lizards belonging to six species from five syntopic localities and analysed using Generalized Mixed-Models (GLMM). Both infestation parameters depended on host-size (like due to longer exposure with age), sex and, for intensity, species. Once accounting for locality and species, males were more parasitized than conspecific females with bisexual species, but no signal of reproductive mode itself on parasitization was recovered. Essentially, male-male interactions increased haemogregarine intensity while females either sexual or asexual had similar reproductive costs when in the same conditions. These findings deviate from the predictions from “Red Queen” dynamics while asymmetric gender costs are here confirmed. Thus, increased parasitization pressure on males adds to other costs, such as higher social interactions and lower fecundity, to explain why parthenogenetic lizards apparently prevail in the short-term evolutionary scale. How this is translated in the long-term requires further phylogenetic analysis.
Arakelyan, M. & Petrosyan, R. & Ilgar, C. & Kumlutaş, S. & Durmus, S.H. & Tayhan, Y. & Danielyan, F. (2013) -
The skeletochronological method has been used to assess age distribution and age-related differences in body size among populations of the parthenogenetic lizards Darevskia sapphirina, D. uzzelli, D. armeniaca and D. unisexualis from Turkey and Armenia. The age distribution between D. armeniaca and D. unisexualis did not significantly differ and ranged from 1 to 8 years. Maximum age for both D. uzzelli and D. sapphirina was 6 years, and 8 years for both D. unisexualis and D. armeniaca. In all the studied species, individuals reached sexual maturity after third hibernation. According to patterns of growth marks resorption, D. sapphirina is distinguished from all other rock lizards of genus Darevskia by a narrowest periosteal bone as result of high rate of endosteal resorption resulting in complete destruction of hatchling line and line of the first hibernation.
Arakelyan, M. & Soghoyan, A. & Ilgac, Ç. & Durmuş, S. & Tayhan, Y. (2010) -
Morphological diversity at 20 pholidotic and 16 meristic characters were studied on three parthenogenetic species Darevskia unisexualis (Darevsky, 1966) D. uzzelli (Darevsky & Danielyan, 1977) and D. sapphirina (Schmidtler, Eiselt & Darevsky, 1994) from Turkey which originated from interspecific hybridization of same parental species D. raddei (Boettger, 1892) and D. valentini (Boettger, 1892) and compared with parthenogenetic species D. armeniaca (Mehely, 1909) which originated from hybridization of D. mixta (Mehely, 1909) and D. valentini (Boettger, 1892) (Murphy et al, 2000, Fu et al, 2000). D. armeniaca significantly differ (ANOVA, Tukey HSD test; p < 0.05) from D. unisexualis and D. sapphirina by 11 pholidotic characters, as well as from D. uzzelli by 9 variables. D. unisexualis has shown significant differences by 8 pholidotic variables from D. sapphrina. Only two pholidotic characters distinguish D. uzzelli from D. sapphirina and D. unisexualis. In addition to meristic characters D. sapphirinia has shown significant less length and height of head as well as length of legs than D. armeniaca, D. unisexualis and D. uzzelli. The average means of snout-vent length of D. sapphirina were less in each age group in comparing with other species. The measurements of bone thickness have revealed the significant high level (p<0.001) of endosteal resorption in femur bones of D. sapphirina removing the first growth layers which usually present in bones of the most Darevskia species. The oldest lizards in our specimens were aged 6 years in D. uzzelli and D. sapphirina and 7 years D. unisexualis and D. armeniaca.
Arakelyan, M. & Stepanyan, I. (2008) -
The cytometrical analyses of diploid bisexual species Darevskia
valentini, and closely relative parthenogenetic species D.
unisexualis and D. armeniaca, as well as their triploid hybrids (D.
valentini × D. unisexualis and D. valentini × D. armeniaca) and
tetraploid hybrid D.valentini × D. unisexualis arising from
hybridization in mixed population in vicinity of Kuchak village
(Aragatsotn region, Armenia) has shown significant differences
(P<0.001) in surface and perimeter of erythrocytes between all of
them according to ANOVAs post-hoc test. The size of red blood
cells of triploid hybrids exceeds that of diploids by 25-30%, but
tetraploid male exceeds diploid males by 22% and by 10 % smaller
than those of triploid male hybrids. Among parental species the
smallest parameters of erythrocytes have D. valentini, when D.
unisexualis have the largest red blood cells. Accordingly, the
erythrocytes of hybrids of D.valentini × D. unisexualis were larger
than D. valentini × D. armeniaca.
Arakelyan, M.S. (2001) -
Arakelyan, M.S. (2007) -
Arakelyan, M.S. (2012) -
Currently among sympatric populations of biparental species of genus Darevskia, which were involved in reticulate evolution,
the processes of hybridization take place only between D. raddei and D. portschinskii species. The successful combination of parental
species with low reproductive isolation may be considered as main factor explaining the origin of parthenogenetic species.
Аракелян, М.C. (2012) -
Среди симпатрических двуполых видов яшериц рода Darevskia, участвующих в сетчатой эволюции, процессы
гибридизации на сегодняшний день интенсивно протекают только между видами D. raddei и D. portschinskii. Удачная
комбинация родительских двуполых видов с неполной репродуктивной изоляциeй является важным фактором появления
партеногенетических видов. Ներկայումս Darevskia ցեղի երկսեռ ժայռային մողեսների սիմպատրիկ պոպուլյացիաներում, որոնք ներգրավված էին եղել
ցանցաձև էվոլյուցիայի գործընթացի մեջ, ընթանում է հիբրիդային տեսակառաջացում միայն D. raddei և D. portschinskii տեսակների
միջև: Ծնողական տեսակների հաջող համադրությունը ցածր վերարտադրողական մեկուսացման դեպքում կուսածին տեսակների
առաջացման կարևոր գործոն է:
Arakelyan, M.S. & Danielyan, F.D. (2000) -
Age determination based on counting the annual layers in the femur was performed for three parthenogenetic species of Armenian rock lizards, Lacerta armeniaca (n = 147), L. unisexualis (n = 68), and L. dahli (n = 18), and for two bisexual species, L. nairensis (n = 29) and L. raddei (n = 32). Comparison of parthenogenetic and bisexual species shows that the growth rate, sexual maturity age, and life longevity in rock lizards are independent of the type of reproduction. The examined Lacerta species reach maturity by the second or third year of life. The maximum age of individuals in the samples studied is 7 and 5 years for L. armeniaca and L. nairensis, respectively, and 6 years for L. unisexualis, L. dahli, and L. raddei.
Arakelyan, M.S. & Danielyan, F.D. (2011) -
Range overlap between species even at local scales is a defining trait in the biogeography of the Caucasian rock lizards of the genus Darevskia. Syntopy between populations of two biparental species is remarkable, especially when they are parental contributors of parthenogenetic species. The detailed analyses of such contact zones revealed that when biparental species D. valentini and D. nairensis are in syntopy, no hybrid forms arise. This is likely due to their different reproductive periods. In contrast, the biparental D. raddei and D. portschinskii often produce hybrids, and the parthenogenetic species D. rostombekovi is also found in low numbers in the same sites which suggests that they are an occasional result of hybridization. Three sympatric zones between D. raddei and D. portschinskii are known for Armenia, Nagorno-Karabakh Republic (NKR) and Azerbaijan. Likely, the reproductive isolation between D. raddei and D. portschinskii is not complete which is contributing to reticular evolution. The hybridization events between syntopic D. raddei and D. portschinskii produce lizards with intermediate scalation and coloration characters when compared to allopatric populations. Among 207 lizards from Northern Armenia 59 (28.5%) D. raddei and 53 (25.5%) of D. portschinskii displayed intermediate morphology, 6 (2.86%) were parthenogenetic D. rostombekovi and 4 (1.93%) triploid hybrid females D. raddei × D. rostombekovi. Of 20 lizards with intermediate morphology 53 eggs and 35 young were obtained; 12 of young were similar to D. rostombekovi according to their external morphology. Among 143 lizards collected in another syntopic locality with D. raddei and D. portschinskii in NKR, 24 were hybrid females of uncertain morphological ascription. Among 66 D. portschinskii 18 individuals displayed modified morphological characters similar to D. raddei while among 43 D. raddei 14 individuals show morphologies close to D. portschinskii. Therefore, the successful combination of parental species with low reproductive isolation may be considered as the main factor explaining the origin of parthenogenetic species. For next steps of reticulate evolution, the combination of biparental and parthenogenetic species is required. Namely, in the sympatric zone of biparental D. raddei (40% of lizards) and parthenogenetic D. rostombekovi (35%), D. armeniaca (20%) and D. dahli (5%), triploid hybrids arise only between D. raddei and D. rostombekovi (18 hybrid males and 6 intersexes with both hemipenises and oviducts found). D. raddei from this locality showed pholidosis and coloration approaching D. rostombekovi.
Arakelyan, M.S. & Danielyan, F.D. (2014) -
Argana, E. & Freitas, S. & Sillero, N. & Corti, C. & Drovetski, S.V. & Garcia-Munoz, E. & Ferreira, D. & Ferreira, S. & Carretero, M.A. (2013) -
Arikan, H. & Göçmen, B. & Yildiz, M.Z. & Ilgaz, Ç. & Kumlutas, Y. (2009) -
The present study is on the morphologies and sizes of peripheral blood cells (erythrocytes, leukocytes, and thrombocytes) on blood smears, stained with Wright`s stain, in some lacertid lizards species [Apathya cappadocica (Werner, 1902), Acanthodactylus boskianus (Daudin, 1802), Acanthodactylus harranensis Baran et al. 2005, Anatolacerta danfordi (Günther, 1876), Darevskia praticola (Eversmann, 1834), D. uzzelli (Darevsky and Danielyan, 1977), D. valentini (Boettger, 1892), Parvilacerta parva (Boulenger, 1887), Lacerta pamphylica Schmidtler, 1975, L. trilineata Bedriaga, 1886, L. viridis (Laurenti, 1768), Ophisops elegans Menetries, 1832, Mesalina brevirostris Blanford, 1876, Podarcis muralis (Laurenti, 1768), P. sicula (Rafinesque-Schmaltz, 1810), Timon princeps (Blanford, 1874)] from Turkey. As a result of our survey, it was determined that the blood cells of the investigated species are shows significant variations in sizes and of leukocytes, agranulocytic leukocytes (lymphocytes and monocytes) are present as predominant cells. Moreover, of granulocytes, neutrophils were no observed in A. danfordi, D. praticola, D. uzzelli, and P. parva.
Arnold, E.N. (1973) -
Algyroides fitzingeri, Algyroides marchi, Algyroides moreoticus, Algyroides nigropunctatus, Lacerta agilis, Lacerta lepida, Lacerta princeps, Lacerta schreiberi, Lacerta strigata, Lacerta trilineata, Lacerta viridis, Lacerta andreanszkyi, Lacerta armeniaca, Lacerta bedriagae, Lacerta brandtii, Lacerta cappadocica, Lacerta caucasica, Lacerta chlorogaster, Lacerta cyanura, Lacerta danfordi, Lacerta derjugini, Lacerta dugesii, Lacerta fraasii, Lacerta graeca, Lacerta horvathi, Lacerta jayakari, Lacerta laevis, Lacerta monticola, Lacerta mosorensis, Lacerta oxycephala, Lacerta parva, Lacerta perspicillata, Lacerta praticola, Lacerta rudis, Lacerta saxicola, Lacerta vivipara, Gallotia atlantica, Gallotia galloti, Gallotia simonyi, Podarcis erhardii, Podarcis filfolensis, Podarcis hispanica, Podarcis lilfordi, Podarcis melisellensis, Podarcis milensis, Podarcis muralis, Podarcis peloponnesiaca, Podarcis pityusensis, Podarcis sicula, Podarcis taurica, Podarcis tiliguerta, Podarcis wagleriana, Psammodromus algirus, Psammodromus blanci, Psammodromus hispanicus, Psammodromus microdactylus.
Arnold, E.N. (1987) -
Twenty-four species of lacertid lizards were examined at 31 sites in western Yogoslavia, Greece and Iberia. Comparative observations were made on over 4500 individual lizards, noting such features as times of activitiy, hunting methods, diet, micro- and macrohabitat, refuges used and body temperature. These data are used as a basis for assessing resource partition in related sympatric species, for find out whether different systematic groups have characteristic types of niche, and for comparing cumminity structures in the peninsulas of southern Europa.
Acanthodactylus erythrurus, Algyroides marchi, Algyroides moreoticus, Algyroides nigropunctatus, Lacerta agilis, Lacerta graeca, Lacerta horvathi, Lacerta lepida, Lacerta monticola, Lacerta oxycephala, Lacerta mosorensis, Lacerta schreiberi, Lacerta trilineata, Lacerta viridis, Lacerta vivipara, Podarcis bocagei, Podarcis erhardii, Podarcis hispanica, Podarcis melisellensis, Podarcis milensis, Podarcis muralis, Podarcis peloponnesiaca, Podarcis sicula, Podarcis taurica, Psammodromus algirus, Psammodromus hispanicus.
Arnold, E.N. (2002) -
Differences in surface structure (ober- hautchen) of body scales of lacertid lizards involve cell size, shape and surface profile, presence or absence of fine pitting, form of cell margins, and the occurrence of longitudinal ridges and pustular projections. Phylogenetic information indicates that the primitive pattern involved narrow strap-shaped cells, with low posteriorly overlapping edges and relatively smooth surfaces. Deviations from this condition produce a more sculptured surface and have developed many times, although subsequent overt reversals are uncommon. Like variations in scale shape, different patterns of dorsal body microornamentation appear to confer different and conflicting performance advantages. The primitive pattern may reduce friction during locomotion and also enhances dirt shedding, especially in ground-dwelling forms from moist habitats. However, this smooth microornamentation generates shine that may compromise cryptic coloration, especially when scales are large. Many derived features show correlation with such large scales and appear to suppress shine. They occur most frequently in forms from dry habitats or forms that climb in vegetation away from the ground, situations where dirt adhesion is less of a problem. Microornamentation differences involving other parts of the body and other squamate groups tend to corroborate this functional interpretation. Microornamentation features can develop on lineages in different orders and appear to act additively in reducing shine. In some cases different combinations may be optimal solutions in particular environments, but lineage effects, such as limited reversibility and different developmental proclivities, may also be important in their genesis. The fine pits often found on cell surfaces are unconnected with shine reduction, as they are smaller than the wavelengths of most visible light.
Arnold, E.N. & Arribas, O. & Carranza, S. (2007) -
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.
Arribas, O. & Candan, K. & Kumluta, Y. & Ilgaz, Ç. (2021) -
All the Turkish populations studied, both those previously assigned to D. dryada (Subaşı and Yoldere villages, near Hopa) and those attributed to D. clarkorum (the largest sample studied so far, 177 specimens in total), are indistinguishable from each other and therefore must all be ascribed to the natural variability of a monotypic D. clarkorum. The Georgian specimens from the Type Locality of D. dryada (Charnaly river gorge, Chevachauri district) are clearly different, so that taxon cannot be considered a simple synonym for D. clarkorum, but as a valid taxon, although its proper status (more probably as a subspecies of D. clarkorum), is yet to be clarified. It is a highly threatened population, so studies should be done in vivo or with as low intrusiveness as possible.
Darevskia dryada is clearly larger (SVL) than any D. clarkorum studied, with strongly longer heads and pilei in adult males (and hence more teeth in dentary bone), and higher dorsalia counts. There also seem to be (but need to be studied in a larger sample) more longitudinal rows of temporal scales between tympanic and parietal plates, a tendency to have more supralabial scales; comparatively smaller values for longitudinal rows of scales on the ventral surface of the thigh between the femoral pores and the outer row of enlarged scales, and higher collaria, and circumanalia scales. Other differences in femoralia and gularia are also reflected in Darevsky & Tuniyev’s (1997) tables and should also be investigated with more Georgian specimens.
Two supposed discriminant characters, the frontonasal index and the presence of developed masseteric, are not valid. The frontonasal index does not discriminate both taxa; D dryada specimens fall inside the variation of D. clarkorum for this character. Also the presence of a developed masseteric plate is supposed to be rare if at all in D. clarkorum but always present in D. dryada; however, it appears in nearly 75% of D. clarkorum studied and in all D. dyada, so is also no longer valid for taxa discrimination.
Although very similar, D. clarkorum and D. dryada are morphologically different, and genetic studies (as the unpublished results mentioned by Fu, 1999) do not make the provenance of the specimens clear, and hence the correct identification of the supposed specimens of D. dryada used.
There are no geographical clines in D. clarkorum. However, as stated by Schmidtler et al. (2002), there is an inverse relationship between altitude and dorsalia values in D. clarkorum. Both the general differentiation between populations and the scalation (dorsalia) appear statistically correlated with the altitude and also with latitude (being both factors not strictly the same). The correlation seems to be stronger with morphology in general (multiple scalation characters and head biometry) than only with dorsalia. In the case of the general differentiation among samples, it is also significantly correlated with temperatures during the activity period (April-September) and with precipitation during incubation (July-August). As these climatic parameters of temperature and precipitation are not directly correlated with the dorsalia variation, the relation with altitude (and perhaps latitude) must be linked to some other climatic parameter not studied here, perhaps solar radiation or evapotranspiration.
Arribas, O. & Candan, K. & Kurnaz, M. & Kumlutas, Y. & Caynak, E-Y. & Ilgaz, Ç. (2022) -
In this study, we re-examine the Darevskia parvula group comprehensively using morphology, osteology and mitochondrial phylogeny, and describe a new endemic species from Turkey: Darevskia tuniyevi sp. nov. A total of 257 adult specimens were evaluated for external morphology (scalation and biometry) with univariate (descriptive statistics and ANOVA with post-hoc tests) and multivariate (Discriminant Analysis and ANOSIM) analyses. In parallel, osteological data and molecular analyses using three DNA markers (mitochondrial 16S rRNA and Cyt-b, nuclear Rag-1) were used to complete the description of the new taxon. The molecular phylogenetic analyses indicated that the D. parvula group is composed of three taxa as D. parvula, D. adjarica and D. tuniyevi sp. nov., and showed that D. adjarica and D. tuniyevi sp. nov. are reciprocal sister taxa. On the other hand, D. adjarica is morphologically very different from other two forms, while D. parvula is hardly distinguishable externally from D. tuniyevi sp. nov. Therefore, we can consider that D. parvula and D. tuniyevi sp. nov. are cryptic species. These two cryptic species retain their primitive morphology within the group, while D. adjarica has changed, perhaps due to different bioclimatic conditions in its Pleistocene refuge and current area.
Arribas, O.J. (1997) -
Arribas, O.J. (1998) -
Die Pyrenäenarten von Archaeolacerta s.l. (A. aranica, A. aurelioi und A. bonnali) unterscheiden sich von den übrigen Arten der Eurasiatischen Radiationslinie (sensu MAYER & BENYR 1994) durch den Besitz knöcherner Rippen im Bereich des dritten Präsakralwirbels. Die drei Pyrenäenarten zeigen in der Morphologie von Postorbitale und Postfrontale jeweils osteologische Verschiedenheiten, die Artniveau erreichen und ihre zweifelsfreie Unterscheidung erlauben. Auch finden sich Unterscheidungsmerkmale im Schultergürtel (Klavikel und Sternalfontanella).
Apomorphe Merkmale innerhalb der kaukasischen ´Lacerta` saxicola – Gruppe stellen deutliche Unterschiede sowohl gegenüber den Archaeolacerten Europas (fast immer einzusätzlicher Präsakralwirbel) als auch gegenüber Apathya (ein zusätzlicher Präsakralwirbel und eine verminderte Anzahl von Prämaxillarzähnen) dar. ´L.` chlorogaster unterscheidet sich von den übrigen kaukasischen Formen klar durch die Zahl der Prämaxillarzähne und Form der Interklavikel. Die parthenogenetischen Taxa zeigen deutliche Ähnlichkeiten (Modalwerte der Wirbelzahl, Morphologie des Postorbitale) zu ihrem Elternarten, besonders zur weiblichen Elternart.
Aufgrund der osteologischen Befunde wird deutlich, daß die europäischen, vorläufig in der Sammelgattung Archaeolacerta s.l. vereinigten Taxa keine natürliche Einheit darstellen. Die Formen der Pyrenäen und des Kaukasus erscheinen osteologisch deutlich differenziert, während die iberischen, balkanischen und tyrrhenischen Taxa in den untersuchten Strukturen nur synplesiomorphe Merkmalsausprägungen gemeinsam haben, die keine Aussagen über ihre Verwandtschaft ermöglichen.
Arribas, O.J. (1999) -
Archaeolacerta s. l., in the currently utilized sense results to be a polyphyletic taxon, composed of three monophyletic and phylogenetically well differentiated taxa: a)-Archaeolacerta (s. str.) Mertens, 1921, which includes three species: A. bedriagae, A. oxycephala, and A. mosorensis, distributed by the northern part of the Central Mediterranean (Corsica, Sardinia, and west Balcanic Peninsula). His most related genera seem to be Teira (his adelphotaxon in our study), Apathya, and Omanosaura. b)-Iberolacerta gen. nov., with two subgenera: Iberolacerta s. str. and Pyrenesaura subgen. nov. includes six species: I. monticola, I. cyreni, I. bonnali, I. aranica, I. aurelioi, and I. horvathi. They are distributed by the mountains of Western Europe (Iberian Peninsula, Pyrenees, Central and Eastern Alps and the north of the Dinaric Chains). His adelphotaxon are the Caucasian and Near East species of Darevskia gen. nov. c)-The third taxa is Darevskia gen. nov., which includes the species of the `L.` saxicola complex besides `L.` derjugini, `L.` praticola, and `L.` chlorogaster. They are distributed by Eastern Europe, Caucasus, and Near East. Other clear relationships among some taxa of the Eurasian Radiation [= `Eurasische linie` from (Mayer and Benyr, 1994)] have been found: Algyroides seems to be the sister group of the species considered of uncertain phylogenetic relationships belonging to the `L.` danfordi-laevis group and of Podarcis. `L.` brandtii seems to be a very primitive species within the Eurasian Radiation. Omanosaura and Apathya appear as sister taxa. `L.` graeca occupies a very basal position in the sister group of Podarcis and relatives (see above). `L.` andreanskyi is the sister species of the Darevskia nov. and Iberolacerta nov. clade. The assimilation of `L.` andreanskyi to Teira is very problematic. Also the `L.` parva group seems to be related to Timon. Some groups of species like `L.` brandtii, `L.` parva-fraasii and `L.` danfordi-laevis probably merit generic rank, but it is necessary a more deep study before to take a decision.
Arribas, O.J. (2012) -
In this paper we deal on the ultraviolet color (invisible to us): where we can find it, the capability of animals to see it and the advantages that this color perception offers to them. As the simplest way to detect it is the photography, we describe and review how to photograph the UV, as a result of 15 years of amateur experience, searching
and testing nearly in complete blindness due to the lack of practical information about “how to do it”. We describe the different kinds of photography (chemical and digital); the cameras and objectives suitable (both astronomically expensive ones and cheap options); what are the best characteristics that the objectives should have for this purpose; the films suitable for their use in chemical photography; the different filters (current or discontinued) manufactured along the years; and the subtle combinations among the different materials to obtain pure UV photographs. This
kind of scientific photography is mainly used in forensics, forgery detection, art dermatology and less in Natural History, despite the fact that a great part of animals see this color and use it in important questions of their biology as the social behavior, mate choice or the food search.
Arribas, O.J. (2016) -
Caucasilacerta Harris, Arnold et Thomas, 1998 has been considered as a nomen nudum for the last 18 years. The
main reason for this was the lack of a diagnosis or reference to it. Now, some authors argue that a paragraph in the
same paper could be a valid diagnosis and thus Caucasilacerta might be an available valid name. In the present
manuscript I demonstrate that Caucasilacerta is a nomen nudum by: a) the lack of diagnosis or reference to it accompanying
(sic! mandatory in the ICZN) to the new name; b) the alleged diagnosis (fide Busack et al., 2016) is
102 lines away (two pages, including a figure and its legend) and there is not a reference to it accompanying the
new name; and c) the subject of the alleged diagnosis are “the Rock-dwelling lacertids in the Caucasus” (sic.!),
different from the group to which the new name supposedly refers to, the “L. saxicola group” (sic!). A relationship
between both names that can only be ascertained by a specialist in the group and the concerned geographic area.
As a result of that, Caucasilacerta is a nomen nudum, unavailable, and thus it is not even a Junior Synonym of
Darevskia Arribas, 1997.
Arribas, O.J. & Ananjeva, N.B. & Carranza, S: & Doronin, I.V. & Harris, D.J. & Orlov, N.L. & Orlova, V.F. (2017) -
Busack et al. (2016) concluded that the name Darevskia should be attributed to Arribas (1999) instead
of Arribas (1997), rendering the latter name a junior synonym of Caucasilacerta Harris et al.
(1998). In this paper, we provide a detailed answer to Busack et al. (2016) and argue that Arribas
(1997) fulfilled the publication requirements of the International Code of Zoological Nomenclature
(ICZN) in place in 1997 and, therefore, Darevskia Arribas, 1997 is a valid name. In order to clarify
this issue, a case has been submitted to the ICZN. To avoid unnecessary confusion in Old World
Lacertid taxonomy, we recommend that until the commission rules on this case, Arribas (1997)
should continue to be considered the authority of both Darevskia and Iberolacerta.
Arribas, O.J. & Ananjeva, N.B. & Carranza, S: & Doronin, I.V. & Orlov, N.L. & Orlova, V.F. (2018) -
The purpose of this application under Articles 78.1 and 81 of the Code is to request the Commission to use its plenary power to choose among two alternatives in order to preserve the genus name Darevskia and nomenclatural stability.
In Alternative A, the Commission is asked to accept the microfiche publication (Arribas,
1997) as a published work in the sense of the Code and made available the names included therein, placing Iberolacerta Arribas, 1997 and Darevskia Arribas, 1997 on the Official List of Generic Names in Zoology, their type species on the Official List of Specific Names in Zoology, and the microfiche edition of Arribas (1997) on the Official List of Works Approved as Available for Zoological Nomenclature. This action will promote stability,
as both names are widely accepted and, if the 1997 publication in microfiche is considered
unavailable, then Darevskia Arribas, 1997 could potentially be threatened by Caucasilacerta Harris et al., 1998 (by authors who would not consider this name a nomen nudum), which would clearly disrupt stability. Not only would this alternative be the best way to preserve nomenclatural stability as stressed in the ICZN Preamble and repeated in several Articles of the Code, but it would also be the only way to ensure that these names (available at the time of their publication) remain available after the retroactive changes that have been introduced between the third and fourth editions of the Code. If Alternative A is chosen, both Iberolacerta and Darevskia become available from Arribas (1997).
In Alternative B, the microfiche publication (Arribas, 1997) is considered invalid and placed on the Official Index of Rejected and Invalid Works in Zoological Nomenclature. Both Iberolacerta and Darevskia would be considered published in Arribas (1999). Also, the Commission is asked to place on the Official Index of Rejected Generic Names in Zoology the name Caucasilacerta Harris et al., 1998 (as a nomen nudum due to the lack of a valid diagnosis) and to place on the Official List of Generic Names in Zoology the names Iberolacerta Arribas, 1999 and Darevskia Arribas, 1999.
Arribas, O.J. & Ilgaz, C. & Kumlutaş, Y. (2018) -
The intraspecific variability of Darevskia parvula (which has two classical subspecies easily identifiable by external characteristics, D. p. parvula and D. p. adjarica), was studied using various approaches including morphology (scalation and biometry), multivariate analyses (PCA, CDA, ANOSIM, UPGMA and MST), osteology, and molecular techniques. High mitochondrial distance, differences at the nuclear level and morphological distinctiveness warrant the specific status of
both taxa, Darevskia parvula (Lantz & Cyrén, 1913) and Darevskia adjarica (Darevsky & Eiselt, 1980) stat. nov. A lectotype for D. parvula, originally described with syntypes of both species -D. parvula and D. adjarica- is designated. The uncorrected genetic distance between D. parvula and D. adjarica in the cytochrome b mitochondrial gene is 14.4% ± 1.9%. Intraspecific variability within D. parvula is very small (1.5% ± 0.5%), and was not detected in our samples of D. adjarica. The two species further differ by two mutations in the nuclear melano-cortin 1 receptor (mc1r) gene. Interestingly, past introgression of D. parvula mitochondrial haplotypes (5% ± 1% different to those currently known) into some D. adjarica has been detected in one locality; all the studied specimens of D. adjarica with mtDNA of D. parvula are unmistakably D. adjarica at the morphological and nuclear levels. Morphologically, there is almost no overlap between D. parvula and D. adjarica. These results are corroborated by CDA, MST and UPGMA trees. Specimens of the inland high mountain population of Ardahan (clearly D. adjarica in
CDA, MST and UPGMA trees) occupy a somewhat intermediate position between both taxa in the PCA (when specimens and not populations as a whole are considered), but this morphological closeness may be attributed to the influence of climatic factors (continental conditions) on scalation of the specimens. Males appear to be more differentiated than females. Overlap among samples within each species is very marked; none can be separated clearly from its conspecifics. This is even more marked in D. parvula, which has a fairly small area compared to D. adjarica.
Darevskia parvula and D. adjarica samples appear to be homogeneously clustered within species and well separated between the two species in the UPGMA trees. In males and females all the D. parvula samples are very similar and moderately differentiated. In males of D. adjarica, the most differentiated seems to be adjBorçka, the others all being clustered together, with adjÇaykara showing slightly more differentiation from the rest (adjOrtacalar, adjArdahan, adjIkizdere and adjÇermik). Darevskia adjarica females are also similarly distributed into two subgroups, one including Borçka, Çermik and Ardahan and the other including Ortacalar, Ikizdere and Çaykara. In both sexes, the inland Ardahan sample clearly belongs to D. adjarica. From the most connected MST samples, speculations can be made about areas of origin and expansion of the different taxa. Ortacalar (D. adjarica) and Hatila (D. parvula) are the most connected (morphologically more “central” in both taxa); in fact, both populations are relatively close, living on the northern (Black Sea) and southern (Anatolian) facing slopes, respectively of the Doğu Karadeniz Mountains (Kaçkar Mountains). This highlights these mountains, which rise from sea level up to nearly 4000 m asl. and have wide buffering possibilities against climate changes, as a zone of refuge and posterior dispersion of this species, and even of the original splitting into two taxa adapted to these different conditions, D. adjarica on the coast and D. parvula on the continental slope. Osteologically D. parvula and D. adjarica are very similar, although Georgian specimens from an isolated population (Atskuri) have closed clavicles not found in Turkish D. adjarica. Also, inland Ardahan D. adjarica have an extra vertebra in both males and females, compared to the other studied specimens from both species. ARRIBAS ET AL. 72 · Zootaxa 4472 (1) © 2018 Magnolia Press The present study indicates that the situation in Turkey is that D. parvula is well differentiated and lives around the
Çoruh River Valley, contoured by D. adjarica populations on the coastal-facing slopes of the Doğu Karadeniz Mountains
on one side, and the Yalnızçam Mountains on the other side, where D. adjarica enters from Georgia as the opposite extreme of a geographic distribution. The attribution of more inland ranges to D. parvula or D. adjarica, as well as the detailed genetic structure of both taxa may be confirmed with more specific studies.
Arribas, O.J. & Ilgaz, C. & Kumlutaş, Y. & Durmus, S.H. & AVci, A. & Üzüm, N., (2013) -
A broad sample of Darevskia rudis from the main part of its range was reviewed with regard to external morphology (dis-criminant, UPGMA, MST and ANOVA analyses) and osteology. Darevskia bithynica is raised to species rank, with two subspecies: D. b. bithynica and D. b. tristis. The other subspecies are fairly similar (D. r. rudis being the most different). Two singular populations are described as subspecies: D. r. mirabilis ssp. nov. from Kaçkar Mountains, geographically adjoins the otherwise different D. r. bischoffi and D. r. bolkardaghica ssp. nov., which is geographically isolated but that seems to be very closely related to D. r. obscura.
Arutyunyan, T.K. (2013) -
The infection of blood parasites rock lizards of genus Darevskia depending on the environment was studied. The bisexual (D. valentini) and parthenogenetic (D. armeniaca) species and their interpopulation differences depending on the infestation were compared.
Арутюнян, Т.К. (2013) -
Изучалась зараженность кровепаразитами скальных ящериц рода Darevskia в зависи- мости от среды обитания. Сравнивались двуполые (D. valentini) и партеногенетические (D. armeniaca) виды и их межпопуляционные различия в зараженности особей указанным паразитом. àõëáõÙÝ3ëÇñí»É ¿ Darevskia ó»ÕÇ Å3Ûé3ÛÇÝ ÙáÕ»ëÝ»ñÇ í3ñ3Ïí3ÍáõÃÛáõÝÝ 3ñÛ3Ý Ù3- Ï3μáõÛÍÝ»ñáía Ï3Ëí3Í μÝ3ÏáõÃÛ3Ý ÙÇç3í3ÛñÇ å3ÛÙ3ÝÝ»ñÇó: Ð3Ù»Ù3ïí»É »Ý »ñÏë»é (D. valentini) ¢ Ïáõë3ÍÇÝ (D. armeniaca) ï»ë3ÏÝ»ñÁ ¢ ïíÛ3É Ù3Ï3μáõÛÍáí í3ñ3Ïí3ÍáõÃÛ3Ý, ÙÇçåáåáõÉÛ3óÇáÝ ï3ñμ»ñáõÃÛáõÝÝ»ñÁ:
Arutyunyan, T.K. & Petrosyan, F.D. & Danielyan, F.D. & Arakelyan, M.S. (2013) -
Aslanyan A.V. & Arakelyan M.S. (1999) -
Асланян А.В. & Аракелян М.С. (1999) -
Badaeva, T.N. (2008) -
Николаевна, Б.T. (2008) -
Badaeva, T.N. & Korchagin, V.I. & Tokarskaya, O.N. & Ryskov, A.P. (2006) -
Badaeva, T.N. & Korchagin, V.I. & Tokarskaya, O.N. & Ryskov, A.P. (2007) -
Badaeva, T.N. & Malyshava, D.N. & Korchagin, V.I. & Ryskov, A.P. (2008) -
Unisexual all-female lizards of the genus Darevskia that are well adapted to various habitats are known to reproduce
normally by true parthenogenesis. Although they consist of unisexual lineages and lack effective genetic recombination,
they are characterized by some level of genetic polymorphism. To reveal the mutational contribution to overall genetic
variability, the most straightforward and conclusive way is the direct detection of mutation events in pedigree genotyping.
Earlier we selected from genomic library of D. unisexualis two polymorphic microsatellite containg loci Du281 and Du215. In
this study, these two loci were analyzed to detect possible de novo mutations in 168 parthenogenetic offspring of 49 D.
unisexualis mothers and in 147 offspring of 50 D. armeniaca mothers . No mutant alleles were detected in D. armeniaca
offspring at both loci, and in D. unisexualis offspring at the Du215 locus. There were a total of seven mutational events in the
germ lines of four of the 49 D. unisexualis mothers at the Du281 locus, yielding the mutation rate of 0.1428 events per germ
line tissue. Sequencing of the mutant alleles has shown that most mutations occur via deletion or insertion of single
microsatellite repeat being identical in all offspring of the family. This indicates that such mutations emerge at the early
stages of embryogenesis. In this study we characterized single highly unstable (GATA)n containing locus in parthenogenetic
lizard species D. unisexualis. Besides, we characterized various types of mutant alleles of this locus found in the D. unisexualis
offspring of the first generation. Our data has shown that microsatellite mutations at highly unstable loci can make a
significant contribution to population variability of parthenogenetic lizards.
Bader, T. & Kopeczky, R. & Riegler, C. & Wieland, F. (2016) -
Die Eidechsenfauna Armeniens setzt sich nach aktuellem Wissensstand aus 17 Arten aus fünf Gattungen zusammen. Die am stärksten vertretene Gattung ist Darevskia mit neun Arten. Davon sind fünf zweigeschlechtlich (D. nairensis, D. portschinskii, D. praticola, D. valentini) und vier Arten parthenogenetisch (D. armeniaca, D. dahli, D. rostombekovi, D. unisexualis). Neben Kurzbeschreibungen der Arten teilen wir unsere Beobachtungen einer Exkursion im Jahr 2015 mit.
Baeckens, S. & Edwards, S. & Huyghe, K. & Van Damme, R. (2015) -
Animals communicate via a variety of sensory channels and signals. Studies on acoustic and visual communication systems suggest that differences in the physical environment contribute to the variety of signalling behaviour, with species investing in those signals that are transmitted best under the local conditions. Whether or not environmental tuning also occurs in chemical communication systems has received much less attention. In the present study, we examined the effect of several aspects of the physical environment on the chemical communication system of lacertid lizards (family Lacertidae). The numbers of femoral pores are used as a proxy reflecting how much a particular species invests in and relies upon chemical signalling. Femoral pores are specialized epidermal structures that function as a secretion channel for the waxy substance produced by glands. In some lacertid species, the secretion carries infochemicals that play an important role in social communication. The number of femoral pores varies considerably among species. We have compiled data on femoral pore numbers for 162 species and tested for the effects of climate and substrate use. After correcting for body size and taking the phylogenetic relationships among the species into account, we found no effect of climate conditions or latitude on species pore numbers. Substrate use did affect pore numbers: shrub-climbing species tended to have fewer femoral pores than species inhabiting other substrates.
Bakradze, M.A. (1976) -
The new subspecies is fouпd in the mauпtain regions of south-eastern Georgia and northern Armenia. It differs from the nominative form Ьу а much darker соlош of the body upper side and Ьу certain peculiarities of the squamous cover.
Bakradze, M.A. (1977) -
Bannikov, A.G. & Darevsky, I.S. & Ishchenko, V.G. & Rustamov, A.K. & Szczerbak, N.N. (1977) -
Банников А.Г. & Даревский И.С. & Ищенко, В.Г. & Рустамов, А.К. & Щербак, Н.Н. (1977) -
Barabanov, A.V. & Doronin, I.V. (2020) -
Ilya S. Darevsky co-described 70 taxa (three genera, 46 species, 21 subspecies) in 44 publications belonging to five orders, eight families of amphibians and reptiles during his career in herpetology. Of this number, three taxa are fossil and 57 taxa are currently considered as valid. By the regions where new taxa were discovered Southeast Asia and Western Asia (includes Caucasus and Asia Minor) dominates. The largest number of descriptions was published in the Russian Journal of Herpetology.
Baran, I. (1977) -
Baran, I. & Atatür, M.K. (1998) -
Baran, I., Kumlutaş, Y., Tok, C.V., Ilgaz, C., Kaska, Y., Olgun, K., Türkozan, O. & F. Iret (2004) -
Die Autotren berichten über zwei Aufsammlungen von Amphibien und Reptilien aus dem Gebiet der ostanatolischen Städte Erzurum, Iğdir, Kars und Ardahan. Das Material umfaßt drei Anurentaxa, eine Schildkrötenform, 11 Eidechsen- und 11 Schlangenarten in 172 Exemplaren.
Bufo viridis, Rana ridibunda complex, Rana camerani, Testudo graeca armeniaca, Phrynocephalus persicus, Eremias pleskei, Eremias strauchi, Lacerta agilis brevicaudata, Parvilacerta parva, Darevskia parvula, Darevskia raddei, Darevskia valentini valentini, Darevskia uzzelli, Lacerta trilineata, Ophisops elegans, Typhlops vermicularis, Eryx jaculus turcicus, Haemorrhois ravergieri, Hierophis schmidti, Platyceps najadum, eirenis modestus, Eirenis punctatolineatus, Coronella austriaca, natrix natrix persa, Natrix tessellata, Vipera wagneri.
Barateli, N. & Tarknishvilii, D. & Iankoshvili, G. & Kokiashvili, L. & Dvali, N. & Janiashvili, Z. (2021) -
Two species of rock lizards, the parthenogenetic D. dahli and the sexually reproducing D. portschinskii, coexist in a rocky outcrop in an area of ca. 1 ha, in the vicinity of Tbilisi, Georgia; the location has been well-known since the middle 1960s. The population density of the parthenogenetic lizard is five times higher than that of the sexual breeder. We studied the distribution of active lizards in space and time over three consecutive years, during the spring and autumn activity periods, to explore spatial and temporal differ-ences between the species on a fine spatial scale. We studied the influence of temperature, humidity, and quantitative characteristics of the surface and the distance from permanent water source on the spatial distribution of D. dahli and D. portschinskii. Darevskia portschinskii was less dependent on the distance from the water source and more evenly distributed in space and time than D. dahli. Despite potential competitive interactions, the species did not avoid each other on the microhabitat scale, suggesting that the observed ecological differences are not caused by a niche shift. More individuals of the sexual breeder than individuals of the parthenogen were found in suboptimal habitats. This feature may increase the evolutionary success of D. portschinskii in a long-term perspective.
Bartenjew, A. & Rjeznikowa, M. (1931) -
Başoğlu, M. & Hellmich, W. (1970) -
Basoglu, M. & I. Baran (1977) -
Bauer, A.M. & Günther, R. (1995) -
A complete catalogue is provided for the type specimens of lacertid lizards in the herpetological collection of the Zoological Museum, Berlin, as of December, 1993. The collection contains a total of 514 type specimens, representing 63 taxa, of which 42 are currently regarded as valid at the specific or subspecific level. Types representing an additional four taxa appear to have been lost from the collection. The collections are especially rich in African lacertid types and in historically significant specimens collected early in the Nineteenth Century by Pallas, Eversmann, and Hemprich & Ehrenberg. Also present are the type series of many subspecies of Podarcis lilfordi and P. pityusensis described by Eisentraut.
Bedriaga, J. von (1878) -
Bedriaga, J. von (1879) -
Bedriaga, J. von (1886) -
Bedriaga, J. von (1895) -
Beshkov, V. & Beron, P. (1964) -
Beshkov, V. & Nanev, K. (2006) -
Beyer, T.V. (1968) -
Beyer, T.V. & Selinova, G.V. (1969) -
Bird, C.G. (1936) -
Birlik, S. & Yildirimhan, H.S. & Ilgaz, C. & Kumlutas, Y. (2018) -
In this study, we assessed the helminth fauna of seventy two Valentin’s Lizard, Darevskia valentini (32♂♂, 35♀♀, 5 subadult). Specimens collected from Kayseri, Ardahan and Van Provinces in Turkey. As a result of the present study, it was detected that forty one hosts are infected with one or more species of helminth. Two species of Cestoda, Oochoristica tuberculata and Mesocestoides spp., and 5 species of Nematoda, Spauligodon aloisei, Skrjabinodon alcaraziensis, Skrjabinodon medinae, Skrjabinelazia hoffmanni and Strongyloides darevsky were found in the hosts. D. valentini represents a new host record for all helminths recorded. Skrjabinodon alcaraziensis is recorded for the first time from Turkey. Van, Kayseri and Ardahan are new locality records for all helminths from D. valentini.
The present study investigated the composition of helminth parasites of Darevskia rudis, Spiny Tailed Lizard from Turkey. One hundred and two samples (49♀♀, 53♂♂) from the Tokat, Trabzon, Rize, Gümüşhane and Artvin Provinces were collected and examined for helminth parasites. New host and locality records were recorded. As a result of the present study, seven species of Nematoda, Skrjabinodon medinae, Spauligodon sp., Spauligodon carbonelli, Spauligodon aloisei, Skrjabinelazia hoffmanni, Strongyloides darevsky, Oswaldocruzia filiformis; one species of Cestoda, Mesocestoides spp. (tetrathyridium) and one species of Acanthocephala Sphaerirostris scanensis were reported from the lizard samples. Sp. carbonelli and Sphaerirostris scanensis are here recorded for the first time in Turkey. D. rudis is the new host recorded for Sk. medinae, Spauligodon sp., Sp. carbonelli, Sp. aloisei from Nematoda, Mesocestoides spp. from Cestoda and Sphaerirostris scanensis from Acanthocephala. This host has been studied for the first time for the helminth parasites from the Tokat and Gümüşhane Provinces. There is, to our knowledge, only one report of helminthes for D. rudis in Turkey.
Bischoff, W. (1973) -
Bischoff, W. (1974) -
Bischoff, W. (1975) -
Bischoff, W. (1976) -
Bischoff, W. (1978) -
The final part of a series of papers on the lizards of the Caucasus, that have been published in previous years.
The paper reports on the life habits, capture and keeping in terraria of several lizards of Soviet Transcaucasia. In these respects the agamid Phrynocephalus helioscopus persi- cus and three species of the lacertid genus Eremias occuring in that area (i. e. strauchi, velox, pleskei) are discussed. Finally a !ist of all lizard species and subspecies actually known to occur in this region is given.
Bischoff, W. (1982) -
Bischoff, W. (1984) -
An analysis based on 55 specimens of Lacerta derjugini from different localities on the southern slope of the Great Caucasus, all situated in the territory of the Georgian SSR/USSR, lead to the following results:
1. The distribution area of Lacerta derjugini boehmei BISCHOFF, 1982, is restricted to the westernmost part of the southern slope of the Caucasus.
2. Apart from the type locality (Suchumi) Lacerta derjugini abchasica BISCHOFF, 1982, also exists in the river valleys of the Kodori and Inguri.
3. The lizards living in the eastern part of the distribution area are described here as Lacerta derjugini orlowae ssp.n. (type locality: Pasanauri).
Bischoff, W. (2000) -
Bischoff, W. (2002) -
Bischoff, W. (2003) -
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.
Bischoff, W. (2006) -
Bischoff, W. (2011) -
Bischoff, W. (2012) -
Bischoff, W. & Deichsel, G. (2002) -
A museum specimen misidentified as a common wall lizard, Podarcis muralis, from Van Wert County, Ohio, USA was re-determined as a Caucasian rock lizard, Darevskia valentini. The specimen displays characteristics of Darevskia, the most notable of which are alternating broad and narrow whorls of scales around the tail. Scalation of the specimen is described and compared to sister taxa in detail.
Bischoff, W. & Franzen, M. & Schmidtler, J.F. (2005) -
We report on new locality records of Darevskia derjugini from the Turkish Pontus Mountains south of Dereli (Giresun province) and near the monastery of Meryemana (Trabzon province). The record from Giresun province extends the Turkish distribution some 130 km to the west. Examination of comparative material questions the taxonomic status of Darevskia derjugini barani to some degree, since characters separating this taxon from D. d. derjugini appear extremely variable and variation of pholidosis may be correlated with climate.
Bischoff, W. & Schmidtler, J.F. (2014) -
From Pallas to Darevsky: The exploration of the Caucasian herpetofauna using the example of the Saurians: The Caucasus, a wild mountain range bordering Europe and Asia, was a cue ball between the Russian, Ottoman and Persian empires up to the beginning of the 20th century. The general history of exploration was mainly initiated by the Russian governments and started at the end of the 18th century. The history of the discovery and exploration of the Caucasus, here especially of the Saurians, is documented. Our report is ending with the researches of Ilya S. Darevsky on
the saxiculous lizards of the genus Darevskia in the 1960ies. From the northern and southern slopes of
this mountain range (“Ciscaucasia” / “Transcaucasia”) 45 Saurian species are currently known. They belong to five families in which exactly two-thirds are members of the lacertids. These are the only ones having
developed numerous species and subspecies in the Caucasian range, and among them, beside Lacerta
agilis, especially the saxicolous lizards of the genus Darevskia are characteristical. 20 scientists are introduced on the basis of their most important publications and their profiles: Peter Simon Pallas (1741-1811), Samuel
Gottlieb Gmelin (1744-1774), Johann Anton von Güldenstädt (1745-1781), Eduard von Eichwald (1795-1876), Edouard Ménétriés (1802-1861), Eduard Eversmann (1794-1860), Alexander von Nordmann (1803-1866), Moritz Wagner (1813-1887),
Karl Kessler (1815-1881), Alexander Strauch (1832-1893), Jacques von Bedriaga (1854-1906), Gustav
Radde (1831-1903), Oskar Boettger (1844-1910), Alexander Mikhailovitch Nikolsky (1858-1942), Ludwig von Méhely (1862-1953), George Albert Boulenger (1858-1937), Louis Amédée Lantz (1886-1953), Otto Cyrén (1878-1946), Ilya Sergeyevitch Darevsky (1924-2009) and Günther Peters (* 1932).
Bischoff, W. & Tarkhnishvili, D. (2002) -
Georgia, its history and the nature are briefly described. This is supplied by a short overview of the country`s herpetological fauna, with particular attention to the especially diverse family of true lizards. At the end, brief description of our common trip to Georgia (May - Juny 2002) is given.
Biserkov, V. (2007) -
Biserkov, V. & Naumov, B. & Tsankov, N. & Stoyanov, A. & Petrov, B. &Dobrev, D. & Stoev , P. (2007) -
The current book provides a synthesis of the information concern-
ing field identification of all the species and subspecies of Amphibia
and Reptilia hitherto registered in Bulgaria. It comprises 61 species
and 23 subspecies, among which 5 species (Pelophylax lessonae, P.
kurtmuelleri, Hemidactylus turcicus, Eremias arguta, Montivipera
xanthina) that have not been found yet on the territory of the
country, but which are known from sites in proximity to Bulgar-
ian frontiers. The frogs and toads are represented with 14 spe-
cies, newts and salamanders with 6 species, turtles and tortoises
with 7 species (Trachemys scripta introduced; Caretta caretta and
Chelonia mydas considered extinct), and snakes with 19 species
(Vipera aspis and V. ursinii considered extinct). The structure of
each species’ description includes the following headings: Name,
Diagnosis, General Distribution, Distribution in Bulgaria, Habitats,
and Biology. Besides in Bulgarian and Latin, all species are sup-
plemented with their English, French, German and Russian names.
All synonyms in the main herpetological literature are listed after
the currently accepted Latin name. Original keys for identification
of the families, genera and species, based entirely on morphologi-
cal characters of adult specimens, as well as on the structure of
amphibians’ eggs and larvae, are elaborated to facilitate the users.
A photo gallery, comprising original colour photos shot in nature,
and numerous line drawings of different key characters are made
for better distinction between the species. For purely conservation-
al reasons, the distribution maps not only indicate the scientifically
confirmed species distribution in the country, but also show the
areas with suitable habitats where a given species is likely to oc-
cur. The degree of suitability is indicated with a different degree
of shading: optimal (black); suboptimal (80% gray); suitable (60%
gray); less suitable (20% gray); white means unsuitable. The guide
is also introducing the readers to some general aspects of the tax-
onomy, systematics, morphology, biology and ecology of herpeto-
fauna. Special chapters are devoted to the methods of observation
and capture of amphibians and reptiles, first aid after a snake’s
bite, and the relationship between herpetofauna – humans. The
status of each species according to main biodiversity conservation
acts and conventions is given in Table 1. Being the first publication
of its kind, the book is intended for professional herpetologists,
students or nature lovers.
Bobyn, M.L. & Darevsky, I.S. & Kupriyanova, L.A. & MacCulloch, R.D. & Upton, D.E. & Danielyan, F.D. & Murphy, R.W. (1996) -
Lacertca raddei and Lacerta nairensis have been recognized as two separate species based on morphology and behavior, and each has been implicated as a sexual parent of different parthenogenetic forms. However, recent mitochondrial DNA work failed to distinguish these two as separate species. We examined genetic diversity at 36 allozyme loci from six populations of L. nairensis and four populations of L. raddei. There were no fixed allelic differences between the two. Mean heterozygosity was slightly higher among populations of L. raddei than among populations of L. nairensis. A Distance Wagner phenogram showed that the northernmost population of L. raddei clustered with the L. nairensis populations; the other L. raddei populations clustered together. We suggest that L. raddei and L. nairensis may not be separate species, a finding which has important implications for determining the origins of some parthenogenetic Lacerta.
Bodenheimer, F.S. (1944) -
Boettger, O. (1880) -
Boettger, O. (1892) -
Boettger, O. (1893) -
Bogdan, H.V. (2012) -
Bogdan, H.V. & Sas-Kovács, I. & Covaciumarov, S.-D. (2014) -
In the southern side of the Lipova Hills we found 11 amphibian species, 9 reptile species and 2 types of hybrid between amphibian species. The herpetofauna of the region contains lowland, hill and mountain related species. Also, in the area are present southern species in the vicinity of the northern limit of their distribution range, and related species in the mountainous areas, found at lower altitudes than usually. The mixture of mountain and lowland species is a consequence of the current aspect of the region, with wide valleys located between hill ridges, valleys that allow the advancement of these lowland species in the hill areas. The second aspect is a consequence of the history of the region. Currently the herpetofauna of the Lipova Hills is greatly advantaged by the relatively large forest covered areas.
Bogdanova, G.O. (1974) -
Bogdanova, G.O. (1976) -
Böhme, W. (2010) -
In the herpetological collection of ZFMK 528 scientific species group names are represented by type materi- al. Of these, 304 names are documented by primary type specimens (onomatophores) while for 224 further names sec- ondary type specimens (typoids) are available, ranging chronologically from 1801 to 2010. The list is a shortened pred- ecessor of a comprehensive type catalogue in progress. It lists name bearing types with their catalogue numbers includ- ing information on further type series members also in other institutions, while secondary types are listed only by pres- ence, both in ZFMK and other collections including holotype repositories. Geographic origin and currently valid names are also provided.
Böhme, W. & Bischoff, W. (1984) -
Böhme, W. & Budak, A. (1977) -
1. Reinvestigation of the syntypes of Lacerta rudis BEDRIAGA 1886 (=nomen novum pro Lacerta depressa CAMERANO 1878, cf. BuDAK & BöHME 1977) and the comparison with a topotypical series show, that only one form is involved. As lectotype for the species, thus for the nominate form, too, the cf` specimen MIZS (Torino) 2737:1 is designated.
2. The populations of NE-Anatolia and Transcaucasia, so far misinterpreted as being the nominate form, represent a distinct form and are described as Lacerta rudis bischoffi n. subsp.
Bonaparte, C.L. (1839) -
Borkin, L.Ya. & Darevsky, I.S. (1987) -
Боркин Л.Я. & Даревский И.С. (1987) -
Bosch, H.A.J. in den (1995) -
L. unisexualis produces one clutch yearly, with 1 - 5 eggs (3.4 on average). Fertile egg measure 7.5 x 12.3 mm and weigh 0.43 g. The eggs grow to 11.1 x 18.9 am and 1.51 g and hatch after 51 days at 25°C und after 40 days at 29°C. The hatchlings measure 27 + 47 mm (snout-vent length + tail) and weigh 0.49 g. In captivity sexual maturity was reached in the second year. Some colour characteristics, especially blue axillary spots, are mentioned.
Bosch, H.A.J. in den (1999) -
Reproduetion of thc parthenogcnctic fonns Lacena armeniaca and Lacerta wzise.rtwlis undcr caplive conditions is documcnted. A short review of parthenogenesis is givcn and thc origin of known parthogcnctic forms with- in the genus Lacerta is discussed. Thc rcccnt- ly described fonns from castem Turkcy. Lacerta henclimalliensis and Lacerta sapphiri- tw. an: illustrated.
Thc current common opinion is that partheno- gcnctic Lacerta fonns found in the eaucasus region are thc rcsult of crosses hetween diffe- rent species. Thc parental species of L. uni- .H`Xtwlis would he Locatanairensis (phenoty- pically vcry similar to L tmisexualis) and utn•rta mlemini. and for L. armeniaca prc- sumahly utcerta mixta and L mlemini.
lt is rcmarkahlc that parthcnogenetic rcptilcs tcnd to he found in arcas wherc major cata- strophes occur regularly. or have occurred in the recent past. Thc ice agcs are mentioned in litcraturc as a possiblc factor driving par- thenogenesis in the eaucasus. The new Turkish uniscxual utcerta forms occur vcry close to a lakc which most prohably has histo- rically had major lluctuations in water leveL Unstahle water levels in rivers along which some L.uni.H`Xtwlis live. is mentioncd in litera- rature in thc samc context. In othcr words. an unstahlc cnvironment secms to fa`our parthe- nogcnetic forms.
Data on capti`e animals rcfute some curious misconccptions conccrning parthcnogenctic lizards. lt is somelimes said that a minimum numhcr of animals is rcquired before repro- duetion occurs hut this has nol proven truc for L. armnziaca and L. tmise.rttalis. Partheno- gcnetic animals are said to die relativcly young. hut at six years of age my L unisexu- alis are still hcalthy and in lïne reproductive condition. Hushandry is reputed to be diftï- cult. hut this is nol so in L. lmisexualis: 68 young have heen reared from 80 eggs - I stop- ped incuhating thc cggs because of a major surplus in young.
lt has heen proposed in literaturc that the suc- ccss of unisexual forms is partially related to time savcd hy nol copulating. Although some time may he saved in not needing to locale a mate. the time gain from nol performing courtship is small. as in closely rclated eaucasus spedes this takes only minutes. The assumption found in some literaturc that onc of the advantages parthenogcnetic forms have over their parental species is a shorter incuba- tion duration. is refutcd for L. unisexualis and partially for L. ttnneniaca (not having onc of the parental species. L mixta).
Although hoth of these parthcnogenetic spe- cies are reputed to occur in high densities. it is often wrongly assumed that they are not aggressive. They are in fact quite aggressive towards other lizards as well as towards con- specitïcs during the reproductivc period. This is particularly ohvious immcdiatcly after ovi- position. when they chase off any intruder that comes too near them or the nest- some at- tackcd lizards have even lost legs in such encounters.
Brecding of L amu•niaca proved dirticuit in that only four young resulted from ..J-3 eggs produced. Possihly a number of dcfects had accumulatcd in thc starting material: these fcmalcs all slmwed major pbolidosis anoma- lics. missing locs. and sornc had short. stuhby tails. Thc young showcd similar dcfects (with up to 15 tocs missing). The adults measured 51-62 rnm (hcadhody) + 57-152 mm (tail). Weigllls varicd from 4.9 g to 5.7 g outside the reproductivc scason (September). Each fenmie oviposited once a ycar (around Aprii-May). An average clutch contained 3.6 eggs (2-5 cggs): healthy cggs rneasured 7.6x 12.5 mm and wcighed 0.42 g at oviposition. The eggs rcached II.XxI6.X 111111 and 1.37 g during dcvclopmcnt. They hatched after 46 and 48 days whcn incuhatcd at 2Y`C. and after 37 days at 29°C. The ncwly hatched juveniles 111easured 25.5 111111 head-body + 25.7 mm tail (and thus had an ahnormal tail length in this sample). and wcighcd 0.43 g. The juveniles resembied thc adults hut for the blueish- greenish tails. and the Jack of bluespots on thc tlanks and on the outer vcntrals. They pro- oueed thcir lirst cggs in their secmld year. Husbandry of L. uni.,·e,rtwlis is remarkahly easy. None has shown the aberrations mentio- ned for L. an11e11iam. Of the eighty eggs incu- hated. 6X resultcd in healthy juveniles. Thc adult /.. tmisexualis measured 52-66 mm (head-hody) + 121-1-1-2 111111 (laiI). Wcights varied from 4.92 g to 7.72 g outside the repro- ductivc season (September). Each female ovi-posited once a year (a sec011d clutch of unfcr- tile eggs was twice found); around Aprii-May. An avcrage clutch contained 4-5 eggs. Healthy cggs measured 7.5x 12.3 mm and weighed 0.42 gat oviposition. These reached 10.9x 16.6 mm and 1.33 g during development. Eggs hatched after 48 days when incubated at 25°C, and after 38 days at 29°C. The newly hatched juveniles measured 27.0 mm (head-body) + 47.6 mm (tail), and weighed 0.46 g. The juve- niles resembied the adults but for the very vaguely blueish-greenish tails; they also lacked the yellowish ventral colour and the blue spots were still very pale. Young pro- duced their tirst fertile eggs in their second ycar. Both the eggs and the resulting juveniles in both species are slightly larger than in sexu- ally rcproducing lacertids of comparable size. This would confer a competitive advantage.
Bosch, H.A.J. in den (2001) -
The phenomenon of twins is very rare in lacertids. This conclusion was reached by looking at a recent instance of twins hatching from a single egg in Lacerta nairensis, an older but comparable occurrence in Algyroides fitzingeri, as well as my personal experience over many years of incubating lizard eggs. It is estimated that twins occur in 1:5000 to 1:500 eggs.
Bosch, H.A.J. in den & Bischoff, W. (1995) -
The courtship of Lacerta chlorogaster includes a hind-leg bite, clearly different from the flank bite of Lacerta laevis. Each female can lay two clutches per year, with 3-6 eggs. The eggs measure 6.9 x 10.7 mm und have a weight of 0.31 g. They grow to 10.9 x 17.0 mm und 1.15 g. Hatchlings appear after 55 days at 25°C und 42 days at 29°C; they measure 24.3 + 40.0 mm (SV+T) und 0.34 g. Adult males change their colors from dull brown dorsally to gaudy green, three to four wecks after reappearing from hibernation. The greenish yellow ventral colour changes into blue-green. At mid June the bright colours fade relatively rapidly. In females und immature subadults the seasonal colour change is much less expressed.
Bosch, H.A.J. in den & Bischoff, W. (2004) -
Many regions of the former USSR still re- ceive little attention from Western visitors, herpetologists not excluded. For the Cau- casus Mountains this is all the more sur- prising since the area forms a zone where northern and Middle Eastern flora and fauna meet, and which is thus of considera- bly zoogeographical interest.
Presently we find three larger states in the Caucasus: Georgia, Azerbaijan and Arme- nia. Turkey borders to the southwest, Iran to the southeast, and in the north it is the Russian federation. Various smaller parts claim independence. The Black Sea forms the natural western boundary, the Caspian Sea the eastern one.
Herpetologically the Caucasus is historically well-known because of the work of Darev- sky and collaborators on the rock lizards (at the time still in the large genus Lacerta, now in Darevskia) and the discovery of the first case of parthenogenesis in vertebrates (DAREVSKY, 1967; DAREVSKII, 1978; DAREVSKY & KULIKOVA, 1961).
Below we will concentrate on the results of our field trip to Georgia during the last two weeks of June 2003. Predictably, because of our common interest in the group, the lacertids will receive the most attention.
Boulenger, G.A. (1904) -
Boulenger, G.A. (1907) -
Boulenger, G.A. (1908) -
Boulenger, G.A. (1910) -
Boulenger, G.A. (1913) -
Boulenger, G.A. (1920) -
Brehm, A.E. (1902) -
Brown, R.P. (2005) -
A phylogenetic-comparative approach was used to assess and refine existing secondary structure models for a frequently studied region of the mitochondrial encoded large subunit (16S) rRNA in two large lizard lineages within the Scincomorpha, namely the Scincidae and the Lacertidae. Potential pairings and mutual information were analyzed to identify site interactions present within each lineage and provide consensus secondary structures. Many of the interactions proposed by previous models were supported, but several refinements were possible. The consensus structures allowed a detailed analysis of rRNA sequence evolution. Phylogenetic trees were inferred from Bayesian analyses of all sites, and the topologies used for maximum likelihood estimation of sequence evolution parameters. Assigning gamma-distributed relative rate categories to all interacting sites that were homologous between lineages revealed substantial differences between helices. In both lineages, sites within helix G2 were mostly conserved, while those within helix E18 evolved rapidly. Clear evidence of substantial site-specific rate variation (covarion-like evolution) was also detected, although this was not strongly associated with specific helices. This study, in conjunction with comparable findings on different, higher-level taxa, supports the ubiquitous nature of site-specific rate variation in this gene and justifies the incorporation of covarion models in phylogenetic inference.
Budak, A. & Böhme, W. (1978) -
New material of Turkish rock-lizards of the rudis-group (Lacerta semcoZa-complex)
can be assigned to the forms r. rudis, r. tristis and r. obscura; thus, the last named form
is recorded for the first time for Turkey. Geographically intermediate samples could be
interpreted as clinal, connecting r. rudis and r. tristis, but it turns out to be more likely,
that there are two sympatric different taxa overlapping in the intermediate zone. This
possibility which could not be proved until now leads to several extraordinaryly difficult
nomenclatorial conclusions. They will be discussed further in a following contribution.
Bülbül, U. & Koc, H. & Orhan, Y. & Odabas, Y. & Kutrup, B. (2019) -
It is a fact that the amphibians and reptiles respond to unfavourable weather conditions by searching an underground refuge to survive during winter. The current literature has shown that some species of reptilians and amphibians may become active before their known end of hibernation periods, especially in lowland areas. In the present study, it has been detected that the changing weather conditions in a highland area in Gümüshane, Turkey can cause similar effects on Lacerta media, Bufo bufo and Rana macrocnemis that share the same habitats. One of the reasons of the early activities of some reptilian and amphibian species before their known end of hibernation periods may be the high sensitivity of these species to the changing temperature conditions. Effects of global warming on reptilians and amphibians obvious, and some members of these animals can even be active during winter. This conclusion brings into mind that L. media, B. bufo and R. macrocnemis can continue to be active at highland areas as long as air temperature values allow them.
Bülbül, U. & Kurnaz, M. & Eroğlu, A.I. & Koç, H. & Kutrup, B. (2016) -
In the present study, we investigated potential effects of some environmental climatic factors on life-history traits of two Darevskia parvula populations located at different elevation sites and evaluated age, body size, SSD and growth rate results in these lizards. The age at maturity and longevity were found to be similar in both populations. Males of the highland population had significantly larger snout-vent length and older age than those of the lowland population. Although the mean ages of females were not significantly different between the two populations, the highland population had significantly larger snout-vent length than the lowland population. Considering all individuals (both males and females), we found that specimens from the population in the higher-elevation site had significantly larger snout-vent length and older age than those of the lower-elevation site. A low level of male-biased sexual size dimorphism was observed in both populations. The growth coefficient was lower in the high-elevation site than the lower elevation site. Growth rates were not significantly different between the populations. Our preliminary results indicate that although our studied populations were located at different elevation sites and had distinct climate, longevity, age at sexual maturity and growth rate results were similar. This may be a reason of the lower male-biased sexual size dimorphism in both populations.
Bülbül, U. & Kurnaz, M. & Eroğlu, A.I. & Koc, H. & Kutrup, B. (2016) -
We investigated age structure, body size and longevity in two breeding populations of Darevskia clarkorum inhabiting altitudes ranging from 450 m a.s.l. (Kamilet) to 2250 m a.s.l. (Basyayla) in Turkey by skeletochronology performed on the phalanges. The mean age was found to be 6 years in the Kamilet population and 7 years in the Basyayla population. The maximum life span was 10 years in the lowland population while it was 12 years in the highland population. Age at sexual maturity of both males and females was 1-2 years in the lowland population while it was 2-3 for both sexes in the highland population. Both age and SVL of specimens from the Kamilet population were signiﬁcantly different between the sexes while age and SVL did not differ signiﬁcantly between the sexes in Basyayla population. As a conclusion, we observed that the mean age, longevity and age at maturity were increased by altitude while there was a decrease based on the mean SVL in the highland population of D. clarkorum. Our data on body size, longevity and age at sexual maturity may contribute to conservation efforts for this endangered species.
Bunyatova, S.N. & Dzhafarova, S.K. (2019) -
Background. From the genus Darevskia in the territory of Azerbaijan Republic noted 11 species (D. armeniaca, D. caucasica, D. chlorogaster, D. daghestanica, D. derjugini, D. portschinskii, D. praticola, D. raddei, D. rostombekovi, D. rudis, D. valentini). Information on the condition of rock lizards in the republic has not been updated for a long time. The aim of the work was to clarify the data on the distribution and abundance of these species in Azerbaijan.
Materials and methods. The research was carried in the Talysh Mountains (Lenkoran, Lerik and Yardimli districts of Azerbaijan in 2008–2011), in the Great Caucasus (Guba, Gusar, Gabala and Zagatala districts in 2013–2016), in the Minor Caucasus (Dashkesen, Gadabay and Tovuz districts in 2017–2018).
Results. On the basis of own data and analysis of literature sources the data on the distribution and number of rock lizards from the genus Darevskia in Azerbaijan are given. The authors note that the areas of forest lizards of the Talysh (D. chlorogaster and D. praticola hyrcanica) are being reduced and these species need legisla- tive protection. The extinct species for the territory of the Republic should be considered D. derjugini.
Бунятова, С.Н. & Джафарова, С.К. (2019) -
Актуальность и цели. Из представителей рода Darevskia на территории Азербайджанской Республики отмечены 11 видов (D. armeniaca, D. caucasica, D. chlorogaster, D. daghestanica, D. derjugini, D. portschinskii, D. praticola, D. raddei, D. rostombekovi, D. rudis, D. valentini). Сведения по состоянию скаль- ных ящериц в стране длительное время не обновлялись. Целью работы явля- лось уточнение данных по распространению и численности этих видов в Азер- байджане.
Материалы и методы. Исследования проводили в Талышских горах (Ленко- ранский, Лерикский и Ярдымлинский районы Азербайджана в 2008–2011 гг.), на Большом Кавказе (Губинский, Гусарский, Габалинский и Закатальский рай- оны в 2013–2016 гг.), на Малом Кавказе (Дашкесанский, Кедабекский и Товуз- ский районы в 2017–2018 гг.).
Результаты. На основании собственных данных и анализа литературных источников приводятся сведения по распространению и численности скальных ящериц рода Darevskia в Азербайджане. Авторы отмечают, что сокращаются ареалы лесных ящериц Талыша (D. chlorogaster и D. praticola hyrcanica) и они нуждаются в законодательной охране. Вымершим видом для территории рес- публики следует считать D. derjugini.
Buresch, I. & Zonkow, J. (1933) -
Busack, S.D. & Salvador, A. & Bauer, A.M. & Kaiser, H. (2016) -
A doctoral thesis presented in the late 1990s by óscar J. Arribas has been frequently cited as the authority for the nomenclatural status of two generic names in the lizard family Lacertidae, Darevskia and Iberolacerta. These names were again introduced, along with the addition of the subgeneric name Pyrenesaura, in 1999 via publication in an international, peer-reviewed journal. We discuss nomenclatural aspects of how these taxon names were presented, detail inconsistencies in the use of publication dates associated with these names, and resolve nomenclatural problems by demonstrating that the date of their first publication, in compliance with the International Code of Zoological Nomenclature, was 1999. This renders the name Darevskia Arribas, 1999 a junior invalid synonym of Caucasilacerta Harris, Arnold & Thomas, 1998, which, contrary to previous assessment, is not a nomen nudum, and must replace Darevskia as the valid name for this genus.
Cakmak, M. & Akman, B. & Yildiz, M.Z. (2017) -
In this investigation, we aimed to determine the amphibian and reptile species distributed in Bartın province (Northwest Blacksea Region of Turkey). As a result of previously and current studies, we recorded two Urodela amphibian species (Ommatotriton ophryticus and Triturus ivanbureschi), six Anura amphibian species (Bufo bufo, Bufotes variabilis, Hyla orientalis, Pelophylax ridibundus, Rana macrocnemis and R. dalmatina), three turtle species (Emys orbicularis, Mauremys rivulata and Testudo graeca), four lizard species (Anguis fragilis, Darevskia rudis, Lacerta viridis and Podarcis muralis) and eight snake species (Coronella austriaca, Dolichophis caspius, Eirenis modestus, Natrix natrix, N. tessellata, N. megalocephala, Vipera transcaucasiana and Zamenis longissimus) in Bartın Province, which totally 23 species. There are no endemic species in our country among them. Natrix megalocephala is also a new record for Bartın province.
Camerano, L. (1877) -
Candan, K. & Gül, S. & Kumlutas, Y. & Yildirim Caynak, E. & Ilgaz, C. (2020) -
Darevskia parvulais a rock lizard that is endemicfor Anatolia.The known distribution range of the species is limited on eastern and northeastern Anatolia. Although many morphological studies have been carried out onthe species, there are also molecular studies to construct its taxonomy in recent years. Four adult lizard specimens were collected from eastern Anatolia in 2016 during a herpetological field survey. We present a summary of a morphological features, and report new locality which is the westernmost record (Çayırlı Village, Erzincan) for D. parvula sensu lato in Turkey. Our finding largely extends the known distribution of the species.
Candan, K. & Kornilios, P. & Ayaz, D. & Kumlutas, Y. & Gül, S, & Yildirim Caynak, E. & Ilgaz, C. (2021) -
Taxonomy of the lacertid genus Darevskia has been complex since the first morphology-based studies. Darevskia valentini, mostly distributed in Turkey, currently includes three subspecies, but its interspecific and intraspecific taxonomic status has so far been shaped only by morphological evaluations, without molecular phylogenetic assessments. In this study, samples from the entire distribution of this species and its closest relatives were evaluated with molecular phylogenetic analyses of a multi-marker dataset, comprised of maternally-inherited mitochondrial and biparentally-inherited nuclear markers. We discovered two previously unrecognized genetically highly divergent lineages. The distinction of these two lineages is supported by gene trees, haplotype networks, divergence time estimation and species-delimitation analyses. We discuss possible taxonomic resolutions given that the phylogenies do not seem to agree with currently accepted taxonomy. Our results advance the knowledge of the ancestry of parthenogenesis as a result of past hybridization events between parental species pairs, because the paternal origin of some parthenogenetic species lies within the new lineages discovered here.
Candan, K. & Kornilios, P. & Yanhukov, A. & Kumlutas, Y. & Ayaz, D. & Tarknishvili, D. & Murtskhvaladze, M. & Kurdaze, S. & Barateli, N. & Gabelaia, M. & Sahin, M.K. & Matur, F. & Gül, S. & Yildirim Caynak, E. & Colak, F. & Ilgaz, C. (2019) -
Darevskia valentini is the most widely distributed species among the Caucasian rock lizard but the phylogenetic relationship among geographically distant populations is poorly known. W sequenced several mitochondrial (12S rRNA, COI an cyt b) andnuclear (MC1R and cmos) genes from 75 individual samples collected across Central and Eastern Turkey. In total, 349 bp of 12S rRNA, 615 bp of COI, 743 bp of cyt b of mtDNA and 620 bp of MC1R, 326 bp of cmos of nDNA were obtained. Additionally, 34 individuals from Georgia and Eastern Turkey were genotyped for 10 microsatellite loci. Phylogenetic analyses (ML and BI) produced trees with similar topologies including well-supported clades. One clade found in the area of Lake Van is clearly separated from others on the phylogenetic tree, and the respective population formed distinct STRUCTURE and PCA clusters in the microsatellite data. Our results may suggest that D. valentini, one of the most complex species in Darevskia genus, needs taxonomical revision at the subspecies level.
Capriglione, T. & Cardone, A. & Odierna, G. & Olmo, E. (1994) -
This paper reports the isolation and characterization of twoHindIII repetitive DNA families from the genome of two lacertid lizards,Podarcis sicula andLacerta saxicola. These satellites did not appear to be related to each other. The consensus sequences of their monomeric units did not show any similarity, though both DNAs were A-T rich. Moreover, each of them was found only in closely related species. The monomeric unit of theHindIII DNA family isolated fromP. sicula (pLHS) showed a close resemblance to pLCS, a centromeric satellite DNA previously isolated from the same species; it was, however, mainly localized at pericentromeric, interstitial and telomeric levels. The results also provide interesting information on the systematics of the lacertids studied.
Carretero, M.A. (2018) -
Carretero, M.A. & Garcia-Munoz, E. & Argana, E. & Freitas, S. & Corti, C. & Arakelyan, M. & Sillero, N. (2016) -
Several Caucasian rock lizards of the genus Darevskia are well-known to reproduce parthenogenetically and have a hybrid origin. While one or more of these all-female parthenogens may monopolize local lizard communities, they may also occur in syntopy with bisexual members of the genus. In several localities, effective reproduction between bisexual and parthenogenetic Darevskia has been reported based on lizard intermediate morphology and karyology (3n, 4n) but frequency of such heterospecific matings is still unknown. In a mixed Darevskia community from Kuchak (Armenia) constituted by two parthenogens (D. armeniaca and D. unisexualis), one bisexual species (D. valentini) and their putative backcrosses, we indirectly quantified the reproductive interactions through the inspection of copulation marks in females. A total of 114 adult females were randomly collected, photographed and later inspected for inguinal marks. Females were measured (SVL) and their marks were ranked twice from 0 (no scars) to 3 (≥ 3 scars). The lizard determination and ploidy was ensured by microsatellites analysis. All female types displayed copulation marks with frequencies varying from 61% in the parthenogenetic D. armeniaca to 30% in the bisexual D. valentini; remarkably, 7 out of 11 (64%) backcross females also had marks. In the most abundant D. armeniaca, the prevalence and intensity of copulation marks increased with size, just as expected for normal female lacertids. These results indicate that copulation of parthenogenetic Darevskia in mixed communities with bisexual species is not an isolate event, thus reinforcing previous suggestions of massive reproductive interaction in syntopy. Evolutionary perspectives of this phenomenon are discussed.
Carretero, M.A. & García-Muñoz, E. & Argaña, E. & Freitas, S. & Corti, C. & Arakelyan, M. & Sillero, N. (2018) -
Several Caucasian rock lizards of the genus Darevskia of hybrid origin are known to reproduce parthenogenetically. Local communities can be composed exclusively of parthenogens, though syntopy with bisexual members of the genus may occur. In some localities, reproduction between bisexual and parthenogenetic Darevskia has been previously reported based on lizard intermediate morphology and karyology (3n, 4n). However, the frequency of such heterospecific matings remains unknown. We indirectly quantified the reproductive interactions through the inspection of copulation marks in females in a mixed Darevskia community from Kuchak (Armenia) composed of two hybrid parthenogens (D. armeniaca and D. unisexualis), one bisexual species (D. valentini) and their putative backcrosses. A total of 139 adults were randomly collected and photographed. Females were later measured (SVL), inspected for inguinal marks and ranked from 0 (no scars) to 3 (≥ 3 scars). The lizard species and ploidy determination was ensured by a parallel microsatellite analysis. Sex-ratio in the community was extremely biased due to the high abundance of parthenogenetic females. All female types displayed copulation marks with frequencies varying from 80% in D. valentini to 64% in D. armeniaca. Remarkably, 7 out of 11 (64%) backcross females also showed marks. In the most abundant D. armeniaca, the prevalence and intensity of copulation marks increased with body size, just as predicted for polygynous female lacertids. These results indicate that copulation between parthenogenetic and bisexual species in Darevskia mixed communities is common and driven by sexual selection, thus reinforcing previous suggestions of reproductive interaction in syntopy. Evolutionary implications of these findings are discussed.
Çavaş, Ç.K. & Kumlutaş, Y. & Candan, K. & Ilgaz, Ç. (2018) -
The aim of the study is to predict the gender of Darevskia bithynica by using a feed-forward back-propagation artificial neural network (ANN). Nine morphological characters were used as an input parameters of the mo¬del. The gender type male or female is the output parameter. The total number of data is 115. In order to train, validate and test the ANN model 70%, 15% and 15% of the total data are randomly selected, respectively. The regression coefficient (R) values are evaluated as prediction performance. The network’s layer with tangent sigmoid activation functions predicts the lizard gender with R values as 0.98, 0.97 and 0.96 for training, tes¬ting and all data, respectively. The mean square error (MSE) values for training and testing data are calculated as 0.0145 and 0.0161, respectively. The obtained results satisfactorily confirm the high ability of the ANNs in predicting the gender of Darevskia bithynica.
Çavaş, Ç.K. & Kumlutaş, Y. & Candan, K. & Ilgaz, Ç. (2019) -
The aim of the study is to predict the gender of Darevskia bithynica by using a feed-forward back-propagation artificial neural network (ANN). Nine morphological characters were used as an input parameters of the mo-del. The gender type male or female is the output parameter. The total number of data is 115. In order to train, validate and test the ANN model 70%, 15% and 15% of the total data are randomly selected, respectively. The regression coefficient (R) values are evaluated as prediction performance. The network’s layer with tangent sigmoid activation functions predicts the lizard gender with R values as 0.98, 0.97 and 0.96 for training, tes-ting and all data, respectively. The mean square error (MSE) values for training and testing data are calculated as 0.0145 and 0.0161, respectively. The obtained results satisfactorily confirm the high ability of the ANNs in predicting the gender of Darevskia bithynica.
Çevic, I.E. (1999) -
In this investigation, lizard species of Turkish Thrace are investigated indetail for the first time. Morphological
characretirstics, range, distribution and subspecific states of the species are given.
Furthermore, some observations concerning their biological and ecological characteristics are also given. In this study a total of 678
specimens and 10 species belonging to 3 families of Lacertilia have been investigated from Turkish Trace.
Çevik, I.E. (1999) -
Araştırmada Trakya Bölgesinin kertenkele türleri ayrıntılı olarak incelenmiştir. İlk önce adı geçen bölgede türlerin tespiti yapılmış ve bunların morfolojik karakterleri, dağılışları ve alttür durumları açıklanmıştır. Ayrıca Biyolojik ve ekolojik özelliklerden gözlenebilenler hakkında bilgiler verilmiştir. Bu çalışmada 3 familya’dan 10 tür ve toplam 678 örnek değerlendirilmiştir.
Çevic, I.E. & Tok, C.V. (1997) -
In the present study forty-four specimens of Lacerta praticola were collected from Dereköy (Kırklareli) and were investigated in detail in the view of morphological features. The results were compared with the existing knowledge of literature. The population was decided to be classified as a subspecies of L.p. pontica. Additionally, some information about the ecology of the species are given.
Choopani, A. & Hojati, V. & Shiravi, A. (2014) -
Spermatogenic cycle in the Green-bellied lizard, Darevskia chlorogaster, was studied in forests of Sari County in Mazandaran Province, northern Iran. Sampling took place periodically every 15 days during the activity period of this species, from 5 April to 20 October 2012. In total, 59 adult males were captured by hand. Testes were removed and processed for histological and morphometric studies. The results show that testes consist of 74.45 seminiferous tubules in average. The mean diameters of seminiferous tubules, lumen, tunica albuginea and germinal layer are 350.14, 162, 17.69 and 64.10 microns, respectively. Spermatogenesis begins in mid March to early April, reaching its peak in late May and early June, and ends between early to mid August. The maximum level of sperm production occurred in early June. The minimum diameter, weight and volume of testes were observed in early August. Four phases were observed during the spermatogenesis of this species: active, transitional, inactive and regenerative. Spermatogenesis of D. chlorogaster in Iran is seasonal and the type of spermatogenic cycle is associated.
Cicek, K. & Candan, K. & Yildirim, E. & Kumlutas, Y. & Ilgaz, C. (2020) -
The study presents data on the food composition of the Unisexual Lizard, Darevskia unisexualis (Darevsky, 1966), from Hanak, Ardahan, Turkey. A total of 172 preys were determined in the digestive systems of 44 individuals examined in the study. Insecta, Aranea, Malacostraca, and Gastropoda constitute food composition of lizards. Major prey groups in the food composition are included in Hymenoptera (46.5%), Coleoptera (13.4%), Diptera (8.1%), Lepidoptera (7.6%), Orthoptera (7%) and Hemiptera (7%) in number, whereas Hymenoptera (70%), Lepidoptera (25%), Coleoptera (23%), and Diptera (20%) in frequency. Our result confirm that the Unisexual Lizard is an insectivorous and active predator and generally feed with poor flying or non-flying invertebrates.
Ciobanu, D.G. & Grechko, V.V. & Darevsky, I.S. (2003) -
The structure and evolution of a satellite DNA family was examined in lizards from the genus Darevskia(family Lacertidae). Comparison of tandem units of repeated DNA (satDNA), CLsat, in all species from the genus Darevskiahas shown that their variability is largely explained by single-nucleotide substitutions, which form about 50 diagnostic positions underlying classification of the family into three subfamilies. Maximum differences between the subfamilies reached 25%. At this level of tandem unit divergence in the subfamilies, no cross-hybridization between them was observed (at 65°C). The individual variability within one subfamily within the species was on average 5% while the variability between species consensuses within a subfamily was 10%. The presence of highly conserved regions in all monomers and some features of their organization show that satellites of all Darevskia species belong to one satDNA family. The organization of unit sequences of satellites CLsat and Agi160 also detected by us in another lizard genus, Lacerta s. str. was compared. Similarity that was found between these satellites suggests their relatedness and common origin. A possible pathway of evolution of these two satDNA families is proposed. The distribution and content of CLsat repeat subfamilies in all species of the genus was examined by Southern hybridization. Seven species had mainly CLsatI (83 to 96%); three species, approximately equal amounts of CLsatI and CLsatIII (the admixture of CLsatII was 2–5%); and five species, a combination of all three subfamilies in highly varying proportions. Based on these results as well as on zoogeographic views on the taxonomy and phylogeny of the Darevskia species, hypotheses on the evolution of molecular-genetic relationships within this genus are advanced.
Чобану Д.Г., Гречко В.В., Даревский И.С. (2003) -
Исследовали структуру и эволюцию семейства сателлитной ДНК ящериц рода Darevskia сем. Lac-ertidae. При сравнительном анализе последовательностей мономеров тандемных повторов ДНК (сатДНК) - CLsat - всех видов ящериц р. Darevskia показано, что вариабельность повторов обусловлена главным образом однонуклеотидными заменами, которые образуют около 50 диагностических позиций, на основании которых выделены три подсемейства повторов. Максимальные значения различий между подсемействами достигают 25%. При такой степени дивергенции мономеров трех подсемейств перекрестной гибридизации (при 65°С) между ними не наблюдается. Индивидуальная вариабельность мономеров одного подсемейства внутри вида в среднем составляет 5%, а вариабельность между видовыми консенсусами внутри подсемейства - 10%. Наличие высококонсервативных областей во всех мономерах и ряд особенностей их организации позволяют рассматривать сателлиты всех видов р. Darevskia как единое семейство сатДНК. Приведено сравнение организации последовательности мономеров сателлита CLsat и обнаруженного нами Agi160 из другого рода ящериц -Lacerta s. str. Выявлено сходство, свидетельствующее о родстве этих сателлитов и об их общем происхождении, предложен вероятный путь эволюции этих двух семейств сатДНК. Методом Саузерн-гибридизации изучено распространение и содержание подсемейств повторов CLsat во всех видах рода. Семь видов содержат в основном CLsatl (от 83 до 96%), три вида - примерно равные количества CLsatl и CLsatIII (примесь CLsatII составляет 2-5%), а пять видов - сумму всех трех подсемейств в сильно отличающихся соотношениях. На основе этих данных, в совокупности с зоогеографически-ми представлениями о родстве видов р. Darevskia, предложены гипотезы эволюции молекулярно-генетического родства видов этого рода.
Ciobanu, D.G. & Grechko, V.V. & Kramerov, D.A. & Darevsky, I.S. (2003) -
Ciobanu, D.G. & Roudykh, I.A. & Ryabinina, N.L. & Grechko, V.V. & Kramerov, D.A. & Darevsky, I.S. (2002) -
The genetic relatedness of several bisexual and of four unisexual Lacerta saxicola complexlizards was studied, using monomer sequences of the complex-specific CLsat tandem repeats and anonymous RAPD markers. Genomes of parthenospecies were shown to include different satellite monomers. The structure of each such monomer is specific for a certain pair of bisexual species. This fact might be interpreted in favor of co-dominant inheritance of these markers in bisexual species hybridogenesis. This idea is supported by the results obtained with RAPD markers; i.e., unisexual species genomes include only the loci characteristic of certain bisexual species. At the same time, in neither case parthenospecies possess specific, autoapomorphic loci that were not present in this or that bisexual species.
Clark, R.J. & Clark, E.D. (1973) -
Clark, R.J. & Clark, E.D. & Anderson, S.C. (1966) -
Cogălniceanu, D. & Rozylowicz, L. & Székely, P. & Samoilă, C. & Stănescu, F. & Tudor, M. & Székely, D. & Iosif, R. (2013) -
e reptile fauna of Romania comprises 23 species, out of which 12 species reach here the limit of their geographic range. We compiled and updated a national database of the reptile species occurrences from a variety of sources including our own eld surveys, personal communication from specialists, museum collections and the scienti c literature. e occurrence records were georeferenced and stored in a geoda- tabase for additional analysis of their spatial patterns. e spatial analysis revealed a biased sampling e ort concentrated in various protected areas, and de cient in the vast agricultural areas of the southern part of Romania. e patterns of species richness showed a higher number of species in the warmer and drier regions, and a relatively low number of species in the rest of the country. Our database provides a starting point for further analyses, and represents a reliable tool for drafting conservation plans.
Corovic, J. & Cogalniceanu, M. & Carretero, M.A. & Crnobrnja-Isailovic, J. (2018) -
One of the most interesting but elusive members of the European herpetofauna is the meadow lizard (Darevskia praticola). It is a small, ground dwelling, predominantly forest living lacertid, for which many ecological and biological data are still missing. Moreover, at least for the western part of the species range (south-eastern Europe) its distribution and environmental associations are insufficiently known, likely due to difficulties in detection even with intensive field sampling. Here, we complemented the available literature records with our personal observations and explored the effects of environmental factors shaping the species ecological niche and distribution in Europe using the ecological niche modelling approach from Maxent software. The new records of the meadow lizard fill a gap in its known distribution. The most suitable habitats are in central and eastern Serbia, south-western Romania, and central, southern and south-western Bulgaria. Mean temperature of the coldest quarter, vegetation and slope had the strongest effect in defining the meadow lizard’s ecological niche. Niche suitability increased with the increase in forest cover and slope, while the temperature showed a bell-shaped response with a rather narrow tolerance range for temperature. Contrary to expectations, precipitation seemed to have no contribution to the species occurrence. Ecological niche model performance increased with a higher resolution of predictor variables despite the lower number of available occurrence records, although the slope variable had a greater predictive power when calculated at a lower resolution. Interestingly, high resolution vegetation variables (30 m) were able to show some level of habitat fragmentation, which likely resulted from deforestation. Overall, our results epitomize the effects of limited sampling on the biogeography inference of elusive species while having significant repercussions on conservation priorities and management of the species.
Corovic, J. & Crnobrnja-Isailovic, J. (2018) -
We studied the thermal biology of the meadow lizard (Darevskia praticola) in the peripheral part of its distribution range (westernmost edge of the distribution area). We assessed whether these lizards actively thermoregulate, estimated the accuracy and effectiveness of thermoregulation, and evaluated the thermal quality of the habitat using the standard thermal parameters: body (Tb), preferred (Tpref) with set-point range (Tset) and operative temperatures (Te). Tset of the meadow lizard under controlled laboratory conditions was between 27.8°C and 31.4°C. In the field Tb and Te averaged 29.0°C and 26.1°C, respectively. A large proportion of Tes fell below the Tset range of the meadow lizard, and lizard Tbs were substantially closer to the species’ Tset range. Obtained values of thermoregulatory indices suggested that the meadow lizard thermoregulated actively, with a rather high accuracy (db = 0.8) and effectiveness (E = 0.8 and de - db = 2.6), and that their habitat at this locality was thermally favourable during the spring. Our results suggest that thermal requirements of the meadow lizard resemble those of alpine lacertids, while their Tbs and Tset are lower than in most lacertid lizards. Further thermoregulation studies could be an important step in predicting the impact of the global climate change on the meadow lizard and the risks of local extinctions of its peripheral populations.
Covaciu-Marcov, S.D. & Cicort-Lucasiu, A.S. & Gaceu, O. & Sas, I. & Ferenti, S. & Bogdan, H. (2009) -
The south-western part of Mehedinţi County comprises many herpetofauna species,
compared to most regions in Romania. This is how we managed to encounter 14 amphibian
species (Salamandra salamandra, Lissotriton vulgaris, Triturus cristatus, Triturus dobrogicus,
Bombina bombina, Bombina variegata, Pelobates fuscus, Bufo bufo, Bufo viridis, Hyla arborea,
Pelophylax ridibundus, Pelophylax lessonae, Rana dalmatina, Rana temporaria) plus Pelophylax kl.
esculentus and 15 reptile species (Emys orbicularis, Testudo hermanni, Ablepharus kitaibelii, Lacerta
agilis, Lacerta viridis, Podarcis muralis, Podarcis taurica, Darevskia praticola, Anguis fragilis,
Zamenis longissimus, Dolichophis caspius, Coronella austriaca, Natrix natrix, Natrix tessellata,
Vipera ammodytes). Salamandra salamandra, Bombina variegata and Rana temporaria were
identified at very low altitudes, lower than any other indication from Romania up to present.
This is explained by the fact that mountain valleys, bearing here a typical aspect, reach all the
way to the Danube, carrying with them the occurring species. In Blahnita Plain, Darevskia
praticola appears in cleared areas, being present in the vegetation girdles bordering the canals
found between agricultural fields.
Covaciu-Marcov, S.D. & Popovici, P.V. & Cicort-Lucaciu, A.S. & Kovács, I.S. & Cupsa, D. & Ferenti, S. (2020) -
Herpetofauna is of interest in protected areas because of the large number of protected species. We studied the herpetofauna of Cozia National Park (CNP) between 2016 and 2018. CNP is situated in the central part of the Southern Romanian Carpathians. We recorded 10 species of amphibian (Salamandra salamandra, Triturus cristatus, Lissotriton vulgaris, Bombina variegata, Hyla arborea, Bufo bufo, Bufotes viridis, Pelophylax ridibundus, Rana dalmatina and R. temporaria), and 11 reptile species (Lacerta agilis, L. viridis, Podarcis muralis, Darevskia praticola, Zootoca vivipara, Anguis colchica, Natrix natrix, N. tessellata, Coronella austriaca, Zamenis longissimus and Vipera ammodytes). Reptiles dominate in number of species, number of individuals and distribution records. CNP is situated at the northern limit of the distribution range of some of these reptiles, notably D. praticola and V. ammodytes. Mountain species associated with a colder, moist climate are very rare or even absent. Zootoca vivipara is restricted to the highest areas of Mount Cozia, above 1 350 m. Although mountain species are well represented in other Carpathian regions, the warmer, drier climate of CNP and its surroundings has limited their distribution in the area, pushing Z. vivipara to higher and higher altitudes. Lacerta agilis is syntopic with all the other lizard species. In some areas, as many as four lizard species cohabitate. The distribution of the herpetofauna in CNP has been negatively influenced by past human activity. The dams on the River Olt have favoured species related to large, stagnant bodies of water, in a region where such habitats were naturally missing. In addition, massive deforestation has decreased the abundance of herpetofauna in many areas of CNP.
Cox, N. & Chanson, J. & Stuart, S. (2006) -
Cyrén, O. (1933) -
Cyrén, O. (1941) -
Czernay (1851) -
Danielyan, F.D. (0000) -
Danielyan, F.D. (1965) -
Даниелян, Ф.Д. (1965) -
Danielyan, F.D. (1967) -
Даниелян, Ф.Д. (1967) -
Danielyan, F.D. (1970) -
Danielyan, F.D. (1971) -
Даниелян, Ф.Д. (1971) -
Danielyan, F.D. (1977) -
Danielyan, F.D. (1981) -
Danielyan, F.D. (1986) -
Danielyan, F.D. (1987) -
Danielyan, F.D. (1989) -
Danielyan, F.D. (2003) -
Danielyan, F.D. (2010) -
Danielyan, F.D. & Arakelyan, M. & Spangenberg, B.E. (2018) -
The study of the sympatric population, located in the Ijevan region of Armenia, where the bisexual species D. raddei coexists with three partenogenetic species - D. rostombekovi, D. dahli and D. armeniaca has shown that in this population a hybridization process is taking place between the D. raddei and D. rostombekowi species. Of the captured 104 D. rostombekowi in the study area, 18 triploid hybrid males have been found. A comparative study of the smears and histological slides prepared from the testicles of 9 hybrid males has detected their possible fertility. The recorded triploid hybrid males could potentially play important role in hybrid speciation.
Danielyan, F.D. & Arakelyan, M. & Stepanyan, I. (2008) -
A unique hybridization zone of rock lizard species of genus Darevskia, with diploid (2n), triploid (3n) and tetraploid (4n) individuals, occurs in the mountain steppe of central Armenia. Our long-term monitoring has showed an increasing birth rate of triploid hybrids in this mixed population. Among these hybrids, the frequency of captured males with fully developed reproductive systems and presumably fertile females also increased significantly. Consequently, intensive microevolution has taken place in this sympatric population. The morphological, ecological, cytological, histological, and parasitological characteristics of the hybrid Darevskia are presented here and compared with parental species.
Along with sterile triploid females, the male hybrids, intersexual individuals,
female hybrids with developing follicles and eggs as well as tetraploid male hybrid
are found in the mixed population of three species of rock lizards’ genus Darevskia
in mountain steppe zone of central Armenia.
Intensive microevolution takes place in this sympatric population. The
evolutionary potential for hybridogenic speciation by Caucasian rock
lizards is discussed.
Հայաստանի կենտրոնական մասի լեռնատափաստանային գոտում,
Darevskia ցեղին պատկանող երկսեռ և կուսածին ժայռային մողեսների երեք
տեսակների խառը պոպուլյացիաներում, ստերիլ տրիպլոիդ էգերի հետ
հայտնաբերվել են տրիպլոիդ հիբրիդային էգեր` զարգացող օօցիտներով և
ձվերեվ, հիբրիդային արու և ինտերսեքսուալ առանձնյակներ, ինչպես նաև
տետրապլոիդ արու առանձնյակ:
Այդ պոպուլյացիայում ընթանում է միկրոէվոլյուցիոն գործընթաց:
Քննարկվում է Կովկասյան ժայռային մողեսների հիբրիդոգեն տեսակառաջացման էվոլյուցիոն պոտենցիալը:
В горно-степной зоне центральной Армении, в смешанной популяции
трех видов двуполых и партеногенетических скальных ящериц рода
Darevskia, наряду со стерильными триплоидными гибридными самками,
обнаружены триплоидные гибридные самки с развивающимися фолликулами
и яйцами, гибридные самцы, интерсексуальные особи, а также тетраплоидный гибридный самец.
В данной смешанной популяции протекает интенсивный микроэволюционный процесс. Обсуждается эволюционный потенциал гибридогенного видообразования у Кавказских скальных ящериц.
Danielyan, F.D. & Grigoryan, A. & Aslanyan, A.V. & Arakelyan, M.S. & Arzumanyan, G. (1998) -
Даниелян, Ф.Д. & Григорян, А. & Асланян А.В. & Даревский И.С. & Арзуманян Г. (1998) -
Danielyan, F.D. & Haikazian, A.K. (1974) -
Даниелян, Ф.Д. & Айказян, А.К. (1974) -
Darevska, I.S. (1978) -
Darevsky, I.S. (1953) -
Даревский И.С. (1953) -
Darevsky, I.S. (1957) -
Даревский И.С. (1957) -
Даревский, И.С. (1957) -
Darevsky, I.S. (1958) -
Даревский И.С. (1958) -
Darevsky, I.S. (1960) -
Darevsky, I.S. (1962) -
Darevsky, I.S. (1963) -
Даревский И.С. (1963) -
Darevsky, I.S. (1965) -
Даревский, И.С. (1965) -
Darevsky, I.S. (1966) -
The polymorphic species Lacerta saxicola Eversmann forms 19 distinct forms in the Caucasus, four of which (armeniaca, dahli, rostombekovi, and unisexualis) are devoid of males and normally reproduce parthenogenetically. These parthenogenetic forms inhabit the mountains on the periphery of the range of the normal bisexual form, L. s. defilippii, thus suggesting a case of geographic parthenogenesis. Since the populations of parthenogenetic lizards are virtually devoid of males, the application of geographic and physiological species criteria to them is a matter of some difficulty. A discussion of this situation with regard to the problem of so-called `agamic species` enables us to regard the parthenogenetic relatives of L. saxicola as distinct species rather than subspecies, and these forms are raised to that rank. The parthenogenetic form of L. s. defilippii is described as a separate species (L. unisexualis). Cytological studies have shown that parthenogenetic species of Rock Lizards are diploid (2n = 38) and that oogenesis proceeds similarly in parthenogenetic and bisexual forms up to the second meiotic division. In regions of sympatry the parthenogenetic and bisexual forms occasionally produce natural hybrids, resulting in sterile triploid (3n = 57) females, the sterility of which is caused by anomalies in the development and structure of the gonads. In the eggs laid by parthenogenetic females, peculiar abnormal embryos, which commonly die in an early stage of incubation, are often developed. Most of them prove to be males, the development of which stops approximately in the middle of incubation. There is reason to suppose that the transition of Rock Lizards to parthenogenesis is associated with natural interspecific hybridization, which in turn favors unisexual reproduction. This tendency could be realized under the influence of selection and apparently took place in forest refuges during Quaternary glaciation of the Caucasus. The transition of Rock Lizards to unisexual reproduction is shown to be accompanied by the appearance of morphologically and biologically distinct species. Natural parthenogenesis may thus be regarded as one of the factors of speciation in the L. saxicola group.
Darevsky, I.S. (1967) -
Four parthenogenetic subspecies of Lacerta . saxicola Eversmann (Transcaucasia) have closely overlapping sympatric ranges. The lizards show no intermediate forms in zones of their mutual habitat. On the border - line of their ranges parthenogenetic lizards can mate with males of neighbouring bisexual forms of Lacerta saxicola, producing hybrids that always are sterile triploid (3n=57) females. Therefore, application of geographical and physiological species criteria to parthenogenetic forms allows to regard them as independent species. However, geographical .and physiofogiical criteria are based on the effect of reproductive isolation mechanisms whose realization is possible upon the occurrence of both sexes in the population. Thus, both criteria should not be unconditionally applied to parthenogenetic and agamic species consisting of females alone. However, the absence of males also plays the role of a peculiar isolation mechanism which maintains the integrity of parthenogenetic species. Due to this, parthenogenetic forms of Lacerta saxicola Eversmann which can be easily distinguished in the morphological respect should be considered as independent agamic species: Lacertra armeniaca Meh., L. dahli Dar., L. rostombekovi Dar. and L. unisexualis Dar.
Darevsky, I.S. (1972) -
Darevsky, I.S. (1978) -
Darevsky, I.S. (1981) -
Darevsky, I.S. (1982) -
Darevsky, I.S. (1984) -
Darevsky, I.S. (1992) -
Darevsky, I.S. (1993) -
Darevsky, I.S. (1997) -
Darevsky, I.S. (2006) -
Darevsky, I.S. & Danielyan, F.D. (1968) -
Darevsky, I.S. & Danielyan, F.D. (1969) -
Darevsky, I.S. & Danielyan, F.D. (1977) -
Darevsky, I.S. & Danielyan, F.D. (1979) -
Darevsky, I.S. & Danielyan, F.D. (2001) -
Darevsky, I.S. & Danielyan, F.D. & Rosanow, J.M. & Sokolowa, T.M. (1991) -
Darevsky, I.S. & Danielyan, F.D. & Sokolova, T.M. & Rozanov, Y.M. (1989) -
Darevsky, I.S. & Eiselt, J. (1967) -
Darevsky, I.S. & Eiselt, J. (1980) -
Präsentation von Lacerta rudis svanetica ssp. nov. vom Südabfall der zentralen Kaukasus-Hauptkette in Nordwest-Grusinien einschließlich einer isolierten Population in Nordwest-Abchasien und von Lacerta parvula adjarica ssp. nov. von der türkischen Schwarzmeerküste samt ihrem Hinterland zwischen Trabzon und Batumi sowie im (Çoruh- (Tschoroch-)tal ab Borçka, in Adjarien und in den angrenzenden Gebieten Grusiniens zwischen Meria (SE Poti) und Borzomi; bei Bakuriani existiert ein isoliertes Vorkommen.
Darevsky, I.S. & Eiselt, J. & Lukina, G.P. (1984) -
Darevsky, I.S. & Gabriel, C.D. (1970) -
Darevsky, I.S. & Grechko, V.V. & Kupriyanova, L.A. (2009) -
Darevsky, I.S. & Kann, N.G. & Ryabinina, N.L. (1988) -
Darevsky, I.S. & Krasilnikov, E.N. (1965) -
Darevsky, I.S. & Kulikova, W.N. (1961) -
Darevsky, I.S. & Kulikova, W.N. (1962) -
Darevsky, I.S. & Kulikova, W.N. (1964) -
Darevsky, I.S. & Kupriyanova, L.A. (1982) -
It is known that karyotypes of parthenogenetic female and male Lacerta armeniaca are similar. This implies that the above males are most likely the results of hormonal inversion. A recently collected male L. armeniaca has also proved to be a diploid individual. This is supported by cytophotometrical measurement of the DNA content of somatic cells in both this male and the bisexual (Control) male of L. portschinskii Kessler, as well as by haploid number of Bivalents at metaphase 1. The DNA content of spermatids in the two males corresponds with the haploid level, and the presence of haploid spermatids argues for normal meiosis. Histological studies supported the above conclusions. The male L. armeniaca appeared to be fertile.
Darevsky, I.S. & Kupriyanova, L.A. & Bakradze, M.A. (1977) -
Darevsky, I.S. & Kupriyanova, L.A. & Bakradze, M.A. (1978) -
Males and intersexes sometimes occur in the progeny of parthenogenetic species of the genus Lacerta, but usually they die during embryogenesis. Their frequency in nature does not exceed 0.1%. Diploid chromosome complements in a male and an intersex of Lacerta armeniaca differ from the female karyotype by the presence of a heteromorphic pair of what are apparently sex chromosomes. These phenomena are discussed in connection with the hypothesis of relictual bisexuality in parthenogenetic species of lizards. All stages of spermatogenesis and mature spermatozoa were discovered in the testes of the male, without a strict order to the rows of maturing cells. The possibility of the existence of fertile males in natural populations of L. armeniaca is supported by the occurrence of natural hybrids between sympatric unisexual species and some other indirect data. The intersex examined, in addition to having hemipenes and testes, had well-developed oviducts. Also, the right gonad was an `ovotestis` consisting of a testis united to several oocytes. The evolutionary importance of fertile diploid males is the possibility of the formation (with their participation) of fertile triploid individuals and even the further transition to tetraploidy: i.e., the theoretical return to bisexuality on a polyploid level.
Darevsky, I.S. & Kupriyanova, L.A. & Danielyan, F.D. (1986) -
Darevsky, I.S. & Kupriyanova, L.A. & Uzzell, T. (1985) -
Darevsky, I.S. & Lukina, G.P. (1977) -
Darevsky, I.S. & Roitberg, E.S. (1999) -
A new subspecies of Lacerta caucasica (s. str.), L. c. vedenica from the northern slope of the Andiiskii Ridge (SE Chechenia) is described. This population appears to be geographically isolated from the main range of L. caucasica, but contacts the northern limits of the range of a closely related species, L. daghestanica. It differs from the other populations of L. caucasica as well as from the related species, L. daghestanica and L. alpina by a very low number of superciliary granules and superciliary scales as well as by the lack of greenish paints in the dorsal coloration.
Darevsky, I.S. & Szczerbak, N.N. (1967) -
Darevsky, I.S. & Tuniyev, B.S. (1997) -
The re-examination of Lacerta clarkorum Darevsky et Vedmederja, 1977 type series as well as analysis of the recently collected new material may suggest that within the frames of Northeastern Turkey and neighboring region of Adzharistan within the Georgia, live two close allopatric species: Lacerta clarkorum proper and a new species Lacerta dryada sp. nov., whose description is made in this article. Evidence is provided on the distribution and comparative ecology of both species, their possible phylogenetic relation being discussed.
Darevsky, I.S. & Uzzell, T. & Kupriyanova, L.A: & Danielyan, F.D. (1973) -
Darevsky, I.S. & Vedmederja, W.I. (1977) -
Davies, R.S. (1984) -
Davoyan, A.G. & Aslanyan, A.V. & Danielyan, F.D. & Darevsky, I.S. & Martirosyan I.A. (2007) -
Locus-specific PCR was used to study the genetic polymorphism in three populations of parthenogenetic lizard species Darevskia dahli. The analysis was carried at the two (GATA) n -containing loci (Du215 and Du281) using the sample of 26 individuals. A total of eight Du215 and three Du281 allelic variants were detected. It was demonstrated that all the lizards examined were heterozygous at these loci. In 12 animals, unusual Du215 allelic variant was revealed, the origin of which was thought to be associated with different types of genomic rearrangements, or segmental duplication. The populations studied were substantially different relative to the levels of allelic polymorphism, which could be explained by different habitation conditions, leading to accumulation of mutations in noncoding genome regions.
Давоян А.Г., Асланян А.В., Даниелян Ф.Д., Даревский И.С., Мартиросян И.А. (2007) -
Локус-специфическая ПЦР использована для определения генетического полиморфизма в трех популяциях партеногенетического вида ящериц Darevskia dahli. Анализ проведен на выборке 26 особей по двум class=`txr-`>(class=`txr-`>GATA)class=`txi-x-x-70`>n-содержащим локусам (Du215 и Du281). Обнаружено восемь аллельных вариантов по локусу Du215 и три по локусу Du281. Показано, что все исследованные особи гетерозиготны по этим локусам. У 12 особей по локусу Du215 обнаружен необычный аллельный вариант, происхождение которого может быть связано с различными типами геномной реорганизации или сегментной дупликации. Выявлены значительные различия по уровню аллельного полиморфизма между исследованными популяциями, что может объясняться различными экологическими условиями обитания популяций, приводящими к накоплению мутаций в некодирующей области генома.
Dehgani, A. & Karamiani, R. & Rastegar-Pouyani, N. & Banan-Khojasteh, S.M. (2016) -
The Arasbaran Protected Area is a forested and mountainous area, which is located in northern part of East Azerbaijan province and bordered to the north by Armenia and Azerbaijan and southern Aras River of the Iranian Plateau confined to 38°40` to 39°08`N and 46°39` to 47°02`E. A study on the status of Sauria in this area was carried out from June to September 2011 and 2013. A total of 141 specimens were collected and identified, which represented four families, 11genera, and 12 species and subspecies, including collected Agamidae: Paralaudakia caucasia, Phrynocephalus persicus, Trapelus lessonae, Anguidae: Pseudopus apodus, Lacertidae: Darevskia raddeii, Iranolacerta brandtii, Eremias strauchi, Lacerta media, Lacerta strigata, Ophisops elegans and Scincidae: Eumeces schneideri Ablepharus bivittatus. The Lacertidae with six species, showed the highest diversity among the families represented here. In this study a new locality was found for Lacerta strigata 30 km north of kalibar, in the Arasbaran.
سید مهدی بانان خجسته امیر دهقانی، رسول کرمیانی،، نصراله رستگار-پویانی، (1396) -
ارسباران یک منطقه کوهستانی با کوههای دارای شیب تند و جنگلی است که در شمال استان آذربایجانشرقی و در همسایگی کشور های جمهوری آذربایجان، ارمنستان و جنوب رود ارس، در محدوده `۴۴ ۳۳° تا `۳ °۳3 شمالی و`۳3 °۴1 تا `۲ °۴۴ شرقی واقع شده است. در طی مطالعه و نمونهبرداری فون خزندگان منطقه ارسباران که از تیر ماه سال 3۴ تا تیر سال 3۲ انجام شد. مجموعا۱۴۱ نمونه از ۴ خانواده، ۱۱ جنس و ۱۲ گونه و زیر گونه سوسمار جمعآوری و
شناسایی شد. که این نمونهها شامل:
Paralaudakia caucasia, Phrynocephalus persicus, Trapelus lessonae, Anguidae: Pseudopus apodus, Lacertidae: Darevskia raddeii, Iranolacerta brandtii, Eremias strauchi, Lacerta media, Lacerta strigata, Ophisops elegans and Scincidae: Eumeces schneideri, Ablepharus bivittatus
که در بین اینها خانواده سرتیده با 1 گونه با ترین تنوع را
نشان میدهد و گونه Darevskia raddeii بیشترین پراکنش را دارد. در این مطالعه گونه Lacerta strigata برای اولین بار از
Dehghani, A. & Hosseinian Yousefkhani, S.S. & Rastegar-Pouyani, N. & Banan-Khojasteh, S.M. & Mohammadpour, A. (2014) -
We examined sexual size dimorphism of the rock-dwelling lizard Darevskia raddei (Boettger, 1892) with the help of 30 specimens that were provided from various sources. Eleven metric and seven meristic features were examined. Seven characters (gulars, length of basal tail, femoral pores, length of head, width of head, length of fore limb and length of hind limb) were identified as dimorphic between the two sexes. Some of these characters have important roles in copulation for males, especially the hind limb and the tail base. The number of femoral pores is important in the release of signal components because females release these components to attract males during the mating season. The length of the hind limb as locomotor performance plays an important role during mating, so that the male can grasp the female and adopt the correct position during copulation.
Dehghani, A. & Rastegar-Pouyani, N. & Mahdi Banan-Khojasteh, S. & Mohammadpour, A. (2014) -
In this article we present the effects of waste water of the Sungun copper mine on Darevskia raddei
habitat in the Arasbaran protected region, a UNESCO registered biosphere reserve since 1976, and
appeal to the Department of Environment of East Azarbaijan province to more consideration on control and
precautions regarding the water waste of the Sungun mine.
Deichsel, G. (2004) -
Deichsel, G. & Bischoff, W. (2002) -
Delmas, C. (2019) -
Dieckmann, M. (2011) -
Diesener, G. & Reichholf, J. (1986) -
Dobrovolskaja, H. (1964) -
Doronin I.V. (2017) -
The paper provides data on the current location of the type specimens of Darevskia (rudis) complex as of July 2017:
Podarcis depressa Camerano, 1878 (syntypes – MZUT R2737.1-3, NHM 19126.96.36.199-32), Lacerta depressa var.
rudis Bedriaga, 1886 (lectotype – MZUT 2737.1, paralectotypes – MZUT 2737.2-3, NHM 19188.8.131.52), Lacerta
depressa var. modesta Bedriaga, 1886 (syntype – NHM 19184.108.40.206), Lacerta saxicola Subsp. bithynica Méhely,
1909 (lectotype [here designated] – ZISP 17128, paralectotype – NMW 11685), Lacerta saxicola tristis Lantz et
Cyrén, 1936 (lectotype – GNM 2481, paralectotypes – GNM 2482-2485, 2487 and GNM one unnumbered specimen),
Lacerta saxicola obscura Lantz et Cyrén, 1936 (lectotype [here designated] – ZISP ZISP 17171.1, paralectotypes
– NHM 1966.769.773, ZISP 14435, 14436, 14437, 14438, 14439, 14440, 14442, 15402, 16290, 17057.1-
3, 17058, 17059.1-3, 17171.2-26, 17172.1-15), Lacerta rudis macromaculata Darevsky, 1967 (holotype – ZISP
17940.6, paratypes – ZISP 17440.1-18, the location of three paratypes is unknown), Lacerta rudis bischoffi Böhme
et Budak, 1977 (holotype – ZDEU 183/977, paratypes – ZDEU 222/976:1, 3-6, 8-9, ZDEU 183/977.2-10, ZFMK
22062.63), Lacerta rudis svanetica Darevsky et Eiselt, 1980 (holotype – ZISP 17875.1, paratypes – ZISP 17875.2-8,
17814.1-18, NMW 24138.1-6), Lacerta rudis chechenica Eiselt et Darevsky, 1991 (holotype – NMW 33504.1, paratypes
– ZISP 17878.1-13, 17882.2-4, ZISP 18374.1-5, 18781.1-6, 19545.1-13, ZFMK 16448.52, 29262, 38037.45,
40431-33, NMW 32619.1,2, 33504.2,4,5), Darevskia rudis mirabilis Arribas, Ilgaz, Kumlutaş, Durmuş, Avci et
Üzüm, 2013 (holotype – ZDEU 145/2002.9, paratypes – ZDEU 145/2002.1-8, 10-18, O. Arribas private collection
(1-6)), Darevskia rudis bolkardaghica Arribas, Ilgaz, Kumlutaş, Durmuş, Avci et Üzüm, 2013 (holotype – ZDEU
144/2009.6, paratypes – ZDEU 144/2009.1-5, 7-8, O. Arribas private collection (1-4)). Strictly following the
ICZN the name Lacerta depressa var. rudis Bedriaga, 1886 should not be considered as the replacement name (nomen
novum) for Podarcis depressa Camerano, 1878, but the only available name. The history of description of all
known forms of the complex is given. Localities, collectors and dates of collection of the type specimens are clarified.
Доронин И.В. (2017) -
В работе приводится информация о современном месте хранения типовых экземпляров ящериц комплекса Darevskia (rudis) по состоянию на июль 2017 г.: Podarcis depressa Camerano, 1878 (синтипы – MZUT R2737.1- 3, NHM 19220.127.116.11-32), Lacerta depressa var. rudis Bedriaga, 1886 (лектотип – MZUT 2737.1, паралектотипы – MZUT 2737.2-3, NHM 1918.104.22.168), Lacerta depressa var. modesta Bedriaga, 1886 (синтип – NHM 1922.214.171.124), Lacerta saxicola Subsp. bithynica Méhely, 1909 (лектотип [обозначается здесь] – ZISP 17128, паралектотип – NMW 11685), Lacerta saxicola tristis Lantz et Cyrén, 1936 (лектотип – GNM 2481, паралектотипы – GNM 2482-2485, 2487 и GNM один экземпляр без номера), Lacerta saxicola obscura Lantz et Cyrén, 1936 (лекто- тип [обозначается здесь] – ZISP 17171.1, паралектотипы – NHM 1966.769.773, ZISP 14435-14440, 14442, 15402, 16290, 17057.1-3, 17058, 17059.1-3, 17171.2-26, 17172.1-15), Lacerta rudis macromaculata Darevsky, 1967 (голотип – ZISP 17940.6, паратипы – ZISP 17440.1-18, местонахождение трех паратипов неизвестно), Lacerta rudis bischoffi Böhme et Budak, 1977 (голотип – ZDEU 183/977, паратипы – ZDEU 222/976:1, 3-6, 8-9, ZDEU 183/977.2-10, ZFMK 22062.63), Lacerta rudis svanetica Darevsky et Eiselt, 1980 (голотип – ZISP 17875.1, паратипы – ZISP 17875.2-8, 17814.1-18, NMW 24138.1-6), Lacerta rudis chechenica Eiselt et Darevsky, 1991 (голотип – NMW 33504.1, паратипы – ZISP 17878.1-13, 17882.2-4, ZISP 18374.1-5, 18781.1-6, 19545.1- 13, ZFMK 16448.52, 29262, 38037.45, 40431-33, NMW 32619.1,2, 33504.2,4,5), Darevskia rudis mirabilis Arribas, Ilgaz, Kumlutaş, Durmuş, Avci et Üzüm, 2013 (голотип – ZDEU 145/2002.9, паратипы – ZDEU 145/2002.1-8, 10-18, личная коллекция О. Аррибаса (1-6)), Darevskia rudis bolkardaghica Arribas, Ilgaz, Kumlutaş, Durmuş, Avci et Üzüm, 2013 (голотип – ZDEU 144/2009.6, паратипы – ZDEU 144/2009.1-5, 7-8, личная коллекция О. Аррибаса (1-4)). Показано что в соответствии с МКЗН, таксон Lacerta depressa var. rudis Bedriaga, 1886 сле- дует считать не замещающим названием (nomen novum) для Podarcis depressa Camerano, 1878, а единствен- ным пригодным названием (available name). Приводится история описания форм комплекса. Уточняются локалитеты, сборщики и даты сбора типовых экземпляров.
Doronin, I. & Garmasheva, N.N. (2015) -
The paper describes a first case of successful hunting of a Parasteatoda tepidariorum (C. L. Koch, 1841) spider for a juvenile individual of Pontic Lizard, Darevskia pontica (Lantz et Cyrén, 1919), recorded in the territory of Slavyansk-on-Kuban town (Krasnodar Region, Russia).
Доронин И.В., Гармашева Н.Н. (2015) -
Впервые описан случай успешной охоты паука Parasteatoda tepidariorum (C. L. Koch, 1841) на ювенильную особь понтийской ящерицы, Darevskia pontica (Lantz et Cyrén, 1919), зафиксированный на территории г. Славянск-на-Кубани (Краснодарский край, Россия).
Doronin, I.V. (2011) -
The description of a new subspecies of rock lizards Darevskia saxicola complex, inhabited the territory of Pitsunda-Myussera Reserve and in the Gagra (Republic of Abkhazia) is given. Myussera lizard, Darevskia brauneri myusserica ssp. nov., differs from other taxa of the complex by the following combination of morphological characters: (1) large or very large masseteris scute; (2) interrupted series of ciliated grains between supraciliar and supraocular scutes; (3) presence of additional scutes lying on either side of occipital and interparietal scutes or splitting of the latter; (4) net color pattern on the back (in females fuzzy); (5) dominance of gray and light gray color in the coloration of the back of females; (6) white throat and belly. In addition, these subspecies differs by some peculiarities of biology: biotopical preference to coastal pebble conglomerates and a relatively low population size. Probably, the taxon was formed during the Pleistocene. Formation of the coastal plains of Pitsunda Peninsula by alluvial and marine accumulation in the Late Neopleistocene – Holocene divided the area of Myussera lizard into the Gagra and Myussera plots. This area is situated within the Black Sea refuge of Eastern Mediterranean species of herpetofauna.
И.В. Доронин (2011) -
В статье приводится описание нового подвида скальной ящерицы комплекса Darevskia saxicola, обитающего на территории Пицундо-Мюссерского заповедника и в районе г. Гагра Республики Абхазия. Мюссерская ящерица, Darevskia brauneri myusserica ssp. nov., отличается от других таксонов комплекса следующей комбинацией морфологических признаков: (1) крупный или очень крупный центральновисочный щиток; (2) прерывистый ряд ресничных зернышек между верхнересничными и надглазничными щитками; (3) наличие дополнительных щитков, лежащих по обе стороны от затылочного и межтеменного щитков, либо дробление последнего; (4) сетчатый рисунок на спине (у самок нечеткий); (5) доминирование у самок серого и светлосерого цвета в окраске дорсальной поверхности тела; (6) белое горло и брюхо. Кроме того, новый подвид отличается некоторыми особенностями биологии: биотопической приуроченностью к прибрежным выходам конгломерата и относительно низкой численностью популяции. Предположительно, формирование таксона протекало в плейстоцене. Образование приморской равнины полуострова Пицунда за счет аллювиальной и морской аккумуляции в позднем неоплейстоцене – голоцене разделило ареал мюссерской ящерицы на гагрский и мюссерский участки. Эта территория расположена в пределах Черноморского рефугиума восточносредиземноморских видов герпетофауны.
Doronin, I.V. (2012) -
The paper provide information on the current status of the type specimens of Darevskia (saxicola) complex as they were in December 2011. I order to stabilize nomenclature of this complex, the neotype of Lacerta saxicola Eversmann, 1834, lectotypes of Lacerta saxicola brauneri Méhely, 1909 and Lacerta saxicola lindholmi Szczerbak, 1962 are designated. History of description of these taxa is discussed.
И.В. Доронин (2012) -
В работе приводится информация о современном месте хранения типовых экземпляров ящериц комплекса Darevskia (saxicola) по состоянию на декабрь 2011 г. С целью стабилизации номенклатуры выделяются и описываются неотип Lacerta saxicola Eversmann, 1834, лектотипы Lacerta saxicola brauneri Mehely, 1909 и Lacerta saxicola lindholmi Szczerbak, 1962. Приводится история описания форм комплекса.
The greatest number of reptiles having melanism (the number of specimens and species) have been recorded in the Caucasus in populations of five Darevskia species: Darevskia alpina, D. brauneri, D. lindholmi, D. pontica and D. rudis. The occurrence of melanistic specimens in reptiles can be explained by the influence of geochemical anomalies, thermoregulatory adaptation, isolation, and occurence in the periphery of their range. Melanism of D. lindholmi appears to depend on geochemical factors. The melanistic specimens of D. rudis, D. pontica and D. brauneri were found in a city on the northeastern boundary of their range. D. alpina was recorded in the mountains on the eastern periphery of the range.
The article is the result of the analysis of the distribution of the Darevskia praticola praticola (Eversmann, D. p. hyrcanica Tuniyev, Doronin, Kidov et Tuniyev, 2011, and D. pontica (Lantz et Cyrén, 1919) in the Caucasus. GIS-modeling was used by the program Maxent 3.3.3e. An analysis of the obtained distribution maps led to the conclusion about the location of centres (zones optimum) of distribution ranges. The zone of optimum for studied forms do not overlap each other. It can be considered as indication of the difference in the location of the centers of their formation. Data on the modern change of borders of ranges of lizard Darevskia (praticola) complex are presented.
To study the habitats and potential spread of the rock lizards from the Darevskia (saxicola) complex, the
Maxent 3.3.3e program was used. Our analysis has found that currently in the Ciscaucasia (on the
Stavropol Upland) there are no favorable climatic conditions for habitation of D. saxicola but it can be
found on the territory of North Ossetia-Alania (in the Digorskoe Gorge), there are no clear boundaries
between the habitats of D. b. brauneri and D. b. darevskii. This is further evidence in favor of their
conspecificity; the habitat of D. [brauneri] szczerbaki has been stable in recent decades and no significant
movement of its borders has taken place, in Abkhazia there is very limited territory suitable for habitation
of D. b. myusserica. Bioclimatic factors affecting the habitat of the lizards from the complex were
Доронин И.В. (2012) -
Для изучения ареалов и потенциального распространения скальных ящериц комплекса Darevskia (saxicola) была использована программа Maxent 3.3.3e. В результате установлено, что в настоящее время в Предкавказье (на Ставропольской возвышенности) отсутствуют благоприятные климатические условия для обитания D. saxicola, но она может быть обнаружена на территории Северной Осетии-Алании (в Дигорском ущелье); нет четких границ между ареалами D. b. brauneri и D. b. darevskii, что является еще одним фактом, свидетельствующим в пользу их отнесения к одному подвиду; ареал D. [brauneri] szczerbaki на протяжении последних десятилетий был стабилен и существенных подвижек его границ не происходило; имеется крайне ограниченная территория в Абхазии, подходящая для обитания D. b. myusserica. Был проведен анализ биоклиматических факторов, влияющих на ареал ящериц комплекса.
Doronin, I.V. (2013) -
New records of Darevskia alpina from the vicinities of Zemo Bachiani Village in South Ossetia are presented and discussed: D. brauneri from the place of the confluence of the Luhunistskale River into the Rion River in Georgia and vicinities of Ertso Lake in South Ossetia, D. pontica from the vicinities of Slavyansk-na-Kubani, Timashevsk, Novotitarovskaya Village, Poltavskaya Village and Burakovsky Village in Krasnodar Region, D. portschinskii nigrita from the vicinities of Kizilkilisa Village in Georgia, D. praticola from the vicinities of Tskhinval, Acrisheu Village, Malaya Gufta Village, Hetagurova Village and the southern slope of the Zonkar Ridge in South Ossetia. Information on an attempt to introduce D. pontica into the territory of Rostov-on-Don is provided.
И.В. Доронин (2013) -
Приводятся и обсуждаются новые находки Darevskia alpina из окрестностей с. Земо Бачиани Южной Осетии, D. brauneri из района впадения р. Лухунисцкале в р. Рион Грузии и окрестностей оз. Эрцо Южной Осетии, D. pontica из окрестностей г. Славянск-на-Кубани, г. Тимашевск, ст-цы Новотитаровская, ст-цы Полтавская и хут. Бураковский Краснодарского края, D. portschinskii nigrita из окрестностей c. Кизилкилиса Грузии, D. praticola из окрестностей г. Цхинвал, с. Ацрисхеу, с. Малая Гуфта, с. Хетагурова и южного склона Зонкарского хребта Южной Осетии. Приводятся данные о попытке интродукции D. pontica на территорию г. Ростов-на-Дону.
Doronin, I.V. (2013) -
Доронин, И.В. (2013) -
Доронин И.В. (2013) -
Doronin, I.V. (2014) -
The paper provides information on the current status of the type specimens of Darevskia (caucasica) complex as they were in August 2014. In order to stabilize nomenclature of this complex, the lectotype of Lacerta saxicola daghestanica Darevsky, 1967 is designated. The description of external morphology of paralektotype of a L. s. daghestanica, lektotype and paralektotype of Lacerta caucasica Méhely, 1909 is given. History of description of these taxa is discussed.
И.В. Доронин (2014) -
В работе приводится информация о современном месте хранения типовых экземпляров ящериц комплекса Darevskia (caucasica) по состоянию на август 2014 г. Для стабилизации номенклатуры выделяется лектотип Lacerta saxicola daghestanica Darevsky, 1967. Дается описание внешней морфологии паралектотипа L. s. daghestanica, лектотипа и паралектотипа Lacerta caucasica Mehely, 1909. Приводится история описания форм комплекса.
Doronin, I.V. (2015) -
The paper presents analysis of the distribution of Darevskia praticola praticola, D. praticola loriensis, D. p. hyrcanica, and D. pontica in the Caucasus. GIS modeling (Maxent 3.3.3k) was used. Analysis of the obtained GIS maps of distribution has allowed us to locate territories with maximum occurrence probabilities of these taxa. Analysis of the bioclimatic factors that affect the lizard habitats from the complex and their zoogeographical position is given; data on the modern changes in the boundaries of the taxa from the Darevskia (praticola) complex are presented.
Доронин И.В. (2015) -
В статье приводятся результаты анализа распространения Darevskia praticola praticola, D. praticola loriensis, D. p. hyrcanica и D. pontica на Кавказе. Была использована ГИС-программа Maxent 3.3.3k. Анализ полученных ГИС-карт распространения позволил определить расположение территорий с максимальной вероятностью нахождения таксонов. Дается анализ биоклиматических факторов, влияющих на ареалы ящериц комплекса, и их зоогеографичеcкого положения; приводятся данные о современном изменении границ распространения представителей комплекса Darevskia (praticola).
1. Sexual dimorphism and geographical variability in signs of external morphology are revealed in the species of the studied complexes of rock lizards. The most important for their diagnosis are the features of folidosis of the head. Interspecific and intraspecific differences in morphometric features, color and pattern of the body were detected.
2. In the phylogenetic analysis, the Darevskia (caucasica) samples were divided into the group of the northern slope of the Central Caucasus (D. caucasica), the East Caucasus (D. daghestanica), the Western Caucasus (D. alpina), the northern slope of the Central and Western Caucasus (D. alpina ). Reliable identification and significant genetic differences of D. caucasica and D. daghestanica samples favor their attribution to different species. Within D. alpina two groups with high genetic differences (distances) were identified. Inside Darevskia (saxicola) two lines are described, one of which is connected with the Crimea and the Western Caucasus (D. lindholmi-D. Brauneri), and the second - with the North and West Caucasus (D. saxicola-D. Szczerbaki).
3. As a result of the nomenclatural revision, the lectotype Lacerta praticola pontica, L. saxicola brauneri, L. saxicola daghestanica, L. saxicola lindholmi, and the neotype L. saxicola; Specific locations, years and authors of the description of a number of taxa have been refined.
4. It has been established that the studied supernumerary complexes include the following valid taxa: Darevskia (praticola) - D. pontica pontica, D. pontica hungarica, D. praticola praticola, D. praticola hyrcanica, D. praticola loriensis; Darevskia (caucasica) - D. alpina, D. caucasica caucasica, D. caucasica vedenica, D. daghestanica; Darevskia (saxicola) - D. brauneri brauneri, D. brauneri myusserica, D. saxicola, D. szczerbaki, D. lindholmi. Lacerta saxicola darevskii is regarded as a minor synonym for Lacerta saxicola brauneri.
5. For the first time accurate inventories of findings (1,443 locations) and GIS maps of species arealways were compiled, reliably characterizing the features of their distribution; The structure of areals, the current trends of their changes and the factors influencing the distribution of rock lizards are revealed. It is recommended to carry out conservation measures of international and national level in order to preserve the populations of D. alpina, D. brauneri myusserica and D. szczerbaki, especially in typical territories.
6. The first and only reliable paleontological find of Darevskia sp. - Darevskia sf. brauneri from the Pleistocene site, the caves of Mezmaiskaya. A hypothesis is proposed for the formation of species of complexes, according to which the isolation of their common ancestor from a hypothetical form
it is logical to associate with the Miocene and Pliocene boundaries, or with the Cimmerian century of the Pliocene, 153
and the radiation of modern species - with the Akchagylian age, corresponding to the Piacenza and Gelazian tiers. The parallel formation of several Darevskia (praticola) and Darevskia (caucasica) lines around the Greater Caucasus is assumed in the early Pliocene. At the end of the Pontian age, the primary area of the ancestor of the modern forms of Darevskia (saxicola) and the subsequent formation in the Mediterranean landscapes of the Black Sea coast, along the surviving cliff, D. szczerbaki, and the cuestas of the North Caucasus - D. saxicola, could have occurred at the periphery of the Caucasian peninsula.
1. У видов изученных комплексов скальных ящериц выявлены половой диморфизм и географическая изменчивость в признаках внешней морфологии. Наиболее важными для их диагностики являются особенности фолидоза головы. Обнаружены меж- и внутривидовые различия по морфометрическим признакам, окраске и рисунку тела.
2. При филогенетическом анализе исследуемые образцы Darevskia (caucasica) распределились на группу северного склона Центрального Кавказа (D. caucasica), Восточного Кавказа (D. daghestanica), Западного Кавказа (D. alpina), северного склона Центрального и Западного Кавказа (D. alpina). Надежная идентификация и значимые генетические различия образцов D. caucasica и D. daghestanica говорят в пользу их отнесения к разным видам. Внутри D. alpina выявлены две группы с высокими генетическими различиями (дистанциями). Внутри Darevskia (saxicola) описаны две линии, одна из которых связана с Крымом и Западным Кавказом (D. lindholmi–D. brauneri), а вторая – с Северным и Западным Кавказом (D. saxicola–D. szczerbaki).
3. В результате проведения номенклатурной ревизии выделены лектотипы Lacerta praticola pontica, L. saxicola brauneri, L. saxicola daghestanica, L. saxicola lindholmi и неотип L. saxicola; уточнены типовые местонахождения, годы и авторы описания ряда таксонов.
4. Установлено, что изученные надвидовые комплексы включают следующие валидные таксоны: Darevskia (praticola) – D. pontica pontica, D. pontica hungarica, D. praticola praticola, D. praticola hyrcanica, D. praticola loriensis; Darevskia (caucasica) – D. alpina, D. caucasica caucasica, D. caucasica vedenica, D. daghestanica; Darevskia (saxicola) – D. brauneri brauneri, D. brauneri myusserica, D. saxicola, D. szczerbaki, D. lindholmi. Lacerta saxicola darevskii рассматривается как младший синоним Lacerta saxicola brauneri.
5. Впервые составлены точные кадастры находок (1443 локалитетов) и ГИС-карты ареалов видов, надежно характеризующие особенности их распространения; выявлена структура ареалов, современные тенденции их изменения и факторы, влияющие на распространение скальных ящериц. Рекомендовано проводить охранные мероприятия международного и национального уровня с целью сохранения популяций D. alpina, D. brauneri myusserica и D. szczerbaki, в особенности на типовых территориях.
6. Обнаружена первая и единственная на сегодняшний день достоверная палеонтологическая находка Darevskia sp. – Darevskia sf. brauneri из плейстоценового местонахождения, пещеры Мезмайской. Предложена гипотеза формирования видов комплексов, согласно которой обособление их общего предка от гипотетической формы
логично связывать с рубежом миоцена и плиоцена, либо с киммерийским веком плиоцена, 153
а радиацию современных видов – с акчагыльским веком, соответствующим пьяченцскому и гелазскому ярусам. Предполагается параллельное становление в раннем плиоцене нескольких линий Darevskia (praticola) и Darevskia (caucasica) вокруг Большого Кавказа. В конце понтического века мог произойти разрыв первичного ареала по периферии Кавказского полуострова предка современных форм Darevskia (saxicola) и последующее становление в средиземноморских ландшафтах Черноморского побережья, вдоль сохранившегося клифа, D. szczerbaki, а по куэстам Северного Кавказа – D. saxicola.
Doronin, I.V. (2016) -
The paper provides data on the current location of the type specimens of Darevskia (praticola) complex as of March
2016: Lacerta praticola Eversmann, 1834 (storage holotype place (on monotype) is unknown), Lacerta vivipara
stenolepis Nikolsky, 1911 (holotype for monotypes – ZISP 7203, location unknown), Lacerta praticola pontica Lantz
et Cyrén, 1918 (lectotype – ZISP 22853, paralectotypes – ZISP 5279, 5280, 22847, 22852.1-2, 22854), Lacerta
praticola hungarica Sobolewsky, 1930 (location lectotype and paralectotypes ZISP 9814 unknown, paralectotypes –
ZMMU R 2538), Lacerta plicata Bartenef et Reznikova, 1931 (holotype (on monotypes) – ZISP 15204), Darevskia
praticola hyrcanica Tuniyev, Doronin, Kidov et Tuniyev, 2011 (holotype – SNP 1473.5, paratypes – SNP 1473.0-19,
ZISP 12301, 12630, 12632-12635), Darevskia praticola loriensis Tuniyev, Doronin, Tuniyev, Aghasyan, Kidov et
Aghasyan, 2013 (holotype – SNP 1568.9, paratypes – SNP 1569.1-19, ZISP 17075). The history of description of
all known forms of the complex is given. L. praticola pontica has been described in 1918, not in 1919.
Доронин, И.В. (2016) -
В работе приводится информация о современном месте хранения типовых экземпляров ящериц комплекса Darevskia (praticola) по состоянию на март 2016 г.: Lacerta praticola Eversmann, 1834 (место хранения голотипа (по монотипии) неизвестно), Lacerta vivipara stenolepis Nikolsky, 1911 (голотип по монотипии – ZISP 7203, место нахождения неизвестно), Lacerta praticola pontica Lantz et Cyrén, 1918 (лектотип – ZISP 22853, паралектотипы – ZISP 5279, 5280, 22847, 22852.1-2, 22854), Lacerta praticola hungarica Sobolewsky, 1930 (место нахождения лектотипа и паралектотипов ZISP 9814 неизвестно, паралектотип – ZMMU R 2538), Lacerta plicata Bartenef et Reznikova, 1931 (голотипа (по монотипии) – ZISP 15204), Darevskia pra- ticola hyrcanica Tuniyev, Doronin, Kidov et Tuniyev, 2011 (голотип – SNP 1473.5, паратипы – SNP 1473.0-19, ZISP 12301, 12630, 12632-12635), Darevskia praticola loriensis Tuniyev, Doronin, Tuniyev, Aghasyan, Kidov et Aghasyan, 2013 (голотип – SNP 1568.9, паратипы – SNP 1569.1-19, ZISP 17075). Приводится история описания форм комплекса. Показано, что L. praticola pontica описана не в 1919 г., а в 1918 г.
Doronin, I.V. (2018) -
The purpose of this work, under Articles 126.96.36.199, 23.9.2, and 24.2 of the International Code of Zoological Nomenclature,
is to conserve the priority of the specific name Lacerta mixta Méhely, 1909. This name is threatened
by the never-used-as-valid name Lacerta composita Méhely, 1909. It is proposed that Lacerta mixta, Méhely, 1909
is given precedence as a nomen protectum over Lacerta composita Méhely, 1909, which will be considered as a
Doronin, I.V. & Doronina, M.A. & Bekoshvili, D. (2019) -
New data on the distribution of four species of lizards (Darevskia brauneri, D. daghestanica, Lacerta agilis ioriensis, L. strigata) in the Caucasus are presented.
Doronin, I.V. & Doroninа, M.A. (2020) -
The finding of a melanist meadow lizard (Darevskia praticola praticola) in the Stavropol Territory is discussed. Information on the clutch and offspring obtained from the female of this aberration is given.
Доронин, И.В. & Доронина, М.А. (2020) -
Обсуждается находка меланиста луговой ящерицы (Darevskia praticola praticola) на территории Ставропольского края. Приводятся сведения о полученных от самки этой аберрации кладке и потомстве.
Doronin, I.V. & Dzhelali, P.A. & Lotiev, K.Yu. & Mazanaeva, L.F. & Mustafaeva, G.A. & Bunyatova S.N. (2021) -
The range of superspecific complex of rock lizards Darevskia (caucasica) (D. alpina, D. c. caucasica, D. c. vedeni- ca, D. daghestanica) includes the Main Caucasian ridge and its spurs on the southern and northern macroslopes within heights from 500 m to 3200 m above sea level. A fragment of the mitochondrial cytochrome b gene (1096 bp) was analyzed in 48 specimens of these lizards. Specimens were collected in 23 localities. We have found 39 unique haplotypes. The minimum genetic diversity was found in D. c. vedenica, maximum in D. c. caucasica. All taxa of the complex, except for D. c. vedenica, demonstrate genetic heterogeneity. Four clades can be distinguished on the dendrogram of phylogenetic relationships. They correspond to D. alpina, D. c. caucasica, D. c. vedenica, D. dagh- estanica. D. c. caucasica forms two distinct haplogroups. On the phylogenetic tree, the sample of D. c. vedenica formed an independent clade close to D. c. caucasica. The Darevskia (caucasica) complex is polyphyletic accord- ing to the analysis of the mitochondrial marker. Nucleotide sequences of D. alpina formed a single clade with Darevskia (saxicola) complex representatives and combined with the sequences of D. brauneri into subclade. On the example of the population of D. daghestanica, found in a forest biotope without rocky outcrops and genetically not differing from individuals from the adjacent territory, the ability of rocky lizards to master biotopes unusual for them in a short period of time was demonstrated. This example demonstrates the ability of rock lizards to adapt to unusual biotopes in a short period of time. Genetic distance between D. c. vedenica and D. c. caucasica is com- parable to the distance between D. c. caucasica and D. daghestanica. We assume the existence of an independent form of D. caucasica ssp. inhabiting South Ossetia and adjacent territories.
И.В. Доронин, П.А. Джелали, К.Ю. Лотиев, Л.Ф. Мазанаева, Г.А. Мустафаева5 и С.Н. Буньятова (2021) -
Ареал представителей комплекса Darevskia (caucasica) (D. alpina, D. с. caucasica, D. c. vedenica, D. daghes- tanica) охватывает Главный Кавказский хребет и его отроги на южном и северном макросклонах в пре- делах высот от 500 м до 3200 м над уровнем моря. В работе проанализирован участок митохондриаль- ного гена цитохрома b (1096 п.н.) от 48 особей этих ящериц, собранных в 23 локалитетах. Обнаружены 39 уникальных гаплотипов. Минимальные значения показателей генетического разнообразия выяв- лены у D. c. vedenica, максимальные – у D. c. caucasica. Все таксоны комплекса, за исключением ве- денской ящерицы, демонстрируют генетическую гетерогенность. На дендрограмме филогенетических отношений можно выделить четыре клады, соответствующие D. alpina, D. с. caucasica, D. c. vedenica, D. daghestanica. D. с. caucasica формирует две обособленные гаплогруппы. Выборка веденской ящерицы сформировала самостоятельную кладу на дереве, близкую по положению к номинативному подвиду кавказской ящерицы. По митохондриальному маркеру комплекс Darevskia (caucasica) полифилетичен: нуклеотидные последовательности D. alpina сформировали единую кладу с Darevskia (saxicola), обра- зовав субкладу с последовательностями D. brauneri. Пример популяции D. daghestanica, обнаруженной в лесном биотопе без скальных выходов и генетически не отличающейся от особей с сопредельной тер- ритории, свидетельствует о способности скальных ящериц осваивать не свойственные им биотопы за короткий период времени. Генетическая дистанция между D. c. vedenica и D. с. caucasica сопоставима с таковой для пары D. с. caucasica и D. daghestanica. Мы предполагаем наличие самостоятельной формы D. caucasica ssp., обитающей в Южной Осетии и на сопредельных территориях.
Doronin, I.V. & Ermolina, L.P. (2012) -
A reptile catalogue of the Zoological Museum, Stavropol State University, is presented. 1,019 specimens of 88 species collected within the territories of Russian Federation, Ukraine, Georgia, Kazakhstan, Uzbekistan, Turkmenistan, and some other areas are listed. The history of the Museum and the formation of its collection are given.
Доронин И.В., Ермолина Л.П. (2012) -
Представлен каталог рептилий Зоологического музея Ставропольского государственного университета, в котором дается информация о 1019 экз. 88 видов, собранных на территориях России, Украины, Грузии, Казахстана, Узбекистана, Туркмении. Рассмотрена история создания музея и формирования коллекции.
Doronin, I.V. & Ljubisavljevic, K. (2014) -
И.В. Доронин, К. Любисавлиевич (2014) -
Doronin, I.V. & Lotiev, K.J. (2013) -
Doronin, I.V. & Melnikov, D.A. & Melnikova, E.N. (2016) -
e results of molecular genetic analysis of phylogenetic relationships in Darevskia (caucasica) on the basis of COI sequences variability is presented. Two main lineages are detected: the Central and Eastern Caucasian (clades A–B — D. caucasica-D. daghestanica), and Central and Western Caucasian (clades C–D — D. alpina). A DNA-barcoding tool can be successfully used for species identi cation of D. alpina, D. caucasica and D. daghestanica.
Доронин, И.В. Мельников Д.А., Мельникова Е.Н. (2016) -
Приведены результаты молекулярно-генетического анализа филогенетических взаимо- отношений скальных ящериц Darevskia (caucasica) на основе изменчивости фрагмента пер- вой субъединицы цитохром оксидазы (COI). На полученных деревьях четко выделяются две линии, одна из которых связана с Центральным и Восточным Кавказом (группы A–B — D. caucasica-D. daghestanica), а вторая — с Центральным и Западным Кавказом (группы С–D — D.alpina). Делается вывод об эффективности ДНК-штрихкодирования для дифференциации D. alpine, D. caucasica и D. daghestanica. Библиогр. 19 назв. Ил. 2. Табл. 2.
Doronin, I.V. & Smirnova N.G. (2022) -
New findings of Darevskia pontica at the northern border of its habitat, in the Kuban-Azov lowland within the administrative boundaries of the Krasnodar and Rostov regions, are presented and discussed. The species was found in the Rostov region for the first time. It is proposed to include D. pontica into the regional Red Data Book of the Rostov region.
И. В. Доронин & Н. Г. Смирнова (2022) -
Приводятся и обсуждаются новые находки понтийской ящерицы (Darevskia pontica) на северной границе ареала в Кубано-Приазовской низменности в администра- тивных границах Краснодарского края и Ростовской области. Для последнего региона вид указан нами впервые. Предложено включить D. pontica в региональную Красную книгу Ростовской области.
Doronin, I.V. & Tuniyev, B.S. (2015) -
Доронин И.В., Туниев Б.С. (2015) -
Doronin, I.V. & Tuniyev, B.S. (2018) -
The article describes the history of the study and data on the biology of the Szczerbak‘s Lizard, Darevskia szczerbaki - the endemic of Russia. The species is listed in the Red Data Book of Krasnodar Krai and is recommended for inclusion in the Red Data Book of Russian Federation. The main reason for this lizard extinction is the construction of coastal cliffs in the Krasnodar krai. Particular concern is caused by the state of its populations in a type locality — the town of Anapa. To protect a rock lizard, the following steps are necessary: optimization of the territory of the Utrish State Natural Reserve, establishing of new protected areas in Anapa, on the coastline between Gelendzhik and Tuapse, and prohibition of catching of lizards.
Доронин И.В. & Туниев Б.С. (2018) -
статье изложена история изучения ящерицы Щербака (Darevskia szczerbaki) и сведения о биологии этого узкоареального эндемика России. Вид занесен в Красную книгу Краснодарского края и рекомендован для включе
Doronin, I.V. & Tuniyev, B.S. & Kukushkin, O.V. (2013) -
The results of statistical analysis of morphological characters and the study of the cytochrome b gene variability (mtDNA) of rock lizards of the Darevskia (saxicola) complex distributed in the Caucasus and the Crimea, have shown significant differentiation within this complex, assuming specific independence of D. szczerbaki (Lukina, 1963) and subspecific status of recently described D. brauneri myusserica Doronin, 2011. According to our data, within D. brauneri (Méhely, 1909) only two forms of subspecific status can be distinguished, namely D. b. brauneri and D. b. myusserica. In our opinion, Lacerta saxicola darevskii Szczerbak, 1962 (= D. brauneri darevskii) should be considered to be a junior synonym of D. b. brauneri. The possible scenarios of range formation in the Darevskia (saxicola) complex are discussed.
И.В. Доронин, Б.С. Туниев и О.В. Кукушкин (2013) -
Результаты статистического анализа морфологических признаков шести распространенных на Кавказе и в Крыму форм скальных ящериц комплекса Darevskia (saxicola), наряду с данными исследования изменчивости фрагмента гена цитохром b митохондриальной ДНК, указывают на глубокую дифференциацию внутри этого комплекса, говорят в пользу видовой самостоятельности D. szczerbaki (Lukina, 1963) и обособленности недавно описанного таксона D. brauneri myusserica Doronin, 2011. Вместе с тем внутривидовая изменчивость D. brauneri (Mehely, 1909) позволяет выделить только две валидные формы подвидового статуса: D. b. brauneri и D. b. myusserica. По нашему мнению, Lacerta saxicola darevskii Szczerbak, 1962 (= D. brauneri darevskii) должна рассматриваться как младший синоним D. b. brauneri. Обсуждаются возможные сценарии происхождения форм комплекса.
Dotsenko, I.B. (2007) -
The information is given on finding of males, having characters similar with Darevskia mixta (settled in 1968) and with D. armeniaca in the structure of the mixed experimental population of rocky lizards D. armeniaca and D.dahli. Data about distribution of the above mentioned species in the territory populated by them, characters of external morphology and others are cited. The possible reasons of not planned occurrence `D.dahli` in the population of D. armeniacа, including by hybridization are considered.
Dotsenko, I.B. & Darevsky, I.S. (2005) -
Specimens of Dahl’s rocky lizard Darevskia dahli (Darevsky, 1957) was found among the introduced on the territory of Ukraine D. armeniaca (Mehely, 1909). The 11 specimens of Dahl’s rocky lizard was compared D. armeniaca from the same location by 16 external features and with 15 specimens of D. dahli from Georgia by 2 features. The results were commented.
Dotsenko, I.B. & Melnichenko, R.K. & Demidova, M.I. (2016) -
Data on biology of the partenogenetic rock lizards of Darevskia which are artificially acclimatized in Ukraine are provided. Biotopes and station of the mixed population of introduced species of D. armeniaca and D. dahli near the Denishi village in the Zhytomyr region are described. Expansion of an area of these species on inflows of the river Teteriv is noted. The new location of rocky lizards on the Bobrovka River near the Buki village is described. The seasonal and daily activity of lizards and density of their settlement in this region are investigated. The review of publications about population of introduced species of rocky lizards is provided. The factors limiting resettlement of parthenogenetic rocky lizards and promoting it are analyzed.
Доценко, И. Б. & Мельниченко, Р.К. & Демидова, М.И. (2016) -
Приводятся данные об особенностях биологии пapтeнoгeнeтических cкальных ящеpиц poда Darevskia, искусственнo aкклимaтизoвaнных в Укpaине. Описаны биотопы и стации смешанной популяции D. armeniaca и D. dahli в окрестностях c. Дeниши в Житoмиpской обл. Отмечается расширение ареала этих видов по притокам р. Тетерев. Детально описана новая локация скальных ящериц на р. Бобровка близ с. Буки. Изучены сезонная и суточная aктивноcть ящеpиц и плотноcть их в исследованном регионе. Приво- дится обзор публикаций о популяции интродуцентов. Анализируются факторы, лимитиру- ющие расселение партеногенетических скальных ящериц и способствующие ему.
Dotsenko, I.B. & Peskov, V.N. & Miropolskaya, M.V. (2008) -
Drohvalenko, M. & Mykhailenko, A. & Rekrotchuk, M. & Shpak, L. & Shuba, V. & Trokhymchuk, R. & Utevsky, S. & Zinenko, O. (2019) -
A part of the COI mitochondrial barcoding gene was sequenced from seven species of diff erent taxonomical groups: Ambystoma mexicanum (Amphibia, Ambystomatidae), Darevskia lindholmi, Lacerta agilis exigua (Reptilia, Lacertidae), Erinaceus roumanicus (Mammalia, Erinaceidae), Macrobiotus sp. 1 and 2 (Eutardigrada, Macrobiotidae) and Cameraria ohridella (Insecta, Gracillariidae). Th e sequences were compared with available sequences from databases and positioned on phylogenetic trees when the taxa had not yet been sequenced. Th e presence of Mexican axolotls in herpetoculture in Ukraine was confi rmed. Th e partial COI genes of the Crimean rock lizard and an eastern sub-species of the sand lizard were sequenced. We demonstrated the presence of two tardigrade mitochondrial lineages of the Macrobiotus hufelandi group in the same sample from the Zeya Natural Reserve in the Far East: one was nearly identical to the Italian M. macrocalix, and the other one is similar to M. persimilis and M. vladimiri. We also confi rmed the presence of the invasive haplotype “A” of the horse chestnut leaf miner in Ukraine, in line with the hypothesized route of invasion from Central Europe.
Drosdov, N.N. (1964) -
Dunaev, E.A. & Orlova, V.F. (2017) -
From the publisher:
Before you a unique atlas-determinant. For the first time in one edition descriptions of all species of amphibians and reptiles living on the territory of Russia are presented, illustrated with beautiful color photographs. The book is written by E.A. Dunaev and V.F. Orlova, the famous Russian herpetologists, researchers of the Zoological Museum of Moscow State University named after MV Lomonosov. Lomonosov. A simple and accessible form of presentation, detailed descriptions of the external appearance, lifestyle and behavior, photographs reflecting all variants of the variability of these animals, will help the curious reader to become more familiar with turtles, snakes, lizards, frogs and other representatives of the herpetofauna. And special keys will allow you to determine who you met on a country walk, in a city park, at a dacha or while traveling to different corners of our country.
The book is of great interest not only for nature lovers, but also for specialists in herpetology and students of biological faculties of universities. It lists the amphibians and reptiles of the native fauna, reflects the latest changes in the taxonomy of these groups, lists species included in the Red Book of Russia and regional Red Books, lists reference and scientific publications on reptiles and amphibians of Russian fauna.
Дунаев, Е.A. & Орлова, B.Ф. (2017) -
Перед вами уникальный атлас-определитель. Впервые в одном издании представлены описания всех видов земноводных и пресмыкающихся, обитающих на территории России, проиллюстрированные прекрасными цветными фотографиями. Книга написана Е.А. Дунаевым и В.Ф. Орловой, известными отечественными герпетологами, научными сотрудниками Зоологического музея Московского государственного университета имени М.В. Ломоносова.Простая и доступная форма изложения, подробные описания внешнего облика, образа жизни и поведения, фотографии, отражающие все варианты изменчивости этих животных, помогут любознательному читателю ближе познакомиться с черепахами, змеями, ящерицами, лягушками и другими представителями герпетофауны. А специальные ключи позволят определить, с кем именно вы встретились на загородной прогулке, в городском парке, на даче или во время путешествий по различным уголкам нашей страны.
Книга представляет большой интерес не только для любителей природы, но и для специалистов-герпетологов и студентов биологических факультетов вузов. В ней приведен полный список амфибий и рептилий отечественной фауны, отражены новейшие изменения в таксономии этих групп, указаны виды, включенные в Красную книгу России и региональные Красные книги, перечислены справочные и научные издания, посвященные рептилиям и амфибиям фауны России.
Dunn, W. (2016) -
Dzhelali, P.A. & Doronin, I.V. & Lotiev, K.Y. & Mazanaeva, L.F. & Mustafifaeva, G.A. & Bunyatova, S.N. (2021) -
Dzukic, G. & Kalezic, M.L. (2001) -
Eiselt, J. (1965) -
Eiselt, J. (1970) -
Als erste Belege für das Vorkommen in der Türkei werden eingehend
besprochen: Lacerta praticola pontica, Varanus griseus griseus, Spalerosophis
diadema cliffordi, Eirenis punctatolineatus kumerloevei ssp. nov., Eirenis persicus
und Eirenis rothii.
Darüber hinaus werden noch Funde folgender Schlangen erwähnt:
Coluber jugularis jugularis, C. jugularis schmidti, Eirenis punctatolineatus
punctatolineatus, Eirenis modestus modestus, Rhynchocalamus satunini.
Eiselt, J. (1995) -
A new species of Archaeolacerta from northeastern Iran is described in comparison with other archaeolacertids - chlorogaster and defilippi from Iran, as well as raddei and valentini from Transcaucasia.
Eiselt, J. & Darevsky, I.S. (1991) -
Conventional and multivariate taxonomic methods allow the description of a new subspecies of
Lacerta rudis BEDRIAGA from the Central Caucasus: L. rudis chechenica ssp. n. - In this connection the
intraspecific relations between the northeastern groupings of Lacerta rudis are demonstrated. - The
data-analyses used are described.
Eiselt, J. & Darevsky, I.S. & Schmidtler, J.F. (1992) -
Series of newly collected specimens of Lacerta valentini BOETTGER in Eastern Turkey were closely
investigated in order to demarcate the areas of the subspecies L. v. valentini and L. v. lantzicyreni.
Intermediary populations were found to exist between them, one was described as a new subspecies,
Lacerta valentini spitzenbergerae, living in the Cilo-Dag Mountains, Vilayet Hakkâri, Turk
Eiselt, J. & Schmidtler, J.F. & Darevsky, I.S. (1993) -
A new subspecies of Lacerta raddei BOETTGER, 1892 is described. Terra typica: castle mountain of
Van City, eastern Turkey;. Distribution: easternmost Turkey. Intermediary populations towards L. raddei
raddei in northwesternmost Iran are still to be defined.
Eksilmez, H. & Altunisik, A. (2020) -
The Artvin Lizard, Darevskia derjugini (Nikolsky, 1898), is distributed in the Black Sea coastal region of Turkey (Provinces of Trabzon, Rize and Artvin), Georgia, Azerbaijan and Russia.The vertical distribution of the species is up to 1700 m a.s.l. It is classified as “near threatened” in the IUCN red List of Threatened species. In this study, we investigated age structure, age at sexual maturity and lifespan (as demographic life-history traits) as well as body size of a Darevskia derjugini population (n=30, 13 males, 17 females) from Murgul, Artvin, using skeletochronology. Our findings indicated that age upon attaining sexual maturity was three years for both sexes in this high-altitude population. The maximum longevity was seven years. There was no statistically significant difference between male and female individuals in terms of average age. Female individuals were significantly larger than male individuals. As reported for many lizards, we found a significant positive relationship between age and body size within each sex of Artvin lizard. The information given about some life history characteristics of a near-threatened species in this study may contribute to effective conservation management in the future.
Eversmann, E. (1834) -
Fedorov, A.N. & Fedorova, L.V. & Grechko, V.V. & Ryabinin, D.M. & Sheremet`eva, V.A. & Bannikova, A.A. & Lomov, A.A. & Ryskov, A.P. & Darevsky, I.S. (1999) -
A specially optimized restriction analysis of
highly repetitive DNA elements, called DNA taxonprint,
was applied for phylogenetic study of primates and lizards.
It was shown that electrophoretic bands of DNA
repeats revealed by the taxonprint technique have valuable
properties for molecular systematics. Approximately
half of taxonprint bands (TB) are invariable and
do not disappear from the genomes during evolution or
change spontaneously. Presumably these invariable
bands are restriction fragments of dispersed DNA repeats.
Another group represents variable taxonprint
bands that differ even between closely related species.
These variable bands are probably represented by tandem
DNA repeats and could be used as species-specific
markers. It was shown that taxonprint bands are independent
characters since the appearance of a new taxonprint
band does not change the previous band pattern.
Phylogenetic reconstruction carried out on taxonprint data
demonstrated that this approach could be of general utility
for molecular systematics and species identification.
Ficetola, G.F. & Falaschi, M. & Bonardi, A. & Padoa-Schioppa, E. & Sindaco, R. (2018) -
The analysis of biogeographical structure and patterns of endemism are central topics of biogeography, but require exhaustive distribution data. A lack of accurate broad-scale information on the distribution of reptiles has so far limited the analyses of biogeographical structure. Here we analysed the distribution of reptiles within the broad-sense Western Palearctic to assess biogeographical regionalization using phylogenetic and non-phylogenetic approaches, identified areas of endemism and evaluated the environmental factors pro- moting community uniqueness and endemism. We gathered distributional records from the literature and from the field, mapping the distribution of all the Western Palearctic reptiles on a 1-degree resolution grid. bsim dissimilarity and hierarchical clustering was used to identify bioregions, analysing data both at the species and at the genus level, and considering phylogenetic dissimilarity. Consensus areas of endemism were identified on the basis of the optimality criterion. We then assessed whether biogeographical structure is related to present-day climate, insularity, orography and velocity of climate change during the Late Qua- ternary. The genus-level analysis identified five main biogeographical regions within the Western Palearctic, in partial agreement with previous proposals, while the species-level analysis identified more bioregions, largely by dividing the ones identified by genera. Phylogenetic bioregions were generally consistent with the non- phylogenetic ones. The strongest community uniqueness was observed in subtropical warm climates with seasonal precipitation and low productivity. We found nine consensus areas of endemism, mostly in regions with limited velocity of Quaternary climate change and warm subtropical climates. The biogeographical structure of Western Palearctic reptiles is comparable to what has been observed in other vertebrates, with a clear distinction between the Saharo-Arabian-Sindian and Euro-Mediterranean herpetofaunas. Unlike other vertebrates, in reptiles the highest uniqueness and endemism is observed in dry climates, but the velocity of climate change during the Quaternary remains a major driver of endemism across all the vertebrates.
Fitch, H.S. (1970) -
Franzen, M. (1990) -
Franzen, M. (1991) -
Franzen, M. (2000) -
During a short visit at the Turkish Black Sea coast between Trabzon and Hopa in December 1989 lizards of the Darevskia rudis complex, D. parvula and D. derjugini were regulary observed being active. At three sites densities of D. rudis complex were similar to those observed at the same localities during the summer months. Lizards seemed to be fully active in such cases, and once feeding was observed. Compared to the Mediterranean zone of southwestern Turkey, where Lacerta oertzeni was observed being fully active only twice during an eight week survey (December, January), winter activity of lizards seerns to occur more regulary at the Black Sea coast. This may be due to an overall low insolation caused by extensive rainfalls, even during the summer months.
Franzen, M. & Heckes, U. (1992) -
Following the first report of Lacerta praticola from Turkey by ErsEL T (1970), two new locali- ties have now been found near Cucurpmar (SE Dereköy), Vil. K1rklareli . In the Istranca moun- tains, Lacerta praticola is an inhabitant of premontane , su bxerophilous oak forests and exposed edge habitats within mountain beech forests . The collected 4 specimens belang to the subspecies L. praticola pontica.
Freise, K. & Müller, G. (1962) -
Freitas, S. & Perera, A. & Harris, D.J. & Arakelyan, M. & Danielyan, F. & Corti, C. & Carretero, M.A. (2010) -
The genus Darevskia is a lacertid group radiating and diversifying in the Caucasus region. The studies on the phylogeny of this group started early, however, the phylogeographic pattern within this genus is far to be complete. Dareskia raddei has been described for Armenia and adjacent S Georgia, E Turkey and N Iran. D. nairensis from Armenia is sometimes considered conspecific although evidences from morphologyand proteins are conflicting. Here, we assess the phylogeographic relationships based on the analysis of an extensive sampling of specimens morphologically assigned to both species throughout most of the range using the Cytochrome b mitochondrial marker. Results suggest that D. raddei is paraphyletic to D. nairensis for mtDNA, confirming previous evidence of proteins. Nevertheless, the group portschinskii-valentini is confirmed as the sister clade of the raddei-nairensis complex. Within this complex, strong phylogeographic structure is observed. Three main clades are observed: one comprising individuals from Georgia and the most northern and western Armenia, the second comprising individuals from Southern Armenia, and the last with one individual from Iran. The Darevskia unisexualis individuals (i.e. D. raddei and D. valentini parthenogenetic hybrids) analysed in this study fall within the first clade. Because of that we can say that the putative parental D. raddei populations may have been individuals present in between Georgia and the most western part of the current range distribution of D. raddei. The overall pattern suggests an expansion of D. raddei from SE to the rest of the range which is correlated with the recent paleogeographic history of the region.
Freitas, S. & Perera, A. & Harris, D.J. & Arakelyan, M. & Danielyan, F. & Corti, C. & Ilgaz, C. & Ahmadzadeh, F. & Garcia-Munoz, E. & Carretero, M.A. (2011) -
Due to its geographic location, the Caucasus region connects species from three diferent continents in a heterogeneous landscape, creating an environment prune to the development of speciation events and multiple endemisms. Among them, the lacertid genus Darevskia is a group of rock lizards radiating and diversifying in the region. Among their highly polymorphic members a number of parthenogenetic forms are present result from the hybridisation between bisexual Darevskia species. However, the origin of these forms lacks an explicit spatiotemporal context. One of the most interesting bisexual species in the group is Darevskia raddei which is responsible for the motherhood of some unisexual forms, namely D. unisexualis, D. uzzeli and D. bendimahiensis and D. sapphirina). Fruthermore, this species is considered conspeciic to D. nairensis and D. (raddei) vanensis, two forms of unclear status which link to the parthenogenetic forms need to be clariied. This, this study aims to reach a global comprehension of the three putative bisexual taxa within of the D. raddei complex (D. raddei, D. nairensis and D. (raddei) vanensis), and their phylogenetic relationships with the parthenogenetic forms they theoretically parented. A phylogenetic analysis was conducted on samples of the D. raddei complex, ranging from Turkey, Armenia, Georgia and Iran. Maximum Likelihood and Bayesian methods were applied on a combination of mitochondrial and nuclearmarkers. Namely, three mtDNA (cytochrome b, ND4 and 12s) and two nDNA (MC1R and c-mos) markers were used. Our results conirm that Darevskia raddei (sensu lato) is the proposed maternal species for at least D. unisexualis, D. uzzeli, D. bendimahiensis, D. sapphirina and D. rostombekowi, as already supported by previous studies. However, it does not support a genetic diferentiation between the three forms described within the complex, which are to be considered conspeciic. Moreover, there is reinforced evidence that the three parthenogenetic forms considered arose more than once in the history of the genus, from diferent lineages of the bisexual parents.
Freitas, S. & Rocha, S. & Campos, J. & Ahmadzadeh, F. & Corti, C. & Sillero, N. & Ilgaz, C. & Kumlutaş, Y. & Arakelyan, M. & Harris, D.J. & Carretero, M.A. (2016) -
Darevskia rock lizards include both sexual and parthenogenetic species, mostly distributed in the heterogeneous and ecologically diverse Caucasus. The parthenogenetic species originated via directional hybridogenesis, with only some of the sexual species known to serve as parentals. However, it remains unclear when and where these events happened and how many parental lineages were involved. A multilocus phylogeographic analysis was performed on the parthenogens D. unisexualis, D. bendimahiensis and D. uzzeli, and their putative maternal species D. raddei. Results show the parthenogenetic species all have relatively recent origins, approximately 200–70 kyr ago, and at least three hybridization events were involved in their formation. Ecological niche models identify the region where hybridization events leading to the formation of D. unisexualis took place, namely in the northeast of the current distribution. Models also suggest that the sexual D. raddei might have undergone a habitat shift between the Last Interglacial and the Last Glacial Maximum.
Freitas, S. & Vavakou, A. & Arakelyan, M. & Drovetski, S.V. & Crnobrnja-Isailovic, J. & Kidov, A.A. & Cogalniceanu, D. & Corti, C. & Lymberakis, P. & Harris, D.J. & Carretero, M.A. (2016) -
Darevskia praticola differs from the other species of the genus in having a large but disjunct distribution, covering the Balkan and the Caucasus regions. Furthermore, most Darevskia species occupy saxicolous habitats, whereas D. praticola inhabits meadows and forest environments. Here we determine the phylogeographic and phylogenetic relationships of Darevskia praticola sensu lato and evaluate the current, morphology-based taxonomy. We sequenced two mtDNA genes (Cyt-b and ND4) and two nuclear loci (MC1R and RELN) for samples collected across the species range. Because our sequences amplified with the Cyt-b primers appear to represent a nuclear pseudogene we excluded this marker from the final analysis. Our results support monophyly of D. praticola and show its division into three clades. The first divergence, dated to the Late Pliocene, is between the Balkans and the Caucasus. The Caucasus lineage is further subdivided in a western Greater Caucasus and a Transcaucasia clade, likely due to subsequent differentiation during the Pleistocene. Our findings do not support the current taxonomic arrangement within D. praticola. The main geographic divergence likely happened due to a vicariance event associated with Plio-Pleistocene climatic and vegetation oscillations.
Freitas, S. & Westram, A.M. & Schwander, T. & Arakelyan, M. & Ilgaz, C. & Kumlutas, Y. & Harris, D.J. & Carretero, M.A. & Butlin, R.K. (2022) -
Hybridization is a common evolutionary process with multiple possible outcomes. In vertebrates, interspecific hybridization has repeatedly generated parthenogenetic hybrid species. However, it is unknown whether the generation of parthenogenetic hybrids is a rare outcome of frequent hybridization between sexual species within a genus or the typical outcome of rare hybridization events. Darevskia is a genus of rock lizards with both hybrid parthenogenetic and sexual species. Using capture sequencing, we estimate phylogenetic relationships and gene flow among the sexual species, to determine how introgressive hybridization relates to the origins of parthenogenetic hybrids. We find evidence for widespread hybridization with gene flow, both between recently diverged species and deep branches. Surprisingly, we find no signal of gene flow between parental species of the parthenogenetic hybrids, suggesting that the parental pairs were either reproductively or geographically isolated early in their divergence. The generation of parthenogenetic hybrids in Darevskia is, then, a rare outcome of the total occurrence of hybridization within the genus, but the typical outcome when specific species pairs hybridize. Our results question the conventional view that parthenogenetic lineages are generated by hybridization in a window of divergence. Instead,
Freitas, S.M.R.N. de (2017) -
Considering all costs associated to sex, sexual reproduction could be expected to be rare. However, we find sexual reproduction to be pervasive in nature, and only a reduced number of taxa are completely asexual. In fact, the variety of sex reproductive systems, mating techniques and traits, reveals that the organisms’ reproduction has been an expressive target of selection and adaptation throughout evolution. Sex is selected because it facilitates adaptation in complex and changing environments (Weismann, 1889), and this has been already shown empirically in several models. However, studies and theories developed concerning the paradox of sex will not be able to clarify the current distribution and abundance of asexual lineages in nature. The understanding of rates of origin and extinction of asexual lineages, together with the putative differential evolution rates in organisms with sexual or asexual reproduction, should be considered in the main “origin of sex” question. The model used here, the genus Darevskia, is the first true parthenogenetic vertebrate described. It is a highly speciose genus, currently including 27 species, of which seven are parthenogenetic and of hybrid origin. Only a few of the sexual species contributed for the hybridization events which originated the parthenogens clonal lineages, and these were directional: the maternal species were always Darevskia raddei or D. mixta, and the paternal always D. valentini or D. portshinskii. Using a set of microsatellite and mtDNA markers, we first start to study the role of hybridization in the origin of vertebrate parthenogens, the extent of gene flow with sexual relatives and the relation between hybridization, asexuality and polyploidy, in the context of the main theories of asexual evolution proposed to date. We focus on the D. unisexualis, D. uzzelli and D. armeniaca parthenogens and polyploid backcrosses found in sympatric locations between the parthenogens and its sexual parentals. We find that only specific parental pairs are responsible for the origin of vertebrate parthenogenesis, regardless of their phylogenetic distance. Despite the recurrent hybridization presently reported in Darevskia, asexuality originated multiple times but only in a single temporal event the past. Parthenogenetic females are capable of backcrossing with sexual males, but can only produce polyploid individuals with reduced fertility that do not contribute for gene flow between parthenogens and sexuals. The sexual reproduction machinery can be lost with time in the parthenogenetic hybrids, which are most likely the result of a post-zygotic reproductive barrier to gene flow in the speciation continuum of Darevskia sexual species. In order to reconstruct the phylogenetic inference of this group, a set of capture sequence probes were designed from a transcriptome de novo assembly. Phylogenetic inference reconstruction has been extensively used to ask several evolutionary biology questions. However, despite the exponential use of high-throughput sequencing technologies and its promising applications in phylogenetics, next generation sequencing (NGS) techniques are still far from being as widely used in this field as in other areas of evolutionary genetics. The workflow developed in this thesis has proved to be not only cost-effective, but also to produce a very high number of cross genome phylogenetically informative markers in this non-model species. Finally, taking advantage of the hundreds of cross genome markers developed, a species tree was constructed and within genus relationships evaluated. Introgression tests showed a widespread pattern of gene flow across the genus, not only recent between recently diverged clades, but also both recent and ancient gene flow between early diverging Darevskia clades. Several evidences of hybridization with gene flow between diverging clades have been found in other systems. However, such widespread patterns where introgression has happened between all major clades, and in a very high number of species pairs, are not frequently found. It is also found that gene flow is completely absent between parthenogens sexual parental pairs. In groups with sexual-parthenogenetic reproducing species, pre- or post-zygotic reproductive barriers could be more difficult to attain and hybrid asexuality can, thus, effectively create reproductive barriers between diverging sexual taxa, that were likely faced with recurrent secondary contact. The work presented in this thesis shows that only a few sexual species pairs were responsible for the origin of the parthenogens, that some species acted always as the maternal species and others always as the paternal, that the phylogenetic distances between each pair are included in a wide range interval, and that introgression is found widespread across the genus but absent between the parental pairs. Regardless of the reproduction type, this study also shows the relevance and depth of hybridization during divergence. As has been shown intensively, gene flow is common during speciation and clades divergence. However, it would be interesting to analyse other groups together and test if deep branch gene flow is found only in some exclusive groups (such as sexual-asexual genera) or if it is something more widespread. This work presented here moves Darevskia towards providing a model that can be used to understand the origin of asexuality in vertebrates. Here the basis for future studies about sex and asexuality is provided, so that a focus on the differential evolution rates genomes of both reproduction mode can be analysed.
Freitas, S.N. & Harris, D.J. & Sillero, N. & Arakelyan, M. & Butloin, R.K. & Carretero, M.A. (2019) -
Freitas, S.N. & Harris, D.J. & Sillero, N., & Arakelyan, M. & Butlin, R.K. & Carretero, M.A. (2019) -
Obligate parthenogenesis is found in only 0.1% of the vertebrate species, is thought to be relatively short lived and is typically of hybrid origin. However, neither the evolutionary persistence of asexuality in vertebrates, nor the conditions that allow the generation of new parthenogenetic lineages are currently well understood. It has been proposed that vertebrate parthenogenetic lineages arise from hybridisation between two divergent taxa within a specific range of phylogenetic distances (the ‘Balance Hypothesis’). Moreover, parthenogenetic species often maintain a certain level of hybridisation with their closest sexual relatives, potentially generating new polyploid hybrid lineages. Here we address the role of hybridisation in the origin and evolutionary lifespan of vertebrate parthenogens. We use a set of microsatellite markers to characterise the origins of parthenogens in the lizard genus Darevskia, to study the distinctiveness of sexual and asexual taxa currently in sympatry, and to analyse the evolutionary consequences of interspecific hybridisation between asexual females and sexual males. We find that parthenogens result from multiple past hybridisation events between species from specific lineages over a range of phylogenetic distances. This suggests that the Balance Hypothesis needs to allow for lineage-specific effects, as envisaged in the Phylogenetic Constraint Hypothesis. Our results show recurrent backcrossing between sexual and parthenogenic Darevskia but neither gene flow nor formation of new asexual lineages. We suggest that, along with their demographic advantage, parthenogens gain additional leverage to outcompete sexuals in nature when the retention of sexual reproductive machinery allows backcrossing with their sexual ancestors.
Frotzler, N. & Bader, T. (2007) -
Frynta, D. & Moravec, J. & Čiháková, J. & Sádlo, J. & Hodková, Z. & Kaftan, M. & Kodym, P. & Král, D. & Pitule, V. & Šejna, V. (1997) -
Fu, J. & Darevsky, I.S. & MacCulloch, R.D. & Kupriyanova, L.A. & Roitberg, E.S. & Sokolova, T.M. & Murphy, R.W. (1995) -
The genetic diversity of 34 loci was surveyed from among five populations of the supposed conspecific taxa Lacerta c. caucasica, L. c. alpina, and L. c. daghestanica. Twelve loci exhibited variation. Fixed or nearly fixed allelic differences between L. c. caucasica and L. c. daghestanica were found at two loci, mannose-6-phosphate isomerase-A and creatine kinase-C. These two taxa differed from L. c. alpina at eight loci. Nei`s (1978) genetic distance values among populations of L. c. daghestanica ranged from 0.000 - 0.029, between L. c. caucasica and L. c. daghestanica from 0.076 - 0.087 and between L. c. alpina and the other taxa from 0.472 - 0.501. Fixed allelic differences and consistent morphological character states support the recognition of these three taxa as separate species. Therefore, we recommend use of the names L. alpina, L. caucasica, and L. daghestanica for these taxa.
Fu, J. & MacCulloch, R.D. & Murphy, R.W. & Darevsky, I.S. (2000) -
Clonal variation in Lacerta armeniaca was investigated using allozyme electrophoresis and morphology. Among the 35 allozyme loci examined, three were variable, which divided L. armeniaca into four clones. One rare clone of L. armeniaca made up the majority of two populations. This contrasts to rare clones in other parthenogenetic Caucasian rock lizards which typically consist of only one or two individuals. Another rare clone, which showed a striking colouration difference, had different allelic composition at two loci. Although mutation is a possible explanation of the origin of the clonal variation, the alternative, multiple origin, is equally likely.
Fu, J. & MacCulloch, R.D. & Murphy, R.W. & Darevsky, I.S. & Kupriyanova, L.A. & Danielyan, F. (1998) -
Protein electrophoresis of Lacerta unisexualis from three populations found that 21 of 36 allozyme loci were homozygous, while 14 expressed fixed heterozygotes and one locus was variable. Three clones were detected at the locus Cat-A. Two individuals represent two rare clones while all others form a common clone. Our favored explanation is the mutation of a preexisting common clone rather than multiple origins.
Fu, J. & MacCulloch, R.D. & Murphy, R.W. & Darevsky, I.S. & Tuniyev, B.S. (2000) -
Allozyme electrophoresis of four sibling parthenogenetic Caucasian rock lizards Darevskia unisexualis, D.uzzelli, D.sapphirina, and D.bendimahiensis found seven clones and five variable loci. The data supported the hypothesis that D.raddei and D.valentini are the parental species of all four parthenogens. Variation patterns in Darevskia were summarized. Species that originated from a single F1 typically consisted of one widespread clone with a few rare clones. Species with multiple origins displayed variation only slightly higher than species with a single origin. This is contrary to other genera of parthenogenetic lizards, in which cases massive clonal variations were observed.
Fu, J. & Murphy, R.W. & Darevsky, I.S. (1997) -
A maximum parsimony phylogeny of 14 Caucasian species of rock lizards, genus Ijicerta, subgenus Archaeolacerta, was constructed from mitochondrial cytochrome b and ATPase 6 partial gene sequences. Congruence analyses were carried out between the two genes. A synthesis of the data sets reveals three well supported monophyletic groups: (1) the caucasica group including (Lacerta derjugini ((Lalpina, L. clarkorum) (L. caucasica, L. daghestanica))); (2) the rudis group including (L. parvula (L. portschinskii (L. vakntini, L. rudis))); and (3) the saxicola group including (L. mixta (L. nairensis (L. saxicola, L. raddeijj). Despite the diagnosis of three groups, the placement of L. praticola as a basal taxon is uncertain, as are the relationships among the three groups. The mitochondrial DNA sequence data suggested prior hybridization between L. mixta and L. alpina and possibly between L. saxicola and L. nairensis. Lacerta raddei was resolved as a paraphyletic species on the mtDNA tree; this may result from either hybridization or random gene sorting.
FU, J. & Murphy, R.W. & Darevsky, I.S. (1999) -
Little mtDNA variation was observed among populations of the bisexual Caucasian rock lizard Lacerta mixta and unisexual L. dahli and L. armeniaca. Three haplotypes were detected in L. mixta and the maximum pairwise difference among the samples was 0.67%. No intra- and interspecific variation was found among populations of either L. armeniaca or L. dahli. Moreover, both unisexual species were identical to one of the three haplotypes of L. mixta. The limited variation in L. mixta is likely the result of bottleneck effect, although the small sample size may also be responsible. The lack of variation in the unisexuals was attributed to the restricted variation among the maternal parents, limited involvement of females in the hybridization, and recent origin.
Fuhn, I.E. (1932) -
Fuhn, I.E. (1970) -
Fuhn, I.E. & Hirsu, M. (1962) -
Gabelaia, M. (2019) -
Gabelaia, M. & Adriaens, D. & Tarkhnishvili, D. (2017) -
The genus Darevskia comprises over 20 species of small-bodied lizards, mainly occurring in the Caucasus Mountain Region. They show differences in body size, scalation and coloration, however, fully diagnostic characters that could separate the species with a high confidence level are still lacking. The early phylogenetic hypothesis of this group was based on ‘traditional’ analysis of morphology, based on multiple body and head measurements and scalation traits. Later, a molecular phylogeny of the genus rejected some of the proposed topology based on morphological traits. In this paper, we used quantitative morphological data (outline-based shape data) to test phylogenetic similarities, as proposed by these earlier hypotheses. We analyzed the pileus shape and the anal area of more than 200 individuals, representing six species of Darevskia, using outline based elliptic Fourier analysis. The analysis did confirm a clustering of the individuals and species (using UPGMA) from the same mitochondrial DNA clade. Hence, the phylogenetic affinity of the major clades in Darevskia could be identified using both molecular methods and outline analysis, whereas a traditional quantitative morphological analysis could not.
Gabelaia, M. & Tarkhnishvili, D. & Murtskhvaladze, M. (2015) -
The Caucasian rock lizard Darevskia mixta was sampled and studied from throughout its range, using mitochondrial cytochrome b sequences and scalation. The populations of the Greater and the Lesser Caucasus are reciprocally monophyletic matrilineally, and the respective lineages have been separated since the mid-Pleistocene. The lizards from the Greater Caucasus commonly have an unpaired preanal scale, whereas the lizards from the Lesser Caucasus have an additional scale behind the central temporal and subdivided interparietal scale more commonly than those from the Greater Caucasus. The Lesser Caucasus populations are further subdivided into two geographically distinct matrilineages, and are more diverse genetically and morphologically than the Greater Caucasus populations. The central part of the Lesser Caucasus is suggested to be the ancestral area for the entire D. mixta lineage. Successive Pleistocene periods of glaciation appear to be responsible both for the isolation of D. mixta from its common relatives, and for subdivision within D. mixta. Presence of D. mixta in NE Turkey is challenged.
Gabelaia, M. & Tarknishvili, D. & Adriaens, D. (2018) -
Geometric morphometrics is a better tool to evaluate the variation of shape than ‘traditional’ morphometrics. In reptiles, it outperforms morphometrics based on linear measurements and scalation. In an earlier study, two-dimensional outline-based geometric morphometrics in six species of rock lizards (Darevskia) showed that their shapes reflected the species divergence pattern. This allowed us to separate species from different clades, but we could not distinguish closely related species within a clade. We hypothesized that three-dimensional (3D) head shape data would be sufficiently discriminative to identify closely related species. To test this hypothesis, we compared head shapes of three closely related species of the ‘rudis’ clade, Darevskia portschinskii, Darevskia valentini and Darevskia rudis, with the last species treated as two distinct groups (D. r. obscura vs. D. r. rudis), using 3D landmark data. The 3D analysis isolated the species of the ‘rudis’ clade from each other and even separated individuals from populations that showed a genetic introgression pattern. The analysis showed that D. r. obscura is morphologically as distinct from D. r. rudis as from the other nominal species. For this reason, we suggest elevating the status of D. r. obscura to species level, i.e. Darevskia obscura Lantz & Cyrén, 1936.
Gabriel, C.D. (1970) -
Gaceu, O. & Josan, I. (2013) -
In the spring of 2013 we identified the species Darevskia pontica in two localities situated on the slope of Metaliferi Mountains, north of the Mureş River. D. pontica inhabits forested areas, being present in the open, grassy sectors near some brooks or on some forest roadsides.
Galoyan, E. (2013) -
According to kin selection theory, competition among monoclonal animals must be lower than between unrelated individuals. Thus, we propose that home range and core area overlap in parthenogenetic lizards will be broader than female range overlap in similar gonochoristic species. To test this hypothesis, we examined home range locations and space use of parthenogenetic Armenian Rock Lizards (Darevskia armeniaca) and compared them with home range locations and space use of gonochoristic species. We demonstrated that parthenogenetic Rock Lizards have a total range structure typical of insectivorous lizards, consisting of a sally zone, a home range, and one or more core areas. Some core areas contained activity centers, associated strongly with key basking sites and shelters. Provisional residents were found within the same range for 1 or 2 years, whereas wanderers visited study sites for 1 or 2 weeks per season. Settlement structure varied greatly among years. Home ranges, core areas, and even activity centers and basking sites of parthenogenetic females overlapped extensively, unlike in females of nonparthenogenic species. Monoclonal origins and high level of relatedness within unisexual species are possible explanations of the extended range overlap among parthenogenetic females.
Galoyan, E. & Bolshakova, A. & Abrahamyan, M. & Petroyan, R. & Komarova, V. & Spangenberg, V. & Arakelyan, M. (2019) -
Valentin’s rock lizard (Darevskia valentini) is suggested to be the parent for several parthenogenetic species (e.g., D. armeniaca, D. bendimahiensis, D. sapphirina, and D. unisexualis) that evolved through hybridization. Complex evolutionary processes (including reticulate evolution) are occurring within the areas where Valentin’s rock lizard coexists with these and other rock lizards. Hence, a detailed biological specification of this species is important for understanding how vertebrates evolve. Valentin’s rock lizard is a long-lived (up to 9 years), small diurnal lizard with larger females than males, which is unlike other species of the genus. Their relatively large eggs and early reproduction period, which occurs just after emergence from winter shelters, are adaptations for living in a high elevation climate (higher than 2 000 m a.s.l.). Their body temperatures (31–32 °С) are comparable to body temperatures of rock lizards living in milder climates, though female body temperature is more dependent on substrate temperature and basking due to their lower activity than that found in males. Population density fluctuates from several individuals to several hundred per hectare and is not affected by parthenogen coexistence, although hybrids do occur in sexually biased populations where males are more common than females. The male home range is larger than that of females, though these home ranges broadly overlap. Prey is not limited in the mountain meadows and Valentin’s rock lizards feed on a great variety of arthropods. Infanticide occurs in high-density populations.
瓦伦汀岩蜥（Darevskia valentini）被认为可能是几种孤雌生殖蜥蜴（如D. armeniaca、D. bendimahiensis、D. sapphirina和D. unisexualis）的祖先，前者通过杂交产生后者。瓦伦汀岩蜥和其它几种蜥蜴的共有的分布区域中，发生了复杂的进化过程（包括网状进化过程），因此，详细了解该物种生物学背景，有利于我们理解脊椎动物的进化过程。瓦伦汀岩蜥是一种寿命较长的（最多可达9年）小型蜥蜴，昼行性，雌性个体体型大于雄性，这一点和同属的其它物种都不一样。该物种的卵相对较大，且繁殖期较早（离开冬眠场所不久以后），这是一种对高海拔生存环境（大于2000米）的适应特征。尽管雌性由于活跃度更低（相较于雄性），其体温更依赖于生活环境的地层温度和取暖行为（晒太阳），但瓦伦汀岩蜥的体温（31–32 °С）和其他生活在温和气候地区的岩蜥区别并不大。瓦伦汀岩蜥的群体密度波动较大，从每公顷几个个体到几百个个体不等，且并不受孤雌生殖的影响。事实上杂交在性别比例偏倚的群体（雄性数量大于雌性）中仍然可能发生。雄性的领域范围比雌性更大，尽管领域间存在广泛的重叠。瓦伦汀岩蜥主要捕食节肢动物，捕食行为发生在包括高山草甸地区的很多地区。群体密度较高时会发生杀婴现象。
Galoyan, E. & Moskalenko, V. & Gabelaia, M. & Tarknishvilii, D. & Spangenberg, V. & Chamkina, A. & Arakelyan, M. (2020) -
The two species of rock lizards, Darevsia raddei and Darevskia portschinskii, belong to two different phylogenetic clades of the same genus. They are supposed ancestors for the hybrid parthenogenetic, Darevskia rostombekowi. The present study aims to identify morphological features of these two species and the potential gene introgression between them in the area of sympatry. External morphological features provided the evidence of specific morphology in D. raddei and D. portschinskii: the species differed in scalation and ventral coloration pattern, however, they had some proportional similarities within both sexes of the two species. Males of both species had relatively larger heads and shorter bodies than females. Males of D. raddei were slightly larger than males of D. portschinskii and had longer hindlimbs. Microsatellite genotyping revealed no traces of hybridization or gene flow among these species. We suggest that the absence of individuals with combined morphological and genetic features of the studied species reflects existing reproductive barriers. The hybridization between two bisexual rock lizard species from distant clades is rare and only takes place under very specific environmental conditions.
Galoyan, E.A. (2007) -
Galoyan, E.A. (2010) -
Genus Darevskia - diverse multispecies group, which in addition includes gay bisexual species forms that reproduce by parthenogenesis. An accurate estimate is given of the number and distribution of parthenogenetic lizards in different habitats. In the case of sympatric habitat bisexual and parthenogenetic species of rock lizards population density of parthenogenetic species is much higher. Meanwhile, in different habitats population density of the same-sex species can vary significantly, as determined by the distribution of shelters, shelter and supportive for basking and hunting places.
Галоян Э.А. (2010) -
Род Darevskia – разнообразная многовидовая группа, в которую помимо бисексуальных видов вхо дят однополые формы, размножающиеся путем партеногенеза. Дана точная оценка численности и распределения партеногенетических ящериц в различных местообитаниях. В случае симпатриче ского обитания обоеполого и партеногенетического видов скальных ящериц популяционная плот ность партеногенетического вида значительно выше. Между тем в различных местообитаниях плот ность популяции одного и того же однополого вида может значительно различаться, что определя ется распределением убежищ, укрытий и благоприятных для баскинга и охоты мест.
Galoyan, E.A. (2013) -
Galoyan, E.A. & Tsellarius, E.Y. & Arakelyan, M.S. (2019) -
The parthenogenetic rock lizard Darevskia rostombekowi is considered to be the product of hybridization between female Darevskia raddei and male Darevskia portschinskii. These two species coexist within several secondary contact areas; however, no trace of their hybridization has been previously reported. We conducted focal observations of individually marked lizards in 2017 and 2018 to establish if there is behavioural isolation between these species. We demonstrate that individuals distinguish between lizards of the same and different species. Individuals of both studied species interacted regularly, but we found no evidence of males of either species using interspecific interference competition for resources or for females. Neutral reactions prevailed in the reactions of males to the individuals of the different species; aggressive or submissive behaviour was more common towards males of the same species. The differences in reactions of males to females of the same and different species were less clear and interspecific social interactions were almost as common as intraspecific interactions. Interspecific male mating behaviour was rare and unsuccessful; only a single female with heterospecific copulation marks on the body was found. The mating period in both species overlapped broadly in late May and early June; body size (SVL) in females of both species was equal. Hence, we suggest that the selection of the sexual partner and preference of the conspecific partner for the social and sexual contacts is the most plausible explanation of the revealed behavioural asymmetry in intra- and interspecific relationship.
Garcia-Porta, J. & Irisarri, I. & Kirchner, M. & Rodríguez, A. & Kirchhof, S. & Brown, J.L. & MacLeod, A. & Turner, A.P. & Ahmadzadeh, F. & Albaladejo, G. & Crnobrnja-Isailovic, J. & Riva, I. de la & Fawzi, A. & Galán, P. & Göçmen, B. & Harris, D.J. & Jiménez-Robles, O. & Joger, U. & Jovanović Glavaš, O. & Karış, M. & Koziel, G. & Künzel, S. & Lyra, M. & Miles, D. & Nogales, M. & Oğuz, M.A. & Paf (2019) -
Climatic conditions changing over time and space shape the evolution of organisms at multiple levels, including temperate lizards in the family Lacertidae. Here we reconstruct a dated phylogenetic tree of 262 lacertid species based on a supermatrix relying on novel phylogenomic datasets and fossil calibrations. Diversification of lacertids was accompanied by an increasing disparity among occupied bioclimatic niches, especially in the last 10 Ma, during a period of progressive global cooling. Temperate species also underwent a genome- wide slowdown in molecular substitution rates compared to tropical and desert-adapted lacertids. Evaporative water loss and preferred temperature are correlated with bioclimatic parameters, indicating physiological adaptations to climate. Tropical, but also some popu- lations of cool-adapted species experience maximum temperatures close to their preferred temperatures. We hypothesize these species-specific physiological preferences may con- stitute a handicap to prevail under rapid global warming, and contribute to explaining local lizard extinctions in cool and humid climates.
Ghaemitalab, V.S. & Kami, H.G. (2016) -
Squamata or scaled Reptiles are the largest order of new Reptiles including Lizards, Snakes and Amphisbaenia. They have formed 95 precent of new Reptiles. Our goal is Lizard Biodiversity of Torbat-e Heydaryeh in Razavi Khorasan Province. The study of Lizards Biodiversity of Torbat-e Heydaryeh in Razavi Khorasan Province was carried out in 2012 until 2013. In this research 132 specimens were collected and observed and identified on the basis of morphological, morphometric and meristic characters. The collected and observated Lizards belonging to 5 families: Agamidae, Lacertidae, Scincidae, Gekkonidae and Varanidae including 8 genera and 12 species. Results showed that Agamidae and Lacertidae families have the most frequency. The Agamidae family has the most genera and Lacertidae family has the most species. laudakia caucasia, laudakia microlepis, Trapelus sanguinolentus and Mesalina watsonana were the dominant species in this region and had a wider distribution rang in coparision with other species.
وجیهه السادات قائمیطلب1، حاجی قلی کمی2* (1396) -
Squamata یا خزندگان فلسدار بزرگترین راسته از خزندگان عصر حاضر هستند که دربرگیرنده همه سوسمارها و مارها و سوسمارهای کرممانند هستند و 33 درصد همه خزندگان امروزی را دربرمیگیرند. هدف ما مطالعه تنوع زیستی مارمولکها در منطقه تربتحیدریه در استان خراسان رضوی است. برای مطالعه تنوع زیستی سوسماران شهرستان تربتحیدریه در استان خراسان رضوی نمونهبرداری در بهار 1931 آغاز و تا تابستان 1932 به طول انجامید. در این پژوهش 192 نمونه جمعآوری و مشاهده گردید و بر اساس ویژگیهای مورفولوژیک، مورفومتریک و مریستیک شناسایی گردیدند. نتایج مطالعات نشان داد سوسمارهای جمعآوری و مشاهده شده متعلق به 3 خانواده آگامیده (Agamidae)، گکونیده (Gekkonidae)، سرتیده (Lacertidae)، سینسیده (Scincidae) و وارانیده (Varanidae)، شامل 8 جنس و 12 گونه میباشند. نتایج نشان داده خانواده آگامیده و سرتیده بیشترین فراوانی را دارا بوده و خانواده آگامیده بیشترین تعداد جنس و خانواده سرتیده بیشترین تعداد گونه را در منطقه دارا میباشند که گونههای laudakia Trapelus ،laudakia microlepis ،caucasia sanguinolentus و Mesalina watsonana نمونههای بارز منطقه میباشند و دامنه پراکندگی گستردهای در مقایسه با سایر
Gherghel, J. & Strugariu, A. & Sahlean, T. Stefanescu, A. (2011) -
Ghira, I. (1994) -
Ghira, I. & Venczel, M. & Covaciu-Marcov, S. & Mara, G. & Tiberiu, P.G. (2002) -
Gholamifard, A. (2011) -
Iran contains 36 named endemic reptile species in 17 genera and seven families. The
most endemic and speciose family is the Gekkonidae, followed by the Lacertidae. These
two families collectively dominate with 22 species (or 60.1%) of the total endemic
herpetofauna. Twenty one endemics are known only from a single location or restricted
area within a single physiographic region. An analysis of endemicity is given in terms of
systematics and distribution.
Girnyk, A. & Korchagin, V. & Vergun, A. (2018) -
Girnyk, A. & Omelchenko, A. & Vergun, A. & Ryskov, A. (2016) -
Nowadays approximately 80 taxa of unisexual vertebrates have been described. Among them, reproduction in the complete absence of males is restricted to reptiles, especially among lizards and is referred to as true parthenogenesis. The Lacertidae was the first family in which this phenomenon was discovered among Squamata. The genus Darevskia is of special significance, because these lizards have been the subject of extensive ecological and biogeographical studies, and because parthenogenesis has probably arisen several times within the group. D. armeniaca, one of the seven parthenogenetic species, arose from the interspecific hybridization of D. valentini and D. mixta, and exhibits low mitochondrial DNA and allozyme variability. In this study, we performed microsatellite genotyping of 127 specimens of D. armeniaca from 14 Armenian and one Ukrainian (introduced about 50 years ago from Armenia) populations. Using 4 microsatellite loci, 15 clonal lineages were detected in D. armeniaca. Some of them were found as major clones, distributed in various populations, and other’s represented rare clones, geographically restricted. It was shown that in the introduced Ukrainian population D. armeniaca, new genotypes (probably clones) were detected after the invasion. Some genetic characteristics of the populations and their phylogenetic relationships were also estimated.
Girnyk, A.E. & Vergun, A.A. & Omelchenko, A.V. & Petrosyan, V.G. & Korchagin, V.I. & Ryskov, A.P. (2017) -
A key issue in the study of unisexual (parthenogenetic) vertebrate species is the determination of their genetic and clonal diversity. In pursuing this aim, various markers of nuclear and mitochondrial genomes can be used. The most effective genetic markers include microsatellite DNA, characterized by high variability. The development and characterization of such markers is a necessary step in the genetic studies of parthenogenetic species. In the present study, using locus-specific PCR, for the first time, an analysis of allelic polymorphism of four microsatellite loci is performed in the populations of parthenogenetic species Darevskia armeniaca. In the studied populations, allelic variants of each locus are identified, and the nucleotide sequences of each allele are determined. It is demonstrated that allele differences are associated with the variation in the structure of microsatellite clusters and single nucleotide substitutions at fixed distances in flanking DNA regions. Structural allele variations form haplotype markers that are specific to each allele and are inherited from their parental bisexual species. It is established which of the parental alleles of each locus were inherited by the parthenogenetic species. The characteristics of the distribution and frequency of the alleles of microsatellite loci in the populations of D. armeniaca determining specific features of each population are obtained. The observed heterozygosity of the populations at the studied loci and the mutation rates in genome regions, as well as Nei’s genetic distances between the studied populations, are determined, and the phylogenetic relationships between them are established.
Гирнык А.Е., Вергун А.А., Омельченко А.В., Петросян В.Г., Корчагин В.И., Рысков А.П. (2017) -
Ключевым вопросом в исследовании однополых (партеногенетических) видов позвоночных является определение их генетического и клонального разнообразия. Для этого могут быть использованы различные маркеры ядерного и митохондриального геномов. К наиболее эффективным генетическим маркерам относятся микросателлитные ДНК, обладающие высокой скоростью изменчивости. Разработка и характеристика таких маркеров являются необходимым этапом в генетических исследованиях партеновидов. В настоящей работе с помощью локус-специфической ПЦР впервые проведен анализ аллельного полиморфизма четырех микросателлитных локусов в популяциях пар-теновида Darevskia armeniaca. В изученных популяциях выявлены аллельные варианты каждого локуса и определена нуклеотидная последовательность каждого аллеля. Показано, что отличия аллелей связаны с вариациями в структуре микросателлитных кластеров и с однонуклеотидными заменами на фиксированных расстояниях в прилежащих к кластеру участках ДНК. Структурные вариации аллелей формируют гаплотипические маркеры, специфические для каждого аллеля и наследуемые от родительских двуполых видов. Установлено, какие из родительских аллелей каждого локуса наследуются партеновидом. Получены характеристики по распределению и встречаемости аллелей микросателлитных локусов в популяциях D. armeniaca, определяющие отличительные особенности каждой популяции. Определена наблюдаемая гетерозиготность популяций по изучаемым локусам, рассчитаны скорость мутаций на данных участках генома и генетические расстояния по Нею между исследуемыми популяциями, а также установлены филогенетические связи между ними.
Girnyk, A.E. & Vergun, A.A. & Semyenova, S.K. & Guliaev, A.S. & Arakelyan, M.S. & Danielyan, F.D. & Martirosyan, I.A. & Murphy, R.W. & Ryskov, A.P. (2018) -
Background: The parthenogenetic Caucasian rock lizard Darevskia armeniaca, like most other parthenogenetic vertebrate species, originated through interspecific hybridization between the closely related sexual Darevskia mixta and Darevskia valentini. Darevskia armeniaca was shown to consist of one widespread allozyme clone and a few rare ones, but notwithstanding the origin of clonal diversity remains unclear. We conduct genomic analysis of D. armeniaca and its parental sexual species using microsatellite and SNP markers to identify the origin of parthenogenetic clonal lineages. Results: Four microsatellite-containing loci were genotyped for 111 specimens of D. armeniaca, 17 D. valentini, and four D. mixta. For these species, a total of 47 alleles were isolated and sequenced. Analysis of the data revealed 13 genotypes or presumptive clones in parthenogenetic D. armeniaca, including one widespread clone, two apparently geographically restricted clones, and ten rare clones. Comparisons of genotype-specific markers in D. armeniaca with those of its parental species revealed three founder-events including a common and two rare clones. All other clones appeared to have originated via post-formation microsatellite mutations in the course of evolutionary history of D. armeniaca. Conclusion: Our new approach to microsatellite genotyping reveals allele-specific microsatellite and SNP markers for each locus studied. Interspecies comparison of these markers identifies alleles inherited by parthenospecies from parental species, and provides new information on origin and evolution of clonal diversity in D. armeniaca. SNP analyses reveal at least three interspecific origins of D. armeniaca, and microsatellite mutations in these initial clones give rise to new clones. Thus, we first establish multiple origins of D. armeniaca. Our study identifies the most effective molecular markers for elucidating the origins of clonal diversity in other unisexual species that arose via interspecific hybridization.
Glandt, D. (2010) -
Glandt, D. (2011) -
Glandt, D. & Trapp, B. (2022) -
Göcmen, B. & Oguz, M.A. & Karis, M. & Kösemen, C.M. (2016) -
During fieldwork excursions on June and September 2015 we obtained new distributional records for three lizard species. Here, we present the first record of Darevskia rudis tristis and Lacerta viridis meridionalis from Turkey’s Ankara province and confirm the occurrence of Anguis fragilis in this locality.
Godakova, S.A. & Koechagin, V.I. & Semeyanova, S.K. & Chernyavskaya, M. & Sevast`yanova, G.A. & Ryskov, A.P. (2015) -
Cloning and sequencing of the partial reverse transcriptase gene (750 bp) of the Bov-B LINE retrotransposon have been held in parthenogenetic lizards Darevskia unisexualis and its assumed parental bisexual species D. nairensis and D. valentini. It was shown that the percentage of transcriptionally active copies of this gene, which does not contain a stop codon, is almost the same in the three species and is about 75%. The intragenomic divergence level of these sequences is low and was found to be 2.6% in D. unisexualis, 1.9% in D. nairensis, and 1.6% in D. valentini. The phylogenetic analysis shows the distribution of copies of D. unisexualis in each of the two clusters of RT sequences characteristic of D. nairensis and D. valentini. This result supports the view of the hybrid origin of D. unisexualis and does not exclude intraspecific hybridization between D. nairensis and D. valentini.
Годакова С.А., Корчагин В.И., Семенова С.К., Чернявская М.М., Севастьянова Г.А., Рысков А.П. (2015) -
Клонированы и секвенированы последовательности участка гена ревертазы (750 п.н.) ретротранспозона Bov-B LINE, составляющие около двух третей его длины, у партеногенетического вида ящериц Darevskia unisexualis и двуполых видов D. nairensis и D. valentini. Показано, что доля транскрипционно активных копий этого гена, не содержащих стоп-кодонов, примерно одинакова у трех видов и составляет около 75% от числа исследованных копий. Уровень внутригеномной дивергенции всех копий невысок и составляет 2.6% у D. unisexualis, 1.9% у D. nairensis и 1.6% у D. valentini. На основании филогенетических реконструкций показано распределение копии D. unisexualis внутри каждого из двух кластеров последовательностей гена ревертазы, характерных для D. nairensis и D. valentini. Этот результат поддерживает мнение о гибридном происхождении D. unisexualis и не исключает возможности участия в этом процессе D. nairensis and D. valentini.
Golynsky, E.A. & Doronin, I.V. (2014) -
Gorman, G.C. (1969) -
Karyotype data are presented for 12 species of lacertid lizards. Of these, 4 Acanthodactylus, 2 Eremias, 1 Ophisops, and 4 Lacerta have n = 19, with all chromosomes acrocentric. This is the typical lacertid karyotype, as reported in previous literature. One species, Lacerta parva, is quite different from all the others. The haploid number is 12, consisting of 7 pairs of metacentric macrochromosomes and 5 pairs of microchromosomes. The fundamental number (number of arms in the karyotype) is the same in L. parva as in all the other species studied. The possibility of female sex chromosomal heteromorphism is raised, but data are not sufficient to confirm this.
Grechko, V.V. & Bannikova, A.A. & Kosushkin, S.A. & Ryabinina, N.L. & Milto, K.D. & Darevsky, I.S. & Kramerov, D.A. (2007) -
To characterize the molecular genetic diversity of the genus Darevskia, several populations were examined by the inter-SINE-PCR method, reporting the number and sizes of the spacers between individual copies of SINE-like interspersed repeats. Examination of 17 D. raddei geographical populations and several reference species revealed unequal genetic differences, measured as Nei and Li’s genetic distances (DNL), for different groups of samples. The highest homogeneity was observed for the apparently panmictic D. raddei nairensis population from the basin of the Hrazdan River: genetic differences within each of the five samples and between them were similarly low (less than 0.1). The difference between ten samples of D. raddei raddei from Armenia and Karabakh (0.2–0.3) was somewhat higher than the interindividual difference within each sample (0.1–0.2), indicating that the samples belonged to different populations. The assumption was supported by the phylogenetic tree topology and multidimensional scaling. The differences between samples from the morphological subspecies D. raddei raddei and D. raddei nairensis ranged 0.3–0.4. The difference of two D. raddei raddei samples of Talysh (Azerbaijan) from other samples of the same subspecies corresponded to the subspecific level. The genetic distances between the good species D. raddei and D. rudis was 0.6–0.7. In terms of DNL, a questionable population from northwestern Turkey (“D. tristis”) was closer to D. rudis (DNL = 0.45), probably representing its subspecies. The phylogeography of D. raddei is discussed.
Grechko, V.V. & Ciobanu, D.G. & Darevsky, I.S. & Kosushkin, S.A. & Kramerov, D.A. (2006) -
Satellite DNA repeats were studied in Caucasian populations of 18 rock lizard species of the genus Darevskia. Four subfamilies (Caucasian Lacerta satellites (CLsat)I–IV) were identified, which shared 70%–75% sequence similarity. The distribution of CLsat subfamilies among the species was studied. All the species could be divided into at least 3 clades, depending on the content of CLsat subfamilies in each genome: “saxicola”, “rudis”, and “mixta” lizards. CLsatI was found in all studied species, but in very different quantities; the “saxicola” group contained this subfamily predominantly. The “rudis” group also contained CLsatIII, and the “mixta” group carried considerable amounts of CLsatII. The highest concentrations of CLsatI and CLsatII were detected in 2 ground lizards — D. derjugini and D. praticola, respectively. D. parvula predominantly carried CLsatIII. CLsatIV was found only in the Crimean species D. lindholmi. The distribution patterns of satellite subfamilies show possible postglacial speciation within the genus Darevskia. A hybrid origin of species that possess 2 or 3 CLsat subfamilies and important clarifications to the systematics of the genus are proposed.
Grechko, V.V. & Darevsky, I.S. (2007) -
Grechko, V.V. & Kataev, M.V. & Melnikova, M.N. & Darevsky, I.S. (1993) -
Due to some suggestion  the parthenogenesis in vertebrates is the result of interspecies hybridization, and some species seem to be the main candidates participating in this process. DNA parts, which may be revealed using polymerase chain reaction in AP-RAPD modification, were compared in parthenogenetic lizards forms and in supposed parental species for them. As it was shown by other authors and confirmed by us, arbitrary primed randomly amplified DNA products are species specific and may be used as a molecular taxonomic criterion. It was shown that each parthenogenetic species studied has the patterns containing only the bands of one or another supposed parents and no more. This is in favour of the hypothesis mentioned above. Investigation of 103 10-nucleotide primers in PCR was done.
Grechko, V.V. & Ryabinin, D.M. & Fedorova, L.V. & Fedorov, A.N. & Ryskov, A.P. & Darevsky, I.S. (1997) -
The Caucasian parthenogenetic rock lizards of the L. saxicola complex probably originated by hybridisation of some bisexual species. We verified this hypothesis using a new approach based on comparison of repetitive DNA characters, which produce species-specific patterns (named `taxonprint`). The method relies on restriction endonuclease hydrolysis of genomic DNA, with the following 32P-end labeling and polyacrylamide gel electrophoretic separation of the fragments. The parthenogenetic species L. armeniaca and L. dahli possess specific features both of putative maternal L. mixta (in some taxonprints), and of paternal L. valentini (in other ones), whereas L. portschinskii and L. rudis also could be the paternal species. Parthenoclones L. unisexualis and L. uzzelli have the specific DNA features in several taxonprints of L. raddei or L. nairensis (which cannot be discriminated by taxonprints) which are supposed to be maternal species for L. unisexualis and paternal ones for L. uzzelli. The specific features of L. valentini (or L. portschinskii) are observed in L. unisexualis; L. uzzelli has not been investigated in this respect. Parthenogenetic L. rostombekovi also possesses properties of L. raddei or L. nairensis, but we have not be able to find any features of supposed paternal L. portschinskii in this parthenogenetic species.
Grechko, V.V. et al. (1997) -
Multiple band patterns of DNA repeats in
the 20–500-nucleotide range can be detected by digesting
genomic DNA with short—cutting restriction endonucleases,
followed by end labeling of the restriction
fragments and fractionation in nondenaturing polyacrylamide
gels. We call such band patterns obtained from
genomic DNA ‘‘taxonprints’’ (Fedorov et al. 1992).
Here we show that taxonprints for the taxonomic groups
studied (mammals, reptiles, fish, insects—altogether
more than 50 species) have the following properties: (1)
All individuals from the same species have identical taxonprints.
(2) Taxonprint bands can be subdivided into
those specific for a single species and those specific for
groups of closely related species, genera, and even families.
(3) Each restriction endonuclease produces unique
band patterns; thus, five to ten restriction enzymes (about
100 bands) may be sufficient for a statistical treatment of
phylogenetic relationships based on polymorphisms of
restriction endinuclease sites. We demonstrate that taxonprint
analysis allows one to distinguish closely related
species and to establish the degree of similarity among
species and among genera. These characteristics make to use this phenomenon for phylogenetic aims (Cooke
1975; Christie and Skinner 1979; Brown and Dover
1979, 1980; Elizur et al. 1982; Hembelen et al. 1982;
Buckland 1983; Dandieu et al. 1984; Lima de Faria et al.
1984; Shubina and Mednikov 1986; Turner et al. 1991),
but this approach has not been extensively applied. In
this paper we have reexamined the problem on the basis
of our own preliminary results, partly published in Russian
journals, confirming the ideas about possible correlations
between evolution of DNA repeats and speciation
(Fedorov et al. 1992; Grechko et al. 1993; Potapov and
We have demonstrated that the modified method designated
‘‘DNA taxonprint’’ (Fedorov et al. 1992) can be
effectively used for revealing species—specific genomic
markers in different taxonomic groups. These taxonprint
markers are found to be helpful for evaluation of the
degree of species similarity between species, genera, and
families. More than 50 species from different taxa were
Gül, S. & Özdemir, N. & Kumlutaş, Y. & Durmus, S.H. & Ilgaz, C. (2015) -
We present the first data of body size and age structure (based on the skeletochronological examination) of Darevskia bithynica from two localities characterised with different altitudes, i.e. Zonguldak (50 m) and Bolu (1200 m). Males from both lowland and highland populations were slightly larger than females. This difference was not significant for the lowland population. The maximum lifespan was found to be six years for both populations. Intersexual differences in snout-vent length were male-biased and this may be linked to the climatic conditions.
Gül, S. & Özdemir, N. & Kumlutaş, Y. & Ilgaz, C. (2014) -
Darevskia rudis (BEDRIAGA, 1886) zeigt im Gebiet der türkischen Schwarzmeerregion eine weite vertikale Vebreitung. Insgesamt wurden 62 Individuen dieser Eidechse von drei Populationen/Fundorten aus unterschiedlichen Höhenlagen (Provinz Artvin: Ardanuç, 2137 m ü.M., Borçka, 1277 m ü.M., Çermik, 700 m ü.M.) skeletochronologisch altersbestimmt und hinsichtlich ihrer Kopf-Rumpf-Länge vermessen. Im Mittel waren Kopf-Rumpf-Länge und Alter in beiden Geschlechtern bei der höchst gelegenen Population geringer als bei den beiden tiefer gelegenene Populationen. Ein signifikant positiver Zusammenhang fand sich zwischen Alter und Kopf-Rumpf-Länge bei den Weibchen der beiden hoch gelegenen Populationen, jedoch nicht für die aus Çermik und die Männchen aller drei Populationen. Der Geschlechtsdimorphismus in der Körpergröße war in der tiefstgelegenenen (Çermik) Population durch größere Männchen und in den höher gelegenen Popultionen (Borçka und Ardanuç) durch größere Weibchen gekennzeichnet. Die skeletochronologischen Untersuchungen zeigten deutliche Unterschiede in der Körpergröße und Alterstruktur zwischen tiefer und höher gelegenen Populationen von D. rudis auf.
Haklová-Kociková, B. & Hiznanová, A. & Majláth, I. & Racka, K. & Földvárui, G. & Tryjanowski, P. & Kokosová, N. & Malceková, B. & Majláthová, V. (2014) -
Background: Blood parasites of the genus Karyolysus Labbé, 1894 (Apicomplexa: Adeleida: Karyolysidae) represent the protozoan haemogregarines found in various genera of lizards, including Lacerta, Podarcis, Darevskia (Lacertidae) and Mabouia (Scincidae). The vectors of parasites are gamasid mites from the genus Ophionyssus. Methods: A total of 557 individuals of lacertid lizards were captured in four different localities in Europe (Hungary, Poland, Romania and Slovakia) and blood was collected. Samples were examined using both microscopic and molecular methods, and phylogenetic relationships of all isolates of Karyolysus sp. were assessed for the first time. Karyolysus sp. 18S rRNA isolates were evaluated using Bayesian and Maximum Likelihood analyses. Results: A total of 520 blood smears were examined microscopically and unicellular protozoan parasites were found in 116 samples (22.3% prevalence). The presence of two Karyolysus species, K. latus and K. lacazei was identified. In total, of 210 samples tested by polymerase chain reaction (PCR), the presence of parasites was observed in 64 individuals (prevalence 30.5%). Results of phylogenetic analyses revealed the existence of four haplotypes, all part of the same lineage, with other parasites identified as belonging to the genus Hepatozoon. Conclusions: Classification of these parasites using current taxonomy is complex - they were identified in both mites and ticks that typically are considered to host Karyolysus and Hepatozoon respectively. Furthermore although distortions to the intermediate host erythrocyte nuclei were observed, the defining characteristic of Karyolysus, the haplotypes were nearly identical to those reported from lizards in the Iberian Peninsula, where such distortions were not reported and which were thus identified as Hepatozoon. Based on the phylogenetic analyses, neither vertebrate host, nor geographical patterns of the studied blood parasites could be established.
Harris, D.J. & Arnold, E.N. & Thomas, R.H. (1998) -
DNA sequences from parts of the 12S, 16S and cytochrome b mitochondrial genes, which totalled 1049 aligned base pairs, were used to estimate the relationships of 49 species of Lacertidae, including representatives of 19 out of the 23 recognized genera and 23 species of the paraphyletic genus Lacerta. These data were used, together with morphological information, to estimate the relationships within the family. Molecular evidence corroborates the monophyletic status of many genera and species groups originally based on morphology. It indicates that Psammodromus forms a clade with Gallotia, which is the sister taxon of all other lacertids. These comprise three units: the primarily Afrotropical armatured group; the largely Oriental Takydromus; and the west Palaearctic Lacerta and its derivatives, Podarcis and Algyroides. Morphology also supports the first three assemblages, but suggests that they are derived from a paraphyletic Lacerta. Within Lacerta and its allies, DNA sequence analysis corroborates the affinity of some members of each of the subgenera Lacerta s. str. and Timon, and of the L. saxicola group. It also supports the relationship of L. monticola, L. bonnali and L. horvathi, and suggests that the L. parva–L. fraasi clade and L. brandti are not related to Psammodromus–Gallotia, as morphology indicates, but instead are associated respectively with the L. danfordi and L. saxicola groups. DNA sequence data provide additional evidence that the eastern Arabian `Lacerta` jayakari and `L.` cyanura are members of the armatured clade and also sister species. Our analysis supports an origin for present lacertids in west Eurasia. The armatured clade invaded Africa, probably in the mid-Miocene, spreading widely and evolving increasingly xeric-adapted forms, one lineage of which later moved back into the Palaearctic. `Lacerta` jayakari and `L.` cyanura are assigned to Omanosaura, Lutz and Mayer 1986. The name Gallotiinae Cano, Baez, Lopez-Jurado and Ortega, 1984 is available for the Gallotia–Psammodromus clade, Eremiainae Shcherbak 1975 for the armatured clade and Lacertinae for Lacerta, Podarcis and Algyroides. Two new subgenera of Lacerta are proposed here: Caucasilacerta for L. saxicola and its allies, and Parvilacerta for L. parva and L. fraasi.
Harris, D.J. & Carretero, M.A. (2003) -
Harutyanyan, T.K. & Danielyan, F.D. & Arakelyan, M.S. (2015) -
Harutyunyan, T. & Arakelyan, M. & Danielyan F. (2010) -
Harutyunyan, T. & Petrosyan, R.K. & Danielyan, F.D. & Arakelyan, M.S. (2013) -
Հարությունյան, Թ.Կ. & Պետրոսյան, Ռ.Կ. & Դանիելյան, Ֆ.Դ. & Առաքելյան, Մ.Ս. (2013) -
Համեմատվել են Երևանի Հրազդանի կիրճի Darevskia nairensis և Գեղարդի տարածքի D. raddei-ի ժայռային մողեսների պոպուլյացիաները: Նկարագրվել է արյան մակաբույծերով և էկտոմակաբույծներով վարակվածությունը` ելնելով սեզոնային և տարածքային առանձնահատկություններից: Сравнивались популяции скальных ящериц Darevskia nairensis из территории Еревана
(ущелье Раздан) и D. raddei из территории Гегарда. Описaнa зараженность экто- и кровепаразитaми в зависимости от сезонных и региональных особенностей.
Harutyunyan, T.K. (2011) -
The infections by blood parasites in sympatric populations of rock lizards
D. raddei and D. portchinskii were studied. 121 adult individuals have been observed.
Infection by blood parasites depends on sex of host. D. raddei species is more infected
than D. portchinskii.
Հարությունյան, Թ.Կ. (2011) -
Ուսումնասիրվել է արյան մակաբույծներով վարակվածությունը Darevskia raddei եւ D. portchinskii ժայռային մողեսների սիմպատրիկ պոպուլյացիաներում` 121 սեռա- հասուն առանձնյակների վրա: Արյան մակաբույծներով վարակունակությունը կարող է կախված լինել տեր-օրգանիզմի սեռից: Մի տեսակի մոտ շատ ավելի վարակված են արուները, մյուսի մոտ՝ էգերը: D. raddei-ի վարակվածության աստիճանն ավելի բարձր է, քան D. portchinskii մոտ: Исследовалась на зараженность кровепаразитами 121 половозрелой особи в симпатрических популяциях скальных ящериц D. raddei и D. portchinskii. Зараженность кровепаразитами может зависеть от пола хозяина. У одного вида больше заражены самцы, у других – самки. У D. raddei уровень зараженности выше, чем у D. portchinskii.
Hellmich, W. (1969) -
Helmer, W. & Strijbosch, H. & Scholte, P. (1988) -
Hemmerling, J. & Meusel, W. & Obst, F.J. (1974) -
This publication on the herpetofauna oft he Caucasus is a compilation oft wo articles published in 1967 and 1970 (Zool. Abh. SStaatl. Mus. Tierk. Dresden 29, pp. 85-93 and Aquarien terrarien 17, pp. 120-123.
Henle, K. (1983) -
A remarkable hindleg regeneration was observed in a Sharp-snouted Rock Lizard (La- certa oxycephala). The hindleg was replaced by a scaled elongate and conical appendage. The moving lizard kept the regenerate motionless. The specimen apparently was able to catch enough food in nature. Comparable regenerations were found in Caucasian Rock Lizards, in two L. saxicola brauneri and in one L. raddei raddei. Always a hindleg was rege- nerated.
Hipsley, C. (2012) -
The evolutionary diversification of many terrestrial vertebrate groups is strongly linked to climatic events in the Cenozoic, the period from 65 Million years ago to today when modern animals first appeared. I investigated the effects of Cenozoic climate change on the taxonomic and morphological diversification of the Old World lizard family Lacertidae, with particular emphasis on the African radiation. African lacertids exhibit an unusual pattern of diversification, in which their highest species richness occurs in deserts north and south of the equator, despite being spread throughout the continent. This disparity is particularly surprising given that desert lacertids are thought to be evolutionarily younger than their mesic-dwelling relatives, suggesting increased diversification rates in arid habitats. To identify the evolutionary factors underlying this pattern, I use a combination of phylogenetic, morphological and ecological techniques. In Chapter 1, I apply Bayesian methods and fossil-based calibrations to molecular sequence data to construct a time-calibrated phylogeny for Lacertidae. I estimate that the family arose in the early Cenozoic, with the majority of their African radiation occurring in the Eocene and Oligocene. In Chapter 2, I describe changes in lacertid body shape across biomes and substrates, and find widespread morphological convergence in similar habitat types. I suggest that in addition to foraging demands, fluctuating and extreme climatic conditions, largely driven by precipitation and temperature, contribute to morphological convergence across independent arid-dwelling clades. Finally, I test if ancestral transitions in ecology, morphology, and rates of diversification temporally coincide with paleoclimatic events in the Cenozoic. I use High Resolution X-ray Computed Tomography to characterize changes in the skull related to life in arid habitats, and apply maximum likelihood methods to test if the origins of those traits temporally coincide with significant shifts in habitat, diversification rates and climatic changes. My results show that African lacertids experienced three major peaks in diversification, accompanied by the evolution of suites of arid-adapted morphological traits. These changes coincide with climatic shifts in Africa, including the transition from closed forests to open grasslands and savanna in the late Oligocene, prior to the peak temperatures of the mid-Miocene Climatic Optimum, and following the formation of the Benguela current leading to hyper-aridity in southern Africa. I conclude that deserts are important centers for reptile evolution, but that expected changes in climate due to global warming may outpace the ability of arid-dwelling species to adapt and persist in the future.
Hipsley, C. & Himmelmann L. & Metzler D. & Müller J. (2009) -
Background: Although current molecular clock methods offer greater flexibility in modelling
evolutionary events, calibration of the clock with dates from the fossil record is still problematic
for many groups. Here we implement several new approaches in molecular dating to estimate the
evolutionary ages of Lacertidae, an Old World family of lizards with a poor fossil record and
uncertain phylogeny. Four different models of rate variation are tested in a new program for
Bayesian phylogenetic analysis called TreeTime, based on a combination of mitochondrial and
nuclear gene sequences. We incorporate paleontological uncertainty into divergence estimates by
expressing multiple calibration dates as a range of probabilistic distributions. We also test the
reliability of our proposed calibrations by exploring effects of individual priors on posterior
Results: According to the most reliable model, as indicated by Bayes factor comparison, modern
lacertids arose shortly after the K/T transition and entered Africa about 45 million years ago, with
the majority of their African radiation occurring in the Eocene and Oligocene. Our findings indicate
much earlier origins for these clades than previously reported, and we discuss our results in light
of paleogeographic trends during the Cenozoic.
Conclusion: This study represents the first attempt to estimate evolutionary ages of a specific
group of reptiles exhibiting uncertain phylogenetic relationships, molecular rate variation and a
poor fossil record. Our results emphasize the sensitivity of molecular divergence dates to fossil
calibrations, and support the use of combined molecular data sets and multiple, well-spaced dates
from the fossil record as minimum node constraints. The bioinformatics program used here,
TreeTime, is publicly available, and we recommend its use for molecular dating of taxa faced with
Hipsley, C.A. & Miles, D.B. & Müller, J. (2014) -
While global variation in taxonomic diversity is strongly linked to latitude, the extent to which morphological disparity follows geographical gradients is less well known. We estimated patterns of lineage diversification, morphological disparity and rates of phenotypic evolution in the Old World lizard family Lacertidae, which displays a nearly inverse latitudinal diversity gradient with decreasing species richness towards the tropics. We found that lacertids exhibit relatively constant rates of lineage accumulation over time, although the majority of morphological variation appears to have originated during recent divergence events, resulting in increased partitioning of disparity within subclades. Among subclades, tropical arboreal taxa exhibited the fastest rates of shape change while temperate European taxa were the slowest, resulting in an inverse relationship between latitudinal diversity and rates of phenotypic evolution. This pattern demonstrates a compelling counterexample to the ecological opportunity theory of diversification, suggesting an uncoupling of the processes generating species diversity and morphological differentiation across spatial scales.
Höricht, W. (1927) -
Hosseinian Yousefkhani, S (2022) -
Hosseinian Yousefkhani, S.S. & Rastegar-Pouyani, E. & Rastegar-Pouyani, N. & Arab, M. (2012) -
During fieldwork on 15 August 2011 in Golestan province, several specimens of Darevskia steineri (EISELT, 1995) were found in the vicinity of Loveh Waterfall at midday. The locality of D. steineri is limited around the Loveh waterfall. In this survey, no specimen of D. steineri was observed in other parts near to the waterfall, but some reptiles such as Darevskia chlorogaster and Gloydius halys were found as sympatric with D. steineri. The unique habitat of D. steineri is now in the risk of destruction due to human activity and tourism affects as well as the constructing of a gas pipe which has traversed the habitat to the west into Semnan province.
Hosseinian Yousefkhani, S.S. & Rastegar-Pouyani, N. & Rastegar-Pouyani, E. & Arab, M. (2013) -
During fieldwork on 15 August 2011 in Golestan province, several specimens of Darevskia steineri (Eiselt, 1995) were found at midday in the vicinity of the Loveh Waterfall. The locality of Darevskia steineri is limited to the area around the Loveh waterfall. During this survey, no specimens of D. steineri could be found in other parts near to the waterfall, but some reptiles such as Darevskia chlorogaster and Gloydius halys were found as sympatric with D. steineri. The unique habitat of D. steineri is now at the risk of destruction due to human activities as well as tourism effects. In addition, the habitat is affected by a gas pipeline that runs through the area west to the province of Semnan.
Iftime, A. & Iftime, O. (2006) -
The preliminary results of herpetological studies in the Cozia massif in Vâlcea county,
Romania, are presented; the 10 reptile species identified in the field (4 of which are new records) are
presented together with data on their habitat association, abundance and impact of human activity.
Iftime, A. & Iftime, O. (2019) -
The results of faunistic surveys upon the amphibian and reptile populations in the Cozia National Park are presented. We recorded (found) 11 amphibian species (including the hybridogenetic Pelophylax kl. esculentus) and 12 reptile species; some species (Bufotes viridis and Emys orbicularis) were found after a long lapse in scientific recording. We observe that the Cozia Massif appears to have a richer herpeto fauna than that of the adjoining Narâțu and Căpățânii massifs.
Ilgaz, Ç & Akkaya, A. & Hürriyet, H. (2019) -
ased on two specimens collected from Çatalca, İstanbul, Turkey we present a new distribution record for Darevskia pontica in Turkey. This find composes the south-easternmost record for the species and extends its range about 80 km. Data based on meristic pholidosis characters, metric measurements and color-patter features of the specimens were given in detail and it was observed that Çatalca population is similar to D. pontica reported in the literature.
Ilgaz, C. (2006) -
Forty-seven Darevskia armeniacaspecimens collected from 2 different localities in Ardahan were examined in terms of
pholidosis characters, morphometric measurements and colour-pattern features. The 2 populations were differentiated from one
another in terms of the number of supraciliar plates (left-right) (SCPa-SCPb), supralabial plates (left-right) (SRLa-SRLb) and
transversal series of ventral plates (TVP) according to the results of the Mann-Whitney U test. An independent t-test based on
morphometric measurements revealed significant differences in terms of the head index (HI) between the 2 populations. Regarding
pholidosis characters, morphometric measurements and colour-pattern features, specimens collected from the 2 localities showed a
similarity to Darevskia armeniacaspecimens from Georgia, Armenia, Azerbaijan and Zigana.
Ilgaz, C. (2007) -
In all, 103 specimens of Clark’s lizard, Darevskia clarkorum(Darevsky and Vedmederja, 1977), from 6 localities in the
eastern Black Sea region of Turkey were examined in terms of pholidosis characteristics, morphometric measurements, and color
and pattern features. Kolmogorov-Smirnov testing across all populations verified sexual dimorphism in the Ortacalar and Hemﬂin
populations, regarding transversal series of ventral plates (P < 0.05). According to the Mann-Whitney U test, 16 pholidosis
characteristics showed differences among the populations (P < 0.05). For comparison of morphometric measurements, 4
morphometric indices and ratios were developed. According to the results of independent t-testing, no differences were found
between males and females among the populations. One-way variance analyses based on 4 morphometric indices and ratios
confirmed that head index, snout-vent length/tail length, and tail length/total body length showed differences among the populations
(P < 0.05). Regarding pholidosis characteristics (except rates of the presence of a masseteric plate and contact between the rostral
and internasal plate), morphometric measurements, and color and pattern features, the specimens collected from 6 different
localities were similar to D. clarkorum specimens mentioned in the literature. Furthermore, the known range of the species in Turkey
has been extended.
Ilgaz, C. (2009) -
This study was designed to determine the degree of morphological differentiation
between the subspecies of Darevskia parvula from northwestern Anatolia. The meristic
pholidolial characters, metric dimensions and color-pattern features of specimens of both
subspecies obtained from 10 different localities in the northeastern Anatolia, Turkey are given in
detail and compared with regard to literature. In addition, thirty-six morphological characters
were compared statistically and 66.7% of them showed differences between two subspecies.
Ilgaz, Ç. & Kumlutaç, Y. (2005) -
Ineich, I. & Doronin, I. (2017) -
Ineich, I. & Doronin, I. & Lescure, J. (2017) -
The life and work of the Alsatian Louis Amédée Lantz (1886-1953), a pioneer of European herpetology. A trained and professional research-chemist, the Alsatian Louis Amédée Lantz was an active and particularly dynamic herpetologist. His residence of nearly twelve years in Russia enabled him to make several field trips to regions such as the Caucasus and Iran. He always carried with him a home made telescopic net that he used to catch lizards. That net became symbolic of Lantz for most people around him. He realized significant herpetological collections mainly deposited in the Museum of the Zoology Institute of the Russian Academy of Sciences in St. Petersburg. Soon after the Russian Revolution in 1917, he had to leave the country and stayed for a few years in France before joining England. During this period, Lantz visited the French Pyrenees and the Mediterranean border (Hyeres Islands, Bandol), enabling him to discover for the first time oviparous populations of common lizard and also to describe a new species of lizard from the French Pyrenees. Lantz is also at the origin of the discovery of unisexual reproduction by parthenogenesis in reptiles. His collaboration with the Swedish amateur herpetologist Otto Cyrén, whom he met through the German herpetologist Willy Wolterstorff, was particularly successful throughout his career. Lantz died at the age of 66 years, in Switzerland, as he was about to join the team of herpetologists at the Natural History Museum in Paris.
Ineich, I. & Doronin, I.V. & Cheylan, M. & Campbell, P.D. (2019) -
Several recent papers have reviewed the life and work of French herpetologist Louis Amédée Lantz. They have detailed the composition of his collections deposited in several museums. However, since then, several other important specimens from his collections deposited at the Natural History Museum (NHM, UK) have come to light and we here present all of them in detail. We discovered paralectotypes of Lacerta saxicola obscura Lantz & Cyrén (BMNH 19188.8.131.52–7), syntypes of Lacerta boemica Sukhov (BMNH 19184.108.40.206–30, BMNH 1965.337–342) and Lacerta viridis media Lantz & Cyrén (BMNH 19220.127.116.11, 1966.512). We also identified numerous specimens from the French Mediterranean islands in Lantz’s collection deposited at the NHM, some of which represent the first reported specimens of their species from certain islands. We here provide data on all these specimens. We also place the Mediterranean island specimens from the Lantz collection in their historical context and emphasize the historical and taxonomic value of these collections.
International Commission on Zoological Nomenclature (2020) -
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.
Kafash, A. & Ashrafi, S. & Ohler, A. (2018) -
Little is known about altitudinal distribution of lizards in Iran. In the present study we studied distribution pattern of members of family Lacertidae along the eleveatinal gradient in Iran. To determine environmental drivers of distribution pattern of 48 lizard species, all known valid members of family Lacertidae, along elevational gradient in Iran. Firstly, we classified Iran digital elevation model (DEM) based on 100m intervals, that resulted in 56 altitudinal bands and number of species in each band was recorded. Secondly, we extracted mean value of following variables; altitude, slop, area, solar radiation index (SRI), normalized differences vegetation index (NDVI), annual precipitation, precipitation of wettest month, precipitation of driest month precipitation, precipitation of wettest quarter, precipitation of driest quarter, and precipitation of warmest quarter, and using VIF measure correlated variables were removed. Finally, we performed a multiple regression and found that area and precipitation of warmest quarter are the most important drivers of distribution pattern of family Lacertidae along elevational gradient in Iran. General distribution pattern of family Lacertidae was unimodal and maximum number of species living from 1475 m to 1675 m.
انوشه کفاش؛ سهراب اشرفی ؛ آنماری اوهلر (1397) -
اطلاعات اندکی در ارتباط با توزیع ارتفاعی سوسماران در ایران در دسترس است. در حالی که تعیین مناطق ارتفاعی با بیشترین تعداد گونه میتواند برای انتخاب مناطق حفاظت شده مفید باشد. در مطالعه حاضر الگوی توزیع ارتفاعی 48 گونه از سوسماران ایران متعلق به خانواده لاسرتیده در امتداد شیبهای ارتفاعی مورد بررسی قرار گرفت. به منظور تعیین الگوی توزیع ارتفاعی اعضای خانواده لاسرتیده مدل رقومی ارتفاع کشور به طبقات ارتفاعی صد متری تقسیم شد و تعداد گونه در هر طبقه ارتفاعی ثبت شد. ارزش عددی متغیرهای شیب، مساحت، ارتفاع، شاخص تشعشع حرارتی، شاخص نرمال شده پوشش گیاهی، بارش سالیانه، بارش در مرطوب ترین ماه سال، بارش در خشک ترین ماه سال، بارش در مرطوب ترین فصل سال، بارش در سردترین فصل سال، بارش در خشک ترین فصل سال و بارش در گرم ترین فصل سال در هر طبقه ارتفاعی از طریق جعبه ابزار Raster استخراج شده و بعد از بررسی میزان همبستگی بین این متغییرها با استفاده از معیار VIF متغیرهای با همبستگی بالا حذف شدند. سپس با اجرای آنالیز رگرسیون چندگانه مهمترین متغییرهای موثر بر توزیع ارتفاعی گونه های خانواده لاسرتیده دو متغییر مساحت و بارش در گرمترین فصل سال شناسایی شد. الگوی کلی توزیع سوسماران خانواده لاسرتیده در امتداد شیب های ارتفاعی به صورت تک اوجی بوده و بیشترین تعداد گونه ها در ارتفاع 1475 تا 1675حضور دارند.
Kan, N.G. & MartyrosyanI, I.A. & Darevsky, I.S. & Danielyan, F.D. & Arakelyan, M.S. & Aslanyan, A.V. & Grechko, V.V. & Tokarskaya, O.N. & Ryskov, A.P. (2000) -
Two parthenogenic families of unisexual species of Caucasian rock lizards of genus Lacerta, L. armeniaca andL. unisexualis, were analyzed by DNA fingerprinting. Inheritance of M13 minisatellite and of (GACA) n , (GATA) n , and (TCC) n microsatellite loci in the first generation of the lizards was studied. M13, (GACA) n , and (TCC) n loci in the families ofL. armeniaca were strictly inherited, as well as M13 and (GACA) n loci in the families ofL. unisexualis: each DNA fragment in the fingerprint patterns of progeny could be detected in the maternal pattern. However, when a (TCC)50 microsatellite probe was applied in the study ofL. unisexualis families, specific DNA fragments with altered mobility were revealed in the progeny patterns, and the frequency of such events was rather high. It might be hypothesized that some of the (TCC) n loci inL. unisexualis genome are highly mutable. Hence, the family analysis allowed us to demonstrate experimentally the presence of genetically unstable loci in genomes of parthenogenic species of vertebrates. The nature and mechanism of the instability of these loci in parthenogenesis remain obscure.
Н.Г. Кан, И.А. Мартиросян, И.С. Даревский, Ф.Д. Даниелян, М.С. Аракелян, А.В. Асланян, В.В. Гречко, О.Н. Токарская, А.П. Рысков (2000) -
Проведен ДНК-фингерпринтный анализ партеногенетических семей двух однополых видов кавказских скальных ящериц рода Lacerta-L. armeniaca и L. unisexualis. Изучено наследование М13-минисателлитных и (GACA)n-, (GATA)n-, (ТСС)п-микросателлитных локусов в первом поколении партеногенетических ящериц исследуемых видов. Показано стабильное наследование М13-, (GАСА)n- и (ТСС)п-локусов в семьях L. armeniaca, а также М13- и (ОАТА)п-локусов в семьях L. unisexualis - любой фрагмент ДНК в ДНК-фингерпринтном паттерне потомка обнаруживается в материнском паттерне. В то же время, при использовании (ТСС)5о-микросателлитной пробы в семьях L. unisexualis у потомков обнаружены измененные по подвижности фрагменты ДНК, не присутствующие в материнском паттерне, причем частота появления таких фрагментов достаточно высока. Предполагается, что некоторые (ТСС)n-локусы в геноме L. unisexualis обладают повышенной мутационной способностью. Таким образом, в настоящей работе при семейном анализе впервые выявлены генетически нестабильные локусы у партеногенетических видов позвоночных. Природа и механизм нестабильности этих локусов при партеногенетическом способе размножения остаются неясными.
Kapli, P. & Poulakakis, N. & Lymberakis, P. & Mylonas, M. (2011) -
the Lacertidae is one of the most diverse and widespread lizard families throughout Eurasia and Africa. Several studies so far have attempted to unravel the phylogeny of Lacertidae using morphological and mole- cular data. However, the intra-family relationships remain unclear. in an effort to explore the phylogenetic relationships within the family Lacertidae, a concatenated dataset of 5727 bp from six genes (two nuclear and four mitochondrial) and 40 genera was assembled based on GenBank database. Phylogenetic inference analy- ses were conducted using Maximum Parsimony (MP), Bayesian inference (Bi) and Maximum Likelihood (ML), revealing that even a combined dataset of both mitochondrial and nuclear genes is not able to resolve the phylogenetic relationships of the Lacertidae family under the tribe level.
Karaman, S. (1939) -
Karmishev, Y. & Yarigin, O. (2013) -
This work presents data on reproductive features of Ukrainian fauna Lacertidae lizards in laboratory conditions and comparative data on lizards clutches in wild nature. Terms of egg incubation of these lizards in the laboratory in a stable, relatively low temperatures can significantly exceed those in nature, which allows to obtain embryonic material with a smooth, long process of development. The most prolific representatives of different major types - green lizard (Lacerta viridis) and nimble (Lacerta agilis). Females sand lizard can be aggressive towards other females clutches, which is manifested in the destruction of nests.
Karyeva, O. & Shammakov, S. & Iazgulyev, A. & Atamuradov, K.I. (1999) -
Kaska, Y. & Kumlutaş, Y. & Avci, A. & Üzüm, N. & Yeniyurt, C. & Akarsu, F. (2009) -
Kaska, Y. & Kumlutaş, Y. & Kaya, U. & Avci, A. & Üzüm, N. & Yeniyurt, C. & Akarsu, F. (2009) -
Kattke, P. (2014) -
Kaya, N. & Özuluğ, O. (2017) -
Amphibia and Reptilia collections at the Zoological Museum, University of Istanbul (ZMUI) were studied and revised. The samples were collected from 1921 until today. The localities of most species are found in the different regions of Turkey. The collection have 75 species of which 21 species are amphibians and 54 species are reptiles. With this study a zoological collection belong to ZMUI was given for the first time.
İstanbul Üniversitesi Zooloji Müzesi (ZMUI) Amphibia ve Reptilia koleksiyonu çalışıldı ve revize edildi. Örnekler 1921 yılından günümüze kadar toplanmıştır. Çoğu türün lokalitesi Türkiye’nin farklı bölgelerindendir. Koleksiyonda 75 tür bulunmakta olup bunların 21 türü amfibi ve 54 türü sürüngenlerdendir. Bu çalışma ile ilk defa ZMUI’a ait bir zoolojik koleksiyon verilmiştir.
Kessler, K.F. (1878) -
Khan, N.G. & Petrosyan, V.G. & Martirosyan, I.A. & Ryskov, A.P. & Darevsky, I.S. & Danielyan, F.D. & Ryabinin, D.M. & Grechko, V.V. & Tolkarskaya, O.N. (1998) -
The variability of mini-and microsatellite DNAs among the individuals of the parthenogenic Caucasian rock lizard, Lacerta dahli, from populations of Armenia was studied by multiloci DNA fingerprinting. The DNA fingerprints of all specimens of the parthenospecies appeared to be practically identical (species-specific), whereas those of the biparental species, L. raddei, proved to be individual-specific. Some of the mini-and microsatellite loci of parthenogenic animals revealed genetic heterogeneity. Quantitative estimates of the genetic variability of mini-and microsatellite DNA fragments, such as the mean index of similarity (S) and heterozygosity (H), for parthenogenic lizards were 0.967 and 0.025, respectively, whereas in bisexual species they were similar to those observed in other outbreeding populations (S = 0.38 and H = 0.695). Computer analysis of 1561 marker fragments of parthenogenic lizard DNAs discerned 12 DNA fingerprinting variants differing in 1-14 fragments. The data on the genetic variability of mini-and microsattelite loci of L. dahli may illustrate the process of spontaneous mutagenesis and the initial stage of intraspecies differentiation in natural populations of parthenogenic species.
Khonyakina, Z.P. (1964) -
Kidov A.A. (2018) -
the purpose of the research: to identify features of the parasite-host relations of the European common tick, Ixodes ricinus (Linnaeus, 1758)
and Hyrcanian meadow lizard, Darevskia praticola hyrcanica (Tuniyev, Doronin, Kidov, et Tuniyev), 2011 in spring period.
materials and methods. The study was carried out in May 2016 in the Gadazyghahi natural boundaries in Astara region of Azerbaijan. The animals were caught on the routes, divided into three mature groups (adult females, males and juvenile lizards) and examined for the presence of ticks. Frequency of occurrence, abundance and location of parasites were determined according to standard methods. Localization of parasites on lizards’ body was defined by the standard scheme.
results and discussion. Only larvae and nymphs of the I. ricinus were found on the lizards. Ticks were found only on 32 lizards of 123 examined. The distribution of ticks in year-old lizards was 10,8%, adult females 28,6%, and adult males 35,3%. The distribution of feeding ticks is aggregated: 26 % of examined lizards were feeders for 100% of ticks. Higher abundance of parasites per one animal was observed in males (on average 3,1±0,78 ticks on one male and 1,6±0,36 on one female). The lowest value of parasite abundance was defined on one-year-old animals (no more than one tick per lizard). The ticks were attached on the forelegs, neck and sides of lizards. 89% of ticks were located under the forelegs. Domestic animals are the main feeders of imago I. ricinus, and the lizards provide feeding for larvae and nymphs.
Keywords: European common tick, Ixodes ricinus, Hyrcanian meadow lizard, Darevskia praticola hyrcanica, parasite-host relationships, Talysh Mountains, Southeastern Azerbaijan.
For citation: Kidov A. A. Parasite-host relationships of the ixodid tick Ixodes Ricinus (Linnaeus, 1758) and Hyrcanian meadow lizard Darevskia practicola hyrcanica (Tuniyev, Doronin, Kidov et Tuniyev, 2011) in Talysh Mountains (Southeastern Azerbaijan).
кидов, A.A. (2018) -
Цель исследований: выявление особенностей паразито-хозяинных отношений европейского лесного клеща Ixodes ricinus (Linnaeus,
1758) и гирканской луговой ящерицы Darevskia praticola hyrcanica (Tuniyev, Doronin, Kidov et Tuniyev, 2011).
материалы и методы. Исследования проводили в мае 2016 г. в урочище Гадазыгахи в Астаринском районе Азербайджана. Живот- ных отлавливали на маршрутах, измеряли длину тела по стандартным методикам, подразделяли на три половозрастные груп- пы (взрослые самки, взрослые самцы и годовики), обследовали на наличие клещей, определяли встречаемость, обилие и локали- зацию паразитов по общепринятым методикам. Локализацию паразитов на теле ящериц определяли по стандартной схеме.
результаты и обсуждение. На ящерицах были обнаружены лишь клещи преимагинальных стадий – личинки и нимфы I. ricinus. Всего клещи были отмечены на 32 ящерицах из 123 исследованных. Встречаемость клещей у годовиков гирканской луговой ящерицы со- ставила 10,8%; у взрослых самок – 28,6; у взрослых самцов – 35,3%. Распределение питающихся иксодовых клещей в выборке вида хозяина носило агрегированный характер: 26% изученных ящериц являлись прокормителями 100% изученных клещей. Более высо- ким обилием паразитов на одном животном характеризовались самцы (в среднем, 3,1±0,78 клещей на одной пораженной ящерице у самцов против 1,6±0,36 у самок). Годовики имели наименьшие значения обилия паразита (не более одного клеща на одну ящерицу). На ящерицах клещи прикреплялись на поясе передних конечностей, шее и боках. Наиболее предпочитаемое место прикрепления паразитов – область под передними конечностями, на которую приходилось 89% всех отмеченных клещей. Домашние животные являются основными прокормителями имаго I. ricinus, а ящерицы обеспечивают прокормление личинок и нимф.
ключевые слова: европейский лесной клещ, Ixodes ricinus, гирканская луговая ящерица, Darevskia praticola hyrcanica, зараженность, паразито-хозяинные отношения, Талышские горы, юго-восточный Азербайджан.
для цитирования: Кидов А. А. Паразито-хозяинные отношения иксодового клеща Ixodes ricinus (Linnaeus, 1758) и гирканской луговой ящерицы Darevskia praticola hyrcanica (Tuniyev, Doronin, Kidov et Tuniyev, 2011) в Талышских горах (юго-восточный Азербайджан). Рос- сийский паразитологический журнал.
Kidov A.A. (2021) -
Data on the reproduction of the rock lizard (Darevskia saxicola (Eversmann 1834)) in the gorges of Bol’shaya Laba (Karachay-Cherkessia) and Malaya Laba (Krasnodar Territory) rivers are presented. Gravid females of this species were captured in nature and transported to the laboratory. The animals were kept in captivity until oviposition. All eggs were obtained between July 20 and August 20. Breeding females had a body length of 42.5–60.1 mm. Hatches comprised 1–4 eggs 11.2–18.0 × 5.8–8.5 mm in size and weighing 0.185–0.535 g. Incubation in the laboratory lasted 38–47 days. Young lizards after hatching had a body length of 21.6–30.8 mm, tail 39.9–60.4 mm long, and weight of 0.375–0.635 g. A comparison of the reproductive characteristics of D. saxicola and syntopic species (D. derjugini (Nikolsky 1898) and D. pontica (Lantz et Cyren 1919)) was carried out. The late oviposition and relatively low fertility in the rock lizards are compen- sated for by the large size of eggs and newborn lizards.
Kidov, A.A. (2011) -
Кидов А.А. (2011) -
Kidov, A.A. (2018) -
The paper presents materials on the reproductive biology of the hyrcanian meadow lizard, Darevskia praticola hyrcanica from a typical locality (the Gadasygahi Natural Boundaries, Talysh Mountains, As- tara administrative district of Azerbaijan). Animals were caught in the first decade of May, 2016. After being caught, the adult lizards were kept in groups (one female and two males) in our laboratory until oviposition. Females with a body length of 45.8 – 60.6 mm were laying eggs from May 28 till June 20. A total of 25 egg clutches with 72 eggs were studied. The resulting clutches contained 1 to 6 eggs. The egg length was 8.71 – 13.4 mm, the width 5.4 – 7.0 mm, and the weight 0.18 – 0.36 g. Incubation at 27 – 30°C lasted 45 – 58 days. The newborn lizards at hatching from eggs had a body length of 19.4 – 25.7 mm, a tail length of 19.8 – 36.2 mm, and a body weight of 0.16 – 0.44 g.
Кидов А.А. (2018) -
Приводятся материалы по репродуктивной биологии гирканской луговой ящерицы, Darevskia praticola hyrcanica из типовой территории (урочище Гадазыгахи, Талышские горы, Астаринский район, Азербайджан). Животных отлавливали в I декаде мая 2016 г. После поимки взрослых ящериц группами (из одной самки и двух самцов) содержали в лаборатории до получения яиц. Самки с длиной тела 45.8 – 60.6 мм откладывали яйца в период с 28 мая по 20 июня. Всего были изучены 25 кладок яиц и 72 яйца. Полученные кладки насчитывали от 1 до 6 яиц. Дли- на яиц 8.71 – 13.4 мм, ширина 5.4 – 7.0 мм, масса 0.18 – 0.36 г. Инкубация при температуре 27 – 30°C составляла 45 – 58 суток. Молодь при вылуплении из яиц имела длину тела 19.4 – 25.7 мм, длину хвоста 19.8 – 36.2 мм, массу тела 0.16 – 0.44 г.
Kidov, A.A. (2019) -
Materials and methods. Observations of Amphibians and Reptiles were carried out during two tourist trips in March 2016 and in May 2018 in the Iranian provinces of Gilan and Ardebil.
Results. New data on the distribution of 6 species of Amphibians (Bufo eichwaldi, Bufotes variabilis, Hyla orientalis, H. savignyi, Pelophylax ridibundus, Rana pseudodalmatina) and 21 species of Reptiles (Emys orbicularis, Paralaudakia caucasia, Trapelus ruderatus, Darevskia chlorogaster, D. defilippii, D. raddei, Eremias strauchi, Iranolacerta brandtii, Lacerta strigata, Ophisops elegans, Ablepharus bivittatus, Heremites auratus, Pseudopus apodus, Eryx jaculus, Eirenis punctatolineatus, Hemorrhois ravergieri, Platyceps najadum, Telescopus fallax, Xerotyphlops vermicularis, Natrix natrix, N. tessellata) in the study area were obtained. The author compares the obtained data with the results of other Herpetological researches in the region.
Conclusions. In general, the herpetofauna of the Caspian slopes of the Talysh Range in Iran is an inseparable part of mesophilic herpetocomplexes with the prevalence of gyrcan elements common from the north-western foothills of Talysh in Azerbaijan to the Western Kopetdag in Turkmenistan.
Кидов, А.А. (2019) -
Актуальность и цели. Изученность герпетофауны иранской части гор Талыша и Ленкоранской низменности остается низкой. Цель настоящей работы – уточнить распространение земноводных и пресмыкающихся на территории иранского Талыша.
Материалы и методы. Наблюдения за земноводными и пресмыкающимися осуществляли в ходе двух туристических поездок в марте 2016 г. и в мае 2018 г. на территории иранских провинций Гилян и Ардебиль.
Результаты. Были получены новые данные о распространении на изучае- мой территории 6 видов земноводных (Bufo eichwaldi, Bufotes variabilis, Hyla orientalis, H. savignyi, Pelophylax ridibundus, Rana pseudodalmatina) и 21 вида пресмыкающихся (Emys orbicularis, Paralaudakia caucasia, Trapelus ruderatus, Darevskia chlorogaster, D. defilippii, D. raddei, Eremias strauchi, Iranolacerta brandtii, Lacerta strigata, Ophisops elegans, Ablepharus bivittatus, Heremites aura- tus, Pseudopus apodus, Eryx jaculus, Eirenis punctatolineatus, Hemorrhois ravergieri, Platyceps najadum, Telescopus fallax, Xerotyphlops vermicularis, Natrix natrix, N. tessellata). Автор проводит сравнение полученных данных с результатами других герпетологических исследований в регионе.
Выводы. В целом герпетофауна прикаспийских склонов Талышского хребта в Иране является неразрывной частью мезофильных герпетокомплексов с превалированием гирканских элементов, распространенных от северо-западных предгорий Талыша в Азербайджане до Западного Копетдага в Туркменистане.
Kidov, A.A. (2020) -
The paper presents materials on growing and winter hibernation of the Hyrcanian meadow lizard, Darevskia praticola hyrcanica, in artificial conditions. Juveniles were obtained from the captive breeding of lizards caught on the typical locality of this subspecies (Gadazyghahi Natural Boundary, the Talysh Mountains, Astara District, Azerbaijan) in the first decade of May, 2016. Immediately after leaving the eggs, newborn lizards were placed in groups of 5 specimens each into plastic 39×28×28 cm containers. Six groups were formed in total, and our research involved 30 lizards. The animals from the first three containers received only two-spotted crickets, Grillus bimaculatus, as feed (the control group). The liz- ards from the second three containers ate speckled cockroaches, Nauphoeta cinerea only (the experimen- tal group). The feed was offered to the lizards every other day. The growing lasted 450 days. The weight gain, survival and feed costs per unit weight of the animal (feed coefficient) were investigated. After growing the animals were placed into wintering rooms, where they were kept without feeding for 60 days at a temperature between 4–11oC. It was noted that during 450 days of growing the cockroach-fed lizards did not differ in weight from the cricket-fed animals. The feed coefficient of young meadow lizard grow- ing with cockroach (11.7–14.3) was similar to that of the animals that ate their traditional food, crickets (12.3–12.7). The survival rate of young lizards in different containers for 450 days of cultivation on both feed types was 100%. The lizards that ate cockroaches showed a better survival rate (100%) in the period of winter hibernation than the animals from the control group (86.6%).
Кидов, А.А. (2020) -
Приводятся материалы по выращиванию и зимовке гирканской луговой ящерицы, Darevskia praticola hyrcanica в искусственных условиях. Молодь была получена от размножения ящериц, пойманных на типовой территории подвида (урочище Гадазыгахи, Талышские горы, Астаринский район, Азербайджан) в I декаде мая 2016 г. Сразу после выхода из яиц новорожденных ящериц рассаживали группами по 5 экз. в пластиковые контейнеры раз- мером 39×28×28 см. Было сформировано 6 групп, а всего в исследованиях принимало участие 30 ящериц. Живот- ные из первых трех контейнеров в качестве корма получали только нимф двупятнистого сверчка, Grillus bimacu- latus (контрольная группа). Ящерицы из вторых трех контейнеров питались только нимфами пепельного тара- кана, Nauphoeta cinerea (опытная группа). Корма предлагали ящерицам через день. Выращивание длилось 450 суток. Исследовали прирост массы, выживаемость и затраты кормов на единицу массы животного (кормовой коэффициент). После выращивания животных помещали в зимовальные помещения, где содержали без кормле- ния 60 сут. при температуре 4 – 11 C. Отмечается, что за 450 сут. выращивания ящерицы, получавшие в качестве корма таракана, не отличались по массе от животных, поедавших двупятнистого сверчка. Кормовой коэффициент при выращивании молоди луговой ящерицы с использованием пепельного таракана (11.7 – 14.3) был схож с тако- вым у животных, питавшихся традиционным кормом – двупятнистым сверчком (12.3 – 12.7). Выживаемость мо- лоди ящериц в разных контейнерах за 450 сут. выращивания на обоих видах кормов составила 100%. За 2 месяца зимнего содержания ящерицы, до этого питавшиеся пепельным тараканом, характеризовались лучшей выживае- мостью (100%), чем потреблявшие двупятнистого сверчка (80%).
Data on the reproduction of the rock lizard (Darevskia saxicola (Eversmann 1834)) in the gorges of Bolshaya Laba (Karachay-Cherkessia) and Malaya Laba (Krasnodar Territory) rivers are presented. Gravid females of this species were captured in nature and transported to the laboratory. The animals were kept in captivity until oviposition. All eggs were obtained between July 20th and August 20th. Breeding females had a body length of 42.5–60.1 mm. Hatches comprised 1–4 eggs sized 11.2–18.0 × 5.8–8.5 mm and weighting 0.185–0.535 g. Incubation in the laboratory lasted 38–47 days. Young lizards after hatching had a body length of 21.6– 30.8 mm, tail 39.9–60.4 mm long, and weight 0.375–0.635 g. A comparison of the reproductive characteris- tics of D. saxicola and the syntopic species (D. derjugini (Nikolsky 1898) and D. pontica (Lantz et Cyren 1919)) was carried out. The late oviposition and relatively low fertility in the rock lizards are compensated for by the large size of eggs and newborn lizards.
Представлены данные о размножении скальной ящерицы (Darevskia saxicola (Eversmann 1834)) в ущельях рек Большая Лаба (Карачаево-Черкесия) и Малая Лаба (Краснодарский край). Беремен- ных самок этого вида отлавливали в природе и перевозили в лабораторию. Животных содержали в неволе до откладки яиц. Отмечается, что все кладки яиц были получены с III декады июля по III де- каду августа. Размножающиеся самки имели длину тела 42.5–60.1 мм. В кладках было от 1 до 4 яиц размерами 11.2–18.0 × 5.8–8.5 мм и массой 0.185–0.535 г. Инкубация в лабораторных условиях дли- лась 38–47 сут. Молодь после вылупления имела длину тела 21.6–30.8 мм, хвоста – 39.9–60.4 мм и массу 0.375–0.635 г. Проводится сравнение репродуктивных показателей D. saxicola с синтопиче- скими видами – D. derjugini (Nikolsky 1898) и D. pontica (Lantz et Cyren 1919). По мнению автора, поздние сроки откладки яиц и относительно невысокая плодовитость у скальной ящерицы воспол- няются крупными размерами яиц и новорожденных.
Kidov, A.A. & Ivanov, A.A. & Kondratova, T.E. & Stolyarova, E.A. & Nemyko, E.A. (2019) -
The paper discusses cases of eggs re-laying in two lizard species from the Darevskia (chlorogaster) com- plex. A male and a female of D. Chlorogaster were captured on 22 March, 2016 in the Gilan province (Iran), and she subsequently laid eggs on 29 May (4 eggs) and 5 July (4 ones). The eggs had length of 10.4–12.8 mm, the width of 6.1–6.5 mm, and the weight of 0.240–0.340 g. Young lizards came out from seven eggs after 52–55 days of incubation. The body length of these newborn lizards was 22.9–27.8 mm, the tail length was 38.5–46.5 mm, and the weight was 0.330–0.460 g. A male and a female of D. caspica were captured on May 22 in the Mazandaran province (Iran). The female laid eggs on June 15 (3 eggs) and July 12 (4 ones). The length of the eggs was 10.0–12.0 mm, the width was 5.0–7.0 mm, and the weight was 0.220–0.310. Young lizards exited from all eggs in 54–68 days. Their body length was 23.1– 26.0 mm, the tail length was 39.0–47.0 mm, and the weight was 0.300–0.390 g. The authors believe that these cases prove the possibility of multiple reproductions per season in D. chlorogaster and D. caspica.
Кидов А.А., Иванов А.А., Кондратова Т.Э., Столярова Е.А. & Немыко Е.А. (2019) -
Обсуждаются случаи повторной откладки яиц у двух видов ящериц из комплекса Darevskia (Chlorogaster). Самец и самка D. chlorogaster, отловленные 22 марта 2016 г. в провинции Гилян, в последующем откладывала яйца 29 мая (4 яйца) и 5 июля (4 яйца). Яйца имели длину 10.4 – 12.8 мм, ширину – 6.1 – 6.5 мм, массу – 0.240 – 0.340 г. Молодые вышли из 7 яиц через 52 – 55 суток инкубации. Длина тела новорожденных ящериц 22.9 – 27.8 мм, хвоста – 38.5 – 46.5 мм, масса – 0.330 – 0.460 г. Самец и самка D. caspica были пойманы 22 мая в провинции Мазендаран (Иран). Самка откладывала яйца 15 июня (3 яйца) и 12 июля (4 яйца). Длина яиц 10.0 – 12.0 мм, ширина – 5.0 – 7.0 мм, масса 0.220 – 0.310. Молодые ящерицы вышли из всех яиц через 54 – 68 суток. Длина их тела равнялась 23.1 – 26.0 мм, хвоста – 39.0 – 47.0 мм, масса – 0.300 – 0.390 г. Авторы считают, что эти случаи доказывают возможность неоднократного размножения за сезон у D. chlorogaster и D. caspica.
Kidov, A.A. & Ivanov, A.A. & Nemyko, E.A. (2019) -
Kidov, A.A. & Kovrina, E.G. & Timoshina, A.L. & Baksheyeva, A.A. & Matushkina, K.A. & Blinova, S.A. & Afrin, K.A. (2014) -
New data on the reproductive biology of the Artvin lizard from an isolated northeastern population in the Northwest Caucasus are presented. These lizards reach sexual maturity after their second wintering. Pregnant females have their body length from 53.4 to 63.7 mm and the mass of 3.8 - 5.9 g. Oviposition occurs from the first decade of June till the third decade of August. Clutches contain from 2 to 8 eggs with sizes of 9.3 - 14.8×5.7 - 7.1 mm and weights 0.23 - 0.28 g. On oviposition a female loses from 14.3 to 37.5% of her weight. The fertility of females depends on their length (r = 0.58) and weight (r = 0.60). The incubation duration at a temperature within 28 - 30°C is 41 - 48 days. The mass of eggs from their oviposition to hatching of young lizards increases by 2.7 - 4.0 times, the length does by 1.2 - 1.3 times, and the width does by 1.6 times. Young lizards with a body length of 22.4 - 26.4 mm and weight 0.29 - 0.45 g are hatched from the second decade of July till the first decade of October.
Кидов А.А., Коврина Е.Г., Тимошина А.Л., Бакшеева А.А., Матушкина К.А., Блинова С.А., Африн К.А. (2014) -
Впервые приводятся материалы по репродуктивной биологии артвинской ящерицы из изолированной северо-восточной популяции на Северо-Западном Кавказе. Половой зрелости ящерицы достигают после второй зимовки. Беременные самки имеют длину тела от 53.4 до 63.7 мм и массу 3.8 - 5.9 г. Откладка яиц происходит с I декады июня по III декаду августа. В кладках от 2 до 8 яиц размером 9.3 - 14.8×5.7 - 7.1 мм и массой 0.23 - 0.28 г. При откладке яиц самки теряют от 14.3 до 37.5% от своего веса. Плодовитость самок зависит от их длины (r = 0.58) и массы (r = 0.60). Длительность инкубации при температуре 28 - 30°C равняется 41 - 48 сут. Масса яиц от их откладки до вылупления молоди увеличивается в 2.7 - 4.0 раз, длина - в 1.2 - 1.3 раз, ширина - в 1.6 раз. Молодые ящерицы с длиной тела 22.4 - 26.4 мм и массой 0.29 - 0.45 г вылупляются со II декады июля по I декаду октября.
Kidov, A.A. & Kovrina, E.G. & Timoshina, A.L. & Matuschkina, K.A. & Blinova, S.A. & Afrin, K.A. (2015) -
Data on reproduction of the Black Sea lizard, Darevskia pontica (Lantz et Cyren, 1919) in the Northwestern Caucasus (a valley of Ubin River in the Seversky District of Krasnodar Krai) are provided. Adult females are larger than males. Oviposition was noted in June. Fecundity of females varied from 1 to 6 eggs. Sizes of eggs were the following: 8.5–11.9×4.3–6.8 mm. The incubation of eggs under the laboratory conditions lasted from 37 to 42 days. Newborn lizards had length of a body (L) 22.0–26.7 mm and mass of 0.2–0.4 g. Strong negative dependence of eggs length and length of newborn lizards from fertility of females is noted.
А.А. КИДОВ, Е.Г. КОВРИНА, А.Л. ТИМОШИНА, К.А. МАТУШКИНА, С.А. БЛИНОВА, К.А. АФРИН (2015) -
Приводятся данные о размножении понтийской ящерицы, Darevskia pontica (Lantz et Cyren, 1919) на Северо-Западном Кавказе (долина реки Убин в Северском районе Краснодар- ского края). Взрослые самки крупнее самцов. Откладка яиц отмечалась в июне. Плодови- тость самок составляла от 1 до 6 яиц. Размеры яиц — 8,5–11,9×4,3–6,8 мм. Инкубация яиц в лабораторных условиях длилась от 37 до 42 сут. Новорожденные ящерицы имели длину туловища (L) 22,0–26,7 мм и массу 0,2–0,4 г. Отмечена сильная отрицательная зависимость длины яиц и длины новорожденных ящериц от количества яиц в кладке.
Kidov, A.A. & Matushkina, K.A. (2016) -
is article provides data on the distribution of lacertid lizards in the region of the Southwest Caspian. e study was performed between 2007 and 2014 in the territory of Jalilabad, Masalli, Yardimli, Len- koran, Masalli and Lerik administrative districts of the Republic of Azerbaijan. On the basis of eld data and the results of studies of museum collections, we noted a change in the range of lizards under the in uence of anthropogenic deforestation. In particular, it is noted that there is a narrowing of the distribution of the green belly lizard, Darevskia chlorogaster and the extension of the habitats of Azer- baijani, D.raddei and striped, Lacerta strigata lizards. Because of the fact that the green belly lizard distribution in the region decreased by more than a third, it is proposed that it be included in the Red Data Book of Azerbaijan.
Kidov, A.A. & Nemyko, E.A. & Ivanov, A.A. & Pykhov, S.G. (2018) -
The article discusses the cases of finding pregnant females and eggs of the Pontic lizard (Darevskia pontica) in Northwest Caucasus in July and August. In the spring and early summer in 2017 in the region there were the temperatures below climate normal. The Pontic lizard began to lay eggs only in late July and in August. Along the Black sea coast of the Caucasus (Tuapse, Krasnodar Krai) there was egg laying in late July, and hatching took place from early August to early September. The Pontic lizards’ hatching in the North Caucasus (valley of the Malaya Laba River, the settlement Burniy, Krasnodar Krai) took place in July – August, and the young lizards hatched out in September. The authors believe that the cases of late laying in nature prove the changes in the reproductive season rather than some repeated parturiating.
А. А. Кидов, Е. А. Немыко, А. А. Иванов, С. Г. Пыхов (2018) -
В статье рассматриваются случаи находок во второй половине лета (в июле и августе) беременных самок и яиц понтийской ящерицы, Darevskia pontica, на Северо-Западном Кавказе. Весна и начало лета 2017 г. в указанном регионе характеризовались температурами ниже климатической нормы. Понтийская ящерица приступила к размножению только в конце июля – в августе. На Черноморском побережье Кавказа (город Туапсе Краснодарского края) кладки яиц от- мечались в конце июля, а молодь вылуплялась с начала августа по начало сентября. На Северном Кавказе (долина реки Малая Лаба, поселок Бурный Краснодарского края) кладки яиц были полу- чены в июле-августе, а молодые ящерицы вышли из яиц только в сентябре.
Kidov, A.A. & Pykhov, S.G. & Dernakov, V.V. (2009) -
Kidov, A.A. & Timoshina, A. (2017) -
We report the investigation results of reproductive biology of the Black Sea lizard Darevskia pontica in the valley of the Malaya Laba River (Northwest Caucasus). Based on allocation of groups in size and color, we assume that these lizards begin to reproduce after the second hibernation. Adult females of the Black Sea lizard are statistically superior to the males in body length and weight. Females collected in nature from the III decade of May to the I decade of June laid eggs in the laboratory in the II – III decades of June. In the nature, eggs were found also in the II–III decade of August. The body length of the females which brought the offspring was equal to 50,4–57,3 mm. Clutches contain 3–7 eggs with a mass of 0,21–0,45 g and size of 9,0–10,8×5,1–6,4 mm. Incubation in the laboratory conditions lasted for 40–47 days. Newborn lizards had a mass of 0,20–0,36 g and a body length of 20,2–26,5 mm.
Кидов А.А. & Тимошина А.Л. (2017) -
статье приводятся результаты изучения репродуктивной биологии понтий- ской ящерицы, Darevskia pontica в долине реки Малая Лаба (Северо-Западный Кавказ). На основании выделения групп по размерам и окраске предполагается, что к размножению эти ящерицы приступают после второй зимовки. Взрослые самки понтийской ящерицы статистически значимо превосходят самцов, как по длине тела, так и по массе. Самки, от- ловленные в природе в III декаде мая – I декаде июня, в лаборатории откладывали яйца во II–III декадах июня. В природе находили кладки также во II–III декадах августа. Длина тела самок, принесших потомство, равнялась 50,4–57,3 мм. Кладки содержали 3–7 яиц массой 0,21–0,45 г и размерами 9,0–10,8×5,1–6,4 мм. Инкубация в лабораторных условиях длилась 40–47 суток. Новорожденные ящерицы имели массу 0,20–0,36 г и длину тела 20,2–26,5 мм.
Kidov, A.A. & Timoshina, A. &Hairutdinov, I. & Matushkina, K. (2016) -
We report data on the age and growth of the Black Sea lizard in the Ubin River valley
(Severskiy District of Krasnodar Krai, Russia, Northwest Caucasus). Using skeletochronological
analysis, we have studied 111 finger phalanxes, including 59 from females and 52 from males.
The average age of females in the studied samples is 5.6±0.12 years (from 4+ to 8+ years).
Males have an average age of 4.58±0.12 (from 3+ to 6+ years). Lizards of different age groups
have a high individual variability in size and body weight. This circumstance makes it impossible
to determine the age of these animals on the basis of allocation of different size`s groups. From studied females, 14 specimens aged 5+ to 8+ years (average age of 5.7±0.28 years) yielded
offspring. Age variability in the reproductive performance has not been observed.
Kidov, A.A. & Timoshina, A.L. & Hayrutdinoiv, I.Z. & Matushkina, K.A. (2017) -
The article provides the data on the age, growth and reproduction of the Derjugin’s lizards on the northeastern periphery of the area – in the gorge of the Malaya Laba River (Northwestern Caucasus). With the help of skeletochronological analysis the age of 29 lizards (16 males and 13 females) was determined. The age of the males was 3–5, the average of which is 3.9. The females were also aged 3–5 with the average of 3.7. The relative body length of the three-year-old animals in comparison with the length of the body of the newborn lizards was 172.0–195.5% (males) and 205.3–258.9% (females). At the age of 4 the length of the body of the lizards was 172.4–206.1% (males) and 215.0254.5% (females). The five-year-old males and females had a relative length of the body of 179.7–213.8% and 188.6–230.5% respectively. Among the thirteen females under consideration, the eight exemplars brought forth. A large part of the breeding lizards were aged 3 (5 exemplars), but there were also a four-year-old (2 exemplars) and one five-year-old female. The authors stress that in the North-East of the area the Derjugin’s lizard has a relatively small size and later onset of puberty in females. The average age and maximum life span of lizards studied are consistent with the literature data.
Kimovna, P.R. (2018) -
Кимовна, П.Р. (2018) -
Klemmer, K. (1957) -
Koc, H. & Bülbül, U. & Kutrup, B. (2018) -
The current literature and our field observations have shown that spiny-tailed lizard may also be active in the winter season. Activity of Darevskia rudis in the February was recorded for the first time in the province of Giresun, Eastern Black Sea Region, Turkey. In conclusion, we assessed that the spiny-tailed lizard could be active throughout the year due to global climate changes.
Koç, H. & Bülbül, U. & Kutrup, B. (2018) -
Koc, H. & Kutrup, B. & Bülbül, U. & Kurnaz, M. (2021) -
The spiny-tailed lizard, which has a series of taxonomic revisions, is one of the most common lizard species in Turkey. In this study, sequence data derived from three microsatellite loci (Du215, Du281, and Du323), two mito chondrial (16S rRNA and Cyt-b) genes and combined data were used to evaluate the taxonomic status of Darevskia rudis and Darevskia bithynica with new samples from all subspecies populations in Turkey. Our results indicated that the genetic variations of microsatellite loci were not specific to populations within species, and only minor differences separated D. rudis and D. bithynica populations. Furthermore, the markers we used for phylo genetic analyses (NJ, ML, MP, and BI) produced topologically similar trees based on 16S rRNA and Cyt-b while the combined data produced conflicting trees with the separate gene analyses. Finally, the basal relationships among the populations in Turkish populations D. rudis and D. bithynica were not resolved with this dataset, and we found a hard polytomy at the basis of the phylogeny.
Koc, H. & Kutrup, B. & Eroglu, O. & Bülbül, U. & Kurnaz, M. & Afan, F. & Eroğlu, A.I. (2016) -
The spiny-tailed lizard of the genus Darevskia have a series of taxonomic revisions, but still their phylogenetic relationships remain uncertain. In the present study, we have assessed taxonomic relationships among Darevskia bithynica and Darevskia rudis populations through estimation of phylogenetic relationships among 96 specimens using microsatellite DNA (Du215, Du281, Du323 and Du418 loci) and 53 specimens using mtDNA (16S rRNA and cytb) from main populations in Turkey. Although D. b. bithynica and D. r. mirabilis were separated based on the PCoA analysis at low level from other D. rudis and D. bithynica populations, the distance values of Nei’s genetic distance, Nei’s unbiased genetic distance, Fst and Linear Fst were not high among taxa in microsatellite DNA. On the other hand, our phylogenetic analyses (NJ, ML, MP and BI) did not separate D. rudis and D. bithynica populations. Finally, most of the topologically identical trees of phylogenetic analyses and microsatellite results showed that the extant populations of D. rudis and D. bithynica were found to be polytomy. Based on our molecular phylogenetic study, D. rudis complex is still ongoing revisions.
Kodym, P. & Frynta, D. & Moravec, J. & Kaftan, M. & Hodková, Z. (1999) -
Komissarov, A. & Korchagin, V. & Kliver, S. & Dobrynin, P. & Semyenova, S. & Vergun, A. & O`Brien, S: & Ryskov, A. (2016) -
The first complete mitochondrial genome sequence of parthenogenetic Caucasian rock lizard Darevskia unisexualis (Lacertidae family) is determined by hybrid assembly with Illumina HiSeq and PacBio RS II platforms. The circular 21.4 kbp mitogenome contains 13 protein-coding genes, 12S and 16S rRNA genes, 20 tRNAs, two pseudogenized tRNAs, and one long tandem repeats with 4.1 kbp length formed by 59 bp monomer repeated x70.6 times located before control region. This finding represents a new example of mitogenome variation in lizards of hybrid origin, providing fundamental data for following study of a unique hybridization system formed by parthenogenetic and bisexual species in the mountain steppe of central Armenia.
Korchagin, V.I. & Badaeva, T.N. & Tokarskaya, O.N. & Martirosyan, I.A. & Darevsky, I.S. & Ryskov, A.P. (2007) -
Populations of parthenogenetic lizards of the genus Darevskia consist of genetically identical animals, and represent a unique model for studying the molecular mechanisms underlying the variability and evolution of hypervariable DNA repeats. As unisexual lineages, parthenogenetic lizards are characterized by some level of genetic diversity at microsatellite loci. We cloned and sequenced a number of (GATA)n microsatellite loci of Darevskia unisexualis. PCR products from these loci were also sequenced and the degree of intraspecific polymorphism was assessed. Among the five (GATA)n loci analysed, two (Du215 and Du281) were polymorphic. Cross-species analysis of Du215 and Du281 indicate that the priming sites at the D. unisexualis loci are conserved in the bisexual parental species, D. raddei and D. valentini. Sequencing the PCR products amplified from Du215 and Du281 and from monomorphic Du323 showed that allelic differences at the polymorphic loci are caused by microsatellite mutations and by point mutations in the flanking regions. The haplotypes identified among the allelic variants of Du281 and among its orthologues in the parental species provide new evidence of the cross-species origin of D. unisexualis. To our knowledge, these data are the first to characterize the nucleotide sequences of allelic variants at microsatellite loci within parthenogenetic vertebrate animals.
Korchagin, V.I. & Churikov, N.A. & Tokarskaya, O.N. & Sevastyanova, G.A. & Filippovich, Y.B. & Ryskov, A.P. (2004) -
Hypervariable mini- and microsatellites represent the most widespread and universal system of molecular markers and are widely used in population and evolutional biology, genomic marking, paternity determination, and medico-biological studies [1, 2]. These repeats are classified with the most instable regions of eukaryotic genomes and are characterized by the highest mutation rates reported for genetic loci [3, 4]. The studies of individual mini- and microsatellite loci showed that changes in them significantly vary depending on the species, repeat type, alleles, age, and sex. However, the genesis of mini- and microsatellite loci, intensively studied in the human  and some other bisexual species , practically has not been studied in species with clonal type of reproduction. Obligate parthenogenetic reptiles, whose reproduction system implies clonal reproducing of maternal genotype in a series of generations, is a unique model system for monitoring of genetic variability in loci with increased mutation activity . Earlier, multilocus DNA fingerprinting was used to study variability of several types of mini- and microsatellite DNA markers in four parthenogenetic species of the genus Darevskia ( D. dahli, D. armeniaca, D. unisexualis , and D. rostombekovi ) [8‐11]. It was shown that genetic heterogeneity of parthenogenetic species with respect to some microsatellite markers may be detected against the background of species-specific DNA fingerprints. The most variable were the loci that contained (TCC) n , (TCT) n , and (GATA) n microsatellites in D. unisexualis genome [8, 9, 12‐14]. DNA fingerprints of parthenogenetic D. unisexualis families demonstrated high frequency of mutations for these loci already in the firstgeneration progeny [9, 14]. Furthermore, mutant (GATA) n fingerprint phenotypes were also detected in the progeny of population D. unisexualis samples. This finding is indicative of the mutant nature of genotypic diversity of D. unisexualis populations with respect to (GATA) n loci. However, the molecular nature and mechanisms of occurrence of these mutations remain obscure. It is conceivable that certain progress in this field may be reached with the use of genetic engineering studies of variable loci. In this study, we for the first time cloned and sequenced one variable locus of D. unisexualis genome and revealed the nature of structural variations of its allele variants. Total cell DNA was isolated form blood of adult D. unisexualis by standard phenol‐chloroform method using protein kinase K. DNA was hydrolyzed with endonuclease Sau 3A (Amersham) until complete degradation and ligated with the pUC12 vector, treated with Bam H1 restriction endonuclease using standard procedure. The ligase mixture was used to transform XL-1 Blue E. coli cells (Stratagene). The clonotheque obtained was then used for searching for recombinant clones containing (GATA) n microsatellites. The screening of approximately 1000 recombinant clones using
Korchagin, V.I. & Martirosyan, I.A. & Omelchenko, A.V. & Darevsky, I.S. & Ryskov, A.P. & Tokarskaya, O.N. (2004) -
The genesis of mini- and microsatellite loci, which is under extensive study in humans and some other bisexual species, has been virtually overlooked in species with clonal mode of reproduction. Earlier, using multilocus DNA fingerprinting, we have examined variability of some mini- and microsatellite DNA markers in parthenogenetic lizards from the genus Darevskia. In particular, mutant (GATA)n-restriction DNA fragments were found in Darevskia unisexualis. In the present study, we examined intraspecific polymorphism of three cloned loci of D. unisexualis—Du323, Du215, and Du281—containing (GATA)7GAT(GATA)2, GAT(GATA)9, and (GATA)10TA(GATA) microsatellite clusters, respectively. Different levels of intrapopulation and interpopulation variability of these loci were found. Locus Du281 showed the highest polymorphism—(six allelic variants in the sample of 68 DNA specimens). Three alleles were found for locus Du215. The Du323 locus was electrophoretically invariant. The primers chosen for loci Du323, Du215, and Du281 were also used for PCR analysis of homologous loci in two presumptive parental bisexual species, D. valentini and D. nairensis. The PCR products of the corresponding loci of the parental species had approximately the same size (200 bp) as their counterparts in D. unisexualis, but the polymorphism levels of the paternal, maternal, and hybrid species were shown to be somewhat different. These data on the structure of the D. unisexualis loci provide a possibility to study genetic diversity in the parthenogenetic species D. unisexualis and other related unisexual and bisexual species of this genus, which can provide new information on the origin of parthenogenetic species and on the phylogenetic relationships in the genus Darevskia. These data can also be used for resolving problems of marking the lizard genome, which is still poorly studied.
Корчагин B.И., Мартиросян И.А., Омельченко А.В., Даревский И.С., Рысков А.П., Токарская О.Н. (2004) -
Проблема генезиса мини- и микросателлитных локусов, интенсивно изучаемая на человеке и некоторых других двуполых видах, практически не исследована на видах с клональным типом размножения. Ранее с помощью мультилокусного ДНК-фингерпринтинга нами впервые была изучена изменчивость некоторых мини- и микросателлитных маркеров ДНК у партеногенетических ящериц рода Darevskia. В частности, у Darevskia unisexualis были обнаружены мутантные (GATA)n-pecrриктные фрагменты ДНК в фингерпринтных спектрах партеногенетических потомков. В настоящей работе проведен анализ внутривидового полиморфизма трех клонированных локусов D. unisexualis - Du323, Du215 и Du281, содержащих (GATA)7GAT(GATA)2, GAT(GATA)9 и (GATA)10TA(GATA) микросателлитные кластеры соответственно. Обнаружен разный уровень внутри- и межпопуляционной изменчивости исследованных локусов. Наиболее полиморфным - шесть аллельных вариантов (выборка из 68 образцов ДНК) - оказался локус Du281. Три аллельных варианта показаны для локуса Du215. Электрофоретически инвариантным оказался локус Du323. Праймеры, подобранные для локусов Du323, Du215 и Du281, были также использованы для ПЦР-анализа гомологичных локусов двух предполагаемых родительских двуполых видов D. valentini и D. nairensis. Показано, что ПЦР-продукты соответствующих локусов родительских видов имеют примерно такие же размеры (~200 пн), что иуО. unisexualis, но по уровню полиморфизма выявлены определенные различия между отцовским, материнским и гибридным видами. Полученные данные о структуре локусов D. unisexualis позволяют изучать генетическое разнообразие у партеновида D. unisexualis и других родственных однополых и двуполых видов ящериц этого рода, что может дать новую информацию о происхождении партеновидов и о филогенетических связях в группе ящериц рода Darevskia. Эти данные также могут быть использованы для решения задач по маркированию мало изученного пока генома ящериц.
Korchagin, V.I. & Martyrosian, I.A. & Malysheva, D.N. & Badaeva, T.N. & Tokarskaya, O.N. (2003) -
Korchagin, V.I. & Tokarskaya, O.N. (2010) -
Microsatellite repeats are one of the most widespread elements of the eukaryotic genome, but are poorly studied in species with clonal reproduction. PCR analysis and DNA sequencing were used to study the molecular structure of the allelic variants of microsatellite locus Du47D in the parthenogenetic species Darevskia unisexualis and its evolutionary ancestors, bisexual species D. raddei and D. valentini, of the genus Darevskia (Lacerta saxicola complex). Sequencing showed that the allelic variants of the D. unisexualis Du47D locus and the alleles of its D. raddei and D. valentini orthologs have a perfect microsatellite cluster structure, differ in number of ATT monomeric units, and have certain species-specific combinations of nucleotide substitutions, deletions, and insertions in the microsatellite-flanking DNA sequences. The Du47D alleles that the parthenogenetic species inherited from D. valentini or from D. raddei were identified.
Корчагин В.И., Токарская О.Н. (2010) -
Микросателлитные повторы, являясь одним из наиболее распространенных элементов генома эукариот, мало изучены у видов с клональным типом размножения. С помощью ПЦР-анализа и секвенирования ДНК изучена молекулярная структура аллельных вариантов тринуклеотидного микросателлитного локуса Du47D у партеновида Darevskia unisexualis и эволюционно-прародительских двуполых видов ящериц D. raddei и D. valentini рода Darevskia (Lacerta saxicola комплекс). Секвенирование аллельных вариантов Du47D у D. unisexualis и аллелей ортологичного локуса у D. raddei и D. valentini показало, что все исследованные аллели характеризуются совершенной структурой микросателлитного кластера, отличаются количеством мономерных (AAT)-звеньев и образуют определенные видоспецифические сочетания нуклеотидных замен, делеций и инсерций в прилежащих к микросателлиту последовательностях ДНК. Установлено, какие аллели Du47D партеновида унаследованы от D. valentini и какие от D. raddei.
Korchagin, V.I. & Vergun, A.A. & Godakova, S.A. & Tokarskaya, O.N. (2013) -
The molecular structure of the allelic variants of (AAT) n of the Du47D microsatellite locus was determined in parthenogenetic lizards Darevskia dahli, D. armeniaca, and D. rostombekovi. Comparative analysis of these alleles showed that they were characterized by perfect structure of microsatellite cluster, and were different in the number of (AAT) monomeric units, as well as in the combinations of species-specific substitutions and deletions in the microsatellite flanking regions. Molecular structure of microsatellite cluster, species-specific single nucleotide polymorphism (SNP), and different representation of alleles Du47 in the samples of parthenogenetic species examined point to the origin of the alleles from different bisexual species, which is consistent with the hybrid nature of unisexual species of the genus Darevskia. In addition, these data reflect different combination patterns of interspecific hybridization events with the participation of the same bisexual species upon the formation of hybrid genomes of parthenogenetic species. Possible application of the allelic variants of microsatellite loci of parthenogenetic lizards as the genetic markers for the analysis of the genomes of parthenogenetic species in the light of evolution, ecology, and parthenogenetic type of reproduction in vertebrates is discussed.
Корчагин В.И., Вергун А.А., Годакова С.А., Токарская О.Н. (2013) -
Определена молекулярная структура аллелей (AAT)n-микросателлитного локуса Du47D у партеногенетических ящериц Darevskia dahli, D. armeniaca и D. rostombekovi. Сравнительный анализ этих аллелей показал, что они характеризуются совершенной структурой микросателлитного кластера, различаются количеством мономерных (AAT)-звеньев, а также сочетаниями видоспецифических нуклеотидных замен и делеций в прилежащих к микросателлиту областях ДНК. Молекулярная структура микросателлитного кластера, видоспецифический однонуклеотидный полиморфизм (SNP single nucleotide polymorphism) и различная представленность аллелей Du47D в исследованных выборках партеновидов указывают на происхождение аллелей от различных двуполых видов, что согласуется с гибридной природой однополых видов рода Darevskia. Кроме того, это отражает комбинаторику межвидовых гибридизаций одних и тех же двуполых видов при образовании гибридных геномов партеновидов. В работе также обсуждается возможность использования аллелей микросателлитных локусов партеногенетических рептилий как генетических маркеров для изучения геномов партеновидов в свете проблем эволюции, экологии и партеногенетического типа размножения у позвоночных животных.
Korkiya, I.K. (1976) -
Kosushkin, S.A. (2003) -
Kosushkin, S.A. & Bannikova, A.A. & Grechko, V.V. & Darevsky, I.S. & Kramerov, D.A. (2007) -
IS-PCR method reveals a set of DNA sequences separating copies of short interspersed repeats (SINE) fBuntjer, 1997]. The resulting electrophoretic patterns possess taxon-specific features at a intra-generic level. Degrees of molecular genetic diversities have been tested by the values of DNL coefficients and roughly range between 0.0 - 0.20 for intrapopulational levels, 0.3 - 0.5 for intersubspecils of one species, and about 0.6 - 0.9 for known good species. These values were obtained in our study of populations of Darevskia raddei complex and their two systematic subspecies in comparison with some other species of Darevskia group (D. rudis, D. chlorogaster and`D. tristis`). On the basis of these features for L7 D. raddei populations we show that the difference of south-western Azerbaijan (Talysh) population from other populations by DNL (0.a) is similar to that befween D. r. raddei and D. r. nairensis. This observation supports the subspecies status for Talysh sample. The same values of molecular genetic diversity were found for D. rudis obscura and`D. tristis` from North- Pontic Ridge of Turkey that could argue forthe subspecies level for `D. tristis` within the D. rudis complex, as was suggested by Bohme and Bischoff . Both groups mentioned differ from D. raddei by the values of DNL of 0.6 - 0.7. Nearly the same correlations were obtained when some populations of D. praticola and D. derjugini were studied. Some of the known systematic subspecies of these species were supported by IS-PCR markers, the others were not: the data will be presented. In another species complex studied by IS-PCR - Lacerta s. str. - 12 populations inhabitating a vast territory (flom Baltic States and Ural Mountains to Caucasus) differed by DNL ranging from 0.02 to 0.2 apparently belong to L. agilis exigua subspecies as was deduced by morphology. The same DNL values characterize intrapopulational similarity inthe L. a. chersonensis, but the differences between these two systematic subspecies reach the values of around 0.6. The samples from Munchen (presumably L. a. argtts) also differs from the first two by 0.6 - 0.7. The most important is that the molecular differences between each of these subspecies and, L. strigata and L. media were characterized also by the same order of values. In other words, the genetic distances between I. strigata, L. media, and three subspecies of L. agilis have the same level and all of them could be considered either as subspecies of L. agilis, or the three subspecies of this species could be evaluated as a separate species.
Kosushkin, S.A. & Grechko, V.V. (2013) -
To study the molecular genetic relationships and correlate them with the taxonomy within the complex of lacertid lizards of the genus Darevskia, the locus analysis of the copies of the SINE-type repeat (Squam1) specific for the order Squamata was used. It was demonstrated that one of the loci (No. 34) contained the Squam1 copy insert in all species and subspecies of the examined genus. SINE allelic copies in some of the loci contained large indels and specific sets of mutations. The allelic variant M (medium, about 340 bp) was found most frequently; it was detected in all subspecies of D. saxicola (saxicola, darevskii, szczerbaki, lindholmi) and in most of the other species of the genus. Two species, D. derjugini and D. praticola, differed from the other species in the presence of long (L) and short (S) alleles. The longest allele was characteristic of the D. derjugini population from the Northern Caucasus (L, 379 bp, ssp. silvatica), while the shortest allele (97 bp) united the derjugini and barani subspecies. The second allele S (279 bp) characterizes the subspecies D. praticola praticola, some individuals of which also carry allele M. The second subspecies, D. p. pontica, contains allele L2, which differs from all other medium alleles in the presence of strictly specific short indel. In addition to apomorphic indels, the specificity and mutation distribution patterns among the Squam1 alleles were also examined. An analysis of the NJ tree indicated the concordance between morphological and molecular genetic characters of the species derjugini, praticola, and saxicola. Furthermore, four subspecies of D. saxicola were much closer to each other than the subspecies within the first two species; D. d. silvatica and the group of D. d. derjugini + barani were clearly separated. It cannot be excluded that populations from Azerbaijan and Serbia can be treated as the independent subspecies of D. praticola.
Косушкин С.А., Гречко В.В. (2013) -
Для изучения молекулярно-генетического родства и сопоставления его с таксономией в комплексе ящериц-лацертид рода Darevskia применили локусный анализ копий повтора типа SINE (Squam1), специфичного для отр. Squamata. Один из локусов (№ 34) содержит копию Squam1 во всех исследованных видах и подвидах рода, некоторые из локусов имеют крупные индели и специфический набор мутаций. Наиболее часто обнаруживается аллельный вариант М (средний, около 340 пн) во всех подвидах D. saxicola (saxicola, darevskii, szczerbaki, lindholmi) и в большинстве других видов рода. Два вида D. derjugini и D. praticola отличаются от всех по содержанию в них длинных (L) и коротких (S) аллелей. Наиболее длинный аллель характеризует популяцию D. derjugini Сев. Кавказа (L, 379 пн, ssp. silvatica), наименьший (97 пн) объединяет подвиды derjugini и barani. Второй S-аллель (279 пн) характеризует подвид D. praticola praticola; некоторые особи обладают, кроме того, аллелем М. Второй подвид D. p. pontica содержит аллель L2, который отличается от всех других средних аллелей по строго специфичной небольшой индели. Помимо апоморфных инделей учитывали также специфичность и паттерны распределения мутаций в аллелях Squam1. NJ-дерево свидетельствует не только о соответствии между морфологическими и молекулярно-генетическими признаками видов derjugini, praticola и saxicola, но и о том, что четыре подвида D. saxicola гораздо более близкородственны, чем подвиды внутри первых двух видов; обособляются также D. d. silvatica и группа D. d. derjugini + barani. Не исключается возможность выделения популяций из Азербайджана и Сербии в качестве самостоятельных подвидов D. praticola.
Kotenko, T.I. (2010) -
This article includes a brief outline of herpetological studies in Crimea over the past 50 years, checklists of taxa of amphibians and reptiles, and a general description of the herpetofauna of Crimea. Some formal issues for the protection of amphibians and reptiles are also considered.
Котенко, Т.И. (2010) -
Два класса холоднокровных позвоночных – земноводные или амфибии1, и пресмыкающиеся или рептилии – по своей морфологии, биологии и образу жизни сильно различаются, однако традиционно рассматриваются вместе и изучаются в рамках специального раздела зоологии – герпетологии. И хотя последнюю часто разделяют на собственно герпетологию или науку о рептилиях, и батрахологию или науку об амфибиях, в этом обзоре мы будем придерживаться широкой трактовки термина «герпетология». Предлагаемая статья включает краткий очерк герпетологических исследований в Крыму за последние 50 лет, контрольные списки таксонов амфибий и рептилий, общую характеристику герпетофауны Крыма. Рассматриваются также некоторые формальные вопросы охраны амфибий и рептилий.
Kovaleva, A.V. & Galoyan, E.A. (2021) -
Krasilnikov, E.N. (1967) -
Kretzschmar, K.G. (1995) -
The dangers of helminth infestations for lizards, particularly infestations with nematodes und cestodes, are reported. Infection routes, diagnoses, und treatments are described.
Kroniger, M. & Zawadzki, M. (2005) -
Kukushkin, O. & Ermakov, O. & Gherghel, I. & Lukonina, S. & Svinin, A. & Doronin, I. & Simonov, E. & Jablonski, D. (2021) -
The Lindholm rock lizard, Darevskia lindholmi, is the only member of the genus Darevskia whose range is restricted solely to Europe, representing a local endemism found only in the Crimean Mountains. In our study, we investigated the cytochrome b gene (mtDNA) of 101 D. lindholmi sequences from 65 Crimean localities, representing its entire range. We found that D. lindholmi is highly genetically structured, and its range is divided into populations belonging to three mitochondrial lineages. The Lindholm rock lizard populations inhabiting the middle part of the Crimean Mountains (further referred to as the Central lineage) are sharply differ entiated from the other two lineages (the Common and the Southwestern lineages), which are present in most of the species range. The genetic distance between the Central lineage and the other two taken together is 4.6%, according to our results, suggesting that the divergence occurred during the Early Pleistocene. The narrowly distributed Southwestern lineage and the widespread Common lineage, on the other hand, are differentiated by 1%. Field observations on the representatives of the main evolutionary groups show that their ecology is also different: the Central lineage is a mesophilic and cold-resistant form, while the other two closely related lineages are more xerophilic and thermophilic. Results of the potential ranges modeling and ecological niche analysis confirm that the genetic lineages occupy different niches of the Crimea. Furthermore, the area of inhabitation of the Central lineage splits the western and eastern parts of the Common lineage range, while the Southwestern lineage is restricted along the coast of the southwestern coast of the peninsula. The long-term co-existence of deeply divergent sister mitochondrial lineages in a relatively small (circa 7,000 km2) isolated mountain system serves as a mesocosm for understanding the speciation process. Our data suggest that the Central lineage warrants further taxonomic investigation.
Kukushkin, O.V. & Doronin, I.V. & Tuniyev, B.S. & Ananjeva, N.B. & Doronina, M.A. (2017) -
An overview of the cases of introduction (both accidental and intentional) of amphibians and reptiles in
the Caucasus and Crimea is provided. The introduction cases are systemized in accordance with the goals
and ways of introduction and with regards to the expected scale of the impact and its nature. No negative
consequences of the introduction of amphibian and reptile species in the Crimea and Caucasus are currently
observed on any of the known examples. No cases of species naturalization, which the “invasion”
term can be applied to, have been revealed. The importance of allochthonous populations as experimental
sites to study microevolutionary processes and ecological adaptations of the species in new environmental
conditions is discussed.
Кукушкин О.В., Доронин И.В., Туниев Б.С., Ананьева Н.Б., Доронина М.А. (2017) -
Представлен обзор случаев интродукции (как случайной, так и преднамеренной) земноводных и пресмыкаю- щихся на Кавказе и в Крыму. Факты интродукции систематизированы по целям и путям проникновения живот- ных с учетом предполагаемого масштаба воздействия и характера влияния на природные комплексы. Негатив- ные последствия интродукции видов земноводных и пресмыкающихся в Крыму и на Кавказе в настоящее время не прослеживаются ни на одном из известных нам примеров; случаи натурализации видов, к которым обосно- ванно может быть применен термин «инвазия», не выявлены. Рассмотрено значение аллохтонных популяций как экспериментальных полигонов для изучения микроэволюционных процессов и экологических адаптаций видов в новых для них условиях.
Kukushkin, O.V. & Lukonina, S.A. & Simonov, E.P. & Doronin, I.V. & Ermakov, O.A. (2021) -
Kukushkin, O.V. & Sviridenko, E.Yu. (2002) -
Kukushkin, O.V. & Trofimov, A.G. & Turbanov, I.S. & Slodkevich, V.Y. (2019) -
This work summarizes information on the distribution and status of the populations of amphibians and reptiles of the city of Sevastopol. Data obtained over a quarter of a century were refined by a targeted herpetological examination of the entire territory of Sevastopol (over 1000 km2) in 2018 and early 2019. Most species of am- phibians and reptiles known in Crimea are recorded from the Sevastopol Region, with the exception of some taxa that inhabit only or mainly plains environments (Pelobates vespertinus, Eremias arguta, and Lacerta agilis exigua). Most taxa included in the Red Book of Sevastopol, to date, retain stable populations. Apparently, Vi- pera renardi has disappeared from the region. Analysis of the taxon chorotypes indicates a dominance of spe- cies of Mediterranean (sensu lato) origin. The mild climate of the southwestern part of the Crimean Peninsula determines the unique spatial distribution of the most thermophilic reptile species (Mediodactylus danilewskii, Pseudopus apodus, and Zamenis situla) and, in particular, their wide distribution on the northern macroslope of the Crimean Mountains and (or) the highest elevations in Crimea. The zoning of the territory of Sevastopol, according to herpetological data, made it possible to identify eight districts that differ clearly in species compo- sition and population density of background and rare species. On a national scale, the territory of Sevastopol is important for the conservation of the genetic diversity of species such as Triturus karelinii, M. danilewskii, Ps. apodus, Z. situla, Dolichophis caspius, and Elaphe sauromates. Currently, the state of the populations of T. karelinii, Emys orbicularis, and El. sauromates is most alarming. Scientifically important natural and some synanthropic (in the Khersonesos of Taurida) populations of M. danilewskii, as well as relic populations of the Crimean endemic Lacerta agilis tauridica, require close attention. The “Baydarskyi”, “Cape Aya” and “Laspi” state regional wildlife sanctuaries play the most significant role in preserving the herpetofauna of the region, covering the upper part of the Chernaya River basin (Main Range) and the extreme southwestern part of the Southern Coast of Crimea, as well as the Mekenzievskoe Forestry in the foothills.
Kukushkin, O.V. & Turbanov, I.S. & Gorelov, R.A. & Trofimov, A.G. (2021) -
New data on the boundaries of the range of the Lindholm lizard (Darevskia lindholmi), an endemic of the Crimean peninsula, are presented. The species is petrophilic and inhabits a wide range of biotopes in various landscape layers of the Crimean Mountains. The upper boundary of the distribution of D. lindholmi in the southwest of the Main Ridge of the Crimean Mountains reaches 1520 m above sea level. m. (Ai-Petrinskaya yayla, Mount Kemal-Egerek), while on other highlands with maximum heights of over 1.5 km, but colder climate (Babugan, Chatyrdag), the species was traced only up to 1250-1320 m above sea level. m. The northern border of the D. lindholmi range in the western part of the Crimean Mountains runs along the Outer piedmont ridge (right bank of the Alma River), while in the eastern one, along the last rock massifs of the Inner piedmont ridge north of 45 ° N. Isolated peripheral populations found in the forest-steppe or frigan-steppe landscapes of the Crimean foothills and the arid South-East coast differ significantly in their remoteness from the main range, the number and density of the lizard population. A hypothetical history of the formation of the current range of D. lindholmi is discussed.
О.В. Кукушкин, И.С. Турбанов , Р.А. Горелов & А.Г. Трофимов (2021) -