Parthenogenetic species, D. armeniaca, originated as a result of the interspecific hybridisation between bisexual species D. valentini (“paternal”) and D. mixta (“maternal”) (Darevsky 1967, Murphy et al. 2000). However, there is a hypothesis that D. armeniaca might be a descendant of the hybridisation between D. valentini males and parthenogenetic D. dahli (Tarkhnishvili et al. 2017). Darevskia armeniaca identification from different SZs was performed in different years by different authors using allozyme loci, mt-DNA, multilocus DNA fingerprinting, mini- and micro-satellite markers and morphological features (Darevsky 1966, Darevsky 1967, Darevsky and Danielyan 1968, Uzzell and Darevsky 1975, Tokarskaya et al. 2001, Malysheva et al. 2007a, Malysheva et al. 2007b, Martirosian et al. 2003, Petrosyan et al. 2003). In the field, D. armeniaca was identified using species identification guides (Darevsky 1967, Arakelyan et al. 2011) (Fig. 3), since it has previously been shown that identification, based on visual observation, did not cause confusion between species (Petrosyan et al. 2019a, Petrosyan et al. 2019b). During our field surveys, we captured and identified 42 individuals in 24 SZs. All individuals were found to be females, determined by visual inspection of the genitals.

Figure 3.  

Distinguishing characters of D. armeniaca. A. Between the central temporal and tympanum shields are two scales of similar sizes or the central temporal shield touches the tympanic scale (indicated by arrow). B. In front of the large anal shield, there are one or two enlarged pre-anal scales of different size than the other pre-anals (indicated by arrow). Photos by I. Kropachev.

Darevskia armeniaca is widespread in the Transcaucasus in the north-west of Armenia, in the western part of Azerbaijan, in the south of Georgia and in the north-eastern part of Turkey (Petrosyan et al. 2019a, Petrosyan et al. 2019b). Species were identified in 24 (20%) SZs of 121 examined sites. In the study region, eight zones of co-existence with the “paternal” species D. valentini were revealed. The numbers of SZs, where there was сo-existence with other parthenogenetic species D. dahli, D. unisexualis and D. rostombekowi, were eleven, four and five, respectively. Hybrid individuals of D. valentini x D. armeniaca were found in three SZs (Lchashen, Kuchak and Tezh), which were previously stated in literature (Danielyan et al. 2008, Carretero et al. 2018). Our data showed that the identified SZs are located in diverse habitats, such as outcrops of rocks, large stones and clay rocks in mountain steppes, mountain meadows, mountain forests and urbanised biotopes of central and eastern parts of Armenia and southern Georgia (Fig. 1).

The parthenogenetic D. dahli lizard is of hybrid origin (Darevsky 1967, Uzzell and Darevsky 1975). The "maternal" species of D. dahli is D. mixta; "paternal" is D. portschinskii (Darevsky 1967, Uzzell and Darevsky 1975, Moritz et al. 1992, Fu et al. 1999, Murphy et al. 2000). Species identification of D. dahli from SZs was carried out by authors in different years using allozyme loci, mt-DNA, multilocus DNA fingerprinting, mini- and micro-satellite markers and morphological features (Darevsky 1966, Darevsky 1967, Uzzell and Darevsky 1975, Fu et al. 1999, Ryabinina et al. 1999, Murphy et al. 2000, Tokarskaya et al. 2001, Davoyan et al. 2007, Vergun et al. 2014, Arakelyan et al. 2011). In the field, D. dahli was identified using species identification guides (Darevsky 1967), (Fig. 4), since it has previously been shown that identification, based on visual observation, did not cause confusion between the species (Tarkhnishvili et al. 2010, Arakelyan et al. 2011). During our field survey, we captured and identified 29 individuals. All individuals were found to be females, determined by visual inspection of the genitals.

Figure 4.  

Distinguishing characters of D. dahli. A. Between the central temporal and tympanum shields in the narrowest place in the same row, there are 2-3 enlarged shields (indicated by arrow); B. In front of the large anal shield, there are always two symmetrical enlarged pre-anal ones (indicated by arrow). The differences between D. dahli and D. armeniaca also exist in number and placement of ciliar granules. Photos by I. Kropachev.

Darevskia dahli is widespread in north-eastern Armenia, western Azerbaijan and southern and central Georgia. Suitable habitats of the species in north-eastern Armenia are divided into seven vast isolated areas assigned to highland forest, meadow and steppe zones (Arakelyan et al. 2011, Petrosyan et al. 2020). During the survey, 15 SZs have been identified, 12 of which being located in the two north-western Provinces of Tavush and Lori (Arakelyan et al. 2011). Darevskia dahli co-exists in three SZs with a "paternal" species D. portschinskii and with D. armeniaca, D. rostombekowi and D. unisexualis in eleven, three and two SZs, respectively. Identified SZs are located in various biotopes: mountain forests, mountain meadows, mountain steppes and urbanised biotopes.

The parthenogenetic lizard D. rostombekowi has a hybrid origin (Darevsky 1967, Uzzell and Darevsky 1975). The "maternal" species for D. rostombekowi is D. r. raddei and "paternal" is D. portschinskii (Darevsky 1967, Uzzell and Darevsky 1975, Murphy et al. 2000, Ryskov et al. 2017). The identification of Darevskia rostombekowi in SZs was undertaken in different years using allozyme loci, mt-DNA, multilocus DNA fingerprinting, mini- and micro-satellite markers and morphological features (Darevsky 1967, Uzzell and Darevsky 1975, Macculloch et al. 1997, Fu et al. 2000a, Murphy et al. 2000, Martirosian et al. 2002, Petrosyan et al. 2003, Arakelyan et al. 2011, Ryskov et al. 2017). In the field, D. rostombekowi was identified using species identification guides according to Darevsky (Darevsky 1967) (Fig. 5), since it has previously been shown that identification, based on visual observation, did not cause confusion between the species (Martirosian et al. 2002, Petrosyan et al. 2003, Arakelyan et al. 2011, Ryskov et al. 2017). We captured and identified 17 individuals in 10 SZs. All individuals were found to be females, determined by visual inspection of the genitals.

Figure 5.  

Distinguishing characters of D. rostombekowi. A. The central temporal shield is large, often divided into two; from the first upper temporal shield in the narrowest place, it is separated by 1-3 and, from the tympanum, by 2-4 transverse rows of enlarged shields (indicated by arrow); B. The anal shield is large, in front of it are symmetrically located four small and approximately equal in size pre-anal shields, the middle of which can be slightly increased (indicated by arrow). Photos by M. Arakelyan.

Darevskia rostombekowi has a relatively small range, consisting of several different isolated areas within northern Armenia, north-western Azerbaijan, the territory of Nagorno-Karabakh and a small alpine relict isolated area from the main range on the south-eastern coast of Lake Sevan (Macculloch et al. 1997, Arakelyan et al. 2011). Darevskia rostombekowi has at least two SZs with both parental species D. portschinskii and D. r. raddei in the two north-western provinces of Tavush and Lori in Armenia. The number of SZs, where there was co-existence with other parthenogenetic forms of D. armeniaca, D. dahli and D. unisexualis, are four, three and two, respectively. In general, SZs with other species of the genus Darevskia located in the forest zone, mountain meadows, mountain steppes and anthropogenic transformed habitats, were identified within the range of the species.

The parthenogenetic lizard of D. unisexualis is formed as a result of interspecific hybridisation between bisexual D. valentini ("paternal") and D. r. nairensis ("maternal") species (Darevsky 1966, Darevsky 1967, Uzzell and Darevsky 1975, Moritz et al. 1992, Murphy et al. 2000). Species identification in studed SZs was carried out using allozyme loci, mt-DNA, multilocus DNA fingerprinting, mini- and micro-satellite markers and morphological traits (Darevsky 1967, Uzzell and Darevsky 1975, Murphy et al. 2000, Ryskov et al. 2000, Tokarskaya et al. 2001, Ryskov et al. 2003, Tokarskaya et al. 2003, Korchagin and Tokarskaya 2010, Omel'chenko et al. 2009a, Omel'chenko et al. 2009b, Arakelyan et al. 2011). In the field, D. unisexualis was identified using species identification guides according to Darevsky (Darevsky 1967), (Fig. 6), since it has previously been shown that the identification using specific features did not cause confusion between the species (Darevsky 1967, Uzzell and Darevsky 1975, Tokarskaya et al. 2001, Tokarskaya et al. 2003, Omel'chenko et al. 2009a, Omel'chenko et al. 2009b, Arakelyan et al. 2011). We captured and identified 18 individuals in 11 SZs. All individuals were found to be females, determined by visual inspection of the genitals.

Figure 6.  

Distinguishing characters of D. unisexualis. A. The central temporal shield is large, often divided into two; from the first upper temporal shield in the narrowest place, it is separated by 1-3 and from the tympanum - by 2-4 transverse rows of enlarged shields (indicated by arrow); B. The anal shield is large, in front of it four small and approximately equal in size pre-anal shields are located symmetrically, the middle of which can be slightly increased. (indicated by arrow); C. Only in this species - the maxillary shield is in contact with the fronto-nasal (indicated by arrow). Photos by I. Kropachev.

The D. unisexualis range covers the territory of northern and central highland Armenia and the adjacent regions of eastern Turkey. In Armenia, the species is found in isolated populations in the five Provinces of Aragatsotn, Gegharkunik, Kotayk, Lori and Shirak. In these Provinces, nine SZs were identified, including four SZs in which D. unisexualis co-exists with the "maternal" species D. r. nairensis and in two SZs with the "paternal" species D. valentini. The number of SZs, where there is co-existence with other parthenogenetic species D. armeniaca, D. dahli and D. rostombekowi, are five, two and two, respectively. Hybrid individuals of D. valentini x D. unisexualis were found in two zones (Artavazd, Kuchak), which were previously noted in literature (Danielyan et al. 2008,Spangenberg et al. 2017, Carretero et al. 2018). In the SZ Artavazd, autotriploid males and sterile intersexual individuals were found from 1984-1988 (Darevsky et al. 1989). The origin of these hybrids can be explained by interclonal mating between parthenogenetic females and rare, conspecific diploid males. Identified SZs are located at rock outcrops, piles of stones and rocky slopes in the mountain-steppe zone.

The bisexual species, D. raddei, is considered as a complex (Darevskia raddei sensu lato) containing four forms (subspecies) “raddei”, “nairensis”, “vanensis” and “chaldoranensis” (Grechko et al. 2007, Rastegar-Pouyani et al. 2012, Omelchenko et al. 2016, Freitas et al. 2016). However, the subspecies “raddei” (D. r. raddei) is the only form widely distributed in Armenia, Nagorno-Karabakh, Azerbaijan and the northern province of Ardabil of Iran (Freitas et al. 2016). Identification of D. r. raddei individuals in different SZs was undertaken by authors using allozyme markers, mt-DNA, multilocus DNA fingerprinting, mini- and micro-satellite markers and morphological traits (Bobyn et al. 1996, Fu et al. 2000a, Grechko et al. 2007, Korchagin et al. 2007, Korchagin and Tokarskaya 2010, Omelchenko et al. 2016, Freitas et al. 2016). In the field, D. r. raddei was identified using identification guides according to Darevsky (Darevsky 1967) (Fig. 7). During the field survey, we captured and identified 11 individuals, including nine adult females and two males.

Figure 7.  

Distinguishing characters of D. r. raddei (female individual on the photographs). A. Between the average size central temporal and small tympanum shield in the narrowest place, there are 2-5 small shields (indicated by arrow); B. The anal shield is large, in front of it, there are two more or less enlarged pre-anals located symmetrically, between which a small third one is often wedged (indicated by arrow). Photos by I. Kropachev.

The bisexual form of D. r. raddei is widespread in southern Armenia, north-eastern Turkey, north-western regions of Nagorno-Karabakh, south and south-western Azerbaijan and in adjacent regions of north-western Iran (Darevsky 1967). During the field survey, five SZs were identified with other parthenogenetic and bisexual species in isolated populations in north-eastern Armenia and Nagorno-Karabakh. Three SZs exist in north-eastern Armenia in the Tavush Province and two zones in Nagorno-Karabakh (Shahumyan Province). This species in the two SZs co-exists with the bisexual species D. portschinskii and in two SZs with the "daughter" species, D. rostombekowi. The identified SZs are located in rocky habitats in forest and mountain steppes, including the stone walls of buildings (e.g. on the walls of churches) and stone ruins.

Although early studies found very low genetic differences between D. r. raddei and D. r. nairensis (Bobyn et al. 1996, Fu et al. 2000a), later studies (Arnold et al. 2007Grechko et al. 2007, Omelchenko et al. 2016), however, considered them as two subspecies. Darevskia r. raddei and D. r. nairensis were considered as different subspecies, since they are reproductively isolated in terms of distinctive periods of mating (Danielyan 1989). In the field, D. r. nairensis was identified using identification guides according to Darevsky (Darevsky 1967), (Fig. 8). During the field survey, we captured and identified 13 individuals, including ten adult females and three males.

Figure 8.  

Distinguishing characters of D. r. nairensis (female individual on the photographs). A. Central temporal shield is small. Between the central temporal and rather large tympanic shields on the sides of the head, there are two or three enlarged shields (indicated by arrow); B. In front of a large, elongated transverse anal shield, there are two large pre-anal ones (indicated by arrow). Photos by I. Kropachev.

The range of D. r. nairensis covers central, south-western Armenia, the western shore of Lake Sevan and is restricted to the north-eastern part of Armenia. This species also inhabits neighbouring regions of north-eastern Turkey and southern Georgia. In Armenia, there are several isolated populations in the north Provinces (Tumanyan and Lori) and in the south-central part (Karmrashen and Vayots Dzor Provinces). During the field survey, six SZs were identified in the three Provinces of Gegharkunik, Kotayk and Shirak. This species has a relatively large number of SZs - four with a "daughter" species D. unisexualis and, with other parthenogenetic species, D. armeniaca and D. dahli, there are two and one SZs, respectively. Our data show that D. r. nairensis has only one SZ with a bisexual species D. valentini. The SZs are mainly located in the rocky habitats of the mountain steppe.