Ablepharus species

4.2. On fused eyelids Ablepharus species distribution

The morphology of the hemipenes and the locality records of some of the examined specimens raise questions regarding the validity of taxonomic assignment of some literature records and, as a result, the known range of these taxa. The general distribution of the genus was presented by Karamiani et al. (2021b), prior to the synonymisation of Asymblepharus with Ablepharus by Mirza et al. (2022). Mapping all published literature records reveals a rather mosaic and scattered distribution of several taxa, as well as potentially overlapping ranges particularly in the region of the Anatolian coast of the Mediterranean (for the “ A. kitaibelii group”) ( Fig. 5 View Fig ). Although interdigitating ranges cannot be dismissed, the presence of so many taxa barely discernible by external morphology in a single region appears odd. We can even speculate that some of the available DNA sequences have originated from incorrectly identified specimens, which can explain the present distribution of these taxa.

4.2.1. Distribution of the “ А. kitaibelii group ” ( Fig. 5 View Fig )

These species occur from southern Slovakia and Hungary (north), across the Balkans (all countries), to Israel and Egypt (south), and to Azerbaijan and western Iran (east) ( Darevsky, 1953; Sofradˇzija, 1978;

Bruno, 1989; Gruber, 1981; Schmidtler, 1997; Haxhiu, 1998; Ljubisavljevic et al., 2002; Modry et al., 2004; Korsos´et al., 2008; Valakos et al., 2008; Szov ¨´enyi and Jeli´c, 2011; Stojanov et al., 2011; Cog˘alniceanu et al., 2013; Roll et al., 2013; Sterijovski et al., 2014; Vergilov et al., 2016; Handal et al., 2016; Karamiani et al., 2018). Ablepharus chernovi is a species that is distributed in northern and central Anatolia, Georgia, Armenia, Iran, and northernmost Syria ( Darevsky, 1953; Schmidtler, 1997; Skourtanioti et al., 2016; Karamiani et al., 2018). The holotype of the nominate subspecies is from Armenia ( Darevsky, 1953) but it is also distributed in West Iran ( Karamiani et al., 2018) and in several Turkish provinces, such as Adana, Hatay, Yozgat and Erzurum (see Schmidtler, 1997). Skourtanioti et al. (2016) reports the presence of A. chernovi close to the Sea of Galilee in Syria, which appears far away in the south. Its western distribution (in Anatolia) is reaching the eastern parts of the distribution of another morphologically similar species – A. kitaibelii ( Schmidtler, 1997) . The subspecies A. chernovi ressli was described from the region of Çamlıyayla (=Namrun), north of Mersin, Turkey, not far from another described subspecies A. ch. isauriensis, which is known from the region of Karaman, Turkey ( Schmidtler, 1997). We should also mention that the subspecies A. b. anatolicus is also known from localities in Namrun and north of Mersin, including the paratype material of this taxon ( Schmidtler, 1997) ( Fig. 5 View Fig ). Ablepharus kitaibelii and A. budaki are species with close and partially overlapping ranges (see Schmidtler, 1997; Skourtanioti et al., 2016). The general distribution of the A. kitaibelii subspecies covers most of the Balkans, West Anatolia, most of the Greek Islands, and also includes sporadic locations in Central Anatolia ( Vergilov et al., 2016). The distribution in the southern mainland of Anatolia reaches as far as Fethye ( Turkey) ( Schmidtler, 1997). The morphological study of Ljubisavljevic et al. (2002) revealed the presence and intergradient zones (from north to south) of A. kitaibelii fitzingeri , A. k. stepaneki and A. k. kitaibelii in the north-western parts of these subspecies’ range. According to published records ( Schmidtler, 1997; Poulakakis et al., 2013; Ozkan ¨ et al., 2019), A. budaki occurs along the coast of the northeastern Mediterranean Sea. At first, no range overlap was evident, however, specimens of A. budaki nested deeply within the A. kitaibelii range and the presence of a specimen from North Turkey (Golcük ¨) ( Fig. 5 View Fig ) hints towards the fact that the actual distribution of the two species may differ from their published ranges. As previously mentioned, the lack of sufficient distributional data and the similarity in external morphology between the two taxa may be a reason for incorrect interpretations of the general distribution and habitat preferences of these taxa. Morphologically, A. budaki and A. kitaibelii are well defined, and hemipenial morphology provides good verification of the taxa ( Vergilov et al., 2017). According to Schmidtler (1997), the subspecies A. b. anatolicus is distributed from Fethye and the region of Kaş, along the Mediterranean, to Adana. Ablepharus b. budaki (described from Cyprus) occurs in the region of West Syria and southeastern Anatolia, reaching the southern distribution area of A. ch. chernovi ( Schmidtler, 1997; Skourtanioti et al., 2016). In general, a glance at the distribution of these taxa reveals a highly mosaic distribution of the group in South and South-East Anatolia. Ablepharus rueppellii is distributed in South Syria, Lebanon, Israel, northeastern Egypt, and West Jordan (see Uetz et al., 2023). The subspecies A. rueppelli festae was described from Jordan but based on the published literature it is also present in Syria and Lebanon, whereas the nominate subspecies has been reported from Israel (and Palestine), Egypt, and possibly Lebanon ( Schmidtler, 1997; Roll et al., 2013; Handal et al., 2016; Uetz et al., 2023). These patterns raise a question regarding the exact distribution of both taxa as apparently both subspecies may co-occur in northern areas. Another morphologically similar taxon, A. budaki , is present in Syria (as well as Lebanon) ( Schmidtler, 1997; Poulakakis et al., 2013). Roll et al. (2013) synthesised the distribution of the species A. rueppellii in Israel. The current general distribution of both subspecies A. r. rueppellii and A. r. festae remains unclear.

4.2.2. Distribution of the “ А. pannonicus group ” ( Fig. 6 View Fig )

The group is distributed from eastern Turkey, Armenia, Syria, Iraq, Iran, Azerbeijdjan, Afghanistan, Tajikistan, Kazakhstan, Kyrgyzstan, Uzbekistan, Turkmenistan, to Pakistan and West India ( Jeriomtshenko and Szczerbak, 1986; Karamiani et al., 2021b). The distribution of A. bivittatus was modelled in northern Iran based on literature data ( Sanchooli, 2016). The species also was reported from Eastern Turkey ( Ilgaz et al., 2007; Bozkurt and Olgun, 2020), Azerbaijan ( Kidov and Kondratova, 2021), and Armenia ( Darevsky, 1953; Karamiani et al., 2021a). A. lindbergi was described from Afghanistan based on specimens from two localities ( Jeriomtshenko and Szczerbak, 1980) but its distribution in the country was recently expanded ( Wagner et al., 2016; Jablonski et al., 2019). A. darvazi was described by Jeriomtshenko and Panfilov (1990) from the mountains of Tajikistan. This species is nestled inside the range of A. pannonicus , another species known from mountainous areas ( Fig. 6 View Fig ). Ablepharus pannonicus has a large distribution, occurring from Syria, Iraq, Iran, Afghanistan, Pakistan, South Tajikistan to the border of northwestern India ( Jeriomtshenko and Szczerbak, 1986). Khan (2002) gave the following distribution (general and for Pakistan) for A. pannonicus : “ Reported from around Quetta, Waziristan hills, Chitral and the Salt Range. Ranges from the Arabian Peninsula and North Arabian Desert, through Iran to circum-Mediterranean region, Tadzhikstan and Afghanistan ”. This species is also mentioned for parts of western India ( Smith, 1935). Karamiani et al. (2019) constructed a model of the present and past distribution of A. pannonicus and A. grayanus in Iran. Based on habitat preferences, the models revealed that A. pannonicus inhabits montane regions with sparse annual grasses and shrubs (i.e., drier areas), while A. grayanus prefers wetter habitats, including palm groves and areas near rivers. According to the models and locations of A. pannonicus and A. bivittatus (by Karamiani et al., 2019; Sanchooli, 2016), it appears that the ranges of these two species overlap in northern Iran. The true distribution of A. grayanus has been poorly investigated with recent data provided only for the Iranian populations ( Karamiani et al., 2019). This species’ distribution also includes northern India and South Pakistan ( Jeriomtshenko and Szczerbak, 1986). In fact, this species was described from Kachh, West India, and later the knowledge of its distribution expands in other, western parts, of the country ( Smith, 1935; Vyas, 2011). In Pakistan, A. grayanus occurs in lowland grasslands throughout the country but also was recorded at 1800 m elevation ( Khan, 2002). According to Jeriomtshenko and Szczerbak (1986), A. deserti is autochthonous to the semi-desert areas of Middle Asia, occurring from South Kazakhstan, Kyrgyzstan, Uzbekistan, to the northern parts of Tajikistan and East Turkmenistan. In northern Tajikistan, the ranges of A. deserti and A. pannonicus partially overlap. Given their similar morphology, the exact species identity of the individuals in these areas appears doubtful and should be interpreted with caution. Based on literature and new locality data, Dujsebayeva (2015) provided a more accurate distribution of A. deserti in southern Kazakhstan.