Species list of Amphibians and Reptiles from Turkey

Turkey is biogeographically diverse and consequently has a rich herpetofauna. As a result of active herpetological research, the number of species has steadily increased in recent years. I present here a new checklist of amphibian and reptile species distributed in Turkey, revising the nomenclature to reflect the latest taxonomic knowledge. In addition, information about the systematics of many species is also given. In total 35 (19.4%) amphibian and 145 (80.6%) reptile species comprise the Turkish herpetofauna. Among amphibians, 16 (45.7%) anurans and 19 urodelans (54.3%) are present. Among reptiles, 11 (7.6%) testudines, 71 (49%) saurians, 3 (2.1%) amphisbaenians and 60 (41.3%) ophidians are considered part of the herpetofauna. The endemism rate in Turkey is considered relatively high with a total of 34 species (12 amphibian species – 34.3% and 22 reptile species – 15.2%) endemic to Turkey, yielding a total herpetofaunal endemism of 18.9%. While 38 species have not been threat-assessed by the IUCN, 92 of the 180 Turkish herpetofaunal species are of Least Concern (LC), 13 are Near Threatened (NT), 10 are Vulnerable (VU), 14 are Endangered (EN), and 7 are Critically Endangered (CR). In addition, 6 species are in the DD (Data Deficient) category.


Introduction
Turkey lies near the intersection of Asia, Europe, and Africa, which contributes to its rich biodiversity (Şekercioğlu et al., 2011;Gür, 2016;Tavşanoğlu, 2016). A key factor, with an important influence on species diversity, is that the country includes significant parts of three biodiversity hotspots, namely, the Mediterranean, Iran-Turan and Caucasian (Mittermeier et al., 2004). Turkey also straddles two major geographical areas (Euro-Siberian and Eastern Mediterranean) from a herpetological perspective (Ficetola et al., 2018). In addition, the isolation created by the Anatolian diagonal, formed by high mountain ranges, and by the mountains separating the northeast and the south (west) play a crucial role in increasing this diversity (Rokas et al., 2003;Gündüz et al., 2007;Mutun, 2010;Vamberger et al., 2013;Korkmaz et al., 2014).
The Turkish herpetofauna is rich, with 180 verified species, or about 60% of the total present in the entire European continent (Speybroeck et al., 2020).
Thirty-five of these 180 species are amphibians and the rest are reptiles (Frost, 2020;Uetz et al., 2020; https://amphibiaweb.org; https://www.lacerta.de). The herpetofaunal list is frequently updated as a result of new expeditions and/or local faunistic surveys and phylogenetic studies.
Herpetological studies in Turkey date back to Linnaeus (1758) with the description of Hemidactylus turcicus (Linnaeus, 1758) and have continued for more than a quarter of a millennium. Surveys were carried out during the first half of the 20th century by herpetologists such as Werner (1902), Bodenheimer (1944) and Mertens (1952), and checklists of Turkish amphibians and reptiles were generated by these initiatives, yielding a list of 85 species. In the following years, the number of amphibian and reptile species in Turkey has almost doubled through the activities of new researchers and their expeditions. Subsequently, separate species lists have been created for amphibians (Başoğlu and Özeti, 1973) and reptiles (Başoğlu and Baran, 1977;1980;Sindaco et al., 2000); the most comprehensive list being that of Baran et al. (2012), including 157 species. However, this list needs to be updated based on the latest explorations and output of biogeographic studies. At the same time, names of many species have changed in accordance with recent molecular-based systematic evaluations. Thus, there is a need for an updated source that provides the currently recognized names of the Turkish herpetofaunal species.
The fundamental aims of this study are: i) to give information about recent changes to the taxonomic status of amphibian and reptile species of Turkey, and ii) to bring together an authoritative list of all amphibians and reptiles in Turkey in one available resource.

Material and Methods
The area considered in this study lies within the political boundaries of Turkey, between 26-45º Eastern Longitudes and 36-42º Northern Latitudes, including the Anatolian and Thracian Peninsulas of Turkey (Fig. 1). A list of all the species in the herpetofauna of Turkey was compiled and evaluated in light of the current literature. Data on taxonomically relevant variation (subspecies), endemism status, chorotypes, and IUCN status (DD-Data Deficient; LC-Least Concern; NT-Near Threatened; VU-Vulnurable; EN-Endangered; CR-Critically Endangered) of all the listed species in Turkey were assembled. The chorotypes and IUCN status of the species are given according to the most recent global assessments (Sindaco et al., 2000(Sindaco et al., , 2013IUCN, 2020).

Results and Discussion
Amphibians Thirty-five amphibian species are distributed in Turkey, comprising 16 (45.7%) in the Order Anura and 19 (54.3%) in the Order Urodela. The endemism rate in amphibian species in Turkey is very high, with 12 amphibian species (34.3%) unique to Turkey.
The genus Bombina Oken, 1816 was previously represented by only a single species, B. bombina (Linnaeus, 1761). A second species, B. variegata (Linnaeus, 1758) was recently reported in Turkey by Bülbül et al. (2016). Although the first record of the species in Turkey was given by Boulenger (1897), he did not provide any information about its exact locality. It was more than a century before locality information in Turkey was obtained. The first precise locality record was given by Bülbül et al. (2016) in Enez, Edirne (Lake Gala) and the species has subsequently been recorded from two more localities in Enez by Bülbül et al. (2018).
Bufo bufo (Linnaeus, 1758) and Bufo verrucosissimus (Pallas, 1814) are two toad species distributed in Turkey. In the literature, it has been stated that the distribution boundaries and the taxonomic status of these two taxa are still uncertain. Although García-Porta et al. (2012) reported that these taxa were two subspecies of B. bufo, Arntzen et al. (2013) rejected this proposal because they downgraded B. verrucosissimus to the subspecies level of B. bufo. According to a recent study based on phylogenetic data, two main clades of common toads are living in Turkey, and morphological data is also compatible with this phylogeny (Özdemir et al., 2020). On this basis, it is accepted that these two taxa should be evaluated as two separate species.
Until 2009, the genus Lissotriton Bell, 1839 was represented in Turkey by a single species, Lissotriton vulgaris (Linnaeus, 1758) with three subspecies (L. v. kosswigi, L. v. lantzi and L. v. schmidtleri). Dubois and Raffaëlli (2009) discussed their reasoning for recognizing Lissotriton vulgaris kosswigi and L. v. lantzi as distinct from L. vulgaris and they have since been considered as different species. Lissotriton lantzi (Wolterstorff, 1914) was accepted on the Turkish amphibian list, because there was a historic locality record in Artvin from Louis Amédée Lantz in 1911. These specimens are still extant as ZISP (Zoological Museum in St. Petersburg) 3187 (Skorinov et al., 2014). Wielstra et al. (2015) suggested that Lissotriton schmidtleri (Raxworthy, 1988) might be a distinct species and discussed its allopatric distribution relative to Lissotriton kosswigi (Freytag, 1955). The latest phylogenetic study has demonstrated that L. schmidtleri is a distinct species (Pabijan et al., 2017). As a result of this taxonomic study, it has been revealed that three different species, belonging to the genus Lissotriton, exist in Turkey.
The genus Neurergus Cope, 1862 is represented by three species in Turkey, Neurergus barani (Öz, 1994), N. crocatus Cope, 1862 andN. strauchii (Steindachner, 1887). However, N. strauchii was classified as a subspecies of N. crocatus for a long time by different authors (Schmidt, 1939;Bodenheimmer, 1944;Başoğlu and Özeti, 1973). Schmidtler and Schmidtler (1975) reported that N. crocatus and N. strauchii were two distinct species in terms of morphological characters. Two subspecies of N. strauchii have been recognized (N. s. barani Öz, 1994 andN. s. munzurensis Olgun, Avcı, Bozkurt, Üzüm, Olgun, and (Öz, 1994;Olgun et al., 2016). The former was treated as a subspecies in the first phylogenetic study following its description (Özdemir et al., 2009), however, Rancilhac et al. (2019) Litvinchuk, Zuiderwijk, Borkin, and Rosanov, 2005. Ommatotriton nesterovi was initially recognized as a distinct species from O. ophryticus, based on morphological data and mitochondrial DNA by Bülbül and Kutrup (2013). Afterwards, this study was expanded, validated, and more precisely documented by Van Riemsdijk et al. (2017). As a result of these studies, it was concluded that there are three different species of Ommatotriton in Turkey. Within these species, O. nesterovi is distributed in southern Anatolia, whereas the other two species occur in northern Anatolia (van Riemsdijk et al., 2017). It is thought that the Kızılırmak River is a geographical barrier between the two species living in the north (Bülbül and Kutrup, 2013).
Genus Triturus Rafinesque, 1815 was represented in Turkey by only one species, Triturus karelinii (Strauch, 1870) until recently. Wielstra et al. (2012) proposed that T. karelinii should be divided into three mitochondrial species groups (western, central, and eastern). As a result, Wielstra et al. (2013) described a morphologically distinct new species as Triturus ivanbureschi Arntzen and Wielstra, 2013 in the western T. karelinii group. Later, Wielstra and Arntzen (2016) described a new species as Triturus anatolicus Wielstra and Arntzen (2016) in the central T. karelinii group. Although Triturus karelinii was separated into three different species as the result of recent phylogenetic studies, the occurence of only two species (T. anatolicus and T. ivanbureschi) was reported from Turkey (Wielstra et al., 2012;Wielstra and Arntzen, 2016). Therefore, T. karelini sensu stricto is not currently verified as being included in the Turkish herpetofauna.
Based on morphological data, three Testudo species are found in Turkey: Testudo graeca Linnaeus, 1758, T. hermanni Gmelin, 1789, and T. perses Perälä, 2002(Baran et al., 2012. Although T. graeca and T. hermanni have been known for a long time in Turkey, T. perses was recorded in 2004 from Hakkari for the first time (Türkozan et al., 2004). In studies including all populations of T. graeca distributed in Turkey, DNA samples were examined phylogenetically, and four clades were identified, T. g. armeniaca Chkhikvadze and Bakradze 1991, T. g. buxtoni Boulenger 1921, T. g. ibera Pallas 1814, and T. g. terrestris Forskål 1775(Fritz et al., 2007Türkozan et al., 2018). In addition, the authors indicated that the taxon previously reported as T. perses was a member of the buxtoni group. For this reason, T. perses is here considered as a synonym of T. g. buxtoni subspecies (Fritz et al., 2007;Türkozan et al., 2018). Thus, only two species (T. graeca and T. hermanni) need to be addressed in the species list of Turkey.
The origin of Trachyemys scripta (Thunberg in Schoepff, 1792) is the American continent. It is a species that came to Turkey with the pet trade, was released to nature by irresponsible owners, and has started to breed in inland waters (Çiçek and Ayaz, 2015). Trachyemys scripta can compete with native species in terms of food sources and this invasive exotic is increasing in population size and expanding its distribution range. It was already considered as one of the worst 100 alien invasive species by the IUCN (IUCN, 2020). Reproduction in a population of T. scripta in the wild was reported from southern Anatolia (Anamur, Mersin) for the first time (Çiçek and Ayaz, 2015), and because of this, this invasive reptile species was added to the species list.
The Family Agamidae is represented by four genera and species in Turkey. The taxonomic status of the genera Stellagama Baig, Wagner, Anajeva and Böhme, 2012 and Paralaudakia Baig, Wagner, Anajeva and Böhme, 2012, previously included within the genus Laudakia Gray, 1845, remains controversial. One of the reasons for this controversy is that the results of a phylogenetic study based on the mitochondrial genome, reported that Laudakia is paraphyletic (Macey et al., 2000). However, other studies, based on both the mitochondrial and nuclear genes, concluded that Laudakia is monophyletic (Melville et al., 2009;Edwards and Melville, 2011). Following these studies, Baig et al. (2012) conducted a morphologically-based study and as a result, Laudakia was divided into three different genera based on morphology: Laudakia, Paralaudakia and Stellagama. Although Pyron et al. (2013) confirmed that Laudakia is monophyletic in their study using a super matrix approach, Stellagama and Paralaudakia are still being used by many herpetologists since 2012. The reptile-database still uses these two names (Uetz et al., 2020). However, Speybroeck et al. (2020) do not follow the split of Laudakia and they accepted it as monophyletic. The genera Stellagama and Paralaudakia continue to accepted, within the scope of the current checklist, until this controversial situation is completely resolved.
Anatololacerta Arnold, Arribas and Carranza, 2007 was represented with three species until recently, based on morphological data (Baran et al., 2012). However, a recent phylogenetic study by Bellati et al. (2015) revealed that Anatololacerta was separated into five different clades, represented by four named species: Anatololacerta anatolica (Werner, 1900), A. budaki (Eiselt and Schmidtler, 1987), A. danfordi (Günther, 1876), and A. pelasgiana (Mertens, 1959), as well as an undescribed species. In addition, the results of Bellati et al. (2015) suggested that A. oertzeni (Werner, 1904) should be considered as a subspecies of A. anatolica rather than a species. Although Darevskia mixta (Méhely, 1909) was recorded by Baran and Atatür (1998) for the first time from Turkey, Gabelaia et al. (2015) reported that previous records from Turkey may have been wrong. The species has only spread in Georgia, and they reported that no D. mixta were found in the previous reported localities in Turkey during their field studies 2007-2013 ( Gabelaia et al., 2015). However, D. mixta was reported from Turkey in the studies of Tuniyev et al. (2014) and Freitas et al. (2019). Also, Freitas et al. (2019) used DNA samples of D. mixta from the eastern Black Sea region in Turkey in their study. Based on this data, it was decided to include D. mixta in the species list, however, it is necessary to explore the new localities from Turkey and to evaluate its morphology.
According to the literature, members of the genus Iranolacerta Arnold, Arribas and Carranza, 2007 were not recorded in Turkey until 2015 (Baran and Atatür, 1998;Sindaco et al., 2000;Baran et al., 2012). Iranolacerta brandtii (De Filippi, 1863) was only known from Iran until recently. The first record of the species from Turkey was given in two independent studies in 2015 (Avcı et al., 2015a;Yıldız and İğci, 2015).
The Anatolian taxon Lacerta pamphylica Schmidtler, 1975, has been classified as L. trilineata Bedriaga, 1886 (Godinho et al., 2005;Ahmadzadeh et al., 2013;Sagonas et al., 2014). More recently, analyses of SNPs and mitochondrial sequences by Kornilios et al. (2019;2020) yielded a sister-group relationship between L. pamphylica and the eastern Aegean populations of L. trilineata. This led to the identification of four specieslevel units: L. trilineata, L. pamphylica, L. citrovittata Werner, 1938, andL. diplochondrodes Wettstein, 1952. Lacerta diplochondroles was earlier known as a subspecies of L. trilineata and the same studies that proposed L. diplochondroles as a distinct species revealed that L trilineata sensu stricto do not occur in Turkey (Kornilios et al. 2019;2020). Lacerta agilis was represented by two subspecies (L. a. brevicaudata ann L. a. grusinica) in Turkey. However, the latest phylogenetic study concluded that they are synonymous with L. a. exigua (Andres et al., 2014). Although there is a difference at the subspecies level based on morphology, this has not manifested itself at the molecular level. So, all L. agilis samples distributed in Turkey are treated as L. a. exigua in the scope of this study.
Mesalina microlepis (Angel, 1936) was recorded for the first time from Akçakale, Şanlıurfa, Southern Anatolia as M. breviostris Blanford, 1874 (Kumlutaş et al., 2002a;2002b). According to a recent phylogenetic study, Mesalina populations distributed in Turkey were revealed to be the same species as that distributed to the south of Turkey (Syria, Lebanon and Jordan) (Šmíd et al., 2017). Therefore, Turkish populations of Mesalina are treated as M. microlepis in the current study.
Timon kurdistanicus (Suchow, 1936) was previously known as a subspecies of T. princeps (Blanford, 1874). A recent phylogenomic study by Ahmadzadeh et al. (2012) proposed that T. kurdistanicus is a full species based on the high genetic distance from T. princeps.
Recent studies changed the taxonomic position of some scincid species in Turkey. For instance, Bozkurt and Olgun (2020) reported that Ablepharus bivittatus has highly divergent genetic and morphological characteristics compared to the others, and it shares similar morphological characteristics with the genus Asymblepharus, with which they share an elliptical tympanum, a hidden upper eye opening under 3 or 4 large shields, light and dark longitudinal stripes on the back, and adpressed hind-limbs reaching to knees, and well-developed limbs with five toes. I follow Bozkurt and Olgun (2020) in transferring Ablepharus bivittatus to Asymblepharus. These authors also proposed that Ablepharus budaki anatolicus Schmidtler 1997 is a distinct species in terms of phylogenetic and morphological traits from the nominate form of A. budaki (Bozkurt and Olgun, 2020). This taxonomic outcome supports the results of the study of Skourtanioti et al. (2016).
Until recently, the Middle Eastern mabuyine species, Heremites auratus, H. septemtaeniatus and H. vittatus, were considered to belong to the genus Trachylepis Fitzinger, 1843. Karin et al. (2016) explained that the Middle Eastern species group formed a phylogenetically distinct clade from the African species group, for which Heremites Gray, 1945 was available (Karin et al., 2016). Also, Ophiomorus kardesi was known as O. punctatissimus (Bibron and Bory de Saint-Vincent, 1833) in Turkey until recently. According to a recent phylogenetic and morphological study, Ophiomorus populations distributed in Turkey were revealed to be a distinct species from O. punctatissimus (Kornilios et al., 2018).

Considerable differences within the genus Anguis
Linnaeus, 1758 were reported by means of genetic and morphological analyses (Cabela and Grillitsch, 1989;Gvoždík et al., 2010). This led to the conclusion that there are four different Anguis species in Europe (Gvoždík et al., 2010). Anguis colchica was known as a subspecies of A. fragilis Linnaeus, 1758, until recently. According to recent phylogenetic studies, A. colchica (Nordmann, 1840) individuals were revealed as a distinct species from A. fragilis (Gvoždík et al., 2010;Gvoždík et al., 2013;Jablonski et al., 2016).
The genus Blanus Wagler, 1830 was represented by three subspecies within one species in Turkey until recently. However, a recent phylogenetic and morphological study revealed that the genus was separated into three different species, namely Blanus alexandri Sindaco, Kornilios, Sacchi and Lymberakis, 2014, B. aporus Werner, 1898, and B. strauchi (Bedriaga, 1884 (Sindaco et al., 2014). A recent study based on ecological niche divergence has shown that these three taxa are also separable in terms of niche (Şahin et al., 2021).
The genus Eirenis Jan, 1863, represented by 14 species in Turkey, has gone through many revisions (Schmidtler, 1993;1997b;Nagy et al., 2003;Sivan and Werner, 2003). As a result of the revision by Nagy et al. (2003), the species previously known as Eirenis coronella (Schlegel, 1837), and distributed in Turkey, has been changed to E. coronelloides (Jan, 1862). Eirenis hakkariensis Schmidtler and Eiselt, 1991 was known as a subspecies of E. thospitis Schmidtler and Lanza, 1990 based on mitochondrial and nuclear genes, until recently (Nagy et al., 2003). However, a recent morphological study revealed that E. hakkariensis is a different species from E. thospitis based on external morphology, having a higher number of teeth and a different body pattern (Mahlow et al., 2013). Populations of Eirenis persicus in Turkey were represented by one taxon until a recent morphological, molecular, and ecological study revealed that some populations represent a different species (Rajabizadeh et al., 2015). The new taxon was described as E. occidentalis by Rajabizadeh, Nagy, Adriaens, Avci, Masroor, Schmidtler, Nazarov, Esmaeili and Christiaens, 2015. In addition, the populations of E. persicus distributed in Turkey should be referred to E. occidentalis (Rajabizadeh et al., 2015).
Elaphe was represented by a single species, Elaphe sauromates (Pallas, 1811) in Turkey until 2004. The first record of the second species, Elaphe dione (Pallas, 1773), was given by Garzoni and Geniez (2004) based on three speciemens from north-eastern Turkey, although there have been no more recent records. The third species, Elaphe urartica Jablonski, Kukushkin, Avci, Bunyatova, Ilgaz, Tuniyev, and Jandzik, 2019, named for the ancient Kingdom of Uratuu, was newly described from among eastern populations of E. sauromates and is discriminated based on molecular phylogenetic and morphological data .
The genus Rhynchocalamus Günther, 1864, until recently comprised three species known from Turkey. One of them, Rhynchocalamus barani, shows different morphological features from the other two . Osteological and phylogenetic results of Avcı et al. (2015b) revealed that R. barani should be placed in its own genus as Muhtarophis barani (Olgun, Avci, Ilgaz, Üzüm and Yilmaz, 2007). However, results of the study of Rajabizadeh et al. (2020) strongly support a sister-group relationship of Muhtarophis Avcı, Ilgaz, Rajabizadeh, Yılmaz, Üzüm, Adriaens, Kumlutaş and Olgun, 2015 and Scaphiophis. According to Rajabizadeh et al. (2020), they form a clade that is the sister group to all the above genera of the Western Palearctic and South Asian colubrids. Since this situation has not yet been resolved, I included barani in the genus Muhtarophis in this study. The genus Natrix Laurenti, 1768 has a broad distribution in Turkey, with three species currently recorded. One of these, Natrix megalocephala (Orlov and Tuniyev, 1987), based on morphological data, has Journal of Animal Diversity (2020), 2 (4): 10-32 | www.jad.lu.ac.ir been considered as a full species (Orlov and Tuniyev, 1987). However, according to a recent phylogenetic study, N. megalocephala is a synonym of N. natrix (Linnaeus, 1758) (Kindler et al., 2013) and it has not been included in the species list for Turkey.
Vipers have an enormous diversity in Turkey, represented by Daboia Gray, 1842, Macrovipera Reuss, 1927, Montivipera Nilson, Tuniyev, Andren, Orlov, Joger, and Herrmann, 1999, and Vipera Laurenti, 1768. The subgenus Pelias Merrem, 1820 was nested within Vipera, however this has been used as a full genus in some studies (Avcı et al., 2010;Tuniyev et al., 2012;. Daboia has one species distributed in Turkey, Daboia palaestinae (Werner, 1938) first recorded from Hatay Province, its only locality in Turkey (Göçmen et al., 2018). Also, Macrovipera has one species, M. lebetinus (Linnaeus, 1758), represented within Turkey. Montivipera is a complex. Although it was represented by five species, there is also the possibility of more species being recognized in the near future. According to the study of Stümpel et al. (2016), Montivipera xanthina (Gray, 1849) has four lineages at the species level, but they did not clarify the morphology of these lineages. In addition, same authors reported that Montivipera albizona (Nilson, Andren and Flärdh, 1990) was phylogenetically within Montivipera bulgardaghica (Nilson and Andren, 1985) (Stümpel and Joger, 2009;Stümpel et al., 2016;Freitas et al., 2020). In this study, M. albizona is considered as a full species. Because this taxon has been evaluated only phylogenetically and it is also important to know about reproductive isolation and the ecological niches of the species, I have adopted a conservative approach in this instance.
In order to clarify this situation, more DNA sequences, including both mitochondrial and nuclear genomes are needed. Until this complicated situation is rectified, P. barani and P. olguni will be treated at the species level. Another reason why P. barani is treated at the species level is that P. berus shows a very different geography; it is only found in Anatolia and therefore allopatric with respect to all other P. berus populations. However, future studies may resolve the situation more clearly. The same situation is valid for the ammodytes-transcaucasiana complex reported by Freitas et al. (2020). Boulenger, 1913 was known as a full species until 2008 when Ursenbacher et al. (2008) reported that, based on their phylogeny, it did not differ from V. ammodytes (Linnaeus, 1758). However, the study required more samples from Turkey, and pending more comprehensive results, two separate species are recognized within the scope of this study.

Vipera transcaucasiana
The Blindsnakes are represented by three different species in Turkey. The genus previously known as Typhlops Oppel, 1811 displays high diversity. Hedges et al. (2014) described a new genus, Xerotyphlops Hedges, Marion, Lipp, Marin, and Vidal, 2014, with the species X. vermicularis as its Turkish representative. The type specimen of Myriopholis macroryncha housed in the Milan museum was lost during World War II (Sindaco et al., 2013). The position of this taxon in Africa is still uncertain because almost all of the examples that were once included in this taxon have been attributed to other taxa. So, the original descriptions and drawings of the type examples are included in the species Leptotyphlops cairi (Trape, 2002). Therefore, Broadley and Wallach (2007) suggested that more studies are needed to determine the taxonomic status of Middle Eastern populations. However, within the scope of this study, this taxon is treated as M. macroryncha.