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Elaphe urartica JABLONSKI, KUKUSHKIN, AVCI, BUNYATOVA, ILGAZ, TUNIYEV ET JANDZIK, 2019

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Higher TaxaColubridae, Colubrinae, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes) 
Subspecies 
Common NamesE: Urartian Rat Snake 
SynonymElaphe urartica JABLONSKI, KUKUSHKIN, AVCI, BUNYATOVA, ILGAZ, TUNIYEV ET JANDZIK 2019
Coluber sp. (No. 15) — HOHENACKER 1831: 374
Coluber sp. (No. 18) — HOHENACKER, 1831: 375
C. cereus — DWIGUBSKY 1832: 27, (nomen dubium)
C. fulvus — DWIGUBSKY 1832: 28, (nomen dubium)
C. taeniothys FISCHER (nomen dubium) 1832: 575
C. taeniothys — HOHENACKER 1837: 145
Tropidonotus sauromates EICHWALD 1841: 140
Elaphis sauromates — DUMÉRIL, 1853: 453 (part.)
Elaphis sauromates — STRAUCH, 1873: 92 (part.)
C. quatuorlineata sauromates — BOULENGER 1894: 47 (part.)
C. dione var. sauromates NIKOLSKY 1905: 257 (part.);
C. quatuorlineata var. sauromates SCHREIBER 1912: 698 (part.)
Elaphe quatuorlineata — NIKOLSKY 1916: 133 (part.)
Elaphe quatuorlineata — ALEKPEROV 1978: 128
Elaphe quatuorlineata sauromates — LINDHOLM 1929: 80 (part.)
Elaphe quatuorlineata sauromates — SOBOLEVSSKY 1929
Elaphe quatuorlineata sauromates — SZCZERBAK 1966: 200 (part.)
Elaphe quatuorlineata sauromates — MUSKHELISHVILI 1970: 163
Elaphe quatuorlineata sauromates — SCHULZ, 1996: 224 (part.)
Elaphe quatuorlineata sauromates — SINDACO et al., 2000: 473 (part.)
Elaphe sauromates — LENK et al. 2001: 329 (part.)
Elaphe sauromates — UTIGER et al. 2002: 105 (part.)
Elaphe sauromates — TUNIYEV et al. 2009: 80 (part.)
Elaphe sauromates — ARAKELYAN et al. 2011: 88
Elaphe sauromates — KORNILIOS et al. 2014: 149 (part.)
Elaphe sauromates — WALLACH et al. 2014: 263 (part.)
Elaphe sauromates — SAFAEI-MAHROO et al. 2015: 279 
DistributionE Turkey, Georgia, Armenia, Azerbaijan, Nagorno-Karabakh, Iran, and Russia; elevation ~25 m below sea level in the Lenkoran foredeep to about 2,600 m a.s.l. in the Shirak Province in Armenia.

Type locality: Bitlis Province, Turkey (Kısıklı Village, Süphan Mts.; 38.93N, 42.91E, 2,394 m elevation; Fig. 5 in Jablonski et al. 2019.  
Reproduction 
TypesHolotype: ZDEU 26/2012 (tissue sample no. 1,124), adult male, collected by Sako B. Tuniyev; July 16, 2012 (see also Tuniyev et al., 2014). Paratypes: 12 specimens (seven males, five females; eight fixed specimens and four alive), ZDEU 114/1970, adult male from Kars Province (Hoşerenler Plate), Turkey, 40.61N, 43.09E; IZANAS T-17, (tissue sample 3,655) adult male from surroundings of Guzdak, Qobustan, Azerbaijan, 40.37N, 49.68E (Fig. 6C); IZANAS 518, adult male from surroundings of Zaqatala, Azerbaijan, 41.63N, 46.65E; IZANAS 529, adult female from surroundings of Zaqatala, Azerbaijan, 41.63N, 46.65E; IZANAS 68, adult male from surroundings of Lenkoran, Azerbaijan, 38.75N, 48.85E; IZANAS 71, adult female from Zivi-Zkaro (=Tsivi-Tskaro; now Soyuqbulaq), Azerbaijan, 41.32N, 45.27E; IZANAS 70, adult male from surroundings of Şamaxi, Azerbaijan, 40.63N, 48.63E; IZANAS 69, adult female from Şamaxi, Azerbaijan; TuRE ES 0000003300 (tissue sample 3,196), live adult female from Mt. Gutanasar, Gegamsky Ridge (=Ahmangan), near Abovyan, Armenia, 40.37N, 44.69E (Fig. 6A); TuRE ES 0000003301 (tissue sample 3,195), live adult male from Mt. Gutanasar, Gegamsky Ridge (=Ahmangan), near Abovyan, Armenia, 40.37N, 44.69E (Fig. 6B); TuRE ES 0000003302 (tissue sample 3,450), live adult male from surroundings of Mt. Atis, Kotayikskoe Plateau, Armenia, 40.36N, 44.61E; TuRE ES 0000003303 (tissue sample 3,449), live adult female from NW slope of the Mt. Atis, Kotayikskoe Plateau, Armenia, 40.31N, 44.73E. (see also Fig. 6; Table 6 in Jablonski et al. 2019). 
DiagnosisDiagnosis. A new species of western Palearctic genus Elaphe, very similar to E. sauromates (Pallas, 1814), characterized by the combination of the following characters: total length usually does not exceed 1,200 mm (796–1,205 mm), snout-vent (SVL) length usually less than 1,000 mm (650–970 mm), tail length less than 250 mm (146–245 mm) (see Tables 4 and 6 in Jablonski et al. 2019). Tail forms about 25% of the SVL in males and about 21% in females. Head relatively large, distinguished from the body. Snout in prefrontal and internasal area is conspicuously convex which usually forms a hook-nosed head profile. Pileus length on average 1.8–1.9 times larger than its width. Frontal plate 1.2–1.3 times longer than wide. Anterior inframaxillar scute relatively large and wide, 1.2–1.3 times longer than the narrow posterior inframaxillar scute. One or two preocular scales, one loreal, two postoculars, two temporals, three or four posttemporals, eight labials, 10–11 sublabials on each side of the head. Eye in contact with fourth and fifth labials (Table 5; Table S3). Variation in head scale counts is relatively low (see Table S3). Usually two gulars located the anterior inframaxillars. The total number of gulars between inframaxillars and first preventral scale exceeds 12. Number of ventrals is 154–211 (154–206 in males, 194–211 in females), 60–74 subcaudal pairs (65–74 in males, 60–72 in females). 23–25 longitudinal rows of scales are around the midbody, with well-developed keels on 18–21 rows of body scales. The background of dorsal surfaces of the body and lateral surfaces of the head are yellowish or whitish, or seldomly bright yellow. The pattern of the dorsal surface of the body is composed of 50–65 rounded brown or black large ellipsoid spots, which may have whitish edges. Spots can be extended transversely in the posterior part of the body. Pileus is dark, often almost black, slightly lighter on the tip of the snout. Upper preoculars and temporals are dark forming a postocular stripe extending toward the mouth corner. This stripe blends with the dark dorsolateral head coloration anterior to the eye.
Pale spots on the labials, only barely visible or lacking on sublabials. Ventral side of the body is whitish to pale yellow, sometimes with pinkish tint. There are marbled patterns of numerous small irregular dark brown and light gray spots with reddish contours that are more pronounced on the lateral sides of ventral plates. Throat is light, with numerous reddish-orange or brownish speckles on the lower jaws and anterior ventral plates. Iris is dark brown or almost black with a thin light rim around the pupil.

Differential diagnosis. Elaphe urartica sp. nov. is closely related to E. sauromates and E. quatuorlineata. The genetic distance between E. urartica sp. nov. and E. sauromates is 7.20% and 6.91% in COI and ND4, respectively, and 0.15%, 0.15%, 0.73%, and 0.24%, in C-MOS, MC1R, PRLR, and RAG1, respectively. The genetic distance between E. urartica sp. nov. and E. quatuorlineata is 8.24% and 5.80% in COI and ND4, respectively, and 0.15%, 0.15%, and 0.20%, in C-MOS, MC1R, and RAG1, respectively. E. urartica sp. nov. is also morphologically very similar to E. sauromates. E. urartica sp. nov. attains shorter lengths than E. sauromates (795 ± 80 mm vs. 937 ± 152 mm in E. sauromates males, 861 ± 97 mm vs. 929 ± 160 mm in E. sauromates females; though E. sauromates from other parts of the range, for example, the Balkans, can be even larger than specimens in our dataset, see Sahlean et al., 2016). Both taxa also differ in some relative head dimensions. E. urartica sp. nov males have relatively (in comparison to the head length) longer pileus, higher rostrum, but shorter frontal plate and anterior inframaxillary scute. Females only differ in the pileus length (Table 4). The upper surface of the head is more convex near orbits, prefrontals, and internasals and the rostrum is more anteriorly pronounced in E. urartica sp. nov. than in E. sauromates. Males of both species also slightly differ in their scalation: E. urartica sp. nov. males have fewer subcaudal pairs (64 ± 4 vs. 75 ± 3 in E. sauromates) and loreal scales (1–2 vs. 1–3 in E. sauromates). Other differences in metric and meristic characters were not found statistically significant. The coloration of E. urartica sp. nov. is generally darker than that of E. sauromates (Figs. 4 and 6–10). The dorsal side of the head is very dark, sometimes almost black and without the whitish area separating two blotches just posterior to the head as seen in E. sauromates (Figs. 4 and 6–10). On the lateral side of the head the dark stripe running from behind the eye toward the corner of the mouth is also less distinguished in E. urartica sp. nov. compared to
E. sauromates, in which it is clearly separated by lighter color from the darker head coloration. E. urartica sp. nov. has more conspicuous dorsal body spots that are more rounded than in E. sauromates, in which transverse elongation of the spots is common. These dorsal spots are typically lined with whitish color in E. urartica sp. nov. rather than yellow or yellowish as in E. sauromates. 
CommentSynonymy: after Jablonski et al. 2019. E. urartica was split off from E. sauromates, hence previous references to E. sauromates may refer to E. urartica.

Sympatry: E. dione in Dagestan, central-eastern Azerbaijan, eastern Georgia, and presumably in north-eastern Turkey, southern Armenia, and northern Iran. All other species of the genus Elaphe have allopatric distribution relative to E. urartica .

Distribution: see map in Jablonski et al. 2019: 4 (Fig. 1). 
EtymologyThe specific epithet is a feminine adjective derived from the name of the ancient kingdom of Urartu that flourished in the Armenian Highlands and around lake Van, an area of recent distribution of E. urartica sp. nov., in the 9th–6th century BCE (Asher & Asher, 2009). We are choosing this name out of respect for Peter Simon Pallas, who proposed the name for E. sauromates, now the sister species of E. urartica, which most likely refers to Sarmatians (Sauromatae; Sauromatai in Greek), a confederation of nomadic peoples inhabiting vast portions of the recent range of E. sauromates between the 5th century BCE and 4th century CE. 
References
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