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Pseudopus apodus (PALLAS, 1775)

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Higher TaxaAnguidae (Anguinae), Diploglossa, Anguimorpha, Sauria, Squamata (lizards)
SubspeciesPseudopus apodus apodus (PALLAS 1775)
Pseudopus apodus thracius (OBST 1978) 
Common NamesE: European Glass Lizard
G: Scheltopusik, Panzerschleiche
Croatian: blavor
Russian: Желтопузик 
SynonymLacerta apoda PALLAS 1775: 435
Chalcida apus — MEYER 1795: 31
Lacerta apus — SHAW & NODDER 1799: plate 411
Sheltopusik didactylus LATREILLE 1802
Pseudopus serpentinus MERREM 1820 (nom. subst.)
Pseudopus durvillii CUVIER 1829 (nomen dubium fide OBST 1980)
Pseudopus Fischeri MÉNESTRIÉS 1832
Pseudopus serpentinus — GERVAIS 1836: 311
Pseudopus Pallasii — DUMÉRIL & BIBRON 1839: 417
Pseudopus serpentinus — BERTHOLD 1840
Pseudopus Pallasii — GRAVENHORST 1851: 384
Pseudopus Pallasii — BRÜHL 1886
Ophisaurus apus — BOULENGER 1885: 280
Pseudopus Pallasii — LEYDIG 1886
Ophisaurus apus —BOETTGER 1889: 269
Pseudopus pallasii — CAMP 1923: 365
Ophisaurus pallasii — CAMP 1923: 368
Ophisaurus apodus — MERTENS & MÜLLER 1928
Ophisaurus apodus — SMITH 1935: 395
Ophiosaurus apodus — LEVITON 1959 (in error)
Pseudopus apodus — KLEMBARA 1979
Ophisaurus apodus — OBST 1981
Ophisaurus apodus — KASPAREK 1990
Ophisaurus apodus — ENGELMANN et al 1993
Ophisaurus apodus — FRYNTA et al. 1997
Ophisaurus apodus — ANDERSON 1999
Anguis apodus — MACEY et al. 1999
Pseudopus apodus — ROITBERG et al. 2000
Ophisaurus apodus — PLESSAS 2002
Ophisaurus apodus — KUMLUTAS et al. 2004
Pseudopus apodus — RUTSCHKE et al. 2005
Pseudopus apodus apodus — JANDZIK et al. 2017
Pseudopus apodus apodus — JABLONSKI et al. 2021

Pseudopus apodus thracius (OBST 1978)
Ophisaurus apodus thracius OBST 1978
Ophisaurus apodus thracius — THIEME 1985
Ophisaurus apodus thracius — ENGELMANN et al. 1993
Ophisarus [sic] apodus thracius — CEVIK 1999
Pseudopus apodus thracius — STOEV 2000
Pseudopus apodus thracius — SCHLÜTER 2005
Pseudopus apodus thracius — JANDZIK et al. 2017
Pseudopus apodus thracius — JABLONSKI et al. 2021

DistributionGreece (incl. Limnos, Rhodos, Lesbos, Corfu = Corfou, Chios, Samos), coastal Croatia, Hercegovina, Monte Negro, Macedonia, Montenegro, Albania, Bulgaria, Turkey, Lebanon,
European Soviet Union (Caucasus, Dagestan), S Russia (Dagestan), S Ukraine, Republic of Georgia, Armenia, Azerbaijan, Turkmenistan, Uzbekistan, Kyrgyzstan, W Tajikistan, S Kazakhstan, Afghanistan, Syria, Iran, Iraq, Syria, Jordan, Israel

thracicus: northern coastal areas of the Black Sea, Bulgaria, Greece (Lesbos etc.)

Type locality: North Caucasus; Naryn Steppe, Russia (N coast of Caspian Sea) fide ANDERSON 1999. Obst 1978:136 considered this unlikely and restricts the type locality to the Terek River region.

levantinus: southern Turkey, western Syria, northern and central Israel; Type locality: Giv’at Ada (גבעת עדה), Israel)32.52°N, 35.01°E.  
TypesHolotype: unlocated fide ANDERSON 1999.
Holotype: MTD (= MTKD); Paratypes: ZISP 19642, ZFMK, MTD (= MTKD) [thracius]
Syntypes: ZISP 2403 [Pseudopus Fischeri]
Holotype. TAU-R 16895 (Fig. 6, Table S7), an adult male, collected on the 16th of May, 2014 by Erez Maza, at Giv’at Ada (גבעת עדה), Israel )32.52°N, 35.01°E, Fig. 7). The molecular-genetic data of the holotype are available for ND2 (MW400924), Cyt b (MW400903), and microsatellites (Fig. 2, Table S1, S2). Paratypes. TAU-R 17076 (MW400927, MW400906), an adult male, th collected on 19 May 2014 by Talia Oron, at Biranit (בירנית), Israel )33.05°N, 35.34°E(; TAU-R 17235 )MW400928, MW400907), an adult male, collected on 6th May 2015 by Ron Elazari, at Hadera (ֲח ֵד ָרה ), Israel )32.47°N, 34.89°E(; TAU-R 17895 )MW400930, MW400909), an adult הר( ’male, collected on 18th April 2016 by Ofer Shimoni, on Mt. Gilboa -Israel )32.45°N, 35.43°E(; TAU-R 19403, an adult male, col ,(הגלבוע th ,)רמת הנדיב( lected on 16 May 2019 by Amir Arnon, at Ramat HaNadiv Israel )32.55°N, 34.95°E( )Figs. 2, 8, Table S1, S2, S7) [levantinus] 
DiagnosisDiagnosis: Ear opening distinct; vestiges ofhind limbs appearing as elongate papillae at sides of vent; pterygoid teeth in two series; dorsal scales keeled. See Obst (1978: 137) for distinctions between subspecies he recognizes (Anderson 1999: 114).

Diagnosis (levantinus). A large Pseudopus (up to 610 mm snout-vent length, 1,367 mm total length and a mass of 1,100 g) that can be distinguished from the other two subspecies by a combination of the following characters (means followed by standard deviations; Table 4): (1) preanal scales (PAN) 10; (2) long body (SVL; 490.22 mm ± 76.15); (3) long tail (TL; 600.70 mm ± 107.04); (4) long head (HL1; from the ear aperture to the tip of the snout, 47.6 mm ± 7.99); pileus length (PL; mean 41.98 mm ± 6.78); and distance from the frontal shield to the tip of the snout (FSL; 17.27 mm ± 2.96); (5) wide head (HW1; maximum width, 32.04 mm ± 6.34); and width at the level of the posterior edge of the eyes orbits (HW2; 28.21 mm ± 4.95); (6) relatively long limb rudiments (RL; 5.55 mm ± 1.53) (see and Tables 3 and 4 for comparisons to P. a. apodus and P. a. thracius).

Comparisons (levantinus). Pseudopus apodus levantinus ssp. nov. can be distinguished from P. a. apodus and P. a. thracius by having 10 preanal scales (PAN) in all morphologically examined specimens (vs. 5–11, with the mean of 8 in P. a. apodus, and 5–8 and mean of 6 in P. a. thracius). Other differences can be used in combination to support differences between P. a. levantinus ssp. nov., P. a. apodus, and P. a. thracius: longer body (SVL; 610 mm maximum length and mean of 490.22 mm in P. a. levantinus ssp. nov., vs. 487 mm maximum and mean of 387.61 mm in P. a. apodus, and 410 mm maximum and mean of 360.06 mm in P. a. thracius); longer distance between the ear aperture and the tip of the snout (HL1; 63.75 mm maximum length and mean of 47.60 in P. a. levantinus ssp. nov., vs. 58 mm maximum and mean of 38.61 mm in P. a. apodus, and 49.50 maximum and mean of 39.55 mm in P. a. thracius); longer pileus (PL; 54.79 mm maximum length and mean of 41.98 in P. a. levantinus ssp. nov., vs. 46.30 mm maximum and mean of 35.82 mm in P. a. apodus, and 28.80 mm maximum and mean of 36.93 mm in P. a. thracius); longer distance between the anterior frontal scale to the tip of snout (FSL; 23.39 mm maximum length and mean of 17.27 mm in P. a. levantinus ssp. nov., vs. 18 mm maximum and mean of 13.90 mm in P. a. apodus, and 15.98 mm maximum and mean of 12.58 mm in P. a. thracius); wider head (HW1; 44.90 mm maximum width and mean of 32.04 mm in P. a. levantinus ssp. nov., vs. 36.5 mm maximum and mean of 24.47 mm in P. a. apodus, and 29.50 mm maximum and mean of 24.35 mm in P. a. thracius); wider distance between the posterior edge of the orbits (HW2; 39.33 mm maximum width and mean of 28.21 mm in P. a. levantinus ssp. nov., vs. 28.76 mm maximum and mean of 19.44 mm in P. a. apo­dus, and 26.30 mm maximum and mean of 22.33 mm in P. a. thracius); and longer limb rudiments (RL; 8.56 mm maximum length and mean of 5.55 mm in P. a. levantinus ssp. nov., vs. 6.55 mm maximum and mean of 3.5 mm in P. a. apodus, and 4.8 mm maximum and mean of 3.31 mm in P. a. thracius(. The remaining differences are presented in Tables 3–4. 
CommentSynonymy: Partly after BRYGOO 1987. KLEMBARA (1979) also transferred Ophisaurus moguntinus and Ophisaurus pannonicus to Pseudopus. Macey et al. (1999) found this species nested within other Ophisaurus species thereby the name Ophisaurus apodus is required to render the genus monophyletic.

Limb morphology: Limbless.

Type Species: Pseudopus serpentinus MERREM 1820 is the type species of the genus Pseudopus MERREM 1820.

Distribution: See map in SMID et al. 2014 for distribution in Iran. See Jandzik et al. 2017 for map of subspecies. These authors also identified a southern lineage that may be described as a separate subspecies. See Bar et al. 2021 for a map.

Fossils: Multiple fossil species of the Pseudopus lineage have been described, including P. rugosus (Klembara 2014) and P. ahnikoviensis (Klembara 2012) in the early Miocene, Pseudopus laurillardi (Klembara et al. 2010) in the middle Miocene, and Pseudopus pannonicus in the late Miocene and Pliocene (Klembara et al. 2010).

Karyotype: 2n = 44 (Altmanová et al. 2022) 
EtymologyNamed after Greek “pus, podos” = leg, pseudo = fake, and a-podus for no leg.
The German name “scheltopusik” is derived from Russian for “yellow belly” (Langner 2019).
P. a. levantinus has been named after the Levantine region. 
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