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Macrovipera lebetinus (LINNAEUS, 1758)

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Higher TaxaViperidae, Viperinae, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
SubspeciesMacrovipera lebetinus lebetinus (LINNAEUS 1758)
Macrovipera lebetinus cernovi (CHIKIN & SZCZERBAK 1992)
Macrovipera lebetinus obtusa (DWIGUBSKY 1832)
Macrovipera lebetinus schweizeri (WERNER 1935)
Macrovipera lebetinus transmediterranea (NILSON & ANDRÉN 1988)
Macrovipera lebetinus turanica (CHERNOV 1939) 
Common NamesE: Levantine Viper
G: Levanteotter
Russian: Гюрза
G: Milosotter, Milosviper, Kykladenotter [schweizeri] 
SynonymColuber Lebetinus LINNAEUS 1758: 218
Vipera lebetina — WALL 1908: 804
Vipera lebetina — SMITH 1943: 486
Vipera xanthina (GRAY 1849) fide KHALIKOV (pers. comm.)
Vipera mauritanica (GRAY 1849) fide KHALIKOV (pers. comm.)
Vipera euphratica — BOETTGER 1880: 167
Vipera euphratica — BOULENGER 1887: 345
Daboia lebetina — ENGELMANN et al 1993
Macrovipera lebetina — MCDIARMID, CAMPBELL & TOURÉ 1999: 383
Vipera lebetina — SHARMA 2004
Macrovipera lebetina — KWET & TRAPP 2015
Macrovipera lebetina — WALLACH et al. 2014: 421
Macrovipera lebetinus — FRETEY 2019
Macrovipera lebetina — TUNIYEV et al. 2019
Macrovipera lebetinus — SPEYBROECK et al. 2020

Macrovipera lebetinus cernovi (CHIKIN & SZCZERBAK 1992)
Macrovipera lebetina cernovi — DAVLETBAKOV et al. 2016
Macrovipera lebetina cernovi — ORAIE et al. 2018

Macrovipera lebetinus lebetinus (LINNAEUS 1758)
Coluber lebetinus LINNAEUS 1758
Macrovipera lebetina cypriensis REUSS 1933
Vipera lebetina euphratica — SCHMIDT 1939
Vipera lebetina lebetina — HARDING & WELCH 1980
Vipera lebetina lebetina — WELCH 1994: 73
Daboia lebetina lebetina — OBST 1983: 232
Macrovipera lebetina lebetina — GOLAY et al. 1993

Macrovipera lebetinus obtusa (DWIGUBSKY 1832)
Vipera obtusa DWIGUBSKY 1832: 30
Vipera euphratica MARTIN 1838
Vipera peilei MURRAY 1892
Vipera lebetina BOULENGER 1920
Vipera lebetina peilei — SCHWARZ 1936: 244
Vipera lebetina euphratica SCHMIDT 1939
Vipera lebetina obtusa — MERTENS & WERMUTH 1960
Vipera lebetina euphratica — HAAS & WERNER 1969
Vipera lebetina obtusa — ANDERSON & LEVITON 1969
Vipera lebetina obtusa — HARDING & WELCH 1980
Daboia lebetina obtusa — HERRMAN, JOGER & NILSON 1989
Daboia lebetina obtusa — ENGELMANN et al 1993
Macrovipera lebetina obtusa — GOLAY et al. 1993
Vipera lebetina obtusa — WELCH 1994: 121
Vipera lebetina obtusa — FRYNTA et al. 1997
Macrovipera lebetina obtusa — SCHWEIGER 2009
Macrovipera lebetina obtusa — ISKENDEROV 2013
Macrovipera lebetina obtusa — ORAIE et al. 2018

Macrovipera lebetinus transmediterranea (NILSON & ANDRÉN 1988)
Vipera lebetina transmediterranea NILSON & ANDRÉN 1988
Macrovipera lebetina transmediterranea — GOLAY et al. 1993
Vipera lebetina transmediterranea — WELCH 1994: 121
Daboia lebetina transmediterranea
Macrovipera lebetina transmediterranea — KUCHARZEWSKI 2011
Macrovipera lebetina transmediterranea — SAVAGE & MCDIARMID 2017

Macrovipera lebetinus schweizeri (WERNER 1935)
Vipera lebetina schweizeri WERNER 1935: 117
Vipera lebetina siphnensis WETTSTEIN 1952
Vipera schweizeri — NILSON & ANDRÉN 1988
Daboia lebetina schweizeri — OBST 1983
Macrovipera schweizeri — HERRMANN, JOGER & NILSON 1992
Daboia lebetina schweizeri — ENGELMANN et al. 1993
Vipera schweizeri — WELCH 1994: 123
Macrovipera schweizeri — MCDIARMID, CAMPBELL & TOURÉ 1999: 386
Macrovipera schweizeri — SCHWEIGER 2009
Macrovipera schweizeri — LIVIGNI 2013: 343
Macrovipera lebetina schweizeri — SCHMID 2015
Macrovipera schweizeri — WALLACH et al. 2014: 421
Macrovipera schweizeri — SPEYBROECK et al. 2020
Macrovipera lebetina schweizeri — FREITAS et al. 2020

Macrovipera lebetinus turanica (CHERNOV 1939)
Vipera lebetina turanica CHERNOV 1939
Vipera lebetina turanica — HARDING & WELCH 1980
Daboia lebetina turanica — OBST 1983: 232
Macrovipera lebetina turanica — GOLAY et al. 1993
Macrovipera lebetina turanica — LAITA 2013
Macrovipera lebetina turanica — DAVLETBAKOV et al. 2016
Macrovipera lebetina turanica — ANANJEVA et al. 2020 
DistributionCyprus, Iraq, Turkey, Afghanistan, Syria, Lebanon, Iran, W Pakistan, NW India (Jammu and Kashmir), Jordan, Yemen ? (single specimen fide Scortecci 1932),
S Russia (Dagestan), Armenia, Azerbaijan, Turkmenistan, Georgia, Uzbekistan, Kyrgyzstan, Kazakhstan, Tajikistan,

cernovi: Turkmenistan, Iran, Uzbekistan, Kyrgyzstan, Afghanistan, India (Kashmir), Pakistan; Type locality: bank of the river Murghab in the vicinity of town Iolotan, Marijsky district, Turkmenistan.

lebetinus: Cyprus, N Iraq, NW Syria; Type locality: Oriente; MERTENS & MÜLLER (1928) suggested to restrict the type locality to Cyprus.

schweizeri: Greece (Milos, Syphnos [= Sifnou = Siphnos], Kimolos, Polynos = Polyaigos = Polinos ?); Type locality: “Insel Milos”, Greece

transmediterranea: Algeria; Type locality: Algiers.

turanica: Tajikistan, Uzbekistan, Kyrgyzstan

obtusa: NE Turkey, Iraq, Afghanistan, Syria, Transcaucasia, Lebanon, Iran, Pakistan, Jordan; neotype locality: “Geok Tapa (Elizabetpol)”, now Gyanja, Azerbaijan.

Type locality: Transcaucasia  
Reproductionoviparous (although sometimes reported to be viviparous). 
TypesHolotype: nondesignated (formerly MAFR), lost fide Andersson (1899: 29)
Holotype: NMNHU (formerly ZIK) Re-19 (given as Zool. Mus., Inst. Zool., Acad. Sc. Ukraine) [cernovi]
Holotype: lost; neotype: ZISP 8595 [obtusa]
Lectotype: ZSM (SLM) 3210, female, “Umgebung von Larnaka, Cypern”, received from Behringwerke, 05.1933 (card index), designation by Krecsák (2007) [cypriensis]
Lectotype: MCZ 37008 (also given as 37000) [schweizeri]
Holotype: USNM 6210, paratypes: ZMUC, BMNH, NMW [transmediterranea]
Holotype: ZISP (also as ZIL) 9080 [turanica] 
Diagnosis

Diagnosis (schweizeri): Werner 1935 diagnoses this subspecies from bornmülleri, xanthina, lebeina (s.str.) and mauritanica by these traits: “Schuppen in 21 — 23 Reihen, unterste Temporalenreihen ungekielt, Supraoculare in kleine Schildchen aufgelöst; Subcaudalia Männchen 40—45, Weibchen 33—41”

Diagnosis (transmediterranea): A subspecies of Vipera lebetina that is geographically separated from other subspecies by the Mediterranean Sea, and which can be characterized as a typical lebetina form having light ground colour with a pattern consisting of 34-41 transverse bars. Midbody scale rows 25 in number and the number of ventrals is between 150 and 164 in females; between 154 and 163 in males.
This new subspecies differs from other taxa of the lebetina complex s. l. (except for V. lebetina and V. schweizeri, Table 1), in having a low ventral number. The mean values for different scalation characteristics indicate a higher fragmentation of head scales in this new taxon compared with other subspecies of lebetina as weil as with
V. schweizeri and V. mauritanica (see Table 2). The mean value for the number of loreals, interoculars and 1st and 2nd circumoculars is higher than in any of the compared taxa. lt is most similar to the nominate subspecies, Vipera l. lebetina from Cyprus, from which, however, it differs in having a higher ventral count (150 to 164 in female, and 154 to 163 in male transmediterranea; 146 to 153 in female, and 147 to 154 in male l. lebetina), and in the number of anterior scale rows which range from 23 to 27 (X = 24) in transmediterranea and from 21 to 23 (X = 22.2) in l. lebetina. From the sympatric (or parapatric) Vipera mauritanica the new subspecies differs in colour pattern (V. mauritanica has a wavy zig-zag band containing 23 to 33 windings; V. l. transmediterranea has 34 to 41 transverse bands), the midbody scale rows are 25 in V. l. transmediterranea, 27 in V. mauritanica; the number of scale rows on the neck is around 24 in V. l. transmediterranea,between 26 and 27 in V. mauritanica; the number of ventrals is 150 to 164 in female and 154 to 163 in male V. l. transmediterranea,157 to 174 in female and 163 to 170 in male V.mauritanica; the number of subcaudals is between 37 and 43 in female V. l. transmediterranea, and between 45 and 48 in female V. mauritanica; corresponding figures for males are between 42 to 51 and 45 to 50 respectively.

Size: Maximum length for both sexes is 98.5 cm. Relative tail length in the two males are 12.6 and 13.2% of total length, while corresponding figures in the two adult females are 10.1 and 13.1 %. (Nilson & Andren 1988: 373)

Comparisons: for a comparison of meristic characters across schweizeri, turanica, obtusa, lebetina, transmediterranea, deserti, and mauritanica, see tables in Nilson & Andren 1988: 374. 
CommentVenomous!

Type species: Coluber Lebetinus LINNAEUS 1758 is the type species of the genus Macrovipera REUSS 1927.

Synonymy: Listed as Vipera lebetina obtusa in Leviton et al. Macrovipera lebetina deserti, V. l. mauritanica have been elevated to full species status. According to molecular studies (Herrmann et al. 1999), Macrovipera lebetina is paraphyletic with respect to Vipera and Pseudocerastes, and Eristicophis. Macrovipera lebetina peilei is not recogized as subspecies by GOLAY et al. and MCDIARMID et al. (1999). Golay 1993 lists peilei as a synonym of obtusa. Kaiser et al. 2013 considered the generic name Laidlawus Hoser 2012 invalid and rejected its use instead of Macrovipera.

Subspecies: the status of Macrovipera lebetinus schweizeri is controversial (or preliminary) and some authors consider it as valid species.

Illustration of M. l. turanica in KHAN (2002), Figure 151; the specimen is NOT M. l. obtusa as indicated by the author (fide TILLACK & KUCHARZEWSKI 2004).

Phylogenetics: for a recent phylogeny of this and related Viperinae see Šmíd & Tolley 2019.

Habitat: partly arboreal (Harrington et al. 2018).

Distribution: For a map see Sindaco et al. 2013. Not in Israel fide Shai Meiri, pers. comm., 29 Apr. 2021. Occurrence in Saudi Arabia unlikely; the record from Yemen is based on a specimen collected in 1932 by Scortecci, with uncertain specimen data (according to Corkill & Cochrane 1966 probably in FMNH or AMNH. Records from Algeria represent the subspecies transmediterranea, which may be an introduced population. The origin of a paratype from “Tunis”[Tunisia] is questionable.

NCBI taxonID: 110205 [schweizeri] 
EtymologyThere are two diferent interpretations of this name: “lebetinus” referring to its origin in the Levant (Schleich et al. 1996), or (2) referring to the Greek kettle drummer of funerals (”lebes” or similar), referring to a warning signification (Lescure & Le Garff 2006, Fretey 2019). The species epithet has to be “lebetinus” fide Fretey 2019. 
References
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