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Trapelus mutabilis (MERREM, 1820)

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Higher TaxaAgamidae (Agaminae), Sauria, Iguania, Squamata (lizards)
SubspeciesTrapelus mutabilis pallidus (REUSS 1834)
Trapelus mutabilis mutabilis (MERREM 1820)
Trapelus mutabilis poppeki WAGNER, MELVILLE, WILMS & SCHMITZ 2011 
Common NamesE: Desert Agama 
SynonymAgama mutabilis MERREM 1820: 50
Trapelus aegyptius CUVIER 1829
Agama inermis REUSS 1834: 33 (fide MERTENS 1922)
Agama gularis REUSS 1834: 36 (fide MERTENS 1922)
Agama mutabilis — DUMÉRIL & BIBRON 1837: 505 (part)
Trapelus aegyptius DUVERNOY in CUVIER 1848 (fide ANDERSON 1898)
Agama agilis — EICHWALD (= inermis fide BOULENGER 1891)
Agama mutabilis — BOULENGER 1885: 338
Agama aspera WERNER 1893: 359
Agama latastii BOULENGER 1885: 344
Agama inermis — BOULENGER 1885: 344
Agama leucostigma — BOULENGER 1885: 348 (nom. subst.)
Agama inermis — DE GRIJS 1899: 401
Agama inermis — ANGEL 1923
Agama inermis — ANGEL 1932: 386
Agama mutabilis — WERMUTH 1967: 19
Agama mutabilis — PAPENFUSS 1969
Agama agnetae — WELCH
Agama mutabilis — JOHANN 1981
Agama mutabilis — BRYGOO 1988: 3
Trapelus mutabilis — SCHLEICH, KÄSTLE & KABISCH 1996: 291
Trapelus mutabilis — PADIAL 2006

Trapelus mutabilis pallidus (REUSS 1834)
Agama pallida REUSS 1834: 38
Agama deserti LICHTENSTEIN 1823 (fide WAGNER et al. 2011)
Agama loricata REUSS 1834: 40 (fide MERTENS 1922)
Agama nigrofasciata REUSS 1834: 42 (fide MERTENS 1922)
Agama leucostygma REUSS 1834: 44 (fide MERTENS 1922; partim)
Agama pallida — BOULENGER 1885: 348
Agama pallida — ANDERSON 1898: 100
Agama pallida — HAAS 1957: 65
Agama mutabilis — WERMUTH 1967: 19
Agama pallida — GORMAN & SHOCHAT 1972
Agama pallida — ESTERBAUER 1985
Trapelus pallidus — DISI et al. 2001: 151
Trapelus mutabilis pallidus — WAGNER et al. 2011
Trapelus pallidus — IBRAHIM 2013

Trapelus mutabilis poppeki WAGNER et al. 2011
Trapelus mutabilis poppeki WAGNER, MELVILLE, WILMS & SCHMITZ 2011: 896
Trapelus mutabilis poppeki — BAUER et al. 2017 
DistributionMauritania, Western Sahara, Algeria, Tunisia, Libya, Egypt, Mali, Iraq, Chad, Sudan (Jumhūriyyat), Saudi Arabia

Type locality: “in Aegypto”.

pallidus: Egypt, Sinai, Syria, Jordan; Type locality: “Ober-Ägypten”.

poppeki: Libya; Type locality: Libya: east of Tarbu, 24.48.27°N, 16°19’11’’E, 490 m elevation.  
TypesNeotype: ZFMK 64395: Egypt, 10 km northwest of Cairo; leg. Hans-Werner Herrmann, 1990.
Holotype: SMF 10007 [pallidus]
Holotype: ZFMK 63678: juvenile, leg. Hemmo Nickel, ix.1996 (Fig. 7). Paratype: ZFMK 20848: female, Libya: Tripolis, leg. W. Schlüter, 1913 [poppeki]
Syntypes: ZMB 767, ZMB 768-769 sowie ZMB 771= Agama ruderata ?, ZMB 55736 (vormals ZMB 771 part.) [deserti] 
DiagnosisDiagnosis (genus). Tympanum small, diameter less than half of the orbit, deeply sunk. Head usually rather high than short. Caudal scales not forming distinct annuli. Only males with callous precloacal scales (Wagner et al. 2011: 893).

Diagnosis (mutabilis). Trapelus mutabilis is a small [largest voucher from Egypt SVL 90 mm (Baha el Din, 2006)] species within the genus, possessing irregular dorsal scalation. This species differs from its probable closest relative, T. pallidus, in having a heterogeneous dorsal scalation with rhomboidal and scattered enlarged scales. In contrast, T. pallidus has a matrix of comparatively uniform, smooth dorsal scales, with some scattered larger keeled scales. In addition, T. mutabilis differs from T. pallidus in having homogeneous scalation on the hindlimbs and base of the tail. Coloration in life is uniformly sandy grey with four to five transverse bands and a barred tail. Male T. mutabilis have nuptial coloration of violet-blue flanks and throat; in contrast male T. pallidus coloration is restricted to having a completely blue head. Trapelus mutabilis differs from two specimens from Libya in not possessing scale rows of enlarged vertebral scales (for details see description in WAGNER et al. 2011). Trapelus mutabilis differs from T. savignii in having smooth ventral scales and no gular pouch, from T. schmitzi in having heterogeneous dorsal scalation and from T. tournevillei in having a shorter tail (average of ratio TL/SVL 1.39 in T. mutabilis instead of 1.60 in T. tournevillei), no gular pouch, a heterogenous dorsal scalation, and the lack of longitudinal stripes on the belly [from WAGNER et al. 2011].

Diagnosis (pallidus): Small to medium-sized species of the genus with an average ratio TL/SVL of 1.1. Gular pouch absent. Ear opening more or less round, usually with a smooth margin, but often spiny scales are present. Dorsal scales small, equal in size, forming a matrix with intermixed larger scales. Ventral scales smooth. Trapelus mutabilis pallidus is distinct from the nominate form in possessing a higher count of scale rows around midbody and in having a matrix of uniform small dorsal scales with some intermixed larger scales (Wagner et al. 2011: 896).

Diagnosis (poppeki): A small to medium sized subspecies of T. mutabilis with all the many other valid Trapelus species and T. m. mutabilis and T. m. pallidus in its typical body scalation of enlarged vertebral scales and much smaller lateral scales. Only T. savignii has a similar body scalation. A second characteristic is the small pineal scale, which is not much larger than the pineal organ. Trapelus m. poppeki ssp. nov. differs from valid African species of Trapelus:
– T. savignii (type locality: Egypt) in not possessing a gular pouch, in having the ventral scales smooth and in a high genetic distance.ypical characteristics of the genus. Trapelus m. poppeki ssp. nov. differs from
– T. m. mutabilis (type locality: Egypt) in possessing enlarged vertebral scales in comparison to much smaller scales on the flanks.
– T. m. pallidus (type locality: southern Egypt) in possessing enlarged vertebral scales in comparison to much smaller scales on the flanks.
– T. tournevillei (type locality: Ouargla, Algeria) in having a shorter head, brownish transverse bands on the body, and smooth ventral scales (instead of keeled ventral scales in T. tournevillei) and in lacking longitudinal lines on the belly.– T. schmitzi (type locality: Ennedi Mts., Chad) in possessing a very heterogenous body scalation with enlarged vertebral scales and smaller lateral scales, and in having brownish transverse bands on the body.
Trapelus m. poppeki ssp. nov. differs from the following synonyms (Agama aspera is maybe a valid taxon, but clearly different to the here-described subspecies and more research is needed to clarify the status of this taxon):
– Agama inermis Reuss, 1833 (type locality: southern Egypt) in possessing enlarged vertebral scales.
– Agama gularis Reuss, 1833 (type locality: southern Egypt) in possessing enlarged vertebral scales.
– Agama latastii Boulenger, 1885 (type locality: Egypt) in lacking equal sized, rhomboidal dorsal scales.
– Agama aspera F. Werner, 1893 (type locality: Algerian Sahara, between Kef-el-Dhor and Chegga; Biskra-Bordj-Saada; Zab-el-Zig south of El Meranyer) in possessing no spiny scales and a heterogeneous dorsal scalation.
(diagnosis from Wagner et al. 2011: 896). 
CommentSynonymy: Possible synonymy between T. mutabilis and Agama pallida is discussed in Leviton et al. (1992) who list Trapelus pallidus as a separate species.
Synonymy: Agama pallida has been synonymized with Agama mutabilis by WERMUTH 1967 and others. Both “species” are very similar or identical. WERMUTH (1967) also listed Agama agnetae as synonym of Agama mutabilis. Synonymy mostly after WERMUTH 1967 and WAGNER et al. 2011.

Distribution: Not in Israel where it is replaced by T. agnetae. Not in Morocco (Martínez et al. 2019, and pers. comm. 18 Jan 2021).

Illustration in BARTS & WILMS 2003.

Type species: Agama mutabilis Merrem, 1820 is the type species of the genus Trapelus CUVIER 1817.

Types: One of the syntypes, ZMB 760, is actually Trapelus pallidus (fide Daniel Melnikov 2012, TILLACK, pers. comm. 22 Jan 2014) 
EtymologyEtymology (pallidus): Named after Latin “pallidus” = pale or yellow(ish). 
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