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Sauromalus ater DUMÉRIL, 1856

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Higher TaxaIguanidae, Iguania, Sauria, Squamata (lizards)
SubspeciesSauromalus ater ater DUMÉRIL 1856
Sauromalus ater shawi CLIFF 1958
Sauromalus ater multiforaminatus (TANNER & AVERY 1964)
Sauromalus ater townsendi DICKERSON 1919
Sauromalus ater tumidus SHAW 1945 
Common NamesE: Common Chuckwalla, Northern Chuckwalla
S: Cachorón de Roca 
SynonymSauromalus ater DUMÉRIL 1856: 536
Euphryne obesus BAIRD 1859: 253
Euphryne obesa — BAIRD 1859 (emendation)
Sauromalus obesus — COPE 1875
Sauromalus ater — BOULENGER 1885: 202
Sauromalus interbrachialis DICKERSON 1919: 640 (part.)
Sauromalus obesus — SCHMIDT 1922
Sauromalus obesus — BURT 1933
Sauromalus obesus — GLOYD 1937
Sauromalus obesus — KLAUBER 1939
Sauromalus australis SHAW 1945: 284
Sauromalus australis — LEVITON & BANTA 1964
Sauromalus australis — BOSTIC 1971
Sauromalus obesus — STEBBINS 1985: 114
Sauromalus ater — LINER 1994
Sauromalus australis — LINER 1994
Sauromalus australis — GRISMER et al. 1994
Sauromalus obesus — QUEIROZ 1995
Sauromalus ater — HOLLINGSWORTH 1998: 113, 144
Sauromalus ater — KÖHLER & ZORN 2014

Sauromalus ater ater DUMÉRIL 1856
Sauromalus ater ater — SOULÉ & SLOAN 1966
Sauromalus ater ater — ETHERIDGE 1982: 32

Sauromalus ater multiforaminatus TANNER & AVERY 1964
Sauromalus obesus multiforaminatus TANNER & AVERY 1964: 38
Sauromalus obesus multiforaminatus — TANNER 1970: 22
Sauromalus obesus multiforminatus — ETHERIDGE 1982: 33
Sauromalus ater multiforminatus —HOLLINGSWORTH 1998 (by implication)

Sauromalus ater shawi CLIFF 1958
Sauromalus shawi CLIFF 1958
Sauromalus ater shawi — SOULÉ & SLOAN 1966
Sauromalus ater shawi — ETHERIDGE 1982: 32
Sauromalus shawi — QUEIROZ 1995

Sauromalus ater townsendi DICKERSON 1919
Sauromalus townsendi DICKERSON 1919
Sauromalus townsendi — TAYLOR 1938: 481
Sauromalus obesus townsendi — SHAW 1945
Sauromalus obesus townsendi — ETHERIDGE 1982: 33
Sauromalus ater townsendi — HOLLINGSWORTH 1998 (by implication)
Sauromalus ater townsendi — REYES et al. 2014

Sauromalus ater tumidus SHAW 1945
Sauromalus ater tumidus SHAW 1945: 292
Sauromalus obesus tumidus — SCHMIDT 1953: 118
Sauromalus ater tumidus — ETHERIDGE 1982: 33
Sauromalus obesus tumidus — PREGILL 1984: 156
Sauromalus ater tumidus — LINER & CASAS-ANDREU 2008: 74 
DistributionUSA (California, S Nevada, S Utah, W Arizona),
Mexico (Baja California, W Sonora; Espiritu Santo, Partida, San Jose, San Francisco, San Diego, Santa Cruz, San Marcos, Santa Catalina, Isla Ballena [HR 27(1)], Isla Pardo [HR 28(1)], Gulf of California, Mexico).

multiforaminatus: Colorado River from Glen Canyon Dam at Page, Arizona to near Hite and the Henry Mountains, Garfiled County, Utah.

obesus: Type locality: Fort Yuma, California.

australis: Mexico (SE Baja California, from Punta San Gabriel southward to La Paz)

townsendi: Mexico (Sonora)

Type locality: restricted to Espiritu Santo Island.(Smith & Taylor 1950).  
TypesHolotype: MNHN-RA 0813
Holotype: SDNHM = SDSNH 30170; paratypes SDNHM = SDSNH [australis]
Holotype: CAS-SUR 16120 [shawi]
Syntypes: UMMZ 3804; USNM 4172 f [obesus]
Holotype: SDNHM = SDSNH 27323; paratypes SDNHM = SDSNH [tumidus]
Holotype: USNM 64442; C. H. Townsend collector. [townsendi]
Holotype: BYU 11376, adult male, collected June 9, 1954, by Wilmer W. Tanner. [multiforaminatus]
Holotype: USNM 64443, given as AMNH 6809, paratype: AMNH 6808 [Sauromalus interbrachialis] 
DiagnosisDiagnosis (genus): differs from all other iguanid genera in possessing the following combination of character states: a) parietal foramen located visibly within frontal, b) splenial relatively small, c) angular does not extend far up labial surface of dentary and is not visible, or is only barely visible, in lateral view, d) angular reduced and narrow posteriorly, e) modal number of premaxillary teeth fewer than seven, f) crowns of posterior marginal teeth with five or more cusps, g) second ceratobranchials of hyoid apparatus short, often less than two-thirds the length of the first ceratobranchials, h) second ceratobranchials not in contact medially for most or all of their lengths, i) neural spines of presacral vertebrae short, less than 50% of total vertebral height, j) fewer than 40 caudal vertebrae, k) postxiphisternal inscriptional ribs never form continuous midventral chevrons, l) suprascapular cartilages situated primarily in a horizontal plane, and each forms an angle rather than a smooth curve with a scapula, m) scapular fenestrae small or absent, n) clavicles narrow with the lateral shelf small or absent, o) posterior process of interclavicle does not extend beyond corners of the sternum, p) lateral processes of the interclavicle form angles of between 75° and 90° with posterior process (interclavicle roughly t-shaped), q) sternal fontanelle small or absent, r) sternum pentagonal with xiphisterna widely separated, s) pelvic girdle short and broad, t) anterior iliac process small, u) heart extends posterior to transverse axillary plane, v) rostral scale divided by a median suture, w) superciliary scales quadrangular and non-overlapping, x)anterior auricular scales enlarged, y) middorsal scale rows absent, z) anterior and posterior keels of subdigital scales approximately equal in size, aa) subdigital scales roughly symmetrical with respect to long axis of toe, bb) body strongly depressed (John Iverson et al., pers. comm.).

Diagnosis (townsendi). General resemblance to S. ater Dumeril but with coarser scutellation throughout, especially on extremities. Size medium, length of type (adult, but probably not full-grown; also tail reproduced) 302 mm; dorsal scale rows in head length, 28; ventrals from gular fold to anus, 125; number of scales around thick part of tail, 80-90; femoral pores, 14. Coloration yellowish, with head, shoulders, brachials, gular region, and anterior breast, also lumbar region and hind legs dorsally and ventrally, very dark brown, and an irregular freckling of the same color dorsally and ventrally over the yellow between (Dickerson 1919: 464). 
CommentSynonymy: Hollingsworth (1998) synonymized Sauromalus obesus with S. ater because both forms (including their subspecies) show a wide range of character overlap and variation and are therefore not clearly distinguishable. Sauromales ater DUMÉRIL 1856 was described earlier and therefore has priority to S. obesus. See also Brown et al. (2001) and the ruling in the Bulletin of Zoological Nomenclature 61(1): 74-75 (2004) which gave the name Sauromalus ater Duméril 1856 precedence over the name Sauromalus obesus (Baird 1858).

Subspecies: Sauromalus ater klauberi has been raised to species status. See also note for S. obesus (which is a synonym of S. ater according to Hollingsworth 1998). Buckley et al. 2016: 32 appear to consider multiforminatus and tumidus as synonyms of ater although they are not explicit about synonymy but rather list them under “other names”.

Type species: Sauromalus ater DUMÉRIL 1856 is the type species of the genus Sauromalus DUMÉRIL 1856.

NCBI taxonID: 65996 [australis] 
EtymologyNamed after its color, Latin “ater, atra, atrum” = dark or black. 
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