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Pituophis catenifer (BLAINVILLE, 1835)

IUCN Red List - Pituophis catenifer - Least Concern, LC
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Higher TaxaColubridae, Colubrinae, Lampropeltini, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
SubspeciesPituophis catenifer affinis HALLOWELL 1852
Pituophis catenifer annectens BAIRD & GIRARD 1853
Pituophis catenifer catenifer (BLAINVILLE 1835)
Pituophis catenifer deserticola STEJNEGER 1893
Pituophis catenifer fulginatus KLAUBER 1946
Pituophis catenifer pumilus KLAUBER 1946
Pituophis catenifer sayi (SCHLEGEL 1837) 
Common NamesE: Gopher Snake
E: Sonoran Gopher Snake [affinis]
E: San Diego Gopher Snake [annectens]
E: Central Baja California Gopher Snake [bimaris]
E: Pacific Gophersnake [catenifer]
E: Great Basin Gopher Snake [deserticola]
E: Santa Cruz Gopher Snake [pumilus]
E: Bullsnake [sayi]
S: Cincuate Casero 
SynonymColuber catenifer BLAINVILLE 1835: 290
Pituophis catenifer — BAIRD & GIRARD 1853: 69
Pituophis wilkesii BAIRD & GIRARD 1853: 71
Pityophis heermanni — HALLOWELL 1853
Elaphis reticulatus DUMÉRIL, BIBRON & DUMÉRIL 1854: 246
Pituophis mexicanus DUMÉRIL, BIBRON & DUMÉRIL 1854: 256
Pityophis catenifer — GARMAN 1884: 52
Pityophis catenifer var. mexicanus — GARMAN 1884: 52
Coluber catenifer — BOULENGER 1894: 67
Pituophis catenifer rutilus — VAN DENBURGH 1920
Pituophis catenifer stejnegeri — VAN DENBURGH 1920
Pituophis catenifer coronalis KLAUBER 1946: 19
Pituophis melanoleucus catenifer — HIRSCHKORN 1986
Pituophis catenifer — LINER 1994
Pituophis catenifer — ERNST & ERNST 2003: 272
Pituophis melanoleucus catenifer — BOL 2007
Pituophis catenifer — WALLACH et al. 2014: 561

Pituophis catenifer affinis HALLOWELL 1852
Pituophis affinis HALLOWELL 1852
Pituophis sayi affinis — TAYLOR 1938: 493
Pituophis sayi affinis — SMITH 1939
Pituophis sayi affinis — STULL 1940
Pituophis catenifer affinis — FUGLER 1956
Pituophis catenifer affinis — FUGLER & DIXON 1961
Pituophis melanoleucus affinis — BOGERT 1966
Pituophis melanoleucus affinis — HARDY 1969: 191
Pituophis melanoleucus affinis — WEBB 1984
Pituophis melanoleucus affinis — STEBBINS 1985: 189
Pituophis melanoleucus affinis — TANNER 1985: 638
Pituophis melanoleucus affinis — CONANT & COLLINS 1991: 204
Pituophis catenifer affinis — COLLINS 1997
Pituophis catenifer affinis — TENNANT & BARTLETT 2000: 371
Pituophis catenifer affinis — CROTHER 2000: 68
Pituophis catenifer affinis — HIRSCHKORN & SKUBOWIUS 2011
Pituophis catenifer affinis — LOVE 2011
Pituophis catenifer affinis — SKUBOWIUS 2012

Pituophis catenifer annectens BAIRD & GIRARD 1853
Pituophis annectens BAIRD & GIRARD 1853: 72
Pituophis catenifer annectens — STULL 1940
Pituophis catenifer annectens — KLAUBER 1946: 25
Pituophis melanoleucus annectens — STEBBINS 1985: 190
Pituophis melanoleucus annectans [sic] — HIRSCHKORN 1986
Pituophis melanoleucus annectens — LINER 1994
Pituophis melanoleucus annectens — GRISMER et al. 1996
Pituophis catenifer annectens — RODRIGUEZ-ROBLES & JESUS-ESCOBAR 2000
Pituophis catenifer annectans [sic] — MATTISON 2007: 29
Pituophis catenifer annectens — HIRSCHKORN & SKUBOWIUS 2011: 94

Pituophis catenifer catenifer (BLAINVILLE 1835)
Pituophis catenifer catenifer — STULL 1940
Pituophis melanoleucus catenifer — STEBBINS 1985: 190
Pituophis catenifer catenifer — CROTHER 2000: 68
Pituophis catenifer catenifer — HIRSCHKORN & SKUBOWIUS 2011

Pituophis catenifer deserticola STEJNEGER 1893: 206
Pituophis catenifer deserticola — VAN DENBURGH & SLEVIN 1921
Pituophis catenifer deserticola — FITCH 1939
Pituophis catenifer deserticola — STULL 1940
Pituophis catenifer deserticola — MITTLERMAN 1942
Pituophis catenifer deserticola — BRADLEY & DEACON 1966
Pituophis melanoleucus deserticola — PARKER 1973
Pituophis melanoleucus deserticola — STEBBINS 1985: 190
Pituophis catenifer deserticola — CROTHER 2000: 68
Pituophis catenifer deserticola — HIRSCHKORN & SKUBOWIUS 2011

Pituophis catenifer fulginatus KLAUBER 1946
Pituophis catenifer fulginatus KLAUBER 1946: 14
Pituophis catenifer fuliginatus — RODRÍGUEZ-ROBLES & DE JESÚS-ESCOBAR 2000
Pituophis catenifer bimaris — LOVICH et al. 2008: 28

Pituophis catenifer pumilus KLAUBER 1946: 41
Pituophis melanoleucus pumilis — STEBBINS 1985: 190
Pituophis catenifer pumilis — CROTHER 2000: 68
Pituophis catenifer pumilis — LOVICH et al. 2008: 28
Pituophis catenifer pumilus — HIRSCHKORN & SKUBOWIUS 2011
Pituophis catenifer pumilis — ALLF et al. 2020 (in error)

Pituophis catenifer sayi (SCHLEGEL 1837)
Coluber Sayi SCHLEGEL 1837: 157
Coluber Sayi DEKAY 1842: 41 (fide DUMÉRIL et al. 1854)
Coronella Sayi — HOLBROOK 1842: 99
Pituophis mcclellanii BAIRD & GIRARD 1853: 68
Pituophis mcclellanii — BAIRD & GIRARD in MARCY & MCCLELLAN 1854: 196
Ophibolus Sayi — BAIRD & GIRARD 1853: 84
Coronella Sayi — DUMÉRIL, BIBRON & DUMÉRIL 1854: 619
Pityophis catenifer var. sayi — GARMAN 1884: 52
Pituophis sayi — FORCE 1928
Pituophis sayi sayi — BURT 1935
Pituophis sayi sayi — SMITH 1939
Pituophis sayi sayi — STULL 1940: 91
Pituophis catenifer sayi — JAMESON & FLURY 1949
Pituophis catenifer sayi — SHANNON & SMITH 1949
Pituophis melanoleucus — CONANT & COLLINS 1991: 203
Pituophis catenifer sayi — LINER et al. 1993
Pituophis melanoleucus sayi — LINER 1994
Pituophis catenifer sayi — COLLINS 1997
Pituophis catenifer sayi — TENNANT & BARTLETT 2000: 369
Pituophis sayi — RODRÍGUEZ-ROBLES & JESUS-ESCOBAR 2000
Pituophis catenifer sayi — MATTISON 2007: 50
Pituophis catenifer sayi — JOHNSON et al. 2020 
DistributionCanada (British Columbia, Alberta, Saskatchewan),
USA (Washington, Oregon, Wisconsin, Illinois, Indiana, Kansas, South Dakota, Iowa, California, New Mexico, Arizona),
Mexico (Baja California, Aguascalientes, Nuevo León, Sonora, Durango, Hidalgo, Zacatecas, N Veracruz, San Luis Potosí, Hidalgo)

affinis: USA (W Texas, New Mexico, Arizona, SE California), Mexico (Zacatecas and S Sinaloa, Sonora, San Luis Potosi, Coahuila, Chihuahua); Type locality: New Mexico.

annectens: USA (California), Mexico (N Baja California); Type locality: San Diego, California.

catenifer: USA (N California, W Oregon, W Washington), Canada (S British Columbia); Type locality: California.

coronalis (invalid): California: Coronado Island; Type locality: Coronado Island

deserticola: USA (E Washington, Oregon, E California, Nevada, Idaho, N Arizona, NW New Mexico, W Colorado, Utah, SW Wyoming); Type locality: Great Basin and southwestern deserts.

fulginatus: Mexico (Baja California, San Martín Island); Type locality: San Martín Island.

mexicanus (invalid): Mexico (Nuevo Leon); Type locality: restricted to Sabinas Hidalgo in northern Nuevo León by Smith & Taylor (1950).

pumilus: USA (California: Santa Cruz Island); Type locality: Santa Cruz Island

sayi: Canada (S Alberta, S Saskatchewan), USA (Montana, SW North Dakota, South Dakota, Illinois, Indiana, Iowa, Colorado, Wyoming, Kansas, E New Mexico, Minnesota, Nebraska, Oklahoma (BURT 1935), Wisconsin, Texas), Mexico (Chihuahua, Coahuila, Tamaulipas); Type locality: Missouri.  
Reproductionoviparous (Van Denburgh 1898: 139). 
TypesHolotype: MNHN-RA 0057
Holotype: CAS 33869 [affinis]
Syntypes USNM 1839 (3, of which 2 are heads) [annectens]
Holotype: SDNHM = SDSNH 32621; paratypes SDNHM = SDSNH [bimaris]
Holotype: SDNHM = SDSNH 20229; paratype SDNHM = SDSNH 11365 [coronalis]
Holotype: SDNHM = SDSNH 17449; paratype SDNHM = SDSNH 17463, 17464 (topotypes) [fuliginatus]
Holotype: USNM 1540 [Pituophis mcclellanii]
Lectotype: USNM 1415 (designated by Schätti et al. 2020), paralectotype: MNHN-RA 3188 [Pituophis mexicanus]
Holotype: SDNHM = SDSNH 17238 [pumilus]
Holotype: unknown fide Smith & Taylor 1945: 107 [sayi]
Holotype: CAS 14203 [stejnegeri]
Holotype: USNM 5471 [Pituophis wilkesii]
Holotype: SDNHM 17449 (also given as SDSNH), collected by Lewis W. Walker on 11 July 1939 [fulginatus]
Holotype: USNM 18070, collected by C. Hart Merriam on 11 May 1891 [deserticola] 
Diagnosis 
CommentSubspecies: Some authors recognize Pituophis melanoleucus annectens (e.g. http://elib.cs.berkeley.edu/photos/fauna/cas-herp.html). COLLINS (1997) doesn’t recognize the subspecies bimaris, coronalis, fulginatus, insulanus, vertebralis but LINER (1994) does. Others consider some of them (like vertebralis) as valid species (e.g. GRISMER 1997). GRISMER 2002 considers P. c. bimaris as pattern variant of P. vertebralis. P. c. sayi is closely related to affinis and P. ruthveni (Rodríguez-Robles & De Jesús-Escobar, 2000). We do not recognize P. c. coronalis following Crother et al. 2012.

Hybridization: Pituophis catenifer sayi × Pantherophis vulpinus hybridize in nature (LeClere et al. 2012).

Distribution: see map in Schätti et al. 2020: 503 for distribution in Mexico (Fig. 14). 
EtymologyProbably named after the chain-like pattern (Latin “catena” = chain, -fer = carrying). P. c. sayi was named after Thomas Say (1787-1834), American naturalist. For biographical notes see Conniff 2011: 120ff. The subspecific name affinis is a Latin word meaning "related," applied in recognition of its relationship to P c. sayi. 
References
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