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Pantherophis alleghaniensis (HOLBROOK, 1836)

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Higher TaxaColubridae, Colubrinae, Lampropeltini, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
Subspecies 
Common NamesE: Eastern Rat Snake (alleghaniensis)
E: Western Rat Snake (obsoleta)
E: Yellow Rat Snake (quadrivittata)
E: Everglades Ratsnake (rossalleni)
G: Erdnatter
E: Midland Rat Snake, Gray Rat Snake (spiloides)
G: Graue Pilotnatter (spiloides) 
SynonymColuber Alleghaniensis HOLBROOK 1836: 111
Coluber reticularis DAUDIN 1803 (part., see comment)
Scotophis alleghaniensis BAIRD & GIRARD 1853: 73
Elaphis spiloides DUMÉRIL, BIBRON & DUMÉRIL 1854: 269
Elaphis holbrookii DUMÉRIL & BIBRON 1854: 272
Elaphis alleghaniensis — HALLOWELL 1856: 243
Elaphis obsoletus var. alleghaniensis — GARMAN 1884: 54
Coluber obsoletus lemniscatus COPE 1888: 386 (part.)
Elaphe quadrivittata deckerti BRADY 1932
Elaphe williamsi BARBOUR & CARR 1940: 340
Elaphe quadrivittata parallela BARBOUR & ENGELS 1942: 103
Elaphe obsoleta rossalleni NEILL 1949
Elaphe obsoleta williamsi — BECHTEL 1978: 521
Elaphe obsoleta deckerti — STANKOWSKI 1986
Elaphe obsoleta obsoleta — CONANT & COLLINS 1991: 197
Elaphe obsoleta rossalleni — CONANT & COLLINS 1991: 198
Elaphe obsoleta spiloides — CONANT & COLLINS 1991: 199
Elaphe obsoleta spiloides — CROTHER 2000: 62
Elaphe obsoleta spiloides — TENNANT & BARTLETT 2000: 386
Elaphe obsoleta obsoleta — CROTHER 2000: 62
Elaphe obsoleta rossalleni — CROTHER 2000: 62
Elaphe obsoleta rossalleni — TENNANT & BARTLETT 2000: 385
Elaphe alleghaniensis — BURBRINK 2001
Elaphe spiloides — BURBRINK 2001
Elaphe obsoleta spiloides — TENNANT 2003: 395
Pantherophis obsoletus spiloides — SCHMIDT & KUNZ 2005: 68
Pantherophis obsoletus spiloides — LOVE 2007
Scotophis spiloides — COLLINS & TAGGART 2008
Elaphe obsoleta spiloides — ENGE 2009
Pantherophis spiloides — PYRON & BURBRINK 2009
Scotophis spiloides — MESHAKA & COLLINS 2009
Scotophis alleghaniensis — MESHAKA & COLLINS 2009
Pantherophis alleghaniensis — PYRON & BURBRINK 2009
Scotophis spiloides — SAVIOLA et al. 2012
Pantherophis spiloides — WALLACH et al. 2014: 530
Pantherophis alleghaniensis — WALLACH et al. 2014: 528
Pantherophis alleghaniensis — BURBRINK et al. 2021
Pantherophis obsoletus alleghaniensis — HILLIS 2022
Pantherophis obsoletus spiloides — HILLIS 2022 
DistributionUSA (east of the Appalachian Mountains, north to Ohio, SE New York, W Vermont, Pennsylvania, Maryland, South Carolina, North Carolina, Georgia [east of the Chattohoochee River], Florida [east of the Apalachicola River]).

Type locality: “summit of the blue Ridge in Virginia”, not “highlands of the Hudson” (see Pyron & Burbrink 2021)

spiloides: Canada (S Ontario), USA (west of the Apalachicola River and the Appalachian Mountains and east of the Mississippi River. South from Louisiana along the gulf Coast of the US to Florida, and north in W New York, Maryland, Pennsylvania, S Michigan, NE Indiana, C Illinois, SW Wisconsin, Tennessee). Type locality: La Nouvelle-Orleans [= New Orleans, Louisiana]  
Reproductionoviparous 
TypesLectotype: ANSP 16792; paralectotype: MNHN-RA 0.829 (this specimen is also one of two primary syntypes of Elaphis holbrookii Duméril, Bibron & Duméril, 854, see Pyron & Burbrink 2021)
Syntypes: MNHN-RA 0.829, MNHN-RA 0.7256 [Elaphis holbrookii]
Lectotype: USNM 248870 (skin only, originally USNM 4710), designated by Burbrink et al. 2021; paralectotypes: ANSP 10893, USNM 8798 [Coluber obsoletus lemniscatus]
Holotype: MCZ 46468 [parallela]
Holotype: ANSP 3773; South Carolina (?) [quadrivittatus]
Holotype: Florida State Museum of Natural History [rossalleni]
Holotype: MCZ 45705, adult male [williamsi]
Holotype: USNM 84295 [Elaphe quadrivittata deckerti]
Holotype: MNHN-RA 0827 [spiloides]
Holotype: USNM 1563 [Scotophis confinis] 
Diagnosis 
CommentHybrids: BRÖER (1977) reports a hybrid between E. guttata guttata and E. obsoleta quadrivittata. Bechtel & Mountain (1960) report hybrids between Elaphe guttata and E. obsoleta rossaleni. A complete genome of Pantherophis alleghaniensis was shown to be a hybrid of P. obsoletus and P. alleghaniensis (Burbrink et al. 2023).

Distribution: see maps in BURBRINK 2001, Burbrink et al. 2021.

The diagnosis of E. alleghaniensis by BURBRINK (2001) is unfortunately not very useful as it gives mostly relative characters (”E. alleghaniensis tends to have a higher number of subcaudals than E. spiloides” etc.) and hardly any comparisons with other species.

Phylogenetics: Burbrink et al. 2020 recognized three species (P. obsoletus (western clade), P. alleghaniensis (eastern), and P. spiloides (central ) but then discovered that they type of P. alleghaniensis actually was within the central clade (only noted in the supplemental material). The correct clade names are thus: P. obsoletus (western; west of the Mississippi River), P. alleghaniensis (central), and P. quadrivittatus (eastern) (T. Taggart, pers. comm., 20 Oct 2021).

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

Synonymy: Coluber reticularis DAUDIN 1803 is a binomical based on La Cépède (1789) name ‘La réticulaire’, but unavailable because the latter was suppressed by ICZN opinion 1463. See Pyron & Burbrink 2021 and Burbrink et al. 2021 for details.

NCBI taxonomy ID: 648198 [spiloides] 
EtymologyNamed after the Allegheny Mountains (a part of the Apalachians) or the Allegheny River in west-central Pennsylvania. The meaning of the word, which comes from the Lenape (Delaware) Native Americans, is not definitively known but is usually translated as "fine river" (Wikipedia). 
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