Pantherophis

Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Cladus: Craniata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Lepidosauromorpha
Superordo: Lepidosauria
Ordo: Squamata
Subordo: Serpentes
Infraordo: Caenophidia
Superfamilia: Colubroidea

Familia: Colubridae
Subfamilia: Colubrinae
Genus: Pantherophis
Species (9): P. alleghaniensis – P. bairdi – P. emoryi – P. guttatus – P. obsoletus – P. ramspotti – P. slowinskii – P. spiloides – P. vulpinus
Name

Pantherophis Fitzinger, 1843: 25

Type species: Coluber guttatus Linnaeus, 1766, by original designation.

Synonyms

Scotophis Baird & Girard, 1853: 73
Type species: Coluber alleghaniensis Holbrook, 1836, by original designation.
Mintonius Collins & Taggart, 2008: 16 [synonymized by Pyron & Burbrink (2009: 528)]
Type species: Scotophis vulpinus Baird & Girard, 1853, by original designation.

References
Primary references

Fitzinger, L. 1843. Systema Reptilium. Fasciculus primus. Amblyglossae. Braumüller et Seidel: Wien. vi + 106 pp. BHL Reference page.
Baird, S.F. & Girard, C. 1853. Catalogue of North American Reptiles in the Museum of the Smithsonian Institution. Part 1.–Serpents. Smithsonian Institution: Washington. xvi + 172 pp. BHL Reference page.

Additional references

Utiger, U., Helfenberger, N., Schätti, B., Schmidt, C., Ruf, M. & Ziswiler, V. 2002. Molecular Systematics and Phylogeny of Old and New World Ratsnakes, Elaphe Auct., and Related Genera (Reptilia, Squamata, Colubridae). Russian Journal of Herpetology 9(2): 105–124. DOI: 10.30906/1026-2296-2002-9-2-105-124 [nonfunctional] Broken access Reference page.

Links

Uetz, P. & Hallermann, J. 2021. Pantherophis . The Reptile Database. Accessed on 27 June 2020.
Pantherophis – Taxon details on Integrated Taxonomic Information System (ITIS).

Vernacular names
English: North American Rat Snakes

Pantherophis is a genus of nonvenomous colubrid snakes endemic to central and eastern regions of North America. They consist of the North American ratsnakes, the foxsnakes, and the cornsnakes. The genus first appeared in the fossil record in the Middle Miocene around 16.3 million years ago. In total there are 10 species recognized to be valid. They are a large terrestrial snake genus that lack subocular scales. Originally classified in the genus Elaphe, phylogenetic studies have found this taxon to be closely related to Pituophis. As with all snakes Pantherophis is an obligate faunivore with a diet that consists of small mammals, birds, reptiles and amphibians, and even insects. While many species conservation status is categorized as "least concern", many local populations in some species have declined where some places have them listed as federally protected. The corn snake (P. guttatus) is a popular pet reptile, due to the availability of captive-bred animals, their low maintenance and calm disposition, and the variety of color morphs. There are other species of Pantherophis that are in the pet trade, though are not as popular as the corn snake.

Field Characteristics

Members in this genus are large terrestrial snakes, built for constriction. Pantherophis is characterized in having a divided cloacal plate, with at least some of the dorsal scales are keeled, which are faint, in more than 30 rows of dorsal scales, and all species lack the subocular scales.[4] Each of the 10 or so species can be further distinguished based on subtle characteristics, such as color markings, scale nuances, and geographic range.[4]
Systematics and Paleontology
Taxonomy

The taxonomy of Pantherophis has been a complicated area of research. The genus was named by the Austrian zoologist Leopold Fitzinger in 1843, with the type species being Pantherophis guttatus. Shortly after the genus was then considered to be a junior synomyn of the otherwise Old World genus Elaphe. However the morphology of the hemipenes, the dentition, and the squamatization found them to be closer to the genus Pituophis (gopher snakes, pine snakes, and bullsnakes). Soon subsequent molecular studies have found support of the sister grouping between Pituophis and the North American Elaphe species, which resulted in the resurrection of the genus Pantherophis.[5] These two genera, along with several other North American endemic taxa like Lampropeltis belong to the tribe Lampropeltini.[6]

There are currently four subgenera of Pantherophis with three extant and one extinct: Scotophis Baird & Girard, 1853 for the ratsnakes; †Palaeoelaphe Gilmore, 1938 for a Miocene fossil species;[2] Mintonius Collins & Taggart, 2008 for the foxsnakes, and Pantherophis for the cornsnake complex. There are few studies that suggested the possibility that Pantherophis is paraphyletic in respect to Pituophis.[7] To maintain taxonomic stability, some of the aforementioned subgenera are reevaluated into proper genera. This, however, has not been supported by larger scale molecular trees concerning the relationships of various snake taxa.[6][8][9]
Phylogeny

The interspecies relationships of Pantherophis usually has the subgenera Pantherophis and Mintonius being sister taxa, with Scotophis at the root of the genus. Below is the widely supported phylogenetic tree of the species in the genus.[6][8][9]

Pantherophis

Scotophis

	Pantherophis bairdi

	Pantherophis obsoletus

	Pantherophis alleghaniensis

	Pantherophis spiloides

Mintonius

	Pantherophis vulpinus

	Pantherophis ramspotti

Pantherophis

Pantherophis guttatus

	Pantherophis slowinskii

	Pantherophis emoryi

 
 
Extant Species

Below is the list of valid species with their distribution range.[3][10]

Scientific Name	English Name	Geographic Distribution
*Genus Pantherophis* Fitzinger, 1843**
*Subgenus Scotophis* Baird & Girard, 1853**
Pantherophis bairdi (Yarrow, 1880)	Baird's Ratsnake	Southwestern United States and adjacent northeastern Mexico.
Pantherophis obsoletus (Say, 1823)	Western Ratsnake or Texas Ratsnake	West of the Mississippi River, from eastern and southern Iowa southward through Missouri and Arkansas to western Louisiana, westward to eastern Texas, northward through Oklahoma and eastern Kansas to southeastern Nebraska.
Pantherophis alleghaniensis (Holbrook, 1836)	Eastern Ratsnake	United States east of the Apalachicola River in Florida, east of the Chattahoochee River in Georgia, east of the Appalachian Mountains, north to southeastern New York and western Vermont, eastern Pennsylvania, Maryland, South Carolina, North Carolina, Georgia, south to the Florida Keys.
Pantherophis spiloides (A.M.C. Duméril, Bibron & A.H.A. Duméril, 1854)	Gray Ratsnake	Eastern and Central United States, west of the Appalachian Mountains and east of the Mississippi River
*Subgenus Mintonius* Collins & Taggart, 2008**
Pantherophis ramspotti Crother, White, Savage, Eckstut, Graham & Gardner, 2011	Western Foxsnake	United States, west of the Mississippi River.
Pantherophis vulpinus (Baird & Girard, 1853)	Eastern Foxsnake	Eastern Great Lakes region of the United States, as well as adjacent western Ontario in Canada.
*Subgenus Pantherophis* Fitzinger, 1843**
Pantherophis guttatus (Linnaeus, 1766)	Red Cornsnake	Southeastern and central United States.
Pantherophis emoryi (Baird & Girard, 1853)	Great Plains Ratsnake	United States, from Missouri to Nebraska, to Colorado, south to Texas, and into northern Mexico Great Plains rat snake
Pantherophis slowinskii (Burbrink, 2002)	Slowinski's Cornsnake	Louisiana, eastern Texas, and Arkansas

Fossil Remains

The fossil record of Pantherophis is the Middle Miocene to Upper Pleistocene.[2] In addition to fossil remains of some of the extant species, there are fragmentary Pantherophis remains throughout the second half of the Cenozoic in North America. Only three fossil species have been described from more complete remains. The Early Pliocene P. buisi which might be a basal species in the Scotophis subgenus,[1] and P. kansensis which is the sole species in the subgenus of Palaeoelaphe, the trunk vertebrae similar to Mintonius subgenus.[1] P. nebraskensis was another valid species, but it is considered to be a junior synonym of P. kansensis.[2]
References

Holman, J.A. (2000). Fossil Snakes of North America: Origin, Evolution, Distribution, Paleoecology. Indiana University Press. ISBN 0253337216.
Wallach, V.; Williams, K.L.; Boundy, J. (2014). Snakes of the World: A Catalogue of Living and Extinct Species. CRC Press.
Crother BI (chair) (2012). Scientific and Standard English Names of Amphibians and Reptiles of North America North of Mexico. 7th ed. SSAR Herpetological Circular 39. 84 pp. PDF at SSAR. Accessed 4 July 2011.
Powell, R.; Collins, J.T.; Hooper, Jr., E.D. (2019). Key to the Herpetofauna of the Continental United States and Canada (3rd ed.). University Press of Kansas. ISBN 978-0700628902.
Utiger, Urs; Notker Helfenberger; Beat Schätti; Catherine Schmidt; Markus Ruf; and Vincent Ziswile (2002). "Molecular Systematics and Phylogeny of Old and New World Ratsnakes ..." Russian Journal of Herpetology 9 (2): 105-124.
Pyron, R. A.; Burbrink, F. T. (2009). "Neogene diversification and taxonomic stability in the snake tribe Lampropeltini (Serpentes: Colubridae)". Molecular Phylogenetics and Evolution. 52 (2): 524–529. doi:10.1016/j.ympev.2009.02.008. PMID 19236930.
Figueroa, A.; McKelvy, A. D.; Grismer, L. L.; Bell, C. D.; Lailvaux, S. P. (2016). "A species-level phylogeny of extant snakes with description of a new colubrid subfamily and genus". PLOS ONE. 11 (9): e0161070. Bibcode:2016PLoSO..1161070F. doi:10.1371/journal.pone.0161070. PMC 5014348. PMID 27603205.
Pyron; Burbrink; Wiens (2013). "A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes". BMC Evolutionary Biology. 13: 93. doi:10.1186/1471-2148-13-93. PMC 3682911. PMID 23627680.
Dahn, H. A.; Strickland, J. L.; Osorio, A.; Colston, T. J.; Parkinson, C. L. (2018). "Hidden diversity within the depauperate genera of the snake tribe Lampropeltini (Serpentes, Colubridae)". Molecular Phylogenetics and Evolution. 129: 214–225. doi:10.1016/j.ympev.2018.08.018. PMID 30189319.

Pantherophis. The Reptile Database. www.reptile-database.org.