Elaphe

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
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: Elaphe
Species (16): E. anomala – E. bimaculata – E. cantoris – E. carinata – E. climacophora – E. davidi – E. dione – E. hodgsonii – E. moellendorffi – E. quadrivirgata – E. quatuorlineata – E. sauromates – E. schrenckii – E. taeniura – E. urartica – E. xiphodonta – E. zoigeensis
Name

Elaphe Fitzinger in Wagler, 1833: unnumbered page

Type species: Elaphe parreyssii Fitzinger in Wagler, 1833, by monotypy.
Gender: feminine.
Placed on the Official List of Generic Names in Zoology by Opinion 490 (1957: 181).

Synonyms

Leptophidium Hallowell, 1861: 497
Type species: Leptophidium dorsale Hallowell, 1860, by monotypy.
Phyllophis Günther, 1864: 295
Type species: Phyllophis carinata Günther, 1864, by monotypy.
Spaniopholis Mocquard, 1897: 216
Type species: Spaniopholis souliei Mocquard, 1897, by monotypy.
Orthriophis Utiger, Helfenberger & Schätti, 2002: 110 [synonymised by Chen et al. (2017: 216)]
Type species: Cynophis moellendorffi Boettger, 1886, by original designation.

References
Primary references

Wagler, J.G. 1828-1833. Descriptiones et icones Amphibiorum. Tres partes cum XXXVI tabulis. J.G. Cottae: Monachii, Stuttgartiae et Tubingae. unnumbered text + 36 pls. BHL Reference page.
Hallowell, E. [1860] 1861. Report upon the Reptilia of the North Pacific Exploring Expedition, under command of Capt. John Rogers, U. S. N. Proceedings of the Academy of Natural Sciences of Philadelphia 12: 480–510. BHL Reference page.
Günther, A.C.L.G. 1864. The Reptiles of British India. Ray Society: London. xxvii + 452 pp. BHL Reference page.
Mocquard, F. 1897. Notes herpétologiques. Bulletin du Muséum national d'Histoire naturelle 3(6): 211–217. BHL
International Commission on Zoological Nomenclature. 1957. Opinion 490. Action under the Plenary Powers to preserve for use in its accustomed sense the generic name Elaphe Fitzinger, 1833 (Class Reptilia). pp. 179–196 In Hemming, F. (ed.). Opinions and declarations rendered by the International Commission on Zoological Nomenclature Volume 17, part 12. International Trust for Zoological Nomenclature: London. 452 pp. BHL Reference page.
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.
Chen, X., Lemmon, A.R., Lemmon, E.M., Pyron, R.A. & Burbrink, F.T. 2017. Using phylogenomics to understand the link between biogeographic origins and regional diversification in ratsnakes. Molecular Phylogenetics and Evolution 111: 206–218. DOI: 10.1016/j.ympev.2017.03.017 Paywall Reference page.

Additional references

Jablonski, D., Kukushkin, O.V., Avcı, A., Bunyatova, S., Kumlutaş, Y., Ilgaz, Ç., Polyakova, E., Shiryaev, K., Tuniyev, B. & Jandzík, D. 2019. The biogeography of Elaphe sauromates (Pallas, 1814), with a description of a new rat snake species. PeerJ 7: e6944. DOI: 10.7717/peerj.6944 Open access Reference page.
Qi, S., Shi, J.-S., Ma, Y.-B., Gao, Y.-F., Bu, S.-H., Grismer, L.L., Li, P.-P. & Wang, Y.-Y. 2021. A sheep in wolf's clothing: Elaphe xiphodonta sp. nov. (Squamata, Colubridae) and its possible mimicry to Protobothrops jerdonii. ZooKeys 1048: 23–47. DOI: 10.3897/zookeys.1048.65650 Open access Reference page.

Links

Uetz, P. & Hallermann, J. 2022. Elaphe . The Reptile Database. Accessed on 28 June 2021.

Vernacular names
Deutsch: Kletternattern
English: Palearctic-Oriental Rat Snakes
հայերեն: Սահնօձ
русский: Лазающие полозы

Elaphe is a genus of snakes in the family Colubridae. Elaphe is one of the main genera of the rat snakes, which are found in many regions of the northern hemisphere. Elaphe species are medium to large constrictors by nature. Although all of the species in Elaphe are nonvenomous, bites from rat snakes are still irritably painful and can potentially cause bacterial infections due to the saliva.[2]

Based on the mitochondrial DNA analysis results, many species of Elaphe have been moved to the genera Bogertophis, Coelognathus, Gonyosoma, Orthriophis, Pantherophis, Rhinechis, Senticolis, Zamenis, and others.[3] Nevertheless, the generic name Elaphe is still widely used.

Physical characteristics

Like most of the rat snakes, Elaphe spp. generally have slender but sturdy bodies, square heads, and extremely flat bellies.[4] Their sizes range from large medium to very large,[1] growing even as large as 108 in. Elaphe spp. have large numbers of vertebrae, ribs, and ventral scales, but few rows of dorsal scales, which are characterized by having slight keels. In cross section, Elaphe spp. are shaped like a loaf of bread, the flat belly meeting the sides of the body at an angle.[5] This special physical characteristic is well observed in Elaphe obsoleta, whose belly scales curve upward. The curving of the ventral scales gives them better traction for tree climbing.[6] The color and pattern of Elaphe spp. are quite variable and hard to generalize.[6]

Internally, Elaphe spp. do not have any observable vestiges of hind limbs or coronoid bones of the lower jaw like any other members of the family Colubridae. Another important characteristic of Elaphe as part of the Colubridae is the presence of Duvernoy's gland, a modified salivary and digestive gland.[7] In addition, Elaphe spp. have equal and smooth maxillary teeth.[1] The teeth are small and slightly curved, and occur in several rows. The curvy teeth help fix the prey and prevent them from escaping once they are caught.[3]
Senses

Elaphe spp. have very developed sensory organs which support their daily activities. They have internal ears that enables them to detect sounds at low frequencies. The ears consist of a single ear bone connected to a jaw bone. Compared to other snakes, they are considered to have highly developed vision that allows them to detect the movement of their predators and prey. The eyes are protected by single transparent brilles. In addition, the analysis of scents is carried out by Jacobson's organ in the palate. To stimulate Jacobson's organ, they use their tongues to transport air and ground molecules to Jacobson's organ. Then, the organ analyzes the molecules to identify the scent. Among these detectable scents are pheromones, which indicate the presence of other snakes and their reproductive readiness.[3]

Lifecycle and behavior

The lifespan of Elaphe spp. is generally 15 years. Males live a little longer than females,[2] but some species, such as E. guttata, live up to 23 years in captivity.[8] However, species such as E. emoryi usually have lifespans around 2 years and 2 months.[1]

Elaphe spp. generally reach sexual maturity after 18–24 months. They usually mate with the opposite sex after emerging from hibernation in the spring. The gestation period of females is around 1.5 months, and females can lay up to 30 eggs in a clutch, the number varying between species. E. guttata and E. obsoleta, among others, are mostly diurnal, but some species are more active in late afternoon or at dusk. Throughout the year, they are usually active from April to October, followed by a period of hibernation.

Hibernation

Elaphe spp. hibernate, especially those that live in cold regions, because snakes are cold-blooded, which makes their body temperature susceptible to the temperature of their environments. Thus, they need to maintain their body energy by switching locations and remaining physiologically inactive when winter comes.[9] The hibernacula sites vary depending on the habitats they live in, yet the most important requirement of a hibernaculum site is that it needs to be frost-free. Some Elaphe spp. hibernate in the rock crevices, rock faults, and burrows.[10] However, other species choose to hibernate in rotting logs, roots of trees, and hollow spaces in elevated tree trunks. Some snakes that live close to human communities even hibernate in old wells and barns.[11]

Most members of the genus Elaphe start their hibernation in October and emerge again the next spring. The length of the hibernation varies by species. Elaphe longissima, or Aesculapian snake, may hibernate from October to May,[10] as it is among the northernmost occurring rat snakes. However, Elaphe bimaculata, or the twin-spotted rat snake, only needs to hibernate for 2 to 3 months.[12] Species such as black rat snakes, or E. obsoleta, hibernate with other rat snakes and/or many other snake species, most notably timber rattlesnakes, racers, and bull snakes.[13]

Reproduction

E. obsoleta generally starts to mate in late April, May, and early June after the winter hibernation. Males try to attract females with pheromones, as the females pass through their territories. Male Aesculapian snakes pursue female snakes until they can coil around them. They continue in such position, which is then followed by dancing for up to an hour before copulation, during which the male snake lines up with the female and holds her in place by wrapping his tail around hers and grasping her with his mouth. Then, the male everts one of his hemipenes into the female's cloaca. The mating process lasts from a few minutes to a few hours.[14]

After copulation, Elaphe spp. seek an appropriate place to lay the developing eggs. They usually lay eggs in the soft heart of a rotten log or in sandy soil under a rock. A good place for laying eggs is one that is damp but not wet, and warm but not hot. After laying eggs, the female snake covers them up with sand or soil, and then she leaves.[15] A few species remain with the eggs until they hatch about 9 weeks later.

Oviparous E. obsoleta lays 12–20 eggs under logs or leaves in late summer, which hatch in the fall. The adult snakes return to their hibernation dens in the late fall.[13] E. guttata breeds from March to May. The mating process is similar to E. obsoleta. E. guttata lays 10–30 eggs in late May to July. The eggs are generally not protected by the adults. After 60–65 days, the eggs hatch in July through September.[8]

Defense mechanisms

Many species of Elaphe are known for being nonaggressive and shy. They are prone to freeze their movements when they are shocked or encountering danger. This motionless response has contributed to many road kills of Elaphe. However, some Elaphe spp. tend to be more defensive if they are continually provoked. For example, Texas rat snakes, Elaphe obsoleta lindheimeri, are well known as one of the most snappy and combative rat snakes. In general, their defense system can be broken down to two levels. The first line of defense involves specific behaviors that they use to warn the intruders. One of the most common ways is by coiling their bodies and vibrating their tails, which simulates a rattle. Fox snakes, Elaphe vulpina and Elaphe gloydi, exemplify this type of defense mechanism; they mimic the rattling vibration of rattlesnakes when they encounter danger.[4] Another common way of defense is by smearing a foul-smelling musk on their predators. The musk is originated from the contents of the anal gland and the intestinal tract.[3] The second line of defense, generally involves striking the intruder with their teeth if they are further provoked.[14]
Locomotion

Elaphe spp. move forward using side-to-side, serpentine movement. They use the curvy ventral scales of their bodies to grab the rough ground surfaces and then push against the ground to generate a forward movement. Thus, the smoother the ground is, the harder it is for them to move across it.[3] Many species apply similar mechanisms when climbing trees.
Predation

Adult Elaphe spp. primarily prey on rodents (i.e., mice and chipmunks), bird eggs, and young birds. Juveniles feed on small lizards, young mice, and occasionally small frogs (i.e., tree frogs). Species such as E. guttata do not feed on a daily basis, but every few days. Elaphe spp. hunt by waiting motionless in a fixed position until the prey comes near enough to attack. Then, they strike the prey and bite it. They use their Jacobson's organ to track and find their prey.[9] Being constrictors, Elaphe spp. kill the prey by creating enormous pressure on the prey's chest. As they continue to coil more tightly, the pressure on the chest prevents the blood from circulating into the heart, which eventually leads to heart failure.[16] They usually bite the prey first to maintain their grip on the prey before they start this deadly mechanism. In addition, they do not chew their food, but swallow it whole.[8]
Habitat and distribution

Elaphe spp. live in a great variety of environments, depending on species and subspecies. Most are terrestrial or semiarboreal, but some burrow in sand or loose soil. For example, the black rat snake, E. obsoleta obsoleta, prefers heavily wooded habitats due to its excellent tree-climbing ability. Their population can be distributed throughout all elevations, usually on rocky hillsides of mountains, or on flat farmlands.[4] However, the yellow rat snakes, E. o. quadrivittata, prefers river swamps as habitats. Similarly, the Everglades rat snake, E. o. rossalleni, lives in the shrubs and trees along the water way.[14] The Elaphe snakes are spread throughout the Northern Hemisphere, but some occur in South America, Africa, Southeast Asia, and Northern Australia.[15] For instance, the black rat snakes are well distributed from Ontario, Canada, to Florida and Texas.[4] The population of E. guttata is widespread in many U.S. states, including Florida, Louisiana, southern New Jersey, and parts of Kentucky.[8]
Species

The following 17 species are recognized as being valid.[17]

Elaphe anomala (Boulenger, 1916)
Elaphe bimaculata Schmidt, 1925
Elaphe cantoris (Boulenger, 1894)
Elaphe carinata (Günther, 1864)
Elaphe climacophora (H. Boie, 1826)
Elaphe davidi (Sauvage, 1884)
Elaphe dione (Pallas, 1773)
Elaphe hodgsoni (Günther, 1860)
Elaphe moellendorffi (Boettger, 1886)
Elaphe quadrivirgata (H. Boie, 1826)
Elaphe quatuorlineata (Bonnaterre, 1790)
Elaphe sauromates (Pallas, 1811)
Elaphe schrenckii Strauch, 1873
Elaphe taeniura (Cope, 1861)
Elaphe urartica Jablonski, Kukushkin, Avcı, Bunyatova, Ilgaz, Tuniyev & Jandzik, 2019
Elaphe xiphodonta Qi, Shi, Ma, Gao, Bu, Grismer, Li & Wang, 2021
Elaphe zoigeensis Huang, Ding, Burbrink, Yang, Huang, Ling, Chen & Zhang, 2012

References

Wright, A.H.; Wright, A.A. (1957). Handbook of Snakes of the United States and Canada. Ithaca and London: Comstock Publishing Associates, a Division of Cornell University Press. 1,105 pp. (in 2 volumes). ("Rat Snakes", p. 209; Elaphe emoryi, longevity, p. 220).
"Rat snakes – lifecycle, predation, diet, bites, hibernation, pictures and videos." Ratsnake.org. (2010-09-23).
Bartlett, Patricia; Bartlett, R.D. (2006-05-26). Corn Snakes and Other Rat Snakes. Complete Pet Owner's Manual (2nd ed.). Hauppauge NY: Barron's Educational Series. ISBN 0-7641-3407-8.
Burton, M.; Burton, R. (1969). "Rat Snake". The International Wildlife Encyclopedia. (pp. 1909–1910). New York: Marshall Cavendish Corp.
Conant, Roger (1975). A Field Guide to Reptiles and Amphibians of Eastern and Central North America. Houghton Mifflin. Boston. xviii + 429 pp. + Plates 1–48. ISBN 0-395-19979-4 (hardcover), ISBN 0-395-19977-8 (paperback).
Wernert, S.J. (1982). Reptiles. Reader's Digest North American Wild Life. Pleasantville, New York: Reader's Digest. p. 174.
Evans, A.V.; Garrison, R.W.; Schlager, N.; Hutchins, M.; Murphy, J. (2004). Vol 7. Grzimeks' Animal life Encyclopedia (2nd ed., p. 467). Detroit: Thomson-Gale.
Barnard, S.M.; Hollinger, T.G.; Romaine, T.A. (1979). "Growth and Food Consumption in the Corn Snake, Elaphe guttata guttata". Copeia. 4 (4): 739–741. doi:10.2307/1443884. JSTOR 1443884.
George, L. (2002). Mating. Rat Snake (1 ed.), Mankato, Minnesota: Capstone Press, pp. 33–34, ISBN 0736809090.
Roots, C. (2006). Uncontrolled Hypothermia. Hibernation, Westport, Conn.: Greenwood Press, p. 88, ISBN 0313335443.
Werler, J.E.; Dixon, J.R. (2000). 39. Texas snakes: identification, distribution, and natural history, p. 118, Austin: University of Texas Press, ISBN 0292791305.
Knight, Rex; Korell, Sacha (1997). "Ratsnakes of the Genus Elaphe – Species." Ratsnakes of the Genus Elaphe – Introduction. OnlineHobbyist.com, Inc. (2010-09-22).
LeClere, Jeff (2005). Iowa Reptiles and Amphibians – Rat Snake – Elaphe obsoleta. HerpNet – Home. Web. 22 Sept. 2010.
Trepanowski, P. (2003). "Elaphe obsoleta", Animal Diversity Web. Accessed September 22, 2010.
Bateman, G. (1981). Snakes. The Encyclopedia of Reptiles and Amphibians (pp. 115, 117).
Ziesmann, Jurgen. (2010). Vertebrate Biology Lecture Notes: Reptiles 09. Azusa Pacific University
Genus Elaphe at The Reptile Database www.reptile-database.org.