Taxonomy: Kingdom: Animalia Phylum: Chordata Class: Amphibia Order: Caudata Family: Cryptobranchidae Genus: Cryptobranchus Species alleganiensis Subspecies alleganiensis, bishopi Information from The Virginia Fish and Wildlife Information Service Image above this paragraph was taken from http://www.rieo.net/amph/saramand/sansyouo/ameandri.htm; permission was asked, but no response given, so permission was assumed. Click image to open up a larger version in new window. Taxonomic history taken directly from Catalogue of American Amphibians and Reptiles. It is account 101 and was written by Harold A. Dundee in 1971. For the full references for the citations in this portion of the website, click here. Cryptobranchus alleganiensis alleganiensis Salamandra alleganiensis Daudin, 1803, 8:231. Type-locality, Allegheny Mountains in Vervinia. Daudin wrote that Sonnini and Latreille, 1801, 2:253, pl. 54, gave the name, "La Salamandre des Monts Alleganis," but there was no technical proposal. Harper, 1940:721 restricted the type-locality to "The vicinity of Davenport's Plantation," North Toe River, Mitchell County, North Carolina, or about four miles ENE of the present-day town of Spruce Pine. No holotype is known to exist. Salamandra horrida Barton, 1808:7-8. Type-locality, "the great lakes of our contry, in the waters of the Ohio and Susquehanna, and other parts of the United States." Schmidt (1953) restricted the type-locality to the Muskingum River, Ohio. Salamandra gigantea Barton, 1808:8. Substitute name for S. horrida Salamandra maxima Baront, 1808:8. Substitute name for S. giantea Molge gigantea: Merrem, 1820:187. Transfer of S. gigantea Barto to Molge Cryptobranchus salamandroides Leuckhart, 1821:260, pl. 9. Substitue name for gigantea In Opinions of the International Congress of Zoological Nomeclature 1956-07, ID 365-388, direction 57, the name gigantea is regarded as a junior synonym of alleganiensis Daudin (1803), as published in combination Salamandra alleganiensis Urotropis mucronata Rafinesque, 1822:3. See Eurycea muscronata, following. Description is based on an animal found in the Kentucky River in 1821. Abranchus alleghaniensis: Harlan, 1825:233. In the Opinions of the International Congress of Zoological Nomenclature 1956-7, ID: 337-364, direction 56, the emendation of alleghaniensis is regarded as invalid although occupied. Protonopsis horridia: Barnes, 1826:233. Credits this name to "Barton (LeConte)" but the name does not appear in Barton's writings of 1808 and 1812. The name Protonopsis was mentioned by LeConte, 1824: 52-58, but not with an accompaying specific ephitet. Salamandrops gigantea: Wagler, 1830:209 (not seen). Eurycea mucronata: Rafinesque, 1832:121. Description and locality suggests that this is the same animal described by Rafinesque in 1822 as Urotropis mucronata. Cryptobranchus alleganiensis: Van der Hoevan, 1837:384. First proper application of generic name and specific epithet, except for altered spelling of specific epithet. Menopoma fuscam Holbrook, 1842:43 Crytobranchus alleghaniensis: Cope, 1887:43. Cryptobranchus alleganiensis: Stejneger and Barbour, 1917:3. First correct combination spelling in current usage. Cryptobranchus terrasodatcylus Wellborn, 1936:63. Based on abberant four-toed specimen. Type-locality "North America." Crytpbranchus alleganiensis alleganiensis: Schmidt, 1953:11. First usage of trinomial. Cryptobranchus alleganiensis bishopi Cryptobranchus bishopi: Grobman, 1943:6. Type-locality, "Current River at Big Spring Park, Carter County, Missouri." Holotype University of Michigan Museum of Zoology 68930, collected by Edwin P. Creaser on August 25, 1930. Cryptobranchus alleganiensis bishopi: Schmidt, 1953:12. New combination. Common names: Hellbender (most common), devil dog, Allegheny alligator, mollyhugger, mud cat Description: 30 - 74 cm in length. Flattened body and head with large and strongly keeled tail that is shorter than the body. Can be distinguished from mudpuppies by the lack of gills in adults; gill slits, though, are retained in adults, and are present on either side of the throat and hidden by flaps of skin. Muscular arms and legs; four digits on front limb and five on the rear. (Cogger and Zweifel 1998). Variable color, ranging from brown or olive as shown to a bright red-orange as seen in this photo or this photo. Top may or may not be marked (see subspecies descriptions below), and patterning tends to fade in preserved specimens (The Virginia Fish and Wildlife Information Service 2001); belly is lighter than top and has fewer markings. Male is smaller than female and has swollen ridge around vent during breeding season. Cryptobranchus alleganiensis alleganiensis: Black spotting variable and throughout body. Cryptobranchus alleganiesis bishopi: Heavily marked with black on back and tail. Gill slits are small and occaisionally closed in adults. The best way to distinguish between alleganiensis and bishopi is geography. All images in this section were described as being of Cryptobranchus alleganiesis alleganiesis. Image above and left was taken by Jeff Humphries and is used with permission. Image above and right was taken by John White and is also used with permission. Image directly to the right of this paragraph is from http://www.rieo.net/amph/saramand/sansyouo/ameandri.htm; permission was asked, but no response given, so permission was assumed. Click any image to open up a larger version in new window. Range: Cryptobranchus alleganiensis alleganiensis is more widely spread, ranging from the Susquehanna River in New York to the northern parts of Alabama and Georgia; it has been found as far east as New York and as far west as Indiana, with a large, isolated population in Missouri and Arkansas. Cryptobranchus alleganiensis bishopi is found only in the Black River and the White River Ozark mountains of Missouri and Arkansas. Map at right indicates range of the hellbender: the range of Cryptobranchus alleganiesis alleganiesis is black, whereas the range of Cryptobranchus alleganiesis bishopi is indicated in light grey. Map edited from the Atlas of Amphibians in Tennessee. Habitat: Completely aquatic. Lives in cool, clean streams with low siltation and rocky bottoms. Humphries (2002) indicates that it is frequently found in trout streams as well as fast rapids. Preferred depth is debated, but all sources indicate that it is less than three meters. Breeding: Breeding season is generally from August to September, but Humphries (2002) reports that breeding may occur in winter in Missouri, and Peterson, Ingersol, and Wilkerson (1989) report it in Arkansas. In all areas, the male will build a nest site underneath rocks, logs, or other river debris, with the hole of the den facing away from the river current (Petranka, 1998); he will then persuade or force gravid females to enter the nest site, and several females may lay their eggs in a given nest within a single Herman 1999) These pale to bright-yellow eggs are paired in "rosary-like" (Ibid.) strings that form an egg mass, and the male will then spread his milt on them. In the course of a breeding season, more than one thousand eggs may be laid or fertilized at a single nest. Although all sources agree that this species provides parental care, various authorities disagree as to the provider and type of care. Whereas our textbook, "Herpetology: An Introductory Biology of Amphibians and Reptiles" (Zug, Vitt and Caldwell 2001) accurately cites Sherman Bishop (1941) in that females remain at the nest and increase oxygen availability by moving eggs around, a 1996 article states that the male remains at the nest site to guard the eggs. (Danch) Even more interestingly, the Hood Department of Biology's Amphibian Ecology class website states that the male guards the eggs, but in the following sentence cites the 1941 Bishop citation that our text does. Incubation lasts for 62-75 days in Pennsylvania and New York and 45 days in Missouri. (The Virginia Fish and Wildlife Information Service 2001) Image of Cryptobranchus eggs above by Joyce Sun. Please ask for permission before usage. Development: At hatching, the larvae are about 30 millimeters long, with stubby forelegs and paddle-shaped hind-legs with a belly that Bishop (1943) describes as "clear, yellow, yolk colored" and gills that appear as "short, flattened filaments." The larvae retain their external gills for approximately eighteen months or until they reach approximately four inches in length; at this point, they undergo incomplete metamorphasis. The external gills are folded into the body, and the adult retains internal gill arches but respires via cutaneous surface; the adult also lacks eyelids. Hellbenders grow approximately two centimeters each year, and may continue to grow for five or six years after they reach sexual maturity between the ages of five and eight. (Herman 1999) Image in this section taken by Jeff Humphries and used with permission. Lifespan: The average age of Cryptobranchus in the wild is unknown, but in captivity, they have lived as long as twenty-nine years. Behavior: The hellbender is a nocturnal animal that is most active approximately two hours after dark and between the months of May and September. (Humphries and Pauley 2001) Extremely territorial, these animals defend an average range of 346 square meters in Pennsylvania (The Virginia Fish and Wildlife Information Service 2001), and investigators in the field report that it is common to see adults missing several fingers from both hands and feet, presumably due to territorial combat; the aberrant four-toed specimen that led Wellburn to conclude the existence of a new species may have been a result of this. Contrary to popular belief, they do not feed on trout or any other fish; rather, they eat a variety of stream invertebrate life, particularly worms and crayfish. All members of the Cryptobranchus genus, though, do exhibit an unusual method of feeding known as asymmetrical suction feeding the animal opens one side of its mouth, then moves the bilateral elements of the mandibles and hyoid separately to direct a unidirectional flow of water into the mouth. (Elwood and Cundall 1994). Conservation status: Population declines through its range caused the hellbender to be listed as a Category 2 species by the US Fish and Wildlife Service in 1989, meaning that they were a candidate for protection under the Endangered Species Act but that insufficient information was known about the species to conclusively move it into candidate status. The USFWS has since discontinued the Category 2 species classification (United States Fish and Wildlife Service 1996), but the hellbender remains on the State of Virginia's "State Special Concern" list. Hellbenders are particularly vulnerable due to their slow sexual maturation and sensitivity to declining water quality. Museums with Cryptobranchus alleganiensis: Cryptobranchus alleganiensis is not a rare species, and it is commonly used as a model in comparative anatomy classes; thus, most museums have Cryptobranchus alleganiensis in their collections. Listed below several institutions that do, the number that they possess, and the number of the bishopi subspecies that they posses; the Yale and the Auburn collections did not specify whether their specimens were alleganiesis or bishopi: The Yale Peabody Museum of Natural History (2; unspecified) The Field Museum (53; 0 bishopi) Auburn University (38; unspecified) Illinois Natural History Survey Amphibian and Reptile Collection (21; 1 bishopi) The Museum of Comparative Zoology at Harvard University(28; 7 bishopi) Selected Papers: Guimond, R and Hutchinson, V. "Aquatic Respiration - Unusual Strategy in Hellbender Cryptobranchus alleganiensis allegiensis (Daudin)." Science 182 (4118): 1263-1265 1973 Humpheries, W. Jeffrey, Thomas K Pauley. "Seasonal changes in nocturnal activity of the hellbender, Cryptobranchus alleganiensis, in West Virginia." Journal of Herpetology 34 (4): Dec 2000 Jerrett DP, Mays CE. "Comparative Hematology of Hellbender, Cryptobranchus Alleganiensis in Missouri." Copeia (2): 1973, pp. 331-337 Peterson, Chris L, Chris A Ingersol, Robert F Wilkerson. "Winter breeding of Cryptobranchus allenganiensis bishopi in Arkansas." Copeia (4): Dec 27 1989. p. 1031-6 Reese, Albert M. "Anatomy of Cryptobranchus allegheniensis." American Naturalist 40 (472): April 1906, pp. 287-326. Routman, Eric, Rosalind Wu, and Alan R Templeton. "Parsimony, molecular evolution, and biogeography: the case of the North American giant salamander." Evolution 48: Dec 1994 p. 1799-809. This species has an extremely extensive history of scholarship, and thus, a full list was impractical. For papers that were used in the preparation of this website, please see the references section. Also see the papers section of The Hellbender Homepage. Useful Links: Members of Cryptobranchus utilize an unusual method of feeding which, as mentioned above, is known as asymmeterical suction feeding. A video of this is shown here: http://socrates.berkeley.edu/~deban/Cryptomovie.html A great deal of the information on this webpage comes from Jeff Humphries, who wrote his master's thesis on these creatures, and his hellbender webpage has an enormous amoutn of information, including a very thorough listing of publications on the hellbender. He's also a great guy and incredibly enthusiastic about the species. :) The Hellbender Homepage Picture list: Hellbender at bottom of river: Courtesy Jeff Humphries Hellbender profile: Courtesy of John White Hellbender juvenile:Courtesy Jeff Humphries Full-body shot of hellbender. Notice strongly flattened body: Photo from http://www.rieo.net/amph/saramand/sansyouo/ameandri.htm . Head shot of an ugly bastard: Photo from http://www.rieo.net/amph/saramand/sansyouo/ameandri.htm . Cryptobranchus eggs: Image by Joyce Sun. References: The Amphibian Ecology Course at Hood College. http://www.hood.edu/academic/biology/hellbender.html Bishop, Sherman C. "Handbook of Salamanders: the Salamanders of the United States, of Canada, and of Lower California." Ithaca, NY: Comstock Publishing Company, Inc., 1943. Bishop, Sherman C. "The salamanders of New York." N. Y. State Museum Bulletin (324): 1941. Cogger, H and R Zweifel. 1998. Encyclopedia of Reptiles and Amphibians, 2nd edition. Academic Press, San Diego Danch, J. July 1996. "The Hellbender." Reptiles 4:48-59. Dundee, Harold A. "Cryptobranchus alleganiensis." Catalogue of American AMphibians and Reptiles. American Society of Icthyologists and Herpetologists: 1971. Account 101. Elwood, J. R. L. and D. Cundall. "Morphology and behavior of the feeding apparatus in Cryptobranchus allegeniensis." Journal of Morphology. 220:47-70. "Endangered and Threatened Wildlife and Plants; Notice of Final Decision on Identification of Candidates for Listing as Endangered or Threatened." Notice of Final Decision by the United States Fish and Wildlife Service, 1996. http://endangered.fws.gov/policy/cand_det.htm Harper, Francis. "Some Works of Bartram, Daudin, Latreille, and Sonnini, and Their Bearing Upon North American Herpetological Nomenclature." American Midland Naturalist 23: May 1940, pp. 720-721. Accessed through subscription service JSTOR at http://links.jstor.org/sici?sici=0003-0031%28194005%2923%3A3%3C692%3ASWOBDL%3E2.0.CO%3B2-T "Hellbender, eastern." The Virginia Fish and Wildlife Information Service. Written December 04, 2001. Last updated December 04, 2001. http://vafwis.org/BOVA/BOOKS/020020.HTM Herman, John. Animal Diversity Web. http://animaldiversity.ummz.umich.edu/accounts/cryptobranchus/c._alleganiensis$narrative.htmLast updated 1999. Humphries, Jeff. The Hellbender Homepage. Last updated February 2002. Peterson, Chris L, Robert F. Wilkinson Jr, and Milton S Topping. Age and growth of the Ozark hellbender (Cryptobranchus alleganiensis bishopi). Copeia: February 1983. pp. 225-31 Petranka, J.W. 1998. Salamanders of the United States and Canada. Smithsonian Institution Press, Washington, DC. Phillips, Chris. Illinois Natural History Survey Amphibian and Reptile Collection. http://www.inhs.uiuc.edu/cbd/collections/AmphReptColln/herps.html Redmond, William H. and Scott, A. Floyd. Atlas of Amphibians and Reptiles in Tennessee. Written 1996. Website updated November 1, 2001. http://www.apsu.edu/amatlas/ Zug, George R, Laurie J. Vitt, and Janalee P. Caldwell. "Herpetology: AnIntroductory Biology of Amphibians and REptiles. Second Edition." San Diego: Academic Press, 2001.