Hellbender Publications 

The Help the Hellbender team collaborates with many partners in the conservation efforts for the hellbender. The list of Purdue Hellbender research publications continues to be shared with scientists and conservation enthusiasts around the world.

(If you find a broken link please email Nick Burgmeier for publication, nburgmei@purdue.edu. Page is still under construction to share live links to the research publications.)

Albanese, B., J.B. Jensen, and S.D. Unger. 2011. Occurrence of the eastern hellbender (Cryptobranchus alleganiensis alleganiensis) in the Coosawattee River system (Mobile River Basin), Georgia. Southeastern Naturalist 9(4):181-184.

Avery, A. 2005. Rachel Carson syndrome: jumping to pesticide conclusions in the global frog crisis. Hudson Institute.

Augustine, L., K.A. Terrell, C. Petzinger, B. Nissen, and M. Maslanka. 2016. Nutritional analysis of diet items available to captive and free-ranging hellbenders (Cryptobranchus alleganiensis). Herpetological Review​ 47(1):63-69. ​  

Barton, B.S. 1812. A memoir concerning an animal of the class of reptilia, and amphibia, which is known, in the United States, by the name of alligator and hell-bender. Private printing. Griggs & Dickinson, Printers. 26 pp.

Beane, J. 2012. Hellbent. Wildlife in North Carolina 19-22.

Beffa, D.A. 1976. Responses of Cryptobranchus alleganiensis alleganiensis to different oxygen concentrations, temperatures and photoperiod. M. S. Thesis, Southwest Missouri State University, Springfield, Missouri. 37 pp.

Blais, D.P. 1996. Movement, home range, and other aspects of the biology of the eastern hellbender (Cryptobranchus alleganiensis alleganiensis): a radio telemetry study. M. S. Thesis, St. Lawrence University, Canton, New York. 63 pp.

Bodinof, C.M. 2010. Translocation and conservation of hellbenders (Cryptobranchus alleganiensis) in Missouri. M. S. Thesis, University of Missouri, Columbia, Missouri. 152 pp.

Bodinof, C.M. 2016. Effects of Land Use on Hellbenders (Cryptobranchus alleganiensis) at Multiple Levels and Efficacy of Artificial Shelters as a Monitoring Tool. Ph.D. Dissertation, Virginia Tech​, Blacksburg, Virginia. 201 pp.

Bodinof, C.M., J.T. Briggler, M.C. Duncan, J. Beringer, and J.J. Millspaugh. 2011. Historic occurrence of the amphibian chytrid fungus Batrachochytrium dendrobatidis in hellbender Cryptobranchus alleganiensis populations from Missouri. Diseases of Aquatic Organisms 96:1-7.

Bodinof, C.M., J.T. Briggler, R.E. Junge, J. Beringer, M.D. Wanner, C.D. Schuette, J. Ettling, R.A. Gitzen, and J.J. Millspaugh. 2012. Postrelease movements of captive-reared Ozark hellbenders (Cryptobranchus alleganiensis bishopi). Herpetologica 68(2):160-173.

Bodinof, C.M., J.T. Briggler, R.E. Junge, J. Beringer, M.D. Wanner, C.D. Schuette, J. Ettling, and J.J. Millspaugh. 2012. Habitat attributes associated with short-term settlement of Ozark hellbender (Cryptobranchus alleganiensis bishopi) salamanders following translocation to the wild. Freshwater Biology 57:178-192.

Bodinof, C.M., J.T. Briggler, R.E. Junge, T. Mong, J. Beringer, M.D. Wanner, C.D. Schuette, J. Ettling, and J.J. Millspaugh. 2012. Survival and body condition of captive-reared juvenile Ozark hellbenders (Cryptobranchus alleganiensis bishopi) following translocation to the wild. Copeia 2012(1):150-159.

Bodinof-Jachowski, C.M., J.J. Millspaugh, and  W.A. Hopkins. 2016. Current land use is a poor predictor of hellbender occurrence: why assumptions matter when predicting distributions of data-deficient species. Diversity and Distributions (2016):1-16. ​

Bodinof-Jachowski, C.M., and W.A. Hopkins. 2018. Loss of catchment-wide riparian forest cover is associated with reduced recruitment in a long-lived amphibian. Biological Conservation 202(2018):215-227.

Bodinof​-Jachowski, C.M., B.E. Ross, and W.A. Hopkins. 2020. Evaluating artificial shelter arrays as a minimally invasive monitoring tool for the hellbender Cryptobranchus alleganiensis. Endangered Species Research 41:167-181.

Bogo, J. 2009. Fighting the current. National Parks 83(2):22-23.

Bothner, R.C., and J.A. Gottlieb. 1991. A study of the New York state populations of the hellbender, Cryptobranchus alleganiensis alleganiensis(Daudin). Proceedings of the Rochester Academy of Science 17:41-54.

Boutilier, R.G., D.G. McDonald, and D.P. Toews. 1980. The effects of enforced activity on ventilation, circulation and blood acid-base balance in the aquatic gill-less urodele, Cryptobranchus alleganiensis; a comparison with the semi-terrestrial anuran, Bufo marinus. Journal of Experimental Biology 84:289–302.

Boutilier, R.G., and D.P. Toews. 1981a. Respiratory, circulatory and acid-base adjustments to hypercapnia in a strictly aquatic and predominantly skin-breathing urodele, Cryptobranchus alleganiensis. Respiration Physiology 46:177–192.

Boutilier, R.G., and D.P. Toews. 1981b. Respiratory properties of blood in a strictly aquatic and predominantly skin-breathing urodele,Cryptobranchus alleganiensis. Respiration Physiology 46:161–176.

Branch, H.E. 1935. A laboratory manual of Cryptobranchus alleganiensis Daudin. Vantage Press, Wichita, 79 pp.

Bredehoeft, K.E. 2010. A re-evaluation of the Pleistocene hellbender, Cryptobranchus guildayi, and an overview ofCryptobranchus remains from caves in the Potomac River region. M. S. Thesis, East Tennessee State University, Johnson City, Tennessee. 82 pp.

Briggler, J.T., and Y. Huang. 2006. Evaluation of health conditions, reproductive hormones, and mercury in adult hellbenders (Cryptobranchus alleganiensis). Missouri Department of Conservation, Jefferson City, Missouri.

Briggler, J.T. 2007. Hellbender recovery actions in Missouri. Missouri Department of Conservation, Jefferson City, Missouri. Science Note 2(15).

Briggler, J., J. Utrup, C. Davidson, J. Humphries, J. Groves, T. Johnson, J. Ettling, M. Wanner, K. Traylor-Holzer, D. Reed, V. Lindgren, and O. Byers (eds.). 2007. Hellbender population and habitat viability assessment. IUCN/SSC Conservation Breeding Specialist Group, Apple Valley, Minnesota. 46 pp.

Brown, R.K., A.J. Silla, R. Upton, G. Della-Togna, R. Marcec-Greaves, N.V. Shishova, V.K. Uteshev, B. Proano, O.D. Perez, N. Mansour, S.A. Kaurova, E.N. Gakhova, J. Cosson, B. Dyzuba, L.I. Kramarova, D. McGinnity, M. Gonzalez, J. Clulow, and S. Cluwlow. 2019. Sperm collection and storage for the sustainable management of amphibian biodiversity. Theriogenology 133:187-200

Browne, R.K., H. Li, Z. Wang, D. McGinnity, ​S. Okada, W. Zhenghuan, C. Bodinof, K. Irwin, A. McMillan, and J.T. Briggler. 2011. Survey techniques for giant salamanders and other aquatic Caudata. Amphibian and Reptile Conservation 5(4):1-16.​

Browne, R.K., H. Li, Z. Wang, P.M. Hime, A. McMillan, M. Wu, R. Diaz, Z. Hongxing, D. McGinnity, and J.T. Briggler. 2012. The giant salamanders (Cryptobranchidae): part A. palaeontology, phylogeny, genetics, and morphology. Amphibian and Reptile Conservation 5(4):17-29.

Browne, R.K., H. Li, Z. Wang, S. Okada, P.M. Hime, A. McMillan, M. Wu, R. Diaz, D. McGinnity, and J.T. Briggler. 2014. The giant salamanders (Cryptobranchidae): part B. ​biogeography, ecology and reproduction. Amphibian and Reptile Conservation 5(4):30-50.​

Browne, R., Z. Wang, S. Okada, D. McGinnity, Q. Luo, Y. Taguchi, D. Kilpatrick, R. Hardman, P. Janzen, Z. Zhang, and Y. Geng. 2020. The Sustainable Management of Giant Salamanders (Cryptobranchoidea). Sustainability America, Belize

Burgmeier, N.G. 2010. Population estimation, spatial ecology, and health status of eastern hellbenders in Indiana. M.S. Thesis, Purdue University, West Lafayette, Indiana. 76 pp.

Burgmeier, N.G., R.N. Williams, and S. Klueh-Mundy. 2016. One hell of a tale: a holistic model for hellbender conservation in Indiana. The Wildlife Professional​ 10(3): 32-35. 

Burgmeier, N.G., S.D. Unger, T.M. Sutton, and R.N. Williams. 2010. The bender board: a new design for the restraint and measurement of hellbenders. Herpetological Review 41(3):319-320.

Burgmeier, N.G., T.M. Sutton, and R.N. Williams. 2011. Spatial ecology of the eastern hellbender (Cryptobranchus alleganiensis alleganiensis) in Indiana. Herpetologica 67(2):135-145.

Burgmeier, N.G., S.D. Unger, J.L. Meyer, T.M. Sutton, and R.N. Williams. 2011. Health and habitat quality assessment for the eastern hellbender (Cryptobranchus alleganiensis alleganiensis) in Indiana, USA. Journal of Wildlife Diseases 47(4):836-848.

Burgmeier, N.G., S.D. Unger, T.M. Sutton, and R.N. Williams. 2011. Population status of eastern hellbender (Cryptobranchus alleganiensis alleganiensis) in Indiana. Journal of Herpetology 45(2):195-201.

Button, S.T., C.M. Bodinof-Jachowski, B.F. Case, J. Groffen, and W.A. Hopkins. 2020. The Influence of Multiscale Habitat Variables and Population Density on Artificial Shelter Use by Hellbenders (Cryptobranchus alleganiensis). Herpetologica 76(4):355-365.

Button, S.T., J.J. Hallagan, C.M. Bodinof-Jachowski, B.F. Case, J. Groffen, and W.A. Hopkins. 2020. Weathering the storm: Improving the availability and stability of artificial shelters for hellbender salamanders. River Research and Applications. 36:1944-1953. https://doi.org/10.1002/rra.3722

Caliando, J.J., Jr. 1975. The effects of the pancreatic hormones, insulin and glucagon, on blood glucose and liver glycogen in the hellbender,Cryptobranchus alleganiensis. M. S. Thesis, Sothwest Missouri State University, Springfield, Missouri. 25 pp.

Canada, B., and A.F. Scott. 2003. Cryptobranchus alleganiensis alleganiensis. Herpetological Review 34(3):255-256.

Cava, Z.A., A.M. McMillan, C. M. Pennuto, and R.J Warren II. 2018. Hellbender Prey Preference is Superseded by Native and Nonnative Prey Behavior. Journal of Herpetology 52(2):162-170.

Chudyk, S., A. McMillan, and C. Lange. 2014.  Using the eastern hellbender salamander in a high school genetics and ecological conservation activity. The American Biology Teacher 76:338-344.

Civiello, J.A., T.J. Bruce, S.J. Brisco, and J.T. Briggler. 2018. Propagation of eastern hellbenders (Cryptobranchus alleganiensis alleganiensis in a recirculating aquaculture system at Shepherd of the Hills State Fish Hatchery. North American Journal of Aquaculture 1-10.

Coatney, C.E., Jr. 1982. Home range and nocturnal activity of the Ozark hellbender. M. S. Thesis, Southwest Missouri State University, Springfield, Missouri. 55 pp.

Collins, J.T. 1981. Cryptobranchus alleganiensis (hellbender) longevity. Herpetological Review 12(2):69.

Connock, J.R., B.F. Case, S.T. Button, J. Groffen, T.M. Galligan, and W.A. Hopkins. 2019. Factors Influencing In-situ Detection of PIT-tagged Hellbenders (Cryptobranchus alleganiensis) Occupying Artificial Shelters Using a Submersible Antenna. Herpetological Conservation and Biology 14(2):429-437.

Conservation Research Reports. 2008. Non-native fishes threatening hellbenders in Missouri. Iguana 15(3):177.

Crane, A.L., and A. Mathis. 2011. Predator-recognition training: a conservation strategy to increase postrelease survival of hellbenders in head-starting programs. Zoo Biology 30:611-622.

Crone, E. 2018. Comparison of habitat and water quality in the Blue River's south and salem forks in southern Indiana to support watershed management. Undergraduate Honors Thesis. Bellarmine University, KY.

Crowhurst, R.S., K.M. Faries, J. Collantes, J.T. Briggler, J.B. Koppelman, and L.S. Eggert. 2011. Genetic relationships of hellbenders in the Ozark highlands of Missouri and conservation implications for the Ozark subspecies (Cryptobranchus alleganiensisbishopi). Conservation Genetics 12:637-646.

Cundall, D, J. Lorenz-Elwood, and J.D. Groves. 1987. Asymmetric suction feeding in primitive salamanders. Experientia 43:1229-1231.

Da Silva Neto, J.G. 2018. Assessing site occupancy and microhabitat use of the hellbender salamander (Cryptobranchus alleganiensis) in Tennessee. Ms Theses. Tennessee State University, TN.

Da Silva Neto, J.G., W.B. Sutton, and M.J. Freake. 2019. Life-stage Differences in Microhabitat Use by Hellbenders (Crypboranchus alleganiensis). Herpetologica 75(1):21-29.

Da Silva Neto, J.G., W.B. Sutton, S.F. Spear, M.J. Freake, M. Kery, and B.R. Schmidt. 2020. Integrating species distribution and occupancy modeling to study hellbender (Cryptobranchus alleganiensis) occurrence based on eDNA surveys. Biological Conservation 251(108787). https://doi.org/10.1016/j.biocon.2020.108787.

Davis, A.K., and W.A. Hopkins. 2012. Widespread trypanosome infections in a population of eastern hellbenders (Cryptobranchus alleganiensis alleganiensis) in Virginia, USA. Parasitology Research. 4 pp.

Davis, A.K., and T.M. Floyd. 2013. Evaluating levels of genotoxic stress in eastern hellbenders (Cryptobranchus alleganiensis alleganiensis) using the erythrocyte micronucleus assay. Comparitive Clinical Pathology DOI 10.1007/s00580-013-1761-1.

Deban, S.M., and J.C. O’Reilly. 2005. The ontogeny of feeding kinematics in a giant salamander Cryptobranchus alleganiensis: does current function or phylogenetic relatedness predict the scaling patterns of movement? Zoology 108:155-167.

Diaz, L., O. Hernandez-Gomez, S.D. Unger, L.A. Williams, and C.M. Bodinof-Jachowski. 2020. Prevalence of Batrachochytrium dendrobatidis in immature eastern hellbenders Cryptobranchus alleganiensis from North Carolina, USA. Diseases of Aquatic Organisms 140:73-78.

Dierenfeld, E.S., K.J. McGraw, K. Fritsche, J.T. Briggler, and J. Ettling. 2009. Nutrient composition of whole crayfish (Orconectes and Procambarusspecies) consumed by hellbender (Cryptobranchus alleganiensis). Herpetological Review 40(3):324-330.

Dundee, H.A., and D.S. Dundee. 1965. Observations on the systematics and ecology of Cryptobranchus from the Ozark Plateaus of Missouri and Arkansas. Copeia 1965(3):369-370.

Ettling, J.A., M.D. Wanner, C.D. Schuette, S.L. Armstrong, A.S. Pedigo, and J.T. Briggler. 2013. Captive reproduction and husbandry of adult Ozark Hellbenders, Cryptobranchus alleganiensis bishopi. Herpetological Review 44:605-610.

Feist, S.M., J.T. Briggler, J.B. Koppelman, and L.S. Eggert. 2014. Within-river gene flow in the hellbender (Cryptobranchus alleganiensis) and implications for restorative release. Conserv Genet DOI 10.1007/s10592-014-0591-2.

Firschein, I.L. 1951. The range of Cryptobranchus bishopi and remarks on the distribution of the genus Cryptobranchus. American Midland Naturalist 45(2):455-459.

Fitch, F. W., Jr. 1947. A record Cryptobranchus alleganiensis. Copeia 1947(3):210.

Floyd, T. M., T.S. M. Stratmann, G.J. Brown, III, and C.S. Pfaff. 2013. Cryptobranchus alleganiensis alleganiensis. Terrestrial Movement. Herpetological Review 44:651.

Floyd, T.M., and S.D. Unger. 2016. Cryptobranchus alleganiensis alleganiensis (Eastern Hellbender). Reproductive Behavior and Habitat. Herpetological Review 47(2).

Fobes, T. M. 1995. Habitat analysis of the Ozark hellbender, Cryptobranchus alleganiensis bishopi, in Missouri. M. S. Thesis, Southwest Missouri State University, Springfield, Missouri. 38 pp.

Foster, R.L., A.M. McMillan, A.R. Breisch, K.J. Roblee, and D. Schranz. 2008. Analysis and Comparison of Three Capture Methods for the Eastern Hellbender (Cryptobranchus alleganiensis alleganiensis). Herpetological Review​​ 39(2):181-186.

Foster, R.L., A.M. McMillan, and K.J. Roblee. 2009. Population status of hellbender salamanders (Cryptobranchus alleganiensis) in the Allegheny River drainage of New York State. Journal of Herpetology 43(4):579-588.

Foster, R. 2018. Lessons from the past: A historical approach to conservation of the eastern hellbender (Cryptobranchus alleganiensis). Ph.D Dissertation, State University of New York at Buffalo.

Freake, M.J., and C.S. DePerno. 2017. Importance of demographic surveys and public lands for the conservation of eastern hellbenders ​Cryptobranchus alleganiensis alleganiensis in the southeast USA. PLoS One 12(6): e0179153​.

Freake, M., E. O'Neill, S. Unger, S. Spear, and E. Routman. 2018. Conservation genetics of eastern hellbenders Cryptobranchus alleganiensis alleganiensis in the Tennessee Valley. Conservation Genetics 19(3): 571-585.

Furniss, L.L. 2003. Conservation assessment for Ozark hellbender (Cryptobranchus alleganiensis bishopi Grobman). USDA Forest Service, Rolla, Missouri.

Gall, B.G. 2008. Predator-prey interactions between hellbenders (Cryptobranchus alleganiensis alleganiensis and C.A. bishopi) and native and nonnative fishes. M.S. Thesis, Missouri State University, Springfield, Missouri. 85 pp.

Gall, B.G., and A. Mathis. 2010. Innate predator recognition and the problem of introduced trout. Ethology 116(2010):47-58.

Gall, B.G., and A. Mathis. 2010. Response of native and introduced fishes to presumed antipredator secretions of Ozark hellbenders (Cryptobranchus alleganiensis bishopi). Behaviour 147:1769-1789.

Gall, B.G., and A. Mathis. 2011. Ontogenetic shift in response to amphibian alarm cues by banded sculpins (Cottus carolinae). Copeia 2011(1):5-8.

Gao, K., and N.H. Shubin. 2003. Earliest known crown-group salamanders. Nature 422:424-428.

Gates, J.E. 1983. The distribution and status of the hellbender (Cryptobranchus alleganiensis) in Maryland: I. The distribution and status ofCryptobranchus alleganiensis in Maryland. II. Movement patterns of translocated Cryptobranchus alleganiensis in a Maryland stream. University of Maryland, Frostburg, Maryland.

Gates, J.E., C.H. Hocutt, and J.R. Stauffer, Jr. 1984. The status of the hellbender (Cryptobranchus alleganiensis) in Maryland. University of Marlyland, Frostburg, Maryland.

Gates, J.E., C.H. Hocutt, J.R. Stauffer, Jr., and G. J. Taylor. 1985. The distribution and status of Cryptobranchus alleganiensis in Maryland. Herp Review 16(1):17-18.

Gates, J.E., R.H. Stouffer, Jr., J.R. Stauffer, Jr., and C.H. Hocutt. 1985. Dispersal patterns of translocated Cryptobranchus alleganiensis in a Maryland stream. Journal of Herpetology 19(3):436-438.

Gonynor, J.L., M.J. Yabsley, and J.B. Jensen. 2011. A preliminary survey of Batrachochytrium dendrobatidis exposure in hellbenders from a stream in Georgia, USA. Herpetological Review 42(1):58-59.

Gottlieb, J.A. 1991. A population study of the hellbender salamander, Cryptobranchus alleganiensis, in the Allegheny River drainage of New York State. M. S. Thesis. St. Bonaventure University, Olean, New York. 54 pp.

Graham, S.P., E.C. Soehren, G.R. Cline, C.M. Schmidt, W.B. Sutton, J.R. Rayburn, S.H. Stiles, and J.A. Stiles. 2011. Conservation status of hellbenders (Cryptobranchus alleganiensis) in Alabama, USA. Herpetological Conservation and Biology 6(2):242-249.

Green, N.B. 1933. Cryptobranchus alleganiensis in West Virginia. Proceedings of the West Virginia Academy of Science, Elkins, West Virginia. 7:28-30.

Greven, H., and G. Clemen. 2009. Early tooth transformation in the paedomorphic hellbender Cryptobranchus alleganiensis (Daudin, 1803) (Amphibia: Urodela). Vertebrate Zoology 59(1):71-79.

Groves, J.D. 2012. eDNA & the search for hellbenders. Alive 12-13.

Groves, J.D., and L.A. Williams. 2011. In search of the hellbender in North Carolina. Connect 12-15.

Groves, J.D., and L.A. Williams. 2014. Cryptobranchus alleganiensis - cannibalism. Hepetological Review 45:108-109.

Groves, J.D., L.A. Willams, and S.P. Graham. 2015. Larval/Adult Association. Herpetological Review 46(1):70-71.

Guimond, R. W., and V. H. Hutchinson. 1973. Aquatic respiration: an unusual strategy in the hellbender Cryptobranchus alleganiensis alleganiensis(Daudin). Science 182(4118):1263-1265.

Hardman, R.H., K.J. Irwin, W.B. Sutton, and D.L. Miller. 2020. Evaluation of Severity and Factors Contributing to Foot Lesions in Endangered Ozark Hellbenders, Cryptobranchus alleganiensis bishopi. Frontiers in Veterinary Science 7(34)

Hardman, R.H., W.B. Sutton, K.J. Irwin, D. McGinnity, S.D. Reinsch, M. Freake, P. Colclough, B.T. Miller, J.G. Da Silva Neto, M. Souza, B. Fitzpatrick, and D.L. Miller. Geographic and Individual Determinants of Important Amphibian Pathogens in Hellbenders (Cryptobranchus alleganiensis) in Tennessee and Arkansas, USA. Journal of Wildlife Diseases 56(4):803-814.

Harlan, R. A., and R. F. Wilkinson. 1981. The effects of progressive hypoxia and rocking activity on blood oxygen tension for hellbenders,Cryptobranchus alleganiensis. Journal of Herpetology 15(4):383-387.

Harshbarger, J. C., and S. E. Trauth. 2002. Squamous cell carcinoma upgrade of the epidermal papilloma reported in an Ozark hellbender (Cryptobranchus alleganiensis bishopi). Pp. 43–48. In: R.G. McKinnell and D.L. Carlson (eds.). Proceeding of the Sixth International Symposium on the Pathology of Reptiles and Amphibians, Saint Paul, Minnesota.

Hartley, J. 2018. Distribution and habitat preferences of eastern hellbenders within the Greenbrier River. Proceedings of the West Virginia Academy of Science 90(1).

Hartzell, S. M., A. L. Pitt., and S. Davis. 2017. Observations of yawning behaviour in the eastern hellbender (Cryptobranchus alleganiensis alleganiensis). The Herpetological Bulletin 142: 46-47.​ 

Hartzell, S. M., A. L. Pitt, S. Davis, C. R. Pavlick, and J. J. Tavano. 2017. ​Cryptobranchus alleganiensis alleganiensis ​(Eastern Hellbender). Temporal and Spatial Niche Partitioning. Herpetological Review 48(1)​.

Hartzell, S.M., A.L. Pitt, and S.M. Davis. 2022. Invasive Rusty Crayfish (Faxonius rusticus) Can Serve as Prey of Eastern Hellbender (Cryptobranchus alleganiensis alleganiensis). Herpetological Conservation and Biology 17(1):22-30.

Hasegawa, H., and Y. Ikeda. 2003. Helminths from the hellbender, Cryptobranchus alleganiensis (Urodela: Cryptobranchidae), in Missouri, U.S.A. Comparative Parasitology 70(1):60–65.

Hecht, K. A., and M. A. Nickerson. 2013. Cryptobranchus alleganiensis (Hellbender). Larval diet. Herpetological Review 44(3): 490.

Hecht, K. A., M. A. Nickerson, and R. Vollbrecht. 2014. Cryptobranchus alleganiensis (Hellbender). Predation. Herpetological Review 45(3): 471.

Hecht, K. A., M. J. Freake, M. A. Nickerson, and P. Colclough. 2019. Hellbender Salamanders (Cryptobranchus alleganiensis) Exhibit an Ontogenetic Shift in Microhabitat Use in a Blue Ridge Physiographic Region Stream. Copeia 107 (1): 152-159.

Hecht-Kardasz, K. A., M. A. Nickerson, M. Freake, and P. Colclough. 2012. Population structure of the hellbender (Cryptobranchus alleganiensis) in a Great Smoky Mountains stream. Bulletin of the Florida Museum of Natural History 51(4):227-241.

Hecht-Kardasz, K. A., M. A. Nickerson, and P. Colclough. 2017. Hellbenders (​Cryptobranchus alleganiensis) May Exhibit an Ontogenetic Dietary Shift. Southeastern Naturalist 16(2):157-162.​ 

Hernandez-Gomez, O., J.T. Hoverman, and R.N. Williams. 2017. Cutaneous Microbrial Community Variation across Populations of Eastern Hellbenders (Cryptobranchus alleganiensis alleganiensis). 2017. Frontiers in Microbiology 8: 1-16.​

Hernandez-Gomez, O., S.J.A. Kimble, J.T. Briggler, and R.N. Williams. 2017. Characterization of the Cutaneous Bacterial Communities of Two Giant Salamander Subspecies. Microbial Ecology 73: 445-454. ​

Hernández-Gómez, O., J.T. Briggler, and R.N. Williams. 2018. Captivity-induced changes in the skin microbial communities of hellbenders (Cryptobranchus alleganiensis). Microbial Ecology 2018: 1-12

Hernández-Gómez, O., J.T. Briggler, and R.N. Williams. 2018. Influence of immunogenetics, sex and body condition on the cutaneous microbial communities of two giant salamanders. Molecular Ecology 27(2): 1915-1929.

Hernández-Gómez, O., J.T. Briggler, and R.N. Williams. 2019. Captivity-Induced Changes in the Skin Microbial Communities of Hellbenders (Cryptobranchus alleganiensis). Microbial Ecology 77: 782-793.  

Hiler, W. R., B. A. Wheeler, and S. E. Trauth. 2005. Abnormalities in the Ozark hellbender (Cryptobranchus alleganiensis bishopi) in Arkansas: a comparison between two rivers with a historical perspective. Journal of Arkansas Academy of Science 59:88–94.

Hill, R. L. 2011. Cryptobranchus alleganiensis alleganiensis (Eastern Hellbender) and Desmognathus quadramaculatus (Black-bellied Salamander). Predator/Prey. Herpetological Review 42(4)​: 580.

Hillis, R. E., and E. D. Bellis. 1971. Some aspects of the ecology of the hellbender, Cryptobranchus alleganiensis alleganiensis, in a Pennsylvania stream. Journal of Herpetology 5(3/4):121-126.

Hime, P.M., J.T. Briggler, J.S. Reece, and D.W. Weisrock. 2019. Genomic Data Reveal Conserved Female Heterogamety in Giant Salamanders with Gigantic Nuclear Genomes. G3 9(10):3467-3476.

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Tonione, M., J. R. Johnson, and E. J. Routman. 2010. Microsatellite analysis supports mitochondrial phylogeography of the hellbender (Cryptobranchus alleganiensis). Genetica 139:209-219.

Topping, M. S. and C. A. Ingersol. 1981. Fecundity in the hellbender, Cryptobranchus alleganiensis. Copeia 1981(4):873-876.

Trauth, S. E., J. D. Wilhide, and P. Daniel. 1992. Status of the Ozark hellbender, Cryptobranchus bishopi (Urodela: Cryptobranchidae), in the Spring River, Fulton Country, Arkansas. Proceedings Arkansas Academy of Science 46:83-86.

Trauth, S. E., J. D. Wilhide, and P. Daniel. 1993. The Ozark hellbender, Cryptobranchus bishopi, in Arkansas: distributional survey for 1992. Bulletin of the Chicago Herpetological Society 28(4):81-85.

Trauth, S. E., J. C. Harshbarger, and P. Daniel. 2002. Epidermal papilloma in an Ozark hellbender (Cryptobranchus alleganiensis bishopi) from the Spring River of northern Arkansas. Journal of the Arkansas Academy of Science 56:190-197.

Ultsch, G. R., and J. T. Duke. 1990. Gas exchange and habitat selection in the aquatic salamanders Necturus maculosus and Cryptobranchus alleganiensis. Oecologia 83:250-258.

Unger, S. D., J. A. Fike, T. Sutton, O. E. Rhodes, Jr., and R. N. Williams. 2010. Isolation and development of 12 polymorphic tetranucleotide microsatellite markers for the eastern hellbender (Cryptobranchus alleganiensis alleganiensis). Conservation Genetics Resources 2:89-91.  

Unger, S. D., N. G. Burgmeier, and R. N. Williams. 2012. Genetic markers reveal high PIT tag retention rates in giant salamanders (Cryptobranchus alleganiensis). Amphibia-Reptilia 33:313-317.

Unger. S. D. and A. Mathis. 2013. Larval growth and the potential for head-starting of eastern and Ozark hellbenders (Cryptobranchus alleganiensis alleganiesis and C. a. bishopi).

Herpetological Review 44: 89-91.

Unger, S. D. A. Mathis, and R. Wilkinson. 2013. A comparison of sperm health in declining and stable populations of hellbenders (Cryptobranchus alleganiensis alleganiensis and C. a. bishopi). American Midland Naturalist 170: 382-392

Unger, S. D., O. E. Rhodes, Jr., T. M. Sutton, and R. N. Williams. 2013. Population genetics of the eastern hellbender (Cryptobranchus alleganiensis alleganiensis) across multiple spatial scales. PLOS ONE 8(10):1-14.

Unger, S. D., T. M. Sutton, and R. N. Williams. 2013. Projected population persistence of eastern hellbenders (Cryptobranchus alleganiensis alleganiensis). Journal for Nature Conservation 21:423-432.

Unger, S.D. and R.N. Williams. 2015. Genetic Analysis Reveals Multiple Parentage in Captive Reared Eastern Hellbender Salamanders (Cryptobranchus alleganiensis). Zoo Biology 34: 525-537​.

Unger, S.D., E.J. Chapman, K.J. Regester, and R.N. Williams. 2016. Genetic Signatures Follow Dendritic Patterns in the Eastern Hellbender (Cryptobranchus allegeniensis allegeniensis). Herpetological Conservation and Biology 11(1): 40-51.

Unger, S. D., L.A. Williams, and J.D. Groves. 2016. Cryptobranchus alleganiensis alleganiensis (Eastern Hellbender). Unusual Mortality. Herpetological Review 47(4): 639​. ​

Unger, S. D., L.A. Williams, J.D. Groves, C.R. Lawson, and W.J. Humphries. 2017. Anthropogenic Associated Mortality in the Eastern hellbender (Cryptobranchus alleganiensis alleganiensis). Southerneastern Naturalist  16(2): 9-13. 

Unger, S. D. 2018. Scavenging Behavior of the Aquatic Eastern Hellbender Salamander (Cryptobranchus alleganiensis) in North Carolina. Journal of North Carolina Academy of Science. 134: 1-2.

Unger, S. D., and R. N. Williams. 2018. Genetic confirmation of filial cannibalism in North America's giant salamander, the Eastern hellbender Cryptobranchus alleganiensis alleganiensis. Ethology Ecology & Evolution​ 30(2): 187-193​.

Unger, S. D., L. A. Williams, L. Diaz, and C. M. Bodinof-Jachowski. 2019. DNA barcoding to assess diet of larval eastern hellbenders in North Carolina. Food Webs 22 (2020).

Unger, S. D., L. A. Williams, G. G. Williams, and C. M. Bodinof-Jachowski. 2019. Cryptobranchus alleganiensis (Hellbender). Egg Predation. Herpetological Review 50(1): 2019.  

Unger, S. D., C. Bodinof-Jachowski, L. Diaz, and L. Williams. 2020. Adult Female Eastern hellbender Cryptobranchus alleganiensis (Cryptobranchidae) conspecific cannibalism confirmed via DNA barcoding. Herpetology Notes 13: 169-170

Unger, S. D., C. Bodinof-Jachowski, L. Diaz, and L. Williams. 2020. Cryptobranchus alleganiensis (Hellbender). Milt Predation. Herpetological Review 51(3)

Unger, S. D., C. Bodinof-Jachowski, L. Diaz, and L. Williams. 2020. Cryptobranchus alleganiensis (Hellbender). Predation Attempt and Anti-Predator Behavior. Herpetological Review 51(1).

Unger, S. D., C. Bodinof-Jachowski, L. Diaz, and L. Williams. 2020. Observations on the habitat preference of juvenile eastern hellbender salamanders (Cryptobranchus alleganiensis). acta ethologica. https://doi.org/10.1007/s10211-020-00344-9.

Unger, S. D., C. Bodinof Jachowski, L. Diaz, and L. Williams. 2020. Shelter Guarding Behavior of the Eastern Hellbender (Cryptobranchus alleganiensis alleganiensis). in North Carolina Streams. Southeastern Naturalist 19(4): 742-758.

Unger, S. D., and C. R. Hickman. 2020. A content analysis from 153 years of print and online media shows positive perceptions of the hellbender salamander follow the conservation biology. Biological Conservation 246(2020).

Unger, S. D., Z. C. Hull, L. Diaz, J. D. Groves, L. A. Williams, C. M. Bodinof-Jachowski. 2020. Underwater video cameras allow for detection of North American giant salamanders (Cryptobranchus alleganiensis alleganiensis) in both captive and wild streams. Aquaculture and Fisheries. https://doi.org/10.1016/j.aaf.2020.03.003.

Unger, S. D., L. A. Williams, C. R. Lawson, and J. D. Groves. 2020. Using Trail Cameras to Assess Recreation in Hellbender Streams of North Carolina National Forests. Journal of the Southeastern Association of Fish and Wildlife Agencies 2020:255-262.

Unger, S. D., L. A. Williams, J. D. Groves, and C. R. Lawson. 2021. Factors Influencing Occupancy and Detection Probability of Larval Cryptobranchus a. alleganiensis in North Carolina, USA. Herpetological Conservation and Biology 16(1):157-172.

Unger, S.D., L.A. Williams, and J.D. Groves. 2021. Cryptobranchus alleganiensis alleganiensis (Eastern Hellbender). Larval Displacement Due to Flooding. Herpetological Review 52(3).

Unger, S., S. Benfield, L. Williams, C. Lawson, and J. Groves. 2021. Macroinvertebrate Communities and Body Condition of Larval Eastern Hellbender Salamanders (Cryptobranchus alleganiensis) in North Carolina. Zoological Studies 60:77

Unger, S.D., L.A. Williams, J. Rash, L. Etchison, D. Owensby, and J.D. Groves. 2021. Cryptobranchus alleganiensis alleganiensis (Eastern Hellbender). Larval Predation. Herpetological Review 52(4).

Wheeler, B. A. 1999. Status of the Ozark hellbender (Cryptobranchus alleganiensis bishopi): a long-term assessment. M. S. Thesis, Southwest Missouri State University, Springfield, Missouri. 32 pp.

Wheeler, B. A., M. L. McCallum, and S. E. Trauth. 2002. Abnormalities in the Ozark hellbender, Cryptobranchus alleganiensis bishopi. Journal of the Arkansas Academy of Science 56:250-252.

Wheeler, B. A., and S. Trauth. 2002. Distributional survey of the Ozark hellbender, Cryptobranchus alleganiensis bishopi: final report for field season 2001. U. S. Fish and Wildlife Service. 28 pp.

Wheeler, B. A., E. Prosen, A. Mathis, and R. F. Wilkinson. 2003. Population declines of a long-lived salamander: a 20 + -year study of hellbenders,Cryptobranchus alleganiensis. Biological Conservation 109:151-156.

Widner, B., and R. Williams. 2018. The Scientific Process of Conservation Biology: Analyze, Design, Debate. Purdue Extension - eduationSTORE.

Wiggs, J. N. 1976. Food habits, starvation and growth in the hellbender, Cryptobranchus alleganiensis. M. S. Thesis, Southwest Missouri State University, Springfield, Missouri. 32 pp.

Wikramanayake, E. D., and G. L. Dryden. 1985. Body length of male Cryptobranchus alleganiensisat sexual maturity. Bulletin Maryland Herpetological Society pp 76-79.

Wineland, S.M., S.M. Welch, T.K. Pauley, J.J. Apodaca, M. Olszack, J.J. Mosher, J.N. Holmes, and J.L. Waldron. 2018. Using environmental DNA and occupancy modelling to identify drivers of eastern hellbender (Cryptobranchus alleganiensis alleganiensis) extirpation. Freshwater Biology 2018: 1-14.

Williams, L. A., and J. D. Groves. 2014. Prevalance of the amphibian pathogen Batrachochytrium Dendrobatidis  in Eastern hellbenders (Cryptobranchus A. Alleganiensis) in Western North Carolina, USA. Herpetological Conservation and Biology 9(3):454-467.

Williams, L. A., J. M. Rash, J. D. Groves, L. L. Stroup, and D. Blatney. 2019. Engaging North Carolina's Trout Anglers and Other Stakeholders to Help Conserve Eastern Hellbenders. Journal of the Southeastern Association of Fish and Wildlife Agencies 6: 166-174. 

Williams, R. D., J. E. Gates, and C. H. Hocutt. 1981. An evaluation of known potential sampling techniques for hellbender, Cryptobranchus alleganiensis. Journal of Herpetology 15(1):23-27.

Williams, R. D., J. E. Gates, C. H. Hocutt, and G. J. Taylor. 1981. A nongame species in need of management. Wildlife Society Bulletin 9(2):94-100.

Yager, V., N. Burgmeier, and R. Williams. In press. Adaptations for Aquatic Amphibians - Lesson Plan, Purdue Extension - educationSTORE.

Zhang, L., Hu Zhao, S. Willard, Q. Wang, W. Jiang, H. Zhang, and A. Kouba. 2019. Spatial distribution and seasonal movement patterns of reintroduced Chinese giant salamanders. BMC Zoology 4:7. https://doi.org/10.1186/s40850-019-0046-9.

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