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Order: Caudata
Family: Proteidae
Proteidae is an ancient family of salamanders comprising just six species, commonly referred to as the olm, mudpuppies and waterdogs. The lineage diverged from their closest relatives 190 million years ago in the early Jurassic period in the era of the dinosaurs. This predates the first fossil bird by 40 million years.
A highly unusual feature of the proteids is that metamorphosis is absent in this family. The young never transition into an adult form, instead retaining their larval characteristics throughout life, which include feathery gills, a tail fin and no eyelids, as appropriate to their permanently aquatic lifestyle. This is termed “neoteny” or “paedomorphosis” because these salamanders achieve reproductive maturity whilst still in the larval form. The family Proteidae is part of a suborder of salamanders that contains all of the internally-fertilising species (known as the “Salamandroidea”). Most salamanders reproduce via external ferlilsation, but in the proteids females take the spermatophone (or sperm packet) deposited by the male into their bodies, allowing fertisation of her eggs to occur within the body cavity.
The proteids comprise two geographically distant genera: Proteus (the European olm) and Necturus (the mudpuppies and waterdogs of eastern North America). In contrast to their closest relative, pale, blind cave-dwelling olm (Proteus anguinus – EDGE rank 18), the mudpuppies and waterdogs have normal skin pigmentation and eyes, and 4 digits on each foot, as opposed to the three fingers and two toes sported by the olm. These two lineages diverged 140 million years ago in the late Jurassic period. In terms of mammal groups, this makes them about as dissimilar from each other as wombats are from humans. These salamanders inhabit streams, rivers and lakes, where they remain concealed under rocks and debris during the day, and forage for invertebrate prey at night.
A highly unusual feature of the proteids is that metamorphosis is absent in this family. The young never transition into an adult form, instead retaining their larval characteristics throughout life, which include feathery gills, a tail fin and no eyelids, as appropriate to their permanently aquatic lifestyle. This is termed “neoteny” or “paedomorphosis” because these salamanders achieve reproductive maturity whilst still in the larval form. The family Proteidae is part of a suborder of salamanders that contains all of the internally-fertilising species (known as the “Salamandroidea”). Most salamanders reproduce via external ferlilsation, but in the proteids females take the spermatophone (or sperm packet) deposited by the male into their bodies, allowing fertisation of her eggs to occur within the body cavity.
The proteids comprise two geographically distant genera: Proteus (the European olm) and Necturus (the mudpuppies and waterdogs of eastern North America). In contrast to their closest relative, pale, blind cave-dwelling olm (Proteus anguinus – EDGE rank 18), the mudpuppies and waterdogs have normal skin pigmentation and eyes, and 4 digits on each foot, as opposed to the three fingers and two toes sported by the olm. These two lineages diverged 140 million years ago in the late Jurassic period. In terms of mammal groups, this makes them about as dissimilar from each other as wombats are from humans. These salamanders inhabit streams, rivers and lakes, where they remain concealed under rocks and debris during the day, and forage for invertebrate prey at night.
The Alabama waterdog, like all proteids, is completely aquatic and gilled throughout life. This is a medium-sized species among the mudpuppies and waterdogs, reaching a total length of 150-220 mm, the tail accounting for around 50 to 70 mm of this measurement. The maximum size reported of an Alabama waterdog was a 248 mm long female. All mudpuppies and waterdogs have bushy external gills, two gill slits, a flattened tail, and four toes on front and hind feet. The body and head are flattened, and sexually mature males can be distinguished from the females by their swollen cloaca (urino-genital reproductive opening) and a pair of enlarged “cloacal papillae” (fleshy projections around the cloaca) that project towards the tail. The limbs are well-developed, although not large. The flattened body of this species may be an adaptation to its habit of hiding under rocks and other cover. The dorsal (or upper) surfaces of this salamander are reddish brown to nearly black. Some populations have spots along the back and tail. The ventral (or lower) surfaces lacks spots in all age classes, and the tips of the toes are light coloured. Hatchlings are mottled dorsally with a few light spots. In some populations, juveniles have light stripes on the head and back, and also have a dark eye stripe running from the nostril, through the eye to the gills. This stripe is retained in adults.
Relatively little is known about the natural history of this species. Alabama waterdogs are most abundant during the winter and are rarely found during summer months. It is thought they potentially undergo aestivation during the summer, which is a prolonged period of inactivity brought on by high temperatures or drought, although these salamanders remain active throughout the winter. The breeding season may start in December, since adults with swollen cloacal lips have been found in this month, which indicates a sexually active condition. However, hatchling larvae have also been collected in December, suggesting late spring or summer nesting. Individuals caught in the autumn may be quite lean in comparison to their winter/spring condition when they are in readiness for reproduction.
Fertilisation is internal in the Alabama waterdog has internal fertilisation. Small egg masses (comprising less than 100 eggs) are laid in water in the spring and early summer. They are often attached to the undersides of rocks and logs, or are deposited within leaf beds or rock crevices. The female guards the eggs until they hatch four to six weeks later, with larvae hatching in the early summer. Alabama waterdogs achieve adult size after about 2.5-3 years, but may require 4-6 years to reach sexual maturity.
This species is active on the bottom of streams at night, most frequently on cold stormy nights when it rains. Their diet includes earthworms, crayfish, aquatic invertebrates, snails and small fish, and they are probably opportunistic feeds taking any prey they can fit into their mouths. The Alabama waterdog is a secretive species, spending the day hidden under rocks or organic debris, especially leaf beds. It inhabits medium to large streams where there is plenty of shelter, and can live in both clear and muddy water.
Fertilisation is internal in the Alabama waterdog has internal fertilisation. Small egg masses (comprising less than 100 eggs) are laid in water in the spring and early summer. They are often attached to the undersides of rocks and logs, or are deposited within leaf beds or rock crevices. The female guards the eggs until they hatch four to six weeks later, with larvae hatching in the early summer. Alabama waterdogs achieve adult size after about 2.5-3 years, but may require 4-6 years to reach sexual maturity.
This species is active on the bottom of streams at night, most frequently on cold stormy nights when it rains. Their diet includes earthworms, crayfish, aquatic invertebrates, snails and small fish, and they are probably opportunistic feeds taking any prey they can fit into their mouths. The Alabama waterdog is a secretive species, spending the day hidden under rocks or organic debris, especially leaf beds. It inhabits medium to large streams where there is plenty of shelter, and can live in both clear and muddy water.
This species is restricted to streams in the Black Warrior River Basin of Alabama. Alabama waterdogs inhabit medium to large streams that have logs, submerged ledges, rocks, and other hiding places on the bottom. They are associated with clay substrates lacking silt, wide and/or narrow stream morphology, increased abundance of snails and larval northern dusky salamanders (Desmognathus fuscus), and decreased Asiatic mussel (Corbicula sp.) occurrence. The presence of leaf beds is important, especially during the breeding season when sexually active adults are found either in rock crevices or near woody of leafy debris, and require cover for their nesting sites.
This species can be found in medium–large streams of the upper (Appalachian) portions of the Black Warrior River drainage system above the Fall Line in Alabama, United States. Populations are known from four counties: : Sipsey Fork and Brushy Creek in Winston County, Locust Fork and Blackburn Fork in Blount County, Mulberry Fork, Blackwater Creek, and Lost Creek in Walker County, and Yellow Creek and North River in Tuscaloosa County. The range of this species includes parts of the North River, Locust Fork, Mulberry Fork, and Sipsey Fork drainages and their tributaries, in Blount, Tuscaloosa, Walker, and Winston counties, although Alabama waterdogs have possibly been eliminated from Blackburn Fork, Blount County. This species can be expected to potentially inhabit the same streams as the threatened flattened musk turtle (Sternotherus depressus), which is also restricted to permanent streams above the Fall Line in the Black Warrior River basin. The Alabama waterdog is known from 9 stream segments within its range, and has highly fragmented populations.
No population estimate currently exists for this species, although numerous studies and surveys have been carried out in an attempt to determine population trends and dynamics. The Alabama waterdog is known to be a rare species with sporadic occurrences within the presumed geographic range. A 1990-1992 survey by Mark A. Bailey found only a few individuals in four localities. Collections included six adults and one larva in the Sipsey Fork, one adult in Lost Creek, one larva in North River, and one sub adult in Yellow Creek. During another survey in 1996-1997 by C. Guyer, 18 individuals were collected from Sipsey Fork and 11 individuals from Brushy Creek. Even though the Alabama waterdog was extensively surveyed from 1990 to 1997, numbers collected are too low to determine population trends. One hundred and twenty sites have been sampled for this species in the Black Warrior River drainage since 1990. The species has been reported recently from only ten sites (8 % success rate) in four counties, despite surveys in 1990, 1991, 1992, 1994, 1996, 1997, and 1998. Sites surveyed included all stream localities within the range of the species that approached or intersected roads and had appropriate habitat. Distribution, even within the best habitat, appears to be patchy, and abundance may fluctuate from year to year depending on the development of submerged leaf beds
The black warrior waterdog is classified as in decline in the IUCN Red List of Threatened Species.
The Alabama waterdog is listed as Endangered in the IUCN Red List of Threatened Species because its area of occupancy is probably less than 500 km sq., its distribution is severely fragmented, and there is continuing decline in the extent and quality of its forest habitat in Alabama.
The primary reasons for the extirpation of this species over much of its historic range in the upper Black Warrior River system are likely to be water quality degradation caused by industrial, mining, agricultural, and urban pollution. The remaining Alabama waterdog populations are isolated from each other by unsuitable habitat created by impoundments, pollution, or other factors. The fragmentation of habitat renders populations vulnerable to catastrophic events such as flood, drought, or chemical spills. In addition, if stream quality improves within areas of the basin, impoundments and polluted reaches will act as barriers to the migration and reestablishment of waterdog populations. This species is also taken for commercial, recreational, scientific, and educational purposes, although this is not considered to represent a serious threat. Disease and predation are not known to be factors in the decline.
The State of Alabama provides no protection for the Alabama waterdog. The Federal Surface Mining Control and Reclamation Act of 1977 and the Clean Water Act of 1972 have been ineffective in preventing the continued decline of species in the Black Warrior basin.
A reserve should be established in an area of the Alabama waterdog’s range that is not severely impacted by population. The State of Alabama should prioritise and fund a conservation programme for this species, and tackle issues of water pollution throughout the Black Warrior drainage basin. Making this species a symbol of local pride in a clean and healthy environment could mobilize public support and drive conservation action across the State.
AmphibiaWeb: Information on amphibian biology and conservation [web application]. 2006. Berkeley, California: AmphibiaWeb. Available: amphibiaweb. Accessed: 08 December 2006.
Ashton Jr., R. E. and Peavy, B. 1985. Tenn-Tom Waterway Necturus project. In: Unpublished report submitted to Alabama Department of Conservation and Natural Resources, Montgomery, Alabama.
Ashton, R. E., Jr. and Peavy, J. 1986. Black Warrior waterdog. In: R.H. Mount (ed.), Vertebrate animals of Alabama in need of special attention, pp. 63-64. Alabama Agricultural Experiment Station, Auburn University, Auburn, Alabama.
Bailey, K.A. and Guyer, C. 1998. Demography and population status of the flattened musk turtle, Sternotherus depressus, in the Black Warrior River basin of Alabama. Chelonian Conservation and Biology 3: 77-83.
Bailey, M.A. 1992. Black Warrior waterdog status survey, Final Report 1990-1992. Unpublished report submitted to Alabama Game and Fish Division, Alabama Department of Conservation and Natural Resources, Montgomery, Alabama 36849. Orihect E-1, Study No. 5.
Bailey, M.A. 1995. Black Warrior waterdog survey 1994-95: Performance report. Unpublished report submitted to Alabama Department of Conservation and Natural Resources, Montgomery, Alabama.
Bailey, M.A. 2005. Family Proteidae. M. Lannoo (ed.). Amphibian Declines: The Conservation Status of United States Species. University of California Press, Ltd. London, England. pp. 648-651.
Bailey, M.A. and Moler, P.E. 2003. Necturus alabamensis Vioscai. Black Warrior waterdog. Catalogue of American Amphibians and Reptiles 761: 1-2.
Bart Jr, H.L., Bailey, M.A., Ashton Jr, R.E. and Moler, P.E. 1997. Taxonomic and nomenclatural status of the upper Black Warrior River waterdog. Journal of Herpetology 31: 192-201.
Behler, J.L. and King, F.W. 1979. The Audubon Society Field Guide to North American Reptiles and Amphibians. Alfred A. Knopf, New York, NY, USA.
Bishop, S.C. 1943. Handbook of Salamanders. Comstock Publishing Company, Inc, Ithaca, New York, USA.
Blackburn, L., Nanjappa, P. and Lannoo, M.J. 2001. An Atlas of the Distribution of U.S. Amphibians. Ball State University, Muncie, IN, USA.
Brode, W.E. 1969. A systematic study of salamanders in the genus, Necturus Rafinesque. Ph.D. Dissertation. University of Southern Mississippi, Hattiesburg.
Conant R. 1975. A Field Guide to Reptiles and Amphibians of Eastern and Central North America. Second Edition. Houghton Mifflin Company, Boston, MA, USA.
Deutsch, W.G., Seesock, W.C., Webber, E.C.and Bayne, D.R. 1990. The impact of poultry rearing operations on water quality and biological communities of second order streams in Cullman and Winston counties, Alabama, 1988-89. Auburn University, Department of Fisheries and Allied Aquacultures, Auburn, Alabama.
Dodd Jr., C.K. 1990. Effects of habitat fragmentation on a stream-dwelling species, the flattened musk turtle Sternotherus depressus. Biological Conservation 54: 33-45.
Dodd, C.K., Enge, K.M. and Stuart, J.N. 1986. The effects of mining siltation on the distribution and abundance of the flattened musk turtle, Sternotherus depressus, in northern Alabama. Denver Wildlife Research Center, Gainesville, Florida.
Frost, Darrel R. 2006. Amphibian Species of the World: an Online Reference. Version 4 (17 August 2006). Electronic Database accessible at:. American Museum of Natural History, New York, USA.
Gunter, G. and Brode, W.E. 1964. Necturus in the state of Mississippi, with notes on adjacent areas. Herpetologica 20: 114-126.
Guttman, S.I., Weight, L.A., Moler, P.A., Ashton, Jr., R.E., Mansell, B.W. and Peavy, J. 1990. An electrophoretic analysis of Necturus from the southeastern United States. Journal of Herpetology 24: 163-175.
Guyer, C. 1997. A status survey of the Black Warrior waterdog (Necturus sp.), Final Report. Unpublished report. Department of Zoology and Wildlife Science, Auburn University, Auburn, Alabama 36849. 334-844-9232.
Guyer, C. 1998. Historical affinities and population biology of the Black Warrior waterdog (Necturus alabamensis). Unpublished report submitted to Alabama Department of Conservation and Natural Resources, Montgomery, Alabama.
Halliday, T. and Adler, C. (eds.). 2002. The new encyclopedia of reptiles and amphibians. Oxford University Press, Oxford.
Hammerson, G. & Bailey, M. 2004. Necturus alabamensis. In: IUCN 2006. 2006 IUCN Red List of Threatened Species. IUCN Red List of Threatened Species. Downloaded on 08 December 2006.
Hartfield, P. 1990. Status survey for mussels in the tributaries of the Black Warrior River, Alabama. U.S. Fish and Wildlife Service, Jackson, Mississippi.
Hecht, M.K. 1958. A synopsis of mud puppies of eastern North America. Proceedings of the Staten Island Institute of Arts and Sciences. 21: 1-38.
IUCN, Conservation International and NatureServe. 2006. Global Amphibian Assessment. Global Amphibian Assessment. Accessed on 08 December 2006.
Maxson, L.R., Moler, P.E. and Mansell,B.W. 1988. Albumin evolution in salamanders of the genus Necturus. Journal of Herpetology 22: 231-235.
Mettee, M.F., O'Neill, P.E., Pierson, J.M. and Suttkus, R.D. 1989. Fishes of the Black Warrior River system in Alabama. Geological Survey of Alabama Bulletin 133: 201 pp.
Mount, R. H. 1975. The Reptiles and Amphibians of Alabama. Auburn University Agricultural Experiment Station, Auburn, AL, USA.
Neill, W.T. 1963. Notes on the Alabama waterdog, Necturus alabamensis Viosca. Herpetologica 19: 166-174.
Obst, F.J., Richter, K. and Jacob, U. 1984. The Completely Illustrated Atlas of Reptiles and Amphibians for the Terrarium. T.F.H. Publication Inc., N.J., U.S.A.
Shepard, T.E., O'Neil, P.E. and McGregor, S.W. 1997. Biological assessment of the Locust Fork system, 1997. Unpublished report submitted to Alabama Department of Conservation and Natural Resources by Geological Survey of Alabama, Tuscaloosa, Alabama. 37 pp.
U.S. Fish and Wildlife Service (USFWS). 1990. Flattened musk turtle recovery plan. Jackson, Mississippi. 15 pp.
U.S. Fish and Wildlife Service (USFWS). 1998. Technical/agency draft Mobile River Basin ecosystem recovery plan.
U.S. Fish and Wildlife Service (USFWS). 1999. Candidate and listing priority assignment forms.
Viosca Jr, P. 1937. A tentative revision of the genus Necturus with descriptions of three new species from the southern Gulf drainage area. Copeia 1937: 120-138.
Ashton Jr., R. E. and Peavy, B. 1985. Tenn-Tom Waterway Necturus project. In: Unpublished report submitted to Alabama Department of Conservation and Natural Resources, Montgomery, Alabama.
Ashton, R. E., Jr. and Peavy, J. 1986. Black Warrior waterdog. In: R.H. Mount (ed.), Vertebrate animals of Alabama in need of special attention, pp. 63-64. Alabama Agricultural Experiment Station, Auburn University, Auburn, Alabama.
Bailey, K.A. and Guyer, C. 1998. Demography and population status of the flattened musk turtle, Sternotherus depressus, in the Black Warrior River basin of Alabama. Chelonian Conservation and Biology 3: 77-83.
Bailey, M.A. 1992. Black Warrior waterdog status survey, Final Report 1990-1992. Unpublished report submitted to Alabama Game and Fish Division, Alabama Department of Conservation and Natural Resources, Montgomery, Alabama 36849. Orihect E-1, Study No. 5.
Bailey, M.A. 1995. Black Warrior waterdog survey 1994-95: Performance report. Unpublished report submitted to Alabama Department of Conservation and Natural Resources, Montgomery, Alabama.
Bailey, M.A. 2005. Family Proteidae. M. Lannoo (ed.). Amphibian Declines: The Conservation Status of United States Species. University of California Press, Ltd. London, England. pp. 648-651.
Bailey, M.A. and Moler, P.E. 2003. Necturus alabamensis Vioscai. Black Warrior waterdog. Catalogue of American Amphibians and Reptiles 761: 1-2.
Bart Jr, H.L., Bailey, M.A., Ashton Jr, R.E. and Moler, P.E. 1997. Taxonomic and nomenclatural status of the upper Black Warrior River waterdog. Journal of Herpetology 31: 192-201.
Behler, J.L. and King, F.W. 1979. The Audubon Society Field Guide to North American Reptiles and Amphibians. Alfred A. Knopf, New York, NY, USA.
Bishop, S.C. 1943. Handbook of Salamanders. Comstock Publishing Company, Inc, Ithaca, New York, USA.
Blackburn, L., Nanjappa, P. and Lannoo, M.J. 2001. An Atlas of the Distribution of U.S. Amphibians. Ball State University, Muncie, IN, USA.
Brode, W.E. 1969. A systematic study of salamanders in the genus, Necturus Rafinesque. Ph.D. Dissertation. University of Southern Mississippi, Hattiesburg.
Conant R. 1975. A Field Guide to Reptiles and Amphibians of Eastern and Central North America. Second Edition. Houghton Mifflin Company, Boston, MA, USA.
Deutsch, W.G., Seesock, W.C., Webber, E.C.and Bayne, D.R. 1990. The impact of poultry rearing operations on water quality and biological communities of second order streams in Cullman and Winston counties, Alabama, 1988-89. Auburn University, Department of Fisheries and Allied Aquacultures, Auburn, Alabama.
Dodd Jr., C.K. 1990. Effects of habitat fragmentation on a stream-dwelling species, the flattened musk turtle Sternotherus depressus. Biological Conservation 54: 33-45.
Dodd, C.K., Enge, K.M. and Stuart, J.N. 1986. The effects of mining siltation on the distribution and abundance of the flattened musk turtle, Sternotherus depressus, in northern Alabama. Denver Wildlife Research Center, Gainesville, Florida.
Frost, Darrel R. 2006. Amphibian Species of the World: an Online Reference. Version 4 (17 August 2006). Electronic Database accessible at:
Gunter, G. and Brode, W.E. 1964. Necturus in the state of Mississippi, with notes on adjacent areas. Herpetologica 20: 114-126.
Guttman, S.I., Weight, L.A., Moler, P.A., Ashton, Jr., R.E., Mansell, B.W. and Peavy, J. 1990. An electrophoretic analysis of Necturus from the southeastern United States. Journal of Herpetology 24: 163-175.
Guyer, C. 1997. A status survey of the Black Warrior waterdog (Necturus sp.), Final Report. Unpublished report. Department of Zoology and Wildlife Science, Auburn University, Auburn, Alabama 36849. 334-844-9232.
Guyer, C. 1998. Historical affinities and population biology of the Black Warrior waterdog (Necturus alabamensis). Unpublished report submitted to Alabama Department of Conservation and Natural Resources, Montgomery, Alabama.
Halliday, T. and Adler, C. (eds.). 2002. The new encyclopedia of reptiles and amphibians. Oxford University Press, Oxford.
Hammerson, G. & Bailey, M. 2004. Necturus alabamensis. In: IUCN 2006. 2006 IUCN Red List of Threatened Species. IUCN Red List of Threatened Species. Downloaded on 08 December 2006.
Hartfield, P. 1990. Status survey for mussels in the tributaries of the Black Warrior River, Alabama. U.S. Fish and Wildlife Service, Jackson, Mississippi.
Hecht, M.K. 1958. A synopsis of mud puppies of eastern North America. Proceedings of the Staten Island Institute of Arts and Sciences. 21: 1-38.
IUCN, Conservation International and NatureServe. 2006. Global Amphibian Assessment. Global Amphibian Assessment. Accessed on 08 December 2006.
Maxson, L.R., Moler, P.E. and Mansell,B.W. 1988. Albumin evolution in salamanders of the genus Necturus. Journal of Herpetology 22: 231-235.
Mettee, M.F., O'Neill, P.E., Pierson, J.M. and Suttkus, R.D. 1989. Fishes of the Black Warrior River system in Alabama. Geological Survey of Alabama Bulletin 133: 201 pp.
Mount, R. H. 1975. The Reptiles and Amphibians of Alabama. Auburn University Agricultural Experiment Station, Auburn, AL, USA.
Neill, W.T. 1963. Notes on the Alabama waterdog, Necturus alabamensis Viosca. Herpetologica 19: 166-174.
Obst, F.J., Richter, K. and Jacob, U. 1984. The Completely Illustrated Atlas of Reptiles and Amphibians for the Terrarium. T.F.H. Publication Inc., N.J., U.S.A.
Shepard, T.E., O'Neil, P.E. and McGregor, S.W. 1997. Biological assessment of the Locust Fork system, 1997. Unpublished report submitted to Alabama Department of Conservation and Natural Resources by Geological Survey of Alabama, Tuscaloosa, Alabama. 37 pp.
U.S. Fish and Wildlife Service (USFWS). 1990. Flattened musk turtle recovery plan. Jackson, Mississippi. 15 pp.
U.S. Fish and Wildlife Service (USFWS). 1998. Technical/agency draft Mobile River Basin ecosystem recovery plan.
U.S. Fish and Wildlife Service (USFWS). 1999. Candidate and listing priority assignment forms.
Viosca Jr, P. 1937. A tentative revision of the genus Necturus with descriptions of three new species from the southern Gulf drainage area. Copeia 1937: 120-138.
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