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USC Vernal Pool Reference List

These are very useful resources available online at the following addresses:

A Field Guide to the Animals of Vernal Pools

This field guide was a cooperative effort between the MA Natural Heritage & Endangered Species Program and the Vernal Pool Association and was written by Leo Kenney and Matthew Burne. For more information click the links below:

in AlA  B  C  D  E  F  G  H  J  K  M  N  P  R  S  T  V  W


Adam, M.D., and M.J. Lacki. 1993. Factors affecting amphibian use of road-rut ponds in Daniel Boone National Forest. Trans. Ky. Acad. Sci. 54:13-16.

Alford, R.A., and R.N. Harris. 1988. Effects of larval growth history on anuran metamorphosis. The American Naturalist 131:91-106.

Alford, R.A., and S.J. Richards. 1999. Global amphibian declines: A problem in applied ecology. Annu. Rev. Ecol. Syst. 30:133-165.


Babbit, K.J., G.W. Tanner 2000. Use of Temporary Wetlands by Anurans in a Hydrologically Modified Landscape. Wetlands, Vol. 20 No 2 pp. 313-322.

Baldauf, R. J. 1952. Climatic factors influencing the breeding migration of the spotted salamander (Ambystoma maculatum). Copeia 1952: 178-181.

Banks, B. and J. C. Beebee. 1987. Factors influencing breeding site choice by the pioneering amphibian Bufo calamita. Holarctic Biology 10: 14-21.

Beck, C.W. 1997. Effect of changes in resource level on age and size at metamorphosis in Hyla squirrella. Oecologia 112:187-192.

Beebee, J. C. Ecology and Conservation of Amphibians. London: Chapman & Hall, 1996.

Beebee, T.J.C. 1997. Changes in dewpond numbers and amphibian diversity over 20 years on chalk downland in Sussex, England. Biological Conservation3 81:215-219.

Beneski, J. T, Jr., E. J. Zalisko, and J. H. Larsen, Jr.1986. Demography and migratory patterns of the eastern long-toed salamander, Ambystoma macrodactylum columbianum. Copeia: 398-

Berven, K.A. 1981. Mate choice in the wood frog, Rana sylvatica. Evolution 35:707-722.

Berven, K.A. 1990. Factors affecting population fluctuations in larval and adult stages of the wood frog (Rana sylvatica). Ecology 71:1599-1608.

Berven, K.A., and T.A. Grudzien. 1990. Dispersal in the wood frog (Rana sylvatica): Implications for genetic population structure. Evolution 44:2047-2056.

Biebighauser, T. R. 2002. A Guide to Creating Vernal Ponds. USDA Forest Service. 33 pages.

Bishop, S. C. 1943. Handbook of Salamanders. Cornell University Press, Ithaca.

Blanchard, F. N. 1930. The stimulus to the breeding migration of the spotted salamander (Ambystoma maculatum). American Naturalist 64: 154-167.

Blaustein, A. R., D. B. Wake, and W. P. Sousa. 1994. Amphibian declines: judging stability, persistence, and susceptibility of populations to local and global extinctions. Conservation Biology 8: 60-71.

Brooks, R.T., M. Hayashi. 2002. Depth-Area-Volume and Hydroperiod relationships of Ephemeral (vernal) Forest Pools in Southern New England. Wetlands 22(2):247-255

Brooks, R.T., J. Stone, P. Lyons. 1998. An Inventory of Seasonal Forest Ponds on the Quabbin Reservoir Watershed, Massachusetts. Northeastern Naturalist 5(3):219-230.

Brooks, Robert T. 2000. Annual and Seasonal Variation and the effects of Hydroperiod on Benthic Macroinvertebrates of seasonal forest ("vernal") ponds in Central Massachusetts, USA. Wetlands Vol. 20 pp. 707-715.

Buhlmann, K.A. and J.W. Gibbons.  2001.  Terrestrial habitat use by aquatic turtles from a seasonally fluctuating wetland: implications for wetland conservation boundaries.  Chelonian Conservation and Biology 4(1):115-127.  [Chelydra serpentina, Clemmys guttata, Kinosternon subrubrum, Sternotherus odoratus]

Burne, M.R. 2000. Conservation of Vernal Pool-Breeding Amphibian Communities: Habitat and Landscape Associations with Community Richness. M.S. Thesis. University of Massachusetts, Amherst.

Burne, M.R. 2001. Massachusetts aerial photo survey of potential vernal pools. Natural Heritage and Endangered Species Program, Massachusetts Department of Fisheries and Wildlife, Westborough, MA.

Burton, T.M., and G.E. Likens. 1975. Salamander populations and biomass in the Hubbard Brook Experimental Forest, New Hampshire. Copeia 1975:541-546.

Bury, R. B. and R. A. Luckenbach. 1976. Introduced amphibians and reptiles in California. Bio. Conserv.10:1-14.

Butterwick, M. 1998. The Hydrogeomorphic Approach and Its Use in Vernal Pool Functional Assessment. Pages 50-55 in: C.W. Witham, Editor. Ecology, Conservation, and Management of Vernal Pool Ecosystems: Proceedings from a 1996 Conference, 19-21 June, 1996, Sacramento, CA. California Native Plant Society. Sacramento, CA. 285 pp.


Calhoun, A. J. K. and P. deMaynadier. 2002. Forestry habitat management guidelines for vernal pool wildlife in Maine. Maine Department of Inland Fisheries and Wildlife.

Calhoun, A.J.K. 1998. Book Review. Northern Forested wetlands: ecology and management. Forest Science 44: 329-330.

Calhoun, A.J.K. 1999. Maine's citizen guide to locating and documenting vernal pools. Maine Audubon Society, Falmouth, Maine.

Calhoun, A.J.K. and M. W. Klemens. 2002. Best development practices for vernal pools: conserving pool-breeding amphibians in commercial and residential developments in the northeastern US. Wildlife Conservation Society Technical Paper #5. Rye, New York.

Calhoun, A.J.K., T. Walls, M. McCollough, and S. Stockwell. In press. Evaluating vernal pools as a basis for conservation strategies: a Maine case study. Wetlands.

Chazal, A. D. and P. H. Niewiaroski. 1998. Responses of the mole salamander, Ambystoma talpoideum, to clear cutting: assessing the consequences of forest management with field experiments. Ecological Applications 8: 1133-1143.

Colburn, E.A. 1997. Certified: A citizen's step-by-step guide to protecting vernal pools. Massachusetts Audubon Society , Lincoln, MA.

Colburn, E.A. 2001. Small Pools Close Up: Examining Vernal Pools of the Northeast. National Wetlands Newsletter 23:1 7-8, 17-18.

Colburn, E.A., ed. 1997. Certified: A Citizen's Step-by-Step Guide to Protecting Vernal Pools, 7th ed. Massachusetts Audubon Society, Lincoln, MA. 109 pages.

Colburn, E.A. Vernal Pools of Glaciated Northeastern North America: A Community Profile," manuscript in preparation.

Conant, R. and J. T. COLLINS. 1998. A Field Guide to Reptiles and Amphibians of Eastern and Central North America. Third Edition Expanded. Houghton Mifflin Co., Boston.


De Weese, June M. Vernal Pool Construction Monitoring Methods and Habitat Replacement Evaluation. Pages 217-223 in: C.W.Witham, E.T. Bauder, D. Belk, W.R.FerrenJr., and R. Ornduff (Editors). Ecology, Conservation, and Management of Vernal Pool Ecosystems - Proceedings from a 1996 conference . California Native Plant Society, Sacramento, CA 1998.

DeMaynadier, P.G. and M.L. Hunter Jr. 1998. Effects of silvicultural edges on the distribution and abundance of amphibians in Maine. Conservation Biology 12:340-352.

Demaynadier, P.G., and M.L. Hunter, Jr. 1995. The relationship between forest management and amphibian ecology: a review of the North American literature. Environmental Reviews 3:230-261.

Demaynadier, P.G., and M.L. Hunter, Jr. 1999. Forest canopy closure and juvenile emigration by pool-breeding amphibians in Maine. Journal of  Wildlife Manage. 63:441-450.

DiMauro, D., M.L. Hunter. 2002. Reproduction of amphibians in natural and anthropogenic temporary pools in managed forests. Forest Science 48 (2) 2002.

Dodd, C.K.Jr.  2001.  North American Box Turtles: A Natural History.  University of Oklahoma Press, Norman, OK.

Donahue, D.F. 1997. A guide to the identification and protection of vernal pool wetlands of Connecticut. State University of Connecticut Cooperative Extension System.

Doty, T.L. 1978. A study of larval amphibian population dynamics in a Rhode Island vernal pond. Ph.D. Dissertation. University of Rhode Island, Kingston.

Douglas , M. E. and B. L. Monroe. 1981. A comparative study of topographic orientation in Ambystoma (Amphibia: Caudata). Copeia :460-463.

Douglas, M. E. 1979. Migration and sexual selection in Ambystoma jeffersonianum. Canadian Journal of Zoology 57: 2303-2310.

Douglas, M.E. 1975. A study of three sympatric ambystomid salamanders in Bernheim Forest, Bullitt County, Kentucky. Abstracts, Am. Soc. of Ichthyologists and Herpetologists:38.

Duellman, W. E. and L. Trueb. Biology of Amphibians. New York, NY: McGraw Hill, 1986.

Dunn, E. R. 1940. The races of Ambystoma tirginum. Copeia 1940(3):154-162.

Dunson, W.A., R.L. Wyman, and E.S. Corbett. 1992. A symposium on amphibian declines and habitat acidification. Journal of Herpetology 26:349-352.


Enge, K.M., and W.R. Marion. 1986. Effects of clear cutting and site preparation on herpetofauna of a north Florida flatwoods. Forest Ecology and Management 14:177-192.


Fahrig, L, J. H. Pedlar, S. E. Pope, P. D. Taylor, and J. F. Wegner. Effect of road traffic on amphibian density. Biological Conservation 73: 177-182.


Gates, E.J., and E.L. Thompson. 1981. Breeding Habitat Association of Spotted Salamanders (Ambystoma maculatum) in Western Maryland. The Journal of the Elisha Mitchell Scientific Society 97: 209-216.

Gibbs, J. P. 1998. Distribution of woodland amphibians along a forest fragmentation gradient. Landscape Ecology 13:263-268.

Gibbs, J.P. 1993. Importance of small wetlands for the persistence of local populations of wetland-associated animals. Wetlands 13:25-31.

Golet, F.C., A.J.K. Calhoun, W.R. DeRagon, D.J. Lowry, and A.J. Gold. 1993. Ecology of red maple swamps in the glaciated northeast: a community profile. U.S. Department of the Interior, Fish and Wildlife Service, Biological Report 12, Washington, D.C.


Harding, J. H. 1997. Amphibians and Reptiles of the Great Lakes Region. The University of Michigan Press, Ann Arbor. 378 pp.

Hulse, A.C., C.J. McCoy, E.J. Censky.  2001.  Amphibians and Reptiles of Pennsylvania and the Northeast.  Cornell University Press, Ithaca, NY.

Husting, E. L. Survivorship and breeding structure of population of Ambystoma maculatum. Copeia 1965: 352-362.


Jackson, M.E., D.E. Scott, and R. Estes. 1989. Determinants of nest success in the marbled salamander (Ambystoma opacum). Canadian Journal of Zoology 67: 2277-2281.

Jaeger, R. G. Density-dependent and density-independent causes of extinction of a salamander population. Evolution 34: 617-621.


Keeley, J.E. and P.H. Zedler. 1998. Characterization and Global Distribution of Vernal Pools. Pages 1-14 in: C.W. Witham, Editor. Ecology, Conservation, and Management of Vernal Pool Ecosystems: Proceedings from a 1996 Conference, 19-21 June, 1996, Sacramento, CA. California Native Plant Society. Sacramento, CA. 285 pp.

Kenney, L. 1996. Student certification of Massachusetts vernal pools. Wetland Journal 8(1):4-7.

Kenney, L. P. 1995. Wicked Big Puddles: A Guide to the Study and Certification of Vernal Pools. Reading Memorial High School-Vernal Pool Association, Reading, MA. 58 pages

Kenney, L.P. and M.R. Burne. 2001. A Field Guide to the Animals of Vernal Pools. Massachusetts Division of Fisheries and Wildlife and Natural Heritage and Endangered Species Program. Westborough, Massachusetts. 77p.

Klefberger, S.R., and J.K. Werner. 1983. Post-breeding migration and summer movement of Ambystoma maculatum. Journal of Herpetology 17:176-177.


McKnight, M. L. and H. B. Shaffer. 1997. Large, rapidly evolving intergenic spacers in the mitochondrial DNA of the salamander family Ambystomidae (Amphibia: Caudata). Molecular Biology and Evolution 14: 1167-1176.

Morin, P. J., S. P. Lawler, and E. A. Johnson. 1990. Ecology and beeding phenology of larval Hyla andersonii: the disadvantage of breeding late. Ecology 71: 1590-1598.


Nunney, L. and K. A. Campbell. 1993. Assessing minimum viable population size: demography meets population genetics. Trends in Ecology and Evolution 8:234 - 239.


Parmar, M.K.B., and D. Macmn. 1995. Survival analysis: A practical approach. Wiley, New York. 255 p.

Paton, P.W.C, W.B. Crouch. 2002. Using the phenology of pond-breeding amphibians to develop conservation strategies. Conservation Biology 16(1):194-204

Pauley, T.K. 1994. The impact of habitat disturbances on amphibian survival in West Virginia. U.S. Environmental Protection Agency Mid-Atlantic Highlands Area Environmental! Monitoring and Assessment Conference. Abstract Booklet. Hershey, PA.

Pechmann, J. H. K., D. E. Scott, R. D. Semlitsch, J. P. Caldwell, L. J. Vitt, and W. Gibbons. 1991. Declining amphibian populations: the problem of separating human impacts from natural populations. Science 253:892-895.

Pechmann, J.H.K., D.E. Scott, J.W. Gibbons and R.D. Semlitsch. 1989. Influence of wetland hydroperiod on diversity and abundance of metamorphosing juvenile amphibians. Wetlands Ecology Management. 1: 3-11.

Pentis, A., M.A. Pentis. 1999. The magic of vernal pools. Coyote Junction Studio, Ramona, CA. 45 pages.

Petranka, J. W. 1998. Salamanders of the United States and Canada. Smithsonian Institute Press, Washington and London.

Pfingsten, R. A. and F. L. DOWNS. 1989. Salamanders of Ohio. Bulletin of the Ohio Biological Survey. Vol. 7 No. 2. College of Biological Sciences The Ohio State University, Columbus.

Phillips, C. A. 1994. Geographic distribution of mitochondrial DNA variants and the historical biogeography of the spotted salamander, Ambystoma maculatum. Evolution 48: 597-607.

Phillips, C. A. and O. J. Sexton. 1989. Orientation and sexual differences during breeding migrations of the spotted salamander, Ambystoma maculatum. Copeia 1989: 17-22.

Pounds, J. A. and M. L. Crump. 1994. Amphibian declines and climate disturbance: the case of the golden toad and the harlequin frog. Conservation Biology 8: 72-85.

Preiller, E.L., J.Y. Kefer, J.D. Lawrence. 2000. Vernal pool conservation in Connecticut: an assessment and recommendations. Environmental Management 26(5):503-513.

Preston, H.Y. 1995. The conservation of vernal pools. Wheaton College, Norton, MA. Thesis.


Raymond, L.R., and L.M. Hardy. 1991. Effects of a clearcut on a population of the mole salamander, Ambystoma talpoideum, in an adjacent unaltered forest. J. Herpetol. 25:509-512.

Rowe, C.L. and W.A. Dunson. 1995. Impacts of hydroperiod on growth and survival of larval amphibians in temporary ponds of Central Pennsylvania, USA. Oecologia 102: 397-403.


Schmidt, K. P. 1938. Herpetological Evidence for the Postglacial Eastward Extension of the Steppe in North America. Ecology 19:396-407.

Schmieder, R. 1996. The North American Amphibian Monitoring Program (NAAMP), http://www.pwrc.usgs.gov/naamp/

Schneider, D.W. and T.M. Frost, "Habitat duration and community structure in temporary ponds," Journal of North American Benthological Society 15 no. 1 (1996): 64-86

Semlitsch, R.D. 1987. Relationship of pond drying to the reproduction success of the salamander Ambystoma tadpodeum. Copeia 1987: 61:69.

Semlitsch, R.D. 1987. Relationship of pond drying to the reproductive success of the salamander Ambystoma talpoideum. Copeia 1987:61-69.

Semlitsch, R.D. 1998. Biological Delineation of Terrestrial Buffer Zones for Pond-Breeding Salamanders. Conservation Biology 12: 1113-1119.

Semlitsch, R.D. and H.W. Wilbur. 1988. Effects of pond drying time on metamorphosis and survival in the salamander Ambystoma tadpoideum. Ecology 55: 440-444.

Semlitsch, R.D. and J.R. Bodie. 1998. Are Small Isolated Wetlands Expendable? Conservation Biology 12: 1129-1133.

Semlitsch, R.D., D.E. Scour, and J.H.K. Pechmann. 1988. Time and size at metamorphosis related to adult fitness in Ambystoma talpoideum. Ecology 69:184-192.

Sexton, O. J., C. Phillips and J. E. Bramble. 1990. The effects of temperature and precipitation on the breeding migration of the spotted salamander (Ambystoma maculatum). Copeia 1990: 781-787.

Shaffer, H. B., and M. L. McKnight. 1996. The polytypic species revisited: differentiation and molecular phylogenetics of the tiger salamander Ambystoma tigrinum (Amphibia: Caudata) complex. Evolution 50: 417-433.

Shoop, C. R. 1965. Orientation of Ambystoma maculatum: movements to and from breeding ponds. Science 149: 558-559.

Shoop, C. R. 1968. Migratory orientation of Ambystoma maculatum: movement near breeding ponds and displacements of migrating individuals. Biological Bulletin 135:230-238.

Shoop, C. R. 1974. Yearly variation in larval survival of Ambystoma maculatum. Ecology 55: 440-444.

Skelly, D.K. 1996. Pond drying, predators, and the distribution of Pseudacris tadpoles. Copeia 1996:599-605.

Skelly, D.K. 1997. Tadpole communities: Pond permanence and predation are powerful forces shaping the structure of tadpole communities. Am. Sci. 85:36-45.

Smith, D.C. 1987. Adult recruitment in chorus frogs: Effects of size and date at metamorphosis. Ecology 68:344-350.

Snodgrass, J.W., J.W. Ackerman, A.L. Bryan Jr., J. Burger. 1999. Influence of Hydroperiod, Isolation, and Heterospecifics on the Distribution of Aquatic Salamanders (Siren and Amphiuma) among Depression Wetlands. Copeia 1999(1) 107-113.

Snodgrass, J.W., M.J. Komoroski, A.L. Bryan, Jr., and J. Burger. 2000b. Relationships among isolated wetland size, hydroperiod, and amphibian species richness: Implications for wetland regulations. Conservation Biology 14: 414-419.

Sobczak, R. "Surface Water Fluctuations in Depressional Wetlands, Cape Cod National Seashore, Eastham, MA, in Realterion to Precipitation and Groundwater, 1982-1997: A Pilot Study" unpublished report to the National Park Service, Cape Cod National Seashore, Wellfleet, Mass. (Cape Cod Commission, Barnstable, Mass., 2000).

Squire, T. and R.A. Newman.  2002.  Fine-scale population structure in the wood frog (Rana sylvatica) in a northern woodland.  Herpetologica  58(1):119-130.

Stenhouse, S. L. Migration, orientation and homing in Ambsytoma maculatum. Copeia 1985: 631-637.

Stine, C. J. 1984. The Life History and Status of the Eastern Tiger Salamander, Ambystoma tigrium. Bulletin of the Maryland Herpetological Society. Vol. 20 No. 3.

Stone, J.S. 1992. Vernal pools in Massachusetts: Aerial photographic identification, biological and physiographic characteristics, and state certification criteria. M.S. Thesis, University of Massachusetts, Amherst, MA.

Stumpel, A.H.P., and H. Van der Voet. 1998. Characterizing the suitability of new ponds for amphibians. Amphibia-Reptilia 19:125-142.


Tappan, A.M. 1997. Identification and documentation of vernal pools in New Hampshire. New Hampshire Fish and Game Department, Concord, NH. 72 pages.

Trenham, P. C. 1998. Demography, migration, and metapopulation structure of pond breeding salamanders. Dissertation, University of California, Davis. 96pp


Van Dusen, B. 1995. Pondwatchers : a guide to ponds and vernal pools of Eastern North America. Massachusetts Audubon Society, Lincoln, MA.

Verde, T. 1997. Hurray for potholes. Wildlife Conservation, March/April.

Vogt, R. C. 1981. Natural History of Amphibians and Reptiles of Wisconsin. The Milwaukee Public Museum, Milwaukee, Wisconsin.


Waldick, R.C. 1994. Implications of forestry-associated habitat conversion on amphibians in the vicinity of Fundy National Park, New Brunswick. M.S. thesis, Dalhousie University, Halifax, Nova Scotia.

Wederkinch, E.W. 1988. Population size, migration barriers, and other features of Rana dalmatina populations near Koge, Zealand, Denmark. Memoranda Societatis Fauna et Flora Fennica 64:101-103.

Weir, B. S. Genetic Data Analysis II. Sunderland, MA: Sinauer Associates Inc., 1996.

Werner, E.E. and K.S. Glennemeier. 1999. Influence of forest canopy cover on the breeding pond distributions of several amphibian species. Copeia 1999: 1-12.

Whitford, W. G. and A. Vinegar. 1966. Homing, survivorship, and overwinter of larvae in spotted salamanders, Ambystoma maculatum. Copeia 1966: 515-519.

Wiggins, G.B., R.J. Mackay, and I.M. Smith. 1980. Evolutionary and ecological strategies of animals in annual temporary pools. Archiv fur Hydrobiologie. Supplement. 58:97-206.

Wilbur, H.M. 1987. Regulation of structure in complex systems: Experimental temporary pond communities. Ecology 68:1437-1452.

Windmiller, B.S. 1996. The pond, the forest, and the city: spotted salamander ecology and conservation in human-dominated landscape. Ph.D. Dissertation, Tufts University, Boston, Massachusetts.

Witham, C.W. 1998. Ecology, conservation, and management of vernal pool ecosystems: proceedings from a 1996 conference California Native Plant Society, Sacramento, CA. 285 pages

Wright, A. H. and A.A. Allen. 1909. The early breeding habits of Amblystoma punctatum. American Naturalist. 43: 687-692.

Wyman, R. L. 1990. What's happening to the amphibians? Conservation Biology 4:3 50-352.

Wyman, R.L. 1998. Experimental assessment of salamanders as predators of detrital food webs: Effects on invertebrates, decomposition and the carbon cycle. Biodiversity and Conservation 7:641-650.













Upper Susquehanna Coalition
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Coordinator: Wendy Walsh - 183 Corporate Drive - Owego, NY 13827 - (607) 687-3553 wwalsh@u-s-c.org