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Montana Field Guides

Western Toad - Anaxyrus boreas
Other Names:  Bufo boreas

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Species of Concern

Global Rank: G4
State Rank: S2

Agency Status
USFWS:
USFS: SENSITIVE
BLM: SENSITIVE
FWP Conservation Tier: 1

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Copyright by Canadian Amphibian and Reptile Conservation Network
 
General Description
The skin of the adult Western Toad is covered with small round or oval warts on a background color that is usually green or brown; the warts may be reddish-brown and encircled by dark pigment. Parotoid glands are oval and larger than the eyes, cranial crests are absent or indistinct. The eyes have horizontal pupils. Usually there is a light stripe down the middle of the back, but this may be absent or inconspicuous in juveniles. The underside of each hind foot has two brown tubercles that lack sharp cutting edges. Mature males have a dark patch on the inner surface of the innermost digit ("thumb") during breeding. Males lack a vocal sac, however, they may produce a repeated chirping sound. Males rarely exceed 9.5 centimeters snout-vent length (SVL), females rarely 11.0 centimeters.

The body and tail of tadpoles is black or dark brown, with the eyes about midway between the dorsal midline and edge of the head. Labial tooth rows are 2/3, oral papillae are restricted to the sides of the mouth, and the anus is on the midline at the front end of the ventral tail fin; maximum total length is about 3.5 centimeters. The eggs are black, about 1.5 to 1.8 millimeters diameter, and are laid in long 5 millimeters-wide strings of double-layered jelly in two rows (sometimes one or three) that appear to be a single zigzag row.

Diagnostic Characteristics
Adult Western Toads lack the prominent cranial crests found on the other species of Montana toads, and have horizontal rather than vertical pupils, as is present in the Plains Spadefoot. Western Toad tadpoles lack visible white or gold flecks on the back that are present in Woodhouse's and Great Plains Toad tadpoles. Woodhouse's Toad ova are enclosed in a single jelly layer, not two, and Great Plains Toad eggs are in strings that are noticeably pinched between each egg. However, eggs and tadpoles of Western and Woodhouse's Toads are very similar and may be indistinguishable in some cases. Distribution is a useful character for all life stages, as the range and habitat of the Western Toad do not overlap any other toad in Montana, with the possible exception of Woodhouse's Toad (Bufo woodhousii) in a very narrow area north of the Beartooth and Absaroka mountains (Maxell et al. 2003).

General Distribution
Montana Range



Western Hemisphere Range

 


Summary of Observations Submitted for Montana
Number of Observations: 2290

(Click on the following maps and charts to see full sized version) Map Help and Descriptions
Relative Density

Recency

 

(Records associated with a range of dates are excluded from time charts)



Migration
No information is available specific to Montana. Elsewhere is it known that the Western Toad migrates between aquatic breeding and terrestrial nonbreeding habitats. In Colorado, movements of 900 meters (with 95 meters change in elevation) to 4 kilometers have been reported (Hammerson 1999), and radio-tracked females in Idaho have been observed to move up to 2.4 kilometers from breeding ponds (Koch and Peterson 1995). Movement patterns are highly variable, with some individuals remaining in the same location for several days, then moving 50 meters or more on several consecutive nights.

Habitat
Habitats used by Western Toads in Montana are similar to those reported for other regions, and include low elevation beaver ponds, reservoirs, streams, marshes, lake shores, potholes, wet meadows, and marshes, to high elevation ponds, fens, and tarns at or near treeline (Rodgers and Jellison 1942, Brunson and Demaree 1951, Miller 1978, Marnell 1997, Werner et al. 1998, Boundy 2001). Forest cover in or near encounter sites is often unreported, but Western Toads have been noted in open-canopy ponderosa pine woodlands and closed-canopy dry conifer forest in Sanders County (Boundy 2001), willow wetland thickets and aspen stands bordering Engelmann spruce stands in Beaverhead County (Jean et al. 2002), and mixed ponderosa pine/cottonwood/willow sites or Douglas-fir/ponderosa pine forest in Ravalli and Missoula counties (Paul Hendricks, personal observation).

Elsewhere the Western Toad is known to utilize a wide variety of habitats, including desert springs and streams, meadows and woodlands, mountain wetlands, beaver ponds, marshes, ditches, and backwater channels of rivers where they prefer shallow areas with mud bottoms (Nussbaum et al. 1983, Baxter and Stone 1985, Russell and Bauer 1993, Koch and Peterson 1995, Hammerson 1999). Forest cover around occupied montane wetlands may include aspen, Douglas-fir, lodgepole pine, Engelmann spruce, and subalpine fir; in local situations it may also be found in ponderosa pine forest. They also occur in urban settings, sometimes congregating under streetlights at night to feed on insects (Hammerson 1999). Normally they remain fairly close to ponds, lakes, reservoirs, and slow-moving rivers and streams during the day, but may range widely at night. Eggs and larvae develop in still, shallow areas of ponds, lakes, or reservoirs or in pools of slow-moving streams, often where there is sparse emergent vegetation. Adult and juvenile Western Toads dig burrows in loose soil or use burrows of small mammals, or occupy shallow shelters under logs or rocks. At least some Western Toads hibernate in terrestrial burrows or cavities, apparently where conditions prevent freezing (Nussbaum et al. 1983, Koch and Peterson 1995, Hammerson 1999).

Ecological Systems Associated with this Species
  • Details on Creation and Suggested Uses and Limitations
    How Associations Were Made
    We associated the use and habitat quality (high, medium, or low) of each of the 82 ecological systems mapped in Montana for vertebrate animal species that regularly breed, overwinter, or migrate through the state by:
    1. Using personal observations and reviewing literature that summarize the breeding, overwintering, or migratory habitat requirements of each species (Dobkin 1992, Hart et al. 1998, Hutto and Young 1999, Maxell 2000, Foresman 2001, Adams 2003, and Werner et al. 2004);
    2. Evaluating structural characteristics and distribution of each ecological system relative to the species’ range and habitat requirements;
    3. Examining the observation records for each species in the state-wide point database associated with each ecological system;
    4. Calculating the percentage of observations associated with each ecological system relative to the percent of Montana covered by each ecological system to get a measure of “observations versus availability of habitat”.
    Species that breed in Montana were only evaluated for breeding habitat use, species that only overwinter in Montana were only evaluated for overwintering habitat use, and species that only migrate through Montana were only evaluated for migratory habitat use.  In general, species were associated as using an ecological system if structural characteristics of used habitat documented in the literature were present in the ecological system or large numbers of point observations were associated with the ecological system.  However, species were not associated with an ecological system if there was no support in the literature for use of structural characteristics in an ecological system, even if point observations were associated with that system.  High, medium, and low habitat quality was assigned based on the degree to which the structural characteristics of an ecological system matched the preferred structural habitat characteristics for each species in the literature.  The percentage of observations associated with each ecological system relative to the percent of Montana covered by each ecological system was also used to guide assignments of habitat quality.  If you have any questions or comments on species associations with ecological systems, please contact Bryce Maxell at bmaxell@mt.gov or (406) 444-3655.

    Suggested Uses and Limitations
    Species associations with ecological systems should be used to generate potential lists of species that may occupy broader landscapes for the purposes of landscape-level planning.  These potential lists of species should not be used in place of documented occurrences of species (this information can be requested at: http://mtnhp.org/requests/default.asp) or systematic surveys for species and evaluations of habitat at a local site level by trained biologists.  Users of this information should be aware that the land cover data used to generate species associations is based on imagery from the late 1990s and early 2000s and was only intended to be used at broader landscape scales.  Land cover mapping accuracy is particularly problematic when the systems occur as small patches or where the land cover types have been altered over the past decade.  Thus, particular caution should be used when using the associations in assessments of smaller areas (e.g., evaluations of public land survey sections).  Finally, although a species may be associated with a particular ecological system within its known geographic range, portions of that ecological system may occur outside of the species’ known geographic range.

    Literature Cited
    • Adams, R.A.  2003.  Bats of the Rocky Mountain West; natural history, ecology, and conservation.  Boulder, CO: University Press of Colorado.  289 p.
    • Dobkin, D. S.  1992.  Neotropical migrant land birds in the Northern Rockies and Great Plains. USDA Forest Service, Northern Region. Publication No. R1-93-34.  Missoula, MT.
    • Foresman, K.R.  2001.  The wild mammals of Montana.  Special Publication No. 12.  Lawrence, KS: The American Society of Mammalogists.  278 p.
    • Hart, M.M., W.A. Williams, P.C. Thornton, K.P. McLaughlin, C.M. Tobalske, B.A. Maxell, D.P. Hendricks, C.R. Peterson, and R.L. Redmond. 1998.  Montana atlas of terrestrial vertebrates.  Montana Cooperative Wildlife Research Unit, University of Montana, Missoula, MT.  1302 p.
    • Hutto, R.L. and J.S. Young.  1999.  Habitat relationships of landbirds in the Northern Region, USDA Forest Service, Rocky Mountain Research Station RMRS-GTR-32.  72 p.
    • Maxell, B.A.  2000.  Management of Montana’s amphibians: a review of factors that may present a risk to population viability and accounts on the identification, distribution, taxonomy, habitat use, natural history, and the status and conservation of individual species.  Report to U.S. Forest Service Region 1.  Missoula, MT: Wildlife Biology Program, University of Montana.  161 p.
    • Werner, J.K., B.A. Maxell, P. Hendricks, and D. Flath.  2004.  Amphibians and reptiles of Montana.  Missoula, MT: Mountain Press Publishing Company. 262 p.

Food Habits
The diet of Montana adults includes five insect orders: spiders, daddy longlegs, and millipeds (Miller 1978), with ants and ground beetles as common items.

Research outside the state shows that metamorphosed individuals feed on various small invertebrates, including seven orders of flying insects: spiders, mites, daddy longlegs, snails, crayfish, sowbugs (terrestrial amphipods), and earthworms (Nussbaum et al. 1983, Hammerson 1999). Larvae filter suspended plant material or feed on bottom detritus.

Ecology
Basking groups of up to 1000 young-of-year have been observed in Waterton National Park; basking sites may be important at high elevations (Black and Black 1969). They are active day/evening in early summer and late evening/night when it's hot (e.g. August) (Miller 1975).

Generally, Western Toads are active during the day and night; juveniles are largely diurnal while adults tend to be nocturnal except in spring (Maxell 2000). The active period typically begins in April or May and extends to September or October, depending on elevation and latitude (Russell and Bauer 1993, Koch and Peterson 1995, Hammerson 1999), although adults may be active in January and February in coastal populations (Nussbaum et al. 1983). In Montana, records extend from late April to early October (Rodgers and Jellison 1942, Brunson and Demaree 1951, Black and Brunson 1971, Hendricks and Reichel 1996, Boundy 2001).

Predators of adult Western Toads include Raccoon, domestic dog, Coyote, Red Fox, Short-tailed Weasel, American Mink, Marten, Badger, American Black Bear, Northern Pygmy-Owl, Black-billed Magpie, Common Raven, American Crow, Steller's Jay, Gray Jay, American Robin, Loggerhead Shrike, and Northern Shrike (Salt 1979, Corn 1993, Olson 1989, Brothers 1994, Koch and Peterson 1995, Hammerson 1999, Jones et al. 1999). Predators of Western Toad tadpoles include Mallard, Spotted Sandpiper, Terrestrial Gartersnake, Tiger Salamander, Wood Frog tadpoles, and diving beetle larvae. Predators of Western Toad in Montana are not documented.

Reproductive Characteristics
The reproductive biology of Western Toads in Montana is poorly described. The breeding period extends from April to mid-July; breeding aggregations of more than 40 adult males have been reported in mid-May (Black and Brunson 1971). Eggs are laid from early May to late June, tadpoles are present from late May to early September, and recently metamorphosed toadlets have been reported from early June to late August (Brunson and Demaree 1951, Werner and Reichel 1994, Reichel 1995, Hendricks and Reichel 1996, Marnell 1997, Boundy 2001, Burton et al. 2002). Size of one clutch in the Bitterroot Valley of Ravalli County was 20,000 eggs; eggs were laid in late May and produced metamorphosed toadlets by July 11, about 40 to 49 days after oviposition (Maxell et al. 2002).

In other areas of the species' range, the breeding period is known to be variable depending on location; in the mountains it follows the melt of winter snowpack, and in some cases eggs may be laid when ponds are still rimmed with ice. Water temperature may be as low as 7.5 but usually more than 9.0 (Salt 1979). At a given site, breeding may extend over several weeks, although a peak usually occurs.

Breeding aggregations may be quite large (more than 2000 adults), but most often are much smaller, with males outnumbering females. Eggs are laid in two jelly strings usually in shallow water often no more than 15 centimeters deep, and often are deposited in the open with little vegetation cover. Typical clutches in Colorado contain 3000 to 9000 eggs (mean of 5200 eggs), although in the Pacific Northwest clutches of 12,000 eggs are considered normal, and may reach 16,000 eggs (Nussbaum et al. 1983, Koch and Peterson 1995, Hammerson 1999). Eggs hatch in 3 to 12 days. Tadpoles are about 1.0 centimeter total length at hatching and grow to about 2.5 to 3.0 centimeters. They sometimes are found in huge aggregations; one in Yellowstone National Park, Wyoming measured 0.5 meter wide and more than 36 meters long, and another in the Oregon Cascades was 1.0 meter wide by 300 meters long (Nussbaum et al. 1983, Koch and Peterson 1995).

Tadpoles metamorphose in first summer and take two months or more to reach metamorphosis, depending on the water temperature. At high elevation sites near treeline in Wyoming and Colorado tadpoles may fail to metamorphose (Baxter and Stone 1985, Hammerson 1999), and persistence of these populations may be dependent on immigration of juveniles and adults from lower elevation; overwintering of tadpoles has never been observed. Metamorphosis usually occurs in August in Colorado and Oregon, but may occur in late July to mid-September. Recently metamorphosed toadlets measure about 1.0 to 1.6 centimeters snout-vent length (SVL), but can be 1.6 to 2.0 centimeters.

Toadlets may overwinter along the borders of the pools where they developed or move to other nearby wetlands. Juveniles 2.0 to 3.5 centimeters SVL often are present in wetlands adjacent to breeding sites (Nussbaum et al. 1983, Hammerson 1999). Breeding adult males are 5.5 to 9.3 centimeters SVL, adult females 7.5 to 11.5 centimeters SVL. The minimum age of breeding males is four years, and six years for breeding females; captive animals have lived up to 35 years (Russell and Bauer 1993, Hammerson 1999).

Management
The Western Toad was considered the most abundant amphibian of the western third of the state in previous decades (Rodgers and Jellison 1942, Brunson 1952, Maxell et al. 2003), and is still encountered widely and frequently though by no means commonly, and is no longer ranked as the most abundant amphibian. Numerous surveys since the early 1990's indicate that this species has experienced regional population declines in the state. Western Toads were documented to breed at only 2 to 5% of more than 2000 standing water bodies surveyed since 1997, and where breeding was documented, fewer than 10 breeding females contributed in a given year (Maxell 2000, Maxell et al. 2003). Thus, range-wide declines for this species in the western United States are also reflected in the Montana results. Because the reasons for declines in Montana remain obscure, it is difficult to suggest management techniques to reverse the trend. Nevertheless, the following should help reduce some immediate impacts. First, reduced access by livestock to known breeding sites within grazing allotments will prevent undue trampling mortality (Bartelt 1998). This can be accomplished by constructing partial or complete livestock exclosures (fencing) at breeding ponds and other sites. Second, use of fertilizers, herbicides, and pesticides within at least a 100 meters buffer zone of breeding sites should be avoided. Third, stocking predatory game fish at sites currently lacking them should be avoided, even though there is evidence that some species of trout do not prey on Western Toad tadpoles and eggs (Jones et al. 1999). And before efforts are enacted to remove undesirable fish by use of lethal chemicals, a survey for toads at the target water bodies should be conducted to prevent unnecessary mortality to any life stages. If Western Toads are present, they can be removed by dipnet and held in captivity (under appropriate conditions) until the effects of the treatment dissipate, then returned to the site. Finally, known breeding sites should not be drained or altered, and water bodies where alteration is planned should first be surveyed for use by Western Toads.

References
  • Literature Cited AboveLegend:   View WorldCat Record   View Online Publication
    • Bartelt, P.E. 1998. Natural history notes: Bufo boreas mortality. Herpetological review 29(2):96.
    • Baxter, G. T. and M. D. Stone. 1985. Amphibians and reptiles of Wyoming. Second edition. Wyoming Game and Fish Department. Cheyenne, WY. 137 p.
    • Black, J. H. and R. B. Brunson. 1971. Breeding behavior of the boreal toad (Bufo boreas boreas) (Baird and Girard) in western Montana. Great Basin Naturalist 31: 109-113.
    • Black, J.H. and J.N. Black. 1969. Postmetamorphic basking aggregations of the boreal toad (Bufo boreas boreas). Canadian Field Naturalist 83: 155-156.
    • Boundy, J. 2001. Herpetofaunal surveys in the Clark Fork Valley region, Montana. Herpetological Natural History 8: 15-26.
    • Brothers, D. R. 1994. Bufo boreas (western toad) predation. Herpetological Review 25(3): 117.
    • Brunson, R. B. 1952. Recent collections of Bufo boreas from western Montana. Proceedings of Montana Academy of Sciences 11: 17-19.
    • Brunson, R. B. and H. A. Demaree, Jr. 1951. The herpetology of the Mission Mountains, Montana. Copeia (4):306-308.
    • Burton, S.R., D.A. Patla, and C.R. Peterson. 2002. Amphibians of Red Rock Lakes National Wildlife Refuge: occurrence, distribution, relative abundance, and habitat associations. Herpetology Laboratory, Department of Biological Sciences, Idaho State University, Pocatello, ID. 66 p.
    • Corn, P.S. 1993. Bufo boreas (boreal toad): predation. Herpetological Review 24(2): 57.
    • Hammerson, G. A. 1999. Amphibians and reptiles in Colorado. University Press of Colorado & Colorado Division of Wildlife. Denver, CO. 484 p.
    • Hendricks, P. and J. D. Reichel. 1996. Amphibian and reptile survey of the Bitterroot National Forest: 1995. Montana Natural Heritage Program. Helena, MT. 95 p.
    • Jean, C., P. Hendricks, M. Jones, S. Cooper, and J. Carlson. 2002. Ecological communities on the Red Rock Lakes National Wildlife Refuge: inventory and review of aspen and wetland systems. Report to Red Rock Lakes National Wildlife Refuge.
    • Jones, M. S., J. P. Goettl, and L. J. Livo. 1999. Bufo boreas (boreal toad): predation. Herpetological Review 30:91.
    • Koch, E. D. and C. R. Peterson. 1995. Amphibians and reptiles of Yellowstone and Grand Teton national parks. University of Utah Press, Salt Lake City, UT. 188 p.
    • Marnell, L. E. 1997. Herpetofauna of Glacier National Park. Northwestern Naturalist 78:17-33.
    • Maxell, B. A. 2000. Management of Montana's amphibians: a review of factors that may present a risk to population viability and accounts on the identification, distribution, taxonomy, habitat use, natural history, and the status and conservation of individual species. Report to USFS Region 1, Order Number 43-0343-0-0224. University of Montana, Wildlife Biology Program. Missoula, MT. 161 p.
    • Maxell, B. A., J. K. Werner, P. Hendricks and D. L. Flath. 2003. Herpetology in Montana: a history, status summary, checklists, dichotomous keys, accounts for native, potentially native, and exotic species, and indexed bibliography. Society for Northwestern Vertebrate Biology, Northwest Fauna Number 5. Olympia, WA. 135 p.
    • Maxell, B.A., K.J. Nelson, and S. Browder. 2002. Record clutch size and observations on breeding and development of the western toad (Bufo boreas) in Montana. Northwestern Naturalist 83(1):27-30.
    • Miller, J. D. 1975. Interspecific food relationships of anurans in northwestern Montana and fluoride accumulation in amphibians and reptiles in northwestern Montana. M.S. thesis. University of Montana, Missoula, MT. 105 p.
    • Miller, J. D. 1978. Observations on the diets of Rana pretiosa, Rana pipiens, and Bufo boreas from western Montana. Northwest Science 52(3): 243-249.
    • Nussbaum, R. A., E. D. Brodie, Jr., and R. M. Storm. 1983. Amphibians and reptiles of the Pacific Northwest. University of Idaho Press. Moscow, ID. 332 p.
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    • Reichel, J. D. 1995. Preliminary amphibian and reptile survey of the Lewis and Clark National Forest: 1994. Montana Natural Heritage Program. Helena, MT. 92 p.
    • Rodgers, T. L. and W. L. Jellison. 1942. A collection of amphibians and reptiles from western Montana. Copeia (1):10-13.
    • Russell, A. P. and A. M. Bauer. 1993. The amphibians and reptiles of Alberta. University of Calgary Press. Calgary, Alberta. 264 p.
    • Salt, J.R. 1979. Some elements of amphibian distribution and biology in the Alberta Rockies. Alberta Naturalist 9(3): 125-136.
    • Werner, J.K. and J.D. Reichel. 1994. Amphibian and reptile survey of the Kootenai National Forest: 1994. Montana Natural Heritage Program. Helena, MT. 104 p.
    • Werner, J.K., T. Plummer, and J. Weaslehead. 1998. Amphibians and reptiles of the Flathead Indian Reservation. Intermountain Journal of Sciences 4(1-2): 33-49.
  • Additional ReferencesLegend:   View WorldCat Record   View Online Publication
    Do you know of a citation we're missing?
    • [PRESI] Powder River Eagle Studies Incorporated. 1998b. Spring Creek Mine 1997 wildlife monitoring studies. Powder River Eagle Studies Incorporated. Gillete, WY.
    • [USDAFS] USDA Forest Service. 1999. Update of U.S. Forest Service Northern Region Sensitive Species list. 12 March, 1999. Region 1 U.S. Forest Service Supervisors Office, Missoula, MT. 20 p.
    • [WWPC] Washington Water Power Company. 1995. 1994 wildlife report Noxon Rapids and Cabinet Gorge Reservoirs. Washington Water Power Company. Spokane, WA.
    • Adams, M.J., B.R. Hossack, and R.A. Knapp. 2005. Distribution patterns of lentic-breeding amphibians in relation to ultraviolet radiation in western North America. Ecosystems 8(5):488-500.
    • Adams, S.B., D.A. Schmetterling, and M.K. Young. 2005. Instream movements by boreal toads (Bufo boreas boreas). Herpetological Review 36(1):27-33.
    • Aguirre, A. 1994. Declining toad populations. Conservation Biology 8(1): 7.
    • Anderson, M.E. 1977. Aspects of the ecology of two sympatric species of Thamnophis and heavy metal accumulation with the species. M.S. thesis, University of Montana, Missoula. 147 pp.
    • Annis, S.L., F.P. Dastoor, H. Ziel, P. Daszak, and J.E. Longcore. 2004. A DNA-based assay identifies Batrachochytrium dendrobatidis in amphibians. Journal of Wildlife Diseases 40(3):420-428.
    • Atkinson, E.C. and M.L. Atkinson. 2003. Status of boreal toads on and adjacent to the Gallatin National Forest, Montana, with special reference to the Hebgen Lake and Bozeman Ranger Districts. Marmot's Edge Conservation. Report to the US Department of Agriculture. Forest Service, Bozeman, MT.
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    • Bartelt, P.E. and C.R. Peterson. 2000. A description and evaluation of a plastic belt for attaching radio transmitters to western toads (Bufo boreas). Northwestern Naturalist 81: 122-128.
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    • Beal, M.D. 1951. The occurrence and seasonal activity of vertebrates in the Norris and Gibbon Geyser Basins of Yellowstone National Park. M.S. Thesis. Utah State Agricultural College. Logan, Utah. 61 pp.
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    • Beiswenger, R.E. 1981. Predation by gray jays on aggregating tadpoles of the boreal toad (Bufo boreas). Copeia 1981(2): 459-460.
    • Belden, L.K., E.L. Wildy, and A.R. Blaustein. 2000b. Juvenile western toads, Bufo boreas, avoid chemical cues of snakes fed juvenile, but not larval, conspecifics. Animal Behaviour 59: 871-875.
    • Benard, M.F. and J.A. Fordyce. 2003. Are induced defenses costly? Consequences of predator-induced defenses in western toads, Bufo boreas. Ecology 84(1):68-78.
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    • Black, J.H. 1970. Some aspects of the distribution, natural history and zoogeography of the toad genus Bufo in Montana. M.S. thesis. University of Montana, Missoula, MT. 70 p.
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    • Blaustein, A.R., J.M. Romansic, and E.A. Scheessele. 2005. Ambient levels of ultraviolet-B radiation cause mortality in juvenile western toads, Bufo boreas. American Midland Naturalist 154(2):375-382.
    • Blaustein, A.R., K.S. Chang, H.G. Lefcort and R.K. O'Hara 1990. Toad tadpole kin recognition: recognition of half siblings and the role of maternal cues. Ethology, Ecology and Evolution 2(2): 215-226.
    • Blaustein, A.R., P.D. Hoffman, D.G. Hokit, J.M. Kiesecker, S.C. Walls, and J.B. Hays. 1994. UV repair and resistance to solar UV-B in amphibian eggs: a link to population declines? Proceedings of the National Academy of Sciences 91: 1791-1795.
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    • Campbell, J.B. 1970b. Hibernacula of a population of Bufo boreas boreas in the Colorado Front Range. Herpetologica 26: 278-282.
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