Ambystoma jeffersonianum - (Green, 1827)
Jefferson Salamander
Other English Common Names: Jefferson salamander
Taxonomic Status: Accepted
Related ITIS Name(s): Ambystoma jeffersonianum (Green, 1827) (TSN 173598)
French Common Names: salamandre de Jefferson
Unique Identifier: ELEMENT_GLOBAL.2.100401
Element Code: AAAAA01050
Informal Taxonomy: Animals, Vertebrates - Amphibians - Salamanders
 
Kingdom Phylum Class Order Family Genus
Animalia Craniata Amphibia Caudata Ambystomatidae Ambystoma
Genus Size: D - Medium to large genus (21+ species)
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Concept Reference
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Concept Reference: Frost, D. R. 1985. Amphibian species of the world. A taxonomic and geographical reference. Allen Press, Inc., and The Association of Systematics Collections, Lawrence, Kansas. v + 732 pp.
Concept Reference Code: B85FRO01HQUS
Name Used in Concept Reference: Ambystoma jeffersonianum
Taxonomic Comments: Ambystoma jeffersonianum and A. laterale comprise a complex that includes pure bisexual diploid populations and populations that may be represented by bisexual diploid individuals, female hybrids with several different combinations of parental chromosomes (including diploids, triploids, tetraploids, and pentaploids), and/or rare male hybrids. Up to at least five different chromosomal combinations have been found at a single site.

In response to the nomenclatural difficulties presented by this complex, an informal system of indicating ploidy and chromosomal makeup has been recommended (Bogart and Klemens 1997). For example, the "LLLJ" designation (or "A. (3) laterale-jeffersonianum") indicates a tetraploid salamander with three sets of laterale chromosomes and one set of jeffersonianum chromosomes. For convenience, all populations including pure laterale (LL) and laterale-dominated genomes (e.g., LLJ) might be included in the species A. laterale (e.g., Klemens 1993).

However, biological complexities confound such a simple treatment. Polyploid hybrid individuals in the complex may produce offspring with diverse chromosomal makeup. For example, an LLJ female can produce offspring that do not have laterale-dominated genomes (e.g., LJJ). And single clutches have been documented with both diploid and triploid embryos. It is even possible for a triploid hybrid to produce pure diploid offspring (e.g., LL or JJ).

Truly intermediate hybrids with equal chromosomal representation (i.e., LJ, LLJJ) are uncommon and are but two of many possible outcomes of hybridization. In New England and New York, hybrids are more common than are the bisexual species, but bisexuals usually (or are presumed to) exist in low numbers in populations composed mostly of hybrids (Bogart and Klemens 1997). Thus, for example, a pond with triploid LLJ should also have at least some LL.

Hybrid populations are maintained by breeding between a hybrid female and a male of one of the diploid bisexual species, and the hybrid offspring are the result of gynogenetic or hybridogenetic reproduction (Bogart and Klemens 1997). In gynogenetic reproduction, male sperm stimulates egg development, but the male genome is not incorporated into the zygote. In hybridogenetic reproduction, the male genome is incorporated, "but upon maturity of the offspring, the paternal genome is eliminated in a meiotic or pre-meiotic event while the rest of the hybrid's genome is passed on to future generations, usually in an altered state" (Bogart and Klemens 1997).

Thus, recent work on hybrid phenomena in these salamanders indicates that the species names "A. platineum" (= LJJ) or "A. tremblayi" (= LLJ) formerly used for the A. jeffersonianum-A. laterale hybrids are inappropriate because LJJ and LLJ are not strictly gynogenetic triploid lineages. Consequently, a recent checklist (Crother et al. 2000) did not recognize either A. platineum or A. tremblayi as valid species.

To accommodate the genetic variation in these salamanders (and the often uncertain genetic composition of particular populations), we have established for this database the following elements: (1) Ambystoma jeffersonianum (Jefferson Salamander), used for pure populations only; (2) Ambystoma laterale (Blue-spotted salamander), used for pure populations only; (3) Ambystoma hybrid pop. 1 (jeffersonianum x laterale; jeffersonianum genome dominates), used for hybrid populations dominated by the jeffersonianum genome (e.g., populations that include mainly LJJ individuals); this element includes "Ambystoma platineum"; (4) Ambystoma hybrid pop. 2 (jeffersonianum x laterale; laterale genome dominates), used for hybrid populations dominated by the laterale genome (e.g., populations that include mainly LLJ individuals); this element includes "Ambystoma tremblayi"; (5) Ambystoma pop. 3 (jeffersonianum/laterale complex - uncertain composition), used for populations for which chromosomal representation is unknown. If needed we could establish additional elements for hybrid populations that involve A. laterale, A. texanum, and/or A. tigrinum.

See Kraus (1985), Bogart and Licht (1987), Bogart et al. (1987), Kraus et al. (1991), Lowcock et al. (1991), and Bogart and Klemens (1997) for information on the involvement of A. jeffersonianum in hybridization with A. texanum, A. tigrinum, and/or A. laterale. See Lowcock et al. (1987) and Bogart and Klemens (1997) for discussions of nomenclatural treatment of hybrid populations.

See Kraus (1988), Shaffer et al. (1991), and Jones et al. (1993) for phylogenetic analyses of North American Ambystoma.
Conservation Status
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NatureServe Status

Global Status: G4
Global Status Last Reviewed: 28Dec2001
Global Status Last Changed: 18Sep2001
Rounded Global Status: G4 - Apparently Secure
Reasons: The Ambystoma jeffersonianum genome is widespread in eastern North America, but it exists mainly in hybrids; the range of pure populations not well documented by apparently is restricted relative to the distribution of the genome.
Nation: United States
National Status: N4 (18Sep2001)
Nation: Canada
National Status: N2 (19Dec2011)

U.S. & Canada State/Province Status
United States Connecticut (S3), Illinois (S2), Indiana (S4), Kentucky (S4), Maryland (S3), Massachusetts (S2S3), New Hampshire (S2S3), New Jersey (S3), New York (S4), Ohio (SNR), Pennsylvania (S3S4), Vermont (S2), Virginia (S4), West Virginia (S2)
Canada Ontario (S2)

Other Statuses

Canadian Species at Risk Act (SARA) Schedule 1/Annexe 1 Status: T (05Jun2003)
Committee on the Status of Endangered Wildlife in Canada (COSEWIC): Endangered (26Nov2010)
Comments on COSEWIC: This salamander has a restricted range within populated and highly modified areas. Over the past three generations, the species has disappeared from many historic locations and the remaining locations are threatened by development, loss of habitat and, potentially, the presence of sperm-stealing unisexual populations of salamanders.
Designated Threatened in November 2000. Status re-examined and designated Endangered in November 2010.

IUCN Red List Category: LC - Least concern

NatureServe Global Conservation Status Factors

Range Extent: 20,000-2,500,000 square km (about 8000-1,000,000 square miles)
Range Extent Comments: Conant and Collins (1991) mapped the range as encompassing the area from southeastern New York through Pennsylvania and eastern and southern Ohio to southern Indiana, and southward to southcentral Kentucky and northern Virginia, with an extensive area of hybridization with A. LATERALE northward of this range to eastern Minnesota, northern Wisconsin, Upper Peninsula of Michigan, southern Ontario, southern Quebec, and eastward to Nova Scotia. However, for most of this range, karyological and electrophoretic data are unavailable, so the precise range of pure JEFFERSONIANUM populations is uncertain (Bogart and Klemens 1997). The core of the range of pure A. JEFFERSONIANUM populations likely extends from Pennsylvania southwestward to Kentucky. The JEFFERSONIANUM genome is widely distributed in eastern North America but exists primarily in hybrids (Bogart and Klemens 1997). Individuals that have solely the A. JEFFERSONIANUM genome occur in many hybridized populations.
Although Klemens (1993) mapped distinct ranges for A. JEFFERSONIANUM and A. LATERALE in Connecticut and adjacent regions, he included in the range of each species populations that were dominated by the pertinent genome, including hybrids. Data presented by Bogart and Klemens (1997) indicate that the few populations in New England and New York represented by only the A. JEFFERSONIANUM genome had sample sizes of only 1-3 individuals, so these actually may have been hybrid populations. Phillips (1991) extended the range of A. JEFFERSONIANUM into eastcentral Illinois, based on one juvenile raised from a larva, but since only one specimen was examined (and he did not indicate what identification criteria were used), it is unclear whether the population represents pure A. JEFFERSONIANUM or a hybrid population. Phillips et al. (1999) indicated the occurrence of both pure A. JEFFERSONIANUM and hybridized A. JEFFERSONIANUM ("A. PLATINEUM") in eastcentral Illinois, and they stated that the hybrids use A. TEXANUM sperm to activate egg development. In northern New Jersey, Nyman et al. (1988) found that triploid hybrids apparently occur wherever A. JEFFERSONIANUM is found. In Indiana and Ohio, JEFFERSONIANUM genomes exist in hybridized individuals that also contain A. TEXANUM and/or A. TIGRINUM genomes (Morris 1985, Morris and Brandon 1984,Selander et al. 1993, Selander 1994).

Area of Occupancy: Unknown 4-km2 grid cells
Area of Occupancy Comments:  

Number of Occurrences: 81 to >300
Number of Occurrences Comments: Number of occurrences of pure A. jeffersonianum is uncertain, but there are many.

Population Size: 10,000 - 1,000,000 individuals
Population Size Comments: Coded value for total adult population size is a guess that pertains only to unhybridized populations. In an extensive survey of New England and New York, Bogart and Klemens (1997) did not find any "sizable" populations. The only populations represented by only the jeffersonianum genome were a few from which only 1-3 specimens were obtained; these may have yielded hybrid individuals if they had been adequately sampled. In Kentucky, where most populations might (but not certainly) represent pure A. jeffersonianum, the species appears to be uncommon, based on the poorly known distribution (Barbour 1971). Similarly, in West Virginia, another area likely represent by pure populations (though this is uncertain), the species is believed to occur statewide but has been documented only from a minority (14) of the state's counties (Green and Pauley 1987), suggesting that A. jeffersonianum is uncommon there.

Number of Occurrences with Good Viability/Integrity: Unknown

Overall Threat Impact: Medium
Overall Threat Impact Comments: Vulnerable to detrimental alteration of vernal pool breeding sites, especially as a result of residential development. Threats to local populations likely include intensive timber harvesting practices that reduce canopy closure, understory vegetation, uncompacted forest litter, or coarse woody debris (moderately to well-decayed) in areas surrounding breeding sites (deMaynadier and Hunter 1999). Some local populations incur heavy road mortality during migrations to and from breeding sites (Klemens 1993). Increased acid deposition is a potential threat.

Short-term Trend: Decline of <30% to relatively stable

Long-term Trend: Decline of <50% to Relatively Stable

Intrinsic Vulnerability: Moderately vulnerable

Environmental Specificity: Narrow. Specialist or community with key requirements common.
Environmental Specificity Comments: Breeding habitat (vernal pools) is moderately to highly specific but common.

Other NatureServe Conservation Status Information

Protection Needs: Conservation goals need to be developed with respect to pure and hybridized populations; should they be assessed separately?

Distribution
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Global Range: (20,000-2,500,000 square km (about 8000-1,000,000 square miles)) Conant and Collins (1991) mapped the range as encompassing the area from southeastern New York through Pennsylvania and eastern and southern Ohio to southern Indiana, and southward to southcentral Kentucky and northern Virginia, with an extensive area of hybridization with A. LATERALE northward of this range to eastern Minnesota, northern Wisconsin, Upper Peninsula of Michigan, southern Ontario, southern Quebec, and eastward to Nova Scotia. However, for most of this range, karyological and electrophoretic data are unavailable, so the precise range of pure JEFFERSONIANUM populations is uncertain (Bogart and Klemens 1997). The core of the range of pure A. JEFFERSONIANUM populations likely extends from Pennsylvania southwestward to Kentucky. The JEFFERSONIANUM genome is widely distributed in eastern North America but exists primarily in hybrids (Bogart and Klemens 1997). Individuals that have solely the A. JEFFERSONIANUM genome occur in many hybridized populations.
Although Klemens (1993) mapped distinct ranges for A. JEFFERSONIANUM and A. LATERALE in Connecticut and adjacent regions, he included in the range of each species populations that were dominated by the pertinent genome, including hybrids. Data presented by Bogart and Klemens (1997) indicate that the few populations in New England and New York represented by only the A. JEFFERSONIANUM genome had sample sizes of only 1-3 individuals, so these actually may have been hybrid populations. Phillips (1991) extended the range of A. JEFFERSONIANUM into eastcentral Illinois, based on one juvenile raised from a larva, but since only one specimen was examined (and he did not indicate what identification criteria were used), it is unclear whether the population represents pure A. JEFFERSONIANUM or a hybrid population. Phillips et al. (1999) indicated the occurrence of both pure A. JEFFERSONIANUM and hybridized A. JEFFERSONIANUM ("A. PLATINEUM") in eastcentral Illinois, and they stated that the hybrids use A. TEXANUM sperm to activate egg development. In northern New Jersey, Nyman et al. (1988) found that triploid hybrids apparently occur wherever A. JEFFERSONIANUM is found. In Indiana and Ohio, JEFFERSONIANUM genomes exist in hybridized individuals that also contain A. TEXANUM and/or A. TIGRINUM genomes (Morris 1985, Morris and Brandon 1984,Selander et al. 1993, Selander 1994).

U.S. States and Canadian Provinces
Color legend for Distribution Map
Endemism: occurs (regularly, as a native taxon) in multiple nations

U.S. & Canada State/Province Distribution
United States CT, IL, IN, KY, MA, MD, NH, NJ, NY, OH, PA, VA, VT, WV
Canada ON

Range Map
Note: Range depicted for New World only. The scale of the maps may cause narrow coastal ranges or ranges on small islands not to appear. Not all vagrant or small disjunct occurrences are depicted. For migratory birds, some individuals occur outside of the passage migrant range depicted. For information on how to obtain shapefiles of species ranges see our Species Mapping pages at www.natureserve.org/conservation-tools/data-maps-tools.

Range Map Compilers: IUCN, Conservation International, NatureServe, and collaborators, 2004


U.S. Distribution by County Help
State County Name (FIPS Code)
CT Fairfield (09001), Hartford (09003), Litchfield (09005), Middlesex (09007), New Haven (09009)
IL Clark (17023), Edgar (17045)
MA Berkshire (25003), Bristol (25005)*, Essex (25009)*, Franklin (25011), Hampden (25013), Hampshire (25015), Middlesex (25017)*
NH Cheshire (33005), Sullivan (33019)
NJ Sussex (34037), Warren (34041)
VT Addison (50001), Bennington (50003), Chittenden (50007), Franklin (50011), Grand Isle (50013), Orange (50017), Rutland (50021), Washington (50023), Windham (50025), Windsor (50027)
WV Berkeley (54003), Cabell (54011), Fayette (54019), Greenbrier (54025)*, Hampshire (54027), Hardy (54031)*, Jefferson (54037)*, McDowell (54047)*, Mercer (54055), Mineral (54057)*, Monroe (54063), Pendleton (54071)*, Pocahontas (54075)*, Raleigh (54081), Randolph (54083), Roane (54087), Summers (54089), Wayne (54099)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
01 Upper Connecticut-Mascoma (01080104)+, White (01080105)+, Black-Ottauquechee (01080106)+, West (01080107)+, Middle Connecticut (01080201)+, Deerfield (01080203)+, Lower Connecticut (01080205)+, Westfield (01080206)+, Farmington (01080207)+, Quinnipiac (01100004)+, Housatonic (01100005)+, Saugatuck (01100006)+
02 Hudson-Hoosic (02020003)+, Middle Hudson (02020006)+, Rondout (02020007)+, Lower Hudson (02030101)+*, Middle Delaware-Mongaup-Brodhead (02040104)+, Middle Delaware-Musconetcong (02040105)+, South Branch Potomac (02070001)+, North Branch Potomac (02070002)+*, Conococheague-Opequon (02070004)+
04 Mettawee River (04150401)+, Otter Creek (04150402)+, Winooski River (04150403)+, Lake Champlain (04150408)+
05 Little Kanawha (05030203)+, Middle New (05050002)+, Greenbrier (05050003)+*, Lower New (05050004)+, Elk (05050007)+, Tug (05070201)+*, Raccoon-Symmes (05090101)+, Twelvepole (05090102)+, Middle Wabash-Busseron (05120111)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Reproduction Comments: Eggs are laid February-April (early to mid-March in Ohio). Clutch of a couple hundred eggs is divided among several clusters. Eggs hatch in about 2 weeks (Green and Pauley 1987) or up to 4-6 weeks. Larvae metamorphose during July-September. In a pond in Ohio, hatching rate over four years was 85-90%, and 51,800 to 84,400 larvae hatched (Brodman, 1995, J. Herpetol. 29:111-113). Aggregates when breeding.

Hybrid offspring are the result of gynogenetic or hybridogenetic reproduction (Bogart and Klemens 1997).

Non-Migrant: N
Locally Migrant: Y
Long Distance Migrant: N
Mobility and Migration Comments: In Kentucky, adults migrated an average of about 250 m from breeding pools to summer range; newly metamorphosed individuals apparently migrated lesser distances (average <100 m in one study) (Douglas and Monroe 1981). In Vermont, 6 radio-tagged individuals migrated 30-205 m (mean 93 m) from their breeding pool; likely a radius of about 175 m around a breeding pool would be needed to encompass 95% of a population (Faccio 2003).
Palustrine Habitat(s): FORESTED WETLAND, Riparian, SCRUB-SHRUB WETLAND, TEMPORARY POOL
Terrestrial Habitat(s): Forest - Hardwood, Forest - Mixed
Special Habitat Factors: Benthic, Burrowing in or using soil, Fallen log/debris
Habitat Comments: Often in well-shaded deciduous forest; hides in rodent burrows and beneath leaf litter, logs, stumps, and other surface objects; hibernates underground or in rotting logs (Faccio 2003). In caves in some areas (e.g., West Virginia) (Green and Pauley 1987). Eggs are attached to sticks and plant stems in ponds and pools with adjacent forest. Presence of fishes and newts reduces reproductive success. Not found in floodplains, swamps or marshes in New Jersey (Nyman et al. 1988). In central Pennsylvania, embryonic mortality was high in ponds below pH 4.5, though this was affected by the availability of other larval amphibians as prey (Sadinski and Dunson 1992).
Adult Food Habits: Carnivore, Invertivore
Immature Food Habits: Carnivore, Invertivore
Food Comments: Adults eat insects, worms, spiders, and other invertebrates, obtained mostly on land. Larvae eat small aquatic invertebrates and smaller amphibian larvae.
Adult Phenology: Hibernates/aestivates, Nocturnal
Immature Phenology: Hibernates/aestivates, Nocturnal
Phenology Comments: Inactive late fall to early winter. Diurnal activity may occur in dark cloudy weather.
Colonial Breeder: Y
Length: 21 centimeters
Economic Attributes Not yet assessed
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Management Summary Not yet assessed
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Population/Occurrence Delineation
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Group Name: Ambystomatid Salamanders

Use Class: Not applicable
Minimum Criteria for an Occurrence: Occurrences are based on evidence of historical presence, or current and likely recurring presence, at a given location. Such evidence minimally includes collection or reliable observation and documentation of one or more individuals (including larvae or eggs) in or near appropriate habitat where the species is presumed to be established and breeding.
Separation Barriers: Heavily traveled road, especially at night during salamander breeding season, such that salamanders almost never successfully traverse the road; road with a barrier that is impermeable to salamanders; wide, fast rivers; areas of intensive development dominated by buildings and pavement.
Separation Distance for Unsuitable Habitat: 1 km
Separation Distance for Suitable Habitat: 3 km
Separation Justification: BARRIERS/UNSUITABLE HABITAT: Rivers may or may not be effective barriers, depending on stream width and hydrodynamics; identification of streams as barriers is a subjective determination. Bodies of water dominated by predatory fishes have been described as barriers but probably should be regarded as unsuitable habitat. For A. barbouri, a stream-pool breeder, predatory fishes appeared to act as a barrier to larval dispersal and gene flow for populations separated by as little as 500-1000 m (Storfer 1999). Highly disturbed land, such as the cleared and bedded soils of some silvicultural site preparation, may serve as an impediment to movement of A. cingulatum (Means et al. 1996), although Ashton (1998) noted the species' use of pine plantations, pastures, and three-year-old clearcuts. Such areas should be treated as unsuitable habitat rather than barriers.

MOVEMENTS: Palis's (1997b) suggested use of 3.2 km between breeding sites to distinguish breeding populations of A. cingulatum was based on Ashton's (1992) finding that individuals may move as much as 1.6 km from their breeding ponds. Ambystoma californiense sometimes migrates up to 2 km between breeding ponds and terrestrial habitat (see USFWS 2004). Funk and Dunlap (1999) found that A. macrodactylum managed to recolonize lakes after trout extirpation despite evidence of low levels of interpopulation dispersal. Based on a review of several Ambystoma species (e.g., Semlitsch 1981, Douglas and Monroe 1981, Kleeberger and Werner 1983, Madison 1997), Semlitsch (1998) concluded that a radius of less than 200 meters around a breeding pond would likely encompass the terrestrial habitat used by more than 95 percent of adults. Faccio's (2003) study of radio-tagged A. maculatum and A. jeffersonianum in Vermont supports this conclusion. In New York, all movements of A. tigrinum occurred in areas within 300 m of the nearest breeding pond (Madison and Farrand 1998). However, most studies of these salamanders had small sample sizes and/or were not designed to detect long-distance movements, so migration distance may be somewhat underestimated.

In summary, ambystomatid salamanders generally stay within a few hundred meters of their breeding pool. Due to high breeding site fidelity and limitation of breeding to pool basins, populations using different breeding sites exhibit little or no interbreeding among adults. Thus one might argue that each pool constitutes a separate occurrence or that the separation distance for suitable habitat should be the nominal minimum of 1 km. However, little is known about how frequently first-time (or experienced) breeders use non-natal pools (pools from which they did not originate) or how far they may move to such sites. Frequent colonization of new and remote habitats by at least some species suggests that dispersal movements sometimes may be longer than typical adult migration distances. It seems unlikely that locations separated by a gap of less than a few kilometers of suitable habitat would represent independent occurrences over the long term.

Inferred Minimum Extent of Habitat Use (when actual extent is unknown): .3 km
Inferred Minimum Extent Justification: Inferred extent distance pertains to breeding sites (with the center of the circle in the center of the breeding site). Most ambystomatids stay within a few hundred meters of their breeding pool (see separation justification section).
Date: 10Sep2004
Author: Hammerson, G.
Population/Occurrence Viability
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U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
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Authors/Contributors
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NatureServe Conservation Status Factors Edition Date: 02May2011
NatureServe Conservation Status Factors Author: Hammerson, G.
Element Ecology & Life History Edition Date: 22Oct2003
Element Ecology & Life History Author(s): Hammerson, G.

Zoological data developed by NatureServe and its network of natural heritage programs (see Local Programs) and other contributors and cooperators (see Sources).

References
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  • Bogart, J.P. and M.W. Klemens. 1997. Hybrids and genetic interactions of mole salamanders (Ambystoma jeffersonianum and A. laterale) (Amphibia: Caudata) in New York and New England. American Museum Novitates (3218):1-78.

  • Bogart, J.P., Lowcock, L.A., Zeyl, C.W. and Mable, B.K. 1987. Genome constitution and reproductive biology of hybrid salamanders, genus Ambystoma, on Kelleys Island in Lake Erie. Canadian Journal of Zoology. 65:2188-2201.

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  • Faccio, S. D. 2003. Postbreeding emigration and habitat use by Jefferson and spotted salamanders in Vermont. Journal of Herpetology 37:479-489.

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  • Lowcock, L. A., L. E. Licht, and J. P. Bogart. 1987. Nomenclature in hybrid complexes of Ambystoma (Urodela: Ambystomatidae): no case for the erection of hybrid "species." Syst. Zool. 36:328-336.

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  • Morris, M. A. 1985. A hybrid AMBYSTOMA PLATINEUM x A. TIGRINUM from Indiana. Herpetologica 41:267-271.

  • Morris, M. A., and R. A. Brandon. 1984. Gynogenesis and hybridization between Ambystoma platineum and Ambystoma texanum in Illinois. Copeia 1984:324-337.

  • Nyman, S., M. J. Ryan, and J. D. Anderson. 1988. The distribution of the AMBYSTOMA JEFFERSONIANUM complex in New Jersey. Journal of Herpetology 22:224-228.

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  • Sadinski, W. J., and W. A. Dunson. 1992. A multilevel study of effects of low pH on amphibians of temporary ponds. J. Herpetol. 26:413-422.

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