Hyla andersonii - Baird, 1854
Pine Barrens Treefrog
Other English Common Names: Pine Barrens treefrog
Taxonomic Status: Accepted
Related ITIS Name(s): Hyla andersonii Baird, 1854 (TSN 173509)
Unique Identifier: ELEMENT_GLOBAL.2.102258
Element Code: AAABC02010
Informal Taxonomy: Animals, Vertebrates - Amphibians - Frogs and Toads
 
Kingdom Phylum Class Order Family Genus
Animalia Craniata Amphibia Anura Hylidae Hyla
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: Hyla andersonii
Taxonomic Comments: Duellman et al. (2016) removed this species from the genus Hyla and included it (and all other U.S./Canada species of Hyla, as well as additional Hyla species in Mexico, Guatemala, and eastern Asia) in the genus Dryophytes (previously recognized as a subgenus).
Conservation Status
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NatureServe Status

Global Status: G4
Global Status Last Reviewed: 18Feb2014
Global Status Last Changed: 18Oct1996
Rounded Global Status: G4 - Apparently Secure
Reasons: Occurs in three widely disjunct areas: New Jersey, the Carolinas, and Florida-Alabama; many occurrences are protected, though most include small numbers of individuals; some habitats need active management to remain suitable; apparently stable and secure overall.
Nation: United States
National Status: N4 (05Nov1996)

U.S. & Canada State/Province Status
United States Alabama (S2), Florida (S3), New Jersey (S2), North Carolina (S3), South Carolina (S2S3)

Other Statuses

IUCN Red List Category: NT - Near threatened

NatureServe Global Conservation Status Factors

Range Extent: 5000-200,000 square km (about 2000-80,000 square miles)
Range Extent Comments: Pine Barrens of New Jersey; upper Coastal Plain and parts of lower Coastal Plain of North and South Carolina; western Florida panhandle and adjacent Alabama, some 750 km southwest of the nearest South Carolina population; known in Georgia from an old record of a single specimen (Means and Mohler 1979, Gosner and Black 1967, Conant and Collins 1991). The largest populations occur in New Jersey (Freda and Morin 1984). Discovery of this species in Florida was fairly recent (Christman 1970), but the population there is large (map in Krysko et al. 2011). Palmer (1977) suggested that the current distribution reflects relicts from a considerably more widespread distribution in the past.

Number of Occurrences:  
Number of Occurrences Comments: Represented by many and/or large occurrences throughout most of the small range.

Population Size: 10,000 - 1,000,000 individuals
Population Size Comments: Total adult population size is unknown but likely exceeds 10,000.

Number of Occurrences with Good Viability/Integrity: Few (4-12)
Viability/Integrity Comments: See Number of Protected EOs Comments.

Overall Threat Impact: Medium
Overall Threat Impact Comments: Apparently secure in most of the range, although relative scarcity and specialized habitat requirements justify continued monitoring and protection. The primary threat in the New Jersey Pine Barrens is habitat destruction or alteration from residential, agricultural, and industrial development (Palmer 1977, Freda and Morin 1984). Development pressures within the Pine Barrens place isolated populations outside preserves at increasing risk of elimination. The early successional shrub bogs, seeps, and sphagnum ponds selected as breeding sites are very acidic and nutrient-poor ecosystems, and any changes in the chemistry of the waters in these habitats (as, for example, from stormwater runoff) would likely cause the disappearance of the characteristic flora and fauna (Ehrenfeld 1983, Morgan et al. 1983, Freda and Morin 1984). The sandy soils of the Pine Barrens are very porous and allow pollutants to quickly enter the ground water, which is the major water source for the wetlands upon which the treefrog depends. Development can also lower the water table, which would have dramatic effects on the hydrology of bog wetlands. Garton and Sill (1979) reported that the specific habitat requirements of the species made it susceptible to local extirpation. Unlike other sympatric treefrog species, HYLA ANDERSONII generally does not breed in temporary water bodies such as natural rain pools or in human-made areas such as roadside ditches and borrow pits. However, Bullard (1965) reported chorusing males along a roadside ditch in North Carolina. As is true for other Sandhills species, plant succession due to fire suppression appears to be a significant threat in South Carolina (Cely and Sorrow 1986).

Short-term Trend: Relatively Stable (<=10% change)
Short-term Trend Comments: Relatively stable overall.

Long-term Trend: Decline of 30-50%
Long-term Trend Comments: Likely relatively stable in extent of occurrence; unknown degree of decline in population size, area of occurrence, and number/condition of occurrences.

Intrinsic Vulnerability: Moderately vulnerable

Environmental Specificity: Narrow. Specialist or community with key requirements common.

Other NatureServe Conservation Status Information

Inventory Needs: As of the mid-1990s, on a range-wide scale, most areas had been adequately surveyed.

Protection Needs: Protect existing populations from habitat destruction/modification. Palmer (1977) suggested that populations might easily be preserved in public areas in North Carolina, including Fort Bragg Military Reservation, Bladen Lakes State Forest (Bladen County), and Sandhills Wildlife Management Area (Moore, Richmond, and Scotland Counties).

Distribution
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Global Range: (5000-200,000 square km (about 2000-80,000 square miles)) Pine Barrens of New Jersey; upper Coastal Plain and parts of lower Coastal Plain of North and South Carolina; western Florida panhandle and adjacent Alabama, some 750 km southwest of the nearest South Carolina population; known in Georgia from an old record of a single specimen (Means and Mohler 1979, Gosner and Black 1967, Conant and Collins 1991). The largest populations occur in New Jersey (Freda and Morin 1984). Discovery of this species in Florida was fairly recent (Christman 1970), but the population there is large (map in Krysko et al. 2011). Palmer (1977) suggested that the current distribution reflects relicts from a considerably more widespread distribution in the past.

U.S. States and Canadian Provinces
Color legend for Distribution Map
Endemism: endemic to a single nation

U.S. & Canada State/Province Distribution
United States AL, FL, NC, NJ, SC

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)
AL Covington (01039), Escambia (01053), Geneva (01061)
FL Holmes (12059), Okaloosa (12091), Santa Rosa (12113), Walton (12131)
NC Bladen (37017), Cumberland (37051), Harnett (37085), Hoke (37093), Johnston (37101)*, Jones (37103), Lee (37105)*, Moore (37125), Onslow (37133), Richmond (37153), Sampson (37163), Scotland (37165), Wayne (37191)*
NJ Atlantic (34001), Burlington (34005), Camden (34007), Cape May (34009), Cumberland (34011), Gloucester (34015), Middlesex (34023), Monmouth (34025), Ocean (34029), Salem (34033)
SC Chesterfield (45025), Kershaw (45055), Marlboro (45069), Richland (45079)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
02 Sandy Hook-Staten Island (02030104)+, Raritan (02030105)+, Crosswicks-Neshaminy (02040201)+, Lower Delaware (02040202)+, Cohansey-Maurice (02040206)+, Mullica-Toms (02040301)+, Great Egg Harbor (02040302)+
03 Upper Neuse (03020201)+*, Middle Neuse (03020202)+*, White Oak River (03020301)+, New River (03020302)+*, Upper Cape Fear (03030004)+, Lower Cape Fear (03030005)+, Black (03030006)+, Northeast Cape Fear (03030007)+, Lower Pee Dee (03040201)+, Lynches (03040202)+, Lumber (03040203)+, Little Pee Dee (03040204)+, Waccamaw (03040206)+*, Wateree (03050104)+*, Lower Broad (03050106)+, Choctawhatchee Bay (03140102)+, Yellow (03140103)+, Blackwater (03140104)+, Pensacola Bay (03140105)+, Pea (03140202)+, Lower Choctawhatchee (03140203)+, Lower Conecuh (03140304)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: A treefrog that reaches a maximum snout-vent length of about 5 cm.
General Description: Dorsum and upper surfaces of legs are pea green to dark olive green . A prominent chocolate-brown to plum-colored or lavender stripe extends along the sides from the nostril to the hind limb; this stripe is bordered dorsally by a narrow white stripe (lemon-yellow in Florida). The venter is white with a light gray chin in the male. Discrete, bright yellow-orange spots and blotches mark the axillae, the inside surfaces of the hindlegs, and the lower sides of the body; these brightly colored areas are concealed when the frog is at rest. The partly webbed toes have expanded terminal disks. Adult snout-vent length usually is 2.9 to 4.4 cm (maximum 5.1 cm) (Conant 1975, Means and Longden 1976). Florida frogs tend to be larger than frogs in New Jersey and the Carolinas and differ in the pitch of the breeding call (Means and Longden 1976).

Descriptions of eggs, larvae, and larval development in New Jersey were provided by Noble and Noble (1923), Gosner and Black (1957), and Gosner (1960). Means and Longden (1976) provided a detailed description of larvae in Florida.

Reproduction Comments: The breeding season in North Carolina appears to be protracted, with choruses heard from April through August (Palmer 1977). Vocalization occurs between mid-April and early September in South Carolina, being most consistent from May through July. Choruses begin at dusk, with periodic choruses of short duration occurring after dark (Garton and Sill 1979, Cely and Sorrow 1986). Locations are generally represented by only a few calling males (Means and Longden 1976, Garton and Sill 1979, Moler 1981, Tardell et al. 1981, Cely and Sorrow 1986).

In New Jersey, males called nightly beginning in May and early June. In July, males called only infrequently and only after rainstorms. Nightly calling ceased by August. Egg laying in New Jersey occurs from late April to mid-July. (Morin et al. 1990). It is uncertain as to whether late clutches represent late breeding subpopulation or are additional clutches deposited by adults that bred earlier (Morin et al. 1990).

In Florida, males called from April to September and larvae were found from mid-June to early September (Means and Longden 1976). Calling activity was strongly correlated with humidity and thundershower activity and no males were heard calling on dry nights, especially after several dry days. Amplexed pairs were observed in mid-June and mid-July. Larvae were found in the field from mid-June through August, but probably occur earlier in May and later into September.

Females oviposit 800 to 900 eggs (50-200 according to Freda and Morin [1984]). Means and Longden (1976) found that a clutch of 206 eggs from one female hatched in 72 h at 25 C. In the field, larvae metamorphose by the end of summer.

Ecology Comments: Individuals are rarely seen. They are disruptively colored, which makes them almost invisible when perched in green vegetation, and call sporadically, which makes them very difficult to locate.

Freda and Morin (1984) used radio-isotope tagging to track eight individual treefrogs for periods ranging from 4 to 30 days. Four of the marked frogs moved more than 50 m from the breeding pond and one frog spent five days more than 100 m from the pond. Frogs were also capable of moving great distances in a short period of time. One tagged frog moved 102 m in a single day. In New Jersey, average estimated daily movements of adult frogs ranged between 0 and 20 m, with longer range movements ranging from 21 to 48 m (Freda and Gonzalez 1986). Of the eight frogs tagged, seven remained within 70 m of the breeding pool near which they were initially captured. Daily movements were not correlated with temperature or daily rainfall.

Habitat Type: Terrestrial
Non-Migrant: N
Locally Migrant: Y
Long Distance Migrant: N
Mobility and Migration Comments: Migrates between breeding pools and adjacent nonbreeding terrestrial habitats. In New Jersey, Freda and Morin (1984) and Freda and Gonzalez (1986) demonstrated that treefrogs often travel distances of 100 m from breeding ponds during the nonbreeding season.
Riverine Habitat(s): Low gradient, Pool
Palustrine Habitat(s): Bog/fen, FORESTED WETLAND, HERBACEOUS WETLAND, Riparian, SCRUB-SHRUB WETLAND
Terrestrial Habitat(s): Woodland - Mixed
Special Habitat Factors: Benthic, Fallen log/debris, Standing snag/hollow tree
Habitat Comments: This species is restricted to localized wetlands such as hillside seepage bogs within dry uplands, pine barrens, and headwater swamps and disperses along drainages within these areas (Cely and Sorrow 1983; Means and Longden 1976; P.Moler, pers. comm., 1998). Nonbreeding habitat generally is in pine-oak areas adjacent to breeding habitat. Activity is terrestrial and arboreal. Important egg-laying and larval habitats include open cedar swamps and sphagnaceous, shrubby, acidic, seepage bogs on hillsides below pine-oak ridges. Intolerant of closed-canopy conditions.

In the Southeast, typical habitat is characterized by the topography, soils, and vegetation of the Carolina Sandhills, with pocosin or evergreen shrub swamps established along seeps and small streams within the surrounding longleaf pine-oak forest (Noble and Noble 1923, Wright and Wright 1949, Gosner and Black 1957, Bullard 1965, Means and Moler 1978, Tardell et al. 1981).

In North Carolina, occurrences are almost exclusively in or near pocosins or shrub bogs with dense growth of pines, bays, and various ericaceous shrubs (Palmer 1977).

Garton and Sill (1979) described breeding habitat in South Carolina as low vegetation with dense growth of SPHAGNUM mosses, similar to those described by Means and Longden (1976) in Florida. The largest population (some 50 calling males) in the Carolina Sandhills National Wildlife Refuge was found in a sphagnaceous seepage slope comprising about 5 acres.

Cely and Sorrow (1986) found that occurrences in South Carolina appeared to be restricted to the Fall Line Sandhills at elevations ranging between 61 and 122 m. The general aspect of 18 inhabited localities was evergreen shrub-herb bog found along small black water tributaries, sometimes upstream from extensive hardwood bottomland forest.

HYLA ANDERSONII is often encountered in disturbed sites associated with utility rights-of-way and recent clearcuts (Tardell et al. 1981, Cely and Sorrow 1986).

During a four-year survey in the South Carolina Sandhills, Cely and Sorrow (1986) found calling males in atypical habitats, including temporary habitats created by exceptionally heavy rainfall, but persistent populations (i.e., sites occupied for at least two years) were generally associated with permanent open water of large stream drainages.

Breeding has been reported in small pools, streams, and ditches (Gerhardt 1974). Eggs sink individually to the bottom (Noble and Noble 1923). Breeding habitats are typically characterized by seepage areas, or small streams, with dense SPHAGNUM cover. Herbaceous or shrubby areas are preferred, but densely forested areas do not support the species (Means and Longden 1976, Garton and Sill 1979).

Freda and Gonzalez (1986) described breeding season habitat in their New Jersey Pine Barrens study site as a small seepage stream, dammed by a sand road, and choked with vegetation (SPHAGNUM spp. and UTRICULARIA spp.).

Breeding habitat in the panhandle of Florida is characterized by acid hillside seepage bogs with copious SPHAGNUM mosses and dense woody vegetation (dominantly CLIFTONIA MONOPHYLLA and CYRILLA RACEMIFLORA) (Means and Longden 1976).

Breeding ponds within the wetland matrix described by Freda and Morin (1984) were typically small (5 to 10 m diameter), sphagnum-filled depressions. Zampella (1994) pointed out that treefrogs were typically found in small, ephemeral ponds, rather than in larger water bodies.

Adult Food Habits: Invertivore
Immature Food Habits: Herbivore
Food Comments: Metamorphosed frogs eat mainly insects and other small invertebrates (Palmer 1977). Bullard (1965:155) described captive frogs consuming flies and crickets, often attacking crickets "almost as large as themselves." Larvae eat organic debris, algae, and plant tissue.
Adult Phenology: Hibernates/aestivates, Nocturnal
Immature Phenology: Hibernates/aestivates, Nocturnal
Phenology Comments: Probably hibernates in north; habits poorly known. Calling occurs most vigorously on warm rainy nights (Palmer 1977).
Colonial Breeder: Y
Length: 5 centimeters
Economic Attributes Not yet assessed
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Management Summary
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Stewardship Overview: HYLA ANDERSONII appears to be relatively secure in most of its disjunct range, although its relative scarcity and specialized habitat requirements justify continued monitoring and protection of the species. Considerable knowledge exists regarding habitat requirements during the breeding season when calling males and amplexed pairs are typically found in small, sometimes ephemeral, ponds within a matrix of shrub and herbaceous bog or pocosin. Successful establishment of breeding colonies appears to depend on an array of specific habitat parameters, including acid pH and low nutrient content in breeding ponds, which are threatened by fire suppression and development of surrounding areas. Data from New Jersey indicate that post-breeding males may move up to 100 m from breeding ponds, so that efforts to protect critical habitat must include protection of suitable buffer zones in upland areas to accommodate these movements. The extreme sensitivity of water quality in breeding habitats makes monitoring of water chemistry important. Use of prescribed fire in shrub and herbaceous seepage bogs and pocosins should be further researched. Successional processes in fire-suppressed systems rapidly alter the sphagnaceous, open-water breeding areas essential to the treefrog's survival.
Restoration Potential: Means and Moler (1978) and Tardell et al. (1981) suggested that the presence of HYLA ANDERSONII in disturbed situations may indicate that the species can readily colonize new areas of suitable habitat and that existing areas may be managed to maintain suitable open shrubby plant communities through judicious use of prescribed burning and selective logging and clearing.
Preserve Selection & Design Considerations: Preserves should include breeding ponds and a surrounding buffer area to accommodate post-breeding movements. A buffer zone of 100 m would include the typical range of dispersing adults and would provide important protection for the sensitive water chemistry of seepage bogs (Freda and Morin 1984).
Management Requirements: The typical shrub bog that supports breeding colonies is an early successional, subclimax community dependent on disturbances such as fire to maintain the appropriate species composition and vegetation structure (Means and Mohler 1979, Freda and Morin 1984). Periodic fire was probably the dominant force in pre-settlement times that created and maintained suitable habitats (Means and Moler 1979). Cely and Sorrow (1986) believed that active fire suppression in the Carolina Sandhills has largely eliminated natural creation of suitable habitat. In New Jersey, the greatest threat to the treefrog on protected lands is vegetation succession due to fire suppression. See Means and Moler (1979) for information on the use of fire in maintaining or creating suitable early successional habitats.

Also of critical importance is the characteristic water chemistry of these wetland systems, particularly a very low pH (Wright and Wright 1949, Gosner and Black 1957, Means and Longden 1976, Means and Mohler 1979, Cely and Sorrow 1982, Freda and Morin 1984) and low nutrient content (Freda and Morin 1984). Protection and management must concentrate on maintaining community integrity as well as water quality within critical habitats.

Monitoring Requirements: Existing populations should be monitored regularly to track trends in numbers and the effects of habitat alterations. As sensitive components of fragile ecosystems, these frogs may indicate environmental quality for an array of associated species.

The most effective survey method for treefrogs appears to be counts of calling males. Calling tends to be affected by ambient conditions, but may be influenced more by local hydrology of seepage bogs than by ambient weather conditions (Moler 1981, Cely and Sorrow 1986). Means and Longden (1976) pointed out that calling, stimulated by high humidities and rainfall, can extend until September, despite the absence of gravid females. Also, locations are generally represented by only a few calling males (Means and Longden 1976, Garton and Sill 1979, Moler 1981, Tardell et al. 1981, Cely and Sorrow 1986). Cely and Sorrow (1986) pointed out that the calling of males can be erratic and revisited sites at least three times to increase the probability of finding the species. Surveys were conducted from late March through mid-September to capture the entire potential breeding season. Males are readily stimulated to call by recorded vocalizations, but synchronous calling can make discrimination of individuals difficult (Means and Longden 1986).

The New Jersey Pinelands Commission monitors some 100 sites on a monthly basis (Zampella 1994). Each site is visited during April to June on warm, humid nights and the numbers of calling males is estimated by the same two person crew, to avoid observer biases. Calling males are recorded in three increments: 2-5, 5-10, and >10. Each site is visited at least three times during the breeding season. In addition, reference ponds have been established with known populations that are sampled at the beginning and end of each night to reduce error due to ambient temperature and weather conditions.

Freda and Morin (1984) and Freda and Gonzalez (1986) presented the results of a short-term (one summer) study of movements using 51-Cr radioactive tags. They were able to follow frogs throughout the breeding and nonbreeding season.

Cely and Sorrow (1986) described the use of soils survey maps and USGS topographic maps to delineate potential habitat in the South Carolina Sandhills.

Management Research Needs: Freda and Morin (1984) suggested that further research was needed on post-breeding movement of females, as well as on the home ranges of males and females during the late summer and fall. Dispersal of newly metamorphosed froglets is also poorly understood. Zampella (1994) stressed that more information is needed on the functional hydrology of ephemeral wetlands used for breeding. Management regimes for shrub bog wetlands, including prescribed fire and woody vegetation control, should be researched (Freda and Morin 1984).
Biological Research Needs: Almost nothing is known of the postlarval development, survival rates, or adult nonbreeding ecology.
Population/Occurrence Delineation
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Group Name: Hylid Frogs (Treefrogs)

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: Busy major highway such that frogs rarely if ever cross successfully; intensive urban development dominated by buildings and pavement and lacking suitable vegetated frog refuges.
Separation Distance for Unsuitable Habitat: 1 km
Separation Distance for Suitable Habitat: 5 km
Separation Justification: Available information is limited but indicates that hylids generally exhibit limited movements on a short-term basis. In New Jersey, Freda and Morin (1984) and Freda and Gonzalez (1986) demonstrated that individual Hyla andersonii often travel distances of 100 m from breeding ponds during the nonbreeding season. In montane Colorado, Spencer (1964) found that Pseudacris triseriata range into wet meadows usually within about 700 m of their breeding sites and sometimes cross a few hundred meters of upland habitat. Kay (1989) determined that most Pseudacris cadaverina individuals range over small segments of streamcourse; 83 percent of movements were less than 25 m in a 1-year study. In Michigan, nonbreeding home range diameters of Pseudacris crucifer, established around forest debris and vegetation, ranged from 1.2 to 5.5 m (Delzell 1958).

Based on this information it appears that 1 km is an appropriate separation distance for unsuitable habitat. Despite limited data suggesting restricted movements, dispersal data are scant, and these frogs are clearly physically capable of long moves. It seems unlikely that occupied locations separated by a gap of less than several kilometers of suitable habitat would represent independent occurrences over the long term.

Inferred Minimum Extent of Habitat Use (when actual extent is unknown): .5 km
Inferred Minimum Extent Justification: Inferred extent distance pertains to distance from breeding sites.
Date: 21Sep2004
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: 18Feb2014
NatureServe Conservation Status Factors Author: Jackson, D. R. (2014); Niese, J., R. Jordan, and G. Hammerson (2002)
Management Information Edition Date: 12Sep1995
Management Information Edition Author: JORDAN, ROBERT A.
Element Ecology & Life History Edition Date: 07Feb2002
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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  • Alabama Department of Conservation and Natural Resources, Division of Wildlife and Freshwater Fisheries. 2005. Conserving Alabama's wildlife: a comprehensive strategy. Alabama Department of Conservation and Natural Resources, Division of Wildlife and Freshwater Fisheries. Montgomery, Alabama. 303 pages. [Available online at http://www.dcnr.state.al.us/research-mgmt/cwcs/outline.cfm ]

  • Baird, S. F. 1854. Description of new genera and species of North American frogs. Proceedings of the Academy of Natural Sci. Phila. 7:59-62.

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  • Bullard, A. J. 1965. Additional records of the treefrog HYLA ANDERSONII from the Coastal Plain of North Carolina. Herpetologica. 21:154-5.

  • Cely, J. E., and J. A. Sorrow, Jr. 1983. Distribution, status and habitat of the pine barrens treefrog in South Carolina. Final report, South Carolina Wildlife and Marine Resources Department, Columbia, South Carolina. 55 pp.

  • Cely, J. E., and J. A. Sorrow, Jr. 1986. Distribution and habitat of HYLA ANDERSONII in South Carolina. J. Herpetol. 20:102-104.

  • Christman, S. P. 1970. HYLA ANDERSONII in Florida. Quarterly Journal of the Florida Academy of Science. 33:80.

  • Conant, R. 1975. A Field Guide to Reptiles and Amphibians of Eastern and Central North America. Second Edition. Houghton Mifflin Company, Boston, Massachusetts. xvii + 429 pp.

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  • Duellman, W. E., A. B. Marion, and S. B. Hedges. 2016. Phylogenetics, classification, and biogeography of the treefrogs (Amphibia: Anura: Arboranae). Zootaxa 4104: 1?109.

  • Ehrenfeld, J. G. 1983. The effects of changes in land-use on swamps of the New Jersey Pine Barrens. Biol. Cons. 25:353-75.

  • Freda, J., and P. J. Morin. 1984. Adult home range of the pine barrens treefrog (Hyla andersonii) and the physical, chemical, and ecological characteristics of its preferred breeding ponds. Final Report to New Jersey Department of Environmental Protection, Division of Fish, Game, and Wildlife, Endangered Nongame Species Program, Trenton, NJ. 33 pp.

  • Freda, J., and R. J. Gonzalez. 1986. Daily movements of the treefrog, Hyla andersoni. Journal of Herpetolology 20:469-471.

  • 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.

  • Garton, J. S. and B. L. Sill. 1979. The status of the pine barrens treefrog, HYLA ANDERSONII Baird, in South Carolina. Pp. 131-2 in D. M. Forsythe and W. B. Ezell Jr. (eds.). Proceedings of the First South Carolina Endangered Species Symposium, South Carolina Wildlife and Marine Res. Department, Columbia, SC.

  • Gerhardt, H. C. 1974. Behavioral isolation of the treefrogs Hyla cinerea and Hyla andersonii. American Midland Naturalist 91:424-433.

  • Godwin, J. C. 1995. Extrinsic factors influencing the breeding of the pine barrens treefrog, HYLA ANDERSONII. Unpublished M.S. thesis, Auburn University, Alabama. 82 pp.

  • Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica. 16:183-90.

  • Gosner, K. L. and I. H. Black. 1956. Notes on amphibians from the upper coastal plain of North Carolina. J. Elisha Mitchell Sci. Soc. 72:40-7.

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  • Gosner, K.L. and Black, I.H. 1967. Hyla andersonii. Catalogue of American Amphibians and Reptiles. 54:1-2.

  • Hulmes, D., P. Hulmes, and R. Zappalorti. 1981. Notes on the ecology and distribution of the pine barrens treefrog, Hyla andersonii, in New Jersey. Bull. New York Herpetol. Soc. 17(1):2-19.

  • Jensen, J. B. 1991. The distribution of the pine barrens treefrog, HYLA ANDERSONII, in Conecuh National Forest, Alabama. Unpublished report.

  • Krysko, K. L., K. M. Enge, and P. E. Moler. 2011. Atlas of amphibians and reptiles in Florida. Final report to Florida Fish and Wildlife Conservation Commission, Tallahassee, Florida. Submitted 15 December 2011.

  • Martof, B. S., W. M. Palmer, J. R. Bailey, and J. R. Harrison, III. 1980. Amphibians and reptiles of the Carolinas and Virginia. University of North Carolina Press, Chapel Hill, North Carolina. 264 pp.

  • Means, D. B. 1983. The enigmatic pine barrens treefrog. Florida Wildlife 37:16-19.

  • Means, D. B. and C. J. Longden. 1976. Aspects of the biology and zoogeography of the pine barrens treefrog (HYLA ANDERSONII) in northern Florida. Herpetologica. 32:117-30.

  • Means, D. B., and P. E. Moler. 1979. The pine barrens treefrog: fire, seepage bogs, and management implications. Pages 77-83 in R. R. Odum and L. Landers, eds. Proceedings.. Georgia Game & Fish Div. Tech. Bull. WL4, Atlanta.

  • Mirarchi, R. E., M. A. Bailey, T. M. Haggerty, and T. L. Best, editors. 2004. Alabama wildlife. Volume 3. Imperiled amphibians, reptiles, birds, and mammals. The University of Alabama Press, Tuscaloosa, Alabama. 225 pages.

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