Overview
Comprehensive Description
Description
- Schaaf, R. T., Jr., and Smith, P. W. (1963). ''Rana palustris (LeConte). Pickerel Frog.'' Catalogue of American Amphibians and Reptiles. American Society of Ichthyologists and Herpetologists, 117.1-117.3.
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Distribution
Range Description
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Geographic Range
The Pickerel frog ranges from the Canadian Maritime Provinces south to the Carolinas and then west to southeast Minnesota and eastern Texas. However, there are many gaps in the distribution of these frogs, especially in the southern parts of Ohio, Illinois, and Indiana (Conant and Collins 1998).
Biogeographic Regions: nearctic (Native )
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National Distribution
Canada
Origin: Native
Regularity: Regularly occurring
Currently: Present
Confidence: Confident
Type of Residency: Year-round
United States
Origin: Native
Regularity: Regularly occurring
Currently: Present
Confidence: Confident
Type of Residency: Year-round
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Global Range: (20,000-2,500,000 square km (about 8000-1,000,000 square miles)) Range extends from the Gaspe Peninsula of Quebec to Wisconsin, and south to southern South Carolina, northern Georgia, southern Mississippi, and southeastern Texas (Conant and Collins 1991). This frog is absent from most of the far southeastern United States (e.g., Florida, southern Georgia) and the prairie region of Illinois and vicinity.
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Distribution and Habitat
North of the coastal plain boundary, it occurs in habitats where the water is cool and clear. On the Coastal Plain, it occurs in floodplain swamp habitat, while in karst topography, it is often confined to the vicinity of the cave mouth.
- Schaaf, R. T., Jr., and Smith, P. W. (1963). ''Rana palustris (LeConte). Pickerel Frog.'' Catalogue of American Amphibians and Reptiles. American Society of Ichthyologists and Herpetologists, 117.1-117.3.
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Physical Description
Morphology
Physical Description
The Pickerel frog is a relatively large frog that is often confused with the Northern Leopard Frog (Rana pipiens). However, the Pickerel frog has chocolate-brown spots arranged in two rows between the dorsolateral folds while the Leopard frog's spots are more irregular and scattered. They can be distinguished by the bright yellow or yellow-orange color on the inside concealed surface of the thigh. Leopard frogs are white in the same area. These frogs range in size from 45 to 75 millimeters as adults. Females are usually larger than males. Male Pickerel frogs have paired vocal sacs, stout forearms and swollen thumbs. These frogs produce toxic skin secretions that are irritating to humans but can be fatal to other small animals, especially other amphibians. Many frog-eating snakes avoid these frogs for this reason (Matson 1999).
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Size
Type Information
Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1933
Locality: Lake Waccamaw, Columbus, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1933
Locality: Lake Waccamaw, Columbus, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1948
Locality: Maxton, Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1948
Locality: Maxton, Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1948
Locality: Maxton, Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Locality: Kinston, Lenoir, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1913
Locality: Edenton, Chowan, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1913
Locality: Edenton, Chowan, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1913
Locality: Edenton, Chowan, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1913
Locality: Edenton, Chowan, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1913
Locality: Edenton, Chowan, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1913
Locality: Edenton, Chowan, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1913
Locality: Edenton, Chowan, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1913
Locality: Edenton, Chowan, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1913
Locality: Edenton, Chowan, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1913
Locality: Edenton, Chowan, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1913
Locality: Edenton, Chowan, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1913
Locality: Edenton, Chowan, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1913
Locality: Edenton, Chowan, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1913
Locality: Edenton, Chowan, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1961
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1961
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1961
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1961
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1961
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1961
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1961
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1961
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1961
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1961
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1961
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Holotype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1961
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1961
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1961
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1961
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1963
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Year Collected: 1963
Locality: Maxton, Hayes Pond (= Maxton Pond), Robeson, North Carolina, United States, North America
- Paratype: Hardy, J. D. 1964. Chesapeake Science. 5 (1-2): 91, figures 2, 4 and 5.
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Ecology
Habitat
Habitat and Ecology
Systems
- Terrestrial
- Freshwater
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Habitat
Pickerel frogs commonly inhabit cool, wooded streams, seeps and springs although they are also found in many other habitats. In the South, it can also be found in the relatively warm, turbid waters of the Coastal Plain and floodplain swamps. These frogs tend to wander far into grassy fields or into weed-covered areas in the summer (Conant and Collins 1998).
Terrestrial Biomes: forest
Aquatic Biomes: rivers and streams
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Comments: Pickerel frogs occur in various freshwater aquatic and wetland habitats in wooded regions, ranging from the vicinity of cool clear streams and ponds in the north to warm, turbid swamps in parts of the south. In summer, they commonly range into fields and woods away from ponds or streams. In winter, they may be hidden at the bottom of a water body or secluded in wet caves.Breeding sites include standing water of woodland ponds, bog ponds, impoundments, stream pools, sloughs, and flooded ditches, often but not always in sites with few or no fishes (e.g., Holomuzki 1995). Males call while floating or submerged. Females attach egg masses to submerged vegetation.
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Migration
Non-Migrant: Yes. At least some populations of this species do not make significant seasonal migrations. Juvenile dispersal is not considered a migration.
Locally Migrant: No. No populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).
Locally Migrant: No. No populations of this species make annual migrations of over 200 km.
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Trophic Strategy
Food Habits
These frogs are carnivorous and their diet consists mostly of small insects and other invertebrates. However, as tadpoles, these frogs are herbivorous (Conant and Collins 1998).
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Comments: Metamorphosed frogs eat various small invertebrates, terrestrial and aquatic. Larvae probably eat mostly suspended matter, algae, plant tissue, organic debris, and minute organisms in water.
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Population Biology
Number of Occurrences
Note: For many non-migratory species, occurrences are roughly equivalent to populations.
Estimated Number of Occurrences: > 300
Comments: Represented by many and/or large occurrences throughout most of the range.
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Global Abundance
100,000 to >1,000,000 individuals
Comments: Total adult population size is unknown but likely exceeds 100,000.
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General Ecology
Pickerel frogs have a distinctive odor and bright colors that are thought to advertise their distasteful skin secretions, and some predators such as shrews, certain fishes, and common garter snakes may reject them as food. Larvae are palatable to some fishes (green sunfish), less palatable to others (longear sunfish) (Holomuzki 1995).
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Life History and Behavior
Cyclicity
Comments: n the northern part of the range, pickerel frogs are mostly inactive during the coldest winter months.
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Reproduction
Reproduction
Pickerel frogs breed in late March to early May. Males have low, snore-like calls to attract females. After fertilization, females lay spherical egg masses attached to tree branches in permanent or temporary ponds. These masses may contain from 700 to 3000 eggs. Each egg has an average diameter of 1.6 millimeters when laid. After the eggs hatch, it takes around 87 to 95 days for the tadpoles to transform into small frogs and leave the water. It requires an additional two years before these frogs reach sexual maturity and are able to reproduce (Matson 1999).
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Breeding occurs in winter in the far south, in spring (mostly April-May) in the north. Larvae hatch in several days or up to three weeks after laying, metamorphose usually within 2-3 months, in summer (often July-August in the north).
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Molecular Biology and Genetics
Molecular Biology
Statistics of barcoding coverage: Rana palustris
Public Records: 0
Species: 11
Species With Barcodes: 1
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Conservation
Conservation Status
IUCN Red List Assessment
Red List Category
Red List Criteria
Version
Year Assessed
Assessor/s
Reviewer/s
Justification
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Conservation Status
The population of Pickerel frogs is listed as stable, and there are no special restrictions on them. However, in many areas populations are declining due to habitat changes.
US Federal List: no special status
CITES: no special status
State of Michigan List: no special status
IUCN Red List of Threatened Species: least concern
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National NatureServe Conservation Status
Canada
Rounded National Status Rank: N5 - Secure
United States
Rounded National Status Rank: N5 - Secure
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NatureServe Conservation Status
Rounded Global Status Rank: G5 - Secure
Intrinsic Vulnerability: Moderately vulnerable
Environmental Specificity: Moderate to broad.
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Trends
Population
Population Trend
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Global Short Term Trend: Relatively stable (=10% change)
Comments: Population trend is unknown but probably stable to slightly declining.
Global Long Term Trend: Increase of 10-25% to decline of 30%
Comments: Likely relatively stable in extent of occurrence, likely less than 25% decline in in population size, area of occurrence, and number/condition of occurrences. In some areas, habitat losses have been offset by habitat augmentation (e.g., creation of breeding sites through damming).
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Life History, Abundance, Activity, and Special Behaviors
- Schaaf, R. T., Jr., and Smith, P. W. (1963). ''Rana palustris (LeConte). Pickerel Frog.'' Catalogue of American Amphibians and Reptiles. American Society of Ichthyologists and Herpetologists, 117.1-117.3.
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Threats
Threats
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Management
Conservation Actions
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Relevance to Humans and Ecosystems
Benefits
Economic Importance for Humans: Negative
This species does not seem to adversely affect humans at all.
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Economic Importance for Humans: Positive
These frogs are not of great economic importance to humans. They are not caught as game and are not kept as pets due to their skin secretions. They are occasionally used as fishing bait for anglers.
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Wikipedia
Pickerel Frog
The Pickerel Frog (Rana palustris)[1][2][3] is a small North American frog, characterized by the appearance of seemingly "hand-drawn" squares on its dorsal surface.
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Distinguishing features
The distinctive rectangular spots of the Pickerel Frog may blend together to form a long rectangle along the back. All leopard frogs have circular spots. In addition, Pickerel Frogs have prominent dorsolateral ridges that are unbroken. Another important distinguishing mark is the orange or yellow flash pattern found on the inner surface of the hind legs of Pickerel Frogs. The frog must be picked up to examine this, as the legs cover the coloration otherwise. The Plains Leopard Frog (Lithobates blairi) exhibits this coloration as well, but the dorsolateral ridges are interrupted and inset medially in that species.
Range
The Pickerel Frog ranges in the west from much of Wisconsin, southeast Minnesota, eastern Iowa, through Missouri and down to eastern Texas. To the east they extend through northern Louisiana, most of Mississippi, northern Alabama, Georgia (U.S. state), and South Carolina to the coast. Their northern range extends into Canada in the southern reaches of Ontario, Quebec, New Brunswick, and Nova Scotia. The range is spotty through the midwestern states and a field guide should be obtained for the specifics on ranges in a particular area.
Other
The skin secretions of a stressed Pickerel Frog are known to be toxic to other frogs, as many a novice frog catcher has found when he finds only the Pickerel Frog still alive in his bucket. These secretions can also be moderately irritating if they come in contact with the eyes, mucous membranes, or broken skin. It is advisable to wash one's hands after handling Pickerel Frogs.
Footnotes
References
- Arnold, K. 2000. "Rana palustris" (On-line), Animal Diversity Web. Accessed September 25, 2007 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Rana_palustris.html.
- Hillis, D.M. & Wilcox, T.P. (2005): Phylogeny of the New World true frogs (Rana). Mol. Phylogenet. Evol. 34(2): 299–314. doi:10.1016/j.ympev.2004.10.007 PMID 15619443 PDF fulltext.
- Hillis, D. M. (2007) Constraints in naming parts of the Tree of Life. Mol. Phylogenet. Evol. 42: 331–338.
- Hammerson (2004). Rana palustris. 2006. IUCN Red List of Threatened Species. IUCN 2006. www.iucnredlist.org. Retrieved on 09 May 2006. Database entry includes a range map and justification for why this species is of least concern
- Pauly, Greg B., Hillis, David M. & Cannatella, David C. (2009): Taxonomic freedom and the role of official lists of species names. Herpetologica 65: 115-128. PDF fulltext
- Redmer, M. and Mierzwa, K.S. (1994). "A review of the distribution and zoogeography of the pickerel frog, Rana palustris, in northern Illinois". Bulletin of the Chicago Herpetological Society 29: 21–30.
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