Overview

Brief Summary

Slim green or brownish, with well-defined, pale-bordered, oval or round dark spots on back, white to cream below. White stripe on upper jaw. Well-defined, pale dorsolateral folds that are continuous and not angled inward. Voice is a low 'motorboat' or snore-like sound interspersed with grunting and chuckling, lasting about 1-5 seconds. Choruses are a medley of moaning, grunting, and chuckling that suggests the sounds made by rubbing a well-inflated rubber balloon. Paired vocal sacs expand over the forelimbs" (Stebbins 1985). There is usually one spot on the head anterior to the eyes. Few or no tubercules on the dorsal and lateral body surface. Mean SVL in males is 68.3 mm (2.7 in) and in females 74.2 mm (2.92 in). The eardrum is without a light center. During breeding season the males have a swollen, darkened thumb base and loose skin between the jaw and the shoulder. Males are usually smaller in size. The tadpole has coarse indistinct mottling on the tail. The distal half of the tail tends to darken approaching metamorphosis.

Color variations include the Burnsi variant, which may be found in either brown or green and does not have any dorsal spots. It has spots or bars on the limbs and may have black stippling on the back and sides. The second variant Kandiyohi, is brown with dashes of white and brown or black. The spots on the back and legs are still discernable, as well as the dorsolateral fold (LeClere).

Found in a variety of habitats, most cold-adapted of all leopard frogs. May forage far from water, when frightened seeks water in a zigzag pattern of jumps. Like most frogs, leopard frogs are sluggish animals, often staying immobile for long periods of time. Sometimes the males call while underwater. They produce a low-pitched snore often followed by a chuckling noise, or a deep urr, urr, urr. They have internal vocal sacs, so their throats do not appear to move when they call. When they move far from a body of water they may absorb dew to keep moist. Hibernates in deep water. Juvenile leopard frogs often cluster together.

Consumes small invertebrates; rarely eats small vertebrates. Larvae eat algae, plant tissue, organic debris, and probably small invertebrates (TNC 1988).

  • Arizona Game and Fish Department. 2002. Lithobates pipiens. Unpublished abstract compiled and edited by the Heritage Data Management System, Arizona Game and Fish Department, Phoenix, AZ
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Comprehensive Description

Description

Rana pipiens is a slim green or brownish frog, varying in length between about 5 and 11 cm. On its back it has dark rounds spots with pale borders. Its underside is white or cream colored. Spotting may be reduced or absent on young. The species has well-defined, continuous dorsolateral folds, which are not angled inward. The upper jaw has a white stripe. The male has a swollen, darkened thumb base and loose skin between jaw and shoulder during breeding season. Tadpoles are dark brown or olive to gray on top, often with gold spots. Their undersides are cream to whitish, light enough that intestines often show through (Stebbins 2003).

See another account at californiaherps.com.

  • Stebbins, R. C. (1985). A Field Guide to Western Reptiles and Amphibians. Houghton Mifflin, Boston.
  • Stebbins, R. C. (2003). Western Reptiles and Amphibians, Third Edition. Houghton Mifflin, Boston.
  • Fellers, G. J., Green, D. E., and Longcore, J. E. (2001). ''Oral chytridiomycosis in the Mountain Yellow-Legged Frog (Rana muscosa).'' Copeia, 2001(4), 945-953.
  • Brodkin, M., Vatcnick, I., Simon, M., Hollyann, H., Butler-Holston, K., and Leonard, M. (2003). ''Effects of acid stress in adult Rana pipiens.'' Journal of Experimental Zoology, 298A(1), 16-22.
  • Gibbs, J. P. (2000). ''Wetland loss and biodiversity conservation.'' Conservation Biology, 14(1), 314-317.
  • Glennemeier, K. A. and Denver, R. J. (2001). ''Sublethal effects of chronic exposure to an organochlorine compound on northern leopard frog (Rana pipiens) tadpoles.'' Environmental Toxicology, 16(4), 287-297.
  • Hayes, T., Haston, K., Tsui, M., Hoang, A., Haeffele, C., and Vonk, A. (2002). ''The feminization of male frogs in the wild.'' Nature, 419, 895-896.
  • Hecnar, S. J. (1995). "Acute and chronic toxicity of ammonium nitrate fertilizer to amphibians from southern Ontario." Environmental Toxicology and Chemistry, 14(12), 2131-2137.
  • Lannoo, M. J., Lang, K., Waltz, T., and Phillips, G. S. (1994). "An altered amphibian assemblage: Dickinson County, Iowa, 70 years after Frank Blanchard's survey." American Midland Naturalist, 131(2), 311-319.
  • Linck, M. (2000). ''Reduction in road mortality in a northern leopard frog population.'' Journal of the Iowa Academy of Science, 107(3-4), 209-211.
  • Peterson, G., Johnson, L. B., Axler, R. P., and Diamond, S. A. (2002). ''Assessment of the risk of solar ultraviolet radiation to amphibians. II. In situ characterization of exposure in amphibian habitats.'' Environmental Science and Technology, 36(13), 2859-2865.
  • Schothoeffer, A. M., and Koehler, A. V., Meteyer, C. U., Cole, R. A. (2003). ''Influence of Ribeiroia ondatrae (Trematoda: Digenea) infection on limb development and survival of northern leopard frogs (Rana pipiens): Effects of host stage and parasite-exposure level.'' Canadian Journal of Zoology, 81(7), 1144-1153.
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Distribution

Global Range: (>2,500,000 square km (greater than 1,000,000 square miles)) Range extends from the Great Slave Lake, Hudson Bay, and Labrador, Canada, south to southern New England, Kentucky, Nebraska, New Mexico, and Arizona, west to southeastern British Columbia, eastern Washington, eastern Oregon, and eastern California (Conant and Collins 1991, Stebbins 2003). Distribution is spotty in the west, where this frog has been introduced in many localities.

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Range Description

This species is known from Great Slave Lake and Hudson Bay, Canada, south to Kentucky and New Mexico, USA (Stebbins 1985, Conant and Collins 1991). It has a spotty distribution in the west, where it has been introduced in many localities. It is also known from Panama where it is endemic to the central cordillera and western Pacific lowlands, although this is most likely an undescribed species (see taxonomic note). It occurs at approximately 100-600m asl in the eastern portion of the Panamanian distribution.
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occurs (regularly, as a native taxon) in multiple nations

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Geographic Range

Leopard frogs are native to the Nearctic region. They are found throughout much of North America, from as far north as the Hudson Bay, along the eastern seaboard to northern Virginia and west to British Columbia, eastern Washington, and Oregon. The western part of the range extends as far south as New Mexico, Arizona, Colorado, Utah, and portions of California and Nevada. Populations in the west are fragmented and some are declining.

Biogeographic Regions: nearctic (Native )

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Distribution and Habitat

Rana pipiens is a wide-ranging species. It can be found across a broad swath of territory from the Hudson Bay south to northern Virginia, and to the west as far as southern British Columbia and eastern Washington and Oregon. In the west its range extends south through Utah, Colorado, New Mexico, and Arizona. It exists in scattered populations in some areas of California and Nevada, and has been introduced in other areas of California. In the western U.S. its range is now greatly fragmented and lacks confirmation in recent years. (Stebbins 2003).

R. pipiens lives in a wide variety of habitats: grassland, brushland, and forest. It is the most cold-adapted of all the leopard frogs, and can be found up to an elevation of about 11,000 feet (Stebbins 2003). It can also be found in agricultural lands and in developed areas such as golf courses (Hayes et al. 2002). It prefers to live where there is a permanent body of standing or slowly flowing water, and among the aquatic vegetation of such places (Stebbins 2003).

  • Stebbins, R. C. (1985). A Field Guide to Western Reptiles and Amphibians. Houghton Mifflin, Boston.
  • Stebbins, R. C. (2003). Western Reptiles and Amphibians, Third Edition. Houghton Mifflin, Boston.
  • Fellers, G. J., Green, D. E., and Longcore, J. E. (2001). ''Oral chytridiomycosis in the Mountain Yellow-Legged Frog (Rana muscosa).'' Copeia, 2001(4), 945-953.
  • Brodkin, M., Vatcnick, I., Simon, M., Hollyann, H., Butler-Holston, K., and Leonard, M. (2003). ''Effects of acid stress in adult Rana pipiens.'' Journal of Experimental Zoology, 298A(1), 16-22.
  • Gibbs, J. P. (2000). ''Wetland loss and biodiversity conservation.'' Conservation Biology, 14(1), 314-317.
  • Glennemeier, K. A. and Denver, R. J. (2001). ''Sublethal effects of chronic exposure to an organochlorine compound on northern leopard frog (Rana pipiens) tadpoles.'' Environmental Toxicology, 16(4), 287-297.
  • Hayes, T., Haston, K., Tsui, M., Hoang, A., Haeffele, C., and Vonk, A. (2002). ''The feminization of male frogs in the wild.'' Nature, 419, 895-896.
  • Hecnar, S. J. (1995). "Acute and chronic toxicity of ammonium nitrate fertilizer to amphibians from southern Ontario." Environmental Toxicology and Chemistry, 14(12), 2131-2137.
  • Lannoo, M. J., Lang, K., Waltz, T., and Phillips, G. S. (1994). "An altered amphibian assemblage: Dickinson County, Iowa, 70 years after Frank Blanchard's survey." American Midland Naturalist, 131(2), 311-319.
  • Linck, M. (2000). ''Reduction in road mortality in a northern leopard frog population.'' Journal of the Iowa Academy of Science, 107(3-4), 209-211.
  • Peterson, G., Johnson, L. B., Axler, R. P., and Diamond, S. A. (2002). ''Assessment of the risk of solar ultraviolet radiation to amphibians. II. In situ characterization of exposure in amphibian habitats.'' Environmental Science and Technology, 36(13), 2859-2865.
  • Schothoeffer, A. M., and Koehler, A. V., Meteyer, C. U., Cole, R. A. (2003). ''Influence of Ribeiroia ondatrae (Trematoda: Digenea) infection on limb development and survival of northern leopard frogs (Rana pipiens): Effects of host stage and parasite-exposure level.'' Canadian Journal of Zoology, 81(7), 1144-1153.
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Geographic Range

Leopard frogs are found throughout much of North America, from as far north as the Hudson Bay, along the eastern seaboard to northern Virginia and west to British Columbia, eastern Washington, and Oregon. The western part of the range extends as far south as New Mexico, Arizona, Colorado, Utah, and portions of California and Nevada. Populations in the west are fragmented and some are declining.

Biogeographic Regions: nearctic (Native )

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Physical Description

Morphology

Physical Description

Leopard frogs are from 5 to 11.1 cm long. They are green or greenish-brown dorsally, with round, brown spots arranged on their back, sides, and legs. These spots usually have a whitish or yellow border. There is a distinct, white dorso-lateral fold along the length of the back extending from each eye. A white line runs on either side of the mouth, from the nose to the shoulder. The underside is white or greenish white.

As with most frogs, males are smaller than females. Males have thickened thumb pads and paired vocal sacs that inflate over their shoulders as they call.

Tadpoles are greenish or brown, with yellow or black speckles frequently visible. The belly is white and somewhat transparent, with the intestinal coils visible through the skin. Tadpoles reach a maximum size of 8.4 cm.

Range length: 5 to 11.1 cm.

Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry

Sexual Dimorphism: female larger

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Physical Description

Leopard frogs are from 5 to 11.1 cm long. They are green or greenish-brown dorsally, with round, brown spots arranged on their back, sides, and legs. These spots usually have a whitish or yellow border. There is a distinct, white dorso-lateral fold along the length of the back extending from each eye. A white line runs on either side of the mouth, from the nose to the shoulder. The underside is white or greenish white.

As with most frogs, males are smaller than females. Males have thickened thumb pads and paired vocal sacs that inflate over their shoulders as they call.

Tadpoles are greenish or brown, with yellow or black speckles frequently visible. The belly is white and somewhat transparent, with the intestinal coils visible through the skin. Tadpoles reach a maximum size of 8.4 cm.

Range length: 5 to 11.1 cm.

Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry

Sexual Dimorphism: female larger

Average mass: 29.0222 g.

Average basal metabolic rate: 0.00576 W.

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Size

Length: 13 cm

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Diagnostic Description

Differs from RANA PALUSTRIS in having rounded rather than squarish dorsal spots and in lacking yellow or orange pigment on the usually concealed surfaces of the hind limbs and groin. Differs from other leopard frogs as follows: RANA BLAIRI is never green, usually has a distinct pale spot on the eardrum, has the posterior end of the dorsolateral ridges inset or angled inward, and lacks vestigial oviducts in males. RANA CHIRICAHUENSIS has a "salt-and-pepper" pattern of small tubercles on the back of the thighs, and stockier proportions (Stebbins 1985). RANA ONCA is smaller, with shorter legs, the spotting toward the head often is reduced, and the underside of the hind limbs is yellow to yellow-orange (Stebbins 1985). RANA YAVAPAIENSIS is stockier and paler (Stebbins 1985). RANA BERLANDIERI is paler and has the dorsolateral ridges inset medially at the rear end.

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Type Information

Syntype for Lithobates pipiens
Catalog Number: USNM 3363
Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Preparation: Ethanol
Locality: Yellowstone River, Locality In Multiple Counties, Montana, United States, North America
  • Syntype: Cope, E. D. 1889. United States National Museum Bulletin. (34): 403, Figure 101.
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Ecology

Habitat

Montana Valley and Foothill Grasslands Habitat

This taxon can be found in the Montana valley and foothill grasslands ecoregions, along with some other North American ecoregions. This ecoregion occupies high valleys and foothill regions in the central Rocky Mountains of Montana in the USA and Alberta, Canada. The ecoregion the uppermost flatland reaches of the Missouri River drainage involving part of the Yellowstone River basin, and extends into the Clark Fork-Bitterroot drainage of the Columbia River system. The ecoregion, consisting of three chief disjunctive units, also extends marginally into a small portion of northern Wyoming. Having moderate vertebrate species richness, 321 different vertebrate taxa have been recorded here.

The dominant vegetation type of this ecoregion consists chiefly of wheatgrass (Agropyron spp.) and fescue (Festuca spp.). Certain valleys, notably the upper Madison, Ruby, and Red Rock drainages of southwestern Montana, are distinguished by extensive sagebrush (Artemisia spp.) communities as well. This is a reflection of semi-arid conditions caused by pronounced rain shadow effects and high elevation. Thus, near the Continental Divide in southwestern Montana, the ecoregion closely resembles the nearby Snake/Columbia shrub steppe.

A number of mammalian species are found in the ecoregion, including: American Pika (Ochotona princeps), a herbivore preferring talus habitat; Bighorn Sheep (Ovis canadensis), Black-tailed Prairie Dog (Cynomys ludovicianus), who live in underground towns that may occupy vast areas; Brown Bear (Ursos arctos); Hoary Marmot (Marmota caligata), a species who selects treeless meadows and talus as habitat; and the Northern River Otter (Lontra canadensis), a species that can tolerate fresh or brackish water and builds its den in the disused burrows of other animals.

There are six distinct anuran species that can be found in the Montana valleys and foothills grasslands, including: Canadian Toad (Anaxyrus hemiophrys); Western Toad (Anaxyrus boreas); Northern Leopard Frog (Lithobates pipiens); Plains Spadefoot Toad (Spea bombifrons); Columbia Spotted Frog (Rana luteiventris), an anuran that typically breeds in shallow quiet ponds; and the Boreal Chorus Frog (Pseudacris maculata).

Exactly two amphibian taxa occurr in the ecoregion: Long-toed Salamander (Ambystoma macrodactylum), a species who prefers lentic waters and spends most of its life hidden under bark or soil; Tiger Salamander (Ambystoma tigrinum).

Reptilian species within the ecoregion are: Milk Snake (Lampropeltis triangulum), an adaptable taxon that can be found on rocky slopes, prairie and near streambeds; Painted Turtle (Chrysemys picta); Western Plains Garter Snake (Thamnophis radix), a taxon that can hibernate in the burrows of rodents or crayfish or even hibernate underwater; Yellow-bellied Racer (Coluber constrictor); Spiny Softshell Turtle (Apalone spinifera); Western Terrestrial Garter Snake (Thamnophis elegans); Rubber Boa (Charina bottae); Western Skink (Plestiodon skiltonianus); and the Western Rattlesnake (Crotalis viridis).

The ecoregion supports endemic and relict fisheries: Westslope Cutthroat Trout (Oncorhynchus clarki lewisi), Yellowstone Cutthroat Trout (Oncorhynchus clarkii bouvieri), and fluvial Arctic Grayling (Thymallus arcticus), a relict species from past glaciation.

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Colorado Plateau Shrublands Habitat

This taxon can be found in the Colorado Plateau shrublands, as one of its North American ecoregions of occurrence. The Plateau is an elevated, northward-tilted saucer landform, characterized by its high elevation and arid to semi-arid climate. Known for the Grand Canyon, it exhibits dramatic topographic relief through the erosive action of high-gradient, swift-flowing rivers that have downcut and incised the plateau. Approximately 90 percent of the plateau is drained by the Colorado River and its tributaries, notably the lower catchment of the Green River.

A pinyon-juniper zone is extensive, dominated by a pygmy forest of Pinyon pine (Pinus edulis) and several species of juniper (Juniperus spp). Between the trees the ground is sparsely covered by grama, other grasses, herbs, and various shrubs, such as Big sagebrush (Artemisia tridentata) and Alder-leaf cercocarpus (Cercocarpus montanus).

A montane zone extends over large areas on the high plateaus and mountains, but is much smaller than the pinyon-juniper zone. The montane vegetation varies considerably, from Ponderosa pine in the south to Lodgepole pine and Aspen further north. Northern Arizona contains four distinct Douglas-fir habitat types. The lowest zone has arid grasslands but with many bare areas, as well as xeric shrubs and sagebrush. Several species of cacti and yucca are common at low elevations in the south.

Numerous mammalian species are found within the Colorado Plateau shrublands ecoregion, including the Black-tailed prairie dog (Cynomys ludovicianus); Long-eared chipmunk (Tamias quadrimaculatus); Utah prairie dog (Cynomys parvidens EN); Yellow-bellied marmot (Marmota flaviventris); and the Uinta chipmunk (Tamias umbrinus), a burrowing omnivore.

A large number of birds are seen in the ecoregion, with representative taxa: Chestnut-collared longspur (Calcarius ornatus NT); Greater sage grouse (Centrocercus urophasianus NT); Northern pygmy owl (Glaucidium gnoma); Cactus wren (Campylorhynchus brunneicapillus).

There are various snakes occurring within the Colorado Plateau, including: Black-necked garter snake (Thamnophis cyrtopsis), usually found in riparian zones; Plains Blackhead snake (Tantilla nigriceps); Black-tailed rattlesnake (Crotalus molossus), who seeks inactivity refuge in rock crevices, animal burrows and even woodrat houses. Other reptiles found here include the Common checkered whiptail (Cnemidophorus tesselatus).

There are only a limited number of anuran taxa on the Colorado Plateau; in fact, the comprehensive occcurrence list for the ecoregion is: Red-spotted toad (Anaxyrus punctatus); Canyon treefrog (Hyla arenicolor); Woodhouse's toad (Anaxyrus woodhousii); Couch's spadefoot toad (Scaphiopus couchii); Northern leopard frog (Lithobates pipiens); Plains spadefoot toad (Spea bombifrons); and Southwestern toad (Anaxyrus microscaphus). The Tiger salamander (Ambystoma tigrinum) is the sole salamander found on the Colorado Plateau shrublands.

The Colorado River fish fauna display distinctive adaptive radiations. The Humpback chub (Gila cypha), for example, is a highly specialized minnow that lives in the upper Colorado. It adapted to the water’s fast current and its extremes of temperature and flow rate. Dams and water diversion, however, have created a series of placid, stillwater lakes and side streams, and the Humpback chub may not be able to adapt to these altered conditions. The species, along with other native Colorado River fishes including the Bonytail (Gila elegans), Squawfish (Ptychocheilus lucius), and the Flannelmouth sucker (Catostomus latipinnis), may not survive much further in time.

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Palouse Grasslands Habitat

This taxon is found in the Palouse grasslands, among other North American ecoregions. The Palouse ecoregion extends over eastern Washington, northwestern Idaho and northeastern Oregon. Grasslands and savannas once covered extensive areas of the inter-mountain west, from southwest Canada into western Montana in the USA. Today, areas like the great Palouse prairie of eastern  are virtually eliminated as natural areas due to conversion to rangeland. The Palouse, formerly a vast expanse of native wheatgrasses (Agropyron spp), Idaho Fescue (Festuca idahoensis), and other grasses, has been mostly plowed and converted to wheat fields or is covered by Drooping Brome (Bromus tectorum) and other alien plant species.

the Palouse historically resembled the mixed-grass vegetation of the Central grasslands, except for the absence of short grasses. Such species as Bluebunch Wheatgrass (Elymus spicatus), Idaho Fescue (Festuca idahoensis) and Giant Wildrye (Elymus condensatus) and the associated species Lassen County Bluegrass (Poa limosa), Crested Hairgrass (Koeleria pyramidata), Bottlebrush Squirrel-tail (Sitanion hystrix), Needle-and-thread (Stipa comata) and Western Wheatgrass (Agropyron smithii) historically dominated the Palouse prairie grassland.

Representative mammals found in the Palouse grasslands include the Yellow-bellied Marmot (Marmota flaviventris), found burrowing in grasslands or beneath rocky scree; American Black Bear (Ursus americanus); American Pika (Ochotona princeps); Coast Mole (Scapanus orarius), who consumes chiefly earthworms and insects; Golden-mantled Ground Squirrel (Spermophilus lateralis); Gray Wolf (Canis lupus); Great Basin Pocket Mouse (Perognathus parvus); Northern River Otter (Lontra canadensis); the Near Threatened Washington Ground Squirrel (Spermophilus washingtoni), a taxon who prefers habitat with dense grass cover and deep soils; and the Northern Flying Squirrel (Glaucomys sabrinus), a mammal that can be either arboreal or fossorial.

There are not a large number of amphibians in this ecoregion. The species present are the Great Basin Spadefoot Toad (Spea intermontana), a fossorial toad that sometimes filches the burrows of small mammals; Long-toed Salamander (Ambystoma macrodactylum); Northern Leopard Frog (Glaucomys sabrinus), typically found near permanent water bodies or marsh; Columbia Spotted Frog (Rana luteiventris), usually found near permanent lotic water; Pacific Treefrog (Pseudacris regilla), who deposits eggs on submerged plant stems or the bottom of water bodies; Tiger Salamander (Ambystoma tigrinum), fossorial species found in burrows or under rocks; Woodhouse's Toad (Anaxyrus woodhousii), found in arid grasslands with deep friable soils; Western Toad (Anaxyrus boreas), who uses woody debris or submerged vegetation to protect its egg-masses.

There are a limited number of reptiles found in the Palouse grasslands, namely only: the Northern Alligator Lizard (Elgaria coerulea), often found in screes, rock outcrops as well as riparian vicinity; the Painted Turtle (Chrysemys picta), who prefers lentic freshwater habitat with a thick mud layer; Yellow-bellied Racer (Chrysemys picta); Ringneck Snake (Diadophis punctatus), often found under loose stones in this ecoregion; Pygmy Short-horned Lizard (Phrynosoma douglasii), a fossorial taxon often found in bunchgrass habitats; Side-blotched Lizard (Uta stansburiana), frequently found in sandy washes with scattered rocks; Southern Alligator Lizard (Elgaria multicarinata), an essentially terrestrial species that prefers riparian areas and other moist habitats; Pacific Pond Turtle (Emys marmorata), a species that usually overwinters in upland habitat; Western Rattlesnake (Crotalus viridis), who, when inactive, may hide under rocks or in animal burrows; Night Snake (Hypsiglena torquata); Western Skink (Plestiodon skiltonianus), who prefers grasslands with rocky areas; Western Terrestrial Garter Snake (Thamnophis elegans), found in rocky grasslands, especially near water; Rubber Boa (Charina bottae).

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Arizona Mountains Forests Habitat

This taxon is found in the Arizona Mountain Forests, which extend from the Kaibab Plateau in northern Arizona to south of the Mogollon Plateau into portions of southwestern Mexico and eastern Arizona, USA. The species richness in this ecoregion is moderate, with vertebrate taxa numbering 375 species. The topography consists chiefly of steep foothills and mountains, but includes some deeply dissected high plateaus. Soil types have not been well defined; however, most soils are entisols, with alfisols and inceptisols in upland areas. Stony terrain and rock outcrops occupy large areas on the mountains and foothills.

The Transition Zone in this region (1980 to 2440 m in elevation) comprises a strong Mexican fasciation, including Chihuahua Pine (Pinus leiophylla) and Apache Pine (P. engelmannii) and unique varieties of Ponderosa Pine (P. ponderosa var. arizonica). Such forests are open and park-like and contain many bird species from Mexico seldom seen in the U.S.. The Canadian Zone (above 2000 m) includes mostly Rocky Mountain species of mixed-conifer communities such as Douglas-fir (Pseudotsuga menziesii), Engelmann Spruce (Picea engelmanni), Subalpine Fir (Abies lasiocarpa), and Corkbark Fir (A. lasiocarpa var. arizonica). Dwarf Juniper (Juniperus communis) is an understory shrubby closely associated with spruce/fir forests. Exposed sites include Chihuahua White Pine (Pinus strobiformis), while disturbed north-facing sites consists primarily of Lodgepole Pine (Pinus contorta) or Quaking Aspen (Populus tremuloides).

There are a variety of mammalian species found in this ecoregion, including the endemic Arizona Gray Squirrel (Sciurus arizonensis), an herbivore who feeds on a wide spectrum of berries, bark and other vegetable material. Non-endemic mammals occurring in the ecoregion include: the Banner-tailed Kangaroo Rat (Dipodomys spectabilis NT); Desert Pocket Gopher (Geomys arenarius NT). In addition, there is great potential for restoring Mexican Wolf (Canis lupus) and Grizzly Bear (Ursus arctos horribilis) populations in the area because of its remoteness and juxtaposition to other ecoregions where these species were formerly prevalent.

There are few amphibians found in the Arizona mountain forests. Anuran species occurring here are: Red-spotted Toad (Anaxyrus punctatus); Southwestern Toad (Anaxyrus microscaphus); New Mexico Spadefoot Toad (Spea multiplicata); Woodhouse's Toad (Anaxyrus woodhousii); Northern Leopard Frog (Lithobates pipiens); Chiricahua Leopard Frog (Lithobates chiricahuensis VU); Madrean Treefrog (Hyla eximia), a montane anuran found at the northern limit of its range in this ecoregion; Boreal Chorus Frog (Anaxyrus woodhousii); Western Chorus Frog (Pseudacris triseriata); and Canyon Treefrog (Hyla arenicolor). The Jemez Mountains Salamander (Plethodon neomexicanus NT) is an ecoregion endemic, found only in the Jemez Mountains of Los Alamos and Sandoval counties, New Mexico. Another salamander occurring in the ecoregion is the Tiger Salamander (Ambystoma tigrinum).

A number of reptilian taxa occur in the Arizona mountains forests, including: Gila Monster (Heloderma suspectum NT), often associated with cacti or desert scrub type vegetation; Narrow-headed Garter Snake (Thamnophis rufipunctatus), a near-endemic found chiefly in the Mogollon Rim area; Sonoran Mud Turtle (Kinosternon sonoriense NT).

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Habitat and Ecology

Habitat and Ecology
Springs, slow streams, marshes, bogs, ponds, canals, flood plains, reservoirs, and lakes; usually permanent water with rooted aquatic vegetation. In summer, commonly inhabits wet meadows and fields. Takes cover underwater, in damp niches, or in caves when inactive. Over winters usually underwater. Eggs are laid and larvae develop in shallow, still, permanent water (typically), generally in areas well exposed to sunlight. Generally eggs are attached to vegetation just below the surface of the water. In northern Minnesota, successful reproduction in acidic bog water either does not occur or is a rare event (Karns 1992).
In Panama, it is a largely terrestrial species of humid lowland and montane forest.

Systems
  • Terrestrial
  • Freshwater
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Comments: Northern leopard frogs live in the vicinity of springs, slow streams, marshes, bogs, ponds, canals, flood plains, reservoirs, and lakes; usually they are in or near permanent water with rooted aquatic vegetation. In summer, they commonly inhabit wet meadows and fields. The frogs take cover underwater, in damp niches, or in caves when inactive. Wintering sites are usually underwater, though some frogs possibly overwinter underground.

Eggs are laid and larvae develop in shallow, still, permanent water (typically), generally in areas well exposed to sunlight. Generally eggs are attached to vegetation just below the surface of the water. In northern Minnesota, successful reproduction in acidic bog water either does not occur or is a rare event (Karns 1992).

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Leopard frogs are found in a wide variety of habitats, including marshlands, brushlands, and forests. They prefer the presence of permanent, slow-moving water, including aquatic vegetation, but can be found in agricultural areas and on golf courses. Leopard frogs are well-adapted to cold and can be found at elevations up to 3,350 meters. They are commonly known as meadow frogs or grass frogs because they tend to stray far from the water when it is not breeding season. They prefer open areas to woods.

Range elevation: 3350 (high) m.

Habitat Regions: temperate ; terrestrial ; freshwater

Terrestrial Biomes: savanna or grassland ; forest ; scrub forest

Aquatic Biomes: lakes and ponds

Wetlands: marsh

Other Habitat Features: agricultural ; riparian

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Leopard frogs are found in a wide variety of habitats, including marshlands, brushlands, and forests. They prefer the presence of permanent, slow-moving water, including aquatic vegetation, but can be found in agricultural areas and on golf courses. Leopard frogs are well-adapted to cold and can be found at elevations up to 3,350 meters. They are commonly known as meadow frogs or grass frogs because they tend to stray far from the water when it is not breeding season. They prefer open areas to woods.

Range elevation: 3350 (high) m.

Habitat Regions: temperate ; terrestrial ; freshwater

Terrestrial Biomes: savanna or grassland ; forest ; scrub forest

Aquatic Biomes: lakes and ponds

Wetlands: marsh

Other Habitat Features: agricultural ; riparian

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Migration

Non-Migrant: No. All populations of this species make significant seasonal migrations.

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

Comments: Metamorphosed frogs eat various small invertebrates obtained along water's edge or in nearby meadows or fields; rarely eats small vertebrates. Larvae eat algae, plant tissue, organic debris, and probably some small invertebrates.

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Food Habits

Leopard frog tadpoles are mainly herbivorous, eating algae, diatoms, and small animal matter filtered from the water or scraped from surfaces. Once they metamorphose into a frog, leopard frogs eat terrestrial invertebrates, including spiders, insects and their larvae, slugs, snails, and earthworms. Large adults may also eat small vertebrates, such as smaller frogs (spring peepers, Pseudacris_crucifer, and chorus frogs, Pseudacris_triseriata).

Animal Foods: amphibians; insects; terrestrial non-insect arthropods; mollusks; terrestrial worms

Plant Foods: algae

Other Foods: detritus

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Food Habits

Leopard frog tadpoles are mainly herbivorous, eating algae, diatoms, and small animal matter filtered from the water or scraped from surfaces. Once they metamorphose into a frog, leopard frogs eat terrestrial invertebrates, including spiders, insects and their larvae, slugs, snails, and earthworms. Large adults may also eat small vertebrates, such as smaller frogs (spring peepers, Pseudacris crucifer, and chorus frogs, Pseudacris triseriata).

Animal Foods: amphibians; insects; terrestrial non-insect arthropods; mollusks; terrestrial worms

Plant Foods: algae

Other Foods: detritus

Primary Diet: carnivore (Insectivore , Molluscivore )

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Associations

Ecosystem Roles

Leopard frogs are important predators of their invertebrate prey and eggs and adults can act as important food sources for small to medium-sized predators.

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Predation

Many predators take advantage of leopard frog prey. Adults are taken by fish (Micropterus and Esox), Ardeidae, Rana clamitans, Rana catesbeiana, Thamnophis, Nerodia, Accipitridae, Laridae, Procyon lotor, Vulpes vulpes, Mustela vison, and Lontra canadensis, as well as other predators. Eggs are eaten by Hirudinea, Salamandridae, and Testudinea. Tadpoles are preyed on by Dysticidae, Belostomatidae, Odonata, and most of the vertebrates that prey on adults.

Leopard frogs do not have distasteful skin secretions, they rely on their quick responses to leap into the water or make erratic hops to escape capture. Their coloration makes them blend into their surroundings when in vegetation. In areas where they co-occur with pickerel frogs (Rana_palustris), leopard frogs have spots that are squarish, like those of pickerel frogs. Because pickerel frogs have distasteful skin secretions, it is thought that perhaps leopard frogs in those areas are mimicing pickerel frogs to avoid predation.

Known Predators:

  • bass (Micropterus)
  • pike (Esox)
  • herons (Ardeidae)
  • green frogs (Rana_clamitans)
  • bullfrogs (Rana_catesbeiana)
  • garter snakes (Thamnophis)
  • water snakes (Nerodia)
  • hawks (Accipitridae)
  • gulls Laridae 
  • raccoons (Procyon_lotor)
  • foxes (Vulpes_vulpes)
  • mink (Mustela_vison)
  • otters (Lontra_canadensis)
  • leeches (Hirudinea)
  • newts (Salamandridae)
  • turtles (Testudines)
  • diving beetles (Dysticidae)
  • giant water bugs (Belostomatidae)
  • dragonfly larvae (Odonata)

Anti-predator Adaptations: mimic; cryptic

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Ecosystem Roles

Leopard frogs are important predators of their invertebrate prey and eggs and adults can act as important food sources for small to medium-sized predators.

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Predation

Many predators take advantage of leopard frog prey. Adults are taken by fish (bass and pike), herons, green frogs, bullfrogs, garter snakes, water snakes, hawks, gulls, raccoons, foxes, mink, and otters, as well as other predators. Eggs are eaten by leeches, newts, and turtles. Tadpoles are preyed on by diving beetles, giant water bugs, dragongfly larvae, and most of the vertebrates that prey on adults.

Leopard frogs do not have distasteful skin secretions, they rely on their quick responses to leap into the water or make erratic hops to escape capture. Their coloration makes them blend into their surroundings when in vegetation. In areas where they co-occur with pickerel frogs (Lithobates palustris), leopard frogs have spots that are squarish, like those of pickerel frogs. Because pickerel frogs have distasteful skin secretions, it is thought that perhaps leopard frogs in those areas are mimicing pickerel frogs to avoid predation.

Known Predators:

Anti-predator Adaptations: mimic; cryptic

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Known predators

Rana pipiens is prey of:
Thamnophis
Actinopterygii

Based on studies in:
Canada: Manitoba (Forest)

This list may not be complete but is based on published studies.
  • R. D. Bird, Biotic communities of the Aspen Parkland of central Canada, Ecology, 11:356-442, from p. 410 (1930).
  • R. D. Bird, Biotic communities of the Aspen Parkland of central Canada, Ecology, 11:356-442, from p. 393 (1930).
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Known prey organisms

Rana pipiens preys on:
Pontania petiolaridis
Insecta
Collembola
Disyonicha quinquevitata
Gastropoda
Araneae
Orthoptera
Elateridae
Noctuidae

Based on studies in:
Canada: Manitoba (Forest)

This list may not be complete but is based on published studies.
  • R. D. Bird, Biotic communities of the Aspen Parkland of central Canada, Ecology, 11:356-442, from p. 410 (1930).
  • R. D. Bird, Biotic communities of the Aspen Parkland of central Canada, Ecology, 11:356-442, from p. 393 (1930).
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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. Ranked S4 or S5 in more than 15 states/provinces.

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Global Abundance

100,000 - 1,000,000 individuals

Comments: Total adult population size likely is in the hundreds of thousands or millions.

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General Ecology

In Michigan, the average nightly movement during rain was 36 m, occasionally moved more than 100 m. See Mazerolle (2001) for information on movement patterns of frogs in fragmented peat bogs in New Brunswick.

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Life History and Behavior

Behavior

Communication and Perception

Leopard frogs use calls to attract mates during breeding season. Male advertisement calls are described as sounding like a low, rumbling snore with occasional clicks and croaks. Males and non-receptive females will give a chuckle-like "release" call when clasped by a male hoping to mate. Outside of breeding season there is little need to communicate with conspecifics. They may scream loudly when they have been seized by a predator or squawk as they jump to avoid a predator. Frogs in general are quite sensitive to movement in detecting prey visually.

Communication Channels: tactile ; acoustic

Perception Channels: visual ; tactile ; acoustic

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Communication and Perception

Leopard frogs use calls to attract mates during breeding season. Male advertisement calls are described as sounding like a low, rumbling snore with occasional clicks and croaks. Males and non-receptive females will give a chuckle-like "release" call when clasped by a male hoping to mate. Outside of breeding season there is little need to communicate with conspecifics. They may scream loudly when they have been seized by a predator or squawk as they jump to avoid a predator. Frogs in general are quite sensitive to movement in detecting prey visually.

Communication Channels: tactile ; acoustic

Perception Channels: visual ; tactile ; acoustic

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Life Cycle

Development

The small, black and white eggs are laid in clusters and attached to submerged vegetation. When Leopard frog eggs are laid they are roughly 1.7 mm in diameter, but swell to 5 mm when they come in contact with water. Clusters of eggs may act to increase heat absorption by the mass or to protect some eggs from predation. Hatching occurs after 1 to 3 weeks, varying with water temperature, and metamorphosis occurs after 70 to 110 days as a tadpole. Froglets are 2 to 3 cm long at metamorphosis.

Development - Life Cycle: metamorphosis

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Development

The small, black and white eggs are laid in clusters and attached to submerged vegetation. When Leopard frog eggs are laid they are roughly 1.7 mm in diameter, but swell to 5 mm when they come in contact with water. Clusters of eggs may act to increase heat absorption by the mass or to protect some eggs from predation. Hatching occurs after 1 to 3 weeks, varying with water temperature, and metamorphosis occurs after 70 to 110 days as a tadpole. Froglets are 2 to 3 cm long at metamorphosis.

Development - Life Cycle: metamorphosis

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Life Expectancy

Lifespan/Longevity

Leopard frogs may live up to 9 years in the wild, although very few leopard frogs will live for this long. Most mortality occurs as a tadpole or newly transformed froglet, when as many as 95% will die.

Range lifespan

Status: wild:
9 (high) years.

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Lifespan/Longevity

Leopard frogs may live up to 9 years in the wild, although very few leopard frogs will live for this long. Most mortality occurs as a tadpole or newly transformed froglet, when as many as 95% will die.

Range lifespan

Status: wild:
9 (high) years.

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Lifespan, longevity, and ageing

Maximum longevity: 9 years (wild) Observations: In the wild, these animals may live up to 5 years (Smirina 1994).
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Reproduction

The time of egg deposition varies with latitude and elevation. Egg deposition occurs typically in April in southern Quebec, New York, and the Great Lakes region, late April to late May farther north in Manitoba and Nova Scotia (see Gilbert et al. 1994). In Colorado, eggs are laid mainly in early spring at low elevations, in late spring in the mountains (Hammerson 1999). Breeding often peaks when water temperatures reach about 10 C. At a particular site, egg deposition generally occurs within a span of about 10 days. Egg masses include several hundred to several thousand ova. Aquatic larvae metamorphose into small frogs in early to late summer, a few months after egg deposition. Females are sexually mature usually in two years in most areas, three years in high elevation populations. Density of egg masses often reaches a few hundred per ha in favorable habitat, sometimes >1000/ha.

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Leopard frogs gather at communal breeding ponds in the spring, where males call to attract females. If successful, a male will hold a female in "amplexus", using his specialized thumbs, and fertilize her eggs as they leave her body. Mating pairs may move to an area of the pond where other pairs have laid their eggs before they add their own.

Mating System: polygynandrous (promiscuous)

Mating occurs from March to June, but peaks in April. Females lay from 300 to 6500 eggs in globular clusters in breeding ponds. After metamorphosis, sexual maturity is reached in 1 to 3 years, depending on conditions.

Breeding interval: Leopard frogs breed once yearly.

Breeding season: Leopard frogs breed from March to June.

Range number of offspring: 300 to 6500.

Range age at sexual or reproductive maturity (female): 1 to 3 years.

Range age at sexual or reproductive maturity (male): 1 to 3 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (External ); oviparous

Most parental investment occurs prior to fertilization. Females will provide the eggs with nourishment to grow and will attempt to attach them to underwater vegetation in a tight cluster, after which the eggs are left to develop on their own.

Parental Investment: pre-fertilization (Provisioning, Protecting: Female)

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Leopard frogs gather at communal breeding ponds in the spring, where males call to attract females. If successful, a male will hold a female in "amplexus", using his specialized thumbs, and fertilize her eggs as they leave her body. Mating pairs may move to an area of the pond where other pairs have laid their eggs before they add their own.

Mating System: polygynandrous (promiscuous)

Mating occurs from March to June, but peaks in April. Females lay from 300 to 6500 eggs in globular clusters in breeding ponds. After metamorphosis, sexual maturity is reached in 1 to 3 years, depending on conditions.

Breeding interval: Leopard frogs breed once yearly.

Breeding season: Leopard frogs breed from March to June.

Range number of offspring: 300 to 6500.

Range age at sexual or reproductive maturity (female): 1 to 3 years.

Range age at sexual or reproductive maturity (male): 1 to 3 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (External ); oviparous

Most parental investment occurs prior to fertilization. Females will provide the eggs with nourishment to grow and will attempt to attach them to underwater vegetation in a tight cluster, after which the eggs are left to develop on their own.

Parental Investment: pre-fertilization (Provisioning, Protecting: Female)

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Molecular Biology and Genetics

Molecular Biology

Barcode data: Rana pipiens

The following is a representative barcode sequence, the centroid of all available sequences for this species.


There are 10 barcode sequences available from BOLD and GenBank.

Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.

See the BOLD taxonomy browser for more complete information about this specimen and other sequences.

NNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNGAATAAATAATATAAGTTTTTGACTTCTTCCCCCATCCTTCTTCCTACTTTTAGCCTCCTCTACAGTTGAAGCAGGAGCCGGAACTGGATGAACAGTTTATCCCCCCCTGGCTGGTAATCTGGCCCATNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNTCTCCCAGTTCTAGCGGCAGGGATTACTATACTCTTAACAGACCGAAATTTAAATACTACCTTTTTCGACCCCGCAGGAGGTGGTGACCCAGTTCTTTATCAACATTTATTC
-- end --

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Statistics of barcoding coverage: Rana pipiens

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 1
Specimens with Barcodes: 5
Species With Barcodes: 1
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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
LC
Least Concern

Red List Criteria

Version
3.1

Year Assessed
2004

Assessor/s
Geoffrey Hammerson, Frank Solís, Roberto Ibáñez, César Jaramillo, Querube Fuenmayor

Reviewer/s
Global Amphibian Assessment Coordinating Team (Simon Stuart, Janice Chanson, Neil Cox and Bruce Young)

Contributor/s

Justification
Listed as Least Concern in view of its wide distribution, tolerance of a degree of habitat modification, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category.
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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

Reasons: Large range throughout much of the U.S. and southern Canada; still common in many areas and in a diverse array of pristine and disturbed habitats; populations have declined in some areas due to habitat loss and degradation, overexploitation, interactions with non-native species, and unknown causes, but the overall range remains essentially undiminished.

Intrinsic Vulnerability: Moderately vulnerable

Environmental Specificity: Moderate to broad.

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Leopard frogs were once common and widespread throughout much of North America. However, some populations have experienced serious declines. In the Great Lakes area, leopard frogs were abundant through the 1970's, after which they experienced a population decline. They remain uncommon in this area, although they can be locally abundant. In the western states, the status of many leopard frog populations remains unstudied. Leopard frogs, along with many other frog species, are at risk of poisoning by pesticides, including atrazine and organochlorines, herbicides, such as nitrates, and other water contaminants. Atrazine has been demonstrated to induce reproductive abnormalities in frogs at levels lower than are found in most North American water sources. Infectious diseases may also pose a significant threat to leopard frogs. Susceptibility to infectious diseases may be exacerbated by water acidification, lowering leopard frog immune responses. Introduced species, including bullfrogs (Rana_catesbeiana) and common carp (Cyprinus_carpio), may be contributing to declining numbers of leopard frogs as well, as they prey extensively on young and adults. Leopard frogs are extensively collected in some areas for use in classrooms, laboratories, and as bait, devastating local populations. Finally, leopard frogs, and other freshwater aquatic species, are at risk because of extensive freshwater habitat loss in North America, estimated at 53% of wetlands lost in the 1980's since 1780.

IUCN Red List of Threatened Species: no special status

US Federal List: no special status

CITES: no special status

State of Michigan List: no special status

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Leopard frogs were once common and widespread throughout much of North America. However, some populations have experienced serious declines. In the Great Lakes area, leopard frogs were abundant through the 1970's, after which they experienced a population decline. They remain uncommon in this area, although they can be locally abundant. In the western states, the status of many leopard frog populations remains unstudied. Leopard frogs, along with many other frog species, are at risk of poisoning by pesticides, including atrazine and organochlorines, herbicides, such as nitrates, and other water contaminants. Atrazine has been demonstrated to induce reproductive abnormalities in frogs at levels lower than are found in most North American water sources. Infectious diseases may also pose a significant threat to leopard frogs. Susceptibility to infectious diseases may be exacerbated by water acidification, lowering leopard frog immune responses. Introduced species, including bullfrogs (Lithobates catesbeianus) and common carp (Cyprinus carpio), may be contributing to declining numbers of leopard frogs as well, as they prey extensively on young and adults. Leopard frogs are extensively collected in some areas for use in classrooms, laboratories, and as bait, devastating local populations. Finally, leopard frogs, and other freshwater aquatic species, are at risk because of extensive freshwater habitat loss in North America, estimated at 53% of wetlands lost in the 1980's since 1780.

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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Population

Population
In North America there are thousands of populations. The total adult population size is probably in the hundreds of thousands or millions. It is still widespread and common in many areas, especially in lowland areas, but many other populations appear to have declined, especially in the Rocky Mountains of Colorado, Wyoming, and Montana, where the species no longer is extant in most localities where historically it occurred (Corn and Fogleman 1984; Corn et al. 1989; Koch and Peterson 1995; J. Reichel, unpublished map, 1996). It has nearly disappeared from the Greater Yellowstone ecosystem, though natural wetland habitats remain apparently undisturbed with acceptable water quality (Koch and Peterson 1995). It is apparently extirpated from most of its historical range in Washington (Leonard et al. 1999). It has not been observed in recent years in the few historical localities in Oregon (Csuti et al. 1997). Local extirpations have been reported for Alberta (Russell and Bauer 1993) and British Columbia (Orchard 1992). In Panama it can be common in some areas but declining in parts of its range.

Population Trend
Decreasing
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Global Short Term Trend: Decline of 10-30%

Comments: Probably declining in population size, area of occupancy, and condition of occurrences.

Global Long Term Trend: Decline of 30-50%

Comments: Still widespread and common in many areas, especially in lowland areas, but many other populations appear to have declined, especially in the Rocky Mountains of Colorado, Wyoming, and Montana, where the species no longer is extant in most localities where historically it occurred (Corn and Fogleman 1984; Corn et al. 1989; Koch and Peterson 1995; J. Reichel, unpublished map, 1996). Has nearly disappeared from the Greater Yellowstone ecosystem, though natural wetland habitats remain apparently undisturbed with acceptable water quality (Koch and Peterson 1995). Apparently extirpated from most of historical range in Washington (Leonard et al. 1999). Not observed in recent years in the few historical localities in Oregon (Csuti et al. 1997). Local extirpations have been reported for Alberta (Russell and Bauer 1993) and British Columbia (Orchard 1992). Declined in northwestern Indiana between the 1930s and 1990s (Brodman et al. 2002).

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Life History, Abundance, Activity, and Special Behaviors

R. pipiens breeds from mid-March to early June (Stebbins 1985). Egg clusters are typically firm and globular, and 2-6 inches in diameter. They are usually attached to vegetation in the calm water of lakes, ponds, canals, and streams. A cluster can contain up to about 6,500 eggs (Stebbins 2003).

The species has a snore-like call, interspersed with grunting and chuckling and lasting from 1 to 5 seconds. Choruses are a mixture of moaning, grunting, and chuckling. Individuals sometimes squawk when jumping into the water, and may scream if caught. (Stebbins 1985).

R. pipiens may forage far from water in damp meadows. If frightened on land, it rushes toward water in a zigzag pattern of jumps. Specimens are most easily found at night by their eyeshine (Stebbins 1985).

  • Stebbins, R. C. (1985). A Field Guide to Western Reptiles and Amphibians. Houghton Mifflin, Boston.
  • Stebbins, R. C. (2003). Western Reptiles and Amphibians, Third Edition. Houghton Mifflin, Boston.
  • Fellers, G. J., Green, D. E., and Longcore, J. E. (2001). ''Oral chytridiomycosis in the Mountain Yellow-Legged Frog (Rana muscosa).'' Copeia, 2001(4), 945-953.
  • Brodkin, M., Vatcnick, I., Simon, M., Hollyann, H., Butler-Holston, K., and Leonard, M. (2003). ''Effects of acid stress in adult Rana pipiens.'' Journal of Experimental Zoology, 298A(1), 16-22.
  • Gibbs, J. P. (2000). ''Wetland loss and biodiversity conservation.'' Conservation Biology, 14(1), 314-317.
  • Glennemeier, K. A. and Denver, R. J. (2001). ''Sublethal effects of chronic exposure to an organochlorine compound on northern leopard frog (Rana pipiens) tadpoles.'' Environmental Toxicology, 16(4), 287-297.
  • Hayes, T., Haston, K., Tsui, M., Hoang, A., Haeffele, C., and Vonk, A. (2002). ''The feminization of male frogs in the wild.'' Nature, 419, 895-896.
  • Hecnar, S. J. (1995). "Acute and chronic toxicity of ammonium nitrate fertilizer to amphibians from southern Ontario." Environmental Toxicology and Chemistry, 14(12), 2131-2137.
  • Lannoo, M. J., Lang, K., Waltz, T., and Phillips, G. S. (1994). "An altered amphibian assemblage: Dickinson County, Iowa, 70 years after Frank Blanchard's survey." American Midland Naturalist, 131(2), 311-319.
  • Linck, M. (2000). ''Reduction in road mortality in a northern leopard frog population.'' Journal of the Iowa Academy of Science, 107(3-4), 209-211.
  • Peterson, G., Johnson, L. B., Axler, R. P., and Diamond, S. A. (2002). ''Assessment of the risk of solar ultraviolet radiation to amphibians. II. In situ characterization of exposure in amphibian habitats.'' Environmental Science and Technology, 36(13), 2859-2865.
  • Schothoeffer, A. M., and Koehler, A. V., Meteyer, C. U., Cole, R. A. (2003). ''Influence of Ribeiroia ondatrae (Trematoda: Digenea) infection on limb development and survival of northern leopard frogs (Rana pipiens): Effects of host stage and parasite-exposure level.'' Canadian Journal of Zoology, 81(7), 1144-1153.
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Threats

Major Threats
Threats and degree of threat vary greatly across its range. Threats include habitat loss, commercial over-exploitation, and in some areas, probably competition/predation by bullfrogs or other introduced species. The decline in Rocky Mountains (Corn et al. 1989) is not due to acidification of breeding habitats (Corn and Vertucci 1992). Laboratory results suggest that there might be an interaction between crowding, temperature, and mortality from bacterial infection (e.g., red-leg disease); there was higher mortality when frogs were subjected to crowding and high temperatures (Brodkin et al. 1992). Agricultural chemicals such as atrazine have caused feminisation of frogs in agricultural areas (Hayes et al. 2002). In Panama it is threatened by general habitat loss due to the destruction of natural forests.
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Degree of Threat: Medium

Comments: Threats and degree of threat vary greatly across the range. Threats include habitat loss, commercial overexploitation, and, in some areas, probably competition/predation by bullfrogs or other introduced species. Exposure to pH 5.5 or lower increases vulnerability to bacterial infection (Simon et al. 2002). Decline in Rocky Mountains (Corn et al. 1989) is not due to acidification of breeding habitats (Corn and Vertucci 1992). Laboratory results suggests that there may be an interaction between crowding, temperature, and mortality from bacterial infection (e.g., red-leg disease); there was higher mortality when frogs were subjected to crowding and high temperatures (Brodkin et al. 1992). In Ontario, Canada, leopard frog population density was negatively affected by vehicular traffic within a radius of 1.5 km (Carr and Fahrig 2001).

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Life History, Abundance, Activity, and Special Behaviors

R. pipiens faces a number of threats.

One identified threat is chemical contamination, particularly from agricultural enterprises.

One of the most extensively cited contaminants is the herbicide atrazine, the most commonly used herbicide in the United States and one of the most widely used throughout the world. The breeding season for R. pipiens follows the annual peak period for the application of atrazine in the U.S., and thus coincides with the annual peak of atrazine contamination of water sources (Hayes et al. 2002). In a laboratory study of the effects of water-borne atrazine on R. pipiens, Hayes et al. found that 10-92% of exposed males developed gonadal abnormalities such as retarded development and hermaphroditism. Hermaphroditism was also found in wild R. pipiens specimens collected in a transect from Utah to Iowa. The Hayes study found males with testicular oocytes in all areas where local atrazine sales exceeded .4 kilograms per square kilometer, and water-borne atrazine exceeded 2 parts per billion.

It is believed that atrazine may induce the production of an enzyme that converts androgens into estrogens, and can cause males to produce estrogens at the expense of androgens. This would explain both the presence of oocytes and the inhibition of spermatogenesis. At one site in the Hayes study, a wild population exposed continuously to atrazine exhibited fewer abnormalities than a population exposed intermittently, leading to the suggestion that adaptive resistance may be occurring in the more frequently exposed populations. Atrazine is a significant threat to R. pipiens, and possibly to amphibians in general, because most water sources in the U.S., including rain, contain atrazine at higher levels than those needed to induce abnormalities in laboratory specimens (Hayes et al 2002).

Another agriculture-related threat to R. pipiens is nitrate contamination of water sources. In a laboratory study, tadpoles of four amphibian species were exposed to levels of nitrate commonly exceeded in agricultural areas around the world. Effects varied across the species, but included reduced activity, lower rates of metamorphosis, and physical abnormalities (Hecnar 1995).

Some environmental conditions do not directly cause death, but may induce behavior that increases the likelihood of predation by other species. Exposure to organochlorines, the remnants of such pollutants as DDT, has been suggested as a possible cause of population declines. While organochlorines do not appear to have the same dramatic effects on amphibians as they do on large animals such as predatory birds, exposure to these compounds does appear to induce behavior that might cause a population to decline. In a laboratory study, R. pipiens tadpoles exposed to organochlorines tended to spend more time resting and less time feeding, behavior that could reduce the ability of tadpoles’ to consume scarce resources. Organochlorines may also affect the production of certain hormones that help R. pipiens respond to local environmental changes such as pond drying. Most products that contain organochlorines are now banned, but their derivatives are common and persistent pollutants even today (Glennemeier et al. 2001).

The decline of R. pipiens in areas where it used to thrive has also been attributed in recent years to infectious diseases, whose prevalence may be exacerbated by environmental stresses such as acidification (Brodkin et al. 2003). Acidification is a problem for this species because the breeding season, during which the frogs spend a great deal of time in the water, coincides with the highest levels of acidity in the lakes and streams of the northeastern United States. The breeding season also directly follows the period of winter hibernation, during which cold-exposure weakens the immune system of frogs. In a laboratory study by Brodkin et al., both the degree of cold exposure and the level of acidity to which frogs were exposed correlated to the health of the frogs’ immune systems. Frogs exposed to acid, and frogs exposed to acid after being exposed to cold, demonstrated higher levels of bacterial colonization of the spleen and higher rates of mortality. Brodkin et al. conclude that acidic conditions of pH 5.5 and below contribute to this contamination by: 1) damaging the intestinal epithelium, thereby allowing bacteria to pass from the intestinal tract to the bloodstream and spleen, and 2) reducing the number and viability of white blood cells. Cold exposure alone did not damage frogs’ immune systems, and pH levels of 6.0 or above, while damaging to the intestinal epithelium, were not sufficient to induce high levels of mortality. While it is fairly certain that acid exposure is a threat to R. pipiens, it is not clear why it has become a problem in the last twenty years or so (Brodkin et al. 2003).

There has been a flurry of reports in recent years on the widespread prevalence of limb deformities among many species of frogs, even those in seemingly pristine environments. R. pipiens is one of the species most commonly reported to exhibit such deformities. A number of hypotheses have emerged to explain this trend.

One suggestion is that increased exposure to ultraviolet light may be responsible, because ultraviolet light is known to cause damage to cellular DNA. The validity of the hypothesis when applied to conditions in the wild requires further study. Preliminary work has been done to assess how UVB light penetrates aquatic environments, and to determine what other environmental factors affect amphibians’ exposure to that light (Peterson et al. 2002).

Another hypothesis suggests that environmental stresses have increased the vulnerability of amphibians to parasites such as trematodes (Schothoeffer et al. 2003). In a laboratory study of the interaction between R. pipiens tadpoles and the larvae of the trematode Ribeiroia ondatrae, infection of the frog tadpoles by R. ondatrae led to a number of different types of malformations. Depending on the stage of development at which the tadpole is infected and the intensity of the infection, the developing R. pipiens may develop extra limbs, digits, or phalanges; its limbs or digits may be smaller than normal; bones may bridge, skin may web, and the ilium may be reduced or misshapen. The timing of infection appeared to be crucial, as R. pipiens exhibits different levels of vulnerability as it develops. It appears to be most vulnerable at the larval stage, pre-limb bud stage, and limb bud stage. Infections at the paddle stage appear to have no effect on limb development. In addition to mortality due to limb malformations, infected tadpoles may also die as a result of the infections themselves. It is not yet known what environmental factors relate to the timing of infections, or what factors affect the length of amphibian larval periods and tadpole vulnerability (Schothoeffer et al. 2003).

It is also possible that R. pipiens is suffering decline due to the type of fungal infections that have been found responsible for the decline of other species. The chytrid fungus Batrachochytrium dendrobatidis, for example, was found to cause oral abnormalities in the species Rana muscosa, a species in the Sierra Nevada that has drastically declined recently (Fellers et al. 2001).

Introduced species may also be contributing to the decline of this species (Lannoo et al. 1994). A 1994 study investigated amphibian populations in Dickinson County, Iowa, and found that since a study of the same area in 1923, several populations had declined and two had disappeared. In Dickinson County around the year 1900, R. pipiens was collected and exported at a rate of about 20 million specimens per year. In their 1994 report, Lannoo et al. reported an estimated population of about 50,000 for Dickinson County. They cite the introduction of the common carp and predatory bullfrogs as a partial reason for the decline, in addition to disturbance and loss of habitat (Lannoo et al.).

On the subject of habitat loss, James P. Gibbs reports that by the late 1980s, the lower 48 states had lost 53% of the wetlands that existed in the 1780s. Naturally, water-dependent species such as R. pipiens are negatively affected (Gibbs 2000).

In a related matter, R. pipiens’ decline has been attributed to vehicular traffic, particularly in areas where wintering areas are separated from breeding areas by roads. The full extent of this risk has not been established (Linck 2000).

  • Stebbins, R. C. (1985). A Field Guide to Western Reptiles and Amphibians. Houghton Mifflin, Boston.
  • Stebbins, R. C. (2003). Western Reptiles and Amphibians, Third Edition. Houghton Mifflin, Boston.
  • Fellers, G. J., Green, D. E., and Longcore, J. E. (2001). ''Oral chytridiomycosis in the Mountain Yellow-Legged Frog (Rana muscosa).'' Copeia, 2001(4), 945-953.
  • Brodkin, M., Vatcnick, I., Simon, M., Hollyann, H., Butler-Holston, K., and Leonard, M. (2003). ''Effects of acid stress in adult Rana pipiens.'' Journal of Experimental Zoology, 298A(1), 16-22.
  • Gibbs, J. P. (2000). ''Wetland loss and biodiversity conservation.'' Conservation Biology, 14(1), 314-317.
  • Glennemeier, K. A. and Denver, R. J. (2001). ''Sublethal effects of chronic exposure to an organochlorine compound on northern leopard frog (Rana pipiens) tadpoles.'' Environmental Toxicology, 16(4), 287-297.
  • Hayes, T., Haston, K., Tsui, M., Hoang, A., Haeffele, C., and Vonk, A. (2002). ''The feminization of male frogs in the wild.'' Nature, 419, 895-896.
  • Hecnar, S. J. (1995). "Acute and chronic toxicity of ammonium nitrate fertilizer to amphibians from southern Ontario." Environmental Toxicology and Chemistry, 14(12), 2131-2137.
  • Lannoo, M. J., Lang, K., Waltz, T., and Phillips, G. S. (1994). "An altered amphibian assemblage: Dickinson County, Iowa, 70 years after Frank Blanchard's survey." American Midland Naturalist, 131(2), 311-319.
  • Linck, M. (2000). ''Reduction in road mortality in a northern leopard frog population.'' Journal of the Iowa Academy of Science, 107(3-4), 209-211.
  • Peterson, G., Johnson, L. B., Axler, R. P., and Diamond, S. A. (2002). ''Assessment of the risk of solar ultraviolet radiation to amphibians. II. In situ characterization of exposure in amphibian habitats.'' Environmental Science and Technology, 36(13), 2859-2865.
  • Schothoeffer, A. M., and Koehler, A. V., Meteyer, C. U., Cole, R. A. (2003). ''Influence of Ribeiroia ondatrae (Trematoda: Digenea) infection on limb development and survival of northern leopard frogs (Rana pipiens): Effects of host stage and parasite-exposure level.'' Canadian Journal of Zoology, 81(7), 1144-1153.
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Management

Conservation Actions

Conservation Actions
Populations exist in dozens or hundreds of protected areas, though management of those areas might not take leopard frogs into consideration. In Panama it has been recorded from Parque Nacional Altos de Campana. Taxonomic research is needed to resolve this species complex.
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Global Protection: Many to very many (13 to >40) occurrences appropriately protected and managed

Comments: Populations exist in dozens of protected areas, though management of those areas may not take leopard frogs into consideration.

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Relevance to Humans and Ecosystems

Benefits

Economic Uses

Comments: In some areas, has been subject to heavy commercial exploitation for research and teaching. For example, a harvest of over 100,000/year in Quebec was reported in the early 1980s (see Gilbert et al. 1994).

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Economic Importance for Humans: Negative

There are no negative impacts of leopard frogs on humans.

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Economic Importance for Humans: Positive

Leopard frogs are eaten by humans (frog legs), and are used as test subjects in many biomedical research projects, both as adults and as tadpoles. Leopard frogs are also taken for use in biology classrooms. Leopard frogs are valuable members of the ecosystems in which they live, controlling invertebrate pests and acting as an important food source to larger predators. They may also act as indicator species for environmental health and water quality.

Positive Impacts: food ; research and education; controls pest population

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Economic Importance for Humans: Negative

There are no negative impacts of leopard frogs on humans.

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Economic Importance for Humans: Positive

Leopard frogs are eaten by humans (frog legs), and are used as test subjects in many biomedical research projects, both as adults and as tadpoles. Leopard frogs are also taken for use in biology classrooms. Leopard frogs are valuable members of the ecosystems in which they live, controlling invertebrate pests and acting as an important food source to larger predators. They may also act as indicator species for environmental health and water quality.

Positive Impacts: food ; research and education; controls pest population

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Risks

Relation to Humans

Many of the threats to R. pipiens are human-induced. Human activity can directly damage the species, as in the case of declines due to vehicular traffic, and can also interfere with its life processes indirectly, as is likely the case where pollutants travel to seemingly pristine environments.

  • Stebbins, R. C. (1985). A Field Guide to Western Reptiles and Amphibians. Houghton Mifflin, Boston.
  • Stebbins, R. C. (2003). Western Reptiles and Amphibians, Third Edition. Houghton Mifflin, Boston.
  • Fellers, G. J., Green, D. E., and Longcore, J. E. (2001). ''Oral chytridiomycosis in the Mountain Yellow-Legged Frog (Rana muscosa).'' Copeia, 2001(4), 945-953.
  • Brodkin, M., Vatcnick, I., Simon, M., Hollyann, H., Butler-Holston, K., and Leonard, M. (2003). ''Effects of acid stress in adult Rana pipiens.'' Journal of Experimental Zoology, 298A(1), 16-22.
  • Gibbs, J. P. (2000). ''Wetland loss and biodiversity conservation.'' Conservation Biology, 14(1), 314-317.
  • Glennemeier, K. A. and Denver, R. J. (2001). ''Sublethal effects of chronic exposure to an organochlorine compound on northern leopard frog (Rana pipiens) tadpoles.'' Environmental Toxicology, 16(4), 287-297.
  • Hayes, T., Haston, K., Tsui, M., Hoang, A., Haeffele, C., and Vonk, A. (2002). ''The feminization of male frogs in the wild.'' Nature, 419, 895-896.
  • Hecnar, S. J. (1995). "Acute and chronic toxicity of ammonium nitrate fertilizer to amphibians from southern Ontario." Environmental Toxicology and Chemistry, 14(12), 2131-2137.
  • Lannoo, M. J., Lang, K., Waltz, T., and Phillips, G. S. (1994). "An altered amphibian assemblage: Dickinson County, Iowa, 70 years after Frank Blanchard's survey." American Midland Naturalist, 131(2), 311-319.
  • Linck, M. (2000). ''Reduction in road mortality in a northern leopard frog population.'' Journal of the Iowa Academy of Science, 107(3-4), 209-211.
  • Peterson, G., Johnson, L. B., Axler, R. P., and Diamond, S. A. (2002). ''Assessment of the risk of solar ultraviolet radiation to amphibians. II. In situ characterization of exposure in amphibian habitats.'' Environmental Science and Technology, 36(13), 2859-2865.
  • Schothoeffer, A. M., and Koehler, A. V., Meteyer, C. U., Cole, R. A. (2003). ''Influence of Ribeiroia ondatrae (Trematoda: Digenea) infection on limb development and survival of northern leopard frogs (Rana pipiens): Effects of host stage and parasite-exposure level.'' Canadian Journal of Zoology, 81(7), 1144-1153.
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Wikipedia

Northern leopard frog

The northern leopard frog (Lithobates pipiens, formerly known as Rana pipiens)[1][2] is a species of leopard frog from the true frog family, native to parts of Canada and United States. It is the state amphibian of Minnesota and Vermont.

Physical description[edit]

Young northern leopard frog

The northern leopard frog is a fairly large species of frog, reaching about 11 cm (4.3 in) in length. It varies from green to brown in dorsal colour, with large, dark, circular spots on its back, sides, and legs. Each spot is normally bordered by a lighter ring. A pair of dorsolateral folds starting from the back of the eye run parallel to each other down the back. These dorsolateral folds are often lighter or occasionally pinkish in colour. There is also a pale stripe running from the nostril, under the eye and tympanum, terminating at the shoulder. The ventral surface is white or pale green. The iris is golden and toes are webbed. Tadpoles are dark brown or grey, with light blotches on the underside. The tail is pale tan.

Color Variations[edit]

Two burnsi morphs, a green morph, and a brown morph of the northern leopard frog

The northern leopard frog has several different color variations. The most common being the green morph and the brown morph. There is another morph known as the burnsi morph. Individuals with the burnsi morph coloration lack spots on their back, but may or may not retain them on their legs. They can be bright green or brown and have yellow dorsal folds. [3] Albinism also appears in this species, but it is very rare.

Ecology and behavior[edit]

Northern Leopard Frog Ontario 1.JPG

Northern leopard frogs have a wide range of habitats. They are found in permanent ponds, swamps, marshes, and slow-moving streams throughout forest, open, and urban areas. They normally inhabit water bodies with abundant aquatic vegetation. They are well adapted to cold and can be found above 3,000 m (9,800 ft) asl. Males make a short snore-like call from water during spring and summer. The northern leopard frog breeds in the spring (March–June). Up to 6500 eggs are laid in water, and tadpoles complete development within the breeding pond. Tadpoles are light brown with black spots, and development takes 70–110 days, depending on conditions. Metamorph frogs are 2–3 cm (0.79–1.18 in) long and resemble the adult.

This species was once quite common through parts of western Canada and the United States until declines started occurring during the 1970s. Although the definitive cause of this decline is unknown, habitat loss and fragmentation, environmental contaminants, introduced fish, drought, and disease have been proposed as mechanisms of decline and are likely preventing species recovery in many areas. Many populations of northern leopard frogs have not yet recovered from these declines.

Northern leopard frogs are preyed upon by many different animals, such as snakes, raccoons, other frogs, and even humans. They do not produce distasteful skin secretions and rely on speed to evade predation.

They eat a wide variety of animals, including crickets, flies, worms, and smaller frogs. Using their large mouths, they can even swallow birds and garter snakes. This species is similar to the pickerel frog (Lithobates palustris) and the southern leopard frog (Lithobates sphenocephala).

Research[edit]

Medical[edit]

The northern leopard frog produces specific ribonucleases in its oocytes. Those enzymes are potential drugs for cancer. One such molecule, called ranpirnase (onconase), is in clinical trials as a treatment for pleural mesothelioma and lung tumours. Another, amphinase, was recently described as a potential treatment for brain tumors.[4]

Neuroscience[edit]

The northern leopard frog has been a preferred species for making discoveries about basic properties of neurons since before the 1950s. The neuromuscular junction of the sciatic nerve fibers of the sartorius muscle of this frog has been the source of a lot of initial data about the nervous system.[5][6][7][8][9][10]

Muscle physiology and biomechanics[edit]

The northern leopard frog is a popular species for in vitro experiments in muscle physiology and biomechanics due to the ease of accessibility for investigators in its native range and the ability of the sartorius muscle to stay alive in vitro for several hours. Furthermore, the reliance of the frog on two major modes of locomotion (jumping and swimming) allows for understanding how muscle properties contribute to organismal performance in each of these modes.

See also[edit]

References[edit]

  1. ^ Integrated Taxonomic Information System [Internet] 2012. Lithobates pipiens [updated 2012 Sept; cited 2012 Dec 26] Available from: www.itis.gov/
  2. ^ Geoffrey Hammerson, Frank Solís, Roberto Ibáñez, César Jaramillo, Querube Fuenmayor 2004. Lithobates pipiens. In: IUCN 2012. IUCN Red List of Threatened Species. Version 2012.2. <www.iucnredlist.org>. Downloaded on 26 December 2012.
  3. ^ "Northern Leopard Frog Rana pipens". HerpNet. Retrieved 2013-10-30. 
  4. ^ Frog molecule could provide drug treatment for brain tumors
  5. ^ Fatt, P; Katz, B (1952). "Spontaneous subthreshold activity at motor nerve endings". The Journal of physiology 117 (1): 109–28. PMC 1392564. PMID 14946732. 
  6. ^ Del Castillo, J; Katz, B (1954). "Quantal components of the end-plate potential". The Journal of physiology 124 (3): 560–73. PMC 1366292. PMID 13175199. 
  7. ^ Katz, B; Miledi, R (1965). "The Measurement of Synaptic Delay, and the Time Course of Acetylcholine Release at the Neuromuscular Junction". Proceedings of the Royal Society of London. Series B 161 (985): 483–95. doi:10.1098/rspb.1965.0016. PMID 14278409. 
  8. ^ Kuffler, SW; Yoshikami, D (1975). "The number of transmitter molecules in a quantum: An estimate from iontophoretic application of acetylcholine at the neuromuscular synapse". The Journal of physiology 251 (2): 465–82. PMC 1348438. PMID 171380. 
  9. ^ Hille, B (1967). "The selective inhibition of delayed potassium currents in nerve by tetraethylammonium ion". The Journal of General Physiology 50 (5): 1287–302. doi:10.1085/jgp.50.5.1287. PMC 2225709. PMID 6033586. 
  10. ^ Anderson, CR; Stevens, CF (1973). "Voltage clamp analysis of acetylcholine produced end-plate current fluctuations at frog neuromuscular junction". The Journal of physiology 235 (3): 655–91. PMC 1350786. PMID 4543940. 

Further reading[edit]

  • Hillis, David M.; Frost, John S.; Wright, David A. (1983). "Phylogeny and Biogeography of the Rana pipiens Complex: A Biochemical Evaluation". Systematic Zoology 32 (2): 132–43. doi:10.1093/sysbio/32.2.132. JSTOR 2413277. 
  • Hillis, D M (1988). "Systematics of the Rana Pipiens Complex: Puzzle and Paradigm". Annual Review of Ecology and Systematics 19: 39–63. doi:10.1146/annurev.es.19.110188.000351. JSTOR 2097147. 
  • Ankley, G. T., Tietge, J. E., DeFoe, D. L., Jensen, K. M., Holcombe, G. W., Durhan, E. J., & Diamond, S. A. (1998). Effects of ultraviolet light and methoprene on survival and development of Rana pipiens. Environmental Toxicology and Chemistry, 17(12), 2530-2542 (abstract)
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Names and Taxonomy

Taxonomy

Comments: Much published information on "Rana pipiens" actually pertains to other species that have been described or recognized since the early 1970s.

Hoffman and Blouin (2004) used mtDNA data to develop a hypothesis regarding the evolutionary history and phylogeography of Rana pipiens.

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