Endangered and Threatened Wildlife and Plants; 12-Month Finding on a Petition To List the Northern Leopard Frog in the Western United States as Threatened, 61896-61931 [2011-25498]
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DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS–R2–ES–2009–0030;
92210–1111–FY08–B2]
Endangered and Threatened Wildlife
and Plants; 12–Month Finding on a
Petition To List the Northern Leopard
Frog in the Western United States as
Threatened
Fish and Wildlife Service,
Interior.
ACTION: Notice of 12-month petition
finding.
AGENCY:
We, the U.S. Fish and
Wildlife Service, announce a 12-month
finding on a petition to list the northern
leopard frog (Lithobates (=Rana)
pipiens) under the Endangered Species
Act of 1973, as amended (Act). After
review of the best scientific and
commercial information, we find that
listing the northern leopard frog is not
warranted at this time. However, we ask
the public to submit to us any new
information that becomes available
concerning threats to the northern
leopard frog or its habitat at any time.
DATES: The finding announced in this
document was made on October 5, 2011.
ADDRESSES: This finding is available on
the Internet at https://
www.regulations.gov at Docket Number
FWS–R2–ES–2009–0030. Supporting
documentation we used in preparing
this finding is available for public
inspection, by appointment, during
normal business hours at the U.S. Fish
and Wildlife Service, Arizona Ecological
Services Office, 2321 West Royal Palm
Road, Suite 103, Phoenix, AZ 85021.
Please submit any new information,
materials, comments, or questions
concerning this finding to the above
street address.
FOR FURTHER INFORMATION CONTACT:
Steven L. Spangle, Field Supervisor,
Arizona Ecological Services Office (see
ADDRESSES); by telephone at (602) 242–
0210; or by facsimile at (602) 242–2513.
If you use a telecommunications device
for the deaf (TDD), please call the
Federal Information Relay Service
(FIRS) at 800–877–8339.
SUPPLEMENTARY INFORMATION:
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SUMMARY:
Background
Section 4(b)(3)(B) of the Act (16
U.S.C. 1531 et seq.), requires that, for
any petition to revise the Federal Lists
of Endangered and Threatened Wildlife
and Plants that contains substantial
scientific or commercial information
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that listing the species may be
warranted, we make a finding within
12 months of the date of receipt of the
petition. In our finding, we are required
to determine if the petitioned action is:
(a) Not warranted, (b) warranted, or (c)
warranted, but the immediate proposal
of a regulation implementing the
petitioned action is precluded by other
pending proposals to determine whether
species are endangered or threatened,
and expeditious progress is being made
to add or remove qualified species from
the Federal Lists of Endangered and
Threatened Wildlife and Plants. Section
4(b)(3)(C) of the Act requires that we
treat a petition for which the requested
action is found to be warranted but
precluded as though resubmitted on the
date of such finding, that is, requiring a
subsequent finding to be made within
12 months. We must publish these 12month findings in the Federal Register.
Previous Federal Actions
On June 5, 2006, we received a
petition from the Center for Native
Ecosystems, Biodiversity Conservation
Alliance, Defenders of Black Hills,
Forest Guardians, Center for Biological
Diversity, The Ark Initiative, Native
Ecosystems Council, Rocky Mountain
Clean Air Action, and Mr. Jeremy
Nichols requesting that the northern
leopard frog (Lithobates (=Rana)
pipiens) occurring in the western United
States (Arizona, California, Colorado,
Idaho, Iowa, Minnesota, Missouri,
Montana, Nebraska, Nevada, New
Mexico, North Dakota, Oregon, South
Dakota, Texas, Utah, Washington, and
Wyoming) be listed as a threatened
distinct population segment (DPS)
under the Act. The petition contained
detailed information on the natural
history, biology, current status, and
distribution of the western population
of the northern leopard frog. It also
contained information on what the
petitioners reported as potential threats
to the western population of the
northern leopard frog such as habitat
loss and degradation, predation and
competition by nonnative species,
disease, water pollution, climate
change, and other factors. We
acknowledged the receipt of the petition
in a letter to the petitioners dated
August 7, 2006. That letter explained
that we would not be able to address
their petition at that time. The reason
for this delay was that responding to
court orders and settlement agreements
for other listing actions required nearly
all of our listing funding.
In reviewing the petition, there were
two issues for which the Service
requested clarification from the
petitioners. We were petitioned to list
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the population west of the Mississippi
River and the Great Lakes region in the
United States and south of the
international boundary between the
United States and Canada. However,
although Wisconsin is located west of
the Great Lakes region, the petition map
did not show Wisconsin as a part of the
petition, and the status of the species is
not mentioned in that State. Therefore,
we requested that the petitioners clarify
whether they intended to include or
exclude Wisconsin from the petitioned
DPS. We also sought clarification as to
whether the petitioners were requesting
that we review only the western U.S.
population of the northern leopard frog
as a DPS or if they were also requesting
us to consider listing the entire species
or a significant portion of the range of
the species. The petitioners responded
to our clarification request in a letter
dated February 8, 2008, requesting we
review whether Wisconsin should be
included in the western U.S. population
of the northern leopard frog. In addition,
the petitioners clarified that, if we find
that listing the western U.S. population
of northern leopard frogs as a DPS is not
warranted, we review whether listing
the entire species is warranted because
of threats in a significant portion of its
range.
On July 1, 2009, we published our
90-day finding (74 FR 31389) that the
petition presented substantial scientific
information indicating that listing the
western population of the northern
leopard frog may be warranted, and we
initiated a status review to determine if
listing the species as a DPS or
throughout all or a significant portion of
its range is warranted. Our July 1, 2009,
90-day finding opened a 60-day period
to send us information for our status
review. On October 28, 2009, we
reopened this information solicitation
period for our status review for an
additional 30 days, ending November
27, 2009 (74 FR 55525). This notice
constitutes our 12-month finding on the
February 8, 2008, petition to list the
northern leopard frog.
Species Information
Below we provide information
relevant to understanding the analysis
of information pertaining to the five
factors. See Rorabaugh (2005) for a more
complete description of the distribution
and life history of the northern leopard
frog.
Taxonomy
The northern leopard frog is in the
family Ranidae (Lannoo 2005, p. 371),
the true frogs, and is one of about 28
species within the genus Lithobates
(formerly Rana (Frost et al. 2006, p. 10;
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Frost et al. 2008, pp. 7–8)) that occur in
North America (Lannoo 2005, p. 371).
For more than a century, nomenclatural
and taxonomic confusion has
surrounded members of the Lithobates
(=Rana) complex (Moore 1944, p. 349;
Pace 1974, pp. 11–16; Merrell 1977, pp.
1–2; Hillis et al. 1983, p. 132 among
others), and there is a wealth of
literature from the late 1800s to present
day that has attempted to accurately
describe the different species and
geographic variation within the
complex. Until recently, all North
American ranid frogs (frogs in the
family Ranidae) were included within
the single genus Rana. However, Frost
et al. (2006, p. 10) placed most of these
species into the genus Lithobates. This
change is recognized by the Committee
on Standard English and Scientific
Names, which is the official names list
of the American Society of
Ichthyologists and Herpetologists, the
Herpetologists’ League, and the Society
for the Study of Amphibians and
Reptiles (Frost et al. 2008, pp. 7–8).
Accordingly, the Service also recognizes
and accepts Frost et al.’s (2008)
Lithobates classification.
Physical Description
The northern leopard frog is a slim,
smooth-skinned green, brown, or
sometimes yellow-green frog with
webbed hind feet. The frog is covered
with large, oval dark spots, each of
which is surrounded by a lighter halo or
border (Stebbins 2003, pp. 234–235).
The snout (nose) is pointed and the
tympanum (eardrum) is round and
approximately equal in diameter to the
eye (Baxter and Stone 1980, p. 41).
Northern leopard frogs have a white
stripe on the upper jaw and the
dorsolateral folds (paired, glandular
ridges that run along each side of the
back from behind the eyes to the rear)
are light cream to yellow and are
continuous (not broken posteriorly). The
belly is white to cream-colored, and the
posterior thigh has a light background
color with dark spots. There are two
different color morphs (variants) of the
northern leopard frog that most often
occur in western Minnesota, eastern
North Dakota, and South Dakota
(Rorabaugh 2005, p. 570; McKinnel1 et
al. 2005, p. 7). These color morphs do
occur in other locations (for example,
see Ammon 2002, p. 11), but they are
most prevalent in Minnesota, North
Dakota, and South Dakota, as described
above. The burnsi morph lacks dorsal
spots and the kandiyohi morph has
mottled pigment patches (speckles)
between the dorsal spots. Adult body
lengths (snout-vent) range from 2 to 4.5
inches (in) (5 to 11 centimeters (cm))
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(Stebbins 2003, p. 234). Females average
slightly larger than males (Leonard et al.
1993, p. 138; Werner et al. 2004, p. 97).
Subadult, or recently metamorphosed
frogs (see Biology section below), range
in length from 1 to 2 in (2 to 5 cm)
(Merrell 1977, pp. 10–11). During the
breeding season, males have enlarged or
swollen thumbs (innermost digit) on
forefeet, and vocal sacs are not apparent
except when the frog is calling (Baxter
and Stone 1980, p. 41; Hammerson
1999, p. 145). The typical breeding call
is a prolonged ‘‘snore’’ followed by a
series of stuttering croaks or chuckles
that tend to accelerate towards the end
(Hammerson 1999, p. 145). These
vocalizations may be interspersed with
chuckling sounds (Stebbins 2003, p.
235).
Northern leopard frogs deposit their
egg masses underwater in clusters,
which they attach to vegetation. Eggs are
laid in a single orange- to grapefruitsized globular clump, and may be laid
individually or communally in groups
(Nussbaum et al. 1983, p. 182). Each egg
mass may contain 645 to 7,648
individual eggs (Rorabaugh 2005, p.
572). The eggs hatch into tadpoles.
Tadpoles (the larval stage in the
lifecycle of the frog) are dark green to
brown above with metallic flecking, and
a cream to white translucent underside
(Werner et al. 2004, p. 97). Tadpoles
metamorphose into young frogs. For a
detailed description of northern leopard
frog tadpoles, see Scott and Jennings
(1985, pp. 4–16).
Distribution
The northern leopard frog historically
ranged from Newfoundland and
southern Quebec, south through the
northeast portions of the United States
to West Virginia, west across the
Canadian provinces and northern and
central portions of the United States to
British Columbia, Oregon, Washington,
and northern California, and south to
Arizona, New Mexico, and extreme
western Texas (Rorabaugh 2005, p. 570).
Current range maps tend to show an
extensive and connected distribution for
the northern leopard frog; however, its
actual distribution is sparse and
fragmented in Washington, Oregon,
Idaho, California, Nevada, Arizona, New
Mexico, Utah, Colorado, western
Montana, and western Wyoming in the
western United States (Rorabaugh 2005,
pp. 570–571), throughout New England
(New Hampshire Fish and Game
Department 2005, pp. A208–A209), and
in British Columbia, Northern
Territories, Alberta, Saskatchewan, and
parts of Manitoba in Canada (Committee
on the Status of Endangered Wildlife in
Canada 2009, p. iii).
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Habitat
The northern leopard frog is an
amphibian (a cold-blooded vertebrate
that spends some time on land, but must
breed and develop into an adult in
water) and as such is ectothermic
(incapable of generating their own body
heat) (Wells 2007, p. 2). They have
highly permeable skin, which allows for
rapid passage of water and gases so that
they can use their external environment
to regulate body temperature and
moisture loss (Wells 2007, pp. 2–3). As
part of its complex life history, the
northern leopard frog requires a mosaic
of habitats, which includes aquatic
overwintering and breeding habitats,
and upland post-breeding habitats, as
well as habitat linkages, to meet the
requirements of all of its life stages
(Pope et al. 2000, p. 2505; Smith 2003,
pp. 6–15; Rorabaugh 2005, pp. 571–
575). Although aquatic breeding habitat
is required for long-term population
survival, upland foraging, dispersal, and
overwintering habitats are critical if
individual leopard frogs are to survive
to reproductive maturity. For example,
researchers noted an area near Chicago
that had low northern leopard frog
abundance, but extensive potential
aquatic breeding habitat. It was not until
habitat surrounding the ponds was
restored from scrub forest to grasslands
that leopard frog numbers increased
dramatically (K.S. Mierzwa, pers.
comm. in Pope et al. 2000, p. 2506).
These complex habitat requirements
make northern leopard frogs particularly
vulnerable to the impacts of habitat loss
and fragmentation. Reduction or
removal of these habitats or loss of
connectivity between habitat
components could reduce the capacity
of the landscape to support the species
(Pope et al. 2000, p. 2505; Green 2005,
p. 31).
Northern leopard frogs breed in a
variety of aquatic habitats that include
slow-moving or still water along streams
and rivers, wetlands, permanent or
temporary pools, beaver ponds, and
human-constructed habitats such as
earthen stock tanks and borrow pits
(Rorabaugh 2005, p. 572). Successful
breeding areas typically do not contain
predaceous fish or other predators
(Merrell 1968, p. 275; Hine et al. 1981,
p. 12; Orr et al. 1998, p. 92; Smith 2003,
pp. 19–21). Emergent vegetation, such
as sedges and rushes, are important
features of breeding and tadpole habitats
(Gilbert et al. 1994, p. 468; Smith 2003,
pp. 8–9), and tadpoles are most often
found in backwaters and still pools
(Rorabaugh 2005, p. 572).
Sub-adult northern leopard frogs
typically move from breeding areas to
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feeding sites along the borders of larger,
more permanent bodies of water, as
smaller frogs are closely tied to water
(Merrell 1970, p. 49). Recently
metamorphosed frogs will move up and
down drainages and across land in an
effort to disperse from breeding areas
(Seburn et al. 1997, p. 69) and may
disperse more than 0.5 mile (mi) (800
meters (m)) from their place of
metamorphosis (Dole 1971, p. 223). Dole
(1971, p. 226) found that dispersal in
Michigan occurred on warm, rainy
nights and that frogs dispersed
overland; however, warm rains are not
common in all parts of the species’
range and other dispersal routes may be
important as well. Streams are an
important corridor for dispersing
juvenile frogs (Seburn et al. 1997, pp.
68–69), and vegetated drainage ditches
may also facilitate connectivity between
seasonal habitats (Pope et al. 2000, p.
2505). In some areas of the western
United States, subadults may remain in
the breeding habitat within which they
metamorphosed (Smith 2003, p. 10).
In addition to the breeding habitats,
adult northern leopard frogs require
stream, pond, lake, or river habitats for
overwintering and upland habitats
adjacent to these areas for summer
feeding. In summer, adults and
juveniles commonly feed in open or
semi-open wet meadows and fields with
shorter vegetation, usually near the
margins of water bodies, and seek
escape cover underwater. Post-breeding
summer habitats do not include barren
ground, open sandy areas, heavily
wooded areas, cultivated fields, heavily
grazed pastures, or mowed lawns
(Rorabaugh 2005, p. 573). Buffer zones
around wetland breeding sites should be
maintained for movement to
surrounding upland foraging habitat.
Rittenhouse and Semlitsch (2007, p.
154) collected data from 13 published
radio telemetry and tagging studies
looking at frog and salamander use of
terrestrial habitat surrounding wetlands.
They found that, on average, a buffer
width of 1,877 ft (572 m) around the
breeding site is needed to encompass
the non-breeding habitat used by 90
percent of the frogs in a given
population (Rittenhouse and Semlitsch
2007, pp. 155–157).
During winter, northern leopard frogs
are thought to hibernate underwater in
ponds, in lakes, or on the bottom of
deeper streams or waters that do not
freeze to the bottom and that are welloxygenated (Nussbaum 1983, p. 181;
Stewart et al. 2004, p. 72). Northern
leopard frogs are intolerant of freezing
and of waters that have severely
reduced or complete loss of dissolved
oxygen. If these conditions occur during
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hibernation, death of northern leopard
frogs is likely (Rorabaugh 2005, p. 574).
Based upon their research in
Wisconsin, Hine et al. (1981) described
the ideal ‘‘breeding pond’’ as having the
following features:
(1) The pond or wetland site should
be located within approximately 1.0
mile (mi) (1.6 kilometers (km)) of
suitable overwintering habitat (larger
bodies of water) so that adults can find
the breeding habitat when they emerge
in the spring and juvenile frogs are able
to find overwintering sites in the fall.
(2) In the spring, the water depth
should be approximately 5 ft (1.5 m) or
more so that there is balance of open
water and vegetation cover.
(3) Emergent vegetation (such as
sedge, bulrush, and cattail) should occur
along at least two-thirds of the pond or
wetland to provide escape cover and
places to attach egg masses.
(4) The slope should be gradual to
promote habitat for emergent vegetation.
(5) Natural terrestrial habitats should
be maintained peripheral to wetlands
summer habitat for adults post-breeding,
for juvenile growth, and for dispersal or
movement corridors.
(6) Water should be relatively
permanent throughout the year, but
should dry every decade or so in order
to eliminate any predaceous fish that
become established.
Water quality and temperature are
important determinants of northern
leopard frog habitat. Because northern
leopard frogs have permeable skin,
which may transfer external
contaminants to its internal organs,
good (i.e., non-polluted) water quality is
important at breeding locations.
Chemical contamination of habitats can
result in malformations, population
declines, decreased growth rates,
reduced activity, and other impacts to
northern leopard frogs (Diana and
Beasley 1998, pp. 267–276).
Temperature plays an important role in
both the springtime migratory and
breeding behaviors of northern leopard
frogs (Merrell 1970, pp. 50–51; Merrell
1977, pp. 5–6, 9). When ambient air
temperature is greater than or equal to
50 degrees Fahrenheit (°F) (10 degrees
Celsius (°C)), northern leopard frogs
move from their overwintering sites to
their breeding sites (Merrell 1970, p.
50). The calling sites and areas where
egg masses are deposited are not
random and appear to be chosen based
upon temperature as these activities
tend to be located in the warmest
portions of breeding ponds (Merrell
1977, p. 6).
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Biology
As soon as males leave overwintering
sites, they travel to breeding ponds and
call in shallow water (Smith 2003, p.
13). Breeding typically occurs during a
short period in the spring beginning in
early April (Pace 1974, p. 92; Corn and
Livo 1989, p. 4); at higher elevations
and more northern latitudes, the onset
of breeding is late April to early May
(Corn and Livo 1989, p. 5; Gilbert et al.
1994, p. 467). Most northern leopard
frogs are sexually mature at age 2,
although the age of sexual maturity may
vary from age 1 to age 3 in any given
population depending upon
environmental conditions (Leclair and
Castanet 1987, p. 368; Gilbert et al.
1994, pp. 468–469). Male frogs attract
females by calling from specific
locations within a breeding pond when
temperatures are close to 68 °F (20 °C)
or more, with several males typically
calling together to form a chorus
(Merrell 1977, p. 7). Eggs are typically
laid within breeding habitats, 2 to 3
days following the onset of chorusing
(Corn and Livo 1989, p. 5). Eggs are laid
in non-acidic, shallow (4 to 26 in (10 to
65 cm)), still water that is exposed to
sunlight, and are usually attached to
emergent vegetation just below the
water surface (Merrell 1977, p. 6; Gilbert
et al. 1994, pp. 467–468; Pope et al.
2000, p. 2505). Egg masses may include
several hundred to several thousand
eggs (Corn and Livo 1989, pp. 6–7) and
are deposited in a tight, oval mass
(Rorabaugh 2005, p. 572). Time to
hatching is correlated with temperature
and ranges from 2 days at 81 °F (27 °C)
to 17 days at approximately 53 °F
(12 °C) (Nussbaum et al. 1983, p. 182).
Tadpoles are the ephemeral, feeding,
non-reproductive, completely aquatic
larvae in the life cycle of the frog
(McDiarmid and Altig 1999, p. 2). The
length of time required for
metamorphosis (the development of the
aquatic tadpole to a frog) is variable, and
depending upon temperature, may take
3 to 6 months from time of egg-laying
(Merrell 1977, p. 10; Hinshaw 1999, p.
105). Northern leopard frog tadpoles are
predominantly generalist herbivores
(plant eaters), typically eating attached
and free-floating algae (Hoff et al.1999,
p. 215); however they may feed on dead
animals (Hendricks 1973, p. 100). Adult
and subadult frogs are generalist
insectivores (insect eaters) that feed on
a variety of terrestrial invertebrates such
as insect adults, larvae, spiders, and
leeches (Merrell 1977, p. 15; Collier et
al. 1998, p. 41; Smith 2003, p. 12;
Rorabaugh 2005, p. 575). In addition,
adult northern leopard frogs have also
been known to prey upon small
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northern leopard frogs, birds, and
snakes (Merrell 1977, p. 15).
Status
Northern leopard frogs, like many
amphibian populations, are dynamic,
and their individual numbers may
naturally fluctuate in size within
populations. However, across the range
of the northern leopard frog,
information suggests that there is an
ongoing loss of populations throughout
the species’ range. The loss of
populations across the landscape is
what results in species’ declines (Green
2005, p. 29). Population declines of
northern leopard frogs are welldocumented in the western United
States and western Canada, but are also
documented rangewide (through the
Midwestern and Eastern United States),
as described below.
The most recent complete summary of
distributional and abundance patterns
of the northern leopard frog is from
Rorabaugh (2005, pp. 570–571), which
documents a substantial contraction of
the species’ range, especially in the
western two-thirds of the United States,
where widespread extirpations have
occurred. Other authors have also
compiled summary data indicating
population declines (e.g., Smith and
Keinath 2007, p. 14). Since the 1960s,
the northern leopard frog has
experienced significant declines and
losses throughout its range (Gibbs et al.
1971, p. 1028), particularly in the
western United States and western
Canada, and tends to become less
abundant the farther west one proceeds
(Corn and Fogelman 1984, p. 150; Hayes
and Jennings 1986, p. 491; Clarkson and
Rorabaugh 1989, p. 534; Corn et al.
1989, pp. 26–29; Koch and Peterson
1995, pp. 84–87; Corn et al. 1997, pp.
37–38; Weller and Green 1997, p. 323;
Casper 1998, p. 199; Hammerson 1999,
pp. 146–147; Leonard et al. 1999, p. 51;
Dixon 2000, p. 77; Smith 2003, pp. 4–
6; Jennings and Fuller 2004, pp. 125–
127; Werner et al. 2004, pp. 97–98;
Committee on the Status of Endangered
Wildlife in Canada 2009, p. v; Germaine
and Hays 2009, p. 537; Johnson et al.
2011, p. 557).
Based upon this and other
information, the northern leopard frog
appears to be declining, is considered
rare, or is locally extirpated from many
historical locations in Arizona,
California, Colorado, Idaho, Iowa,
Minnesota, Missouri, Montana,
Nebraska, Nevada, New Mexico,
Oregon, Texas, Utah, Washington,
Wisconsin, and Wyoming (Hayes and
Jennings 1986, p. 491; Stebbins and
Cohen 1995, p. 220; Johnson and Batie
1996; Bowers et al. 1998, p. 372; Casper
1998, p. 199; Lannoo 1998, p. xvi;
Mossman et al. 1998, p. 198; Smith
2003, pp. 4–6; Smith and Keinath 2004,
pp. 57–60; McCleod 2005, pp. 292–294;
Rorabaugh 2005, p. 571; Johnson et al.
61899
2011, p. 561). The species is nearly
extirpated from almost 100 percent of its
historical range in Texas, California,
Oregon, and Washington (Stebbins and
Cohen 1995, p. 220; McAllister et al.
1999, p. 15; Stebbins 2003, p. 235;
Germaine and Hays 2009, p. 537).
Table 1 lists current NatureServe
ranks for States and provinces in which
the northern leopard frog is known to
occur. NatureServe conservation status
assessment procedures have different
criteria, evidence requirements,
purposes, and taxonomic coverage than
the Federal Lists of Endangered and
Threatened Wildlife and Plants, and
therefore, these rankings may not
coincide with legal listing processes
(NatureServe 2008, p. 1). However, for
a species as widespread as the northern
leopard frog, the NatureServe rankings
aid in summarizing the relative risks
facing the northern leopard frog
throughout its range and are provided
here for this reason.
NatureServe lists Maryland and New
Jersey as States where the northern
leopard frog occurs. However, the
Maryland Department of Natural
Resources lists the northern leopard frog
as an introduced species that occurs in
one county (Maryland Department of
Natural Resources 2011, p. 2), and the
frog does not occur in New Jersey
(Gessner and Stiles 2001, pp. 1–9; New
Jersey Division of Fish and Wildlife
2006, pp. 1–2).
TABLE 1—NATURESERVE AND STATE, PROVINCE, AND TERRITORY RANKS FOR NORTHERN LEOPARD FROGS IN STATES
AND PROVINCES IT IS KNOWN TO OCCUR
[NatureServe 2011, p. 1]
Natural heritage program rank *
State, province, territory rank
Arizona ...............................................................
California ............................................................
Colorado .............................................................
S2 (Imperiled) ..................................................
S2 (Imperiled) ..................................................
S3 (Vulnerable) ................................................
Connecticut .........................................................
Idaho ...................................................................
Illinois ..................................................................
Indiana ................................................................
Iowa ....................................................................
Kentucky .............................................................
Maine ..................................................................
S2
S3
S5
S2
S5
S3
S3
Maryland .............................................................
S4 (Apparently Secure), introduced spp .........
Massachusetts ....................................................
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State, province, territory or sovereign nation
S3/S4 (Vulnerable/Apparently Secure) ............
Michigan .............................................................
Minnesota ...........................................................
Missouri ..............................................................
Montana ..............................................................
S5 (Secure) ......................................................
S4 (Apparently Secure) ...................................
S2 (Imperiled) ..................................................
S1/S3 (Critically Imperiled/Vulnerable) ............
Navajo Nation (NE Arizona, NW New Mexico,
SE Utah).
Nebraska ............................................................
Nevada ...............................................................
New Hampshire ..................................................
New Jersey .........................................................
S2 (Imperiled) ..................................................
Species of Greatest Conservation Need.
Species of Greatest Conservation Need.
Species of Greatest Conservation Need, Species of Special Concern.
Special Concern Species.
Species of Greatest Conservation Need.
Non-game Indicator Species.
Species of Greatest Conservation Need.
No ranking or status.
Species of Greatest Conservation Need.
Species of Greatest Conservation Need (Priority 3).
No ranking or status (considered an introduced species).
Species of Special Concern, Species of
Greatest Conservation Need.
Species of Greatest Conservation Need.
No ranking or status.
Species of Conservation Concern.
Species of Concern, Species of Greatest
Conservation Need.
Endangered.
S5 (Secure) ......................................................
S2/S3 (Imperiled/Vulnerable) ...........................
S3 (Vulnerable) ................................................
SNR (Unranked), species not present .............
At-Risk Species (Tier II).
Species of Conservation Priority.
Species of Concern.
Species not present.
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(Imperiled) ..................................................
(Vulnerable) ................................................
(Secure) ......................................................
(Imperiled) ..................................................
(Secure) ......................................................
(Vulnerable) ................................................
(Vulnerable) ................................................
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TABLE 1—NATURESERVE AND STATE, PROVINCE, AND TERRITORY RANKS FOR NORTHERN LEOPARD FROGS IN STATES
AND PROVINCES IT IS KNOWN TO OCCUR—Continued
[NatureServe 2011, p. 1]
State, province, territory or sovereign nation
Natural heritage program rank *
State, province, territory rank
New Mexico ........................................................
New York ............................................................
North Dakota ......................................................
Ohio ....................................................................
Oregon ................................................................
S1 (Critically Imperiled) ....................................
S5 (Secure) ......................................................
SNR (Unranked) ..............................................
SNR (Unranked) ..............................................
S1/S2 (Critically Imperiled/Imperiled) ..............
Pennsylvania ......................................................
Rhode Island ......................................................
South Dakota ......................................................
Texas ..................................................................
Utah ....................................................................
Vermont ..............................................................
Washington .........................................................
West Virginia ......................................................
Wisconsin ...........................................................
Wyoming .............................................................
Alberta ................................................................
British Columbia .................................................
Labrador and Newfoundland ..............................
Manitoba .............................................................
New Brunswick ...................................................
Northwest Territories ..........................................
Nova Scotia ........................................................
Ontario ................................................................
Prince Edward Island .........................................
Quebec ...............................................................
Saskatchewan ....................................................
S2/S3 (Imperiled/Vulnerable) ...........................
S2 (Imperiled) ..................................................
S5 (Secure) ......................................................
S1 (Critically Imperiled) ....................................
S3/S4 (Vulnerable/Apparently Secure) ............
S4 (Vulnerable) ................................................
S1 (Critically Imperiled) ....................................
S2 (Imperiled) ..................................................
S4 (Vulnerable) ................................................
S3 (Vulnerable) ................................................
S2/S3 (Imperiled/Vulnerable) ...........................
S1 (Critically Imperiled) ....................................
S3/S4 (Vulnerable/Apparently Secure) ............
S4 (Vulnerable) ................................................
S5 (Secure) ......................................................
SNR (Unranked) ..............................................
S5 (Secure) ......................................................
S5 (Secure) ......................................................
S4/S5 (Apparently Secure/Secure) ..................
S5 (Secure) ......................................................
S3 (Vulnerable) ................................................
Species of Greatest Conservation Need.
No ranking or status.
No ranking or status.
No ranking or status.
Sensitive Critical, List 2 Species (threatened
with extinction or presumed extinct).
Priority Conservation Species (Tier 5).
Species of Greatest Conservation Need.
No ranking or status.
No ranking or status (likely extirpated).
Species of Concern (Tier III).
No ranking or status.
Endangered.
Species in Greatest Need of Conservation.
No ranking or status.
Species of Greatest Conservation Need.
Threatened.
Endangered.
No ranking or status.
No ranking or status.
No ranking or status.
No ranking or status.
No ranking or status.
Not at risk.
No ranking or status.
No ranking or status.
Interim Species at Risk.
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* S1 = Critically Imperiled: At very high risk of extinction due to extreme rarity (often 5 or fewer populations), very steep declines, or other factors.
S2 = Imperiled: At high risk of extinction due to restricted range, few populations (often 20 or fewer), steep declines, or other factors.
S3 = Vulnerable: At moderate risk of extinction due to a restricted range, relatively few populations (often 80 or fewer), recent and widespread
declines, or other factors. Such species are often rare or found locally in a restricted range.
S4 = Apparently Secure: Uncommon but not rare; some cause for long-term concern due to declines or other factors. Such species are likely
to be quite rare in parts of their range, especially at the periphery.
S5 = Secure: Common; widespread and abundant. Such species are potentially rare in parts of their range, especially at the periphery.
SNR = Unranked. State or Province conservation status not yet assessed.
The International Union for the
Conservation of Nature’s ‘‘Red List
Categories and Criteria’’ were developed
for classifying species at high risk of
global extinction (IUCN 2003, p. 1), and
as such have different criteria, evidence
requirements, purposes, and taxonomic
coverage than the Federal Lists of
Endangered and Threatened Wildlife
and Plants. However, just as with the
NatureServe data, because we are
reviewing the entire range of the
northern leopard frog, the International
Union for the Conservation of Nature
assessment is useful in summarizing the
current status of the northern leopard
frog throughout its range.
The International Union for the
Conservation of Nature currently lists
the northern leopard frog as a species of
‘least concern’ in view of its wide
distribution, tolerance to degree of
habitat modification, and presumed
large population (Hammerson et al.
2004, p. 2). The International Union for
the Conservation of Nature states that
the population trend is decreasing
(Hammerson et al. 2004, p. 3), but the
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authors believe that the northern
leopard frog is not declining fast enough
to qualify for listing in a more
threatened category (Hammerson et al.
2004, p. 2). The International Union for
the Conservation of Nature reviewed
Hammerson et al. (2004, pp. 1–6) in
2011, and no updates were made to the
2004 review. Since 2004, Rorabaugh
(2005, pp. 570–577) completed a status
review for the northern leopard frog in
the United States (Rorabaugh 2005, pp.
570–577), and the Committee on the
Status of Endangered Wildlife in Canada
published the Assessment and Update
Status Report for the Northern Leopard
Frog in Canada (Committee on the
Status of Endangered Wildlife in Canada
2009, pp. 1–76). The Rorabaugh (2005,
pp. 570–577) status review found that
for a variety of reasons the northern
leopard frog is declining throughout its
range, but particularly in the western
United States. The Committee on the
Status of Endangered Wildlife in Canada
(2009, pp. iii) assessment notes that
there are continued declines for the
northern leopard frog throughout the
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western provinces and evidence of
declines in eastern Canada. The current
International Union for the
Conservation of Nature review does not
cite either of these documents or
provide any current threats assessment.
The International Union for the
Conservation of Nature analysis for the
northern leopard frog also includes
leopard frogs in Panama, which likely
belong to the Lithobates complex, but
do not belong to the same species as the
northern leopard frog. Therefore, we do
not consider the International Union for
the Conservation of Nature review for
the northern leopard frog a current
assessment of the species’ status in
North America.
Western States
Until the late 1970s, northern leopard
frogs were widespread and abundant in
much of northern Arizona (Apache,
Coconino, Greenlee, Mohave, Navajo,
and Yavapai Counties) in springs,
streams, rivers, stock tanks, and lakes
throughout northern Arizona (Arizona
Game and Fish Department 2009, p. 1).
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Currently, there is one northern leopard
frog population located near Seligman,
Arizona; a metapopulation (several
breeding locations in close proximity to
one another) located south of Flagstaff,
Arizona; and three refugial sites
developed by the State and Service (and
other partners) to assist in stocking
northern leopard frogs to other locations
in Arizona, north of the Colorado River.
All of these locations are located in
Coconino County. Outside of these
locations, fairly rigorous visual
encounter surveys conducted within the
species’ historical range, including
Grand Canyon National Park and the
Kaibab National Forest, have not located
northern leopard frogs (Kaibab National
Forest 2007, p. 1; Kaibab National Forest
2008, p. 1; Drost et al. 2008, p. 7). The
species is listed as a Species of Greatest
Conservation Need in the Arizona State
Wildlife Action Plan (Arizona Game and
Fish Department 2006, Appendix M, p.
153) and has a NatureServe rank of S2
(Imperiled) (NatureServe 2011, p. 1). In
Arizona, there is no open season for
northern leopard frog, and collecting is
illegal except as authorized by State
permit, effective January 1, 1993
(Commission Order 41). The northern
leopard frog has also significantly
declined on the Navajo Nation (which is
situated in southeastern Utah,
northeastern Arizona, and northwestern
New Mexico) in the last century. Most
remote desert populations of northern
leopard frogs were lost between the
1920s and 1970s, and mountain
populations were lost in the late 1980s.
The Navajo Nation has listed the
northern leopard frog as a ‘‘Group 2—
Endangered Species’’ on the Navajo
Endangered Species List, which means
its prospects of survival or recruitment
on the Navajo Nation are in jeopardy
(Navajo Nation Department of Fish and
Wildlife 2009, p. 3).
The northern leopard frog is a State of
California species of special concern
and is listed as a Species of Special
Concern (native populations only)
(California Department of Fish and
Game, Natural Diversity Database, 2009)
and as a Species of Greatest
Conservation Need in California
Department of Fish and Game’s State
Wildlife Action Plan (California
Department of Fish and Game 2007);
however, the northern leopard frog is
not listed under the California
Endangered Species Act. The northern
leopard frog may be taken under the
authority of a sport fishing license,
subject to restrictions (California Code
of Regulations, Title 14, Section 5.05).
The frog is ranked S2 (Imperiled) by
NatureServe (NatureServe 2011, p. 1).
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Northern leopard frogs are likely native
to the region east of the Sierra NevadaCascade crest in the following areas of
California: upper Pit River basin
(Shasta, Lassen, and Modoc counties),
Surprise Valley (Modoc County), lower
Klamath Lake basin (Siskiyou County),
Lake Tahoe region (El Dorado County),
Carson River drainage (Alpine County)
and Owens River Valley (Mono and
Inyo counties) (Jennings and Fuller
2004, p. 122). The northern leopard frog
was introduced to at least 15 other sites
in California, but most of these
introductions have not resulted in
naturalized populations that continue to
exist today (Jennings and Hayes 1994, p.
80; Jennings and Fuller 2004, p. 119).
There is a small, introduced population
in Merced County, near the Merced
National Wildlife Refuge (NWR) that
persisted as recently as 2007 (Jennings
and Fuller 2004, pp. 119, 127;
Woolington 2009, pers. comm.). Since
the 1970s, northern leopard frogs have
disappeared from most (approximately
95 percent) of their historic range in
California, (Jennings and Fuller 2004, p.
119; Rorabaugh 2005, p. 571) and may
be completely extirpated from these
areas of the State as we are not aware
of any recent confirmed sightings.
Jennings and Hayes (1994, p. 82) knew
of only two extant, native northern
leopard frog populations as of the 1990s:
one adult was observed at Tule Lake
National NWR (Siskiyou County) in
1990, and 8 to 10 juveniles were found
near Pine Creek in Round Valley near
Bishop (Inyo County) in 1994. Northern
leopard frogs are no longer found on
Tule Lake NWR (Adams 2011, pers.
comm.), and no northern leopard frogs
have been observed during amphibian
surveys conducted on the Klamath Falls
NWR Complex, including Tule Lake
NWR (Austin 2009, pers. comm.).
Recent surveys conducted by the
California Department of Fish and Game
did not locate any northern leopard
frogs in the Owens River Area (Becker
2011, pers. comm.). In addition, surveys
found that sites previously considered
to be northern leopard frog habitat now
contain nonnative aquatic species, and
the habitat has been extensively
modified such that there are likely few
areas of suitable habitat left in the
Owens Valley (Becker 2011, pers.
comm.). Northern leopard frogs have not
been found in the Lake Tahoe basin for
over 20 years, and the species is
presumed to be extirpated from the area
(Jennings and Fuller 2004, p. 125).
Jennings and Fuller (2004, p. 126) also
report that a formerly isolated native
northern leopard frog population on Hat
Creek, Shasta County, is now apparently
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extirpated as well. Modoc NWR in
northeastern California reported no
known occurrences of northern leopard
frogs on the refuge in recent times, and
no northern leopard frogs were reported
during numerous hours of amphibian
survey time in 2004, 2005, and 2010
(Bachman 2011, pers. comm.).
The northern leopard frog was
historically quite common throughout
Colorado, but over the last 30 to 40
years, populations have declined and
even been locally extirpated from
portions of eastern and north-central
Colorado, including Rocky Mountain
and Mesa Verde National Parks (Corn
and Fogleman 1984, p. 148; Corn et al.
1989, p. 15; Stebbins and Cohen 1995,
p. 220; Corn et al. 1997, pp. 37–38;
Hammerson 1999, pp. 146–147; Mesa
Verde National Park 2009, p. 1; Johnson
et al. 2011, p. 561). The Colorado
Division of Wildlife has designated the
northern leopard frog a Species of
Greatest Conservation Need as well as a
Species of Special Concern due to low
population status and a declining
population trend (Colorado Division of
Wildlife 2006, pp. 2, 28, 305). These are
not statutory categories; however, the
northern leopard frog is classified as
‘‘nongame’’ wildlife and their
harassment, taking, or possession is
prohibited without a permit (Colorado
Division of Wildlife 2009, p. 3).
NatureServe ranks the northern leopard
frog as S3 (Vulnerable) in Colorado
(NatureServe 2011, p. 1). Intensive
surveys conducted from 2007 through
2009 in the Front Range of Colorado
indicate that northern leopard frogs
there have become rare and documented
losses are widespread (Johnson and
McKenzie 2009, p. 9; Keeley 2009, pp.
5–6; Johnson et al. 2011, p. 562).
Historically, northern leopard frogs
were found at high densities in this
region (Johnson et al. 2011, p. 562).
Along the Western Slope (the area west
of the continental divide in Colorado),
data suggest that northern leopard frog
populations remain viable, especially in
the northern region (Johnson and
McKenzie 2009, p. 10). This supports
information from Arapaho and Browns
Park NWRs, both located in
northwestern Colorado, that continue to
support northern leopard frogs (Johnson
2009, pers. comm.; Smart 2009, pers.
comm.). Northern leopard frogs were the
most common amphibian in southwest
Colorado until the late 1960s, but now
they are rare (San Miguel 2009, pers.
comm.). Despite conducting amphibian
surveys for 15 years with an emphasis
on locating northern leopard frogs, none
have been detected within Mesa Verde
National Park, Colorado. Historically,
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this species was found abundantly along
the Mancos River in the park and
adjacent lands (San Miguel 2009, pers.
comm.). However, the overall status of
the northern leopard frog in western
Colorado is not currently known
(Johnson et al. 2011, p. 563).
The Idaho Department of Fish and
Game designated the northern leopard
frog a Type 2 Species of Greatest
Conservation Need (Idaho Department
of Fish and Game 2005, Appendix B p.
6). A Type 2 species of greatest
conservation need is defined as a
rangewide or globally imperiled species
that is experiencing significant declines
throughout its range with a high
likelihood of being listed in the
foreseeable future due to its rarity
(Idaho Department of Fish and Game
2005, Appendix B, p. 4). Reduced
distribution and a declining population
trend are noted in the Idaho
Comprehensive Wildlife Conservation
Strategy as reasons for the designation
(Idaho Department of Fish and Game
2005, Species Account, p. 1). The
northern leopard frog is also a protected
nongame species, which means take or
possession of the species is prohibited
without a permit (Idaho Administrative
Code 13.01.06–300.02). NatureServe
ranks the northern leopard frog in Idaho
as S3 (Vulnerable) (NatureServe 2011, p.
1). Both the Targhee National Forest and
Kootenai NWR have records of northern
leopard frogs from the 1970s (Service
1972, p. 11; Stebbins and Cohen 1995,
p. 220). However, surveys in 1992 at 98
sites on the Targhee National Forest did
not locate northern leopard frogs
(Stebbins and Cohen 1995, p. 220), and
Kootenai NWR has no records of frogs
for the last 30 years (Rose 2009, pers.
comm.). Deer Flat NWR amphibian
surveys have only detected American
bullfrogs (Lithobates catesbeiana).
Northern leopard frogs are known to be
present on Bear Lake, Grays Lake, and
Minidoka NWRs, and presumed to be
present on Camas NWR and Oxford
Slough Wetland Protection Area (WPA)
(Fisher and Mitchell 2009, p. 1).
Localized declines of northern
leopard frogs are documented in Iowa
(Lannoo et al. 1994, pp. 317–318;
Hemesath 1998, p. 216). Lannoo et al.
1994 (p. 311) states, ‘‘From descriptions
of the turn-of-the-century commercial
‘‘frogging’’ industry in Dickinson
County (Iowa), we estimate that the
number of leopard frogs has declined by
at least two, and probably three orders
of magnitude.’’ However, the northern
leopard frog is ranked as Secure (S5) in
Iowa by NatureServe (2011, p. 1) and is
not considered a Species of Greatest
Conservation Need (Iowa Department of
Natural Resources 2006, p. 42).
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Currently, there is a continuous open
season for northern leopard frogs in
inland and boundary waters in Iowa,
and up to 48 frogs can be collected per
day (Iowa Department of Natural
Resources 2011, p. 1). In 1991, the Iowa
Department of Natural Resources
initiated an annual anuran (frog and
toad) survey. The survey is conducted
by volunteers, and until 2007,
volunteers were not required to
distinguish between species of leopard
frogs on the report forms (Iowa
Department of Natural Resources 2009,
p. 1). Survey data from 2007 and 2008
(when the species were separated) and
older data from counties where it was
thought only the northern leopard frog
occurred were reviewed by the State.
The analyses of this information suggest
a possible downward trend in northern
leopard frog presence, but the trend was
not statistically significant (Iowa
Department of Natural Resources 2009,
p. 1).
Northern leopard frog populations
began declining in Minnesota in the late
1960s or early 1970s (Rittschof 1975, p.
103; Minnesota Department of Natural
Resources 2011a, pp. 1–2). The declines
of northern leopard frog populations
from the past are thought to have been
substantial, but information is not
detailed enough to know if the
population is now stable or if it is still
declining in Minnesota (Moriarty 1998,
p. 168). However, because the species is
still considered to be fairly common, it
is not considered a Species of Greatest
Conservation Need in Minnesota’s
Comprehensive Wildlife Strategy
(Minnesota Department of Natural
Resources 2006, Appendix B p. 9). The
Minnesota Department of Natural
Resources’ northern leopard frog fact
page does indicate that the northern
leopard frog is still declining
(Minnesota Department of Natural
Resources 2011a, p. 2). The species is
ranked S4 (Apparently Secure) by
NatureServe (NatureServe 2011, p. 1). In
Minnesota, from May 16 to March 31,
licensed anglers and children under age
16 may take, use, buy, and sell an
unlimited number of northern leopard
frogs up to 6 inches long for bait
(Minnesota Department of Natural
Resources 2011b, p. 70). A Minnesota
Department of Natural Resources
commercial license is required to take
northern leopard frogs for purposes
other than bait.
Missouri is located on the periphery
of the range for northern leopard frogs
and the frog is currently only known to
occur in two counties (Atchison and
Mercer) that border Iowa (Missouri
Department of Conservation 2009, p. 1).
The northern leopard frog is listed as a
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Species of Conservation Concern by the
Missouri Department of Conservation
and NatureServe ranks it as Imperiled
(S2) (Missouri Department of
Conservation 2009, p. 1; NatureServe
2011, p. 1). This ranking is based upon
the low number of known occurrences
in Missouri and not based upon
declining population trends (Missouri
Department of Conservation 2009, p. 1).
The Missouri Department of
Conservation noted that it is likely that
more populations are present in
northern Missouri, but further surveys
need to be completed to affirm this
assumption (Missouri Department of
Conservation 2009, p. 1). In Missouri,
northern leopard frogs have regulatory
protection from commercial take and
non-resident collection. Missouri
residents are allowed to possess up to
five northern leopard frogs for education
use (Wildlife Code Missouri 3CSR10–
9.110); however, these five individuals
cannot be sold, traded, shipped over
State lines, or taken from public lands
(Missouri Department of Conservation
2009, p. 2). Northern leopard frogs also
cannot be used as live bait in Missouri
(Wildlife Code Missouri 3CSR10–6.605).
Montana Fish, Wildlife, and Parks
classified the northern leopard frog as a
Species of Concern in Montana and it is
considered a Species of Greatest
Conservation Need in their Wildlife
Conservation Strategy (Montana Fish,
Wildlife, and Parks 2009, p. 1). Northern
leopard frogs are protected from
commercial collection in Montana
(Montana Code Annotated 2009 87–5–
116). Historically, northern leopard
frogs occurred across the eastern plains
of Montana and in the mountain valleys
on both sides of the Continental Divide
(Montana Fish, Wildlife, and Parks
2009, p. 1). However, since the 1990s,
most previously known northern
leopard frog populations on the west
side of the Continental Divide in
Montana are considered extirpated, and
there has been a clear range contraction
of northern leopard frogs (Werner 2003,
p. 26; Montana Fish, Wildlife, and Parks
2009, p. 1). Currently, only two
populations exist in western Montana.
Surveys in the mid-1990s of historically
occupied sites in central Montana, east
of the Continental Divide, found only 19
percent of the sites to be occupied by
northern leopard frogs (Montana Fish,
Wildlife, and Parks 2009, p. 1).
NatureServe provides a split rank for the
State that reflects the difference in
status between western (S1 Critically
Imperiled) and eastern (S3 Vulnerable)
Montana (NatureServe 2011, p. 1).
Habitat restoration and survey efforts
are being planned Statewide to provide
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a current assessment of northern
leopard frog distribution (Montana Fish,
Wildlife, and Parks, 2009, p. 2).
The northern leopard frog occurs
commonly in the State of Nebraska
(McLeod 2005, p. 292) and has a
NatureServe rank of S5 (Secure)
(NatureServe 2011, p. 1). However,
surveys conducted in 1997 and 1998
indicated a significant decline in
northern leopard frog occurrences at the
State level (McLeod 2005, p. 292). It is
difficult to ascertain if this information
represents a real decline or is
representative of normal stochastic
events, but data indicated significant
differences from location data collected
in the 1970s (McLeod 2005, p. 292). The
Nebraska Game and Parks Commission
identified the northern leopard frog as a
Tier II At-Risk Species during
development of the Nebraska Natural
Legacy Project (2005, p. 319). Tier II
species are typically those that are not
at-risk from a global or national
perspective, but are rare or imperiled
within Nebraska. As of 2011, northern
leopard frogs can no longer be
commercially harvested or sold for bait
in Nebraska; however, anglers can still
collect them as bait for personal use
(Nebraska Game and Parks Commission
2011, p. 5).
In Nevada, northern leopard frogs are
currently ranked S2/S3 (Imperiled/
Vulnerable) by NatureServe
(NatureServe 2011, p. 1) and are on the
Nevada Natural Heritage Program’s
Animal and Plant Watch List, which
means they could be declining in
Nevada or across much of their range, or
may be less common than currently
thought and could become at-risk in the
future. The northern leopard frog is
identified as a Species of Conservation
Priority in the Nevada Wildlife Action
Plan (Wildlife Action Plan Team 2006,
p. 61). In addition, the northern leopard
frog is a protected amphibian by Nevada
statute (NAC 503.075) and cannot be
collected for commercial, recreational,
or educational purposes without a
permit (Nevada Department of Wildlife
2009, p. 5). The Nevada Department of
Wildlife notes that there is little
historical or current information
available to accurately assess the
distribution and status of the northern
leopard frog in Nevada (Nevada
Department of Wildlife 2009, p. 1).
However, recent surveys suggest that
northern leopard frogs may no longer be
abundant in Nevada and that there have
been numerous local extirpations, for
example, along the Truckee and Carson
rivers in western Nevada and in springs
of southern and eastern Nevada (Panik
and Barrett 1993, p. 203; Hitchcock
2001, pp. 9, 109–110). While historical
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records and anecdotal evidence
indicated that northern leopard frogs
were once widely distributed in the
State, the current species distribution is
much smaller than the historical
distribution (Hitchcock 2001, pp. 9, 38,
48). In addition, suitable northern
leopard frog habitat is patchily
distributed in the State due to the
aridity and isolated nature of many
wetland systems, which results in a
discontinuous and limited distribution
(Nevada Department of Wildlife 2009, p.
1). Recent Nevada Department of
Wildlife records document northern
leopard frog populations in Ruby Valley
(including Ruby Lakes NWR) and Lower
Mary’s River in Elko and White Pine
Counties; Spring Valley and Lake Valley
in White Pine County; Lake Valley and
Pahranagat Valley (including Pahranagat
NWR) in Lincoln County; Carson River
near Carson City; the lower Truckee
River and Truckee meadows in Washoe
County; and a small number of
additional sites in western and
northeastern Nevada (Hitchcock 2001,
pp. 96–102; Service 2009, pp. 1–2;
Nevada Department of Wildlife 2009, p.
2). Efforts to restore northern leopard
frog habitat and re-establish the species
have occurred along the lower Truckee
River in western Nevada and on
Pahranagat NWR (Horton 2010, pers.
comm.; Rogers 2010, p. 7).
Historically, the northern leopard frog
was documented from a large area in the
northern and western part of New
Mexico and along the entire length of
the Rio Grande River valley, except
southern Elephant Butte and northern
Caballo Reservoirs (New Mexico
Department of Game and Fish 2009, p.
1). Declines in northern leopard frogs
have been reported from the Lower Rio
Grande (below Caballo Reservoir), in the
Jemez Mountains, and in the Chuska
Mountains (Christman 2009, p. 5; New
Mexico Department of Game and Fish
2009, p. 2). The species is believed to
be extirpated from the Rio Grande
Valley, south of Albuquerque (New
Mexico Department of Game and Fish
2009, p. 3). Recent survey efforts
indicate that northern leopard frogs are
persisting in northern New Mexico, but
most occupied sites contained small
numbers of frogs with very few robust
populations (Christman 2009, p. 13).
The northern leopard frog is not listed
as endangered or threatened in New
Mexico under the Wildlife Conservation
Act, but was designated a Species of
Greatest Conservation Need by the New
Mexico Department of Game and Fish,
and NatureServe ranks it as S1
(Critically Imperiled) in New Mexico
(New Mexico Department of Game and
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Fish 2006, p. 540; NatureServe 2011, p.
1). The northern leopard frog is
protected from commercial take (Section
17–1–14 NMSA); however, take by New
Mexico State residents for pets or other
uses are uncontrolled (New Mexico
Department of Game and Fish 2009, p.
2).
Historically, the northern leopard frog
ranged Statewide in North Dakota and is
still quite common today (North Dakota
Game and Fish Department 2009, p. 1).
Northern leopard frogs are widely
distributed throughout the State and
locally abundant in some locations
(Newman 2009, p. 1; Scherr 2009, pers.
comm.) but surveys conducted by
Bowers et al. (1998, p. 372) found that
the range of the northern leopard frog
was less extensive in the prairie
potholes region of North Dakota than
previously described. Because of its
distribution and local abundance, the
northern leopard frog has no special
status in the State, and there are no
conservation programs that specifically
target the northern leopard frog (North
Dakota Game and Fish Department
2009, p. 1). Commercial frog licenses are
available for unlimited collection of
northern leopard frogs (North Dakota
Administrative Code 30–03–04).
NatureServe does not have a current
ranking for North Dakota as it is
currently under review (NatureServe
2011, p. 1).
The Oregon Department of Fish and
Wildlife ranks the northern leopard frog
as a ‘‘Sensitive Critical’’ species,
meaning that it is imperiled with
extirpation from a specific geographic
area of the State due to small population
sizes, habitat loss or degradation, or
immediate threats (Oregon Biodiversity
Information Center 2010, p. 7, 13). The
sensitive species list is primarily a nonregulatory tool designed to provide a
voluntary, proactive approach to
conservation (Oregon Department of
Fish and Wildlife 2008, p. 1). The
Oregon Biodiversity Information Center
lists the northern leopard frog as a ‘‘List
2 Species’’ meaning that it is threatened
with extirpation or presumed to be
extirpated from the State of Oregon
(Oregon Biodiversity Information Center
2010, pp. 4, 13) and it is ranked S1/S2
(Critically Imperiled/Imperiled) by
NatureServe (NatureServe 2011, p. 1).
The Oregon Biodiversity Information
Center (2010, p. 13), lists the following
counties as containing historical
locations for the northern leopard frog:
Hood River, Wasco, Sherman, Gilliam,
Morrow, Umatilla, Jefferson, Crook,
Grant, Baker, Malheur, Klamath, and
Jackson Counties. Rorabaugh (2005, p.
571) reported that northern leopard
frogs are extirpated from most historical
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localities in Oregon. The six records we
have from the Oregon Natural Heritage
Information Center are observations
from 1975, 1980, 1990, 1995, 1996, and
2003. We have found no records,
current or historical, to indicate the
presence of northern leopard frogs on
either the Hart Mountain National
Antelope Refuge (southern Oregon) or
Sheldon NWR (northern Nevada)
(Harper Collins 2009, pers. comm.). Frog
surveys were conducted at Sheldon
NWR in summer 2009, but they detected
only nonnative American bullfrogs.
The status of the northern leopard
frog in South Dakota is thought to be
stable and NatureServe lists the frog as
secure (S5) (South Dakota Department of
Game, Fish, and Parks 2009, p. 1;
NatureServe 2011, p. 1). The northern
leopard has no specific protection in
South Dakota and can be collected for
commercial and non-commercial bait
(South Dakota Laws and Regulations for
Commercial Bait Dealers 2009, p. 1;
South Dakota Department of Game,
Fish, and Parks 2011, p. 23). The
species’ range includes almost the entire
State based upon historical and current
distribution maps (Fischer et al. 1999, p.
12; Naugle et al. 2005, p. 285). Smith et
al. (2005, p. 9) found northern leopard
frogs to be common in the Black Hills,
and a Statewide herpetology (amphibian
and reptile) survey report indicates that
the distribution of the northern leopard
frog in the State is stable (Backlund
2004, p. 8). However, there is no
historical or recent abundance data to
compare current survey data that would
indicate population trend (Backlund
2004, p. 9). Information received from
Lacreek and Waubay NWRs and the
Huron Wetland Management District
indicate northern leopard frogs are
prevalent (Flannders-Wanner 2009,
pers. comm.; Hubers 2009, pers. comm.;
Koerner 2009, pers. comm.). Anuran
auditory surveys (1997–1998) found
northern leopard frogs to be one of the
most widespread and wetland-abundant
species in eastern South Dakota (Naugle
et al. 2005, p. 290).
The northern leopard frog’s historic
range in Texas was in the Rio Grande
Valley, El Paso County (a relatively
small portion of the State). However,
extensive efforts to locate the frog have
been unsuccessful (Dixon 2000, pp. 42,
77). The northern leopard frog is ranked
S1 (Critically Imperiled) by NatureServe
(NatureServe 2011, p. 1), but is not
listed as a species of conservation
concern in the Texas Comprehensive
Wildlife Conservation Strategy (Texas
Parks and Wildlife Department 2005,
pp. 748–751). The Texas Parks and
Wildlife Department webpage (Texas
Parks and Wildlife Department 2011a, p.
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11) lists the species as occurring in
Texas, but the most current field guide
for amphibians and reptiles of Texas
indicates the species is likely extirpated
(Dixon 2000, p. 77). The Texas Parks
and Wildlife Department requires that
anyone who captures a wild animal,
including frogs, be licensed or permitted
(Texas Parks and Wildlife Department
2011b, p. 1).
The Utah Division of Wildlife
Resources considers northern leopard
frog populations in Utah to be secure
(Utah Division of Wildlife Resources
2009, p. 1). NatureServe ranks the
northern leopard frog as S3/S4
(Vulnerable/Apparently Secure)
(NatureServe 2011, p. 1). In Utah, the
northern leopard frog is classified as
‘‘controlled’’ for collection, importation,
and possession, and may only be
collected with a certificate of
registration (Administrative Rule R657–
53: Amphibian and Reptile Collection,
Importation, Transportation, and
Possession). Historically the northern
leopard frog is considered to be a wideranging species in Utah and is verified
to have occurred in all but Davis and
Wayne Counties (Utah Division of
Wildlife Resources 2009, p. 2). Utah’s
Wildlife Action Plan lists the northern
leopard frog as a Tier III Species of
Concern (Sutter et al. 2005, p. 5–6). Tier
III species are of conservation concern
because they are linked to at-risk
habitats, they have suffered significant
population declines, or there is little
information regarding the species. The
northern leopard frog was listed as a
species of concern due to lack of
information, water development, and
disease. In 2006, the Utah Division of
Wildlife Resources began compiling
survey information and conducting
surveys to determine the current
distribution of northern leopard frogs in
Utah. Recent surveys have documented
northern leopard frogs at 97 new sites
(not historical sites), for a total of 683
known sites in Utah (Utah Division of
Wildlife Resources 2009, p. 2). Of these
sites, 75 percent (512) are extant, and 25
percent (171) are considered historical,
as the observations occurred prior to
1989 (Utah Division of Wildlife
Resources 2009, p. 2). We do not have
information regarding how many of
these sites are breeding sites versus
other observations (such as dispersing
frogs).
The northern leopard frog was listed
in 2000 as an endangered species under
the Endangered, Threatened, and
Sensitive Species Classification
(Washington Administrative Code, Title
232, Chapter 12, Section 014) in
Washington State after surveys of 17
known historic locations confirmed
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occupancy at only two sites (Leonard et
al. 1999, p. 52; Germaine and Hays
2009, p. 537). ‘‘Endangered’’ in this
context means any wildlife species
native to the State of Washington that is
threatened with extinction throughout
all or a significant portion of its range
within the State. The northern leopard
frog is ranked S1 (Critically Imperiled)
in Washington State by NatureServe
(NatureServe 2011, p. 1). Historically,
the northern leopard frog occurred in
six major watersheds in eastern
Washington (Germaine and Hays 2009,
p. 537). However, extensive surveys
conducted at Gloyd Seeps and Potholes
Reservoir in 2002–2005 indicate that the
Gloyd Seeps population is likely no
longer a functional breeding population
and the Potholes Reservoir population is
in sharp decline (Germaine and Hays
2009, p. 542). Although inclement
weather prevented Washington
Department of Fish and Wildlife from
completing surveys in 2009, no
observations of northern leopard frogs
were made during what limited field
time was available (Washington
Department of Fish and Wildlife 2009,
p. 32).
The northern leopard frog is not
currently listed in Wisconsin, but over
the past several decades, declines have
been documented (Hine et al. 1981, pp.
2–3; Mossman et al. 1998, pp. 191–192,
198; Wisconsin Department of Natural
Resources 2009, p. 1). In 1981, the
Wisconsin Frog and Toad Survey began
to monitor several species, including the
northern leopard frog. The occurrence of
a species is determined by whether or
not the species is heard calling, and the
abundance is ranked by the relative
number of individuals heard calling at
a site (Kitchell and Hay 2007, p. 1).
Survey results from 1984 to 2007
indicate an overall decrease in the
estimated population trend for northern
leopard frogs (Kitchell and Hay 2007, p.
7). NatureServe ranks the northern
leopard frog as S4 (Secure) (NatureServe
2011, p. 1). In Wisconsin, northern
leopard frogs may be collected and
possessed in unlimited numbers if the
collector or possessor has a valid Class
A Captive Wild Animal Farm License or
a Commercial Bait License (Wisconsin
Department of Natural Resources 2011,
p. 13).
The northern leopard frog is
considered to be widely distributed in
Wyoming (Wyoming Game and Fish
Department 2009, p. 1). The Wyoming
Game and Fish Department identified
the species as a Species of Greatest
Conservation Need due to potential
habitat degradation and loss, disease,
absence of data, and contaminants
(Wyoming Game and Fish Department
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2005, p. 13). NatureServe ranks it as S3
(Vulnerable) (NatureServe 2011, p. 1).
Population declines have been
documented from the Laramie Plains,
Targhee National Forest, and Grand
Teton National Park (Baxter and Stone
1980, p. 44; Lewis et al. 1985, p. 167;
Koch and Peterson 1995, p. 85). No
population trend data are available for
northern leopard frogs in Wyoming.
Anecdotal reports and local survey
information indicate that the frog may
be common throughout eastern and
southwestern Wyoming (Wyoming
Game and Fish Department 2009, p. 1);
however, others reports indicate that the
present abundance of northern leopard
frogs in Wyoming is unknown and the
population trend is declining (Smith
and Keinath 2007, p. 14). The Wyoming
Game and Fish Department manages
commercial, scientific, and education
activities through their collection
permitting system (Wyoming Game and
Fish Department 2009, p. 3).
Eastern States
The northern leopard frog still occurs
throughout the eastern States it is
historically known from (Connecticut,
Illinois, Indiana, Kentucky, Maine,
Massachusetts, Michigan, New
Hampshire, New York, Ohio,
Pennsylvania, Rhode Island, Vermont,
and West Virginia) (Rorabaugh 2005, pp.
571–572). However, the frog currently
has a very disjunct distribution
throughout the northeast (New
Hampshire Fish and Game Department
2005, pp. A208–A209); some
populations are thought to be both
locally and regionally declining (Smith
and Keinath 2007, p. 14; Spriggs 2009,
p. 29), and, in some cases, local
extirpations have occurred (Rorabaugh
2005, p. 571; Spriggs 2009, p. 26). For
example, habitat loss from urban
development has resulted in local
extirpations in Connecticut,
Massachusetts, and Rhode Island
(Klemens 2000, p. 41; Rorabaugh 2005,
p. 571). Northern leopard frog declines
also occurred in the Midwest in
Michigan, Minnesota, and northeastern
Illinois in the late 1960s or early 1970s
(Rittschof, 1975, p. 103; Moriarty 1998,
p. 168; Mierzwa 1998, p. 117), and
although some populations have
recovered, others have not (Mierzwa
1998, p. 117; Moriarty 1998, p. 168).
In 1999, the Northeast Endangered
Species and Wildlife Diversity
Technical Committee published a list of
regional species of conservation
concern, which included the northern
leopard frog. The northern leopard frog
was added to the list based upon
declining populations or high risk of
disappearing from the Northeast, lack of
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data with suspicion of risk of
disappearing from the region, and
special circumstances (such as
vulnerability to collecting pressures)
(Therres 1999, p. 97).
Northeast Partners in Amphibian and
Reptile Conservation, using information
from State wildlife action plans and
other sources, developed the Northeast
Amphibian and Reptile Species of
Regional Responsibility and
Conservation Concern (Northeast
Partners in Amphibian and Reptile
Conservation 2010, pp. 2–3). Based
upon their analysis, the Northeast
Partners in Amphibian and Reptile
Conservation ranked the northern
leopard frog as a species of High
Concern and Regional Responsibility
that should be considered a target for
habitat and landscape-based
conservation initiatives (such as land
protection), may be an appropriate
indicator for long-term monitoring to
detect changes in distribution due to
climate change, and should be among
the highest priority species for
Northeast Partners in Amphibian and
Reptile Conservation to target
conservation efforts (e.g., create a
regional species working group)
(Northeast Partners in Amphibian and
Reptile Conservation 2010, pp. 3–5).
The ranking is based upon the number
of northeastern States that comprise a
species’ U.S. distribution and the
number of States that listed the species
in their Wildlife Action Plans. Based
upon their analysis, the northeastern
States make up less than 50 percent of
the northern leopard frog’s U.S.
distribution (occurs in 9 of 14
northeastern States), and it is listed as
a Species of Greatest Conservation
Concern in 6 of the 9 States it inhabits
(Northeast Partners in Amphibian and
Reptile Conservation 2010, p. 5).
In Connecticut, the northern leopard
frog is locally common along sections of
the Connecticut River and its tributaries
(the Farmington, Scantic, and
Coginchaug Rivers) (Klemens 2000, p.
40). Historical records of northern
leopard frog distribution indicate that
the frog was once widespread; current
information indicates that the northern
leopard frog no longer is found in some
of these areas (Klemens 2000, p. 41).
The northern leopard frog is considered
a ‘‘Special Concern’’ species under
Connecticut’s State Endangered Species
Act (Connecticut Department of
Environmental Protection 2005,
Appendix 1–b p. 18), and the
NatureServe rank is S2 (Imperiled)
(NatureServe 2011, p. 1). There is no
open season for taking northern leopard
frogs in Connecticut (Title 26 Fisheries
and Game, Department of
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13).
Northern leopard frogs experienced a
die-off in the 1960s or early 1970s in
northeastern Illinois, but have since
recovered in localized areas where
extensive wetland habitat still occurs
(Mierzwa 1998, p. 117). The northern
leopard frog is less common in areas
where significant wetland loss has
occurred (Mierzwa 1998, p. 117).
Statewide, the northern leopard frog is
considered to be abundant with a stable
and secure population trend in Illinois
(S5 (Secure) ranking from NatureServe)
(Smith and Keinath 2007, p. 14;
NatureServe 2011, p. 1). However, most
amphibian sampling efforts in Illinois
have been largely opportunistic, and
data are likely insufficient to accurately
determine changes in distribution and
abundance of species such as the
northern leopard frog (Illinois
Department of Natural Resources 2005,
p. 102). The Illinois Comprehensive
Wildlife Conservation Plan and Strategy
identified the northern leopard frog as a
non-game indicator species for
improving wetland habitat (Illinois
Department of Natural Resources 2005,
p. 172). It is unlawful to take, possess,
buy, sell, offer to buy or sell or barter
any reptile, amphibian, or their eggs or
parts taken from the wild in Illinois for
commercial purposes unless otherwise
authorized by statute (17 Illinois Adm.
Code Section 880–10). If a person
possesses a valid fishing license, they
may take up to eight northern leopard
frogs per day (17 Illinois Adm. Code
Section 880–20, 880–30).
The northern leopard frog’s range in
Indiana includes northern and eastern
Indiana. Minton (1998, pp. 217–220)
noted significant declines in the
northern leopard frogs populations
based on observations he made from
1948 to 1993 throughout Indiana. The
species is listed as a Species of Greatest
Conservation Need in the Indiana
Comprehensive Wildlife Strategy, listed
as a Species of Special Concern by the
Indiana Department of Natural
Resources, and is ranked as Imperiled
(S2) by NatureServe (Indiana
Department of Natural Resources 2006,
p. 30; NatureServe 2011, p. 1). In
Indiana, an individual with a valid
hunting or fishing license may collect
up to four northern leopard frogs for
non-commercial purposes (Indiana
Department of Natural Resources 2011,
p. 11).
The northern leopard frog is known
historically from 22 counties in
northern Kentucky (Kentucky
Department of Fish and Wildlife
Resources 2010, Amphibian Species
Accounts, Northern leopard frog).
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However, the species is considered to be
decreasing in Kentucky, and
populations have declined throughout
the frog’s historical State range.
Kentucky Department of Fish and
Wildlife Resources’ recent survey
records (1984–2004) show northern
leopard frogs persisting in 10 counties,
and no longer present in 12 counties
(Kentucky Department of Fish and
Wildlife Resources 2010, Amphibian
Species Accounts, Northern leopard
frog). The species is considered to be a
Species of Greatest Conservation Need
and ranked by NatureServe as
Vulnerable (S3) (Kentucky Department
of Fish and Wildlife Resources 2010,
Appendix 1–1 p. 6; NatureServe 2011,
p. 1). The northern leopard frog may be
collected for personal bait use in
Kentucky (301 Kentucky Administrative
Regulations 1:130).
The northern leopard frog is a Species
of Special Concern in Maine (Maine
Department of Inland Fisheries and
Wildlife 2005, p. 28) and is listed as a
Priority 3 Species of Greatest
Conservation Need in the
Comprehensive Wildlife Conservation
Strategy (Maine Department of Inland
Fisheries and Wildlife 2005, p. 90). The
Maine Department of Inland Fisheries
and Wildlife chose this ranking due to
the low to moderate potential for the
northern leopard frog to become
extirpated in the State, but concerns
remain regarding restricted distribution,
status, or extreme habitat specialization.
Currently, the present abundance and
population trend for the northern
leopard frog in Maine are unknown
(Smith and Keinath 2007, p. 14), and
NatureServe ranks the species as S3
(Vulnerable) (NatureServe 2011, p. 1). A
wildlife or fish possession permit is
required from the Commissioner to take,
possess, or hold in captivity northern
leopard frogs (Maine Department of
Inland Fisheries and Wildlife 2009,
p. 1).
The northern leopard frog occurs
Statewide in Massachusetts, except in
Barnstable, Dukes, and Nantucket
Counties (Massachusetts Division of
Fisheries and Wildlife 2006, p. 406).
Due to the widespread release of captive
northern leopard frogs, their historical
distribution and native status in
Massachusetts is uncertain (Cardoza and
Mirick (2002) in Massachusetts Division
of Fisheries and Wildlife 2006, p. 406).
As part of the Massachusetts Audubon
Herp Atlas Project (1992 through 1998),
the northern leopard frog was reported
to be well-distributed and confirmed
from approximately 13 percent of the
quadrants (Massachusetts Division of
Fisheries and Wildlife 2006, p. 406).
Though the northern leopard frog is not
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listed in Massachusetts (Massachusetts
Division of Fisheries and Wildlife 2006,
p. 107), because its status in the State is
unclear, it is a species of regional
conservation concern, a Species of
Special Concern, and a Species of
Greatest Conservation Need in the
Massachusetts Comprehensive Wildlife
Conservation Strategy (Massachusetts
Division of Fisheries and Wildlife 2006,
pp. 137, 274, 292, 343, 348). There is a
closed season on the hunting, fishing,
taking and possession of northern
leopard frogs in Massachusetts
(Massachusetts Division of Fisheries
and Wildlife 2002, p. 1). NatureServe
ranks the northern leopard frog in
Massachusetts as S3/S4 (Vulnerable/
Apparently Secure) (NatureServe 2011,
p. 1).
The Michigan Department of Natural
Resources describes the northern
leopard frog’s distribution in Michigan
as unknown, but considered patchy, and
notes that it appears to be declining
based upon the lack of reports compared
to historical records from the current
Frog and Toad Surveys (Eagle et al.
2005, Species of Greatest Conservation
Need, p. 152; Smith and Keinath 2007,
p. 14). The northern leopard frog is a
Species of Greatest Conservation Need
in Michigan’s Wildlife Action Plan
(Eagle et al. 2005, p. 20 in Aquatic
Threats by Species of Greatest
Conservation Need), but is ranked by
NatureServe as S5 (Secure) (NatureServe
2011, p. 1). In Michigan, an all-species
fishing license is required to take
northern leopard frogs for personal bait
use (Michigan Department of Natural
Resources 2011, p. 9).
The northern leopard frog is a Species
of Concern in New Hampshire and
ranked as S3 (Vulnerable) by
NatureServe (2011, p. 1). Possession of
northern leopard frogs in New
Hampshire is prohibited without a
permit (New Hampshire Fish and Game
Department 2011, p. 1). Distribution
records from 1992 to 2004 were verified
for Coos, Merrimack, Rockingham, and
Sullivan Counties; reports from a
number of other towns have not been
verified with a voucher photograph or
specimen (New Hampshire Fish and
Game Department 2005, p. A–209).
Throughout the area that the ranges of
northern leopard frogs and pickerel
frogs (Lithobates palustrus) overlap, it is
important to verify distribution records
via a photograph or a specimen as
northern leopard frogs are commonly
confused with pickerel frogs. New
Hampshire is the only State we found
that appears to require this information
for distribution records. Based upon this
information, it is likely that the current
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distribution of northern leopard frogs in
New Hampshire is unknown.
The northern leopard frog is not
identified as species of greatest
conservation need or a species of
concern in the Comprehensive Wildlife
Conservation Strategy for New York
(New York Department of
Environmental Conservation 2005, p.
73), and NatureServe (2011, p. 1) ranks
the northern leopard frog as S5 (Secure).
Persons holding a freshwater fishing
license or combined hunting and fishing
license (including those entitled to fish
without a license) may take northern
leopard frogs for personal bait use
(except in New York City, Suffolk
County, and Nassau County), and frogs
may be imported, bought, and sold at
any time (New York Department of
Environmental Conservation 2010, pp.
10–11, 16). The northern leopard frog
distribution map for New York shows it
having a very wide distribution
throughout the State (New York
Department of Environmental
Conservation 2011, p. 1), but local
herpetologists have reported declines
throughout New York (O’Donnell 2011,
pers. comm.). It is likely that the current
abundance and population trends for
northern leopard frogs in New York are
unknown (Smith and Keinath 2007,
p. 14).
The northern leopard frog is broadly
distributed throughout Ohio and is
considered to be secure by the Ohio
Department of Natural Resources,
Division of Wildlife (2005, pp. 125, 138,
143) and other sources (Smith and
Keinath 2007, p. 14). Currently,
NatureServe does not have a ranking for
Ohio (NatureServe 2011, p. 1). In Ohio,
a permit is required to possess northern
leopard frogs (Ohio Revised Code
1531.02). Walker (1946, p. 88) described
the northern leopard frog as being one
of the most abundant frogs in Ohio. It
is still considered to be locally
abundant, but it does appear to be
declining where wetlands have been
drained. The range appears to be
contracting in the southeastern counties
where extensive field efforts have
yielded few recent records (Ohio Frog
and Toad Calling Survey 2011, p. 1).
The current distribution, abundance,
and population trend for northern
leopard frogs in Pennsylvania is
unknown (Smith and Keinath 2007, p.
14; Gipe 2011, pers. comm.). The
Comprehensive Wildlife Conservation
Strategy states that there has been a
reduction in the northern leopard frog’s
range, and although it was previously
common in Pennsylvania and the
northeast, it is suspected that it has
significantly declined in recent years
(Pennsylvania Game Commission and
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Federal Register / Vol. 76, No. 193 / Wednesday, October 5, 2011 / Proposed Rules
Pennsylvania Fish and Boat
Commission 2005, p. 10–41). The
northern leopard frog is considered a
Priority Conservation Tier 5 Species,
and the need for a long-term monitoring
program is identified (Pennsylvania
Game Commission and Pennsylvania
Fish and Boat Commission 2005, p. 10–
41). This conservation priority tier
represents species that are fairly secure
in Pennsylvania, but for which the
Pennsylvania Biological Survey
recommends some level of management
attention. NatureServe (2011, p. 1) ranks
the northern leopard frog in
Pennsylvania as S2/S3 (Imperiled/
Vulnerable). The collection of one
northern leopard frog per day from
Pennsylvania waters requires a fishing
license, but a license is not required to
take a frog from land (Pennsylvania Fish
and Boat Commission 2011, pp. 1–2).
The northern leopard frog is a Species
of Greatest Conservation Need and
ranked by NatureServe as S2 (Imperiled)
in Rhode Island (Rhode Island
Department of Environmental
Management, Division of Fish and
Wildlife 2005, p. 24; NatureServe 2011,
p. 1). Rhode Island currently has one
small population of northern leopard
frogs on an island; several other
populations have been extirpated in
recent years (O’Donnell 2011, pers.
comm.). The removal from the wild, for
any purposes, of northern leopard frogs
is prohibited in Rhode Island, except by
special permit (Rhode Island
Department of Environmental
Management, Division of Fish and
Wildlife 2011, p. 38).
The Vermont Fish and Wildlife
Department considers the northern
leopard frog to be secure in Vermont
(Kart et al. 2005, p. 1 Secure Species
Summary; NatureServe 2011, p. 1). The
species is distributed along the western
edge of Vermont and then scattered
populations are documented throughout
the rest of the State (Kart et al. 2005,
Distribution Map). Collection of
northern leopard frogs for scientific
research, education purposes, or for the
purpose of using them as the subjects of
art or photography is authorized
through issuance of a scientific
collection permit; other collections or
take are authorized by Commissioner
Letter with a valid hunting license
(Vermont Fish and Wildlife Regulations
Title 10, Chapter 1, Section 25).
The West Virginia Natural Heritage
Program and NatureServe list a State
rank of S2 (Imperiled) for the northern
leopard frog (West Virginia Natural
Heritage Program 2007, p. 11;
NatureServe 2011, p. 1). The species is
also listed as a Species in Greatest Need
of Conservation (West Virginia Division
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of Natural Resources 2005, pp. 4F–
Habitats-20, 5F–49, 5F–56). Statewide
surveys were conducted between March
2008 and April 2009 to determine the
status and distribution of northern
leopard frogs in West Virginia (Spriggs
2009, p. 17). Surveys of 70 sites found
only four occupied sites and only one of
the sites constituted a breeding
population (only single adult or juvenile
frogs were located at the three other
locations) (Spriggs 2009, pp. 38–39). In
2010, surveyors searched for northern
leopard frogs at the known breeding
population at Greenbottom Wildlife
Management Area, West Virginia
(including one day with four
experienced surveyors), and found only
one dead northern leopard frog
(O’Donnell 2011, pers. comm.). Based
upon Statewide survey data collected,
Spriggs (2009, p. 29) recommended that
the northern leopard frog NatureServe
rank be changed to S1 (Critically
Imperiled).
Canada
Historically, the northern leopard frog
ranged across Canada from British
Columbia to Nova Scotia. Canada
represents approximately half of the
current range of the northern leopard
frog based on an estimation of land area
in the United States and Canada. Within
Canada, the northern leopard frog’s
range includes small to large portions of
the area within the Northwest
Territories, British Columbia, Alberta,
Saskatchewan, Manitoba, Ontario,
Quebec, New Brunswick, Nova Scotia,
Prince Edward Island, and
Newfoundland. The distribution of
northern leopard frogs in western
Canada is more closely tied to major
river drainages than is the species’
distribution in eastern Canada (Seburn
and Seburn 1998, p. 9).
The northern leopard frog is
uncommon in the Northwest Territories
and is historically known from nine
sites (Fournier 1997, p. 104). These
historical locations encompass a small
area between the northern borders of
Alberta and Saskatchewan and the
southern border of Great Slave Lake
(Weller and Green 1997, p. 323). Since
1980, a few frogs have been reported
from three sites (Seburn and Seburn
1998, p. 6). The northern leopard frog is
considered rare within this restricted
range, and a lack of data precludes any
determination of a population trend
(Fournier 1997, p. 104). The northern
leopard frog is not ranked in the
Northwest Territories by NatureServe
(NatureServe 2011, p. 1).
In British Columbia, the northern
leopard frog historically occurred in the
Kootenay and Columbia River valleys
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and in the Rocky Mountains east of
Fernie (Seburn and Seburn 1998, p. 6).
Currently, there is one native northern
leopard frog population remaining at the
Creston Valley Wildlife Management
Area (estimated population less than 60
adults), plus one introduced population
that has likely been extirpated
(Committee on the Status of Endangered
Wildlife in Canada 2009, pp. 42–43).
The British Columbia (or Rocky
Mountain) population is listed as
Endangered under the Species at Risk
Act (Statues of Canada 2002, c.29),
which provides protection similar to
that of the Endangered Species Act in
the United States. The northern leopard
frog is also on the provincial Red List
and is listed as Endangered under
British Columbia’s Wildlife Act
(Revised Statutes of British Columbia
1996, c. 488). The northern leopard frog
is ranked as critically imperiled (S1)
(NatureServe 2011, p. 1) in British
Columbia.
Historically, northern leopard frogs
were widely distributed and locally
abundant in central and southern
Alberta, and in the extreme northeastern
region of the province (Alberta Northern
Leopard Frog Recovery Team 2005, p.
3). Beginning in 1979, the northern
leopard frog disappeared suddenly from
much of its range in Alberta (Roberts
1992, p. 14; Seburn and Seburn 1998, p.
10). All previously known populations
in central Alberta are no longer present,
and to the south, populations have
disappeared or are restricted to small,
fragmented habitats with limited
opportunity for dispersal (Roberts 1992,
p. 14). In 1990–1991 and 2000–2001,
province-wide surveys were conducted
to determine the distribution of
northern leopard frogs in Alberta. In the
first survey, 24 sites were found to be
occupied; the more recent survey found
that of 269 historical sites surveyed,
only 54 supported northern leopard
frogs (Alberta Northern Leopard Frog
Recovery Team 2005, p. 4). Currently,
the northern leopard frog is thought to
occur in about 20 percent of historically
occupied areas in Alberta (Wilson et al.
2008, p. 864), and the NatureServe
ranking is S2/S3 (imperiled/vulnerable)
(NatureServe 2011, p. 1). The species is
listed as Threatened under Alberta’s
Wildlife Act (Revised Statutes of Alberta
2000, Chapter W–10), and a recovery
plan was prepared in 2005 (Alberta
Northern Leopard Frog Recovery Team
2005).
Historically, northern leopard frogs
were considered to be widespread and
abundant in Saskatchewan (Seburn
1992, p. 18). However, the northern
leopard frog experienced significant
declines in the 1970s and is now absent
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throughout most of its historical range
(Didiuk 1997, p. 112; Weller and Green
1997, p. 323). Currently, the number of
northern leopard frog populations in
Saskatchewan is unknown, and there is
no data to evaluate the population
trends (Didiuk 1997, p. 112). Anecdotal
information indicates that populations
may be recovering (Seburn 1992, pp.
17–18), but declines and die-offs have
also been reported and the overall
population status is unknown
(Committee on the Status of Endangered
Wildlife in Canada 2009, p. 29). The
current range of the northern leopard
frog within Saskatchewan is thought to
be discontinuous, and the majority of
occurrences are in the very southern
portion of the province (Saskatchewan
Conservation Data Center 2006, p. 1).
The northern leopard frog is currently
on Saskatchewan’s Interim Species at
Risk List (Wildlife Act 1998, Chapter
W–13.12), and is protected in provincial
and national parks (Committee on the
Status of Endangered Wildlife in Canada
2009, p. vi). The NatureServe rank for
the northern leopard frog in
Saskatchewan is S3 (Vulnerable)
(NatureServe 2011, p. 1).
In Manitoba, northern leopard frogs
suffered a significant die-off from 1975–
1976, and within a year were absent
from previously known population
cores (Koonz 1992, p. 19; Committee on
the Status of Endangered Wildlife in
Canada 2009, p. 29). Since this time,
populations have increased in some
areas and remained extremely low in
others (Koonz 1992, p. 20). Northern
leopard frogs are not monitored in
Manitoba and the current number and
distribution of extant populations is not
known (Committee on the Status of
Endangered Wildlife in Canada 2009, p.
29). The current NatureServe rank for
the northern leopard frog in Manitoba is
S4 (secure) (NatureServe 2011, p. 1).
The northern leopard frog is thought
to be common, widespread, and secure
throughout southern and central
Ontario, with sparse distribution in the
north (Weller and Green 1997, p. 323;
NatureServe 2011, p. 1). The species is
currently listed as ‘‘Not at Risk’’ under
the Ontario Endangered Species Act of
2007 (Statutes of Ontario 2007, Chapter
6) and under the Canadian Species at
Risk Act (Ontario Nature 2011, p. 2).
However, as with many parts of Canada,
northern leopard frog populations have
declined precipitously, particularly in
northern and southwestern Ontario
(Hecnar 1997, p. 9; Seburn and Seburn
1998, p. 10; Committee on the Status of
Endangered Wildlife in Canada 2009, p.
29; Desroches et al. 2010, pp. 308–309).
Although the widespread declines of the
1970s did not occur in Ontario as they
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did in the provinces to the west,
relatively recent mass mortality events
resulting from ranavirus have been
documented in Ontario (Greer et al.
2005, p. 11; Committee on the Status of
Endangered Wildlife in Canada 2009, p.
29). A 4-year study in the eastern and
central regions of the province found
declines of 23 percent (1992–1993) and
5 percent (1993–1994) in abundance of
northern leopard frogs (Hecnar 1997,
pp. 9, 11; Committee on the Status of
Endangered Wildlife in Canada 2009, p.
29). Regional declines of northern
leopard frogs have also been
documented in southern Ontario,
including the southern Great Lakes
Region (Committee on the Status of
Endangered Wildlife in Canada 2009,
pp. 29–30). Hecnar (1997, p. 11) notes,
‘‘Anecdotal reports suggest that R.
pipiens is the most abundant frog in the
Essex Plain. During this study (1992–
1993), R. pipiens declined in occurrence
across all regions of southwestern
Ontario.’’
The northern leopard frog is widely
distributed throughout the southern
region of Quebec, with sparse
populations in the central region of the
province (Weller and Green 1997, p.
323). Weller and Green (1997, p. 323)
note that there is no evidence of historic
or recent declines in Quebec, but Gilbert
et al. (1994, p. 468) found lower
densities of northern leopard frog egg
masses than reported in Wisconsin and
anecdotal declines of northern leopard
frogs in the Richelieu River system of
Quebec. Bonin (1992, p. 24) states that
trends in northern leopard frog
populations in Quebec are not known
based upon data collected for the
Amphibian and Reptile Atlas. In
addition, Desroches et al. (2010, pp.
308–309) found that the northern
leopard frog was uncommon on the
Quebec side of James Bay.
In New Brunswick, the northern
leopard frog is distributed throughout
the province and populations are
thought to be secure (S5 NatureServe
rank) (McAlpine 1997, p. 123; Weller
and Green 1997, p. 323; NatureServe
2011, p. 1). The northern leopard frog
occurs throughout mainland Nova
Scotia and Cape Breton Island and is
considered to be secure (S5 NatureServe
rank) with no evidence of declines
(Weller and Green 1997, p. 323;
NatureServe 2011, p. 1). On Prince
Edward Island, the northern leopard
frog status is apparently secure (S4) or
secure (S5) (NatureServe 2011, p. 1).
In Newfoundland, the northern
leopard frog was introduced to the
western side of the island on several
occasions, but is no longer present
(Buckle 1971, p. 74; Maunder 1997, p.
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94). The species is at the edge of its
range in Labrador, but occurs in a few,
discrete locations that are apparently
secure (Committee on the Status of
Endangered Wildlife in Canada 2009, p.
30; NatureServe 2011, p. 1).
Summary
In summary, the northern leopard frog
appears to be absent or declining
throughout a large portion of its
historical and current range in the
western United States and western
Canada (Rorabaugh 2005, pp. 570–571).
The species generally tends to be more
abundant and more secure in the eastern
portion of its range, but there are
indications that local, and possibly
regional, declines may also be occurring
in the eastern United States (such as in
Connecticut, Indiana, Kentucky, Maine,
Massachusetts, Michigan, New
Hampshire, Rhode Island, and West
Virginia) as well. Historically, regional
declines in the western United States
and Canada occurred in the 1960s
through 1970s, and since this time the
northern leopard frog has either not
recovered in many of these areas (such
as in Alberta, Arizona, British
Columbia, Colorado, Idaho, western
Montana, Nevada, New Mexico, Oregon,
Texas, Washington, and western
Wyoming) or the status of that recovery
is unknown due to a lack of information
regarding changes in the number of sites
occupied across the species’ range over
time (such as in Manitoba, Minnesota,
Saskatchewan, and Utah). Occupancy
trend data are also lacking throughout
much of the western and eastern
portions of the northern leopard frog’s
range where the northern leopard frog’s
status appears to be stable or where it
is unknown (such as in Iowa, Illinois,
Nebraska, New York, North Dakota,
Ontario, Pennsylvania, South Dakota,
and Wisconsin), and as such, the overall
range status is likely unknown.
However, despite the lack of occupancy
trend data, information indicates that in
the eastern United States and eastern
Canada, the northern leopard frog is still
widespread and relatively common.
Distinct Vertebrate Population Segment
We consider a species for listing
under the Act if available information
indicates such an action might be
warranted. ‘‘Species’’ is defined by the
Act as including any subspecies of fish
or wildlife or plants, and any distinct
population segment (DPS) of any
species of vertebrate fish or wildlife that
interbreeds when mature (16 U.S.C.
1532(16)). We, along with the National
Marine Fisheries Service (now the
National Oceanic and Atmospheric
Administration—Fisheries), developed
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the Policy Regarding the Recognition of
Distinct Vertebrate Population Segments
(61 FR 4722; February 7, 1996), to help
us in determining what constitutes a
DPS. The policy identifies three
elements that are to be considered
regarding the status of a possible DPS.
These elements include: (1) The
discreteness of the population segment
in relation to the remainder of the
species to which it belongs; (2) the
significance of the population segment
to the species to which it belongs; and
(3) the population segment’s
conservation status in relation to the
Act’s standards for listing (i.e., is the
population segment, when treated as if
it were a species, is endangered or
threatened?) (61 FR 4722; February 7,
1996). The first two elements are used
to determine if a population segment
constitutes a valid DPS. If it does, then
the third element is used to consider
whether such DPS warrants listing. In
this section, we will consider the first
two criteria (discreteness and
significance) to determine if the western
northern leopard frog is a valid DPS
(i.e., a valid listable entity). Our policy
further recognizes it may be appropriate
to assign different classifications (i.e.,
threatened or endangered) to different
DPSes of the same vertebrate taxon (61
FR 4722; February 7, 1996).
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Discreteness
Under the DPS policy, a population
segment of a vertebrate species may be
considered discrete if it satisfies either
one of the following two conditions:
(1) It is markedly separated from other
populations of the same taxon as a
consequence of physical, physiological,
ecological, or behavioral factors.
Quantitative measures of genetic or
morphological discontinuity (separation
based on genetic or morphological
characters) may provide evidence of this
separation.
(2) It is delimited by international
governmental boundaries within which
differences in control of exploitation,
management of habitat, conservation
status, or regulatory mechanisms exist
that are significant in light of section
4(a)(1)(D) of the Act.
Marked Separation
In our evaluation of discreteness
under the DPS policy, we primarily
used the results of two recent genetic
studies (Hoffman and Blouin 2004a, pp.
145–159; O’Donnell et al. 2011, pp. 1–
11) to evaluate whether any populations
of the northern leopard frog should be
considered markedly separate. We based
our determination on these two studies
because they provided comprehensive
data on the genetic variation across the
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range of the species. The petition to list
a ‘‘western DPS’’ of the northern leopard
frog was mainly based on the genetic
information and conclusions from the
study by Hoffman and Blouin (2004a).
There has since been an additional
genetic study conducted on the species
by O’Donnell et al. (2011) that we also
used in this 12-month finding. We
found no other relevant information
regarding the other factors to consider in
evaluating population discreteness,
such as physical, physiological,
ecological, or behavioral factors, or
morphological characters. We therefore
focused our analysis on these two
genetic studies in determining whether
the best available information supports
that there are discrete populations of the
northern leopard frog that would be
considered markedly separate under our
DPS policy.
Hoffman and Blouin (2004a) reported
two different lineages (lines of descent
from a common ancestor) of
mitochondrial DNA (mtDNA)
haplotypes in northern leopard frogs.
Analyzing mtDNA data is one way to
measure the genetic variation within a
species. When mtDNA lineages are
geographically localized and separated
by geographic barriers, this information
can be used to identify evolutionarily
separate units when it is used in
combination with patterns displayed by
other genetic markers (Avise 2004, p.
301). A haplotype refers to a set of
closely linked genetic markers present
on one chromosome that tend to be
inherited together. The more similar
these genetic markers, or haplotypes, are
in a given sample of frogs, the more
closely related those frogs are likely to
be (with the opposite also being the
case). This study (Hoffman and Blouin
2004a, p. 152) showed haplotypes of
mtDNA genetic markers grouping into a
‘‘western’’ lineage, occurring mostly
west of the Mississippi River and Great
Lakes region in the United States and
Canada, and an ‘‘eastern’’ lineage,
occurring to the east of this area.
The initial study by Hoffman and
Blouin (2004a, pp. 146, 150) found that
on a broad scale the eastern and western
haploypes have diverged for
approximately 2 million years,
indicating that the western and eastern
lineages have likely been separate to
some degree for a long time period, with
secondary contact following Pleistocene
glaciation events that occurred in North
America (Hoffman and Blouin 2004a, p.
152). The overall differences were
measured at approximately 4 percent
sequence divergence, and this amount
of mtDNA divergence is considered to
be relatively high and is comparable to
the differences found between some
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other recognized ranid frog species
(Jaeger et al. 2001, p. 344; Hoffman and
Blouin 2004a, p. 152). Hoffman and
Blouin (2004a, p. 152) note that mtDNA
divergence alone is not enough evidence
to split eastern and western lineages
into separate species and that more
taxonomic work (such as research
regarding nuclear genetic markers,
morphology, and behavior) is needed
before such a taxonomic revision would
be justified. The results of this study
indicated important genetic differences
broadly between northern leopard frogs
in the eastern and western portions of
North America. However, additional
data were needed to determine if the
‘‘western’’ lineage represented a
separate population of the species.
Although a preliminary
administrative report, the recent
O’Donnell et al. (2011) study report by
the U.S. Geological Survey was peerreviewed and presents the findings of a
robust analysis of the genetic variation
of the northern leopard frog across its
range in North America. The study
replicated the earlier mtDNA analysis
but had larger sample sizes (20–24
individuals per sample compared with
12 individuals per sample at most
sample localities) and had more sample
locations in the area of contact between
the eastern and western lineages. In
addition, it also included nuclear gene
sequencing as well. Nuclear genetic
sequences provided an additional way
to measure genetic variation in
populations of the northern leopard
frog. Because of its maternal (mother to
daughter) pattern of inheritance, mtDNA
is inherited only as a single genetic unit
and has some limits in value for
evaluating recent and localized
relationships within a species. However,
DNA sequences from multiple nuclear
genes provided more information from
additional genetic makers. This is an
important distinction because
identification of geographic
subdivisions, like judging population
distinction in the case of this analysis of
the northern leopard frog, depends on
the related geographic patterns of
different genetic markers (Avise 2004,
p. 303).
The study by O’Donnell et al. (2011)
was specifically designed to look at the
genetic relationships of the species and
to supplement the results of Hoffman
and Blouin (2004) by increasing the
number of samples in the area of
probable overlap of the two lineages in
the upper Midwest of the United States.
The analysis for one mtDNA gene
produced similar results to that of the
earlier study—with strong divergence
between east and west lineages and a
narrow area of overlap (O’Donnell et al.
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2011, pp. 2–3). However, the study also
analyzed DNA from four nuclear genes.
These nuclear genetic data still
indicated deeply divergent eastern and
western lineages of the northern leopard
frog. However, and most importantly for
our DPS analysis, the results of the
nuclear data showed a broad zone of
introgression between the two areas (in
other words, a mixing of haplotypes)
(O’Donnell et al. 2011, p. 10). We
considered this large zone of
introgression as the primary reason that
a potential western population of the
northern leopard frog is not considered
markedly separate from other
populations of the species.
So to determine whether these two
lineages should be considered markedly
separate populations and be considered
discreet under our DPS policy, we
looked at the relative amount of overlap
in the distribution of northern leopard
frogs that contain haplotypes from the
eastern and western lineages. Hoffman
and Blouin (2004a, pp. 147, 152, 155)
found that the distributions of eastern
and western haplotypes meet roughly at
the Mississippi River and Great Lakes
region, initially indicating that these
geographic features may serve as
physical barriers separating the eastern
and western lineages. However, the
additional nuclear genetic data from
O’Donnell et al. (2011, p. 10) discussed
above indicate the eastern and western
lineages are not separated along these
geographic features. Hoffman and
Blouin (2004a, pp. 147, 152) also found
some areas of co-occurrence of
haplotypes of both lineages in Ontario,
Canada, and indicated that this is likely
the result of more recent (during the
current interglacial period in North
America) secondary contact between
eastern and western lineages that were
formerly separated. In addition,
O’Donnell et al. (2011) reveal that the
haplotype mixing evident in the nuclear
analyses is more likely associated with
introgression and that more research is
needed to clearly explain the pattern of
haplotype mixing. The full extent of
current contact (and presumably gene
flow from interbreeding) between
northern leopard frogs with eastern and
western haplotypes could not be
evaluated in detail as a part of earlier
study because there were only a few
sample sites from the likely areas of
contact in Wisconsin, Michigan, and
western Ontario and limitations due to
small sample sizes. Further, there are
multiple factors that may be responsible
for the co-occurrence of frogs with
eastern and western haplotypes, for
example, it is possible that the mixing
of haplotypes between the east and west
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in the overlap zone may be attributable
in part to the anthropogenic movement
of individuals associated with the trade
in northern leopard frogs that has taken
place in this area since at least the 1950s
(Gibbs et al. 1971, p. 1027; Collins and
Wilbur 1979, p. 17).
Hoffman and Blouin (2004a, pp. 150–
151) also found one individual frog
(from a sample of 10) from Arizona with
an eastern haplotype. They suggested
this haplotype is likely not from a native
frog, but from a released pet or
laboratory animal. It is reasonable to
believe it was a released eastern frog, or
a descendant of one, because there is
commercial trade in leopard frogs and
tadpoles transported to pet stores,
laboratories, and schools throughout the
United States and Canada for
recreational and scientific uses (Fisher
and Garner 2007, p. 3). Their
supposition is also supported by
specific genetic research regarding this
Arizona population of northern leopard
frogs, which found haplotypes of
mtDNA consistent with frogs from
extreme eastern North America (from
New York, New England, and adjacent
areas of Quebec and Ontario)
widespread in the Stoneman Lake area
of northern Arizona (Theimer et al.
2011, p. 32).
The relatively small sample sizes
(about 12 individuals were used for
most sample localities) were a
disadvantage of the Hoffman and Blouin
(2004a, Appendix pp. 1–8) study in
evaluating genetic variation across a
narrow part of the range. While these
sample sizes were useful for looking at
broad patterns of geographic variation
(which was the object of the study), they
were less useful in answering our
question of separation, because of their
limited power for detecting haplotypes
that may occur at low frequencies and
there were few sample sites in the area
of suspected overlap. The small
differences in the amount of genetic
variation at specific locations are
important because even haplotypes at
low frequencies can help us understand
the relationships between the eastern
and western lineages of northern
leopard frogs and inform our
determination of whether the western
lineage is a markedly separate
population. The O’Donnell et al. (2011,
pp. 2–9) study utilized larger sample
sizes and provides a level of detail more
appropriate and helpful to evaluate
similarities and differences in western
and eastern lineages.
The results of O’Donnell et al. (2011,
pp. 2–9) indicated that neither the
Mississippi River nor the Great Lakes
are acting as a physical barrier between
western and eastern lineages of northern
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leopard frogs. The existence of western
haplotypes in northern leopard frog
populations located east of the
Mississippi River and of eastern
haplotypes in northern leopard frog
populations located both north and
south of the Great Lakes does not
support a marked separation between
eastern and western northern leopard
frogs. Although the nuclear genetic
sequences continue to show east-west
trends in different haplotypes
(supporting the mtDNA data of eastwest differences), these nuclear data
also indicate that western haplotypes
(from frogs in the west) occur in frogs
much farther to the east than the
mtDNA data indicated. Western
haplotypes of some of the nuclear genes
were found extending east of the
Mississippi River to the eastern end of
the Great Lakes in New York (O’Donnell
et al. 2011, pp. 6–8), and eastern
haplotypes of some of the nuclear genes
were found as far west as Nebraska
(O’Donnell et al. 2011, p. 9). This area
of overlap of haplotypes spans roughly
1,900 km (1,200 mi) from east to west
across North America.
This broad co-occurrence of
haplotypes of nuclear genes, as well as
the more gradual geographic trends in
haplotype distributions (O’Donnell et al.
2011, pp. 4–9), indicates there is not a
marked separation between eastern and
western lineages of the northern leopard
frogs. The overlap in genetic markers
across the midwestern United States
leads us to conclude that there is no
physical barrier or other processes
keeping northern leopard frogs in the
western part of the range discrete from
the frogs in the eastern part of the range.
Ongoing genetic analyses (such as
microsatellite allele frequency analyses)
will likely provide additional
information regarding geographic
patterns of genetic variation in northern
leopard frogs (O’Donnell et al. 2011, p.
10), but these data are not currently
available. Therefore, based upon the
genetic information presented above
(Hoffman and Blouin 2004a, pp. 145–
159; O’Donnell et al. 2011, pp. 1–10),
there does not appear to be marked
separation between possible eastern and
western populations of northern leopard
frogs. We do recognize that this lack of
a marked separation between the eastern
and western populations may be a result
of a variety of factors, including the
anthropogenic movement of individuals
for the trade in northern leopard frogs,
but at this time, we do not have data
supporting this claim. Because the
potential eastern and western
populations are not markedly separate,
they are not considered discrete under
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the DPS policy. Based upon the best
available information, we conclude that
the potential western U.S. population of
northern leopard frog is not genetically
discrete, in other words not markedly
separate, from other northern leopard
frogs.
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International Border
In order to determine that the
populations of northern leopard frog in
the western United States are a DPS, we
must have found that the western
United States populations were discrete
from populations in the eastern United
States and that the western United
States populations were discrete from
population in Canada. The DPS policy
allows us to use international borders to
delineate the boundaries of a DPS if
there are differences in control of
exploitation, conservation status, or
regulatory mechanisms between the
countries. However, because we do not
have a discrete east-west boundary of
the potential DPS, we did not conduct
further analysis regarding the northern
boundary of the potential DPS between
Canada and the United States.
Evaluation of Discreteness
The information discussed in the
preceding section provides information
on the geographic patterns that we
evaluated to determine that the genetic
information does not indicate that
northern leopard frogs from the western
United States are markedly separate
from other populations of the northern
leopard frog.
We note that our application of the
DPS policy does not require absolute
reproductive isolation as a prerequisite
to recognizing the discreteness of a
population segment. The presence of a
small degree of sharing of genetic
markers would not necessarily preclude
us from concluding that there is
discontinuity between populations and
that they were markedly separated.
However, in this case of the northern
leopard frog, we do not have the
information to make such an evaluation
of whether or not the two populations
are actually reproductively isolated.
Although the genetic patterns indicate
discontinuity in eastern and western
mtDNA and nuclear haplotypes, the
available genetic data do indicate there
is more than a small degree of sharing
of genetic markers. Rather than a small
degree of shared markers, we found a
broad extent of introgression that has
western haplotypes of some nuclear
genes occurring in samples of northern
leopard frogs as far as New York.
Therefore, because of the large area of
overlap in haplotypes indicating no
apparent barrier between the two
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lineages, we conclude at this time based
on the best available scientific data that
there is not marked separation between
the western and eastern U.S.
populations. This does not mean that
the western and eastern populations of
northern leopard frogs, as has been
suspected for many years, are not
unique and do not have significant
conservation value. It simply means
that, per our policy, the best available
data at this time do not support a
marked separation between the two
populations, based on genetics and
other information available to us.
In conclusion, based on our review of
the best available information and
pursuant to our DPS policy, we find that
the western U.S. populations of
northern leopard frog are not discrete
from other populations of northern
leopard frogs.
Significance
Under our DPS Policy, once we have
determined that a population segment is
not discrete, we do not need to consider
whether that population segment is
significant.
Conclusion
On the basis of the best available
information, we determined that the
western U.S. population of the northern
leopard frog is not discrete in relation to
the other populations of northern
leopard frog. Therefore, we find that the
western U.S. populations of northern
leopard frog do not represent a valid
DPS.
Having determined that the western
U.S. populations of northern leopard
frog are not a valid DPS, we proceed
below with an analysis of threats for the
northern leopard frog throughout its
range.
Summary of Information Pertaining to
Five Factors
Section 4 of the Act (16 U.S.C. 1533)
and implementing regulations (50 CFR
part 424) set forth procedures for adding
species to, removing species from, or
reclassifying species on the Federal
Lists of Endangered and Threatened
Wildlife and Plants. Under section
4(a)(1) of the Act, a species may be
determined to be endangered or
threatened based on any of the
following five factors:
(A) The present or threatened
destruction, modification, or
curtailment of its habitat or range;
(B) Overutilization for commercial,
recreational, scientific, or educational
purposes;
(C) Disease or predation;
(D) The inadequacy of existing
regulatory mechanisms; or
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(E) Other natural or manmade factors
affecting its continued existence.
In making this finding, information
pertaining to the northern leopard frog
in relation to the five factors provided
in section 4(a)(1) of the Act is discussed
below.
In considering what factors might
constitute threats to a species we must
look beyond the exposure of the species
to a particular factor to evaluate whether
the species may respond to that factor
in a way that causes actual impacts to
the species. If there is exposure to a
factor and the species responds
negatively, the factor may be a threat
and, during the status review, we
attempt to determine how significant a
threat it is. The threat is significant if it
drives, or contributes to, the risk of
extinction of the species such that the
species warrants listing as endangered
or threatened as those terms are defined
in the Act. However, the identification
of factors that could impact a species
negatively may not be sufficient to
compel a finding that the species
warrants listing. The information must
include evidence sufficient to suggest
that these factors are operative threats
that act on the species to the point that
the species may meet the definition of
endangered or threatened under the Act.
Due to the wide geographic range of
the northern leopard frog, and the
diversity of habitat types which it
occupies throughout its range, there are
a wide variety and relatively large
number of factors that have the potential
to impact the species. However, these
factors may result in impacts at the
individual, population, or species scale,
and may have a variety of effects from
minor habitat degradation to complete
habitat loss and mortality. As such, it is
important to consider the magnitude
and extent of impacts when assessing
the factors affecting a species, and we
attempt to provide this context
throughout our discussions below.
Factor A. The Present or Threatened
Destruction, Modification, or
Curtailment of Its Habitat or Range
A number of hypotheses, including
habitat loss, have been proposed for
global amphibian declines (Blaustein et
al. 1994, p. 61; Collins and Storfer 2003,
pp. 90–94; Stuart et al. 2004, p. 1783;
Green 2005, p. 28). In our review of the
best scientific and commercial data
available, impacts that are potentially
affecting northern leopard frogs and
their habitats throughout their range
include habitat destruction, habitat
fragmentation, and habitat degradation
resulting from development,
modification, and loss of wetland
habitat. Because the northern leopard
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frog, an amphibian, depends upon
breeding ponds, upland foraging areas,
overwintering aquatic habitats, and
connectivity between these habitats
across the landscape, it is very
susceptible to the destruction (defined
as complete loss of all or part of the
frog’s necessary habitat), fragmentation
(isolation of all or part of the frog’s
necessary habitat without its alteration
or destruction), and degradation (the
deleterious alteration of all or part of the
frog’s necessary habitat) of its habitat
(Green 2005, p. 28).
The destruction and degradation of
northern leopard frog habitat has been
widespread and has affected, and
continues to affect, the species to some
extent throughout its range (Maxell
2000, p. 15; Hitchcock 2001, pp. 64–66;
Rorabaugh 2005, p. 576; Clarkson and
Rorabaugh 1989, p. 535; Smith 2003, pp.
26–31). Habitat destruction and
degradation is reported to be the
primary threat to all ranid and
lithobatid frogs in the United States
(Bradford 2005, p. 923) and a principal
cause of decline of northern leopard
frogs in the western United States and
Canada (Smith 2003, p. 4; Alberta
Northern Leopard Frog Recovery Team
2005, p. 6; Rorabaugh 2005, p. 571;
Committee on the Status of Endangered
Wildlife in Canada 2009, p. 32). Factors
with the potential to impact northern
leopard frog habitat include wetland
loss, agricultural development, livestock
grazing, urban development, oil and gas
development, forest management, roads,
groundwater withdrawal, and air
pollution. Below we present
information about these factors and
discuss the magnitude and extent of the
impacts from these factors on the
northern leopard frog.
Wetland Loss
As a species with aquatic and semiaquatic life-history phases, freshwater
wetland habitat is an extremely
important component of northern
leopard frog habitat. In order to discuss
the different actions that result in
destruction or modification of northern
leopard frog habitat, it is important to
understand what is known about the
current overall status of wetlands
throughout the range of the northern
leopard frog.
It has been estimated that 53 percent
of the Nation’s former wetland area was
lost from the 1780s to the 1980s (Dahl
1990, p. 5). In terms of States where the
northern leopard frog occurs, Minnesota
(42 percent loss), Maine (20 percent
loss), Michigan (50 percent loss), and
Wisconsin (46 percent loss) have the
most remaining wetland area compared
to historical times (Dahl 1990, p. 5).
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New Hampshire (9 percent loss) was the
only State in the range of the northern
leopard frog that lost less than 20
percent of its original wetland acreage
(Dahl 1990, p. 5). California (91 percent
loss), Connecticut (74 percent loss),
Illinois (85 percent loss), Indiana (87
percent loss), Iowa (89 percent loss),
Kentucky (81 percent loss), Missouri (87
percent loss), and Ohio (90 percent loss)
lost over 70 percent of their original
wetland acreage (Dahl 1990, pp. 5–6).
The remaining States within the range
of the northern leopard frog had
estimated wetland losses ranging from
20 percent to 60 percent (Dahl 1990, p.
6).
Dahl (1990, p. 10) noted that wetland
area in the lower 48 States had declined
to the point that ‘‘environmental, and
even socio-economic benefits (ground
water supply, water quality, shoreline
erosion, floodwater storage, trapping of
sediments, and climatic change) are
now seriously threatened.’’ The
destruction and degradation of wetland
and riparian habitat is thought to
represent the most widespread impact
to northern leopard frog populations in
Arizona (Arizona Game and Fish
Department 2009, p. 1), Colorado
(Colorado Division of Wildlife 2009, p.
2), Idaho (Idaho Department of Fish and
Game 2005), Montana (Montana Fish
Wildlife and Parks 2009, p. 2), Nevada
(Nevada Department of Wildlife 2009, p.
4), New Mexico (New Mexico
Department of Game and Fish 2009, p.
3), North Dakota (North Dakota Game
and Fish Department 2009, p. 2), Utah
(Utah Department of Wildlife Resources
2009, pp. 2–3), Wisconsin (Wisconsin
Department of Natural Resources 2009,
p. 1), Connecticut (Klemens 2000, p. 1),
Indiana (Indiana Department of Natural
Resources 2006, p. 113), Kentucky
(Kentucky Department of Fish and
Wildlife Resources 2010, p. 27), Maine
(Maine Department of Natural Resources
2005, p. 90), Massachusetts
(Massachusetts Department of Fish and
Wildlife 2006, pp. 276, 292, 328),
Michigan (Eagle et al. 2005, Threats p.
20), New Hampshire (New Hampshire
Fish and Game Department, p. A–210),
New York (New York Department of
Environmental Conservation 2005, pp.
57–58), and Rhode Island (Rhode Island
Department of Environmental
Management, Division of Fish and
Wildlife 2005, p. 22).
While the total wetland losses in the
United States are significant, the
information regarding status and trend
of wetlands only looks at total losses
and gains of wetland area; there is no
comprehensive data assessing trends in
the quality or function of lost wetlands
(Dahl 2006, p. 74). Therefore, we do not
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know how much of the lost wetland
habitat would have naturally functioned
as northern leopard frog habitat. In
short, while the extent of wetland losses
is broad and widespread throughout the
range of the species, we are unable to
assess the magnitude or severity of
impact of these losses at the species
scale. There have most likely been
losses of northern leopard frog habitat
concurrent with these wetland losses,
but large areas of wetland remain intact
in many States, particularly in the
eastern portion of its range in the United
States. Further, the data above address
total change in wetland area without
reference to the causes of the losses;
thus it is difficult to relate past losses to
future losses in this context. Ongoing
impacts to northern leopard frog
habitats will be discussed more
specifically in the following sections.
Since the late 1980s, creation of new
wetland area has occurred, although the
rate of replacement area is much slower
than the historical loss rate (Dahl 1990,
p. 5). Data collected from 1998 to 2004
indicate that for the first time since
uniform monitoring began, wetland
creation actions resulted in a larger net
gain of wetlands than net loss of
wetlands during this time period (Dahl
2006, p. 15). However, the location and
types of wetlands that represent this
gain in wetland acres has not
necessarily resulted in the creation of
northern leopard frog habitat. In terms
of location, a majority of the wetland
areas gained were created in the
southeast, particularly in Florida, which
is outside the range of the northern
leopard frog (Dahl 2006, p. 62). Further,
review of created ponds from 1986 to
1997 indicates that only 2 percent of
these ponds were reclassified as
vegetated wetlands; most created ponds
are designed and maintained to function
as open water basins—deep waters with
little vegetated shoreline and steep
slopes—that are not conducive to
northern leopard frog breeding, foraging,
or dispersal (Hine et al. 1981, p. 12; Leja
1998, p. 351; Semlitsch 2000, p. 624).
All of the created ponds that Dahl (2006,
pp. 76–78) noted were manmade farm
ponds, freshwater fishing ponds,
detention ponds, and aquaculture
ponds. Deepwater lakes and reservoirs
also increased in area over this time
period (typically associated with urban
development) (Dahl 2006, p. 78). Many
of these ponds or open water bodies are
not an equivalent replacement for
vegetated wetlands (Dahl 2006, p. 76),
and although they count towards the
total of wetland area in the
conterminous United States, they do not
necessarily indicate a gain in northern
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leopard frog habitat, particularly if
water quality, vegetation, and native
species are not objectives for the created
wetland.
In Canada, wetland loss has also
occurred throughout the range of the
northern leopard frog. Wetland habitat
quality is considered to be a limiting
factor for the one remaining northern
leopard frog population in British
Columbia (Committee on the Status of
Endangered Species in Canada 2009, p.
16). It is estimated that approximately
60 percent of basins and 80 percent of
wetland margins in the 1980s in
southern Alberta were degraded and
that local extirpations of northern
leopard frogs likely occurred as a result
(Alberta Northern Leopard Frog
Recovery Team 2005, p. 6). By 1990,
approximately 20 percent of prairie
wetlands that likely functioned as
northern leopard frog habitat in
Manitoba were lost (Committee on the
Status of Endangered Species in Canada
2009, p. 17). Similar patterns of
significant wetland loss have occurred
in southern Ontario and southern
Quebec. Historically, 69 percent of
southwestern Ontario consisted of
wetlands; however, it is estimated that
as much as 90 percent of southwestern
Ontario wetlands no longer exist
(Committee on the Status of Endangered
Species in Canada 2009, p. 17). Again,
similar to the situation in the United
States, we do not have information
assessing how much of this lost habitat
may have functioned as northern
leopard frog habitat or if any mitigation
(such as created wetlands) has resulted
in replacement habitat. While it is likely
there have been losses of northern
leopard frog habitat concurrent with
these wetland losses, large areas of
wetland remain intact, particularly in
the eastern portion of Canada.
Across the range of the species, it is
clear that significant total wetland area
has been lost since colonial times. It is
logically certain that some of these areas
represented historic habitat for northern
leopard frogs; however, it is not possible
to assess the extent of loss of actual
northern leopard frog habitats based on
a generalized review of loss of wetlands.
Further, while wetland losses have
occurred, large areas of wetland remain,
particularly in the eastern portion of the
United States and Canada.
Agricultural Development
Agricultural development has
occurred across the range of the
northern leopard frog, but particularly
in the Midwestern States of the United
States (Leja 1998, p. 349). The U.S.
Department of Agriculture, Natural
Resource Conservation Service (USDA
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NRCS) has a broad land cover and use
map that shows by State the amount of
land in cropland, pastureland,
rangeland, forest land, developed land,
Federal lands, and other lands. Data
from this map shows that greater than
80 percent of the total land area (outside
Federal lands) in Iowa, Nebraska, North
Dakota, and South Dakota is used for
agricultural purposes, such as cropland,
pastureland, and rangeland (USDA
NRCS 2001). In addition, many other
western and Midwestern States also
have significant amounts of land
identified as agricultural within the
range of the northern leopard frog
(USDA NRCS 2001). While agricultural
development continues to be a large
land-use practice in South Dakota (57
percent cropland), North Dakota (35
percent cropland), and Ohio (45 percent
cropland) (USDA NRCS 2001), the
northern leopard frog appears to be
relatively stable in these States (Hossack
et al. 2005, p. 428; Rorabaugh 2005, p.
571), despite this level of usage.
Agricultural development may
fragment, destroy, or degrade northern
leopard frog habitat directly due to
conversion of native habitats to
cropland and de-watering of adjacent
habitats, or indirectly through the
introduction of contaminants and
invasive species into habitats (Wang et
al. 1997, p. 10; Leonard et al. 1999, p.
58; Leja 1998, pp. 345–353; Knutson et
al. 2004, p. 675; Rorabaugh 2005, p.
576). Most of the historic wetland loss
discussed above is thought to be due to
conversion to agriculture (Leja 1998, p.
349). Agricultural development can
result in modification of river valley
habitat, including draining of wetlands,
channelization and damming of rivers,
and development of irrigation systems
(Wang et al. 1997, p. 11; Findlay and
Houlahan 1997, p. 1001), all of which
may modify breeding, overwintering,
and dispersal habitat for northern
leopard frogs (Scott and Jennings 1985,
p. 19; Lannoo et al. 1994, pp. 317–318;
Leja 1998, pp. 345–353; Knutson et al.
2000, p. 139; Ammon 2002, p. 2; Idaho
Department of Fish and Game 2005,
Northern leopard frog species account;
Colorado Division of Wildlife 2009, p. 1;
Rogers 2010, p. 8). For example, in
Idaho, Camas NWR is losing wetlands to
groundwater depletion by nearby
agriculture, and Grays Lake NWR and
Minidoka NWR cannot control water
levels because of senior water rights
assigned to other agencies, and their use
for agriculture (Fisher and Mitchell
2009, pers. comm.). In Canada, the past
conversion of large areas of grassland to
agriculture has also likely resulted in
the loss of northern leopard frog habitat,
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particularly foraging and overwintering
habitats near breeding sites (Didiuk
1997, p. 113; Hecnar 1997, p. 13). In
southern Alberta, drainage of wetlands
for agricultural use in the 1980s was
extensive and is thought to have
contributed to local extirpations of
northern leopard frogs (Alberta
Northern Leopard Frog Recovery Team
2005, p. 6). The land being used for
agriculture in the prairies has lately
increased by 62 million acres (25
million hectares), and there is pressure
to alter remaining wetland areas
(Committee on the Status of Endangered
Wildlife in Canada 2009, p. 32).
Geographically isolated (or
depressional) wetlands surrounded by
upland watersheds (such as the prairie
potholes region) make up a large
proportion of the wetland resource in
arid and semi-arid regions of the
northern leopard frog’s range (Skagen et
al. 2008, p. 594). However, although the
‘‘wet’’ (surface water) portion of the
wetland is vitally important for northern
leopard frog breeding, the upland
terrestrial habitat adjacent to the
wetland is also a critical component of
their habitat needs (Semlitsch 2000, p.
620; Pope et al. 2000, p. 2506; Gibbons
2003, p. 630; Semlitsch and Bodie 2003,
p. 1223). Although agricultural
development may result in the
maintenance or creation of actual ‘‘wet’’
wetland habitat (Leja 1998, p. 350),
crops and pastures—areas that provide
poor or no habitat for northern leopard
frog—typically occur on the immediate
edge of the water (Guerry and Hunter
2002, p. 752; Committee on the Status
of Endangered Species in Canada 2009,
p. 32). Research indicates that land use
practices around the wetland may be as
important as the size of the wetland
itself (Findlay and Houlahan 1997, p.
1007). Amphibian species richness
increases with wetland area, and
herpetofauna abundance, including the
northern leopard frog, show a strong
positive correlation with the proportion
of forest cover on lands within 1.2 mi
(2 km) of wetlands (Findlay and
Houlahan 1997, pp. 1006–1007).
Northern leopard frogs breeding in
active agricultural lands may end up
crossing roads and tilled agricultural
fields which would increase the
likelihood of mortality, and northern
leopard frogs that breed in active
agricultural lands require larger home
ranges than do frogs that breed in intact
wetlands and grasslands (Pember et al.
2002, p. 4.9)
Habitat fragmentation caused by
agriculture has also likely limited
northern leopard frog dispersal, as frogs
may have difficulty moving through
active croplands (Didiuk 1997, p. 113;
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Saskatchewan Conservation Data Centre
2006, p. 2). Agricultural development
also tends to result in disturbed ground,
which can impact the distance and the
quality of habitat between habitat
patches (Didiuk 1997, p. 113; Pember et
al. 2002, p. 4.9; Alberta Northern
Leopard Frog Recovery Team 2005, p. 6;
Mazerolle and Desrochers 2005, p. 455;
Committee on the Status of Endangered
Wildlife in Canada 2009, p. 32). Barren
land, agricultural lands, and recently
cut forests increase the resistance of the
landscape to northern leopard frog
movement (Mazerolle and Desrochers
2005, p. 462). Vegetation on
undisturbed sites likely reduces
evaporative water loss in dispersing or
moving frogs through protection from
the wind and sun (reduced
dehydration), while surfaces with no
vegetative cover likely endanger
individual frogs and constitute barriers
to frog movement (Mazerolle and
Desrochers 2005, p. 462). In addition,
agriculturally induced habitat
fragmentation can increase the role of
genetic drift, which may hamper
adaptive responses to local
environments (Johansson et al. 2007, p.
2699). Research regarding the European
common frog (Rana temporia) found
that populations in fragmented
agricultural habitats were smaller and
had lower genetic diversity compared to
populations in a more continuous
landscape. More genetic diversity leads
to healthier populations. Breeding pond
isolation, resulting from fragmented
landscapes, has also been shown to
negatively affect population persistence
and recolonization of ranid and
lithobatid frogs to suitable habitats
(Witte et al. 2008, p. 381).
Agriculture is also the primary source
of water pollution throughout the
western range of the northern leopard
frog and occurs primarily through
sedimentation, nutrient pollution,
pesticide pollution, and mineral
pollution (Ribaudo 2000, pp. 5–11). On
many NWRs, pesticide and herbicide
use are regulated by Service Pesticide
Use Plans, but these plans may not
adequately account for toxicity to
northern leopard frogs, and thus
pesticide and herbicide use may result
in impacts to individuals or populations
of the species (Dickerson and Ramirez
1993, pp. 1–2; Fisher and Mitchell 2009,
pers. comm.). Overwintering northern
leopard frogs in permanent waters are
likely to be in close contact with
sediments on the pond bottom that may
contain agricultural chemicals resulting
from run-off (Didiuk 2007, p. 113). This
close contact with chemicals may make
the northern leopard frog more
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susceptible to potential adverse
chemical effects in these areas.
Leopard frogs that inhabit agricultural
wetlands and landscapes are also
vulnerable to pesticide exposure (King
et al. 2008, p. 13) (see Pesticides under
Factor E for further discussion). In
addition, ‘‘hotspots’’ of amphibian
malformations, including northern
leopard frog malformations, tend to
occur in altered wetlands (Lannoo 2008,
p. 200) (see Malformations under Factor
E for further discussion).
As described above, agricultural
development has been shown to result
in adverse effects to northern leopard
frogs in some portions of its range. The
above review of the best available
information indicates that large areas of
historical habitat have likely been lost
due to agricultural development and
that current habitats may continue to be
subject to ongoing impacts of
agricultural development. The most
significant impacts associated with
agricultural development are likely the
loss of historical habitats due to
conversion to agricultural lands.
Ongoing impacts to areas currently
associated with agriculture likely
negatively impact local populations
through reduced breeding success and
individual survival. However, even
States with a significant land base in
agriculture (such as South Dakota, North
Dakota, and Ohio) appear to be
maintaining stable populations of
northern leopard frogs. Therefore,
though research indicates that
agricultural development can have a
negative impact on local populations of
northern leopard frogs, the best
available information does not indicate
the ongoing impacts are significant at
the species level. Based upon the best
available information, agricultural
development does not constitute a
significant threat to the northern
leopard frog at the species level now,
nor do we have indication that it will in
the future.
Livestock Grazing
Approximately 70 percent of the land
surface in the western United States
(including Montana, Wyoming,
Colorado, New Mexico, Arizona, Utah,
Nevada, California, Idaho, Oregon, and
Washington) is or has been grazed by
livestock (Fleischner 1994, p. 630;
Krausman et al. 2009, p. 15). Historical
and ongoing livestock grazing are
specifically identified as being
responsible for the loss and degradation
of northern leopard frog habitats, and
for negatively affecting northern leopard
frog populations at sites in Arizona
(Clarkson and Rorabaugh 1989, p. 535;
Sredl 1998, pp. 573–574), California
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(California Department of Fish and
Game 2007), Idaho (Idaho Department of
Fish and Game 2005, Appendix F),
Montana (Maxell 2000, p. 15), Nevada
(Hitchcock 2001, p. 66), North Dakota
(Euliss, Jr. and Mushet 2004, p. 82),
South Dakota (Smith 2003, p. 27), and
Wyoming (BLM 2009, p. 3). For
example, most of the habitat in the Pit
River-Modoc Plateau area and the
Owens Valley of California, where the
northern leopard frog occurred
historically, has been severely altered
and fragmented largely because of
livestock grazing practices. The
essential habitats bordering riparian
zones are either no longer present or so
fragmented that the habitat can no
longer support northern leopard frog
populations (Jennings and Hayes 1994,
p. 82). Although management may be
changing in some areas, many wetland
habitats are likely still recovering from
historical grazing impacts (Krausman et
al. 2009, p. 16). This is particularly true
because the western United States has a
relatively arid climate, which can result
in longer habitat recovery intervals, and
perennial waters tend to be rarer and
more disjunct from other waters than in
the eastern United States.
Livestock select riparian habitats for
water, shade, and cooler temperatures.
They tend to spend a disproportionate
amount of their time in riparian zones,
and they can adversely affect these
systems in a number of important ways
(Fleischner 1994, pp. 633–635; Belsky et
al. 1999, pp. 420–424; Jones 2000, pp.
159–161). Because of this
disproportionate use of mesic and
riparian habitats by livestock, northern
leopard frog populations are vulnerable
to the effects of poorly managed
livestock grazing (Maxell 2000, pp. 15–
16; Smith 2003, p. 30). Specifically,
trampling by livestock may result in the
death of individual frogs (Bartlet 1998,
p. 96; Maxell 2000, p. 15; Smith 2003,
p. 30), and the compaction of soils
around aquatic habitats, thereby
decreasing infiltration of water into the
soil, increasing soil erosion, and
contributing to stream channel down
cutting (Kauffman and Kreuger 1984,
pp. 432–434; Belsky et al. 1999, pp.
419–431). These impacts could hinder
or prevent movements of northern
leopard frogs by reducing and
eliminating riparian vegetation that
provides cover.
Impacts to water quality through
increased sedimentation (Belsky et al.
1999, pp. 420–424; Alberta Northern
Leopard Frog Recovery Team 2005, p. 7)
may reduce the depth of breeding ponds
or overwintering habitats, increase
water temperatures, and create favorable
environments for diseases and parasites
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known to contribute to mortality in
northern leopard frogs (Maxell 2000, pp.
15–16; Johnson and Lunde 2005, pp.
133–136; Ouellet et al. 2005, p. 1435).
Increased watershed erosion caused by
livestock grazing can accelerate
sedimentation of deep pools used by
frogs (Gunderson 1968, p. 510). The
indirect effects of grazing on northern
leopard frog habitat may also include
increases in sedimentation generated by
grazing. Sediment can alter primary
productivity and fill interstitial spaces
in drainage materials with fine
particulates that impede water flow,
reduce oxygen levels, and restrict waste
removal (Chapman 1988, pp. 5–10).
Disturbance from livestock wading
and defecating in northern leopard frog
habitat has been found to have negative
effects on the reproductive success of
northern leopard frogs and to result in
negative impacts to habitat (Knutson et
al. 2004, p. 677). The significant input
of urine and manure and the turbidity
caused by livestock disturbance was
found to lead to poor water quality
(such as increased nitrates) and low
oxygen concentrations, which can result
in reduced development and survival of
egg masses and tadpoles (Marco et al.
1999, p. 2837; Rouse et al. 1999, pp.
800–802; Ortiz et al. 2004, pp. 235–236;
Alberta Northern Leopard Frog
Recovery Team 2005, p. 7; Earl and
Whiteman 2009, p. 1336). In addition,
Knutson et al. (2004 p. 675) found that
the grazed ponds had little or no aquatic
or emergent vegetation, and that this
was a result of livestock wading in the
pond.
In contrast, there is information from
some portions of the range of the species
that indicates leopard frog species can
persist, and even benefit from, wellmanaged livestock grazing (Hitchcock
2001, p. 62; Service 2007, pp. 32–34;
Alberta Northern Leopard Frog
Recovery Plan 2005, p. 7; Arizona Game
and Fish Department 2009, pp. 2–3;
New Mexico Department of Fish and
Game 2009, p. 3). Limited grazing
around riparian areas can create open
foraging areas for leopard frogs, and
livestock management can result in the
creation of stock tanks (ponds or
impoundments that function as
waterholes) that can provide breeding
and dispersal habitat for northern
leopard frogs, particularly in arid
western landscapes (Sredl et al. 1997,
pp. 46, 49; Theimer et al. 2011, p. 11).
Historically, livestock grazing has
likely resulted in degraded habitats and
local declines and extirpations of
northern leopard frogs in some portions
of their range. However, the information
reviewed above suggests that livestock
grazing has only resulted in substantive
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impacts in the western portions of the
United States and Canada, with very
little to no information suggesting how
livestock grazing has or is adversely
impacting northern leopard frog
populations in the eastern United States
or eastern Canada. Further, declines and
extirpations associated with livestock
grazing are likely historical impacts in
most areas, with ongoing impacts
manifesting primarily through effects
associated with degraded habitats.
Finally, there is no evidence that
livestock grazing use is spreading to
areas that are not already subject to
those uses. Therefore, the best available
scientific information indicates that
livestock grazing does not constitute a
significant threat to the northern
leopard frog at the species level now,
nor do we have indication that it will in
the future.
Urban Development
Urbanization refers to the
development of areas for human uses.
Areas subject to urbanization tend to be
correlated to areas with increased
human population growth. This
development is resulting in impacts to
northern leopard frog habitat across its
range (Hitchcock 2001, pp. 64–66;
Smith and Keinath 2007, p. 29;
Connecticut Department of
Environmental Protection 2005, pp. 2–
16–2–18; Maine Department of Inland
Fisheries and Wildlife 2005, Chapter 5
p. 109; New Hampshire Fish and Game
Department 2005, p. A210–212;
Wisconsin Department of Natural
Resources 2009, p. 1). The 2010 Census
reported that the human population in
the United States has increased almost
10 percent since 2000. The only State
within the range of the northern leopard
frog that did not have an increase in
population is Michigan (Mackun and
Wilson 2011, pp. 1–2). Nevada, Arizona,
Utah, Texas, and Idaho were the fastest
growing States, and New Hampshire
and South Dakota were the fastest
growing States in the northeast and
Midwest, respectively. Pennsylvania
ranks fifth in the nation in the amount
of open space it loses to development
every day and it has lost over half of its
wetlands to development (Pennsylvania
Game Commission and Pennsylvania
Fish and Boat Commission 2005, pp.
10–34). In Canada, Ontario and Quebec
are the largest provinces in terms of
numbers of people; larger numbers of
people typically contribute more to
increases in urban development and
modification of northern leopard frog
habitats. Projected human population
growth is also expected to result in
increased needs for water (surface
diversions and groundwater pumping)
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to support this growth (Deacon et al.
2007, p. 688). This could decrease water
availability for northern leopard frogs
and thereby impact the amount and
extent of habitat for northern leopard
frogs. Reexamination of historic
northern leopard frogs sites in
northeastern Ohio (Orr et al. 1998, p.
92) found that two sites had been
destroyed by development and three
had been eliminated by high-intensity
agriculture. A study in Iowa and
Wisconsin found a negative association
with urban land use and relative
abundance of northern leopard frogs
(Knutson et al. 1999, p. 1441; Knutson
et al. 2000, p. 140). From 1998 to 2004,
140,400 ac (56,800 ha) or 61 percent of
wetland losses in the United States
occurred due to urban and rural
development (Dahl 2006, p. 47). These
wetland losses are considered to be
irreversible as they are the result of
permanent construction (such as houses
and roads) that alters wetland hydrology
(Dahl 2006, pp. 47, 63). Urban
development often results in conversion
of natural habitats to homes, roads, and
industrial uses, which can result in
direct mortality from traffic (Mazerolle
2004, p. 47; Bouchard et al. 2009, p. 23),
chemical contamination of wetlands
(Fahrig et al. 1995, p. 177), and
modification of existing wetland
habitats to benefit sport fish rather than
native amphibians (Knutson et al. 1999,
p. 1444).
Based upon the above information,
urban development has likely resulted
in the historical and continued loss of
northern leopard frogs and their habitat
throughout their range. While the
magnitude of these impacts is
conceivably high in localized areas,
urbanization is not ubiquitous
throughout the range of the northern
leopard frog. General information about
human population growth and
associated urbanization cannot be
extrapolated to support high magnitude
threats throughout all portions of the
range of the northern leopard frog.
Further, despite urbanization trends, the
northern leopard frog is apparently still
considered to be widespread and
common in the eastern United States
and eastern Canada. Therefore, the best
available scientific information
indicates that urbanization does not
constitute a significant threat to the
northern leopard frog at the species
level now, nor do we have indication
that it will in the future.
Oil and Gas Development
Natural gas drilling is currently
occurring in at least 25 States that have
populations of northern leopard frogs.
In 2007, there were 449,000 natural gas
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wells in 32 States, which was a 30
percent increase from 2000; it is
estimated that 32,000 new natural gas
wells per year could be drilled by 2012
(Lustgarten 2008, p. 2). Examples of the
increase in magnitude of drilling in the
United States can be observed by the
increase in approved permits in
Wyoming and Pennsylvania. The first
natural gas well in Sublette County,
Wyoming, was drilled in 1939, and by
2008, 700 gas wells were producing
natural gas on the Pinedale Anticline (a
major gas field in Sublette County). In
2008, the Bureau of Land Management
approved 4,400 more natural gas wells
in Sublette County (Lustgarten 2008, p.
3). In Susquehanna County,
Pennsylvania, there was a 27-fold
increase in natural gas well permits
from 2007 to 2009. Natural gas mining
is also occurring in Canada, the world’s
third-largest producer and exporter of
natural gas (Natural Resources Canada
2011, p. 1). However, we have minimal
specific information assessing the
overlap of occupied northern leopard
frog habitats with planned oil and gas
development operations for most of the
range of the species.
The Powder River Basin in Wyoming
and Montana and the San Juan Basin in
Colorado and New Mexico, areas within
the range of the northern leopard frog,
currently have the highest coalbed
methane (a natural gas) productions in
the United States (Environmental
Protection Agency 2004, p. 1–1).
Possible impacts to northern leopard
frogs associated with coalbed methane
development may include discharge of
contaminated water into breeding
ponds, loss of spring flows related to
groundwater withdrawals, discharge of
extremely cold water into breeding
habitats, discharge of water containing
nonnative predatory fish in these same
areas, and road-related mortality
associated with increased use of roads
or new roads to support the coalbed
methane development (Allan 2002, pp.
5–8; Gore 2002, pp. 1–14; Noss and
Wuethner 2002, pp. 1–20). Mining and
oil and gas development may also lead
to contamination of habitats (Spengler
2002, pp. 7–26; Smith 2003, pp. 26, 31).
Domestic and stock tank waters have
dried or become contaminated with gas
in Wyoming’s Powder Basin (Powder
River Basin Resource Council 2009, p.
1). Although some States that have
populations of the northern leopard frog
are implementing wetland and riparian
protections in connection with oil and
gas drilling (Colorado Division of
Wildlife 2009, p. 5), it is unclear if all
States are implementing such measures
and whether or not these measures have
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resulted in decreased impacts to
northern leopard frogs.
Another area where there is
information about oil and gas
development activities in northern
leopard frog habitats is the Marcellus
Shale. The Marcellus Shale is a black
shale formation extending underground
from Ohio and West Virginia northeast
into Pennsylvania and southern New
York that contains natural gas reserves.
Although there are areas where the
Marcellus Shale is exposed at the
surface, it is as deep as 7,000 ft (2,134
m) or more below the ground surface
along the Pennsylvania border. Natural
gas drilling operations have proliferated
in Pennsylvania over the past years, and
at least 1,415 new wells were drilled in
2010 (Goldberg 2011, p. 2). The drilling
is expected to expand into Ohio and
West Virginia. New York is currently
conducting a comprehensive review of
the potential environmental impacts
associated with natural gas development
and Ohio’s State government approved
drilling in Ohio’s State parks on June
15, 2011.
Hydraulic fracturing is a method used
to extract natural gas from the earth.
Environmental concerns with hydraulic
fracturing include water use and
management, and the composition of
the fluids used (Environmental
Protection Agency 2011, p. 1).
Hydraulic fracturing consists of
pumping chemicals (such as benzene)
and high volumes of water and sand
down the well under high pressure to
create fractures in the gas-bearing rock
(New York Department of
Environmental Conservation 2011, p. 1).
The propping material holds the
fractures open allowing more gas to flow
into the well. The hydraulic fracturing
of the Marcellus Shale will require large
volumes of water to fracture the rocks
and produce natural gas. In 2008, oil
and gas wells disgorged approximately
9 million gallons of wastewater a day in
Pennsylvania, and water use is expected
to increase to at least 19 million gallons
per day (Sapien 2009, p. 2).
The wastewater is a product of the
hydraulic fracturing which pumps about
1 million gallons of water mixed with
sand and chemicals into each well to
withdraw the natural gas. When it
comes back out, the water contains
toxins and dissolved solids. Wastewater
contains enough dissolved solids that
the water can be five times as salty as
sea water. Recent research found
methane contamination of drinking
water in Pennsylvania and New York
from natural gas extraction on the
Marcellus Shale (Osborn et al. 2011, p.
2). In addition, water contamination has
been documented near drilling areas in
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Sublette County, Wyoming, and Santa
Fe, New Mexico; chemical spills of
hydraulic fracturing chemicals have
occurred in Colorado (Lustgarten 2008,
pp. 2–9).
The rate, timing, and location of water
withdrawals could result in negative
impacts to streams, downstream riverine
and riparian resources, wetlands, and
aquifer supplies where hydraulic
fracturing to mine natural gas occurs
(New York Department of
Environmental Conservation 2009, p. 6–
4). The draft environmental impact
statement for natural gas drilling in New
York states, ‘‘Water for hydraulic
fracturing may be obtained by
withdrawing it from surface water
bodies away from the well site or
through wells drilled into groundwater
aquifers’’ (New York Department of
Environmental Conservation 2009, p. 6–
4). The existence and sustainability of
wetland habitats directly depend on the
presence of water at or near the surface
of the soil. The functioning of a wetland
is driven by the inflow and outflow of
surface water and groundwater. As a
result, withdrawal of surface water or
groundwater for high volume hydraulic
fracturing could impact wetland
resources and northern leopard frog
habitat. These potential impacts depend
on the amount of water within the
wetland, the amount of water
withdrawn from the catchment area of
the wetland, and the dynamics of water
flowing into and out of the wetland.
Even small changes in the hydrology of
the wetland can have significant
impacts on the wetland plant
community and on the wildlife, such as
the northern leopard frog, that depend
on the wetland. As discussed in the
Biology section, wintering northern
leopard frogs are intolerant of freezing,
and withdrawals that reduce water
depths in overwintering habitat could
lead to high levels of winter kill if water
levels are reduced so much that these
areas freeze.
In summary, some northern leopard
frog populations could be impacted by
oil and gas development activities
through changes to water quantity or
quality (due to chemical pollution or
increased salinity) and through
insufficient water flow to maintain
wetland and stream habitat. Natural gas
drilling and hydraulic fracturing may
occur across the range of the northern
leopard frog; however, the impacts are
expected to be localized population and
habitat losses rather than regional or
species-level effects. Pennsylvania,
Ohio, West Virginia, Kentucky,
Wyoming, Colorado, Montana, and New
Mexico all have oil and gas
development occurring within their
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boundaries; however, we have little to
no information about oil and gas
development activities in northern
leopard frog habitats throughout the rest
of the range of the species, notably the
Midwestern United States and Canada.
Therefore, the best available scientific
information indicates that oil and gas
development does not constitute a
significant threat to the northern
leopard frog at the species level now,
nor do we have indication that it will in
the future.
Roads
Roads have been shown to pose
barriers to northern leopard frog
dispersal, to contribute to nonpoint
source pollution, and to result in direct
mortality of northern leopard frogs
(Smith 2003, pp. 27, 38; Maxell 2000, p.
25; Fahrig et al. 1995, pp. 177–182). The
movements of adult northern leopard
frogs to breeding habitats during spring
rains and the extensive dispersal of
juveniles from breeding ponds in late
summer make this species vulnerable to
highway traffic (Orr et al. 1998, p. 93;
Langen et al. 2009, p. 111), and there are
many reports of large amounts of
leopard frog road mortality (see
references in Carr and Fahrig 2001, p.
1075; Glista et al. 2008, pp. 81–82;
Langen et al. 2009, p. 111). Road
building is often tied to other activities
such as urban, agricultural, and oil and
gas development, so roads may impact
leopard frogs directly and indirectly.
Bouchard et al. (2009, pp. 5–6) found
that the northern leopard frog’s inability
to avoid roads and their slow movement
make them particularly vulnerable to
road mortality and that roads could thus
result in negative effects to local
population abundance. Other studies
did not find any decreasing trends in
abundance for amphibian roadside
populations (Mazerolle 2004, p. 51).
Traffic density within 0.9 mi (1.5 km) of
occupied northern leopard frog habitat
may have negatively affected local frog
abundance, but it was unclear if results
were due to the observed road mortality,
pollution (e.g., vehicle emissions, road
runoff), or increased urbanization (Carr
and Fahrig 2001, p. 1074). Other studies
have also documented smaller
amphibian populations in the vicinity of
major roads and within landscapes with
high road densities than populations
where roads are distant and few (Langen
et al. 2009, p. 104). ‘‘Hotspots’’ for
northern leopard frog road mortality
tend to occur along causeways (road
segments with water on either side)
with wetland sites within 328 ft (100 m)
of the road (Langen et al. 2009, p. 110).
In summary, although research
indicates that roadside populations of
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northern leopard frogs may be adversely
impacted by roads and evidence shows
that individual frogs are certainly
impacted through road mortality, the
information assessed indicates these
impacts are localized and result in
effects to local frog abundance, not
population level impacts. While roads
occur throughout the range of the
northern leopard frog, the best available
information does not suggest that roads
constitute a significant threat to the
northern leopard frog at the species
level now, nor do we have indication
that they will in the future.
Forest Management
The northern leopard frog is
associated with forested as well as
grassland or open areas (Blomquist and
Hunter 2009, p. 150). Based upon broad
land cover and use, forest management
occurs in forested areas throughout the
range of the northern leopard frog
(USDA NRCS 2001). Timber harvest
activities may impact northern leopard
frog populations in several ways.
Clearcuts (areas where all trees are
removed) at breeding sites can result in
enhanced tadpole development through
increased water temperatures and food
production (Semlitsch et al. 2009, p.
859). However, clearcuts can also result
in negative effects to juvenile and adult
northern leopard frog movement due to
higher surface temperatures (from
canopy removal), and loss of soil-litter
moisture in upland habitats surrounding
breeding ponds, which affects the
species’ ability to move through these
areas into post-breeding habitat (Maxell
2000, pp. 12–14; Smith 2003, p. 29;
Semlitsch et al. 2009, p. 860). Research
on timber management and northern
leopard frog seasonal habitat
requirements found that northern
leopard frogs in the late spring and
summer used open, wet areas; frogs
used unharvested forest for longer
movements (Blomquist and Hunter
2009, p. 153). Forest management may
affect local populations of northern
leopard frogs by fragmenting habitats
and reducing landscape connectivity.
Forest management has the potential
to impact northern leopard frog
breeding, dispersal, and foraging
habitats in forested areas throughout its
range. However, the information we
reviewed does not indicate that forest
management, clearcutting in particular,
is occurring at a level or extent that
would result in impacts at the species
level. Therefore, the best available
information indicates forest
management is not a significant threat to
the northern leopard frog at the species
level now, nor do we have indication
that it will in the future.
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Groundwater Withdrawal
Throughout the range of the northern
leopard frog, particularly in the western
United States and Canada, naturally
geographically isolated (or depressional)
wetlands completely surrounded by
upland plant communities (such as the
prairie pothole wetlands in the upper
Midwestern United States and Canada)
and human-caused isolated wetlands
(such as natural wetlands that are no
longer connected to streams due to
roads or other development) are
important habitats for the northern
leopard frog. Many of these ‘‘isolated’’
wetlands appear to be disconnected
from other water sources, but are
hydrologically connected to other
wetlands or waters through sub-surface
or groundwater connections (Tiner
2003, p. 495). Because of this hydrologic
connection, groundwater withdrawal
can result in significant impacts to
wetland habitats and may result in
decreased surface water, decreased
recharge, and reduced water levels in
wetland and spring habitats (Alley et al.
1999, pp. 33–44; Alberta Northern
Leopard Frog Recovery Plan 2005, p. 7;
Wirt et al. 2005, pp. G1–11; Patten et al.
2008, p. 279). Specifically, groundwater
withdrawal can result in loss of
northern leopard frog breeding ponds
and spring- and riparian-associated
vegetation, and thus the loss or
modification of northern leopard frog
habitat (Alberta Northern Leopard Frog
Recovery Plan 2005, p. 7; Patten et al.
2008, p. 286). In addition, decreased
surface water levels may reduce the
water level in overwintering habitats,
which may result in the area freezing
and an increased risk of mortality as
wintering northern leopard frogs are
intolerant of freezing (see Biology
section).
Across the range of the northern
leopard frog, these habitats occur in the
prairie potholes region (see above), the
playas and springs of the Southwest, the
Sandhills wetlands in northern
Nebraska, channeled scablands in
eastern Washington, woodland vernal
pools of the northeastern United States,
and many other natural ponds
throughout the United States (Tiner
2003, p. 497). Within these areas, there
is regional and local information to
indicate that current and proposed
groundwater pumping may result in
reduced habitat for northern leopard
frogs, particularly in the arid West
(Tiner 2003, p. 513; Deacon et al. 2007).
Specifically, the BLM recently released
the Draft Environmental Impact
Statement for the Clark, Lincoln, and
White Pine Counties Groundwater
Development Project in Nevada (BLM
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2011). Based upon the modeling
analysis, the BLM predicts that northern
leopard frog habitat (for all life stages)
will be reduced in currently occupied
areas of central-eastern Nevada as a
result of the proposed action (BLM
2011, p. 3.7–45). This information
indicates that isolated wetland habitats
such as those in Spring Valley, Nevada,
may be significantly impacted by these
proposed groundwater withdrawals.
Groundwater depletion has been a
concern in the Southwest and High
Plains for many years due to the arid
climate and a lack of water resources;
however, increased demands on
groundwater resources have
overstressed aquifers in many areas of
the United States (Bartolino and
Cunningham 2003, p. 2). The Southwest
United States has experienced rapid
human population growth over the last
two decades in conjunction with longterm drought. This situation has
resulted in increased demand for water
resulting in impacts to wetland and
spring habitats from groundwater
pumping (Levick et al. 2008, pp. 70–71).
Brussard et al. (1998, pp. 505–542)
found that pumping of groundwater
from gold mines impacted spring
communities in the north-central region
of Nevada. Groundwater pumping by
the City of Albuquerque, New Mexico,
contributed to the loss of wetland
habitat in the Rio Grande valley as well
(Bogan 1998, pp. 562–563). In addition,
groundwater modeling studies indicate
that aquifers in eastern and southern
Nevada that supply water to springs
currently occupied by northern leopard
frogs may decline in response to
pumping in these areas to meet human
water demands (Schaefer and Harrill
1995, p. 46). However, streams and
wetlands in the Northeast, the High
Plains, the Pacific Northwest, and other
regions of the United States have also
been impacted by groundwater pumping
(Bartolino and Cunningham 2003, p. 2).
Impacts have included lowered water
tables, reduced surface flows,
desiccation of springs, and decreased
lengths of perennial streams as a result
of groundwater pumping (Bartolino and
Cunningham 2003, pp. 2–4). Currently,
there are many ongoing discussions
throughout the Southwest regarding
water supplies and how groundwater
pumping may be used to meet human
water demands. While specific plans
regarding how these future plans may
impact northern leopard frogs are
limited at this time in many areas, as
described above, the recently proposed
Clark, Lincoln, and White Pine Counties
Groundwater Development Project
(Bureau of Land Management 2011) is
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expected to reduce occupied northern
leopard frog habitat in Spring Valley,
Nevada.
As described above in the Oil and Gas
Development section, an increase in
natural gas mining (using hydraulic
fracturing) may also result in increases
in groundwater pumping throughout
Pennsylvania, Ohio, West Virginia,
Kentucky, Wyoming, Colorado,
Montana, and New Mexico (see Oil and
Gas Development above for further
discussion).
In summary, groundwater pumping
has likely contributed to localized and
possibly regional declines of northern
leopard frog habitat, particularly in
isolated wetlands and arid areas.
However, in assessing the impacts of
groundwater pumping on current
northern leopard frog populations,
impacts are most usually described as
potential effects to habitat availability.
These impacts are further described as
occurring at local and regional, rather
than species-wide, scales. Impacts to
isolated wetlands in particular are likely
to be localized. Further, impacts to
water resources in the arid West cannot
be extrapolated to the eastern United
States and eastern Canada due to
differences in climate and geography.
Finally, there is little to no information
about groundwater withdrawals in
Canada, and the northern leopard frog is
apparently still considered to be
widespread and relatively common in
the eastern United States and eastern
Canada. Therefore, the best available
information indicates groundwater
withdrawal is not a significant threat to
the northern leopard frog at the species
level now, nor do we have indication
that it will in the future.
Air Pollution
Acid precipitation may be affecting
northern leopard frog habitat in the
western United States, including the
Rocky Mountain region of Colorado,
New Mexico, and Wyoming. Acidic
water is an environmental stressor for
northern leopard frogs (Simon et al.
2002, p. 697), and leopard frog
abundance may be reduced in areas
where water acidification has occurred
(Pope et al. 2000, p. 2505). In the last
few decades, high-elevation aquatic
habitats have become more acidic (Corn
and Vertucci 1992, p. 363; Simon et al.
2002, p. 697), which may be a result of
air pollution. The emissions of certain
gases (principally sulfur dioxide and
nitrogen oxides) into the air may lead to
acid precipitation and the acidification
of aquatic habitats. Acidification of
aquatic habitats may result in decreased
reproductive capabilities of adult
northern leopard frogs, and mortality
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and developmental abnormalities in
northern leopard frog tadpoles (Simon
et al. 2002, p. 697). In addition, acid
precipitation can result in the direct
destruction of vegetation needed for
habitat (Environmental Protection
Agency 2000, pp. 48699–48701; Jezouit
2004, pp. 423–445). Nitrogen dioxide,
which also contributes to the formation
of acid rain (Baron et al. 2000, p. 352;
Fenn et al. 2003, p. 404; Jezouit 2004,
pp. 423–445; Environmental Protection
Agency 2005, p. 59594), can increase
the acidity of soils and aquatic
ecosystems; may contribute to
eutrophication (a process whereby
increased nutrients lead to decreased
dissolved oxygen); and may possibly
change plant community composition
(e.g., enhanced growth of invasive
species and shifts in phytoplankton
productivity) (Baron et al. 2000, p. 358;
Fenn et al. 2003, pp. 404–418).
However, effects from air pollution (in
the form of acid precipitation) are
currently only a consideration in highelevation habitats in the western United
States. Additionally, at this time, the
potential impacts are theoretical and
have not been shown to result in
population-level impacts to the species.
Therefore, the best available information
does not indicate that air pollution
constitutes a significant threat to
northern leopard frogs at the species
level now, nor do we have indication
that it will in the future.
Summary of Factor A
The northern leopard frog occupies a
wide geographic range across the United
States and Canada. Because it occurs
across such a large area, the habitats it
uses are subject to a number of impacts
that represent potential threats at
various scales. As discussed above,
these factors generally have been
historical in impact or are occurring
now and into the future at scales below
the species level, both individually and
in combination. Further, while there
have been declines noted in portions of
the range of the species, the frog is
apparently still considered to be
widespread and relatively common in
the eastern United States and eastern
Canada. Therefore, the best available
information indicates that the present or
threatened destruction, modification, or
curtailment of its habitat or range is not
a significant threat to the northern
leopard frog at the species level now,
nor do we have indication that it will in
the future.
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Factor B. Overutilization for
Commercial, Recreational, Scientific, or
Educational Purposes
Overutilization of the northern
leopard frog for commercial,
recreational, scientific, or educational
purposes is not reported to be a current
threat to the species in most of its range
(Woolington 2011, pers. comm.; Smith
2003, p. 21; Arizona Game and Fish
Department 2009, p. 2); however,
northern leopard frogs are harvested for
bait and for use in biology laboratories
in some portions of its range (Smith
2003, p. 21; Quinn 2009, pers. comm.;
Minnesota Department of Natural
Resources 2011a, p. 2). Northern
leopard frogs are collected for
commercial purposes in Nebraska,
Minnesota, and Wisconsin, and
historical collection in other States
likely contributed to long-term
population declines in some areas
(Lannoo et al. 1994, p. 317; Moriarty
1998, p. 168; Smith 2003, p. 21). From
1995–1999, approximately 174,772
northern leopard frogs were collected in
Nebraska to supply two biological
supply houses (Smith 2003, p. 21).
Northern leopard frogs in Minnesota
have been heavily collected for fish bait
and for the biological supply trade, and
there is little regulation on the
collection of frogs there (Moriarty 1998,
p. 168). Other States that have identified
overutilization as a potential effect to
the northern leopard frog include
Connecticut (Connecticut Department of
Environmental Protection 2005, p. 4–4–
4–5), Maine (Maine Department of
Inland Fisheries and Wildlife 2005, p.
109), Massachusetts (Massachusetts
Department of Fish and Wildlife 2006,
p. 407), and Michigan (Eagle et al. 2005,
Species of Greatest Conservation Need
152 of 242). However, other than
naming collection as a potential concern
or including conservation measures to
guard against overutilization in their
State wildlife action plans, we have no
information regarding the magnitude of
the potential threat of collection in these
States.
As noted earlier in the Status section,
northern leopard frog populations
crashed in 1973 in Minnesota, which
halted the commercial collections for
uses other than bait from 1974 to 1987.
Harvest records from the 1990s report
collections of 1,000 to 2,000 pounds of
frog per year, compared to reports in the
early 1970s that were in the 100,000pound-per-year range (Moriarty 1998, p.
168). According to North Dakota Game
and Fish Department records, 31,683
leopard frogs were collected by
wholesalers from 1996–2008. That is an
average of 2,463 frogs per year. The
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North Dakota Game and Fish
Department does not believe that this
level of use has impacted the population
(North Dakota Game and Fish
Department 2009, p. 2). There are no
restrictions in South Dakota regarding
the collection of northern leopard frogs,
and they are a legal bait species (limit
of 24 per day) (South Dakota
Department of Game, Fish, and Parks
2011, p. 23) and some South Dakota
tribal members collect and sell northern
leopard frogs to educational suppliers in
Minnesota (Quinn 2009, pers. comm.).
The northern leopard frog may also be
legally used for bait or other personal
uses (typically with a permit or license)
in Iowa, Missouri, Nebraska, New
Mexico, Illinois, Indiana, Kentucky,
Michigan, New York, Pennsylvania, and
Vermont (as identified in the Status
section above).
In 1971, Gibbs et al. (p. 1027)
described the frog trade and the decline
of northern leopard frogs throughout
most of their range. Due to the declines
noted by Gibbs et al. (1971), many States
began establishing laws to prevent
uncontrolled collecting. Today, many
State wildlife agencies, including those
in the western United States, use
commercial and collection regulations
to control human actions that may harm
wildlife populations, such as collection
of amphibians (Adams et al. 1995, p.
394; see also discussion in Status
section describing State collection laws
and under Factor D describing
regulatory mechanisms).
Though many States have established
regulations regarding the collection of
northern leopard frogs, wild-caught
amphibians are still traded on the global
market, and there is some concern as to
whether the take of wild-caught
individuals is biologically sustainable
(Schlaepfer et al. 2005, p. 257). Recent
research found that millions of
individuals, millions of body parts and
products, and more than 2,204,623
pounds (lbs) (1,000,000 kilograms (kg))
of amphibians and reptiles are shipped
across U.S. borders each year for
commercial purposes (Schlaepfer et al.
2005, p. 257). Greater than 2.5 million
whole, wild-caught amphibians and
reptiles were imported into the United
States between 1998 and 2002, but these
animals were not tracked by species
(Schlaepfer et al. 2005, p. 257).
Information tracked by the Service’s
Law Enforcement Management
Information System indicates that
249,233 lbs (113,050 kg) of northern
leopard frog were imported into the
United States between 1998 and 2002,
for food and research (Schlaepfer et al.
2005, p. 259). An additional 112,289
body parts and products and 1,177,970
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lbs (534,318 kg) of Lithobates frogs (not
identified to species), which likely
consisted in part of wild-caught
northern leopard frogs, were imported
into the United States during this same
timeframe. There were 361,858
Lithobates frogs imported or exported
from the United States with no species
specific identification (Schlaepfer et al.
2005, p. 261). We can conclude from
this information that the U.S. trade in
amphibians and reptiles, which is a
fraction of the world trade in terms of
wild-caught amphibians and reptiles
(Schlaepfer et al. 2005, p. 263), is
importing large numbers of northern
leopard frogs from Canada. There are no
data to indicate if this trade in wildcaught northern leopard frogs is
sustainable, and it may partially explain
why the frog continues to decline in
Ontario and other portions of eastern
Canada. Schloegel et al. (2009, p. 1424)
found that an average of 5.1 million
Ranid (= Lithobatid) frogs per year,
including live animals and their parts,
were imported into the United States
between 2000 and 2005. However, based
upon the reported origin of the frogs
(China and Taiwan), it is likely that
most of these imports were American
bullfrogs. However, there is evidence
that the commercial trade in
amphibians, particularly in American
bullfrogs, does result in the spread of
disease (such as ranaviral disease and
Batrachochytrium dendrobatidis, which
can cause the amphibian disease,
chytridiomycosis), and aids in the
spread of invasive species (Fisher and
Garner 2007, pp. 3–4; Picco and Collins
2008, p. 1588; Schloegel et al. 2009, pp.
1424–1425). In Arizona, northern
leopard frogs do appear in the pet trade,
either in local pet stores or through online suppliers (Arizona Game and Fish
Department 2009, p. 3), and
documented releases of eastern northern
leopard frogs into existing populations
have occurred (Hoffman and Blouin
2004a, pp. 150–151; Theimer et al.
2011, pp. 3, 30; O’Donnell et al. 2011,
p. 3), which may have genetic
implications for the ongoing
conservation of the species.
Summary of Factor B
Despite historic population and
regional declines, we do not have any
evidence of impacts to northern leopard
frogs at the species level from
overutilization for commercial,
recreational, scientific, or educational
purposes, and we have no information
that indicates this factor will become a
threat to the species in the future. The
significant declines and extirpations
within the range of the species have
occurred in areas other than those that
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have traditionally been subject to the
highest collection pressures. Further,
the collections appear to be occurring in
portions of the range that have
apparently stable populations.
Therefore, the best scientific and
commercial information available
indicates that overutilization for
commercial, recreational, scientific, or
educational purposes does not
constitute a significant threat to the
northern leopard frog at the species
level now, nor do we have indication
that it will in the future.
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Factor C. Disease or Predation
Disease
Fungal, viral, and bacterial diseases
may cause mass mortality and
contribute to population declines of
northern leopard frogs (Rorabaugh 2005,
pp. 575–577). Disease has caused mass
mortality in ranid and lithobatid frogs in
almost every western State in the United
States (Bradley et al. 2002; Muths et al.
2003; Briggs et al. 2005). There are
several fungal diseases that affect the
northern leopard frog (Faeh et al. 1998,
p. 263); of those, amphibian
chytridiomycosis caused by the fungus
Batrachochytrium dendrobatidis (Bd)
has likely had a large impact on
northern leopard frogs in the western
United States (Johnson et al. 2011, p.
564). Mortality from chytridiomycosis is
reported for several leopard frog species,
including the northern leopard frog, in
Arizona, British Columbia, California,
and Colorado (Bradley et al. 2002, pp.
206–212; Muths et al. 2003, p. 361;
Briggs et al. 2005, p. 3149; Committee
on the Status of Endangered Wildlife in
Canada 2009, p. 26; Johnson et al. 2011,
p. 564). Information in Muths et al.
(2003, p. 364) notes a northern leopard
frog museum specimen from Colorado
preserved in 1974 was examined
histologically and tested positive for Bd,
which means the presence of Bd in
Colorado can be traced back to the
1970s and is a possible contributing
factor to the extensive mortalities that
occurred there (Carey et al. 1999, p.
461). This time period is also when
extensive declines of northern leopard
frogs occurred throughout the western
United States and Canada, in places
such as Wisconsin, Alberta,
Saskatchewan, and Manitoba. Longcore
et al. (2006, p. 440) found that Bd is
widespread in the Northeast and the
highest prevalence of Bd in a Maine
species was the northern leopard frog.
However, there was no observed decline
in northern leopard frog populations
despite the significantly high infection
rate (Longcore et al. 2006, p. 441). It is
possible that northern leopard frogs in
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the eastern United States have
developed some resistance to Bd, or that
thermoregulatory behavior (such as
basking on a sunny day) may slow the
growth of the fungus (Longcore et al.
2006, pp. 441–442). It is currently not
known under what circumstances the
northern leopard frog is susceptible to
the lethal effects of chytridiomycosis,
but it remains a concern as the fungus
appears to be prevalent in the East and
in the West (Ellis 2011, pers. comm.;
Van Stralen 2011, pers. comm.), and
mortality in wild frogs in British
Columbia is thought to be the result of
chytridiomycosis.
Recent studies indicate that factors
such as habitat degradation, habitat
fragmentation, and climate change may
exacerbate the lethal effects of
chytridiomycosis on amphibian
populations (Carey et al. 1999, pp. 459–
472; Ouellet et al. 2005, p. 1437).
Habitat fragmentation may prevent
populations from recovering after lethal
outbreaks of chytridiomycosis (Ouellet
et al. 2005, p. 1437), and other stressors
such as water pollution may make
northern leopard frogs more susceptible
to chytridiomycosis (Carey et al. 1999,
pp. 459–472; Kiesecker et al. 2004, p.
138).
Saprolegniasis, a water-borne fungal
disease, may also affect populations of
northern leopard frogs (Faeh et al. 1998,
p. 263). However, this fungal disease is
usually secondary to other stressors
such as bacterial infections or trauma
(Faeh et al. 1998, p. 263). Saprolegnia
has been associated with embryonic dieoffs of ranid frogs in Oregon, and is
found in Columbia spotted frog (Rana
luteiventris) eggs in Idaho and Montana
(Patla and Keinath 2005, p. 43), but
there is no other information provided
to indicate that this disease is currently
impacting northern leopard frogs.
Faeh et al. (1998, pp. 260–261)
provided information regarding five
viral diseases that have and could
potentially affect the northern leopard
frog. These include the iridoviruses,
which include ranavirus, polyhedral
cytoplasmic amphibian virus, tadpole
edema virus, and frog erythrocytic virus.
Ranavirus may be extremely lethal, and
all life stages of frogs may acquire the
disease, although tadpoles are the most
susceptible to the disease (Daszak et al.
1999, p. 744). The loss of 80 to 90
percent of tadpoles in a population from
ranavirus may result in an 80 percent
loss of adult recruitment (survival of
individuals to sexual maturity and
joining the reproductive population),
which may negatively affect population
viability (Daszak et al. 1999, pp. 742–
745). The introduction of bullfrogs and
spread of tiger salamanders throughout
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the U.S. range of the northern leopard
frog may increase the potential of
ranavirus infection as both American
bullfrogs and tiger salamanders are
hosts for the ranavirus (Picco and
Collins 2008, p. 1588; Schloegel et al.
2009, p. 1424). Relatively recent mass
mortality events of northern leopard
frog metamorphs resulting from
ranavirus have been documented in
Ontario (Greer et al. 2005, p. 11).
Septicemia or ‘‘red leg’’ involves one
or a combination of hemolytic
(destructive to blood cells) bacteria that
enter the body via wounds or abrasions
(Faeh et al. 1998, p. 261). Septicemia
often results in death in individuals and
often results in mass mortality.
Septicemia may also have contributed to
northern leopard frog declines in the
Midwestern United States in the early
1970s (Koonz 1992, p. 20) and caused
declines in Colorado between 1974 and
1982 (Carey 1993, pp. 356–358).
However, ‘‘red leg’’ may be triggered by
a variety of environmental factors, and
it is unclear how it may be influencing
northern leopard frog declines in the
United States and Canada (McAllister et
al. 1999, p. 19).
Significant mortality events of
northern leopard frogs have been
attributable to disease (Rorabaugh 2005,
p. 575). However, with the exception of
chytridiomycosis, impacts to northern
leopard frogs associated with these
diseases appear to be localized.
Chytridiomycosis may be having
significant effects to northern leopard
frogs in the West, but does not appear
to be significantly affecting frogs in
other portions of its range as the frog is
apparently still considered to be
widespread and stable in the eastern
United States and eastern Canada.
Therefore, the best available information
does not indicate that disease is a
significant threat to the northern
leopard frog at the species level now,
nor do we have indication that it will in
the future.
Nonnative Species
The introduction of nonnative aquatic
animals, particularly American bullfrogs
and predatory fishes, has resulted in the
loss and decline of northern leopard
frogs throughout their range, but
particularly in the western United States
and Canada (Merrell 1968, p. 275; Hine
et al. 1981, p. 12; Hammerson 1982, pp.
115–116; Hayes and Jennings 1986, p.
491; Hecnar and M’Closkey 1997, p.
126; Livo et al. 1998, p. 4; Orr et al.
1998, p. 92; Maxell 2000, p. 144;
Hitchcock 2001, p. 63; Smith 2003, pp.
19–21; Alberta Northern Leopard Frog
Recovery Team 2005, p. 8; Rorabaugh
2005, p. 574; Smith and Keinath 2007,
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p. 24; Committee on the Status of
Endangered Wildlife in Canada 2009, p.
35). Northern leopard frogs typically
breed in waters without fish or aquatic
predators (Merrell 1977, p. 16; Hine et
al. 1981, p. 12). Nonnative animals
(including crayfish, American bullfrogs,
and fish) displace northern leopard
frogs by degrading habitat (e.g.,
destroying emergent vegetation,
increasing turbidity, reducing algal or
invertebrate populations) or through
direct predation on eggs, tadpoles, and
adult leopard frogs (Green 1997, p. 300).
American bullfrogs, which compete
with and prey on northern leopard
frogs, are thought to be a primary cause
of the widespread decline of northern
leopard frogs throughout the western
United States (Bury and Luckenbach
1976, p. 10; Hammerson 1982, pp. 115–
116; Kupferberg 1997, p. 1749; Livo et
al. 1998, p. 4). The American bullfrog is
native to the eastern and Midwestern
United States and historically had a
very wide native distribution that
excluded much of the western United
States. American bullfrogs currently are
not present in most of eastern Montana,
North Dakota, South Dakota, southern
Idaho, central and western Wyoming,
most of Utah, and a small portion of
northern Arizona and White Pine
County, Nevada (Casper and Hendricks
2005, p. 541). These areas where the
American bullfrog has yet to invade
coincide with some areas where the
northern leopard frog still occurs and, in
some cases, appears to be stable (such
as Nebraska, North Dakota, South
Dakota, and eastern Montana).
American bullfrogs have also been
introduced into British Columbia
(Weller and Green 1997, p. 320).
As previously described, northern
leopard frogs typically breed in fishless
waters (Merrell 1968, p. 275) and likely
have little natural defense against
predation by introduced fish (Smith and
Keinath 2007, p. 25). In Canada,
research shows that introduced
predaceous fish reduce the abundance
and diversity of frog populations,
including the northern leopard frog
(Hecnar and M’Closkey 1997, pp. 126–
127). Common carp (Cyprinus carpio)
cause increased turbidity and the
destruction of emergent vegetation,
which can displace northern leopard
frogs by modifying habitat, reducing
invertebrates, and eliminating algae
(McAllister et al. 1999, pp. 6–7).
Information from Bradford (2005, pp.
922–923) indicates that lithobatid frogs
in the western United States may be
more adversely affected than lithobatid
frogs in the eastern United States due to
their greater exposure to exotic,
introduced species. Because northern
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leopard frogs in the western United
States evolved in permanent or semipermanent waters without large aquatic
predators (Merrell 1968, p. 275), they
may be more vulnerable to predation by
introduced sport fish, bullfrogs, and
crayfish (Bradford 2005, p. 923). In
addition, literature studying the habitat
preferences of northern leopard frogs
from Ohio and Wisconsin indicates that
across the range of the northern leopard
frog, successful breeding habitats tend
to be free of predaceous fish due to
periodic drying (Merrell 1977, p. 16;
Hine et al. 1981, p. 12). This implies
that when nonnative species are
present, it is more likely that northern
leopard frogs will not successfully
reproduce.
Invasive plants may also impact
northern leopard frog habitat in the
western United States (Maxell 2000, pp.
21–22; Hitchcock 2001, pp. 5–6).
Tamarisk and other nonnative aquatic
and terrestrial plants alter riparian
habitats by forming dense stands that
exclude native amphibians (Maxell
2000, p. 21) and enhance the survival of
other introduced species, such as
American bullfrogs (Adams et al. 2003,
pp. 343–351; Maxell 2000, p. 21;
Hitchcock 2001, pp. 5–6, 62–66).
Effects to northern leopard frogs from
nonnative species are likely significant
in the western United States and
Canada, but information we reviewed
does not indicate nonnative species are
having significant impacts on northern
leopard frog populations in the eastern
portion of their range. Further, northern
leopard frogs are apparently considered
to be widespread and relatively
common in the eastern United States
and eastern Canada. Therefore, the best
available information indicates that
impacts associated with nonnative
species do not constitute a significant
threat to the northern leopard frog at the
species level now, nor do we have
indication that it will in the future.
61921
species level now, nor do we have
indication that it will in the future.
Summary of Factor C
Factor D. The Inadequacy of Existing
Regulatory Mechanisms
Under this factor, we examine
whether existing regulatory mechanisms
are inadequate to address the threats to
the northern leopard frog discussed
under Factors A, B, C, and E. The
Service considers regulatory
mechanisms to mean all regulatory and
statutory mechanisms that are related to
a comprehensive regime designed to
maintain a conserved wildlife
population. Section 4(b)(1)(A) of the Act
requires the Service to take into
account, ‘‘those efforts, if any, being
made by any State or foreign nation, or
any political subdivision of a State or
foreign nation, to protect such species
* * *.’’ We consider these efforts when
developing our threat analyses under all
five factors, and in particular under
Factor D. Therefore, under Factor D we
consider not only laws and regulations,
but other mechanisms that are part of a
regulatory process, such as management
plans, agreements, and conservation
practices.
Regulatory mechanisms, if they exist,
may preclude the need for listing if such
mechanisms are judged to adequately
address the threat to the species such
that listing is not warranted. Conversely,
threats are not ameliorated when not
addressed by existing applicable
regulatory mechanisms, or when the
existing mechanisms are not adequate
(or not adequately implemented or
enforced). Within its distribution in the
United States, the northern leopard frog
occurs on lands managed by a myriad of
Federal and State agencies, Native
American tribes, and private lands. In
Canada, the northern leopard frog
occurs on a similar variety of
jurisdictions. In this section, we review
actions taken by State and Federal
entities that effectively reduce or
remove threats to the northern leopard
frog.
Disease and predation have
undoubtedly contributed to the loss of
northern leopard frog populations
historically, particularly in the western
United States, and will likely continue
to impact northern leopard frogs in
some portions of its range at local or
regional scales. However, despite these
impacts, the frog is apparently still
considered to be widespread and
relatively common in the eastern United
States and eastern Canada. Therefore,
the best available information indicates
that impacts due to disease and
predation do not constitute a significant
threat to the northern leopard frog at the
Federal Laws and Regulations
The northern leopard frog is not
specifically covered by the provisions of
any Federal law or regulation. However,
there are Federal agencies that manage
lands occupied by northern leopard
frogs and laws that are applicable to the
management and conservation of the
species and its habitat.
All Federal agencies are required to
adhere to the National Environmental
Policy Act (NEPA) of 1969, as amended
(42 U.S.C. 4321 et seq.) for projects they
fund, authorize, or carry out. The
Council on Environmental Quality’s
regulations for implementing NEPA (40
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CFR parts 1500–1518) state that
environmental impact statements shall
include a discussion on the
environmental impacts of the various
project alternatives (including the
proposed action), any adverse
environmental effects that cannot be
avoided, and any irreversible or
irretrievable commitments of resources
involved (40 CFR part 1502). NEPA
itself is a disclosure law that provides
an opportunity for the public to submit
comments on the particular project and
propose other conservation measures
that may directly benefit listed or
sensitive fish and wildlife species;
however, it does not require subsequent
minimization or mitigation measures by
the Federal agency involved. Although
Federal agencies may include
conservation measures for listed species
as a result of the NEPA process, there is
no requirement that impacts to the
northern leopard frog from action
analyzed under NEPA would be
precluded. Any such measures are
typically voluntary in nature and are not
required by the statute. Additionally,
activities on non-Federal lands are
subject to NEPA if there is a Federal
nexus, such as permitting by the U.S.
Army Corps of Engineers or the Federal
Energy Regulatory Commission.
The Environmental Protection
Agency’s mission is to protect human
health and the environment. The agency
implements this mission by setting
standards for clean air, and regulating
pesticide use, chemical use, and water
pollution, among other actions. There
are a number of laws that are central to
this mission; however, the most
important in terms of preventing
impacts to northern leopard frogs are
likely the Clean Air Act of 1970 (42
U.S.C. 7401 et seq.), the Clean Water Act
of 1972 (33 U.S.C. 1251 et seq.), and the
Safe Drinking Water Act of 1974 (42
U.S.C. 300f et seq.). However, as
previously discussed, we have
determined that the adverse effects to
habitat for the northern leopard frog is
not nor is likely to have a species-level
impact.
The Clean Air Act is the Federal law
that regulates air emissions from
stationary and mobile sources. Among
other things, this law authorizes the
Environmental Protection Agency to
establish National Ambient Air Quality
Standards to protect public health and
public welfare and to regulate emissions
of hazardous air pollutants. The
Environmental Protection Agency is
required under the Clean Air Act to set
National Ambient Air Quality Standards
for six air pollutants (ozone, particulate
matter, carbon monoxide, nitrogen
oxides, sulfur dioxides, and lead).
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Evidence indicates that the National
Ambient Air Quality Standards for
sulfur dioxide, which contributes to the
formation of acid precipitation, may not
be adequate to protect aquatic
ecosystems from the impacts of acid
precipitation and acidification impacts,
and continued acid precipitation may
cause vegetation damage under the
current sulfur dioxide National Ambient
Air Quality Standards. Under the
current National Ambient Air Quality
Standards, acid precipitation is likely to
continue and may result in adverse
habitat effects from nitrogen deposition
(Baron et al. 2000, p. 365; Fenn et al.
2003, pp. 417–418).
The Clean Water Act establishes the
basic structure for surface water quality
protection in the United States. The
Environmental Protection Agency
employs a variety of regulatory and nonregulatory tools to reduce direct
pollutant discharges into waterways,
finance municipal wastewater treatment
facilities, and manage polluted runoff.
The Clean Water Act made it unlawful
to discharge any pollutant from a point
source into navigable waters, unless a
permit was obtained. The overall
objective of the Clean Water Act is to
restore and maintain the chemical,
physical, and biological integrity of the
nation’s waters so that they can support
‘‘the protection and propagation of fish,
shellfish, and wildlife and recreation in
and on the water.’’
The Safe Drinking Water Act is the
main Federal law that ensures the
quality of Americans’ drinking water.
Under the Safe Drinking Water Act, the
Environmental Protection Agency sets
standards for drinking water quality and
oversees the States, localities, and water
suppliers who implement those
standards. Section 1421 of the Safe
Drinking Water Act tasks the
Environmental Protection Agency with
protecting underground sources of
drinking water for all current and future
drinking water supplies across the
country.
The Service, Bureau of Land
Management (BLM), National Park
Service (NPS), and U.S. Forest Service
(Forest Service) are the primary Federal
agencies that manage lands that provide
habitat for the northern leopard frog.
The northern leopard frog occurs on
the Service’s National Wildlife Refuges
and Wetland Management Areas in
States throughout the northern leopard
frog’s U.S. range. The mission of the
National Wildlife Refuge System is to
administer a national network of lands
and waters for the conservation,
management, and, where appropriate,
restoration of the fish, wildlife, and
plant resources and their habitats within
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the United States for the benefit of
present and future generations of
Americans. Management on these
National Wildlife Refuges largely results
in the enhancement of northern leopard
frog habitat (Hultberg 2009, pers.
comm.; South Dakota Department of
Game, Fish and Parks 2009, pp. 2–3).
The northern leopard frog occurs on
BLM lands in Colorado, Idaho,
Montana, New Mexico, Nevada, and
Wyoming, and may also inhabit BLM
lands in North Dakota and South
Dakota. The frog has declined or is
absent from BLM lands in Arizona
(Clarkson and Rorabaugh 1989, p. 534),
Idaho (Makela 1998, pp. 8–9), Montana
(Maxell 2000, p. 144), Nevada
(Hitchcock 2001, p. 9), Washington
(McAllister et al. 1999, pp. 1–4), and
Wyoming (Smith and Keinath 2004, p.
57), based upon current ranges. BLM
lists the northern leopard frog as a
sensitive species in Colorado, Nevada,
Wyoming, and Montana; the species is
not listed as sensitive on BLM lands
elsewhere.
The Federal Land Policy and
Management Act of 1976 (FLPMA) (43
U.S.C. 1701 et seq.) is the primary
Federal law governing most land uses
on BLM-administered lands. Section
102(a)(8) of FLPMA (43 U.S.C.
1701(a)(8)) specifically recognizes the
public lands are to be managed to
provide food and habitat for fish and
wildlife.
BLM Manual section 6840 guides the
management of sensitive species in a
manner consistent with species and
habitat management objectives in land
use and implementation plans to
promote their conservation and to
minimize the likelihood and need for
listing under the Act (BLM 2008, p.
05V). This manual also requires that
resource management plans (RMPs)
should address sensitive species, and
that implementation ‘‘should consider
all site-specific methods and procedures
needed to bring species and their
habitats to the condition under which
management under the Bureau sensitive
species policies would no longer be
necessary’’ (BLM 2008, p. 2A1).
Where it has been designated as a
sensitive species under BLM Manual
6840, northern leopard frog
conservation must be addressed in the
development and implementation of
RMPs on BLM lands. RMPs are the basis
for all actions and authorizations
involving BLM-administered lands and
resources. Resource management plans
that include areas of northern leopard
frog habitat were completed beginning
in the 1980s. RMPs have been
developed or amended to incorporate
State or regionally developed rangeland
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health standards and guidelines, which
BLM developed beginning in 1995 (60
FR 9894, February 22, 1995). Standards
describe the specific conditions needed
for public land health, such as the
presence of streambank vegetation;
guidelines are the rangeland
management techniques designed to
achieve or maintain healthy public
lands, as defined by the standards.
Standards and guidelines must be
consistent with the fundamentals of
rangeland health, which include
watersheds that are in, or are making
significant progress toward, properly
functioning physical condition,
including their riparian-wetland and
aquatic components, and water quality
that complies with State water quality
standards. Areas and activities are
assessed to determine if the standards
are being achieved, and if not, actions
must be taken towards fulfilling the
standards (43 CFR 4180.1).
The Service has no specific
documentation of how implementation
of the rangeland health standards have
maintained or improved riparian or
wetland conditions within northern
leopard frog habitat on BLMadministered lands. The latest Public
Land Statistics report available (2010)
lists 23,618 acres (ac) (9,558 hectares
(ha)) of wetlands either in properly
functioning condition or functioning-atrisk with an upward trend, out of 49,764
total wetland ac (20,139 ha) on BLM
lands in Colorado, Idaho, Montana,
Nevada, New Mexico, North and South
Dakota, and Wyoming. The same report
lists 12,215 mi (19,658 km) of riparian
areas either in properly functioning
condition or functioning-at-risk with an
upward trend, out of 19,759 total miles
(31,799 km) on BLM lands in the same
States.
The BLM has regulatory authority for
oil and gas leasing on Federal lands and
on private lands with a Federal mineral
estate, as provided at subpart 3100
(Onshore Oil and Gas Leasing: General)
of Title 43 of the CFR, and they are
authorized to require stipulations as a
condition of issuing a lease. The BLM
has developed best management
practices to reduce habitat
fragmentation, loss, and degradation
from energy development. However, use
of these conditions is discretionary, and
the Service does not have information as
to how this authority has been applied.
The NPS manages portions of habitat
throughout the range of the northern
leopard frog. The NPS carries out its
responsibilities in parks and programs
under the authority of the National Park
Service Organic Act of 1916 (16 U.S.C.
1 et seq.). As defined in the National
Park Service Organic Act, the purpose of
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national parks is to conserve the scenery
and the natural and historic objects and
the wildlife therein and to provide for
the enjoyment of the same in such
manner and by such means as will leave
them unimpaired for the enjoyment of
future generations.
The Forest Service manages habitat
for northern leopard frogs in the western
United States on National Forests and
National Grasslands in several States,
including Arizona, Colorado, Idaho,
Minnesota, Montana, New Mexico,
North Dakota, South Dakota, Utah, and
Wyoming. Management of National
Forest System lands is guided
principally by the National Forest
Management Act (NFMA) (16 U.S.C.
1600 et seq.). The NFMA specifies that
all National Forests must have a Land
and Resource Management Plan (LRMP)
(16 U.S.C. 1604) to guide and set
standards for all natural resource
management activities on each National
Forest or National Grassland. The
NFMA requires the Forest Service to
incorporate standards and guidelines
into LRMPs (16 U.S.C. 1604(c)). The
Forest Service conducts NEPA analyses
on its LRMPs, which include provisions
to manage plant and animal
communities for diversity, based on the
suitability and capability of the specific
land area in order to meet overall
multiple-use objectives. The Forest
Service planning process is similar to
that of the BLM.
As described in the Status section,
populations of northern leopard frogs
have declined across most of the
western States on lands with
populations under Forest Service
jurisdiction. The northern leopard frog
is designated a ‘‘sensitive species’’ in
Forest Service Regions 1 (Northern
Region—northern Idaho, Montana,
North Dakota, and northwest South
Dakota), 2 (Rocky Mountain Region—
Colorado, Nebraska, most of South
Dakota and Wyoming), 3 (Southwest
Region—Arizona and New Mexico), 5
(Pacific Southwest Region—California),
and 6 (Pacific Northwest—Oregon and
Washington), but not in Regions 4
(Intermountain Region—southern Idaho,
Nevada, Utah, and western Wyoming)
and 9 (Eastern Region—includes all
eastern States and Minnesota). Sensitive
species status does not provide special
protection but requires, ‘‘an analysis of
the significance of adverse effects on the
population, its habitat, and on the
viability of the species as a whole’’
(Forest Service’s Manual at 2672.1).
Tribal Laws
Of the hundreds of tribal nations
located throughout the range of the
northern leopard frog in the United
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States and Canada, we only received
information regarding the northern
leopard frog from the Navajo Nation
(Arizona, New Mexico, and Utah), the
Fort Peck Tribes (Montana), the
Confederated Salish and Kootenai
Tribes of the Flathead Nation
(Montana), and the Sisseton-Wahpeton
Oyate (South Dakota). The Navajo
Nation provided us with specific
information regarding tribal laws. We
will continue to welcome any additional
information regarding the northern
leopard frog from tribal nations.
Navajo Endangered Species List
Group 2 species are protected under
Navajo Nation law. The Navajo Nation
Code (17 Navajo Nation Code section
507) makes it ‘‘unlawful for any person
to take, possess, transport, export,
process, sell or offer for sale or ship’’ a
Group 2 species. Under this Code,
‘‘take’’ means ‘‘the hunting, capturing,
killing in any manner or the attempt to
hunt, capture or kill in any manner
* * *.’’ Habitat protection, per se, is not
afforded under the Navajo Nation Code.
The Navajo Nation government,
pursuant to 2 Navajo Nation Code
section 164, reviews actions involving
the use of natural resources for
compliance with Navajo Nation law,
including the Navajo Endangered
Species Code. The Navajo Nation Fish
and Wildlife Department, through the
section 164 review process, advises the
tribal Resources Committee and the
Navajo Nation Council whether
proposed natural resources projects are
in compliance with the Navajo
Endangered Species Code. The
Resources Committee has the power to
give final approval for any land
exchanges, non-mineral leases, right-ofways, permits, and other licenses and
interests on Navajo land in accordance
with applicable and Federal and Navajo
Nation laws. The Resources Committee
recommends all actions involving the
approval of mineral agreements, land
acquisitions, and energy development
agreements to the Navajo Nation
Council. Some protection for northern
leopard frog habitat may be provided
through this review.
State Laws and Regulations
Only 1 of the 33 States assessed in the
Status section above has listed the
northern leopard frog under a State
wildlife conservation law. In 2000, the
Washington Department of Fish and
Wildlife listed the northern leopard frog
as an endangered species under the
Endangered, Threatened, and Sensitive
Species Classification (Washington
Administrative Code, Title 232, Chapter
12, Section 014). However, because
northern leopard frogs are currently
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known from only two sites (Germaine
and Hays 2009, p. 537) in Washington
State, this regulatory mechanism
protects relatively few individuals.
Arizona, California, Colorado,
Connecticut, Idaho, Indiana, Kentucky,
Maine, Massachusetts, Michigan,
Missouri, Montana, Nebraska, Nevada,
New Hampshire, New Mexico, Oregon,
Pennsylvania, Rhode Island, Utah, West
Virginia, and Wyoming included the
northern leopard frog specifically as a
species of greatest conservation need or
species of concern in their State wildlife
action plans (designations vary by State
as described in Status section above);
however, this designation provides no
regulatory protection to the species or
its habitat. The northern leopard frog is
not considered a species of concern in
Illinois, Iowa, Minnesota, New York,
North Dakota, Ohio, South Dakota,
Texas, Vermont, and Wisconsin.
Several States have laws that provide
some protection of northern leopard
frogs in regards to collection, as
discussed in the Status section above.
These laws and regulations generally
preclude or limit collection without a
permit, but do not preclude impacts to
habitat.
In summary, State wildlife
conservation laws generally provide for
an inconsistent network of protections
for the northern leopard frog. While take
is prohibited in some States, and the
species is afforded some management
consideration in project planning, the
laws generally do not preclude impacts
to habitat. However, 23 of the 33 States
within the range of the northern leopard
frog have indicated commitment
through their State wildlife action plans
to implementing conservation actions
and habitat enhancement projects to
benefit the northern leopard frog.
International Laws and Regulations
The northern leopard frog, Rocky
Mountain population, is listed as
endangered under the Federal Species at
Risk Act (Statues of Canada 2002, c.29)
in Canada. The Species at Risk Act,
passed December 12, 2002, is a
commitment by the Canadian
government to prevent the extinction of
wildlife and provide the necessary
actions for the recovery of the species
deemed endangered. Wildlife species
listed under the Species at Risk Act are
provided with legal protection to avoid
extinction resulting from human
activities (Government of Canada
Species at Risk Public Registry 2011).
The northern leopard frog is also Red
Listed as endangered under the British
Columbia Wildlife Act (Revised Statutes
of British Columbia 1996, c. 488), which
prohibits the killing or collecting of
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amphibians or keeping them in captivity
without a permit. In British Columbia,
the one remaining northern leopard frog
population is located in the Creston
Valley Wildlife Management Area
(Committee on the Status of Endangered
Wildlife in Canada 2009, p. 42). The
Creston Valley Wildlife Management
area is protected by the British
Columbia government and by the
Convention on Wetlands of
International Importance (‘‘Ramsar
Convention,’’ Ramsar, Iran 1971), where
Creston Valley was designated a
Wetland of International Importance on
February 21, 1994. The Convention on
Wetlands is an intergovernmental treaty
that provides the framework for national
action and international cooperation for
the conservation and wise use of
wetlands and their resources. In
addition, other provincial legislation,
including the Fish Protection Act (Bill
25–1997), the Creston Valley Wildlife
Act (Revised Statutes of British
Columbia 1996, c. 84), the Integrated
Pest Management Act (Statues of British
Columbia 2003, c. 58), and the Riparian
Areas Regulation (Fish Protection Act,
British Columbia Regulation 376/2004)
provide habitat protection and
enhancement to the remaining northern
leopard frog population (Committee on
the Status of Endangered Species in
Canada 2009, p. vi).
The northern leopard frog was listed
as threatened in Schedule 6 of Alberta’s
Wildlife Act (Revised Statutes of Alberta
2000, Chapter W–10), based on a
decline in the number of populations,
the fragmentation of occupied habitats,
and limited population dispersal
capabilities of the species (Alberta
Northern Leopard Frog Recovery Team
2005, p. 1). As a result of the listing, the
Alberta Northern Leopard Frog
Recovery Plan was created and is
currently being implemented (Alberta
Northern Leopard Frog Recovery Team
2005). In Saskatchewan, the northern
leopard frog is currently on the
province’s Interim Species at Risk List
(Wildlife Act 1998, Chapter W–13.12)
and is protected in provincial and
national parks (Committee on the Status
of Endangered Wildlife in Canada 2009,
p. vi). The national status of the western
boreal and prairie population (which
includes Alberta, Saskatchewan,
Manitoba, and the Northwest
Territories) was evaluated in 1998 and
2002, and the northern leopard frog was
designated a Species of Special Concern
(Committee on the Status of Endangered
Wildlife in Canada 2004, p. 20). As a
result of the national designation, a
management plan was required to be
developed for the western boreal and
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prairie population. Although the
northern leopard frog has no national or
provincial status in Eastern Canada, the
species is protected on Federal lands
managed by Parks Canada (national
parks and historic sites), Environment
Canada (national wildlife areas), and the
Department of Defense (Committee on
the Status of Endangered Wildlife in
Canada 2009, p. vi).
As noted in the BACKGROUND section
above, the northern leopard frog
population in western Canada is small
and fragmented, but as one proceeds
east, the number of northern leopard
frog populations and their known status,
based on the best available information,
improves. Where the northern leopard
frog has and likely continues to decline
in western Canada, there is no
information to indicate that the species
is threatened by the inadequacy of
existing regulatory mechanisms in
Canada.
Summary of Factor D
While northern leopard frog
conservation has been addressed in
some State, Federal, and international
plans, laws, regulations, and policies,
none of these have applicability
throughout the range of the northern
leopard frog sufficient to provide
effective population-level conservation.
However, we have found in the analysis
of the other four factors (A, B, C, and E)
that there are no threats that currently
rise to a level such that they
significantly impact the northern
leopard frog at the species level.
Therefore, we conclude that the best
scientific and commercial information
available indicates that the inadequacy
of existing regulatory mechanisms is not
a significant threat to the northern
leopard frog at the species level now,
nor do we have indication that it will in
the future.
Factor E. Other Natural or Manmade
Factors Affecting Its Continued
Existence
Pesticides
Even at low levels, pesticides can
contribute to local declines or
extirpation of northern leopard frog
populations, particularly in areas that
are in close proximity to heavy or
frequent pesticide use because tadpole
and larval stages are sensitive to even
low-level pesticide contamination
(Berrill et al. 1997, p. 244). The effects
to northern leopard frogs from
pesticides, including herbicides,
piscicides (chemical substances
poisonous to fish), and insecticides,
vary, but information indicates that the
species is negatively affected both
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acutely and via sublethal symptoms by
several pesticides and chemicals
(rotenone, Roundup, atrazine,
malathion, copper sulfate, and fenthion)
that are commonly used in the United
States (Stebbins and Cohen 1995, pp.
215–216; Fordham 1999, p. 125; Hayes
et al. 2002, pp. 895–896; Beasley et al.
2005, p. 86; Patla 2005, p. 275; Relyea
2005, p. 353; Rorabaugh 2005, p. 576).
Pesticide contamination of surface
waters in the United States is extensive,
and concentrations of pesticides are
frequently greater than water-quality
benchmarks for aquatic life and fisheating wildlife (Gilliom et al. 2006, p. 8).
Of the streams analyzed as part of the
National Water Quality Assessment
Program, 57 percent contained one or
more pesticides that exceeded at least
one aquatic life protection benchmark
(Gilliom et al. 2006, p. 8), which may
result in decreased habitat quality,
malformations, and decreased fitness of
northern leopard frogs (Rorabaugh 2005,
p. 576).
While northern leopard frogs may be
exposed to pesticides in a number of
ways, they are most significantly
exposed to pesticides when run-off from
agricultural and urban areas reaches
occupied habitats. Exposure to pesticide
run-off can influence parasitic
community structure and seasonal
recruitment in northern leopard frogs
(King et al. 2008, p. 20). Berrill et al.
(1997, p. 243) found that tadpoles
(including northern leopard frog
tadpoles) are extremely sensitive (i.e.,
they experience paralysis and death) to
exposure of one pesticide at a time;
pesticides in combination likely have
more severe effects. Ouellet et al. (1997,
p. 97) examined northern leopard frogs
in agricultural and non-agricultural
ponds in Quebec and found that frogs in
the agricultural ponds had a variety of
hind limb malformations. The authors
identified agricultural pesticides as a
potential causal agent. Pesticide
exposure not only can cause
malformations in frogs (Lannoo 2008,
pp. 142–144), but contact with
pesticides has been found to increase
amphibians vulnerability to Ribeiroia
(trematode) and other parasitic
infections, which are also known to
cause frog malformations (Kiesecker
2002, p. 9903; Lannoo 2008; Rohr et al.
2008, p. 1237). In addition, increased
nitrates from fertilizers can also result in
adverse effects to amphibian
development and survival (Marco et al.
1999, p. 2837; Rouse et al. 1999, pp.
800–802). Therefore, although northern
leopard frogs were not specifically
tested for pesticides in the examples
from Washington or Quebec, it is
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plausible that the habitat alteration and
subsequent contamination of aquatic
habitats with pesticides contributed to
the decline of northern leopard frogs in
these areas. Agrichemical pollution is
also thought to be a factor in declining
amphibian populations in Nebraska and
Quebec (Beasley et al. 2005, p. 86;
McCleod 2005, p. 293; King et al. 2008,
p. 20).
Based upon the above information,
exposure to pesticides has likely
contributed to northern leopard frog
population extirpations throughout their
range. While the magnitude of these
impacts is conceivably high in localized
areas, pesticide use is not ubiquitous
throughout the range of the northern
leopard frog; thus pesticide use is likely
not resulting in impacts at regional and
species-level scales. Further, despite
ongoing exposure to pesticides, the
northern leopard frog is apparently still
considered to be widespread and
common in the eastern United States
and eastern Canada. Therefore, the best
available scientific information
indicates that pesticide use does not
constitute a significant threat to the
northern leopard frog at the species
level now, nor do we have indication
that it will in the future.
Malformations
Within the last 15 to 20 years,
malformed northern leopard frogs have
been reported with increasing frequency
in the United States, particularly in
Minnesota, North Dakota, South Dakota,
and Vermont (Helgen et al. 1998, p. 288;
Sessions 2003, p. 168; Johnson and
Lunde 2005, p. 124; Rorabaugh 2005, p.
576). Malformations are also reported
from Colorado, Indiana, Iowa, Michigan,
Missouri, Montana, Ohio, Quebec, and
Wisconsin (Converse et al. 2000, p. 163;
Johnson and Lunde 2005, pp. 124–128;
Rorabaugh 2005, p. 575; North
American Center for Reporting
Amphibian Malformations 2006). Noted
malformations have included limb
deformities, multiple or missing limbs,
jaw deformities, stunted growth,
multiple eyes, missing eyes, and various
other growths (Helgen et al. 1998, pp.
288–297; Hoppe 2005, p. 104).
Malformations are believed to be caused
by a variety of natural and manmade
factors, including trematode parasites,
pesticides, ultraviolet-B radiation,
predation attempts, and water
contamination (Helgen et al. 1998, pp.
294–297; Blaustein and Johnson 2003,
pp. 87–91; Sessions 2003, p. 168;
Johnson and Lunde 2005, pp. 124–138;),
but are generally linked to humaninduced changes in aquatic habitats
(Meteyer et al. 2000, pp. 151–171;
Johnson and Lunde 2005, pp. 130–136;
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Lannoo 2008, pp. 105–110, 197). These
malformations typically lead to
mortality as behavior and physical
mobility (such as swimming, hopping,
and feeding) are compromised to the
point of affecting individual fitness
(Helgen et al. 1998, p. 289; Hoppe 2005,
pp. 105–108). Northern leopard frogs
tend to be one of the most common
species found with malformations
(Lannoo 2008, p. 207).
Malformations are a concern because
they affect the ability of individual and
local populations of northern leopard
frogs to survive, and because they are a
likely indicator of decreased water
quality and of decreased overall habitat
quality. However, as stated above, there
are likely many causes of malformations
in northern leopard frogs that have to do
with local, site-specific conditions and
are likely not the result of the same
causal agent throughout the range of the
northern leopard frog (Lannoo 2008, p.
200). Further, the diversity of habitat
used by northern leopard frogs may
provide some protection against the
variety of agents that seem to result in
malformation at the local scale. The rate
of malformations in some local
populations of northern leopard frogs
may result in significant effects to these
populations; however, the impact of
malformations on the northern leopard
frog at the species level is not known to
be significant. Therefore, based on the
best available information, we conclude
that malformations are not a significant
threat to northern leopard frogs at the
species level now, nor do we have
indication that it will in the future.
Climate Change
‘‘Climate’’ refers to an area’s long-term
average weather statistics (typically for
at least 20- or 30-year periods),
including the mean and variation of
surface variables such as temperature,
precipitation, and wind, whereas
‘‘climate change’’ refers to a change in
the mean and/or variability of climate
properties that persists for an extended
period (typically decades or longer),
whether due to natural processes or
human activity (Intergovernmental
Panel on Climate Change (IPCC) 2007a,
p. 78). Although changes in climate
occur continuously over geological time,
changes are now occurring at an
accelerated rate. For example, at
continental, regional and ocean basin
scales, recent observed changes in longterm trends include: A substantial
increase in precipitation in eastern parts
of North American and South America,
northern Europe, and northern and
central Asia, and an increase in intense
tropical cyclone activity in the North
Atlantic since about 1970 (IPCC 2007a,
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p. 30); and an increase in annual
average temperature of more than 2 °F
(1.1 °C) across US since 1960 (Global
Climate Change Impacts in the United
States (GCCIUS) 2009, p. 27). Examples
of observed changes in the physical
environment include: An increase in
global average sea level, and declines in
mountain glaciers and average snow
cover in both the northern and southern
hemispheres (IPCC 2007a, p. 30);
substantial and accelerating reductions
in Arctic sea-ice (e.g., Comiso et al.
2008, p. 1), and a variety of changes in
ecosystem processes, the distribution of
species, and the timing of seasonal
events (e.g., GCCIUS 2009, pp. 79–88).
The IPCC used Atmosphere-Ocean
General Circulation Models and various
greenhouse gas emissions scenarios to
make projections of climate change
globally and for broad regions through
the 21st century (Meehl et al. 2007, p.
753; Randall et al. 2007, pp. 596–599),
and reported these projections using a
framework for characterizing certainty
(Solomon et al. 2007, pp. 22–23).
Examples include: (1) It is virtually
certain there will be warmer and more
frequent hot days and nights over most
of the earth’s land areas; (2) it is very
likely there will be increased frequency
of warm spells and heat waves over
most land areas, and the frequency of
heavy precipitation events will increase
over most areas; and (3) it is likely that
increases will occur in the incidence of
extreme high sea level (excludes
tsunamis), intense tropical cyclone
activity, and the area affected by
droughts (IPCC 2007b, p. 8, Table
SPM.2). More recent analyses using a
different global model and comparing
other emissions scenarios resulted in
similar projections of global temperature
change across the different approaches
(Prinn et al. 2011, pp. 527, 529).
All models (not just those involving
climate change) have some uncertainty
associated with projections due to
assumptions used, data available, and
features of the models; with regard to
climate change this includes factors
such as assumptions related to
emissions scenarios, internal climate
variability and differences among
models. Despite this, however, under all
global models and emissions scenarios,
the overall projected trajectory of
surface air temperature is one of
increased warming compared to current
conditions (Meehl et al. 2007, p. 762;
Prinn et al. 2011, p. 527). Climate
models, emissions scenarios, and
associated assumptions, data, and
analytical techniques will continue to
be refined, as will interpretations of
projections, as more information
becomes available. For instance, some
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changes in conditions are occurring
more rapidly than initially projected,
such as melting of Arctic sea ice
(Comiso et al. 2008, p. 1; Polyak et al.
2010, p. 1797), and since 2000 the
observed emissions of greenhouse gases,
which are a key influence on climate
change, have been occurring at the midto higher levels of the various emissions
scenarios developed in the late 1990’s
and used by the IPPC for making
projections (e.g., Raupach et al. 2007,
Figure 1, p. 10289; Manning et al. 2010,
Figure 1, p. 377; Pielke et al. 2008,
entire). Also, the best scientific and
commercial data available indicates that
average global surface air temperature is
increasing and several climate-related
changes are occurring and will continue
for many decades even if emissions are
stabilized soon (e.g. Meehl et al. 2007,
pp. 822–829; Church et al. 2010, pp.
411–412; Gillett et al. 2011, entire).
Changes in climate can have a variety
of direct and indirect impacts on
species, and can exacerbate the effects
of other threats. Rather than assessing
‘‘climate change’’ as a single threat in
and of itself, we examine the potential
consequences to species and their
habitats that arise from changes in
environmental conditions associated
with various aspects of climate change.
For example, climate-related changes to
habitats, predator-prey relationships,
disease and disease vectors, or
conditions that exceed the physiological
tolerances of a species, occurring
individually or in combination, may
affect the status of a species.
Vulnerability to climate change impacts
is a function of sensitivity to those
changes, exposure to those changes, and
adaptive capacity (IPCC 2007, p. 89;
Glick et al 2011, pp. 19–22). As
described above, in evaluating the status
of a species, the Service uses the best
scientific and commercial data
available, and this includes
consideration of direct and indirect
effects of climate change. As is the case
with all potential threats, if a species is
currently affected or is expected to be
affected by one or more climate-related
impacts, this does not necessarily mean
the species is a threatened or
endangered species as defined under the
Act. If a species is listed as threatened
or endangered, this knowledge
regarding its vulnerability to, and
impacts from, climate-associated
changes in environmental conditions
can be used to help devise appropriate
strategies for its recovery.
While projections from global climate
model simulations are informative and
in some cases are the only or the best
scientific information available, various
downscaling methods are being used to
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provide higher-resolution projections
that are more relevant to the spatial
scales used to assess impacts to a given
species (see Glick et al, 2011, pp. 58–
61). With regard to the area of analysis
for the northern leopard frog, specific
downscaled projections are not
available for all the parts of its range,
but we do have more generalized
information. In North America, climate
change is likely to constrain already
over-allocated water resources, resulting
in increased competition among
agricultural, municipal, industrial, and
ecological uses of water (Bates et al.
2008, p. 102). Of particular note are the
expected changes in surface and
groundwater hydrology. As the rate of
warming accelerates, the timing,
volume, quality, and spatial distribution
of fresh water available to most areas in
North America will change (Bates et al.
2008, p. 102; Johnson et al. 2010, p.
138). These changes will likely affect
the quality and quantity of northern
leopard frog habitat. Some areas,
especially in the arid West, will likely
see decreases in habitat, while other
areas may experience stable or
increasing available habitat. The
freshwater wetland habitats the
northern leopard frog depends upon for
breeding and overwintering, particularly
in the arid Southwest (Arizona,
Colorado, New Mexico, Nevada, and
Utah) and the prairie potholes region
(Alberta, Iowa, Manitoba, Minnesota,
Montana, North Dakota, Saskatchewan,
and South Dakota) are expected to be
particularly sensitive to climate change
(Johnson et al. 2010, p. 128). Increases
in drought and seasonal precipitation
may have profound impacts to habitat;
however, we are unable to reliably
predict how changes in precipitation
will affect current and future northern
leopard frog habitat throughout the
species’ range.
Many experts expect that amphibians
may be among the first vertebrates to
exhibit broad-scale changes in response
to global climate change (Reaser and
Blaustein 2005, p. 61). The northern
leopard frog is at the upper limit of its
physiological tolerance to temperature
and dryness throughout the arid and
semi-arid habitats in the western United
States (Hammerson 1999, pp. 146–147;
Hitchcock 2001, pp. 18–19; Rorabaugh
2005, p. 577). As such, if the predictions
for temperature increases are realized,
these arid areas may no longer support
the species. In addition, the northern
leopard frog frequently depends upon
small, ephemeral wetlands for breeding
habitats (Merrell 1968, p. 275), and due
to habitat fragmentation, the presence of
nonnative aquatic species, and other
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factors (such as agricultural and urban
development, and roads), the leopard
frog is bounded by dispersal barriers
throughout its range (Rorabaugh 2005,
p. 577). Species persistence is greater for
species occupying larger patches of their
historical range (Channell and Lomolino
2000, pp. 84–86). Because northern
leopard frogs occupy relatively small
patches of habitat compared to their
historical distribution in some portions
of their range, we may expect that
climate change could result in further
fragmentation of those populations in
those portions of its range. In other
words, the frogs may exist in smaller
and smaller patches that are more
remote from the core of their historical
range.
As described above, changes in the
quality and quantity of habitat are likely
to occur throughout the range of the
northern leopard frog. There are likely
to be additional impacts to frogs in some
portions of it range because of these
changes. Climate change impacts in the
arid and semi-arid areas could include
earlier reproduction and more rapid
development of larva due to more a
more advanced spring, decreased
mobility due to drier conditions, and
shorter hibernation periods due to
longer ice-free periods in the winter
(Carey and Alexander 2003, pp. 111–
121; Patla and Keinath 2005, pp. 44–46;
Johnson et al. 2010, p. 133). Higher
summer temperatures may result in high
egg mortality (in response to freezing
temperatures that may follow earlier
breeding times) and in drying of
breeding habitats prior to
metamorphosis (in response to
increased evaporation rate) (Smith 2003,
p. 34). Climate change may also cause
frogs to experience increased
physiological stress and decreased
immune system function, possibly
leading to disease outbreaks (Carey and
Alexander 2003, pp. 111–121; Pounds et
al. 2006, pp. 161–167). Northern
leopard frog populations at lower
elevations are likely to show changes in
phenology sooner than those at higher
elevations (Corn 2003, pp. 622–625).
Based upon the extended droughts in
the Southwest and changes the Service
has noted to northern leopard frog
habitats in Arizona and New Mexico
(Service 2007, pp. 38–41), it is likely
that climate change may continue to
reduce the amount of habitat available
for northern leopard frogs, particularly
in the western United States.
Climate change may result in
significant impacts to some portions of
the range of the northern leopard frog
and may synergistically result in
increased impacts from disease and
other factors discussed above. The
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overall impacts of climate change will
likely be very different across the range
of the northern leopard frog, and it is
difficult to predict how these effects
will manifest themselves in terms of
species-level impacts. There may be
decreases in habitat in some areas, and
increases in other portions of the range.
As a result, it is possible that the
species’ range could expand, contract,
or shift. However, we do not know
enough about the capacity of this
species to adapt to changing
environmental conditions to make
reliable predictions about future largescale range contractions or shifts in
response to climate change. In the arid
West, it is likely that the predictions for
greater variability in temperature and
precipitation will result in further
decreases in wetland habitats, which
may exacerbate the negative interactions
of native and nonnative species using
wetted habitats. However, we expect
that there may be portions of the
species’ range that may experience more
favorable conditions, such as increased
precipitation and temperature, that will
positively affect habitat for the northern
leopard frog. In conclusion, although we
believe climate change will impact some
northern leopard frog habitats in the
future, the information we reviewed
does not indicate that climate change
will adversely impact northern leopard
frogs at the species level. Therefore,
based on the best available information,
we conclude that climate change is not
a significant threat individually or in
combination to the northern leopard
frog at the species level now, nor do we
have indication that it will in the future.
Summary of Factor E
The northern leopard frog occupies a
wide geographic range across the United
States and Canada. As we have stated
earlier, because it occurs across such a
large area, the habitats it uses are subject
to a number of impacts from pesticide
use and climate change, and the species
is subject to malformations that will
impact local, and possibly even
regional, populations. However, the
wide diversity of wetland and upland
habitats that are currently used by the
northern leopard frog across its range
may provide some protection in the
future from changing climates and
possibly from the variety of potential
agents that cause malformations.
Therefore, the best available information
indicates that other natural and
manmade factors do not constitute a
significant threat to the northern
leopard frog at the species level now,
nor do we have indication that it will in
the future.
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Finding
As required by the Act, we considered
the five factors in assessing whether the
northern leopard frog is endangered or
threatened throughout all of its range
(i.e., in danger of extinction now or in
the foreseeable future). We examined
the best scientific and commercial
information available regarding the past,
present, and future threats faced by the
northern leopard frog. We reviewed the
petition, information available in our
files, and other available published and
unpublished information, and we
consulted with other Federal, State, and
tribal agencies.
There have been historical impacts to
the northern leopard frog, in particular.
The loss and degradation of wetland
habitat, introduction of nonnative
species, and disease, have resulted in
local and regional extirpations of the
species throughout its range, but
particularly in the western United States
and Canada, as described in the
Background section above. Further,
some of the threats discussed in this
finding work in concert with one
another to cumulatively create
situations that potentially impact the
northern leopard frog beyond the scope
of each individual threat. It is likely that
for such a widespread species as the
northern leopard frog, causes of decline
are dependent upon multiple factors or
causes. This is particularly true since
the northern leopard frog uses both
terrestrial and aquatic habitats. For
example, as discussed under Factor A,
degradation of wetland habitats,
resulting from agricultural use and the
application of pesticides, results in
increased immunosuppression and risk
of parasitic infection in northern
leopard frogs (Christin et al. 2003, pp.
1129–1130). These factors can also
enhance the potential for
malformations, which can result in
decreased fitness, and subsequent
declines of northern leopard frog
populations. Malformations (discussed
under Factor E) are likely the result of
multiple causes. Lannoo (2008)
describes the search for ‘‘the’’ cause of
amphibian malformations, but
eloquently determines in his
comprehensive review that there is
likely no one cause, but many factors
that can result in malformations.
Similarly, Thiemann and Wassersug
(2000) found that the presence of
predators and parasites also increased
the susceptibility of Rana (=Lithobates)
tadpoles to trematode infection by
causing tadpoles to decrease their
activity levels. They found that the
combination of such stressors as
increased predator loads (such as from
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widespread predator introductions as
discussed under Factor C), parasite
infection, and pesticide pollution may
synergistically result in increased
impacts to tadpoles, which could be
another factor in declining populations.
However, even where these factors may
work cumulatively to impact northern
leopard frogs, the best available
information does not indicate that
current populations are being impacted
significantly at scales above the
population or regional levels.
In summary, in order to determine
that the northern leopard frog warrants
listing throughout its range, we must
find that the best available information
indicates it is in danger of extinction
now or in the foreseeable future. The
phrase ‘‘in danger of extinction’’
requires a showing that the species is
actually likely in danger of extinction
now, or likely to become so in the
foreseeable future, not merely a showing
that the species is facing threats. We
must show that the threats are operative
on the species such that the species
meets the definition of threatened or
endangered (i.e., in danger of extinction
now or in the foreseeable future). The
northern leopard frog occupies a wide
geographic range across the United
States and Canada. Because it occurs
across such a large area, it is subject to
a number of impacts that represent
potential threats at various scales. The
number of threats the species has faced
and continues to face may appear
significant; however, as discussed
above, the factors affecting the northern
leopard frog have generally been
historical in impact or are occurring
now and into the future at scales below
the species level as indicated by the
presence of apparently stable
populations in large areas of its range.
Further, while there have been regional
declines noted in the range of the
species, particularly in the western
portions of the United States and
Canada, the frog is apparently still
considered to be widespread and
relatively common in the eastern United
States and eastern Canada.
Based on our review of the best
available scientific and commercial
information pertaining to the five
factors, we find that threats, alone or
cumulatively, are not of sufficient
magnitude at the species level to
indicate that the northern leopard frog
is in danger of extinction, or likely to
become in danger of extinction within
the foreseeable future, throughout all of
its range.
Significant Portion of Its Range
The Act defines ‘‘endangered species’’
as any species which is ‘‘in danger of
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extinction throughout all or a significant
portion of its range,’’ and ‘‘threatened
species’’ as any species which is ‘‘likely
to become an endangered species within
the foreseeable future throughout all or
a significant portion of its range.’’ The
definition of ‘‘species’’ is also relevant
to this discussion. The Act defines the
term ‘‘species’’ as follows: ‘‘The term
‘species’ includes any subspecies of fish
or wildlife or plants, and any distinct
population segment of any species of
vertebrate fish or wildlife which
interbreeds when mature.’’ The phrase
‘‘significant portion of its range’’ is not
defined by the statute, and we have
never addressed in our regulations: (1)
The consequences of a determination
that a species is either endangered or
likely to become so throughout a
significant portion of its range, but not
throughout all of its range; or (2) what
qualifies a portion of a range as
‘‘significant.’’
Two recent district court decisions
have addressed whether the ‘‘significant
portion of its range’’ language allows the
Service to list or protect less than all
members of a defined ‘‘species’’:
Defenders of Wildlife v. Salazar, 729 F.
Supp. 2d 1207 (D. Mont. 2010),
concerning the Service’s delisting of the
Northern Rocky Mountain gray wolf (74
FR 15123, Apr. 2, 2009); and WildEarth
Guardians v. Salazar, 2010 U.S. Dist.
LEXIS 105253 (D. Ariz. Sept. 30, 2010),
concerning the Service’s 2008 finding
on a petition to list the Gunnison’s
prairie dog (73 FR 6660, February 5,
2008). The Service had asserted in both
of these determinations that it had
authority, in effect, to protect only some
members of a ‘‘species,’’ as defined by
the Act (i.e., species, subspecies, or
DPS), under the Act. Both courts ruled
that the determinations were arbitrary
and capricious on the grounds that this
approach violated the plain and
unambiguous language of the Act. The
courts concluded that reading the
‘‘significant portion of its range’’
language to allow protecting only a
portion of a species’ range is
inconsistent with the Act’s definition of
‘‘species.’’ The courts concluded that
once a determination is made that a
species (i.e., species, subspecies, or
DPS) meets the definition of
‘‘endangered species’’ or ‘‘threatened
species,’’ it must be placed on the list
in its entirety and the Act’s protections
applied consistently to all members of
that species (subject to modification of
protections through special rules under
sections 4(d) and 10(j) of the Act).
Consistent with that interpretation,
and for the purposes of this finding, we
interpret the phrase ‘‘significant portion
of its range’’ in the Act’s definitions of
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‘‘endangered species’’ and ‘‘threatened
species’’ to provide an independent
basis for listing; thus there are two
situations (or factual bases) under which
a species would qualify for listing: a
species may be endangered or
threatened throughout all of its range; or
a species may be endangered or
threatened in only a significant portion
of its range. If a species is in danger of
extinction throughout a significant
portion of its range, it, the species, is an
‘‘endangered species.’’ The same
analysis applies to ‘‘threatened species.’’
Therefore, the consequence of finding
that a species is endangered or
threatened in only a significant portion
of its range is that the entire species will
be listed as endangered or threatened,
respectively, and the Act’s protections
will be applied across the species’ entire
range.
We conclude, for the purposes of this
finding, that interpreting the
‘‘significant portion of its range’’ phrase
as providing an independent basis for
listing is the best interpretation of the
Act because it is consistent with the
purposes and the plain meaning of the
key definitions of the Act; it does not
conflict with established past agency
practice, as no consistent, long-term
agency practice has been established;
and it is consistent with the judicial
opinions that have most closely
examined this issue. Having concluded
that the phrase ‘‘significant portion of
its range’’ provides an independent
basis for listing and protecting the entire
species, we next turn to the meaning of
‘‘significant’’ to determine the threshold
for when such an independent basis for
listing exists.
Although there are potentially many
ways to determine whether a portion of
a species’ range is ‘‘significant,’’ we
conclude, for the purposes of this
finding, that the significance of the
portion of the range should be
determined based on its biological
contribution to the conservation of the
species. For this reason, we describe the
threshold for ‘‘significant’’ in terms of
an increase in the risk of extinction for
the species. We conclude that a
biologically based definition of
‘‘significant’’ best conforms to the
purposes of the Act, is consistent with
judicial interpretations, and best
ensures species’ conservation. Thus, for
the purposes of this finding, a portion
of the range of a species is ‘‘significant’’
if its contribution to the viability of the
species is so important that, without
that portion, the species would be in
danger of extinction.
We evaluate biological significance
based on the principles of conservation
biology using the concepts of
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redundancy, resiliency, and
representation. Resiliency describes the
characteristics of a species that allow it
to recover from periodic disturbance.
Redundancy (having multiple
populations distributed across the
landscape) may be needed to provide a
margin of safety for the species to
withstand catastrophic events.
Representation (the range of variation
found in a species) ensures that the
species’ adaptive capabilities are
conserved. Redundancy, resiliency, and
representation are not independent of
each other, and some characteristics of
a species or area may contribute to all
three. For example, distribution across a
wide variety of habitats is an indicator
of representation, but it may also
indicate a broad geographic distribution
contributing to redundancy (decreasing
the chance that any one event affects the
entire species), and the likelihood that
some habitat types are less susceptible
to certain threats, contributing to
resiliency (the ability of the species to
recover from disturbance). None of these
concepts is intended to be mutually
exclusive, and a portion of a species’
range may be determined to be
‘‘significant’’ due to its contributions
under any one of these concepts.
For the purposes of this finding, we
determine if a portion’s biological
contribution is so important that the
portion qualifies as ‘‘significant’’ by
asking whether, without that portion,
the representation, redundancy, or
resiliency of the species would be so
impaired that the species would have an
increased vulnerability to threats to the
point that the overall species would be
in danger of extinction (i.e., would be
‘‘endangered’’). Conversely, we would
not consider the portion of the range at
issue to be ‘‘significant’’ if there is
sufficient resiliency, redundancy, and
representation elsewhere in the species’
range that the species would not be in
danger of extinction throughout its
range if the population in that portion
of the range in question became
extirpated (extinct locally).
We recognize that this definition of
‘‘significant’’ establishes a threshold
that is relatively high. On the one hand,
given that the consequences of finding
a species to be endangered or threatened
in a significant portion of its range
would be listing the species throughout
its entire range, it is important to use a
threshold for ‘‘significant’’ that is
robust. It would not be meaningful or
appropriate to establish a very low
threshold whereby a portion of the
range can be considered ‘‘significant’’
even if only a negligible increase in
extinction risk would result from its
loss. Because nearly any portion of a
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species’ range can be said to contribute
some increment to a species’ viability,
use of such a low threshold would
require us to impose restrictions and
expend conservation resources
disproportionately to conservation
benefit: listing would be rangewide,
even if only a portion of the range of
minor conservation importance to the
species is imperiled. On the other hand,
it would be inappropriate to establish a
threshold for ‘‘significant’’ that is too
high. This would be the case if the
standard were, for example, that a
portion of the range can be considered
‘‘significant’’ only if threats in that
portion result in the entire species’
being currently endangered or
threatened. Such a high bar would not
give the ‘‘significant portion of its
range’’ phrase independent meaning, as
the Ninth Circuit held in Defenders of
Wildlife v. Norton, 258 F.3d 1136 (9th
Cir. 2001).
The definition of ‘‘significant’’ used in
this finding carefully balances these
concerns. By setting a relatively high
threshold, we minimize the degree to
which restrictions will be imposed or
resources expended that do not
contribute substantially to species
conservation. But we have not set the
threshold so high that the phrase ‘‘in a
significant portion of its range’’ loses
independent meaning. Specifically, we
have not set the threshold as high as it
was under the interpretation presented
by the Service in the Defenders
litigation. Under that interpretation, the
portion of the range would have to be
so important that current imperilment
there would mean that the species
would be currently imperiled
everywhere. Under the definition of
‘‘significant’’ used in this finding, the
portion of the range need not rise to
such an exceptionally high level of
biological significance. We recognize
that if the species is imperiled in a
portion that rises to that level of
biological significance, then we should
conclude that the species is in fact
imperiled throughout all of its range,
and that we would not need to rely on
the ‘‘significant portion of its range’’
language for such a listing. Rather,
under this interpretation we ask
whether the species would be in danger
of extinction everywhere without that
portion, i.e., if that portion were
completely extirpated.
The range of a species can
theoretically be divided into portions in
an infinite number of ways. However,
there is no purpose to analyzing
portions of the range that have no
reasonable potential to be significant or
to analyzing portions of the range in
which there is no reasonable potential
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for the species to be endangered or
threatened. To identify only those
portions that warrant further
consideration, we determine whether
there is substantial information
indicating that: (1) The portions may be
‘‘significant,’’ and (2) the species may be
in danger of extinction there or likely to
become so within the foreseeable future.
Depending on the biology of the species,
its range, and the threats it faces, it
might be more efficient for us to address
the significance question first or the
status question first. Thus, if we
determine that a portion of the range is
not ‘‘significant,’’ we do not need to
determine whether the species is
endangered or threatened there; if we
determine that the species is not
endangered or threatened in a portion of
its range, we do not need to determine
if that portion is ‘‘significant.’’ In
practice, a key part of the portion status
analysis is whether the threats are
geographically concentrated in some
way. If the threats to the species are
essentially uniform throughout its
range, no portion is likely to warrant
further consideration. Moreover, if any
concentration of threats applies only to
portions of the species’ range that
clearly would not meet the biologically
based definition of ‘‘significant,’’ such
portions will not warrant further
consideration.
After reviewing the potential threats
throughout the range of the northern
leopard frog, we determine that there is
a portion of the range that could be
considered to have concentrated threats.
We defined this area, which we are
calling the westernmost portion, as
including the current range of the
northern leopard frog within British
Columbia and Alberta, Canada, and
Washington, eastern Oregon (if any
native populations remain), Idaho,
California (if any native populations
remain), Nevada, Utah, Arizona, New
Mexico, Colorado, and the portions of
Wyoming and Montana that are west of
the Continental Divide. Below, we
outline the elevated threats found
within this westernmost portion of the
northern leopard frog’s range (see
‘‘Summary of Information Pertaining to
the Five Factors’’ for complete
discussion). We then assess whether
this portion of the species’ range may
meet the biologically based definition of
‘‘significant,’’ that is, whether the
contribution of this portion of the
northern leopard frog’s range to the
viability of the species is so important
that without this westernmost portion of
the range, the species would be in
danger of extinction.
This westernmost portion of the
northern leopard frog’s range has
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experienced significant declines and
continues to experience impacts, likely
resulting from the influence of multiple
contributing factors, but primarily
resulting from the combination of
habitat loss, the spread of American
bullfrogs and predaceous fish into
otherwise suitable breeding habitats,
disease, and increased variability in
temperature and precipitation
(Rorabaugh 2005, pp. 575–577; Smith
and Keinath 2007, pp. 29–31;
Committee on the Status of Endangered
Species in Canada 2009, pp. 31–35;
Johnson et al. 2011, p. 557). As
described above in Species Information,
the northern leopard frog depends upon
a landscape that includes breeding
ponds, upland foraging areas,
overwintering aquatic habitats, and
connectivity among habitats and
between populations (Pope et al. 2000,
p. 2505; Smith 2003, pp. 6–15;
Rorabaugh 2005, pp. 571–575). The
destruction and degradation of wetland
and riparian habitat is thought to
represent the most widespread impact
to northern leopard frog populations in
Arizona (Arizona Game and Fish
Department 2009, p. 1), Colorado
(Colorado Division of Wildlife 2009, p.
2), Idaho (Idaho Department of Fish and
Game 2005, Northern leopard frog
species account), Montana (Montana
Fish Wildlife and Parks 2009, p. 2),
Nevada (Nevada Department of Wildlife
2009, p. 4), New Mexico (New Mexico
Department of Game and Fish 2009, p.
3), and Alberta, Canada (Alberta
Northern Leopard Frog Recovery Team
2005, p. 6). The loss of aquatic habitats
has been compounded by the spread of
the American bullfrog and nonnative
fish in the West. These species predate
on and compete with all life stages of
northern leopard frogs and have further
stressed northern leopard frog
populations in this westernmost
portion, likely contributing to
population declines. Based upon the
extended droughts in the Southwest and
changes the Service has noted to
northern leopard frog habitats in
Arizona and New Mexico (Service 2007,
pp. 38–41), it is likely that increased
variability in temperature and
precipitation will continue to reduce the
amount of breeding and wintering
habitat available for northern leopard
frogs, particularly in the western United
States.
After identifying elevated threats in
the westernmost portion of the range of
the northern leopard frog, we next
consider whether this portion of the
range should be considered a
‘‘significant portion of its range’’ based
on the framework laid out above. In
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order for the westernmost portion of the
range to be considered significant, we
consider whether there is sufficient
resiliency, redundancy, and
representation in the remaining portion
of the range (which includes the species
in the rest of its range; hereafter referred
to as the eastern portion of the range)
such that the northern leopard frog
would not be in danger of extinction if
the westernmost portion of the range in
question became extirpated (extinct
locally). Our analysis, described below,
finds that the westernmost portion of
the range does not meet this definition
of significant, because even without that
portion of the range the species,
rangewide, would not be in danger of
extinction.
To determine whether or not the
westernmost portion of the range is
‘‘significant,’’ we considered the
species’ resiliency, redundancy, and
representation in the remainder (i.e., the
eastern portion) of its range. For
resiliency, we evaluated whether the
eastern portion of the range of the
northern leopard frog, without the
westernmost portion, would maintain
the characteristics necessary to allow
the species to recover from periodic
disturbance. The eastern portion we
refer to here includes Saskatchewan,
eastern Montana, and eastern Wyoming,
and continues east through Canada and
the United States through the rest of the
range of the northern leopard frog. This
area encompasses a large proportion of
the range of the species and contains a
variety of wetland and upland habitats
necessary to provide breeding and
overwintering habitats, and habitat
linkages. This area is also sufficiently
large as to provide a margin of safety for
the species to withstand disturbance
events. We conclude that the eastern
portion of the range of the northern
leopard frog is sufficiently resilient that
even without the westernmost portion
of its range, the species would not be in
danger of extinction.
As part of our evaluation of
redundancy, we evaluated whether the
eastern portion of the range of the
northern leopard frog, without the
westernmost portion, would have
enough populations sufficiently
distributed across the landscape to
allow the species to withstand
catastrophic events. Based upon what
we know of the current population
status in the eastern portion of the
range, there are multiple areas (such as
South Dakota, North Dakota, Ohio,
Ontario, Vermont, New York, and
Quebec) where the northern leopard
frog is currently maintaining stable,
widespread populations. These areas are
sufficient in size and apparent
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distribution to serve as core areas from
which northern leopard frogs can
recolonize areas that could be subject to
catastrophic future events (such as
widespread flooding or drought). We
conclude that the eastern portion of the
range of the northern leopard frog is
sufficiently redundant that even without
the westernmost portion of its range, the
species would not be in danger of
extinction.
In our evaluation of representation,
we considered whether the eastern
portion of the range of the northern
leopard frog, without the westernmost
portion, contains enough variation to
ensure that the species’ adaptive
capabilities are conserved (such that the
genetic, morphological, physiological,
behavioral, or ecological diversity of the
species overall is maintained). Based
upon our current knowledge of the
northern leopard frog, we do not have
evidence of morphological,
physiological, or behavioral differences
between individuals from the
westernmost portion of the range and
individuals in the eastern portion of the
range. Although the westernmost
portion of the range is located on the
periphery of the species’ overall range,
the eastern portion contains large areas
that represent an important genetic
evolutionary history between eastern
and western northern leopard frogs
(Hoffman and Blouin 2004a, 2004b;
Wilson et al. 2008). This important
genetic information is represented
within the defined eastern area and
would not be lost if the westernmost
portion of the range were extirpated. In
addition, although not well studied,
there are likely broad ecological
differences between northern leopard
frogs in the westernmost portion of the
range compared to those in the eastern
portion of the range that result from the
geographical differences in habitat,
climate, and species interactions. We
recognize the ecological importance of
conserving peripheral, as well as
interior, populations of wide-ranging
species. However, due to the diversity of
areas the northern leopard frog occupies
in the large eastern portion of its range,
it is likely that sufficient ecological
adaptation potential would be
maintained to ensure ecological
representation. We conclude that the
eastern portion of the range of the
northern leopard frog is sufficiently
representative that even without the
westernmost portion of its range, the
species would not be in danger of
extinction.
Based on our analysis, we find that
the eastern portion of the range of the
northern leopard frog contains sufficient
redundancy, resiliency, and
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representation that, even without the
contribution of the westernmost portion
of the species’ range, the northern
leopard frog would not be in danger of
extinction. Therefore, we find that the
westernmost portion of the northern
leopard frog does not constitute a
significant portion of the species’ range.
In conclusion, based on a review of
the best available information, we find
the northern leopard frog is not in
danger of extinction now or in the
foreseeable future throughout all or a
significant portion of its range and,
therefore, does not warrant listing at this
time.
We request that you submit any new
information concerning the distribution
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and status of, or threats to, the northern
leopard frog to our Arizona Ecological
Services Office (see ADDRESSES)
whenever it becomes available. New
information will help us monitor the
northern leopard frog and encourage its
conservation. If an emergency situation
develops for the northern leopard frog
or any other species, we will act to
provide immediate protection.
References Cited
A complete list of references cited is
available on the Internet at https://
www.regulations.gov and upon request
from the Arizona Ecological Services
Office (see ADDRESSES section).
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Authors
The primary authors of this notice are
the staff members of the Arizona
Ecological Services Office.
Authority
The authority for this section is
section 4 of the Endangered Species Act
of 1973, as amended (16 U.S.C. 1531 et
seq.).
Dated: September 26, 2011.
Gregory E. Siekaniec,
Acting Director, Fish and Wildlife Service.
[FR Doc. 2011–25498 Filed 10–4–11; 8:45 am]
BILLING CODE 4310–55–P
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Agencies
[Federal Register Volume 76, Number 193 (Wednesday, October 5, 2011)]
[Proposed Rules]
[Pages 61896-61931]
From the Federal Register Online via the Government Printing Office [www.gpo.gov]
[FR Doc No: 2011-25498]
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Vol. 76
Wednesday,
No. 193
October 5, 2011
Part V
Department of the Interior
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Fish and Wildlife Service
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50 CFR Part 17
Endangered and Threatened Wildlife and Plants; 12-Month Finding on a
Petition To List the Northern Leopard Frog in the Western United States
as Threatened; Proposed Rule
Federal Register / Vol. 76 , No. 193 / Wednesday, October 5, 2011 /
Proposed Rules
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DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS-R2-ES-2009-0030; 92210-1111-FY08-B2]
Endangered and Threatened Wildlife and Plants; 12-Month Finding
on a Petition To List the Northern Leopard Frog in the Western United
States as Threatened
AGENCY: Fish and Wildlife Service, Interior.
ACTION: Notice of 12-month petition finding.
-----------------------------------------------------------------------
SUMMARY: We, the U.S. Fish and Wildlife Service, announce a 12-month
finding on a petition to list the northern leopard frog (Lithobates
(=Rana) pipiens) under the Endangered Species Act of 1973, as amended
(Act). After review of the best scientific and commercial information,
we find that listing the northern leopard frog is not warranted at this
time. However, we ask the public to submit to us any new information
that becomes available concerning threats to the northern leopard frog
or its habitat at any time.
DATES: The finding announced in this document was made on October 5,
2011.
ADDRESSES: This finding is available on the Internet at https://www.regulations.gov at Docket Number FWS-R2-ES-2009-0030. Supporting
documentation we used in preparing this finding is available for public
inspection, by appointment, during normal business hours at the U.S.
Fish and Wildlife Service, Arizona Ecological Services Office, 2321
West Royal Palm Road, Suite 103, Phoenix, AZ 85021. Please submit any
new information, materials, comments, or questions concerning this
finding to the above street address.
FOR FURTHER INFORMATION CONTACT: Steven L. Spangle, Field Supervisor,
Arizona Ecological Services Office (see ADDRESSES); by telephone at
(602) 242-0210; or by facsimile at (602) 242-2513. If you use a
telecommunications device for the deaf (TDD), please call the Federal
Information Relay Service (FIRS) at 800-877-8339.
SUPPLEMENTARY INFORMATION:
Background
Section 4(b)(3)(B) of the Act (16 U.S.C. 1531 et seq.), requires
that, for any petition to revise the Federal Lists of Endangered and
Threatened Wildlife and Plants that contains substantial scientific or
commercial information that listing the species may be warranted, we
make a finding within 12 months of the date of receipt of the petition.
In our finding, we are required to determine if the petitioned action
is: (a) Not warranted, (b) warranted, or (c) warranted, but the
immediate proposal of a regulation implementing the petitioned action
is precluded by other pending proposals to determine whether species
are endangered or threatened, and expeditious progress is being made to
add or remove qualified species from the Federal Lists of Endangered
and Threatened Wildlife and Plants. Section 4(b)(3)(C) of the Act
requires that we treat a petition for which the requested action is
found to be warranted but precluded as though resubmitted on the date
of such finding, that is, requiring a subsequent finding to be made
within 12 months. We must publish these 12-month findings in the
Federal Register.
Previous Federal Actions
On June 5, 2006, we received a petition from the Center for Native
Ecosystems, Biodiversity Conservation Alliance, Defenders of Black
Hills, Forest Guardians, Center for Biological Diversity, The Ark
Initiative, Native Ecosystems Council, Rocky Mountain Clean Air Action,
and Mr. Jeremy Nichols requesting that the northern leopard frog
(Lithobates (=Rana) pipiens) occurring in the western United States
(Arizona, California, Colorado, Idaho, Iowa, Minnesota, Missouri,
Montana, Nebraska, Nevada, New Mexico, North Dakota, Oregon, South
Dakota, Texas, Utah, Washington, and Wyoming) be listed as a threatened
distinct population segment (DPS) under the Act. The petition contained
detailed information on the natural history, biology, current status,
and distribution of the western population of the northern leopard
frog. It also contained information on what the petitioners reported as
potential threats to the western population of the northern leopard
frog such as habitat loss and degradation, predation and competition by
nonnative species, disease, water pollution, climate change, and other
factors. We acknowledged the receipt of the petition in a letter to the
petitioners dated August 7, 2006. That letter explained that we would
not be able to address their petition at that time. The reason for this
delay was that responding to court orders and settlement agreements for
other listing actions required nearly all of our listing funding.
In reviewing the petition, there were two issues for which the
Service requested clarification from the petitioners. We were
petitioned to list the population west of the Mississippi River and the
Great Lakes region in the United States and south of the international
boundary between the United States and Canada. However, although
Wisconsin is located west of the Great Lakes region, the petition map
did not show Wisconsin as a part of the petition, and the status of the
species is not mentioned in that State. Therefore, we requested that
the petitioners clarify whether they intended to include or exclude
Wisconsin from the petitioned DPS. We also sought clarification as to
whether the petitioners were requesting that we review only the western
U.S. population of the northern leopard frog as a DPS or if they were
also requesting us to consider listing the entire species or a
significant portion of the range of the species. The petitioners
responded to our clarification request in a letter dated February 8,
2008, requesting we review whether Wisconsin should be included in the
western U.S. population of the northern leopard frog. In addition, the
petitioners clarified that, if we find that listing the western U.S.
population of northern leopard frogs as a DPS is not warranted, we
review whether listing the entire species is warranted because of
threats in a significant portion of its range.
On July 1, 2009, we published our 90-day finding (74 FR 31389) that
the petition presented substantial scientific information indicating
that listing the western population of the northern leopard frog may be
warranted, and we initiated a status review to determine if listing the
species as a DPS or throughout all or a significant portion of its
range is warranted. Our July 1, 2009, 90-day finding opened a 60-day
period to send us information for our status review. On October 28,
2009, we reopened this information solicitation period for our status
review for an additional 30 days, ending November 27, 2009 (74 FR
55525). This notice constitutes our 12-month finding on the February 8,
2008, petition to list the northern leopard frog.
Species Information
Below we provide information relevant to understanding the analysis
of information pertaining to the five factors. See Rorabaugh (2005) for
a more complete description of the distribution and life history of the
northern leopard frog.
Taxonomy
The northern leopard frog is in the family Ranidae (Lannoo 2005, p.
371), the true frogs, and is one of about 28 species within the genus
Lithobates (formerly Rana (Frost et al. 2006, p. 10;
[[Page 61897]]
Frost et al. 2008, pp. 7-8)) that occur in North America (Lannoo 2005,
p. 371). For more than a century, nomenclatural and taxonomic confusion
has surrounded members of the Lithobates (=Rana) complex (Moore 1944,
p. 349; Pace 1974, pp. 11-16; Merrell 1977, pp. 1-2; Hillis et al.
1983, p. 132 among others), and there is a wealth of literature from
the late 1800s to present day that has attempted to accurately describe
the different species and geographic variation within the complex.
Until recently, all North American ranid frogs (frogs in the family
Ranidae) were included within the single genus Rana. However, Frost et
al. (2006, p. 10) placed most of these species into the genus
Lithobates. This change is recognized by the Committee on Standard
English and Scientific Names, which is the official names list of the
American Society of Ichthyologists and Herpetologists, the
Herpetologists' League, and the Society for the Study of Amphibians and
Reptiles (Frost et al. 2008, pp. 7-8). Accordingly, the Service also
recognizes and accepts Frost et al.'s (2008) Lithobates classification.
Physical Description
The northern leopard frog is a slim, smooth-skinned green, brown,
or sometimes yellow-green frog with webbed hind feet. The frog is
covered with large, oval dark spots, each of which is surrounded by a
lighter halo or border (Stebbins 2003, pp. 234-235). The snout (nose)
is pointed and the tympanum (eardrum) is round and approximately equal
in diameter to the eye (Baxter and Stone 1980, p. 41). Northern leopard
frogs have a white stripe on the upper jaw and the dorsolateral folds
(paired, glandular ridges that run along each side of the back from
behind the eyes to the rear) are light cream to yellow and are
continuous (not broken posteriorly). The belly is white to cream-
colored, and the posterior thigh has a light background color with dark
spots. There are two different color morphs (variants) of the northern
leopard frog that most often occur in western Minnesota, eastern North
Dakota, and South Dakota (Rorabaugh 2005, p. 570; McKinnel1 et al.
2005, p. 7). These color morphs do occur in other locations (for
example, see Ammon 2002, p. 11), but they are most prevalent in
Minnesota, North Dakota, and South Dakota, as described above. The
burnsi morph lacks dorsal spots and the kandiyohi morph has mottled
pigment patches (speckles) between the dorsal spots. Adult body lengths
(snout-vent) range from 2 to 4.5 inches (in) (5 to 11 centimeters (cm))
(Stebbins 2003, p. 234). Females average slightly larger than males
(Leonard et al. 1993, p. 138; Werner et al. 2004, p. 97). Subadult, or
recently metamorphosed frogs (see Biology section below), range in
length from 1 to 2 in (2 to 5 cm) (Merrell 1977, pp. 10-11). During the
breeding season, males have enlarged or swollen thumbs (innermost
digit) on forefeet, and vocal sacs are not apparent except when the
frog is calling (Baxter and Stone 1980, p. 41; Hammerson 1999, p. 145).
The typical breeding call is a prolonged ``snore'' followed by a series
of stuttering croaks or chuckles that tend to accelerate towards the
end (Hammerson 1999, p. 145). These vocalizations may be interspersed
with chuckling sounds (Stebbins 2003, p. 235).
Northern leopard frogs deposit their egg masses underwater in
clusters, which they attach to vegetation. Eggs are laid in a single
orange- to grapefruit-sized globular clump, and may be laid
individually or communally in groups (Nussbaum et al. 1983, p. 182).
Each egg mass may contain 645 to 7,648 individual eggs (Rorabaugh 2005,
p. 572). The eggs hatch into tadpoles. Tadpoles (the larval stage in
the lifecycle of the frog) are dark green to brown above with metallic
flecking, and a cream to white translucent underside (Werner et al.
2004, p. 97). Tadpoles metamorphose into young frogs. For a detailed
description of northern leopard frog tadpoles, see Scott and Jennings
(1985, pp. 4-16).
Distribution
The northern leopard frog historically ranged from Newfoundland and
southern Quebec, south through the northeast portions of the United
States to West Virginia, west across the Canadian provinces and
northern and central portions of the United States to British Columbia,
Oregon, Washington, and northern California, and south to Arizona, New
Mexico, and extreme western Texas (Rorabaugh 2005, p. 570).
Current range maps tend to show an extensive and connected
distribution for the northern leopard frog; however, its actual
distribution is sparse and fragmented in Washington, Oregon, Idaho,
California, Nevada, Arizona, New Mexico, Utah, Colorado, western
Montana, and western Wyoming in the western United States (Rorabaugh
2005, pp. 570-571), throughout New England (New Hampshire Fish and Game
Department 2005, pp. A208-A209), and in British Columbia, Northern
Territories, Alberta, Saskatchewan, and parts of Manitoba in Canada
(Committee on the Status of Endangered Wildlife in Canada 2009, p.
iii).
Habitat
The northern leopard frog is an amphibian (a cold-blooded
vertebrate that spends some time on land, but must breed and develop
into an adult in water) and as such is ectothermic (incapable of
generating their own body heat) (Wells 2007, p. 2). They have highly
permeable skin, which allows for rapid passage of water and gases so
that they can use their external environment to regulate body
temperature and moisture loss (Wells 2007, pp. 2-3). As part of its
complex life history, the northern leopard frog requires a mosaic of
habitats, which includes aquatic overwintering and breeding habitats,
and upland post-breeding habitats, as well as habitat linkages, to meet
the requirements of all of its life stages (Pope et al. 2000, p. 2505;
Smith 2003, pp. 6-15; Rorabaugh 2005, pp. 571-575). Although aquatic
breeding habitat is required for long-term population survival, upland
foraging, dispersal, and overwintering habitats are critical if
individual leopard frogs are to survive to reproductive maturity. For
example, researchers noted an area near Chicago that had low northern
leopard frog abundance, but extensive potential aquatic breeding
habitat. It was not until habitat surrounding the ponds was restored
from scrub forest to grasslands that leopard frog numbers increased
dramatically (K.S. Mierzwa, pers. comm. in Pope et al. 2000, p. 2506).
These complex habitat requirements make northern leopard frogs
particularly vulnerable to the impacts of habitat loss and
fragmentation. Reduction or removal of these habitats or loss of
connectivity between habitat components could reduce the capacity of
the landscape to support the species (Pope et al. 2000, p. 2505; Green
2005, p. 31).
Northern leopard frogs breed in a variety of aquatic habitats that
include slow-moving or still water along streams and rivers, wetlands,
permanent or temporary pools, beaver ponds, and human-constructed
habitats such as earthen stock tanks and borrow pits (Rorabaugh 2005,
p. 572). Successful breeding areas typically do not contain predaceous
fish or other predators (Merrell 1968, p. 275; Hine et al. 1981, p. 12;
Orr et al. 1998, p. 92; Smith 2003, pp. 19-21). Emergent vegetation,
such as sedges and rushes, are important features of breeding and
tadpole habitats (Gilbert et al. 1994, p. 468; Smith 2003, pp. 8-9),
and tadpoles are most often found in backwaters and still pools
(Rorabaugh 2005, p. 572).
Sub-adult northern leopard frogs typically move from breeding areas
to
[[Page 61898]]
feeding sites along the borders of larger, more permanent bodies of
water, as smaller frogs are closely tied to water (Merrell 1970, p.
49). Recently metamorphosed frogs will move up and down drainages and
across land in an effort to disperse from breeding areas (Seburn et al.
1997, p. 69) and may disperse more than 0.5 mile (mi) (800 meters (m))
from their place of metamorphosis (Dole 1971, p. 223). Dole (1971, p.
226) found that dispersal in Michigan occurred on warm, rainy nights
and that frogs dispersed overland; however, warm rains are not common
in all parts of the species' range and other dispersal routes may be
important as well. Streams are an important corridor for dispersing
juvenile frogs (Seburn et al. 1997, pp. 68-69), and vegetated drainage
ditches may also facilitate connectivity between seasonal habitats
(Pope et al. 2000, p. 2505). In some areas of the western United
States, subadults may remain in the breeding habitat within which they
metamorphosed (Smith 2003, p. 10).
In addition to the breeding habitats, adult northern leopard frogs
require stream, pond, lake, or river habitats for overwintering and
upland habitats adjacent to these areas for summer feeding. In summer,
adults and juveniles commonly feed in open or semi-open wet meadows and
fields with shorter vegetation, usually near the margins of water
bodies, and seek escape cover underwater. Post-breeding summer habitats
do not include barren ground, open sandy areas, heavily wooded areas,
cultivated fields, heavily grazed pastures, or mowed lawns (Rorabaugh
2005, p. 573). Buffer zones around wetland breeding sites should be
maintained for movement to surrounding upland foraging habitat.
Rittenhouse and Semlitsch (2007, p. 154) collected data from 13
published radio telemetry and tagging studies looking at frog and
salamander use of terrestrial habitat surrounding wetlands. They found
that, on average, a buffer width of 1,877 ft (572 m) around the
breeding site is needed to encompass the non-breeding habitat used by
90 percent of the frogs in a given population (Rittenhouse and
Semlitsch 2007, pp. 155-157).
During winter, northern leopard frogs are thought to hibernate
underwater in ponds, in lakes, or on the bottom of deeper streams or
waters that do not freeze to the bottom and that are well-oxygenated
(Nussbaum 1983, p. 181; Stewart et al. 2004, p. 72). Northern leopard
frogs are intolerant of freezing and of waters that have severely
reduced or complete loss of dissolved oxygen. If these conditions occur
during hibernation, death of northern leopard frogs is likely
(Rorabaugh 2005, p. 574).
Based upon their research in Wisconsin, Hine et al. (1981)
described the ideal ``breeding pond'' as having the following features:
(1) The pond or wetland site should be located within approximately
1.0 mile (mi) (1.6 kilometers (km)) of suitable overwintering habitat
(larger bodies of water) so that adults can find the breeding habitat
when they emerge in the spring and juvenile frogs are able to find
overwintering sites in the fall.
(2) In the spring, the water depth should be approximately 5 ft
(1.5 m) or more so that there is balance of open water and vegetation
cover.
(3) Emergent vegetation (such as sedge, bulrush, and cattail)
should occur along at least two-thirds of the pond or wetland to
provide escape cover and places to attach egg masses.
(4) The slope should be gradual to promote habitat for emergent
vegetation.
(5) Natural terrestrial habitats should be maintained peripheral to
wetlands summer habitat for adults post-breeding, for juvenile growth,
and for dispersal or movement corridors.
(6) Water should be relatively permanent throughout the year, but
should dry every decade or so in order to eliminate any predaceous fish
that become established.
Water quality and temperature are important determinants of
northern leopard frog habitat. Because northern leopard frogs have
permeable skin, which may transfer external contaminants to its
internal organs, good (i.e., non-polluted) water quality is important
at breeding locations. Chemical contamination of habitats can result in
malformations, population declines, decreased growth rates, reduced
activity, and other impacts to northern leopard frogs (Diana and
Beasley 1998, pp. 267-276). Temperature plays an important role in both
the springtime migratory and breeding behaviors of northern leopard
frogs (Merrell 1970, pp. 50-51; Merrell 1977, pp. 5-6, 9). When ambient
air temperature is greater than or equal to 50 degrees Fahrenheit
([deg]F) (10 degrees Celsius ([deg]C)), northern leopard frogs move
from their overwintering sites to their breeding sites (Merrell 1970,
p. 50). The calling sites and areas where egg masses are deposited are
not random and appear to be chosen based upon temperature as these
activities tend to be located in the warmest portions of breeding ponds
(Merrell 1977, p. 6).
Biology
As soon as males leave overwintering sites, they travel to breeding
ponds and call in shallow water (Smith 2003, p. 13). Breeding typically
occurs during a short period in the spring beginning in early April
(Pace 1974, p. 92; Corn and Livo 1989, p. 4); at higher elevations and
more northern latitudes, the onset of breeding is late April to early
May (Corn and Livo 1989, p. 5; Gilbert et al. 1994, p. 467). Most
northern leopard frogs are sexually mature at age 2, although the age
of sexual maturity may vary from age 1 to age 3 in any given population
depending upon environmental conditions (Leclair and Castanet 1987, p.
368; Gilbert et al. 1994, pp. 468-469). Male frogs attract females by
calling from specific locations within a breeding pond when
temperatures are close to 68 [deg]F (20 [deg]C) or more, with several
males typically calling together to form a chorus (Merrell 1977, p. 7).
Eggs are typically laid within breeding habitats, 2 to 3 days following
the onset of chorusing (Corn and Livo 1989, p. 5). Eggs are laid in
non-acidic, shallow (4 to 26 in (10 to 65 cm)), still water that is
exposed to sunlight, and are usually attached to emergent vegetation
just below the water surface (Merrell 1977, p. 6; Gilbert et al. 1994,
pp. 467-468; Pope et al. 2000, p. 2505). Egg masses may include several
hundred to several thousand eggs (Corn and Livo 1989, pp. 6-7) and are
deposited in a tight, oval mass (Rorabaugh 2005, p. 572). Time to
hatching is correlated with temperature and ranges from 2 days at 81
[deg]F (27 [deg]C) to 17 days at approximately 53 [deg]F (12 [deg]C)
(Nussbaum et al. 1983, p. 182).
Tadpoles are the ephemeral, feeding, non-reproductive, completely
aquatic larvae in the life cycle of the frog (McDiarmid and Altig 1999,
p. 2). The length of time required for metamorphosis (the development
of the aquatic tadpole to a frog) is variable, and depending upon
temperature, may take 3 to 6 months from time of egg-laying (Merrell
1977, p. 10; Hinshaw 1999, p. 105). Northern leopard frog tadpoles are
predominantly generalist herbivores (plant eaters), typically eating
attached and free-floating algae (Hoff et al.1999, p. 215); however
they may feed on dead animals (Hendricks 1973, p. 100). Adult and
subadult frogs are generalist insectivores (insect eaters) that feed on
a variety of terrestrial invertebrates such as insect adults, larvae,
spiders, and leeches (Merrell 1977, p. 15; Collier et al. 1998, p. 41;
Smith 2003, p. 12; Rorabaugh 2005, p. 575). In addition, adult northern
leopard frogs have also been known to prey upon small
[[Page 61899]]
northern leopard frogs, birds, and snakes (Merrell 1977, p. 15).
Status
Northern leopard frogs, like many amphibian populations, are
dynamic, and their individual numbers may naturally fluctuate in size
within populations. However, across the range of the northern leopard
frog, information suggests that there is an ongoing loss of populations
throughout the species' range. The loss of populations across the
landscape is what results in species' declines (Green 2005, p. 29).
Population declines of northern leopard frogs are well-documented in
the western United States and western Canada, but are also documented
rangewide (through the Midwestern and Eastern United States), as
described below.
The most recent complete summary of distributional and abundance
patterns of the northern leopard frog is from Rorabaugh (2005, pp. 570-
571), which documents a substantial contraction of the species' range,
especially in the western two-thirds of the United States, where
widespread extirpations have occurred. Other authors have also compiled
summary data indicating population declines (e.g., Smith and Keinath
2007, p. 14). Since the 1960s, the northern leopard frog has
experienced significant declines and losses throughout its range (Gibbs
et al. 1971, p. 1028), particularly in the western United States and
western Canada, and tends to become less abundant the farther west one
proceeds (Corn and Fogelman 1984, p. 150; Hayes and Jennings 1986, p.
491; Clarkson and Rorabaugh 1989, p. 534; Corn et al. 1989, pp. 26-29;
Koch and Peterson 1995, pp. 84-87; Corn et al. 1997, pp. 37-38; Weller
and Green 1997, p. 323; Casper 1998, p. 199; Hammerson 1999, pp. 146-
147; Leonard et al. 1999, p. 51; Dixon 2000, p. 77; Smith 2003, pp. 4-
6; Jennings and Fuller 2004, pp. 125-127; Werner et al. 2004, pp. 97-
98; Committee on the Status of Endangered Wildlife in Canada 2009, p.
v; Germaine and Hays 2009, p. 537; Johnson et al. 2011, p. 557).
Based upon this and other information, the northern leopard frog
appears to be declining, is considered rare, or is locally extirpated
from many historical locations in Arizona, California, Colorado, Idaho,
Iowa, Minnesota, Missouri, Montana, Nebraska, Nevada, New Mexico,
Oregon, Texas, Utah, Washington, Wisconsin, and Wyoming (Hayes and
Jennings 1986, p. 491; Stebbins and Cohen 1995, p. 220; Johnson and
Batie 1996; Bowers et al. 1998, p. 372; Casper 1998, p. 199; Lannoo
1998, p. xvi; Mossman et al. 1998, p. 198; Smith 2003, pp. 4-6; Smith
and Keinath 2004, pp. 57-60; McCleod 2005, pp. 292-294; Rorabaugh 2005,
p. 571; Johnson et al. 2011, p. 561). The species is nearly extirpated
from almost 100 percent of its historical range in Texas, California,
Oregon, and Washington (Stebbins and Cohen 1995, p. 220; McAllister et
al. 1999, p. 15; Stebbins 2003, p. 235; Germaine and Hays 2009, p.
537).
Table 1 lists current NatureServe ranks for States and provinces in
which the northern leopard frog is known to occur. NatureServe
conservation status assessment procedures have different criteria,
evidence requirements, purposes, and taxonomic coverage than the
Federal Lists of Endangered and Threatened Wildlife and Plants, and
therefore, these rankings may not coincide with legal listing processes
(NatureServe 2008, p. 1). However, for a species as widespread as the
northern leopard frog, the NatureServe rankings aid in summarizing the
relative risks facing the northern leopard frog throughout its range
and are provided here for this reason.
NatureServe lists Maryland and New Jersey as States where the
northern leopard frog occurs. However, the Maryland Department of
Natural Resources lists the northern leopard frog as an introduced
species that occurs in one county (Maryland Department of Natural
Resources 2011, p. 2), and the frog does not occur in New Jersey
(Gessner and Stiles 2001, pp. 1-9; New Jersey Division of Fish and
Wildlife 2006, pp. 1-2).
Table 1--NatureServe and State, Province, and Territory Ranks for
Northern Leopard Frogs in States and Provinces It Is Known To Occur
[NatureServe 2011, p. 1]
------------------------------------------------------------------------
State, province, territory Natural heritage State, province,
or sovereign nation program rank * territory rank
------------------------------------------------------------------------
Arizona..................... S2 (Imperiled)...... Species of Greatest
Conservation Need.
California.................. S2 (Imperiled)...... Species of Greatest
Conservation Need.
Colorado.................... S3 (Vulnerable)..... Species of Greatest
Conservation Need,
Species of Special
Concern.
Connecticut................. S2 (Imperiled)...... Special Concern
Species.
Idaho....................... S3 (Vulnerable)..... Species of Greatest
Conservation Need.
Illinois.................... S5 (Secure)......... Non-game Indicator
Species.
Indiana..................... S2 (Imperiled)...... Species of Greatest
Conservation Need.
Iowa........................ S5 (Secure)......... No ranking or
status.
Kentucky.................... S3 (Vulnerable)..... Species of Greatest
Conservation Need.
Maine....................... S3 (Vulnerable)..... Species of Greatest
Conservation Need
(Priority 3).
Maryland.................... S4 (Apparently No ranking or status
Secure), introduced (considered an
spp. introduced
species).
Massachusetts............... S3/S4 (Vulnerable/ Species of Special
Apparently Secure). Concern, Species of
Greatest
Conservation Need.
Michigan.................... S5 (Secure)......... Species of Greatest
Conservation Need.
Minnesota................... S4 (Apparently No ranking or
Secure). status.
Missouri.................... S2 (Imperiled)...... Species of
Conservation
Concern.
Montana..................... S1/S3 (Critically Species of Concern,
Imperiled/ Species of Greatest
Vulnerable). Conservation Need.
Navajo Nation (NE Arizona, S2 (Imperiled)...... Endangered.
NW New Mexico, SE Utah).
Nebraska.................... S5 (Secure)......... At-Risk Species
(Tier II).
Nevada...................... S2/S3 (Imperiled/ Species of
Vulnerable). Conservation
Priority.
New Hampshire............... S3 (Vulnerable)..... Species of Concern.
New Jersey.................. SNR (Unranked), Species not present.
species not present.
[[Page 61900]]
New Mexico.................. S1 (Critically Species of Greatest
Imperiled). Conservation Need.
New York.................... S5 (Secure)......... No ranking or
status.
North Dakota................ SNR (Unranked)...... No ranking or
status.
Ohio........................ SNR (Unranked)...... No ranking or
status.
Oregon...................... S1/S2 (Critically Sensitive Critical,
Imperiled/ List 2 Species
Imperiled). (threatened with
extinction or
presumed extinct).
Pennsylvania................ S2/S3 (Imperiled/ Priority
Vulnerable). Conservation
Species (Tier 5).
Rhode Island................ S2 (Imperiled)...... Species of Greatest
Conservation Need.
South Dakota................ S5 (Secure)......... No ranking or
status.
Texas....................... S1 (Critically No ranking or status
Imperiled). (likely
extirpated).
Utah........................ S3/S4 (Vulnerable/ Species of Concern
Apparently Secure). (Tier III).
Vermont..................... S4 (Vulnerable)..... No ranking or
status.
Washington.................. S1 (Critically Endangered.
Imperiled).
West Virginia............... S2 (Imperiled)...... Species in Greatest
Need of
Conservation.
Wisconsin................... S4 (Vulnerable)..... No ranking or
status.
Wyoming..................... S3 (Vulnerable)..... Species of Greatest
Conservation Need.
Alberta..................... S2/S3 (Imperiled/ Threatened.
Vulnerable).
British Columbia............ S1 (Critically Endangered.
Imperiled).
Labrador and Newfoundland... S3/S4 (Vulnerable/ No ranking or
Apparently Secure). status.
Manitoba.................... S4 (Vulnerable)..... No ranking or
status.
New Brunswick............... S5 (Secure)......... No ranking or
status.
Northwest Territories....... SNR (Unranked)...... No ranking or
status.
Nova Scotia................. S5 (Secure)......... No ranking or
status.
Ontario..................... S5 (Secure)......... Not at risk.
Prince Edward Island........ S4/S5 (Apparently No ranking or
Secure/Secure). status.
Quebec...................... S5 (Secure)......... No ranking or
status.
Saskatchewan................ S3 (Vulnerable)..... Interim Species at
Risk.
------------------------------------------------------------------------
* S1 = Critically Imperiled: At very high risk of extinction due to
extreme rarity (often 5 or fewer populations), very steep declines, or
other factors.
S2 = Imperiled: At high risk of extinction due to restricted range, few
populations (often 20 or fewer), steep declines, or other factors.
S3 = Vulnerable: At moderate risk of extinction due to a restricted
range, relatively few populations (often 80 or fewer), recent and
widespread declines, or other factors. Such species are often rare or
found locally in a restricted range.
S4 = Apparently Secure: Uncommon but not rare; some cause for long-term
concern due to declines or other factors. Such species are likely to
be quite rare in parts of their range, especially at the periphery.
S5 = Secure: Common; widespread and abundant. Such species are
potentially rare in parts of their range, especially at the periphery.
SNR = Unranked. State or Province conservation status not yet assessed.
The International Union for the Conservation of Nature's ``Red List
Categories and Criteria'' were developed for classifying species at
high risk of global extinction (IUCN 2003, p. 1), and as such have
different criteria, evidence requirements, purposes, and taxonomic
coverage than the Federal Lists of Endangered and Threatened Wildlife
and Plants. However, just as with the NatureServe data, because we are
reviewing the entire range of the northern leopard frog, the
International Union for the Conservation of Nature assessment is useful
in summarizing the current status of the northern leopard frog
throughout its range.
The International Union for the Conservation of Nature currently
lists the northern leopard frog as a species of `least concern' in view
of its wide distribution, tolerance to degree of habitat modification,
and presumed large population (Hammerson et al. 2004, p. 2). The
International Union for the Conservation of Nature states that the
population trend is decreasing (Hammerson et al. 2004, p. 3), but the
authors believe that the northern leopard frog is not declining fast
enough to qualify for listing in a more threatened category (Hammerson
et al. 2004, p. 2). The International Union for the Conservation of
Nature reviewed Hammerson et al. (2004, pp. 1-6) in 2011, and no
updates were made to the 2004 review. Since 2004, Rorabaugh (2005, pp.
570-577) completed a status review for the northern leopard frog in the
United States (Rorabaugh 2005, pp. 570-577), and the Committee on the
Status of Endangered Wildlife in Canada published the Assessment and
Update Status Report for the Northern Leopard Frog in Canada (Committee
on the Status of Endangered Wildlife in Canada 2009, pp. 1-76). The
Rorabaugh (2005, pp. 570-577) status review found that for a variety of
reasons the northern leopard frog is declining throughout its range,
but particularly in the western United States. The Committee on the
Status of Endangered Wildlife in Canada (2009, pp. iii) assessment
notes that there are continued declines for the northern leopard frog
throughout the western provinces and evidence of declines in eastern
Canada. The current International Union for the Conservation of Nature
review does not cite either of these documents or provide any current
threats assessment. The International Union for the Conservation of
Nature analysis for the northern leopard frog also includes leopard
frogs in Panama, which likely belong to the Lithobates complex, but do
not belong to the same species as the northern leopard frog. Therefore,
we do not consider the International Union for the Conservation of
Nature review for the northern leopard frog a current assessment of the
species' status in North America.
Western States
Until the late 1970s, northern leopard frogs were widespread and
abundant in much of northern Arizona (Apache, Coconino, Greenlee,
Mohave, Navajo, and Yavapai Counties) in springs, streams, rivers,
stock tanks, and lakes throughout northern Arizona (Arizona Game and
Fish Department 2009, p. 1).
[[Page 61901]]
Currently, there is one northern leopard frog population located near
Seligman, Arizona; a metapopulation (several breeding locations in
close proximity to one another) located south of Flagstaff, Arizona;
and three refugial sites developed by the State and Service (and other
partners) to assist in stocking northern leopard frogs to other
locations in Arizona, north of the Colorado River. All of these
locations are located in Coconino County. Outside of these locations,
fairly rigorous visual encounter surveys conducted within the species'
historical range, including Grand Canyon National Park and the Kaibab
National Forest, have not located northern leopard frogs (Kaibab
National Forest 2007, p. 1; Kaibab National Forest 2008, p. 1; Drost et
al. 2008, p. 7). The species is listed as a Species of Greatest
Conservation Need in the Arizona State Wildlife Action Plan (Arizona
Game and Fish Department 2006, Appendix M, p. 153) and has a
NatureServe rank of S2 (Imperiled) (NatureServe 2011, p. 1). In
Arizona, there is no open season for northern leopard frog, and
collecting is illegal except as authorized by State permit, effective
January 1, 1993 (Commission Order 41). The northern leopard frog has
also significantly declined on the Navajo Nation (which is situated in
southeastern Utah, northeastern Arizona, and northwestern New Mexico)
in the last century. Most remote desert populations of northern leopard
frogs were lost between the 1920s and 1970s, and mountain populations
were lost in the late 1980s. The Navajo Nation has listed the northern
leopard frog as a ``Group 2--Endangered Species'' on the Navajo
Endangered Species List, which means its prospects of survival or
recruitment on the Navajo Nation are in jeopardy (Navajo Nation
Department of Fish and Wildlife 2009, p. 3).
The northern leopard frog is a State of California species of
special concern and is listed as a Species of Special Concern (native
populations only) (California Department of Fish and Game, Natural
Diversity Database, 2009) and as a Species of Greatest Conservation
Need in California Department of Fish and Game's State Wildlife Action
Plan (California Department of Fish and Game 2007); however, the
northern leopard frog is not listed under the California Endangered
Species Act. The northern leopard frog may be taken under the authority
of a sport fishing license, subject to restrictions (California Code of
Regulations, Title 14, Section 5.05). The frog is ranked S2 (Imperiled)
by NatureServe (NatureServe 2011, p. 1). Northern leopard frogs are
likely native to the region east of the Sierra Nevada-Cascade crest in
the following areas of California: upper Pit River basin (Shasta,
Lassen, and Modoc counties), Surprise Valley (Modoc County), lower
Klamath Lake basin (Siskiyou County), Lake Tahoe region (El Dorado
County), Carson River drainage (Alpine County) and Owens River Valley
(Mono and Inyo counties) (Jennings and Fuller 2004, p. 122). The
northern leopard frog was introduced to at least 15 other sites in
California, but most of these introductions have not resulted in
naturalized populations that continue to exist today (Jennings and
Hayes 1994, p. 80; Jennings and Fuller 2004, p. 119). There is a small,
introduced population in Merced County, near the Merced National
Wildlife Refuge (NWR) that persisted as recently as 2007 (Jennings and
Fuller 2004, pp. 119, 127; Woolington 2009, pers. comm.). Since the
1970s, northern leopard frogs have disappeared from most (approximately
95 percent) of their historic range in California, (Jennings and Fuller
2004, p. 119; Rorabaugh 2005, p. 571) and may be completely extirpated
from these areas of the State as we are not aware of any recent
confirmed sightings. Jennings and Hayes (1994, p. 82) knew of only two
extant, native northern leopard frog populations as of the 1990s: one
adult was observed at Tule Lake National NWR (Siskiyou County) in 1990,
and 8 to 10 juveniles were found near Pine Creek in Round Valley near
Bishop (Inyo County) in 1994. Northern leopard frogs are no longer
found on Tule Lake NWR (Adams 2011, pers. comm.), and no northern
leopard frogs have been observed during amphibian surveys conducted on
the Klamath Falls NWR Complex, including Tule Lake NWR (Austin 2009,
pers. comm.). Recent surveys conducted by the California Department of
Fish and Game did not locate any northern leopard frogs in the Owens
River Area (Becker 2011, pers. comm.). In addition, surveys found that
sites previously considered to be northern leopard frog habitat now
contain nonnative aquatic species, and the habitat has been extensively
modified such that there are likely few areas of suitable habitat left
in the Owens Valley (Becker 2011, pers. comm.). Northern leopard frogs
have not been found in the Lake Tahoe basin for over 20 years, and the
species is presumed to be extirpated from the area (Jennings and Fuller
2004, p. 125). Jennings and Fuller (2004, p. 126) also report that a
formerly isolated native northern leopard frog population on Hat Creek,
Shasta County, is now apparently extirpated as well. Modoc NWR in
northeastern California reported no known occurrences of northern
leopard frogs on the refuge in recent times, and no northern leopard
frogs were reported during numerous hours of amphibian survey time in
2004, 2005, and 2010 (Bachman 2011, pers. comm.).
The northern leopard frog was historically quite common throughout
Colorado, but over the last 30 to 40 years, populations have declined
and even been locally extirpated from portions of eastern and north-
central Colorado, including Rocky Mountain and Mesa Verde National
Parks (Corn and Fogleman 1984, p. 148; Corn et al. 1989, p. 15;
Stebbins and Cohen 1995, p. 220; Corn et al. 1997, pp. 37-38; Hammerson
1999, pp. 146-147; Mesa Verde National Park 2009, p. 1; Johnson et al.
2011, p. 561). The Colorado Division of Wildlife has designated the
northern leopard frog a Species of Greatest Conservation Need as well
as a Species of Special Concern due to low population status and a
declining population trend (Colorado Division of Wildlife 2006, pp. 2,
28, 305). These are not statutory categories; however, the northern
leopard frog is classified as ``nongame'' wildlife and their
harassment, taking, or possession is prohibited without a permit
(Colorado Division of Wildlife 2009, p. 3). NatureServe ranks the
northern leopard frog as S3 (Vulnerable) in Colorado (NatureServe 2011,
p. 1). Intensive surveys conducted from 2007 through 2009 in the Front
Range of Colorado indicate that northern leopard frogs there have
become rare and documented losses are widespread (Johnson and McKenzie
2009, p. 9; Keeley 2009, pp. 5-6; Johnson et al. 2011, p. 562).
Historically, northern leopard frogs were found at high densities in
this region (Johnson et al. 2011, p. 562). Along the Western Slope (the
area west of the continental divide in Colorado), data suggest that
northern leopard frog populations remain viable, especially in the
northern region (Johnson and McKenzie 2009, p. 10). This supports
information from Arapaho and Browns Park NWRs, both located in
northwestern Colorado, that continue to support northern leopard frogs
(Johnson 2009, pers. comm.; Smart 2009, pers. comm.). Northern leopard
frogs were the most common amphibian in southwest Colorado until the
late 1960s, but now they are rare (San Miguel 2009, pers. comm.).
Despite conducting amphibian surveys for 15 years with an emphasis on
locating northern leopard frogs, none have been detected within Mesa
Verde National Park, Colorado. Historically,
[[Page 61902]]
this species was found abundantly along the Mancos River in the park
and adjacent lands (San Miguel 2009, pers. comm.). However, the overall
status of the northern leopard frog in western Colorado is not
currently known (Johnson et al. 2011, p. 563).
The Idaho Department of Fish and Game designated the northern
leopard frog a Type 2 Species of Greatest Conservation Need (Idaho
Department of Fish and Game 2005, Appendix B p. 6). A Type 2 species of
greatest conservation need is defined as a rangewide or globally
imperiled species that is experiencing significant declines throughout
its range with a high likelihood of being listed in the foreseeable
future due to its rarity (Idaho Department of Fish and Game 2005,
Appendix B, p. 4). Reduced distribution and a declining population
trend are noted in the Idaho Comprehensive Wildlife Conservation
Strategy as reasons for the designation (Idaho Department of Fish and
Game 2005, Species Account, p. 1). The northern leopard frog is also a
protected nongame species, which means take or possession of the
species is prohibited without a permit (Idaho Administrative Code
13.01.06-300.02). NatureServe ranks the northern leopard frog in Idaho
as S3 (Vulnerable) (NatureServe 2011, p. 1). Both the Targhee National
Forest and Kootenai NWR have records of northern leopard frogs from the
1970s (Service 1972, p. 11; Stebbins and Cohen 1995, p. 220). However,
surveys in 1992 at 98 sites on the Targhee National Forest did not
locate northern leopard frogs (Stebbins and Cohen 1995, p. 220), and
Kootenai NWR has no records of frogs for the last 30 years (Rose 2009,
pers. comm.). Deer Flat NWR amphibian surveys have only detected
American bullfrogs (Lithobates catesbeiana). Northern leopard frogs are
known to be present on Bear Lake, Grays Lake, and Minidoka NWRs, and
presumed to be present on Camas NWR and Oxford Slough Wetland
Protection Area (WPA) (Fisher and Mitchell 2009, p. 1).
Localized declines of northern leopard frogs are documented in Iowa
(Lannoo et al. 1994, pp. 317-318; Hemesath 1998, p. 216). Lannoo et al.
1994 (p. 311) states, ``From descriptions of the turn-of-the-century
commercial ``frogging'' industry in Dickinson County (Iowa), we
estimate that the number of leopard frogs has declined by at least two,
and probably three orders of magnitude.'' However, the northern leopard
frog is ranked as Secure (S5) in Iowa by NatureServe (2011, p. 1) and
is not considered a Species of Greatest Conservation Need (Iowa
Department of Natural Resources 2006, p. 42). Currently, there is a
continuous open season for northern leopard frogs in inland and
boundary waters in Iowa, and up to 48 frogs can be collected per day
(Iowa Department of Natural Resources 2011, p. 1). In 1991, the Iowa
Department of Natural Resources initiated an annual anuran (frog and
toad) survey. The survey is conducted by volunteers, and until 2007,
volunteers were not required to distinguish between species of leopard
frogs on the report forms (Iowa Department of Natural Resources 2009,
p. 1). Survey data from 2007 and 2008 (when the species were separated)
and older data from counties where it was thought only the northern
leopard frog occurred were reviewed by the State. The analyses of this
information suggest a possible downward trend in northern leopard frog
presence, but the trend was not statistically significant (Iowa
Department of Natural Resources 2009, p. 1).
Northern leopard frog populations began declining in Minnesota in
the late 1960s or early 1970s (Rittschof 1975, p. 103; Minnesota
Department of Natural Resources 2011a, pp. 1-2). The declines of
northern leopard frog populations from the past are thought to have
been substantial, but information is not detailed enough to know if the
population is now stable or if it is still declining in Minnesota
(Moriarty 1998, p. 168). However, because the species is still
considered to be fairly common, it is not considered a Species of
Greatest Conservation Need in Minnesota's Comprehensive Wildlife
Strategy (Minnesota Department of Natural Resources 2006, Appendix B p.
9). The Minnesota Department of Natural Resources' northern leopard
frog fact page does indicate that the northern leopard frog is still
declining (Minnesota Department of Natural Resources 2011a, p. 2). The
species is ranked S4 (Apparently Secure) by NatureServe (NatureServe
2011, p. 1). In Minnesota, from May 16 to March 31, licensed anglers
and children under age 16 may take, use, buy, and sell an unlimited
number of northern leopard frogs up to 6 inches long for bait
(Minnesota Department of Natural Resources 2011b, p. 70). A Minnesota
Department of Natural Resources commercial license is required to take
northern leopard frogs for purposes other than bait.
Missouri is located on the periphery of the range for northern
leopard frogs and the frog is currently only known to occur in two
counties (Atchison and Mercer) that border Iowa (Missouri Department of
Conservation 2009, p. 1). The northern leopard frog is listed as a
Species of Conservation Concern by the Missouri Department of
Conservation and NatureServe ranks it as Imperiled (S2) (Missouri
Department of Conservation 2009, p. 1; NatureServe 2011, p. 1). This
ranking is based upon the low number of known occurrences in Missouri
and not based upon declining population trends (Missouri Department of
Conservation 2009, p. 1). The Missouri Department of Conservation noted
that it is likely that more populations are present in northern
Missouri, but further surveys need to be completed to affirm this
assumption (Missouri Department of Conservation 2009, p. 1). In
Missouri, northern leopard frogs have regulatory protection from
commercial take and non-resident collection. Missouri residents are
allowed to possess up to five northern leopard frogs for education use
(Wildlife Code Missouri 3CSR10-9.110); however, these five individuals
cannot be sold, traded, shipped over State lines, or taken from public
lands (Missouri Department of Conservation 2009, p. 2). Northern
leopard frogs also cannot be used as live bait in Missouri (Wildlife
Code Missouri 3CSR10-6.605).
Montana Fish, Wildlife, and Parks classified the northern leopard
frog as a Species of Concern in Montana and it is considered a Species
of Greatest Conservation Need in their Wildlife Conservation Strategy
(Montana Fish, Wildlife, and Parks 2009, p. 1). Northern leopard frogs
are protected from commercial collection in Montana (Montana Code
Annotated 2009 87-5-116). Historically, northern leopard frogs occurred
across the eastern plains of Montana and in the mountain valleys on
both sides of the Continental Divide (Montana Fish, Wildlife, and Parks
2009, p. 1). However, since the 1990s, most previously known northern
leopard frog populations on the west side of the Continental Divide in
Montana are considered extirpated, and there has been a clear range
contraction of northern leopard frogs (Werner 2003, p. 26; Montana
Fish, Wildlife, and Parks 2009, p. 1). Currently, only two populations
exist in western Montana. Surveys in the mid-1990s of historically
occupied sites in central Montana, east of the Continental Divide,
found only 19 percent of the sites to be occupied by northern leopard
frogs (Montana Fish, Wildlife, and Parks 2009, p. 1). NatureServe
provides a split rank for the State that reflects the difference in
status between western (S1 Critically Imperiled) and eastern (S3
Vulnerable) Montana (NatureServe 2011, p. 1). Habitat restoration and
survey efforts are being planned Statewide to provide
[[Page 61903]]
a current assessment of northern leopard frog distribution (Montana
Fish, Wildlife, and Parks, 2009, p. 2).
The northern leopard frog occurs commonly in the State of Nebraska
(McLeod 2005, p. 292) and has a NatureServe rank of S5 (Secure)
(NatureServe 2011, p. 1). However, surveys conducted in 1997 and 1998
indicated a significant decline in northern leopard frog occurrences at
the State level (McLeod 2005, p. 292). It is difficult to ascertain if
this information represents a real decline or is representative of
normal stochastic events, but data indicated significant differences
from location data collected in the 1970s (McLeod 2005, p. 292). The
Nebraska Game and Parks Commission identified the northern leopard frog
as a Tier II At-Risk Species during development of the Nebraska Natural
Legacy Project (2005, p. 319). Tier II species are typically those that
are not at-risk from a global or national perspective, but are rare or
imperiled within Nebraska. As of 2011, northern leopard frogs can no
longer be commercially harvested or sold for bait in Nebraska; however,
anglers can still collect them as bait for personal use (Nebraska Game
and Parks Commission 2011, p. 5).
In Nevada, northern leopard frogs are currently ranked S2/S3
(Imperiled/Vulnerable) by NatureServe (NatureServe 2011, p. 1) and are
on the Nevada Natural Heritage Program's Animal and Plant Watch List,
which means they could be declining in Nevada or across much of their
range, or may be less common than currently thought and could become
at-risk in the future. The northern leopard frog is identified as a
Species of Conservation Priority in the Nevada Wildlife Action Plan
(Wildlife Action Plan Team 2006, p. 61). In addition, the northern
leopard frog is a protected amphibian by Nevada statute (NAC 503.075)
and cannot be collected for commercial, recreational, or educational
purposes without a permit (Nevada Department of Wildlife 2009, p. 5).
The Nevada Department of Wildlife notes that there is little historical
or current information available to accurately assess the distribution
and status of the northern leopard frog in Nevada (Nevada Department of
Wildlife 2009, p. 1). However, recent surveys suggest that northern
leopard frogs may no longer be abundant in Nevada and that there have
been numerous local extirpations, for example, along the Truckee and
Carson rivers in western Nevada and in springs of southern and eastern
Nevada (Panik and Barrett 1993, p. 203; Hitchcock 2001, pp. 9, 109-
110). While historical records and anecdotal evidence indicated that
northern leopard frogs were once widely distributed in the State, the
current species distribution is much smaller than the historical
distribution (Hitchcock 2001, pp. 9, 38, 48). In addition, suitable
northern leopard frog habitat is patchily distributed in the State due
to the aridity and isolated nature of many wetland systems, which
results in a discontinuous and limited distribution (Nevada Department
of Wildlife 2009, p. 1). Recent Nevada Department of Wildlife records
document northern leopard frog populations in Ruby Valley (including
Ruby Lakes NWR) and Lower Mary's River in Elko and White Pine Counties;
Spring Valley and Lake Valley in White Pine County; Lake Valley and
Pahranagat Valley (including Pahranagat NWR) in Lincoln County; Carson
River near Carson City; the lower Truckee River and Truckee meadows in
Washoe County; and a small number of additional sites in western and
northeastern Nevada (Hitchcock 2001, pp. 96-102; Service 2009, pp. 1-2;
Nevada Department of Wildlife 2009, p. 2). Efforts to restore northern
leopard frog habitat and re-establish the species have occurred along
the lower Truckee River in western Nevada and on Pahranagat NWR (Horton
2010, pers. comm.; Rogers 2010, p. 7).
Historically, the northern leopard frog was documented from a large
area in the northern and western part of New Mexico and along the
entire length of the Rio Grande River valley, except southern Elephant
Butte and northern Caballo Reservoirs (New Mexico Department of Game
and Fish 2009, p. 1). Declines in northern leopard frogs have been
reported from the Lower Rio Grande (below Caballo Reservoir), in the
Jemez Mountains, and in the Chuska Mountains (Christman 2009, p. 5; New
Mexico Department of Game and Fish 2009, p. 2). The species is believed
to be extirpated from the Rio Grande Valley, south of Albuquerque (New
Mexico Department of Game and Fish 2009, p. 3). Recent survey efforts
indicate that northern leopard frogs are persisting in northern New
Mexico, but most occupied sites contained small numbers of frogs with
very few robust populations (Christman 2009, p. 13). The northern
leopard frog is not listed as endangered or threatened in New Mexico
under the Wildlife Conservation Act, but was designated a Species of
Greatest Conservation Need by the New Mexico Department of Game and
Fish, and NatureServe ranks it as S1 (Critically Imperiled) in New
Mexico (New Mexico Department of Game and Fish 2006, p. 540;
NatureServe 2011, p. 1). The northern leopard frog is protected from
commercial take (Section 17-1-14 NMSA); however, take by New Mexico
State residents for pets or other uses are uncontrolled (New Mexico
Department of Game and Fish 2009, p. 2).
Historically, the northern leopard frog ranged Statewide in North
Dakota and is still quite common today (North Dakota Game and Fish
Department 2009, p. 1). Northern leopard frogs are widely distributed
throughout the State and locally abundant in some locations (Newman
2009, p. 1; Scherr 2009, pers. comm.) but surveys conducted by Bowers
et al. (1998, p. 372) found that the range of the northern leopard frog
was less extensive in the prairie potholes region of North Dakota than
previously described. Because of its distribution and local abundance,
the northern leopard frog has no special status in the State, and there
are no conservation programs that specifically target the northern
leopard frog (North Dakota Game and Fish Department 2009, p. 1).
Commercial frog licenses are available for unlimited collection of
northern leopard frogs (North Dakota Administrative Code 30-03-04).
NatureServe does not have a current ranking for North Dakota as it is
currently under review (NatureServe 2011, p. 1).
The Oregon Department of Fish and Wildlife ranks the northern
leopard frog as a ``Sensitive Critical'' species, meaning that it is
imperiled with extirpation from a specific geographic area of the State
due to small population sizes, habitat loss or degradation, or
immediate threats (Oregon Biodiversity Information Center 2010, p. 7,
13). The sensitive species list is primarily a non-regulatory tool
designed to provide a voluntary, proactive approach to conservation
(Oregon Department of Fish and Wildlife 2008, p. 1). The Oregon
Biodiversity Information Center lists the northern leopard frog as a
``List 2 Species'' meaning that it is threatened with extirpation or
presumed to be extirpated from the State of Oregon (Oregon Biodiversity
Information Center 2010, pp. 4, 13) and it is ranked S1/S2 (Critically
Imperiled/Imperiled) by NatureServe (NatureServe 2011, p. 1). The
Oregon Biodiversity Information Center (2010, p. 13), lists the
following counties as containing historical locations for the northern
leopard frog: Hood River, Wasco, Sherman, Gilliam, Morrow, Umatilla,
Jefferson, Crook, Grant, Baker, Malheur, Klamath, and Jackson Counties.
Rorabaugh (2005, p. 571) reported that northern leopard frogs are
extirpated from most historical
[[Page 61904]]
localities in Oregon. The six records we have from the Oregon Natural
Heritage Information Center are observations from 1975, 1980, 1990,
1995, 1996, and 2003. We have found no records, current or historical,
to indicate the presence of northern leopard frogs on either the Hart
Mountain National Antelope Refuge (southern Oregon) or Sheldon NWR
(northern Nevada) (Harper Collins 2009, pers. comm.). Frog surveys were
conducted at Sheldon NWR in summer 2009, but they detected only
nonnative American bullfrogs.
The status of the northern leopard frog in South Dakota is thought
to be stable and NatureServe lists the frog as secure (S5) (South
Dakota Department of Game, Fish, and Parks 2009, p. 1; NatureServe
2011, p. 1). The northern leopard has no specific protection in South
Dakota and can be collected for commercial and non-commercial bait
(South Dakota Laws and Regulations for Commercial Bait Dealers 2009, p.
1; South Dakota Department of Game, Fish, and Parks 2011, p. 23). The
species' range includes almost the entire State based upon historical
and current distribution maps (Fischer et al. 1999, p. 12; Naugle et
al. 2005, p. 285). Smith et al. (2005, p. 9) found northern leopard
frogs to be common in the Black Hills, and a Statewide herpetology
(amphibian and reptile) survey report indicates that the distribution
of the northern leopard frog in the State is stable (Backlund 2004, p.
8). However, there is no historical or recent abundance data to compare
current survey data that would indicate population trend (Backlund
2004, p. 9). Information received from Lacreek and Waubay NWRs and the
Huron Wetland Management District indicate northern leopard frogs are
prevalent (Flannders-Wanner 2009, pers. comm.; Hubers 2009, pers.
comm.; Koerner 2009, pers. comm.). Anuran auditory surveys (1997-1998)
found northern leopard frogs to be one of the most widespread and
wetland-abundant species in eastern South Dakota (Naugle et al. 2005,
p. 290).
The northern leopard frog's historic range in Texas was in the Rio
Grande Valley, El Paso County (a relatively small portion of the
State). However, extensive efforts to locate the frog have been
unsuccessful (Dixon 2000, pp. 42, 77). The northern leopard frog is
ranked S1 (Critically Imperiled) by NatureServe (NatureServe 2011, p.
1), but is not listed as a species of conservation concern in the Texas
Comprehensive Wildlife Conservation Strategy (Texas Parks and Wildlife
Department 2005, pp. 748-751). The Texas Parks and Wildlife Department
webpage (Texas Parks and Wildlife Department 2011a, p. 11) lists the
species as occurring in Texas, but the most current field guide for
amphibians and reptiles of Texas indicates the species is likely
extirpated (Dixon 2000, p. 77). The Texas Parks and Wildlife Department
requires that anyone who captures a wild animal, including frogs, be
licensed or permitted (Texas Parks and Wildlife Department 2011b, p.
1).
The Utah Division of Wildlife Resource