Endangered and Threatened Wildlife and Plants; Endangered Species Status for Sierra Nevada Yellow-Legged Frog and Northern Distinct Population Segment of the Mountain Yellow-Legged Frog, and Threatened Species Status for Yosemite Toad, 24255-24310 [2014-09488]
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Vol. 79
Tuesday,
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April 29, 2014
Part IV
Department of the Interior
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Fish and Wildlife Service
50 CFR Part 17
Endangered and Threatened Wildlife and Plants; Endangered Species
Status for Sierra Nevada Yellow-Legged Frog and Northern Distinct
Population Segment of the Mountain Yellow-Legged Frog, and Threatened
Species Status for Yosemite Toad; Final Rule
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Federal Register / Vol. 79, No. 82 / Tuesday, April 29, 2014 / Rules and Regulations
Executive Summary
DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS–R8–ES–2012–0100;
4500030113]
RIN 1018–AZ21
Endangered and Threatened Wildlife
and Plants; Endangered Species
Status for Sierra Nevada YellowLegged Frog and Northern Distinct
Population Segment of the Mountain
Yellow-Legged Frog, and Threatened
Species Status for Yosemite Toad
Fish and Wildlife Service,
Interior.
ACTION: Final rule.
AGENCY:
We, the U.S. Fish and
Wildlife Service (Service), determine
endangered species status under the
Endangered Species Act of 1973 (Act),
as amended, for the Sierra Nevada
yellow-legged frog and the northern
distinct population segment (DPS) of the
mountain yellow-legged frog (mountain
yellow-legged frog populations that
occur north of the Tehachapi
Mountains), and determine threatened
species status under the Act for the
Yosemite toad. The effect of this
regulation will be to add these species
to the List of Endangered and
Threatened Wildlife.
DATES: This rule becomes effective June
30, 2014.
ADDRESSES: This final rule is available
on the Internet at https://
www.regulations.gov and at the
Sacramento Fish and Wildlife Office.
Comments and materials we received, as
well as supporting documentation used
in preparing this rule, are available for
public inspection at https://
www.regulations.gov. All of the
comments, materials, and
documentation that we considered in
this rulemaking are available by
appointment, during normal business
hours at: U.S. Fish and Wildlife Service,
Sacramento Fish and Wildlife Office,
2800 Cottage Way, Room W–2605,
Sacramento, CA 95825; 916–414–6600
(telephone); 916–414–6712 (facsimile).
FOR FURTHER INFORMATION CONTACT:
Jennifer Norris, Field Supervisor, U.S.
Fish and Wildlife Service, Sacramento
Fish and Wildlife Office, 2800 Cottage
Way, Room W–2605, Sacramento, CA
95825; 916–414–6600 (telephone); 916–
414–6712 (facsimile). Persons who use a
telecommunications device for the deaf
(TDD) may call the Federal Information
Relay Service (FIRS) at 800–877–8339.
SUPPLEMENTARY INFORMATION:
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SUMMARY:
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Why we need to publish a rule. Under
the Endangered Species Act, a species
may warrant protection through listing
if it is endangered or threatened
throughout all or a significant portion of
its range. Listing a species as an
endangered or threatened species can be
only completed by issuing a rule.
This rule will finalize the listing of the
Sierra Nevada yellow-legged frog (Rana
sierrae) as an endangered species, the
northern DPS of the mountain yellowlegged frog (Rana muscosa) as an
endangered species, and the Yosemite
toad (Anaxyrus canorus) as a threatened
species.
The basis for our action. Under the
Endangered Species Act, we can
determine that a species is an
endangered or threatened species based
on any of 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 (E) Other natural or
manmade factors affecting its continued
existence.
We have determined that both the
Sierra Nevada yellow-legged frog and
the northern DPS of the mountain
yellow-legged frog are presently in
danger of extinction throughout their
entire ranges, based on the immediacy,
severity, and scope of the threats to their
continued existence. These include
habitat degradation and fragmentation,
predation and disease, climate change,
inadequate regulatory protections, and
the interaction of these various stressors
impacting small remnant populations. A
rangewide reduction in abundance and
geographic extent of surviving
populations of frogs has occurred
following decades of fish stocking,
habitat fragmentation, and most recently
a disease epidemic. Surviving
populations are smaller and more
isolated, and recruitment in diseased
populations is much reduced relative to
historic norms. This combination of
population stressors makes persistence
of these species precarious throughout
the currently occupied range in the
Sierra Nevada.
We have also determined that the
Yosemite toad is likely to become
endangered throughout its range within
the foreseeable future, based on the
immediacy, severity, and scope of the
threats to its continued existence. These
include habitat loss associated with
degradation of meadow hydrology
following stream incision consequent to
the cumulative effects of historical land
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management activities, notably livestock
grazing, and also the anticipated
hydrologic effects upon habitat from
climate change. We also find that the
Yosemite toad is likely to become
endangered through the direct effects of
climate change impacting small remnant
populations, likely compounded with
the cumulative effect of other threat
factors (such as disease).
Peer review and public comment. We
sought comments from independent
specialists to ensure that our
designations are based on scientifically
sound data, assumptions, and analyses.
We invited these peer reviewers to
comment on our listing proposal. We
also considered all comments and
information received during the
comment period.
Previous Federal Actions
Please refer to the proposed listing
rule for the Sierra Nevada yellow-legged
frog, the northern DPS of the mountain
yellow-legged frog, and the Yosemite
toad (78 FR 24472, April 25, 2013) for
a detailed description of previous
Federal actions concerning these
species.
We will also be finalizing critical
habitat designations for the Sierra
Nevada yellow-legged frog, the northern
DPS of the mountain yellow-legged, and
the Yosemite toad under the Act in the
near future.
Summary of Biological Status and
Threats for the Sierra Nevada YellowLegged Frog and the Northern DPS of
the Mountain Yellow-Legged Frog
Background
Please refer to the proposed listing
rule for the Sierra Nevada yellow-legged
frog and the northern DPS of the
mountain yellow-legged frog under the
Act (16 U.S.C. 1531 et seq.) for
additional species information. In the
proposed rule, we described two
separate species of yellow-legged frogs,
Rana sierrae and Rana muscosa, that
resulted from the recent taxonomic split
(see Taxonomy section below) of the
previously known Rana muscosa,
which we referred to in our proposed
rule as the mountain yellow-legged frog
‘‘species complex.’’ For clarity and in
order to maintain consistency with our
previous treatment of the southern DPS
of the mountain yellow legged frog in
southern California (67 FR 44382, July
2, 2002) as well as with our proposed
rule, and for the purposes of this
document, we retain the common name
of mountain yellow-legged frog for Rana
muscosa, as opposed to the new
common name, southern mountain
yellow-legged frog, as published by
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Crother et al. (2008, p. 11). We also note
that the California Department of Fish
and Game (CDFG) was recently renamed
the California Department of Fish and
Wildlife (CDFW). We refer to the
California Department of Fish and
Wildlife in all cases when discussing
the agency in the text. Where citations
are from CDFG documents, we include
CDFW in parentheses for clarification.
Taxonomy
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Please refer to the proposed listing
rule for the Sierra Nevada yellow-legged
frog and the northern DPS of the
mountain yellow-legged frog under the
Act (16 U.S.C. 1531 et seq.) for detailed
species information on taxonomy (78 FR
24472, April 25, 2013).
Vredenburg et al. (2007, p. 371)
determined that Rana sierrae occurs in
the Sierra Nevada north of the South
Fork Kings River watershed, along the
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east slope of the Sierra Nevada south
into Inyo County at the southern extent
of its range, and in the Glass Mountains
just south of Mono Lake; and that R.
muscosa occurs in the southern portion
of the Sierra Nevada within and south
of the South Fork Kings River watershed
to the west of the Sierra Nevada crest
(along with those populations
inhabiting southern California)
(Vredenburg et al. 2007, pp. 370–371).
The Monarch Divide separates these
species in the western Sierra Nevada,
while they are separated by the Cirque
Crest to the east (Knapp 2013,
unpaginated).
For purposes of this rule, we
recognize the species differentiation as
presented in Vredenburg et al. (2007, p.
371) and adopted by the official
societies mentioned above (Crother et
al. 2008, p. 11), and in this final rule we
refer to Rana sierrae as the Sierra
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Nevada yellow-legged frog, and we refer
to the Sierra Nevada populations of R.
muscosa as the northern DPS of the
mountain yellow-legged frog. In
California and Nevada, the Sierra
Nevada yellow-legged frogs occupy the
western Sierra Nevada north of the
Monarch Divide (in Fresno County) and
the eastern slope of the Sierra Nevada
(east of the crest) from Inyo County
through Mono County (including the
Glass Mountains), to areas north of Lake
Tahoe. The northern DPS of the
mountain yellow-legged frog occurs
only in California in the western Sierra
Nevada and extends from south of the
Monarch Divide in Fresno County
through portions of the Kern River
drainage. Figure 1 shows the
approximate species boundaries within
their historical ranges as determined by
Knapp (unpubl. data).
BILLING CODE 4310–55–P
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Species Description
Many studies cited in the rest of this
document include articles and reports
that were published prior to the official
species reclassification, where the
researchers may reference either one or
both species. Where possible and
appropriate, information will be
referenced specifically (either as Sierra
Nevada yellow-legged frog or the
northern DPS of the mountain yellowlegged frog) to reflect the split of the
species. Where information applies to
both species, the two species will be
referred to collectively as mountain
yellow-legged frog or mountain yellowlegged frog species complex.
Please refer to the proposed listing
rule for the Sierra Nevada yellow-legged
frog and the northern DPS of the
mountain yellow-legged frog under the
Act (16 U.S.C. 1531 et seq.) for
additional information about species
descriptions (78 FR 24472, April 25,
2013). The body lengths (snout to vent)
of the mountain yellow-legged frogs
range from 40 to 80 millimeters (mm)
(1.5 to 3.25 inches (in)) (Jennings and
Hayes 1994, p. 74). Females average
slightly larger than males, and males
have a swollen, darkened thumb base
(Wright and Wright 1949, pp. 424–430;
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Stebbins 1951, pp. 330–335; Zweifel
1955, p. 235; Zweifel 1968, p. 65.1).
Dorsal (upper) coloration in adults is
variable, exhibiting a mix of brown and
yellow, but also can be grey, red, or
green-brown, and is usually patterned
with dark spots (Jennings and Hayes
1994, p. 74; Stebbins 2003, p. 233).
These spots may be large (6 mm (0.25
in)) and few, smaller and more
numerous, or a mixture of both (Zweifel
1955, p. 230). Irregular lichen- or mosslike patches (to which the name
muscosa refers) may also be present on
the dorsal surface (Zweifel 1955, pp.
230, 235; Stebbins 2003, p. 233).
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The belly and undersurfaces of the
hind limbs are yellow or orange, and
this pigmentation may extend forward
from the abdomen to the forelimbs
(Wright and Wright 1949, pp. 424–429;
Stebbins 2003, p. 233). Mountain
yellow-legged frogs may produce a
distinctive mink or garlic-like odor
when disturbed (Wright and Wright
1949, p. 432; Stebbins 2003, p. 233).
Although these species lack vocal sacs,
they can vocalize in or out of water,
producing what has been described as a
faint clicking sound (Zweifel 1955, p.
234; Ziesmer 1997, pp. 46–47; Stebbins
2003, p. 233). Mountain yellow-legged
frogs have smoother skin, generally with
heavier spotting and mottling dorsally,
darker toe tips (Zweifel 1955, p. 234),
and more opaque ventral coloration
(Stebbins 2003, p. 233) than the foothill
yellow-legged frog.
The Sierra Nevada yellow-legged frog
and the northern DPS of the mountain
yellow-legged frog are similar
morphologically and behaviorally
(hence their shared taxonomic
designation until recently). However,
these two species can be distinguished
from each other physically by the ratio
of the lower leg (fibulotibia) length to
snout vent length. The northern DPS of
the mountain yellow-legged frog has
longer limbs (Vredenburg et al. 2007, p.
368). Typically, this ratio is greater than
or equal to 0.55 in the northern DPS of
the mountain yellow-legged frog and
less than 0.55 in the Sierra Nevada
yellow-legged frog.
Mountain yellow-legged frogs deposit
their eggs in globular clumps, which are
often somewhat flattened and roughly
2.5 to 5 centimeters (cm) (1 to 2 in) in
diameter (Stebbins 2003, p. 444). When
eggs are close to hatching, egg mass
volume averages 198 cubic cm (78 cubic
in) (Pope 1999, p. 30). Eggs have three
firm, jelly-like, transparent envelopes
surrounding a grey-tan or black vitelline
(egg yolk) capsule (Wright and Wright
1949, pp. 431–433). Clutch size varies
from 15 to 350 eggs per egg mass
(Livezey and Wright 1945, p. 703;
Vredenburg et al. 2005, p. 565). Egg
development is temperature dependent.
In laboratory breeding experiments, egg
hatching time ranged from 18 to 21 days
at temperatures of 5 to 13.5 degrees
Celsius (°C) (41 to 56 degrees Fahrenheit
(°F)) (Zweifel 1955, pp. 262–264). Field
observations show similar results (Pope
1999, p. 31).
The tadpoles of mountain yellowlegged frogs generally are mottled brown
on the dorsal side with a faintly yellow
venter (underside) (Zweifel 1955, p.
231; Stebbins 2003, p. 460). Total
tadpole length reaches 72 mm (2.8 in),
the body is flattened, and the tail
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musculature is wide (about 2.5 cm (1 in)
or more) before tapering into a rounded
tip (Wright and Wright 1949, p. 431).
The mouth has a maximum of eight
labial (lip) tooth rows (two to four upper
and four lower) (Stebbins 2003, p. 460).
Tadpoles may take more than 1 year
(Wright and Wright 1949, p. 431), and
often require 2 to 4 years, to reach
metamorphosis (transformation from
tadpoles to frogs) (Cory 1962b, p. 515;
Bradford 1983, pp. 1171, 1182; Bradford
et al. 1993, p. 883; Knapp and Matthews
2000, p. 435), depending on local
climate conditions and site-specific
variables.
The time required to reach
reproductive maturity in mountain
yellow-legged frogs is thought to vary
between 3 and 4 years post
metamorphosis (Zweifel 1955, p. 254).
This information, in combination with
the extended amount of time as a
tadpole before metamorphosis, means
that it may take 5 to 8 years for
mountain yellow-legged frogs to begin
reproducing. While the typical lifespan
of mountain yellow-legged frogs is
largely unknown, Matthews and Miaud
(2007, p. 991) estimated that the total
lifespan (including tadpole and adult
life stages) ranges up to 14 years, with
other documented estimates of up to 16
years of age for the Sierra Nevada
yellow-legged frog (Fellers et al. 2013, p.
155), suggesting that mountain yellowlegged frogs are long-lived amphibians.
Habitat and Life History
Mountain yellow-legged frogs
currently exist in montane regions of the
Sierra Nevada of California. Throughout
their range, these species historically
inhabited lakes, ponds, marshes,
meadows, and streams at elevations
typically ranging from 1,370 to 3,660
meters (m) (4,500 to 12,000 feet (ft))
((CDFG (CDFW)) 2011, pp. A–1–A–5),
but can occur as low as 1,067 m (3,500
ft) in the northern portions of their
range (USFS 2011, geospatial data;
USFS 2013, p. 4). Mountain yellowlegged frogs are highly aquatic; they are
generally not found more than 1 m (3.3
ft) from water (Stebbins 1951, p. 340;
Mullally and Cunningham 1956a, p.
191; Bradford et al. 1993, p. 886).
Mullally and Cunningham (1956a, p.
191) found adults sitting on rocks along
the shoreline, where there was little or
no vegetation. Although mountain
yellow-legged frogs may use a variety of
shoreline habitats, both tadpoles and
adults are observed less frequently at
shorelines that drop abruptly to a depth
of 60 cm (2 ft) than at open shorelines
that gently slope up to shallow waters
of only 5 to 8 cm (2 to 3 in) in depth
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(Mullally and Cunningham 1956a, p.
191; Jennings and Hayes 1994, p. 77).
At lower elevations within their
historical range, these species have been
associated with rocky streambeds and
wet meadows surrounded by coniferous
forest (Zweifel 1955, p. 237; Zeiner et al.
1988, p. 88), although, in general, little
is known about the ecology of mountain
yellow-legged frogs in Sierra Nevada
stream habitats (Brown 2013,
unpaginated). Zweifel (1955, p. 237)
found that streams utilized by adults
varied from streams having high
gradients and numerous pools, rapids,
and small waterfalls, to streams with
low gradients and slow flows, marshy
edges, and sod banks, while aquatic
substrates varied from bedrock to fine
sand, rubble (rock fragments), and
boulders. Jennings and Hayes (1994, p.
77) have indicated that mountain
yellow-legged frogs appear absent from
the smallest creeks, and suggest that it
is probably because these creeks have
insufficient depth for adequate refuge
and overwintering habitat. However,
Brown (2013, unpaginated) reports that
the frogs are found in small creeks,
although she notes that the extent to
which these are remnant populations
now excluded from preferred habitat is
not known. In the northern portion of
the Sierra Nevada yellow-legged frog
range, the remnant populations
primarily occur in stream habitats.
At higher elevations, these species
occupy lakes, ponds, tarns (small steepbanked mountain lakes or pools,
generally of glacial origin), and streams
(Zweifel 1955, p. 237; Mullally and
Cunningham 1956a, p. 191). Mountain
yellow-legged frogs in the Sierra Nevada
are most abundant in high-elevation
lakes and slow-moving portions of
streams (Zweifel 1955, p. 237; Mullally
and Cunningham 1956a, p. 191). The
borders of alpine (above the tree line)
lakes and mountain meadow streams
used by mountain yellow-legged frogs
are frequently grassy or muddy,
although many are bordered by exposed
glaciated bedrock. Zweifel (1955, pp.
237–238) suggested that alpine
lakeshores differ from the sandy or
rocky shores inhabited by mountain
yellow-legged frogs in lower elevation
streams.
Adult mountain yellow-legged frogs
breed in a variety of habitats including
the shallows of stillwater habitat (lakes
or ponds) and flowing inlet streams
(Zweifel 1955, p. 243; Pope 1999, p. 30).
Adults emerge from overwintering sites
immediately following snowmelt, and
will even move over ice to reach
breeding sites (Pope 1999, pp. 46–47;
Vredenburg et al. 2005, p. 565).
Mountain yellow-legged frogs deposit
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their eggs underwater in clusters, which
they attach to rocks, gravel, or
vegetation, or which they deposit under
banks (Wright and Wright 1949, p. 431;
Stebbins 1951, p. 341; Zweifel 1955, p.
243; Pope 1999, p. 30).
Lake depth is an important attribute
defining habitat suitability for mountain
yellow-legged frogs. At high elevations,
both frogs and tadpoles overwinter
under ice in lakes and streams. As
tadpoles must overwinter multiple years
before metamorphosis, successful
breeding sites are located in (or
connected to) lakes and ponds that do
not dry out in the summer, and also are
deep enough that they do not
completely freeze or become oxygendepleted (anoxic) in winter. Both adults
and tadpole mountain yellow-legged
frogs overwinter for up to 9 months in
the bottoms of lakes that are at least 1.7
m (5.6 ft) deep; however, overwinter
survival may be greater in lakes that are
at least 2.5 m (8.2 ft) deep (Bradford
1983, p. 1179; Vredenburg et al. 2005,
p. 565).
Bradford (1983, pp. 1173, 1178–1179)
found that, in years with exceptional
precipitation (61 percent above average)
and greater than normal ice-depths,
mountain yellow-legged frog die-offs
sometimes result from oxygen depletion
during winter in lakes less than 4 m (13
ft) in depth, finding that in ice-covered
lakes, oxygen depletion occurs most
rapidly in shallow lakes relative to
deeper lakes. However, tadpoles may
survive for months in nearly anoxic
conditions when shallow lakes are
frozen to the bottom. More recent work
reported populations of mountain
yellow-legged frogs overwintering in
lakes less than 1.5 m (5 ft) deep that
were assumed to have frozen to the
bottom, and yet healthy frogs emerged
the following July (Matthews and Pope
1999, pp. 622–623; Pope 1999, pp. 42–
43). Matthews and Pope 1999, p. 619)
used radio telemetry to find that, when
lakes had begun to freeze over, the frogs
were utilizing rock crevices, holes, and
ledges near shore, where water depths
ranged from 0.2 m (0.7 ft) to 1.5 m (5
ft). Vredenburg et al. (2005, p. 565)
noted that such behavior may be a
response to presence of introduced fish.
Matthews and Pope (1999, p. 622)
suggested that the granite surrounding
these overwintering habitats probably
insulates mountain yellow-legged frogs
from extreme winter temperatures, and
that they can survive, provided there is
an adequate supply of oxygen.
Mountain yellow-legged frog tadpoles
maintain a relatively high body
temperature by selecting warmer
microhabitats (Bradford 1984, p. 973).
During winter, tadpoles remain in
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warmer water below the thermocline
(the transition layer between thermally
stratified water). After spring overturn
(thaw and thermal mixing of the water),
they behaviorally modulate their body
temperature by moving to shallow, nearshore water when warmer days raise
surface water temperatures. During the
late afternoon and evening, mountain
yellow-legged frogs retreat to offshore
waters that are less subject to night
cooling (Bradford 1984, p. 974).
Available evidence suggests that adult
mountain yellow-legged frogs display
strong site fidelity and return to the
same overwintering and summer
habitats from year to year (Pope 1999, p.
45; Matthews and Preisler 2010, p. 252).
Matthews and Pope (1999, pp. 618–623)
observed that the frogs’ movement
patterns and habitat associations shifted
seasonally. Frogs were well-distributed
in most lakes, ponds, and creeks during
August, but moved to only a few lakes
by October. Matthews and Pope (1999,
pp. 618–623) established home-range
areas for 10 frogs and found that frogs
remained through August in the lake or
creek where they’d been captured, with
movement confined to areas ranging
from 19.4 to 1,028 square meters (m2)
(23.20 to 1,229 square yards (y2)). In
September, movements increased, with
home-ranges varying from 53 to 9,807
m2 in size (63.4 to 11,729 y2); six of nine
frogs tagged in September moved from
that lake by the end of the month,
suggesting a pattern in which adult
mountain yellow-legged frogs move
among overwintering, breeding, and
feeding sites during the year, with
narrow distributions in early spring and
late fall due to restricted overwintering
habitat (Pope and Matthews 2001, p.
791). Although terrestrial movements of
more than two or three hops from water
were previously undocumented,
overland movements exceeding 66 m
(217 ft) were observed in 17 percent of
tagged frogs, demonstrating that
mountain yellow-legged frogs move
overland as well as along aquatic
pathways (Pope and Matthews 2001, p.
791). Pope and Matthews (2001, p. 791)
also recorded a movement distance of
over 1 km (including a minimum of 420
m (0.26 miles) overland movement and
movement through a stream course).
The farthest reported distance of a
mountain yellow-legged frog from water
is 400 m (1,300 ft) (Vredenburg 2002, p.
4).
Within stream systems, Sierra Nevada
yellow-legged frogs have been
documented to move 1,032 m (3,385 ft)
over a 29-day period (Fellers et al. 2013,
p. 159). Wengert (2008, p. 18) conducted
a telemetry study that documented
single-season movement distances for
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Sierra Nevada yellow-legged frog of up
to 3.3 kilometers (km) (2.05 miles (mi))
along streams. Along stream habitats,
adults have been observed greater than
22 m (71 ft) from the water during the
overwintering period (Wengert 2008, p.
20). Additionally, during the duration of
the study, Wengert (2008, p. 13) found
that 14 percent of the documented frog
locations occurred greater than 0.2 m
(0.66 ft) from the stream edge. While
recent information suggests that the
frogs in the Wengert study may have
actually been foothill yellow-legged frog
(Rana boylii) (Poorten et al., 2013, p. 4),
we expect that the movement distances
recorded are applicable to the Sierra
Nevada yellow-legged frog within a
stream-based system, as the ecology is
comparable between the two sister taxa
in regard to stream systems.
Almost no data exist on the dispersal
of juvenile mountain yellow-legged
frogs away from breeding sites;
however, juveniles that may be
dispersing have been observed in small
intermittent streams (Bradford 1991, p.
176). Regionally, mountain yellowlegged frogs are thought to exhibit a
metapopulation structure (Bradford et
al. 1993, p. 886; Drost and Fellers 1996,
p. 424). Metapopulations are spatially
separated population subunits within
migratory distance of one another such
that individuals may interbreed among
subunits and populations may become
reestablished if they are extirpated
(Hanski and Simberloff 1997, p. 6).
Historical Range and Distribution
Mountain yellow-legged frogs were
historically abundant and ubiquitous
across many of the higher elevations
within the Sierra Nevada. Grinnell and
Storer (1924, p. 664) reported the Sierra
Nevada yellow-legged frog to be the
most common amphibian surveyed in
the Yosemite area. It is difficult to know
the precise historical ranges of the
Sierra Nevada yellow-legged frog and
the northern DPS of the mountain
yellow-legged frog, because projections
must be inferred from museum
collections that do not reflect systematic
surveys, and survey information
predating significant rangewide
reduction is very limited. However,
projections of historical ranges are
available using predictive habitat
modeling based on recent research
(Knapp, unpubl. data).
Historically, the range of the Sierra
Nevada yellow-legged frog extended in
California from north of the Feather
River, in Butte and Plumas Counties,
south to the Monarch Divide on the
west side of the Sierra Nevada crest in
Fresno County. East of the Sierra
Nevada crest in California, the historical
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range of the Sierra Nevada yellowlegged frog extends from areas north of
Lake Tahoe, through Mono County
(including the Glass Mountains) to Inyo
County. Historical records indicate that
the Sierra Nevada yellow-legged frog
also occurred at locations within the
Carson Range of Nevada, including
Mount Rose in Washoe County, and also
occurred in the vicinity of Lake Tahoe
in Douglas County, Nevada (Linsdale
1940, pp. 208–210; Zweifel 1955, p. 231;
Jennings 1984, p. 52; Knapp 2013,
unpaginated).
Historically, the northern DPS of the
mountain yellow-legged frog ranged
from the Monarch Divide in Fresno
County as far southward as
Breckenridge Mountain, in Kern County
(Vredenburg et al. 2007, p. 371). The
historical ranges of the two frog species
within the mountain yellow-legged
complex, therefore, meet each other
roughly along the Monarch Divide to the
north, and along the crest of the Sierra
Nevada to the east. Because we have
determined that the historic range of R.
muscosa is entirely within the State of
California, in this final rule we correct
the listing for the southern DPS of the
mountain yellow-legged frog to remove
Nevada from its historic range.
Park, Kings Canyon National Park, and
Sequoia National Park.
The most pronounced declines within
the mountain yellow-legged frog
complex have occurred north of Lake
Tahoe in the northernmost 125-km (78mi) portion of the range (Sierra Nevada
yellow-legged frog) and south of Kings
Canyon National Park in Tulare County
(the northern DPS of the mountain
yellow-legged frog). In the southernmost
50-km (31-mi) portion of the range, only
a few populations of the northern DPS
of the mountain yellow-legged frog
remain (Fellers 1994, p. 5; Jennings and
Hayes 1994, pp. 74–78); except for a few
small populations in the Kern River
drainage, the northern DPS of the
mountain yellow-legged frog is entirely
extirpated from all of Sequoia National
Park (Knapp 2013, unpaginated). As of
2000, mountain yellow-legged frog
populations were known to have
persisted in greater density in the
National Parks of the Sierra Nevada as
compared to the surrounding U.S.
Forest Service (USFS) lands, and the
populations that did occur in the
National Parks generally exhibited
higher abundances than those on USFS
lands (Bradford et al. 1994, p. 323;
Knapp and Matthews 2000, p. 430).
Current Range and Distribution
Since the time of the mountain
yellow-legged frog observations of
Grinnell and Storer (1924, pp. 664–665),
a number of researchers have reported
disappearances of these species from a
large fraction of their historical ranges
in the Sierra Nevada (Hayes and
Jennings 1986, p. 490; Bradford 1989, p.
775; Bradford et al. 1994, pp. 323–327;
Jennings and Hayes 1994, p. 78;
Jennings 1995, p. 133; Stebbins and
Cohen 1995, pp. 225–226; Drost and
Fellers 1996, p. 414; Jennings 1996, pp.
934–935; Knapp and Matthews 2000, p.
428; Vredenburg et al. 2005, p. 564).
The current distributions of the Sierra
Nevada yellow-legged frog and the
northern DPS of the mountain yellowlegged frog are restricted primarily to
publicly managed lands at high
elevations, including streams, lakes,
ponds, and meadow wetlands located
within National Forests and National
Parks. National Forests with extant
(surviving) populations of mountain
yellow-legged frogs include the Plumas
National Forest, Tahoe National Forest,
Humboldt-Toiyabe National Forest,
Lake Tahoe Basin Management Unit,
Eldorado National Forest, Stanislaus
National Forest, Sierra National Forest,
Sequoia National Forest, and Inyo
National Forest. National Parks with
extant populations of mountain yellowlegged frogs include Yosemite National
Population Estimates and Status
Monitoring efforts and research
studies have documented substantial
declines of mountain yellow-legged frog
populations in the Sierra Nevada. The
number of extant populations has
declined greatly over the last few
decades. Remaining populations are
patchily scattered throughout the
historical range (Jennings and Hayes
1994, pp. 74–78; Jennings 1995, p. 133;
Jennings 1996, p. 936). In the
northernmost portion of the range (Butte
and Plumas Counties), only a few Sierra
Nevada yellow-legged frog populations
have been documented since 1970
(Jennings and Hayes 1994, pp. 74–78;
CDFG (CDFW) et al., unpubl. data).
Declines of both species have also been
noted in the central and southern Sierra
Nevada (Drost and Fellers 1996, p. 420;
Knapp and Matthews 2001, pp. 433–
437; Knapp 2013, unpaginated). In the
southern Sierra Nevada (Sierra, Sequoia,
and Inyo National Forests; and Kings
Canyon and Yosemite National Parks),
modest to relatively large populations
(for example, breeding populations of
approximately 40 to more than 200
adults) of mountain yellow-legged frogs
do remain; however, in recent years
some large populations have been
extirpated in this area (Bradford 1991, p.
176; Bradford et al. 1994, pp. 325–326;
Knapp 2002a, p. 10, Wake and
Vredenburg 2009, pp. 11467–11470).
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Davidson et al. (2002, p. 1591)
reviewed 255 previously documented
mountain yellow-legged frog locations
(based on Jennings and Hayes 1994, pp.
74–78) throughout the historical range
and concluded that 83 percent of these
sites no longer support frog populations.
Vredenburg et al. (2007, pp. 369–371)
compared recent survey records (1995–
2004) with museum records from 1899–
1994 and reported that 92.5 percent of
historical Sierra Nevada yellow-legged
frog populations and 92.3 percent of
populations of the northern DPS of
mountain yellow-legged frog are now
extirpated.
CDFW (CDFG (CDFW) 2011, pp. 17–
20) used historical localities from
museum records covering the same time
interval (1899–1994), but updated
recent locality information with
additional survey data (1995–2010) to
significantly increase proportional
coverage from the Vredenburg et al.
(2007) study. These more recent surveys
failed to detect any extant frog
populations (within 1 km (0.63 mi), a
metric used to capture interbreeding
individuals within metapopulations) at
220 of 318 historical Sierra Nevada
yellow-legged frog localities and 94 of
109 historical northern DPS of the
mountain yellow-legged frog localities
(in the Sierran portion of their range).
This calculates to an estimated loss of
69 percent of Sierra Nevada yellowlegged frog metapopulations and 86
percent of northern DPS of the
mountain yellow-legged frog
metapopulations from historical
occurrences.
In addition to comparisons based on
individual localities, CDFW (CDFG
2011, pp. 20–25) compared historical
and recent population status at the
watershed scale. This is a rough index
of the geographic extent of the species
through their respective ranges. Within
the Sierra Nevada, 44 percent of
watersheds historically utilized by
Sierra Nevada yellow-legged frogs, and
59 percent of watersheds historically
utilized by northern DPS mountain
yellow-legged frogs, no longer support
extant populations. However, this
watershed-level survey methodology is
not a good indicator of population
changes because a watershed is counted
as recently occupied if a single
individual (at any life stage) is observed
within the entire watershed even though
several individual populations may
have been lost (CDFG (CDFW) 2011b, p.
20). Therefore, these surveys likely
underestimate population declines.
Many watersheds support only a single
extant metapopulation, which occupies
one to several adjacent water bodies
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(CDFG (CDFW) 2011, p. 20). Remaining
populations are generally very small.
Rangewide, declines of mountain
yellow-legged frog populations were
estimated at around one-half of
historical populations by the end of the
1980s (Bradford et al. 1994, p. 323).
Between 1988 and 1991, Bradford et al.
(1994a, pp. 323–327) resurveyed sites
known historically (1955 through 1979
surveys) to support mountain yellowlegged frogs. They did not detect frogs
at 27 historical sites on the Kaweah
River, and they detected frogs at 52
percent of historical sites within
Sequoia and Kings Canyon National
Parks and 12.5 percent of historical sites
outside of Sequoia and Kings Canyon
National Parks. Because this work was
completed before the taxonomic
division of mountain yellow-legged
frogs, we have not differentiated
between the two species here. When
both species are combined, this resurvey
effort detected mountain yellow-legged
frogs at 19.4 percent of historical sites
(Bradford et al. 1994, pp. 324–325).
Available information discussed
below indicates that the rates of
population decline have not abated, and
they have likely accelerated during the
1990s into the 2000s. Drost and Fellers
(1996, p. 417) repeated Grinnell and
Storer’s early 20th century surveys in
Yosemite National Park, and reported
frog presence at 2 of 14 historical sites
where what is now known as Sierra
Nevada yellow-legged frogs occurred.
The two positive sightings consisted of
a single tadpole at one site and a single
adult female at another. They identified
17 additional sites with suitable
mountain yellow-legged frog habitat,
and in those surveys, they detected 3
additional populations. In 2002, Knapp
(2002a, p. 10) resurveyed 302 water
bodies known to be occupied by
mountain yellow-legged frogs between
1995 and 1997, and 744 sites where
frogs were not previously detected.
Knapp found frogs at 59 percent of the
previously occupied sites, whereas 8
percent of previously unoccupied sites
were colonized. These data suggest an
extirpation rate five to six times higher
than the colonization rate within this
study area. The documented
extirpations appeared to occur nonrandomly across the landscape, were
typically spatially clumped, and
involved the disappearance of all or
nearly all of the mountain yellow-legged
frog populations in a watershed (Knapp
2002a, p. 9). CDFW (CDFG 2011, p. 20)
assessed data from sites where multiple
surveys were completed after 1995 (at
least 5 years apart). They found that the
Sierra Nevada yellow-legged frog was
not detected at 45 percent of sites where
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they previously had been confirmed,
while the mountain yellow-legged frog
(rangewide, including southern
California) was no longer detectable at
81 percent of historically occupied sites.
The USFS has been conducting a
rangewide, long-term monitoring
program for the Sierra Nevada yellowlegged frog and the northern DPS of the
mountain yellow-legged frog on
National Forest lands in the Sierra
Nevada, known as the Sierra Nevada
Amphibian Monitoring Program
(SNAMPH). This monitoring effort
provides unbiased estimates by using an
integrated unequal probability design,
and it provides numbers for robust
statistical comparisons across 5-year
monitoring cycles spanning 208
watersheds (Brown et al. 2011, pp. 3–4).
The results of this assessment indicate
that the species have declined in both
distribution and abundance. Based on
surveys conducted from 2002 through
2009, breeding activity was found in
about half (48 percent) of the
watersheds where the species were
found in the decade prior to SNAMPH
monitoring (1990 and 2001) (Brown et
al. 2011, p. 4). Breeding was found in
3 percent of watersheds where species
had been found prior to 1990.
Rangewide, breeding was found in 4
percent of watersheds. Moreover,
relative abundances were low; an
estimated 9 percent of populations were
large (numbering more than 100 frogs or
500 tadpoles); about 90 percent of the
watersheds had fewer than 10 adults,
while 80 percent had fewer than 10
subadults and 100 tadpoles (Brown et
al. 2011, p. 24).
To summarize population trends over
the available historical record, estimates
range from losses between 69 to 93
percent of Sierra Nevada yellow-legged
frog populations and 86 to 92 percent of
the northern DPS of the mountain
yellow-legged frog. Rangewide
reduction has diminished the number of
watersheds that support mountain
yellow-legged frogs somewhere between
the conservative estimates of 44 percent
in the case of Sierra Nevada yellowlegged frogs and at least 59 percent in
the case of the northern DPS of the
mountain yellow-legged frogs, to as high
as 97 percent of watersheds for the
mountain yellow-legged frog complex
across the Sierra Nevada. Remaining
populations are much smaller than
historical norms, and the density of
populations per watershed has declined
substantially; as a result, many
watersheds currently support single
metapopulations at low abundances.
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Distinct Vertebrate Population Segment
Analysis
Under the Act, we must consider for
listing any species, subspecies, or, for
vertebrates, any DPS of these taxa if
there is sufficient information to
indicate that such action may be
warranted. To implement the measures
prescribed by the Act, we, along with
the National Marine Fisheries Service
(National Oceanic and Atmospheric
Administration–Fisheries), developed a
joint policy that addresses the
recognition of DPSs for potential listing
actions (61 FR 4722). The policy allows
for a more refined application of the Act
that better reflects the biological needs
of the taxon being considered and
avoids the inclusion of entities that do
not require the Act’s protective
measures.
Under our DPS policy, three elements
are considered in a decision regarding
the status of a possible DPS as
endangered or threatened under the Act.
The elements are: (1) 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. In
other words, if we determine that a
population segment of a vertebrate
species being considered for listing is
both discrete and significant, we would
conclude that it represents a DPS, and
thus a ‘‘species’’ under section 3(16) of
the Act, whereupon we would evaluate
the level of threat to the DPS based on
the five listing factors established under
section 4(a)(1) of the Act to determine
whether listing the DPS as an
‘‘endangered species’’ or a ‘‘threatened
species’’ is warranted.
Please refer to the proposed listing
rule for detailed information about the
distinct vertebrate population segment
analysis for the northern DPS of the
mountain yellow-legged frog (78 FR
24472, April 25, 2013). We previously
confirmed the status of the southern
California population of the mountain
yellow-legged frog as a DPS at the time
that it was listed as endangered under
the Act (67 FR 44382, pp. 44384–
44385). We summarize below the
analysis for discreteness and
significance for the northern California
population of the mountain yellowlegged frog (in the Sierra Nevada); this
summary includes changes from the
proposed rule to address comments
received from the public (78 FR 24472,
April 25, 2013).
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Discreteness
Under our DPS Policy, a population
segment of a vertebrate species may be
considered discrete if it satisfies either
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 may
provide evidence of this separation); or
(2) it is delimited by international
governmental boundaries within which
significant differences in control of
exploitation, management of habitat,
conservation, status, or regulatory
mechanisms exist.
The analysis of the northern
population segment of the mountain
yellow-legged frog (Rana muscosa) (in
the Sierra Nevada) is based on the
marked separation from other
populations. The range of the mountain
yellow-legged frog is divided by a
natural geographic barrier, the
Tehachapi Mountains, which physically
isolates the populations in the southern
Sierra Nevada from those in the
mountains of southern California. The
distance of the geographic separation is
about 225 km (140 mi). The geographic
separation of the Sierra Nevada and
southern California frogs was
recognized in the earliest description of
the species by Camp (1917), who treated
frogs from the two areas as separate
subspecies within the R. boylii group
(see more on classification of the
mountain yellow-legged frogs in
Taxonomy). There is no contiguous
habitat that provides connectivity
between the two populations that is
sufficient for the migration, growth,
rearing, or reproduction of dispersing
frogs. Genetic differences wellsupported in the scientific literature
also provide evidence of this separation
(see Taxonomy). Therefore, we find that
the northern population segment of the
mountain yellow-legged frog (Rana
muscosa) (in the Sierra Nevada) is
discrete from the remainder of the
species.
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Significance
Under our DPS Policy, once we have
determined that a population segment is
discrete, we consider its biological and
ecological significance to the larger
taxon to which it belongs. Our DPS
policy provides several potential
considerations that may demonstrate the
significance of a population segment to
the remainder of its taxon, including: (1)
Evidence of the persistence of the
discrete population segment in an
ecological setting unusual or unique for
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the taxon, (2) evidence that loss of the
discrete population segment would
result in a significant gap in the range
of the taxon, (3) evidence that the
population segment represents the only
surviving natural occurrence of a taxon
that may be more abundant elsewhere as
an introduced population outside its
historic range, or (4) evidence that the
discrete population segment differs
markedly from the remainder of the
species in its genetic characteristics.
We have found substantial evidence
that three of the four significance
criteria are met by the discrete northern
population segment of the mountain
yellow-legged frog that occurs in the
Sierra Nevada. These include its
persistence in an ecological setting that
is unique for the taxon, evidence that its
loss would result in a significant gap in
the range of the taxon, and its genetic
uniqueness (reflecting significant
reproductive isolation over time). To
establish the significance of the discrete
northern population segment, we rely
on the effect that the loss of this
population segment would have on the
range of the taxon, and supplement that
with evidence that the population
segment persists in an ecological setting
unusual or unique for the taxon and also
differs from other population segments
in its genetic characteristics. There are
no introduced populations of the
northern DPS of the mountain yellowlegged frog outside of the species’
historical range.
Evidence indicates that loss of the
northern population segment of the
mountain yellow-legged frog (in the
Sierra Nevada) would result in a
significant gap in the range of the taxon.
The Sierran mountain yellow-legged
frogs comprise the entire distribution of
the species in approximately the
northern half of the species’ range, and
loss of the distinct population segment
in the northern portion of the range
could have significant conservation
implications for the species.
Furthermore, loss of the northern
population segment of the mountain
yellow-legged frog (in the Sierra
Nevada) would reduce the species to the
remaining small, isolated sites in the
streams of southern California (USFWS,
Jul 2012, pp. 11–12). Loss of the
northern population segment of the
mountain yellow-legged frog would
leave an area of the southern Sierra
Nevada over 150 km (93 mi) in length
without any ranid (frogs in the genus
Ranidae) frogs, which were once
abundant and widespread in the higher
elevation Sierra Nevada (Cory 1962b, p.
515; Fellers 1994, p. 5). The potential
loss of the northern population segment
of the mountain yellow-legged frog
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24263
would constitute a significant gap in the
range of the species.
One of the most striking differences
between the northern population
segment and the southern population
segment of the mountain yellow-legged
frogs is the difference in the ecological
setting in which they each persist.
Zweifel (1955, pp. 237–241) observed
that the frogs in southern California are
typically found in steep-gradient
streams in the chaparral belt at low
elevations (370 m (1,220 ft)), even
though they may range into small
meadow streams at higher elevations up
to 2,290 m (7,560 ft). In contrast, frogs
from the northern population segment
of mountain yellow-legged frogs are
most abundant in high-elevation lakes
and slow-moving portions of streams
where winter conditions are extreme.
David Bradford’s (1989) southern Sierra
Nevada study of mountain yellowlegged frogs, for example, was
conducted in Sequoia and Kings Canyon
National Parks at high elevations
between 2,910 and 3,430 m (9,600 to
11,319 ft). The rugged canyons of the
arid mountain ranges of southern
California, where waters seldom freeze,
bear little resemblance to the alpine
lakes and streams of the Sierra Nevada
where adult frogs and tadpoles must
overwinter at the bottoms of ice and
snow-covered lakes for up to 9 months
of the year. The significantly different
ecological settings between mountain
yellow-legged frogs in southern
California and those in the northern
population segment (in the Sierra
Nevada) distinguish these populations
from each other.
Finally, the northern population
segment of the mountain yellow-legged
frog is biologically significant based on
genetic differences. Vredenburg et al.
(2007, p. 361) identified that two of
three distinct genetic clades (groups of
distinct lineage) constitute the northern
range of the mountain yellow-legged
frog found in the Sierra Nevada, with
the remaining clade represented by the
endangered southern California DPS of
the mountain yellow-legged frog. Macey
et al. (2001, p. 141) estimated the
genetic divergence between the northern
population of mountain yellow-legged
frogs (in the Sierra Nevada) and the
southern population of mountain
yellow-legged frogs (in southern
California) to have occurred 1.4 million
years before present (mybp), thereby
indicating functional isolation.
The loss of the northern population of
the mountain yellow-legged frog would
result in a significant gap in the range
of the mountain yellow-legged frog
species. The differences between the
ecological settings for the southern
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population of mountain yellow-legged
frogs (steep-gradient streams that
seldom freeze) and the northern
population of mountain yellow-legged
frogs (high-elevation lakes and slowmoving portions of streams where frogs
overwinter under ice and snow for up
to 75 percent of the year) are significant.
Additionally, the genetic distinction
between these two populations reflects
isolation for over a million years.
Therefore based on the information
discussed above, we find that northern
population of the mountain yellowlegged frog (in the Sierra Nevada
mountains) meets the significance
criteria under our Policy Regarding the
Recognition of Distinct Vertebrate
Population Segments (61 FR 4722).
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DPS Conclusion
Based on the best scientific and
commercial data available on
distribution as well as ecological setting
and genetic characteristics of the
species, we have determined that the
northern population segment of the
mountain yellow-legged frog (in the
Sierra Nevada) is both discrete and
significant per our DPS policy.
Therefore, we conclude that the
northern discrete population segment of
the mountain yellow-legged frog is a
DPS, and thus a ‘‘species’’ under section
3(16) of the Act. Our determination of
biological and ecological significance is
appropriate because the population
segment has a geographical distribution
that is biologically meaningful.
Summary of Changes From the
Proposed Rule for the Sierra Nevada
Yellow-Legged Frog and the Northern
DPS of the Mountain Yellow-Legged
Frog
Based on peer review, Federal and
State, and public comments (see
comments in the Summary of
Comments and Recommendations
section below), we have clarified
information in the sections provided for
the Sierra Nevada yellow-legged frog
and the northern DPS of the mountain
yellow-legged frog to better characterize
our knowledge of the species’ habitat
requirements, correcting some
information based on peer review
(vocalizations (Species Description),
species ranges (Taxonomy and Historic
and Current Ranges and Distribution
sections), current distribution in
Sequoia National Park (Historic and
Current Ranges and Distribution), and
clarifying the basis for our
determination of significance for the
northern population of the mountain
yellow-legged frog in response to public
comments (Distinct Vertebrate
Population Segment)), occasionally
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adding additional information where
needed. In the Summary of Factors
Affecting the Species section, we have
re-ordered threats in Factor A so that the
primary activity that has modified the
habitat of the mountain yellow-legged
frog complex is addressed first, while
activities with potential only for
localized effects are addressed later.
Based on peer review, and Federal,
State, county, and public comments, we
have added information where needed
and clarified our findings on the role of
current activities, such as grazing,
recreation, packstock use, etc., in
species declines. We reviewed the
analysis of dams and diversions that we
presented in the proposed rule and
determined that most large reservoir
facilities are below the current range of
the mountain yellow-legged frogs. We
revised the dams and water diversions
threat magnitude from moderate
prevalent in the proposed rule to minor
localized where such structures occur in
this final rule.
In the proposed rule, we stated that
grazing presented a minor prevalent
threat. We reworded this final rule to
more accurately reflect the contribution
of legacy effects of past grazing levels to
this threat assessment. We found that
current livestock grazing that complies
with forest standards and guidelines is
not expected to negatively affect
mountain yellow-legged frog
populations in most cases, although
limited exceptions could occur (where
extant habitat is limited and legacy
effects to meadows still require
restoration, where habitat is limited
such as in stream riparian zones or
small meadows, or where grazing
standards are exceeded). Rangewide,
livestock grazing is not a substantial
threat to the species.
In response to information provided
during the public comment period, we
added a discussion of mining activities
in the Factor A discussion. In this final
rule, we determine that, while most
mining activities take place below the
extant ranges of the species, where some
types of mining activities occur,
localized habitat-related effects may
result.
We added new information available
on packstock grazing, retaining our
finding that packstock grazing is only
likely to be a threat to mountain yellowlegged frogs in limited situations. We
also added more information on roads
and timber harvests, and we clarified
that these activities primarily do not
occur where there are extant
populations (except where frogs occur
in the northern or lower elevation
portions of the range), and that USFS
standards are generally designed to limit
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potential effects of such activities. We
clarified the threat magnitude for roads
and timber harvest from minor
prevalence rangewide to not a threat to
extant populations across much of the
species’ ranges (although they may pose
important habitat-related effects to the
species in localized areas). We reviewed
information provided by the U.S. Forest
Service (USFS), the National Park
Service (NPS), CDFW, and others on
recreation activities, and we changed
our conclusion on the recreation threat
magnitude from low significance to the
species overall to not considered a
threat to populations over much of their
range. However, we recognize that there
may be localized effects, especially
outside of backcountry areas where use
is high or where motorized and
mechanical use occurs in extant frog
habitat.
We added a brief discussion of
bullfrogs (Lithobates catesbeiana) under
Factor C for mountain yellow-legged
frogs noting that bullfrog predation and
competition is expected to have
population-level effects to mountain
yellow-legged frog populations in those
low elevation areas, or in the Lake
Tahoe Basin, where the two species may
co-occur. We slightly revised our
characterization of the recent
population declines of the mountain
yellow-legged frogs due to
Batrachochytrium dendrobatidis (Bd),
identifying the fungus as one of the
primary drivers of recent declines, and
adding information provided by peer
reviewers and agencies. We also added
information to our discussion under
Factor D, including information about
the National Park Service Organic Act,
information on the provision in the
Wilderness Act about withdrawing
minerals, and information on the status
of the Sierra Nevada yellow-legged frog
and the mountain yellow-legged frog
under the California Endangered
Species Act (CESA). We also moved
discussion of current CDFW fisheries
management to the ‘‘Habitat
Modification Due to Introduction of
Trout to Historically Fishless Areas’’
section under Factor A.
We removed the discussion of
contaminants under Factor E and refer
readers to the proposed rule. Although
we received additional information that
clarified some text and provided
additional references regarding
contaminants, the clarifications
supported our conclusions in the
proposed rule that the best available
information indicates that contaminants
do not pose a current or continuing
threat to the species. We also added
additional information either available
in our files, or provided by commenters,
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to clarify and support our finding on the
threat of climate change. We revised the
explanation in the determinations for
each species to reflect the above
changes.
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Summary of Factors Affecting the
Species
Section 4 of the Act (16 U.S.C. 1533),
and its implementing regulations at 50
CFR part 424, set forth the procedures
for adding species to the Federal Lists
of Endangered and Threatened Wildlife
and Plants. Under section 4(a)(1) of the
Act, we may list a species 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; and (E)
other natural or manmade factors
affecting its continued existence. Listing
actions may be warranted based on any
of the above threat factors, singly or in
combination. Each of these factors is
discussed below, and changes from the
proposed rule (78 FR 24472, April 25,
2013) are reflected in these discussions.
The following analysis is applicable to
both the Sierra Nevada yellow-legged
frog (Rana sierrae) and the northern
distinct population segment of the
mountain yellow-legged frog (Rana
muscosa).
Factor A. The Present or Threatened
Destruction, Modification, or
Curtailment of Its Habitat or Range
A number of hypotheses, including
habitat modification (including loss of
vegetation, loss of wetlands, habitat
modification for urban development,
and degradation of upland habitats)
have been proposed for recent global
amphibian declines (Bradford et al.
1993, p. 883; Corn 1994, p. 62; Alford
and Richards 1999, p. 134). However,
physical habitat modification has not
been associated with the rangewide
decline of mountain yellow-legged
frogs. Mountain yellow-legged frogs
occur primarily at high elevations in the
Sierra Nevada, which have not had the
types or extent of large-scale habitat
conversion and physical disturbance
that have occurred at lower elevations
(Knapp and Matthews 2000, p. 429).
Thus, direct habitat destruction or
modification associated with intensive
human activities has not been
implicated in the decline of this species
(Davidson et al. 2002, p. 1597).
However, other human activities may
have played a role in the modification
of mountain yellow-legged frog habitat.
We have identified the following
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habitat-related activities as potentially
relevant to the conservation status of the
mountain yellow-legged frog complex:
Fish introductions (see also Factor C,
below), dams and water diversions,
livestock grazing, timber management,
road construction and maintenance,
packstock use, recreational activities,
and fire management activities. Such
activities may have degraded habitat in
ways that have reduced its capacity to
sustain viable populations and may
have fragmented and isolated mountain
yellow-legged frog populations from
each other.
Habitat Modification Due to
Introduction of Trout to Historically
Fishless Areas
One habitat feature that is
documented to have a significant
detrimental impact to mountain yellowlegged frog populations is the presence
of introduced trout resulting from
stocking programs for the creation and
maintenance of a recreational fishery.
To further angling success and
opportunity, trout stocking programs in
the Sierra Nevada started in the late
19th century (Bahls 1992, p. 185; Pister
2001, p. 280). This anthropogenic
activity has community-level effects and
is one of the primary threats to
mountain yellow-legged frog habitat and
species viability.
Prior to extensive trout planting
programs, almost all streams and lakes
in the Sierra Nevada at elevations above
1,800 m (6,000 ft) were fishless. Several
native fish species occur naturally in
aquatic habitats below this elevation in
the Sierra Nevada (Knapp 1996, pp. 12–
14; Moyle et al. 1996, p. 354; Moyle
2002, p. 25), but natural barriers
prevented fish from colonizing the
higher-elevation waters of the Sierra
Nevada watershed (Moyle et al. 1996, p.
354). The upper reaches of the Kern
River, where native fish such as the
Little Kern golden trout (Oncorhynchus
mykiss whitei) and California golden
trout (O. m. aguabonita) evolved,
represent the only major exception to
the 1,800-m (6,000-ft) elevation limit for
fishes within the range of the mountain
yellow-legged frog in the Sierra Nevada
(Moyle 2002, p. 25). Additionally, prior
to extensive planting, native Paiute
cutthroat (O. clarki seleneris) and
Lahontan cutthroat (O. c. henshawi)
were limited in their distribution to
several rivers, streams, and limited large
lakes in the eastern Sierra Nevada
(Knapp 1996, p. 369; Moyle 1996 et al.,
pp. 954–958), indicating some overlap
with the range of the Sierra Nevada
yellow-legged frog.
Some of the first practitioners of trout
stocking in the Sierra Nevada were the
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Sierra Club, local sportsmen’s clubs,
private citizens, and the U.S. military
(Knapp 1996, p. 8; Pister 2001, p. 280).
As more hatcheries were built and the
management of the trout fishery became
better organized, fish planting
continued for the purpose of increased
angler opportunities and success (Pister
2001, p. 281). After World War II, the
method of transporting trout to highelevation areas changed from packstock
to aircraft, which allowed stocking in
more remote lakes and in greater
numbers. With the advent of aerial
stocking, trout planting expanded to
new areas, with higher efficiency.
Brook trout (Salvelinus fontinalis),
brown trout (Salmo trutta), rainbow
trout (Oncorhynchus mykiss), and other
trout species assemblages have been
planted in most streams and lakes of the
Sierra Nevada (Knapp 1996, p. 8; Moyle
2002, p. 25). Since the advent of aerial
stocking, backcountry areas not
accessible by truck are stocked by air
(Pert 2002, pers. comm.), which limits
stocking to lakes. National Forests in the
Sierra Nevada have a higher proportion
of lakes with fish occupancy than do
National Parks (Knapp 1996, p. 3),
primarily because the National Park
Service (NPS) began phasing out fish
stocking within their jurisdictional
boundaries in 1969, with limited
stocking occurring until it was
terminated altogether in Sierra Nevada
National Parks in 1991 (Knapp 1996, p.
9). California Department of Fish and
Wildlife (CDFW) continues to stock
trout in National Forest water bodies,
but in 2001 reduced the number of
stocked water bodies to reduce impacts
to native amphibians (ICF Jones &
Stokes 2010, pp. ES–1–ES–16). Current
stocking decisions are based on criteria
outlined in the Environmental Impact
Report for the Hatchery and Stocking
Program (ICF Jones & Stokes 2010,
Appendix K).
Fish stocking as a practice has been
widespread throughout the range of
both species of mountain yellow-legged
frogs. Knapp and Matthews (2000, p.
428) indicated that 65 percent of the
water bodies that were 1 ha (2.5 ac) or
larger in National Forests they studied
were stocked with fish on a regular
basis. Over 90 percent of the total water
body surface area in the John Muir
Wilderness was occupied by nonnative
trout (Knapp and Matthews 2000, p.
434).
Another detrimental feature of fish
stocking is that, in the Sierra Nevada,
fish often persist in water bodies even
after stocking ceases. Thirty-five to 50
percent of lakes larger than 1 ha (2.5 ac)
within Sierra Nevada National Parks are
occupied by nonnative fish, which is
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only a 29 to 44 percent decrease in fish
occupancy since fish stocking was
terminated around 2 decades before the
estimate was made (Knapp 1996, p. 1).
Though data on fish occupancy in
streams are lacking throughout the
Sierra Nevada, Knapp (1996, pp. 9–11)
estimated that 60 percent of the streams
in Yosemite National Park were still
occupied by introduced trout because
trout readily move out of lakes to
colonize both inlet and outlet streams.
The presence of trout in these once
fishless waters has modified the habitat
at a landscape scale.
Thus, the frog’s habitat has been
modified due to the introduction of a
nonnative predator that both competes
for limited food resources and directly
preys on mountain yellow-legged frog
tadpoles and adults (see Factor C
below). Presence of nonnative trout in
naturally fishless ecosystems has had
profound effects on the structure and
composition of faunal assemblages,
severely reducing not only amphibians,
but also zooplankton and large
invertebrate species (see Knapp 1996, p.
6; Bradford et al. 1998, p. 2489; Finlay
and Vredenburg 2007, pp. 2194–2197).
Within the frog’s historical range, past
trout introductions and the continuing
presence of fish in most lakes resulted
in the elimination of frogs from most
waters that were suitable for fish. Across
the range of these species in the Sierra
Nevada, the presence of fish in most of
the deeper lakes has altered the aquatic
habitat that mountain yellow-legged
frogs rely on for overwintering and
breeding, and has also reduced
connectivity among frog populations.
Fish now populate the deeper lakes and
connecting streams and largely separate
and increase the distance between the
current sites inhabited by the highlyaquatic frogs (the connectivity of
occupied sites in present versus former
fishless conditions differs by
approximately 10-fold) (Bradford et al.
1993, pp. 884–887; Knapp 1996, pp.
373–379). Where reservoirs harbor
introduced fish, successful reproduction
of mountain yellow-legged frogs may be
reduced if there are no shallow side
channels or separate pools (Jennings
1996, p. 939). Most reservoirs do not
overlap significantly with the current
extant range of the species (CDFW 2013,
p. 3) (see Dams and Water Diversions
below); however, a number of reservoirs
were constructed in the mid-1900s at
mid-elevations within lower edges of
the species’ historic range (for example,
Sierra Nevada yellow-legged frogs were
taken from Bear River Reservoir
(Eldorado National Forest), Union
Reservoir (Stanislaus National Forest),
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and several others). With the exception
of one 1999 record from Faggs Reservoir
on the Plumas National Forest, all of
several dozen records of the species
from reservoirs are pre-1975, and at
least half pre-date the water
development projects at those locations
(Brown et al. 2009, p. 78). All of these
reservoirs now harbor introduced fish
species, and at least two also harbor
bullfrogs, suggesting that subsequent
introductions may have played a role in
past declines in those areas (see Brown
et al. 2009, p. 78).
The body of scientific research has
demonstrated that introduced trout have
negatively impacted mountain yellowlegged frogs over much of the Sierra
Nevada (Grinnell and Storer 1924, p.
664; Bradford 1989, pp. 775–778;
Bradford et al. 1993, pp. 882–888;
Knapp 1994, p. 3; Drost and Fellers
1996, p. 422; Knapp 1996, pp. 13–15;
Bradford et al. 1998, pp. 2482, 2489;
Knapp and Matthews 2000, p. 428;
Knapp et al. 2001, p. 401). Fish stocking
programs have negative ecological
implications because fish eat aquatic
fauna, including amphibians and
invertebrates (Bahls 1992, p. 191; Erman
1996, p. 992; Jennings 1996, p. 939;
Knapp 1996, pp. 373–379; Matthews et
al. 2001, pp. 1135–1136; Pilliod and
Peterson 2001, p. 329; Schindler et al.
2001, p. 309; Moyle 2002, p. 58;
Epanchin et al. 2010, p. 2406). Finlay
and Vredenburg (2007, p. 2187)
documented that the same benthic
(bottom-dwelling) invertebrate resource
base sustains the growth of both frogs
and trout, suggesting that competition
with trout for prey is an important factor
that may contribute to the decline of the
mountain yellow-legged frog.
Introductions of salmonids to fishless
lakes have also been associated with
alteration of nutrient cycles and primary
productivity in mountain lakes,
including those in the Sierra Nevada
(Schindler et al. 2001, pp. 308, 313–
319).
Knapp and Matthews (2000, p. 428)
surveyed more than 1,700 water bodies,
and concluded that a strong negative
correlation exists between introduced
trout and mountain yellow-legged frogs
(Knapp and Matthews 2000, p. 435).
Consistent with this finding are the
results of an analysis of the distribution
of mountain yellow-legged frog
tadpoles, which indicate that the
presence and abundance of this life
stage are reduced dramatically in fishstocked lakes (Knapp et al. 2001, p.
408). Knapp (2005a, pp. 265–279) also
compared the distribution of nonnative
trout with the distributions of several
amphibian and reptile species in 2,239
lakes and ponds in Yosemite National
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Park, and found that mountain yellowlegged frogs were five times less likely
to be detected in waters where trout
were present. Even though stocking
within the National Park ceased in 1991,
more than 50 percent of water bodies
deeper than 4 m (13 ft) and 75 percent
deeper than 16 m (52 ft) still contained
trout populations in 2000–2002 (Knapp
2005a, p. 270). Both trout and mountain
yellow-legged frogs utilize deeper water
bodies. Based on the results from Knapp
(2005a), the reduced detection of frogs
in trout-occupied waters indicates that
trout are excluding mountain yellowlegged frogs from some of the best
aquatic habitat.
Several aspects of the mountain
yellow-legged frog’s life history are
thought to exacerbate its vulnerability to
extirpation by trout (Bradford 1989, pp.
777–778; Bradford et al. 1993, pp. 886–
888; Knapp 1996, p. 14; Knapp and
Matthews 2000, p. 435). Mountain
yellow-legged frogs are highly aquatic
and are found primarily in lakes, most
of which now contain trout (Knapp
1996, p. 14). In comparison to other
Sierran frogs, mountain yellow-legged
frog tadpoles generally need at least 2
years to reach metamorphosis, which
restricts breeding to waters that are deep
enough to avoid depletion of oxygen
when ice-covered (Knapp 1996, p.14).
Overwintering adults must also avoid
oxygen depletion when the water is
covered by ice, generally limiting
overwintering to deeper waters that do
not become anoxic (Mullally and
Cunningham 1956a, p. 194; Bradford
1983, p. 1179; Knapp and Matthews
2000, pp. 435–436). At high elevations,
both tadpoles and adults overwinter
under ice for up to 9 months (Bradford
1983, p. 1171). These habitat
requirements appear to restrict
successful breeding and overwintering
to the deeper water bodies where the
chances of summer drying and winter
freezing are reduced, the same water
bodies that are most suitable for fishes;
fishes also need deeper water bodies
where the chances of summer drying
and winter freezing are reduced
(Bradford 1983, pp. 1172–1179; Knapp
1996, p. 14; Knapp and Matthews 2000,
pp. 429, 435–436). Past fish-stocking
practices targeted the deeper lakes, so
the percentage of water bodies
containing fish has increased with water
depth, resulting in elimination of
mountain yellow-legged frogs from once
suitable habitats in which they were
historically most common, and thereby
generally isolating populations to the
shallower, marginal habitats that do not
have fish (Bradford 1983, pp. 1172–
1179; Bradford et al. 1993, pp. 884, 886–
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887; Knapp and Matthews 2000, pp.
435–436).
Mountain yellow-legged frogs and
trout (native and nonnative) do co-occur
at some sites, but these co-occurrences
are generally thought to represent
mountain yellow-legged frog ‘‘sink’’
populations (areas with negative
population growth rates in the absence
of immigration) (Bradford et al. 1998, p.
2489; Knapp and Matthews 2000, p.
436). Mountain yellow-legged frogs have
also been extirpated at some fishless
bodies of water (Bradford 1991, p. 176;
Drost and Fellers 1996, p. 422). A
possible explanation is the isolation and
fragmentation of remaining populations
due to introduced fishes in the streams
that once provided mountain yellowlegged frogs with dispersal and
recolonization routes; these remote
populations are now non-functional as
metapopulations (Bradford 1991, p. 176;
Bradford et al. 1993, p. 887). Based on
a survey of 95 basins within Sequoia
and Kings Canyon National Parks,
Bradford et al. (1993, pp. 885–886)
estimated that the introduction of fishes
into the study area resulted in an
approximately 10-fold increase in
habitat fragmentation between
populations of mountain yellow-legged
frogs. Knapp and Matthews (2000, p.
436) believe that this fragmentation has
further isolated mountain yellow-legged
frogs within the already marginal
habitat left unused by fishes.
Fragmentation of mountain yellowlegged frog habitat renders populations
more vulnerable to extirpation from
random events (such as disease) (Wilcox
1980, pp. 114–115; Bradford et al. 1993,
p. 887; Hanski and Simberloff 1997, p.
21; Knapp and Matthews 2000, p. 436).
Isolated population locations may have
higher extinction rates because trout
prevent successful recolonization and
dispersal to and from these sites
(Bradford et al. 1993, p. 887; Blaustein
et al. 1994a, p. 7; Knapp and Matthews
2000, p. 436). If the distance between
sites is too great, amphibians may not
readily recolonize unoccupied sites
following local extinctions because of
physiological constraints, the tendency
to move only short distances, and high
site fidelity. Finally, frogs that do
attempt recolonization may emigrate
into fish-occupied habitat and perish,
rendering sites with such
metapopulation dynamics less able to
sustain frog populations.
In 2001, CDFW revised fish stocking
practices and implemented an informal
policy on fish stocking in the range of
the Sierra Nevada yellow-legged frog
and northern DPS of the mountain
yellow-legged frog. This policy directs
that: (1) Fish will not be stocked in lakes
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with known populations of mountain
yellow-legged frogs, nor in lakes that
have not yet been surveyed for
mountain yellow-legged frog presence;
(2) waters will be stocked only with a
fisheries management justification; and
(3) the number of stocked lakes will be
reduced over time. In 2001, the number
of lakes stocked with fish within the
range of the mountain yellow-legged
frog in the Sierra Nevada was reduced
by 75 percent (Milliron 2002, pp. 6–7;
Pert et al. 2002, pers. comm.). Current
CDFW guidelines stipulate that water
bodies within the same basin and 2 km
(1.25 mi) from a known mountain
yellow-legged frog population will not
be stocked with fish unless stocking is
justified through a management plan
that considers all the aquatic resources
in the basin, or unless there is heavy
angler use and no opportunity to
improve the mountain yellow-legged
frog habitat (Milliron 2002a, p. 5). The
Hatchery and Stocking Program
Environmental Impact Report/
Environmental Impact Statement,
finalized in 2010 (ICF Jones & Stokes
2010, Appendix K), outlines a decision
approach to mitigate fish stocking
effects on Sierra amphibians that
prohibits fish stocking in lakes with
confirmed presence of a limited number
of designated species, including the
mountain yellow-legged frogs (see ICF
Jones & Stokes 2010, Appendix E) using
recognized survey protocols. Large
reservoirs generally continue to be
stocked to provide a put-and-take
fishery for recreational angling.
As part of the High Mountain Lakes
Project, CDFW is in the process of
developing management plans for
basins within the range of the Sierra
Nevada yellow-legged frog and the
northern DPS of mountain yellowlegged frog (CDFG (CDFW) 2001, p. 1;
Lockhart 2011, pers. comm.). CDFW
states that objectives of the basin plans
specific to the mountain yellow-legged
frog include management in a manner
that maintains or restores native
biodiversity and habitat quality,
supports viable populations of native
species, and provides for recreational
opportunities that consider historical
use patterns (CDFG (CDFW) 2001, p. 3).
They state that, under this approach,
lakes that support mountain yellowlegged populations in breeding,
foraging, or dispersal, and/or present
opportunities to restore or expand
habitat, are managed for the
conservation of the species. Lakes that
do not support mountain yellow-legged
frogs are not viable restoration
opportunities, and lakes that support
trout populations are managed primarily
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for recreational angling (CDFG (CDFW)
2001, p. 3). They further note that lakes
managed for recreational angling may be
stocked if CDFW determines that
stocking the lake will achieve a
desirable fisheries management
objective and is not otherwise precluded
by stocking decision guidelines and
agreements (for stocking decision
documents, see CDFW 2013, pp. 1, 2).
Since the mid-1990s, various parties,
including researchers, CDFW, NPS, and
the USFS, have implemented a variety
of projects to actively restore habitat for
the mountain yellow-legged frog via the
removal of nonnative trout (USFS 2011,
pp. 128–130; NPS 2013, pp. 3–5).
Although fish stocking has been
curtailed within many occupied basins,
the impacts to frog populations persist
due to the presence of self-sustaining
fish populations in some of the best
habitat that normally would have
sustained mountain yellow-legged frogs.
The fragmentation that persists across
the range of these frog species renders
them more vulnerable to other
population stressors, and recovery is
slow, if not impossible, without costly
and physically difficult direct human
intervention (such as physical and
chemical trout removal) (see Knapp et
al. 2007a, pp. 11–19). While most of the
impacts occurred historically, the
impact upon the biogeographic
(population/metapopulation) integrity
of the species will be long-lasting.
Currently, habitat degradation and
fragmentation by fish is considered a
highly significant and prevalent threat
to persistence and recovery of the
species.
Dams and Water Diversions
While a majority of dams and water
diversions within the Sierra Nevada are
located at lower elevations (USFS 2011,
p. 83), some large reservoirs have been
constructed within the historic range of
the mountain yellow-legged frog
complex. These large reservoirs include,
but are not limited to Huntington Lake,
Florence Lake, Lake Thomas A. Edison,
Saddlebag Lake, Cherry Lake, Hetch
Hetchy, Upper and Lower Blue Lakes,
Lake Aloha, Silver Lake, Hell Hole
Reservoir, French Meadow Reservoir,
Lake Spaulding, Alpine Lake, Loon
Lake, and Ice House Reservoir. A
number of these occur at elevations
below the current range of the species,
indicating that the network of large
water and power projects found at lower
elevations does not overlap significantly
with the current accepted distribution of
the mountain yellow-legged frogs in the
Sierra Nevada (CDFW 2013, p. 3).
Kondolf et al. (1996, p. 1014) report
that dams can have direct effects to
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riparian habitat through permanent
removal of habitat to construct roads,
penstocks, powerhouses, canals, and
dams. Impacts of reservoirs include
flooding of riparian vegetation and
impediments to establishment of new
shoreline vegetation by fluctuating
water levels. Dams can alter the
temperature and sediment load of the
rivers they impound (Cole and Landres
1996, p. 175). Dams, water diversions,
and their associated structures can also
alter the natural flow regime with
unseasonal and fluctuating releases of
water (Kondolf et al. 1996, p. 1014). We
expect most such effects to occur in
stream systems below the extant range
of the mountain yellow-legged frogs,
although it is possible that stream
localities at the northern extent of the
range or at low elevations may be
affected (see also CDFW 2013, pp. 2–4).
The extent of past impacts to
mountain yellow-legged frog
populations from habitat loss or
modification due to reservoir projects
has not been quantified. CDFW (2013, p.
3) has noted that there are locations
where the habitat inundated as the
result of dam construction (for example,
Lake Aloha in the Desolation
Wilderness) may have been of higher
quality for mountain yellow-legged frogs
than the created impoundment.
Reservoirs can provide habitat for
introduced predators, including fish,
bullfrogs, and crayfish, and in some
cases, the past construction of reservoirs
has facilitated the spread of nonnative
fish (CDFW 2013, pp. 3, 4). In such
cases, reservoirs may function as
barriers to movement of mountain
yellow-legged frogs. However, CDFW
reported observing mountain yellowlegged frogs dispersing through fishless
reservoirs (CDFW 2013, p. 4). (For a
complete discussion of the impacts of
fish stocking see Habitat Modification
Due to Introduction of Trout to
Historically Fishless Areas above and
the discussion under Factor C.).
Most of the dams constructed within
the historic range of the mountain
yellow-legged frogs are small
streamflow-maintenance dams (CDFW
2013, p. 13) at the outflows of highelevation lakes. These small dams may
create additional habitat for the species
and can act as barriers to fish migration
from downstream tributaries into
fishless habitats, although they do not
impede frog movement (CDFW 2013, p.
3). CDFW staff (2013, p. 13) have
observed that extant frog populations
may have persisted where such dams
have helped to preserve a fishless
environment behind the dam.
Based on comments from CDFW and
others and the provision of additional
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information, we have reviewed the
analysis of dams and diversions that we
presented in the proposed rule. We find
that most large facilities are below the
current range of the mountain yellowlegged frogs and have revised our
finding. In the proposed rule, we stated
that dams and diversions presented a
moderate, prevalent threat to
persistence and recovery of the species.
In this final rule, we find that dams and
water diversions present a minor,
localized threat to persistence and
recovery of the species where structures
occur.
Livestock Use (Grazing)
The combined effect of legacy
conditions from historically excessive
grazing use and current livestock
grazing activities has the potential to
impact habitat in the range of the
mountain yellow-legged frog. The
following subsections discuss the effects
of excessive historical grazing, current
extent of grazing, and current grazing
management practices. As discussed
below, grazing has the potential to
reduce the suitability of habitat for
mountain yellow-legged frogs by
reducing its capability to sustain frogs
and facilitate dispersal and migration,
especially in stream areas.
Grazing of livestock in riparian areas
impacts the function of the aquatic
system in multiple ways, including soil
compaction, which increases runoff and
decreases water availability to plants;
vegetation removal, which promotes
increased soil temperatures and
evaporation rates at the soil surface; and
direct physical damage to the vegetation
(Kauffman and Krueger 1984, pp. 433–
434; Cole and Landres 1996, pp. 171–
172; Knapp and Matthews 1996, pp.
816–817). Streamside vegetation
protects and stabilizes streambanks by
binding soils to resist erosion and trap
sediment (Kauffman et al. 1983, p. 683;
Chaney et al. 1990, p. 2). Grazing within
mountain yellow-legged frog habitat has
been observed to remove vegetative
cover, potentially exposing frogs to
predation and increased desiccation
(Knapp 1993b, p. 1; Jennings 1996, p.
539), and to lead to erosion which may
silt in ponds and thereby reduce the
water depth needed for overwinter
survival (Knapp 1993b, p. 1). However,
an appropriately managed grazing
regime (including timing and intensity)
can enhance primary riparian vegetation
attributes that are strongly correlated to
stream channel and riparian soil
stability conditions necessary to
maintain a functioning riparian system
(George et al. 2011, p. 227). Although,
where highly degraded conditions such
as downcut channels exist, grazing
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management alone may not be sufficient
to restore former riparian conditions
(George et al. 2011, p. 227).
Aquatic habitat can also be degraded
by grazing. Mass erosion from trampling
and hoof slide causes streambank
collapse and an accelerated rate of soil
transport to streams (Meehan and Platts
1978, p. 274). Accelerated rates of
erosion lead to elevated instream
sediment loads and depositions, and
changes in stream-channel morphology
(Meehan and Platts 1978, pp. 275–276;
Kauffman and Krueger 1984, p. 432).
Livestock grazing may lead to
diminished perennial streamflows
(Armour et al. 1994, p. 10). Livestock
can increase nutrient-loading in water
bodies due to urination and defecation
in or near the water, and can cause
elevated bacteria levels in areas where
cattle are concentrated (Meehan and
Platts 1978, p. 276; Stephenson and
Street 1978, p. 156; Kauffman and
Krueger 1984, p. 432). With increased
grazing intensity, these adverse effects
to the aquatic ecosystem increase
proportionately (Meehan and Platts
1978, p. 275; Clary and Kinney 2000, p.
294).
Observational data indicate that
livestock can negatively impact
mountain yellow-legged frogs by
altering riparian habitat (Knapp 1993a,
p. 1; 1993b, p. 1; 1994, p. 3; Jennings
1996, p. 938; Carlson 2002, pers. comm.;
Knapp 2002a, p. 29). Livestock tend to
concentrate along streams and wet areas
where there is water and herbaceous
vegetation; grazing impacts are,
therefore, most pronounced in these
habitats (Meehan and Platts 1978, p.
274; U.S. Government Accounting
Office (GAO) 1988, pp. 10–11;
Fleischner 1994, p. 635; Menke et al.
1996, p. 17). This concentration of
livestock contributes to the
destabilization of streambanks, causing
undercuts and bank failures (Kauffman
et al. 1983, p. 684; Marlow and Pogacnik
1985, pp. 282–283; Knapp and
Matthews 1996, p. 816; Moyle 2002, p.
55). Grazing activity can contribute to
the downcutting of streambeds and
lower the water table. The degree of
erosion caused by livestock grazing can
vary with slope gradient, aspect, soil
condition, vegetation density, and
accessibility to livestock, with soil
disturbance greater in areas overused by
livestock (Meehan and Platts 1978, pp.
275–276; Kauffman et al. 1983, p. 685;
Kauffman and Krueger 1984, p. 432;
Bohn and Buckhouse 1985, p. 378; GAO
1988, p. 11; Armour et al. 1994, pp. 9–
11; Moyle 2002, p. 55).
Livestock grazing may impact other
wetland systems, including ponds that
can serve as mountain yellow-legged
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frog habitat. Grazing can modify
shoreline habitats by removing
overhanging banks that provide shelter,
and grazing contributes to the siltation
of breeding ponds. Bradford (1983, p.
1179) and Pope (1999, pp. 43–44) have
documented the importance of deep
lakes to overwinter survival of these
species. We expect that pond siltation
due to grazing may reduce the depth of
breeding ponds and cover underwater
crevices in some circumstances where
grazing is heavy and where soils are
highly erodable, thereby making the
ponds less suitable, or unsuitable, as
overwintering habitat for tadpoles and
adult mountain yellow-legged frogs.
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Effects of Excessive Historical Grazing
In general, historical livestock grazing
within the range of the mountain
yellow-legged frog was at a high
(although undocumented), unregulated
and unsustainable level until the
establishment of National Parks
(beginning in 1890) and National
Forests (beginning in 1905) (UC 1996a,
p. 114; Menke et al. 1996, p. 14).
Historical evidence indicates that heavy
livestock use in the Sierra Nevada has
resulted in widespread damage to
rangelands and riparian systems due to
sod destruction in meadows, vegetation
destruction, and gully erosion (see
review in Brown et al. 2009, pp. 56–58).
Within the newly established National
Parks, grazing by cattle and sheep was
eliminated, although grazing by
packstock, such as horses and mules,
continued. Within the National Forests,
the amount of livestock grazing was
gradually reduced, and the types of
animals shifted away from sheep and
toward cattle and packstock, with cattle
becoming the dominant livestock.
During World Wars I and II, increased
livestock use occurred on National
Forests in the west, causing overuse in
the periods 1914–1920 and 1939–1946.
Between 1950 and 1970 livestock
numbers were permanently reduced due
to allotment closures and uneconomical
operations, with increased emphasis on
resource protection and riparian
enhancement. Further reductions in
livestock use began again in the 1990s,
due in part to USFS reductions in
permitted livestock numbers, seasons of
use, implementation of rest-rotation
grazing systems, and to responses to
drought (Menke et al. 1996, pp. 7, 8).
Between 1981 and 1998, livestock
numbers on National Forests in the
Sierra Nevada decreased from 163,000
to approximately 97,000 head,
concurrent with Forest Service
implementation of standards and
guidelines for grazing and other
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resource management (USFS 2001, pp.
399–416).
Effects of Current Grazing
Yosemite, Sequoia, and Kings Canyon
National Parks remain closed to
livestock grazing. On USFSadministered lands that overlap the
historical ranges of the mountain
yellow-legged frog in the Sierra Nevada,
there are currently 161 active Rangeland
Management Unit Allotments for
livestock grazing. However, based on
frog surveys performed since 2005, only
27 of these allotments have extant
mountain yellow-legged frog
populations, while some allotments that
were located in sensitive areas have
been closed (USFS 2008, unpubl. data;
CDFW (CDFG) unpubl. data). As of
2009, USFS data indicated that grazing
occurs on about 65 percent of National
Forest lands within the range of the
mountain yellow-legged frog; that
livestock numbers remain greatly
reduced from historical levels; and that
numerous watershed restoration
projects have been implemented,
although grazing may still impact many
meadows above mid-elevation and
restoration efforts are far from complete
(Brown et al. 2009, pp. 56, 57).
However, Brown et al. (2009, p. 56)
report that livestock grazing is more
likely to occur in certain habitat types
used by mountain yellow-legged frogs
than others, indicating that populations
found in meadows, stream riparian
zones, and lakes in meadows are more
likely to encounter habitat effects of
grazing than populations found in the
deeper alpine lakes that the species
more likely inhabit (Brown et al. 2009,
p. 56).
USFS standards and guidelines in
forest land and resource management
plans have been implemented to protect
water quality, sensitive species,
vegetation, and stream morphology.
Further, USFS standards have been
implemented in remaining allotments to
protect aquatic habitats (see discussion
of the aquatic management strategy
under Factor D for examples). USFS
data from long-term meadow monitoring
collected from 1999 to 2006 indicate
that most meadows appear to be in an
intermediate quality condition class,
with seeming limited change in
condition class over the first 6 years of
monitoring. In addition, USFS grazing
standards and guidelines are based on
current science and are designed to
improve or maintain range ecological
conditions, and standards for managing
habitat for threatened, endangered, and
sensitive species have also been
incorporated (Brown et al. 2009, pp. 56–
58). The seasonal turn-out dates (dates
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at which livestock are permitted to
move onto USFS allotments) are set
yearly based on factors such as
elevation, annual precipitation, soil
moisture, and forage plant phenology,
and meadow readiness dates are also set
for montane meadows. However,
animals turned out to graze on lowelevation range (until higher elevation
meadows are ready) may reach upper
portions of allotments before the
meadows have reached range readiness
(Brown et al. 2009, p. 58).
Menke et al. (1996) have reported that
grazing livestock in numbers that are
consistent with grazing capacity and use
of sustainable methods led to better
range management in the Sierra Nevada
over the 20 years prior to development
of the report. They also noted that
moderate livestock grazing has the
potential to increase native species
diversity in wet and mesic meadows by
allowing native plant cover to increase
on site. Brown et al. (2009, p. 58) expect
proper livestock management, such as
proper timing, intensity, and duration,
to result in a trend towards increased
riparian species and a trend towards
restored wet and mesic meadows on
National Forests. To date, the scientific
and commercial information available to
us does not include descriptive or
cause-effect research that establishes a
causal link between habitat effects of
livestock grazing and mountain yellowlegged frog populations; however,
anecdotal information of specific habitat
effects suggests that, in specific
locations, the current grazing levels may
have population-level effects (see Knapp
1993b, p. 1; Brown et al. 2009, p. 56).
In addition, where low-elevation
populations occur in meadows,
additional conservation measures may
be required for recovery (USFS 2013, p.
5).
In summary, the legacy effects to
habitat from historical grazing levels,
such as increased erosion, stream
downcutting and headcutting, lowered
water tables, and increased siltation, are
a threat to mountain yellow-legged frogs
in those areas where such conditions
still occur and may need active
restoration. In the proposed rule, we
stated that grazing presented a minor
prevalent threat. Based on USFS and
public comments, we have reevaluated
our analysis of grazing to clarify effects
of past versus current grazing. We have
reworded the finding to more accurately
reflect the contribution of legacy effects
of past grazing levels to this threat
assessment, as follows: Current
livestock grazing activities may present
an ongoing, localized threat to
individual populations in locations
where the populations occur in stream
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riparian zones and in small waters
within meadow systems, where active
grazing co-occurs with extant frog
populations. Livestock grazing that
complies with forest standards and
guidelines is not expected to negatively
affect mountain yellow-legged frog
populations in most cases, although
limited exceptions could occur,
especially where extant habitat is
limited. In addition, mountain yellowlegged frogs may be negatively affected
where grazing standards are exceeded.
Rangewide, current livestock grazing is
not a substantial threat to the species.
Mining
Several types of mining activities
have occurred, or may currently occur,
on National Forests, including aggregate
mining (the extraction of materials from
streams or stream terraces for use in
construction), hardrock mining (the
extraction of minerals by drilling or
digging into solid rock), hydraulic
mining (a historical practice using
pressurized water to erode hillsides,
outlawed in 1884), placer mining
(mining in sand or gravel, or on the
surface, without resorting to
mechanically assisted means or
explosives), and suction-dredge mining
(the extraction of gold from riverine
materials, in which water, sediment,
and rocks are vacuumed from portions
of streams and rivers, sorted to obtain
gold, and the spoils redeposited in the
stream (see review in Brown et al. 2009,
pp. 62–64).
Aggregate mining can alter sediment
transport in streams, altering and
incising stream channels, and can cause
downstream deposition of sediment,
altering or eliminating habitat.
Aggregate mining typically occurs in
large riverine channels that are
downstream of much of the range of the
mountain yellow-legged frog complex
(see review in Brown et al. 2009, pp.
62–64). However, Brown et al. (2009,
pp. 62–64) note that effects of aggregate
mining may occur in some portions of
the Feather River system where such
operations occur within the historic
range of the Sierra Nevada yellowlegged frog, and potentially in localized
areas within the range of both species,
where the USFS maintains small
quarries for road work. They note that,
although effects of aggregate mining on
mountain yellow-legged frogs are
unstudied, impacts are probably slight.
Hardrock mining can be a source of
pollution where potentially toxic metals
are solubilized by waters that are
slightly acidic. Past mining activities
have resulted in the existence of many
shaft or tunnel mines on the forest in
the Sierra Nevada, although most are
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thought to occur below the range of the
species. Most operations that are
thought to have the potential to impact
the mountain yellow-legged frogs occur
in the lower elevation portions of the
Sierra Nevada yellow-legged frog range
on the Plumas National Forest and in
the ranges of both species on the Inyo
National Forest (see review in Brown et
al. 2009, pp. 62–64).
Hydraulic mining has exposed
previously concealed rocks that can
increase pollutants such as acid,
cadmium, mercury, and asbestos, and
its effect on water pollution may still be
apparent on the Feather River. However,
most of the area that was mined in this
way is below the elevation where Sierra
Nevada yellow-legged frogs are present,
so effects are likely highly localized (see
review in Brown et al. 2009, pp. 63, 64).
Although placer mining was dominant
historically, today it’s almost
exclusively recreational and is not
expected to have habitat-related effects.
Brown et al. (2009, p. 64) report that
suction-dredge mining is also primarily
recreational noting that, because nozzles
are currently restricted to 6 inches or
smaller, CDFW (CDFG, 1994) expects
disturbed areas to recover quickly
(although CDFW notes that such
dredging may increase suspended
sediments, change stream
geomorphology, and bury or suffocate
larvae). Suction dredge mining occurs
primarily in the foothills of the Sierra
Nevada, thus presenting a risk primarily
to mountain yellow-legged frog
populations at the lower elevations of
the species’ range. Suction dredging is
highly regulated by the CDFW, and in
the past, many streams have been
seasonally or permanently closed (see
review in Brown et al. 2009, p. 64).
Currently CDFW has imposed a
moratorium on suction dredging.
The high-elevation areas where most
Sierra Nevada yellow-legged frogs and
mountain yellow-legged frogs occur are
within designated wilderness, where
mechanical uses are prohibited by the
Wilderness Act. Designated wilderness
was withdrawn for new mining claims
on January 1, 1984, although a limited
number of active mines that predated
the withdrawal still occur within
wilderness (see Wilderness Act under
Factor D, below). Therefore, we expect
that mining activities may pose local
habitat-related impacts to the species at
specific localities where mining occurs.
Packstock Use
Similar to cattle, horses and mules
may significantly overgraze, trample, or
pollute riparian and aquatic habitat if
too many are concentrated in riparian
areas too often or for too long.
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Commercial packstock trips are
permitted in National Forests and
National Parks within the Sierra
Nevada, often providing transport
services into wilderness areas through
the use of horses or mules. Use of
packstock in the Sierra Nevada
increased after World War II as road
access, leisure time, and disposable
income increased (Menke et al. 1996, p.
919). Packstock grazing is the only
grazing currently permitted in the
National Parks of the Sierra Nevada.
Since the mid-1970s, National Forests
and National Parks have generally
implemented regulations to manage
visitor use and group sizes, including
measures to reduce packstock impacts
to vegetation and soils in order to
protect wilderness resources. For
example, Sequoia and Kings Canyon
National Parks have the backcountry
area with the longest history of research
and management of packstock impacts
(Hendee et al. 1990, p. 461). Hendee et
al. (1990, p. 461) report that the
extensive and long-term monitoring for
Sequoia, Kings Canyon, and Yosemite
National Parks makes it possible to
quantify impacts of packstock use,
showing that the vast majority of Sierra
Nevada yellow-legged frog and
mountain yellow-legged frog
populations in the Parks show no to
negligible impacts from packstock use
(National Park Service 2013, p. 3). In the
Sixty-Lakes Basin of Kings Canyon
National Park, packstock use is
regulated in wet meadows to protect
mountain yellow-legged frog breeding
habitat in bogs and along lake shores
from trampling and associated
degradation (Vredenburg 2002, p. 11;
Werner 2002, p. 2; National Park Service
2013, p. 3). Packstock use is also
regulated in designated wilderness in
National Forests within the Sierra
Nevada.
Packstock use is likely a threat of low
significance to mountain yellow-legged
frogs at the current time, except on a
limited, site-specific basis. As
California’s human population
increases, the impact of recreational
activities, including packstock use and
riding on the National Forests in the
Sierra Nevada, are projected to increase
(USDA 2001a, pp. 473–474). However,
on the Inyo National Forest, current
commercial packstock use is
approximately 27 percent of the level of
use in the 1980s reflecting a decline in
the public’s need and demand for
packstock trips. From 2001 to 2005,
commercial packstock outfitters within
the Golden Trout and South Sierra
Wilderness Areas averaged 28 percent of
their current authorized use (USFS
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2006, p. 3–18). Similarly, long-term
permitting data for administrative,
commercial, and recreational packstock
use in the three National Parks indicates
that packstock use is declining in the
Parks, providing no evidence to suggest
that packstock use will increase in the
future in the Parks (National Park
Service 2013, pp. 3, 4). Habitat changes
due to packstock grazing may pose a
risk to some remnant populations of
frogs and, in certain circumstances, a
hindrance to recovery of populations in
heavily used areas.
Roads and Timber Harvest
Activities that alter the terrestrial
environment (such as road construction
and timber harvest) may impact
amphibian populations in the Sierra
Nevada (Jennings 1996, p. 938) at
locations where these activities occur.
Historically, road construction and
timber harvest may have acted to reduce
the species’ range prior to the more
recent detailed studies and systematic
monitoring that have quantified and
documented species losses. Prior to the
formation of National Parks in 1890 and
National Forests in 1905, timber harvest
was widespread and unregulated, but
primarily took place at elevations on the
western slope of the Sierra Nevada
below the range of the mountain yellowlegged frog (University of California
(UC) 1996b, pp. 24–25). Between 1900
and 1950, the majority of timber harvest
occurred in old-growth forests on
private land (UC 1996b, p. 25). Between
1950 and the early 1990s, timber harvest
on National Forests increased, and the
majority of timber harvest-associated
impacts on mountain yellow-legged
frogs may therefore have taken place
during this period in lower elevation
locations where timber harvest and
species occurrences overlapped.
Currently, these activities are expected
to occur outside National Parks or
National Forest wilderness areas, with
limited exceptions.
Timber harvest activities (including
vegetation management and fuels
management) remove vegetation and
cause ground disturbance and
compaction, making the ground more
susceptible to erosion (Helms and
Tappeiner 1996, p. 446). This erosion
can increase siltation downstream and
potentially damage mountain yellowlegged frog breeding habitat. Timber
harvest may alter the annual hydrograph
(timing and volume of surface flows) in
areas where harvests occur. The
majority of erosion caused by timber
harvests is from logging roads (Helms
and Tappeiner 1996, p. 447). A recent
monitoring effort, which was conducted
by the USFS in stream habitats in the
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northern part of the Sierra Nevada
yellow-legged frog’s range, attempted to
assess the impact of vegetation
management activities, which would
include activities similar to timber
harvest, on mountain yellow-legged frog
populations (Foote et al. 2013, p. 2).
However, given the timing of project
implementation, the results were
limited to the impacts of these
management activities on mountain
yellow-legged frog habitat. The results
of the monitoring suggest these
activities did not significantly impact
perennial stream habitat for the
mountain yellow-legged frog, although
there were instances of habitat
degradation attributed to sedimentation
resulting from road decommissioning
and culvert replacement (Foote et al.
2013, p. 32).
Roadways have the potential to affect
riparian habitat by altering the physical
and chemical environment, including
alteration of surface-water run-off, with
potential changes to hydrology in highmountain lake and stream systems
(Brown et al. 2009, pp. 71–72). Roads,
including those associated with timber
harvests, have also been found to
contribute to habitat fragmentation and
limit amphibian movement, thus having
a negative effect on amphibian species
richness. Therefore, road construction
could fragment mountain yellow-legged
frog habitat if a road bisects habitat
consisting of water bodies in close
proximity. In the prairies and forests of
Minnesota, Lehtinen et al. (1999, pp. 8–
9) found that increased road density
reduced amphibian species richness.
DeMaynadier and Hunter (2000, p. 56)
found similar results in a study of eight
amphibian species in Maine, although
results varied with road type and width.
Results showed that anuran (true frogs,
the group of frogs that includes the
mountain yellow-legged frogs) habitat
use and movement were not affected
even by a wide, heavily used logging
road (deMaynadier and Hunter 2000, p.
56); this finding suggests that forest
roads may not fragment populations
where such roads occur.
Currently, most of the mountain
yellow-legged frog populations occur in
National Parks or designated wilderness
areas where timber is not harvested
(Bradford et al. 1994, p. 323; Drost and
Fellers 1996, p. 421; Knapp and
Matthews 2000, p. 430) and where
motorized access (and roads) does not
occur. Mountain yellow-legged frog
populations outside of these areas are
most often located above the timberline,
so timber harvest activity is not
expected to affect the majority of extant
mountain yellow-legged frog
populations. There is a higher potential
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overlap of timber harvest activities with
the species in the northern and lower
elevation portions of the species’ ranges
where the frogs occur in streams and
meadows in forested environments; in
these areas, populations are very small
and fragmented (Brown 2013,
unpaginated). Likewise, at lower
elevations of the Sierra Nevada, forest
roads and logging roads are more
common (Brown et al. 2009, p. 71).
Habitat effects associated with roads are
most likely to occur where existing
roadways occur (for example, see Knapp
1993b, unpaginated). Although
additional roads may be constructed
within the range of the mountain
yellow-legged frogs, we are not aware of
any proposals to build new roads at this
time.
In riparian areas, the USFS generally
maintains standards and guidelines for
land management activities, such as
timber harvests, that are designed to
maintain the hydrologic, geomorphic,
and ecologic processes that directly
affect streams, stream processes, and
aquatic habitats, and which can limit
potential effects of such activities (Foote
et al. 2013, pp. 4, 32). In general, we
expect the standards to be effective in
preventing habitat-related effects to
these species. Additionally, neither
timber harvests nor roads have been
implicated as important contributors to
the decline of this species (Jennings
1996, pp. 921–941), although habitat
alterations due to these activities may,
in site-specific, localized cases, have
population-level effects to mountain
yellow-legged frogs. We expect that
such cases would be more likely at
lower elevations or in the more northern
portion of the species’ range where
limited extant populations occur in
close proximity to timber harvest, or
where populations occur in drainages
adjacent to roadways. In the proposed
rule, we stated that roads and timber
harvest likely present minor prevalent
threats to the mountain yellow-legged
frogs factored across the range of the
species. We are clarifying that language,
noting that they may pose important
habitat-related effects to the species in
localized areas, but are not likely threats
across most of the species’ ranges.
Fire and Fire Management Activities
Mountain yellow-legged frogs are
generally found at high elevations in
wilderness areas and National Parks
where vegetation is sparse and where
fire may have historically played a
limited role in the ecosystem. However,
at lower elevations and in the northern
portion of the range, mountain yellowlegged frogs occur in stream or lake
environments within areas that are
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forested to various extents. In some
areas within the current range of the
mountain yellow-legged frog, long-term
fire suppression has changed the forest
structure and created conditions that
increase fire severity and intensity
(McKelvey et al. 1996, pp. 1934–1935).
Excessive erosion and siltation of
mountain yellow-legged frog habitats
following wildfire is a concern where
shallow, lower elevation aquatic areas
occur below forested stands. However,
prescribed fire has been used by land
managers to achieve various
silvicultural objectives, including fuel
load reduction. In some systems, fire is
thought to be important in maintaining
open aquatic and riparian habitats for
amphibians (Russell et al. 1999, p. 378),
although severe and intense wildfires
may reduce amphibian survival, as the
moist and permeable skin of amphibians
increases their susceptibility to heat and
desiccation (Russell et al. 1999, p. 374).
Amphibians may avoid direct mortality
from fire by retreating to wet habitats or
sheltering in subterranean burrows.
The effects of past fire and fire
management activities on historical
populations of mountain yellow-legged
frogs are not known. Neither the direct
nor indirect effects of prescribed fire or
wildfire on the mountain yellow-legged
frog have been studied. Hossack et al.
(2012, pp. 221, 226), in a study of the
effects of six stand-replacing fires on
three amphibians that breed in
temporary ponds in low-elevation dense
coniferous forests or in high-elevation
open, subalpine forests in Glacier
National Park, found that effects of
wildfire on amphibians may not be
evident for several years post-fire with
time-lagged declines. The decline in
populations was presumably due to the
proximity of high-severity fires to
important breeding habitats, which
resulted in low recruitment of juveniles
into the breeding population. They
cautioned, however, that amphibian
responses to fire are context specific and
cannot be generalized too broadly; they
found no change in occupancy after
wildfire at high elevations where
wetlands were in sparse forest or open
meadows where there was less change
in canopy cover and insolation after
wildfire. Where fire has occurred in the
steep canyons of southern California
where the southern DPS of the
mountain yellow-legged frog occurs, the
character of the habitat has been
significantly altered, leading to erosive
scouring and flooding of creeks after
surface vegetation is denuded (North
2012, pers. comm.). North (2012, pers.
comm.) reported that at least one
population of the federally endangered
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southern DPS of the mountain yellowlegged frog, which occurs in streams,
declined substantially after fire on the
East Fork City Creek (San Bernardino
Mountains) in 2003 and, by 2012, was
approaching extirpation. Although most
populations of mountain yellow-legged
frogs are in alpine habitat that differs
from the habitat in southern California,
when they occur in lower-elevation
stream habitats, they could be similarly
affected by large wildfires. When a large
fire does occur in occupied habitat,
mountain yellow-legged frogs can be
susceptible to both direct mortality
(leading to significantly reduced
population sizes) and indirect effects
(habitat alteration and reduced breeding
habitat). It is possible that fire has
caused localized extirpations in the
past. However, because these species
generally occupy high-elevation habitat,
we have determined that fire is not a
significant threat to the mountain
yellow-legged frog complex over much
of its current range, although where the
species occur at lower elevations or in
the most northerly portion of their
ranges, fire-related changes to habitat
may have population-level effects to the
species.
Recreation
Recreational activities that include
hiking, camping, and backpacking take
place throughout the Sierra Nevada,
whereas off-road vehicle (ORV) use
takes place in areas outside of
designated wilderness. These activities
can have significant negative impacts on
many plant and animal species and
their habitats (U.S. Department of
Agriculture (USDA) 2001a, pp. 483–
493). Extant populations of the
mountain yellow-legged frog complex
are primarily located at high elevations
in sub-alpine and alpine habitat within
designated wilderness. High-elevation
wilderness areas are ecosystems that are
subject to intense solar exposure;
extremes in temperatures, precipitation
levels, and wind; short growing seasons;
and shallow, nutrient-poor soil. Such
habitats are typically not resilient to
disturbance (Schoenherr 1992, p. 167;
Cole and Landres 1996, p. 170).
In easily accessible areas, heavy foot
traffic in riparian areas can trample
vegetation, compact soils, and
physically damage stream banks
(Kondolf et al. 1996, pp. 1014, 1019).
Human foot, horse, bicycle, or offhighway motor vehicle trails can replace
riparian habitat with compacted soil
(Kondolph et al. 1996, pp. 1014, 1017,
1019), lower the water table, and cause
increased erosion where such activities
occur. Bahls (1992, p. 190) reported that
the recreational activity of anglers at
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high mountain lakes can be locally
intense in western wilderness areas,
with most regions reporting a level of
use greater than the fragile lakeshore
environments can withstand. Heavy
recreation use has been associated with
changes in the basic ecology of lakes. In
the 1970s, Silverman and Erman (1979)
found that the most heavily used backcountry lakes in their study had less
nitrate and more iron and aquatic plants
than other lakes. These researchers
suggested that erosion at trails and
campsites, improper waste disposal,
destruction of vegetation, and campsites
might cause an increase in elements that
formerly limited plant growth (Hendee
et al. 1990, pp. 435, 436). The NPS
considers hiking and backpacking to be
a negligible risk for the mountain
yellow-legged frogs within the Parks,
noting that, while hiking and
backpacking occur adjacent to many
populations, evidence indicates that risk
to habitat is slight to none. For example,
monitoring of a high-use trail that
allows thousands of hikers annually to
come into close contact with several
populations of mountain yellow-legged
frogs, whose habitat is immediately
adjacent to the trail, shows that the
populations have grown substantially
over the last decade (NPS 2013, p. 6). In
one location where high hiking levels
may be having an impact due to access
via an adjacent road, Yosemite National
Park personnel have restricted access
(NPS 2013, p. 6). Although recreation
was noted in 1998 as the fastest growing
use of National Forests (USFS 2001a, p.
453), to our knowledge, no studies to
date have identified a correlation
between such recreation-related impacts
to habitat and effects to populations of
the mountain yellow-legged frog
complex.
Because of demand for wilderness
recreational experiences and concern
about wilderness resource conditions,
wilderness land management now
includes standards for wilderness
conditions, implementing permit
systems and group-size limits for
visitors and packstock, prohibitions on
camping and packstock use close to
water, and other visitor management
techniques to reduce impacts to habitat,
including riparian habitat (Cole 2001,
pp. 4–5). These wilderness land
management techniques are currently
being used in National Forest
Wilderness areas in the Sierra Nevada
and in backcountry areas of Yosemite,
Sequoia, and Kings Canyon National
Parks. In the proposed rule, we stated
that current recreation activities were
considered a threat of low significance
to the species’ habitat overall. Based on
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comments from the National Park
Service, the USFS, CDFW, and the
public, we have reevaluated the
previous analysis and have revised our
finding. Therefore, current habitat
effects of recreational activities are not
considered to have population-level
effects to mountain yellow-legged frogs
over much of their respective ranges,
although there may be localized effects
especially outside of backcountry areas
where use levels are not limited, or
where motorized use occurs in extant
frog habitat.
In summary, based on the best
available scientific and commercial
information, we consider the
modification of habitat and curtailment
of the species’ ranges to be a significant
and ongoing threat to the Sierra Nevada
yellow-legged frog and northern DPS of
the mountain yellow-legged frog.
Habitat fragmentation and degradation
(loss of habitat through competitive
exclusion) from stocking and the
continued presence of introduced trout
across the majority of the species’ range
is a threat of high prevalence. This
threat is a significant limiting factor to
persistence and recovery of the species
rangewide. Threats of low prevalence
(threats that may be important limiting
factors in some areas, but not across a
large part of the mountain yellow-legged
frog complex’s range) include dams and
water diversions, grazing, packstock
use, timber harvest and roads,
recreation, and fire management
activities.
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Factor B. Overutilization for
Commercial, Recreational, Scientific, or
Educational Purposes
No commercial market for mountain
yellow-legged frogs exists, nor any
documented recreational or educational
uses for these species. Scientific
research may cause stress to mountain
yellow-legged frogs through
disturbance, including disruption of the
species’ behavior, handling of
individual frogs, and injuries associated
with marking and tracking individuals.
However, this is a relatively minor
nuisance and not likely a negative
impact to the survival and reproduction
of individuals or the viability of the
populations.
Based on the best available scientific
and commercial information, we do not
consider overutilization for commercial,
recreational, scientific, or educational
purposes to be a threat to the mountain
yellow-legged frog complex now or in
the future.
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Factor C. Disease or Predation
Predation
Researchers have observed predation
of mountain yellow-legged frogs by the
mountain garter snake (Thamnophis
elegans elegans), Brewer’s blackbird
(Euphagus cyanocephalus), Clark’s
nutcracker (Nucifraga columbiana),
coyote (Canis latrans), and black bear
(Ursus americanus) (Mullally and
Cunningham 1956a, p. 193; Bradford
1991, pp. 176–177; Jennings et al. 1992,
p. 505; Feldman and Wilkinson 2000, p.
102; Vredenburg et al. 2005, p. 565).
However, none of these has been
implicated as a driver of population
dynamics, and we expect that such
predation events do not generally have
population-level impacts except where
so few individuals remain that such
predation is associated with loss of a
population (Bradford 1991, pp 174–177;
Jennings 1996, p. 938).
The American bullfrog (Lithobates
catesbeiana) is native to the United
States east of the Rocky Mountains, but
was introduced to California about a
century ago. The American bullfrog has
become common in California in most
permanent lakes and ponds below 1,829
m (6,000 ft) and is implicated in the
declines of a number of native frog
species (Jennings 1996, p. 931).
Mountain yellow-legged frogs are
thought to be particularly vulnerable to
bullfrogs and introduced crayfish,
potentially because the frogs did not
evolve with a predator (Jennings 1996,
p. 939). In addition, research indicates
that bullfrogs may outcompete other
species of amphibians where fish are
present because bullfrogs are both
unpalatable to fish and are naturally
vulnerable to invertebrate predators
such as dragonfly (Anisoptera) nymphs,
which fish preferentially consume.
Bullfrogs may co-occur with mountain
yellow-legged frogs at lower elevations.
On the Plumas National Forest, sites
created as a result of restoration
activities have been invaded by
bullfrogs (Brown et al. 2009, pp. 48, 49).
Bullfrogs also occur in the Lake Tahoe
Basin (USFS 2000, pp. 530, G–12) in the
vicinity of Fallen Leaf Lake. Bullfrog
predation and competition is expected
to have population-level effects where
bullfrog populations occupy the same
areas as extant mountain yellow-legged
frog populations.
The most prominent predator of
mountain yellow-legged frogs is
introduced trout, whose significance is
well-established because it has been
repeatedly observed that the frogs rarely
coexist with fish, and it is known that
introduced trout can and do prey on all
frog life stages except for eggs (Grinnell
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and Storer 1924, p. 664; Mullally and
Cunningham 1956a, p. 190; Cory 1962a,
p. 401; 1963, p. 172; Bradford 1989, pp.
775–778; Bradford and Gordon 1992, p.
65; Bradford et al. 1993, pp. 882–888;
1994a, p. 326; Drost and Fellers 1996, p.
422; Jennings 1996, p. 940; Knapp 1996,
p. 14; Knapp and Matthews 2000, p.
428; Knapp et al. 2001, p. 401;
Vredenburg 2004, p. 7649; Knapp 2013,
unpaginated). Knapp (1996, pp. 1–44)
estimated that 63 percent of lakes larger
than 1 ha (2.5 ac) in the Sierra Nevada
contain one or more nonnative trout
species, and that greater than 60 percent
of streams contain nonnative trout. In
some areas, trout-occupied waters
comprise greater than 90 percent of total
water body surface area (Knapp and
Matthews 2000, p. 434).
The multiple-year tadpole stage of the
mountain yellow-legged frog requires
submersion in the aquatic habitat yearround until metamorphosis. Moreover,
all life stages are highly aquatic,
increasing the frog’s susceptibility to
predation by trout (where they co-occur)
throughout its lifespan. Overwinter
mortality due to predation is especially
significant because, when water bodies
ice over in winter, adults and tadpoles
move from shallow margins of lakes and
ponds into deeper unfrozen water where
they are more vulnerable to predation;
fish encounters in such areas increase,
while refuge is less available.
The predation of mountain yellowlegged frogs by fishes observed in the
early 20th century by Grinnell and
Storer and the documented population
declines of the 1970s (Bradford 1991,
pp. 174–177; Bradford et al. 1994, pp.
323–327; Stebbins and Cohen 1995, pp.
226–227) were not the beginning of the
mountain yellow-legged frog’s decline,
but rather the continuation of a long
decline that started soon after fish
introductions to the Sierra Nevada
began in the mid-1800s (Knapp and
Matthews 2000, p. 436). Metapopulation
theory (Hanski 1997, pp. 85–86)
predicts this type of time lag from
habitat modification to population
extinction (Knapp and Matthews 2000,
p. 436). In 2004, Vredenburg (2004, p.
7647) concluded that introduced trout
are effective predators on mountain
yellow-legged frog tadpoles and
suggested that the introduction of trout
is the most likely reason for the decline
of the mountain yellow-legged frog
complex. This threat due to predation
by introduced trout is a significant,
prevalent (rangewide) risk to mountain
yellow-legged frogs, and it will persist
into the future in those locations where
fish are present. The effect of introduced
bullfrogs is expected to be a substantial
continuing threat in those locations
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where bullfrogs are known to occur
presently, but may present more of a
future threat if bullfrogs expand their
elevational range as a result of climate
change.
Disease
Over roughly the last 2 decades,
pathogens have been associated with
amphibian population declines, mass
die-offs, and even extinctions
worldwide (Bradford 1991, pp. 174–177;
Blaustein et al. 1994b, pp. 251–254;
Alford and Richards 1999, pp. 506;
Muths et al. 2003, p. 357; Weldon et al.
2004, p. 2100; Rachowicz et al. 2005, p.
1446; Fisher et al. 2009, p. 292). One
pathogen strongly associated with
dramatic declines on all continents that
harbor amphibians (all continents
except Antarctica) is the chytrid fungus,
Batrachochytrium dendrobatidis (Bd)
(Rachowicz et al. 2005, p. 1442). This
chytrid fungus has now been reported in
amphibian species worldwide (Fellers et
al. 2001, p. 945; Rachowicz et al. 2005,
p. 1442). Early doubt that this particular
pathogen was responsible for worldwide
die-offs has largely been overcome by
the weight of evidence documenting the
appearance, spread, and detrimental
effects to affected populations
(Vredenburg et al. 2010, p. 9689). The
correlation of notable recent amphibian
declines with reports of outbreaks of
fatal chytridiomycosis (the disease
caused by Bd) in montane areas has led
to a general association between high
altitude, cooler climates, and population
extirpations associated with Bd (Fisher
et al. 2009, p. 298).
Bd affects the mouth parts and
epidermal (skin) tissue of tadpoles and
metamorphosed frogs (Fellers et al.
2001, pp. 950–951). The fungus can
reproduce asexually, and can generally
withstand adverse conditions such as
freezing or drought (Briggs et al. 2002,
p. 38). It also may reproduce sexually,
leading to thick-walled sporangia that
would be capable of long-term survival
(for distant transport and persistence in
sites even after all susceptible host
animal populations are extirpated)
(Morgan et al. 2007, p. 13849). Adult
frogs can acquire this fungus from
tadpoles, and it can also be transmitted
between tadpoles (Rachowicz and
Vredenburg 2004, p. 80).
In California, chytridiomycosis has
been detected in many amphibian
species, including mountain yellowlegged frogs (Briggs et al. 2002, p. 38;
Knapp 2002b, p. 1). The earliest
documented case in the mountain
yellow-legged frog complex was in
1998, at Yosemite National Park (Fellers
et al. 2001, p. 945); however, more
recent literature shows Bd occurring in
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mountain yellow-legged frogs as early as
1975 (Ouellet et al. (2005, p. 1436;
Vredenberg et al. 2010, p. 9689). It is
unclear how Bd was originally
transmitted to the frogs (Briggs et al.
2002, p. 39). Visual examination of 43
tadpole specimens collected between
1955 and 1976 revealed no evidence of
Bd infection, yet 14 of 36 specimens
preserved between 1993 and 1999 did
have abnormalities attributable to Bd
(Fellers et al. 2001, p. 947). The earliest
recorded case of Bd in mountain yellowlegged frogs is from 1975, and Bd was
also identified on two adult Yosemite
toads among over 50 dead, dying, or
healthy Yosemite toads collected during
a die-off in 1976 (Green and Kagarise
Sherman 2001, p. 92), although it was
not thought to be the cause of the dieoff in the population. Given these
records, it is possible that this pathogen
has affected all three amphibian species
covered in this final rule since at least
the mid-1970s. Mountain yellow-legged
frogs may be especially vulnerable to Bd
infections because all life stages share
the same aquatic habitat nearly year
round, facilitating the transmission of
this fungus among individuals at
different life stages (Fellers et al. 2001,
p. 951).
During the epidemic phase of chytrid
infection into unexposed populations,
rapid die-offs of adult and subadult
lifestages are observed (Vredenburg et
al. 2010, p. 9691), with metamorphs
being extremely sensitive to Bd
infection (Kilpatrick et al. 2009, p. 113;
Vredenburg et al. 2010, p. 9691; see also
Vredenburg 2013, unpaginated). Field
and laboratory experiments indicate that
Bd infection is generally lethal to
mountain yellow-legged frogs (Knapp
2005b; Rachowicz 2005, pers. comm.),
and is likely responsible for declines in
sites that were occupied as recently as
2002, but where frogs were absent by
2005 (Knapp 2005b). Rachowicz et al.
(2006, p. 1671) monitored several
infected and uninfected populations in
Sequoia and Kings Canyon National
Parks over multiple years, documenting
dramatic declines and extirpations in
only the infected populations. Rapid
die-offs of mountain yellow-legged frogs
from chytridiomycosis have been
observed in more than 50 water bodies
in the southern Sierra Nevada in recent
years (Briggs et al. 2005, p. 3151).
Studies of the microscopic structure of
tissue and other evidence suggests Bd
caused many of the recent extinctions in
the Sierra National Forest’s John Muir
Wilderness Area and in Kings Canyon
National Park, where 41 percent of the
populations went extinct between 1995
and 2002 (Knapp 2002a, p. 10).
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In several areas where detailed
studies of the effects of Bd on the
mountain yellow-legged frog are
ongoing, substantial declines have been
observed following the course of the
disease infection and spread. Survey
results from 2000 in Yosemite and
Sequoia and Kings Canyon National
Parks indicated that 17 percent of frog
populations in Yosemite and 27 percent
of the mountain yellow-legged frog
populations sampled across both
Sequoia and Kings Canyon National
Parks showed evidence of Bd infection,
although the proportion of infected frogs
at each site varied greatly and disease
incidence varied within each Park
(Briggs et al. 2002, p. 40) (In the
proposed rule, these two figures were
averaged across all three parks; these
numbers reflect the text presented in
Briggs et al. 2002). In both 2003 and
2004, 19 percent of the populations that
were sampled in Sequoia and Kings
Canyon National Parks were infected
with Bd (Rachowicz 2005, pp. 2–3). By
2005, 91 percent of assayed populations
in Yosemite National Park showed
evidence of Bd infection (Knapp 2005b,
pp. 1–2), and the number of occupied
sites in Sequoia and Kings Canyon
National Parks had decreased by 47
percent from those known to be
occupied 3 to 8 years previously (Knapp
2005b, pers. comm). Currently, it is
believed that all populations in
Yosemite Park are infected with Bd
(Knapp et al. 2011, p. 9).
The effects of Bd on host populations
of the mountain yellow-legged frog are
variable, ranging from extirpation to
persistence with a low level of infection
(Briggs et al. 2002, pp. 40–41). When Bd
infection first occurs in a population,
the most common outcome is epidemic
spread of the disease and population
extirpation (Briggs et al. 2010, p. 9699).
Die-offs are characterized by rapid onset
of high-level Bd infections, followed by
death due to chytridiomycosis.
Although most populations that are
newly exposed to Bd are driven to
extirpation following the arrival of Bd,
some populations that experience Bdcaused population crashes are not
extirpated, and some may even recover
despite ongoing chytridiomycosis
(Briggs et al. 2010, pp. 9695–9696).
However, it is apparent that even at sites
exhibiting population persistence with
Bd, high mortality of metamorphosing
frogs persists, and this phenomenon
may explain the lower abundances
observed in such populations (Briggs et
al. 2010, p. 9699).
Vredenburg et al. (2010a, pp. 2–4)
studied frog populations before, during,
and after the infection and spread of Bd
in three study basins constituting 13, 33,
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and 42 frog populations, respectively,
then comprising the most intact
metapopulations remaining for these
species throughout their range. The
spread of Bd averaged 688 m/year (yr)
(2,257 ft/yr), reaching all areas of the
smaller basin in 1 year, and taking 3 to
5 years to completely infect the larger
basins, progressing like a wave across
the landscape. The researchers
documented die-offs following the
spread of Bd, with decreased population
growth rates evident within the first
year of infection. Basinwide,
metapopulations crashed from 1,680 to
22 individuals (northern DPS of the
mountain yellow-legged frog) in
Milestone Basin, with 9 of 13
populations extirpated; from 2,193 to 47
individuals (northern DPS of the
mountain yellow-legged frog) in Sixty
Lakes Basin, with 27 of 33 populations
extirpated; and from 5,588 to 436
individuals (Sierra Nevada yellowlegged frog) in Barrett Lakes Basin, with
33 of 42 populations extirpated. The
evidence is clear that Bd can and does
decimate newly infected frog
populations. Moreover, this rangewide
population threat is acting upon a
landscape already impacted by habitat
modification and degradation by
introduced fishes (see Factor A
discussion, above). As a result, remnant
populations in fishless lakes are now
affected by Bd.
Vredenburg et al. (2010a, p. 3)
projected that, at current extinction
rates, and given the disease dynamics of
Bd (infected tadpoles succumb to
chytridiomycosis at metamorphosis),
most if not all, extant populations
within the recently infected basins they
studied would go extinct within the
next 3 years. Available data (CDFW,
unpubl. data; Knapp 2005b; Rachowicz
2005, pers. comm.; Rachowicz et al.
2006, p. 1671) indicate that Bd is now
widespread throughout the Sierra
Nevada and, although it has not infected
all populations at this time, it is a
serious and substantial threat rangewide
to the mountain yellow-legged frog
complex.
Other diseases have also been
reported as adversely affecting
amphibian species, and these may be
present within the range of the
mountain yellow-legged frog. Bradford
(1991, pp. 174–177) reported an
outbreak of red-leg disease in Kings
Canyon National Park, and suggested
this was a result of overcrowding within
a mountain yellow-legged frog
population. Red-leg disease is caused by
the bacterial pathogen Aeromonas
hydrophila, along with other pathogens.
Red-leg disease is opportunistic and
successfully attacks immune-suppressed
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individuals, and this pathogen appears
to be highly contagious, affecting the
epidermis and digestive tract of
otherwise healthy amphibians (Shotts
1984, pp. 51–52; Carey 1993, p. 358;
Carey and Bryant 1995, pp. 14–15).
Although it has been correlated with
decline of a frog population in at least
one case, red-leg disease is not thought
to be a significant contributor to
observed frog population declines
rangewide, based on the available
literature.
Saprolegnia is a globally distributed
fungus that commonly attacks all life
stages of fishes (especially hatcheryreared fishes), and has recently been
documented to attack and kill egg
masses of western toads (Bufo boreas)
(Blaustein et al. 1994b, p. 252). This
pathogen may be introduced through
fish stocking, or it may already be
established in the aquatic ecosystem.
Fishes and migrating or dispersing
amphibians may be vectors for this
fungus (Blaustein et al. 1994b, p. 253;
Kiesecker et al. 2001, p. 1068).
Saprolegnia has been reported in the
southern DPS of the mountain yellowlegged frog (North 2012, pers. comm.);
however, its occurrence within the
Sierran range of the mountain yellowlegged frog complex and associated
influence on population dynamics (if
any) are unknown.
Other pathogens of concern for
amphibian species include ranaviruses
(Family Iridoviridae). Mao et al. (1999,
pp. 49–50) isolated identical
iridoviruses from co-occurring
populations of the threespine
stickleback (Gasterosteus aculeatus) and
the red-legged frog (Rana aurora),
indicating that infection by a given virus
is not limited to a single species, and
that iridoviruses can infect animals of
different taxonomic classes. This
suggests that virus-hosting trout
introduced into mountain yellow-legged
frog habitat may be a vector for
amphibian viruses. However, definitive
mechanisms for the transmission to the
mountain yellow-legged frog remain
unknown. No viruses were detected in
the mountain yellow-legged frogs that
Fellers et al. (2001, p. 950) analyzed for
Bd. In Kings Canyon National Park,
Knapp (2002a, p. 20) found mountain
yellow-legged frogs showing symptoms
attributed to a ranavirus (Knapp 2013,
unpaginated). To date, ranaviruses
remain a concern for the mountain
yellow-legged frog complex, but the
available information does not indicate
they are negatively affecting
populations.
It is unknown whether amphibian
pathogens in the high Sierra Nevada
have always coexisted with amphibian
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populations or if the presence of such
pathogens is a recent phenomenon.
However, it has been suggested that the
susceptibility of amphibians to
pathogens may have recently increased
in response to anthropogenic
environmental disruption (Carey 1993,
pp. 355–360; Blaustein et al. 1994b, p.
253; Carey et al. 1999, p. 7). This
hypothesis suggests that environmental
changes may be indirectly responsible
for certain amphibian die-offs due to
immune system suppression of tadpoles
or post-metamorphic amphibians (Carey
1993, p. 358; Blaustein et al. 1994b, p.
253; Carey et al. 1999, pp. 7–8).
Pathogens such as Aeromonas
hydrophila, which are present in fresh
water and in healthy organisms, may
become more of a threat, potentially
causing localized amphibian population
die-offs when the immune systems of
individuals within the host population
are suppressed (Carey 1993, p. 358;
Carey and Bryant 1995, p. 14).
The contribution of Bd as an
environmental stressor and limiting
factor on mountain yellow-legged frog
population dynamics is currently
extremely high, and it poses a
significant current and continuing threat
to remnant uninfected populations in
the southern Sierra Nevada. Its effects
are most dramatic following the
epidemic stage as it spreads across
newly infected habitats; massive die-off
events follow the spread of the fungus,
and it is likely that survival of mountain
yellow-legged frogs through the
metamorphosis stage is substantially
reduced even years after the initial
epidemic (Rachowicz et al. 2006, pp.
1679–1680). The relative impact from
other diseases and the interaction of
other stressors and disease on the
immune systems of mountain yellowlegged frogs remains poorly documented
to date.
In summary, based on the best
available scientific and commercial
information, we consider the threats of
predation and disease to be significant,
ongoing threats to the Sierra Nevada
yellow-legged frog and the northern DPS
of the mountain yellow-legged frog.
These threats include predation by
bullfrogs and introduced fishes, and
amphibian pathogens (most specifically,
the chytrid fungus), two primary driving
forces leading to population declines in
the mountain yellow-legged frog
complex. These are highly prevalent
threats, and they are predominant
limiting factors hindering population
viability and precluding recovery across
the ranges of the mountain yellowlegged frog complex.
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Factor D. The Inadequacy of Existing
Regulatory Mechanisms
In determining whether the
inadequacy of regulatory mechanisms
constitutes a threat to the mountain
yellow-legged frog complex, we
analyzed the existing Federal and State
laws and regulations that may address
the threats to these species or contain
relevant protective measures. Regulatory
mechanisms are typically
nondiscretionary and enforceable, and
may preclude the need for listing if such
mechanisms are judged to adequately
address the threat(s) to the species such
that listing is not warranted. Conversely,
threats on the landscape are not
ameliorated where existing regulatory
mechanisms are not adequate (or when
existing mechanisms are not adequately
implemented or enforced).
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Federal Wilderness Act
The Wilderness Act of 1964 (16 U.S.C.
1131 et seq.) established a National
Wilderness Preservation System made
up of federally owned areas designated
by Congress as ‘‘wilderness’’ for the
purpose of preserving and protecting
designated areas in their natural
condition. The Wilderness Act states the
use of these areas with limited
exception are subject to the following
restrictions: (1) New or temporary roads
cannot be built; (2) motor vehicles,
motorized equipment, or motorboats
cannot be used; (3) aircraft cannot land;
(4) no form of mechanical transport can
occur; and (5) no structure or
installation may be built. In addition, a
special provision within the Wilderness
Act stipulated that, except for valid
existing rights, effective January 1, 1984,
the minerals within designated
wilderness areas would be withdrawn
from all forms of appropriation under
mining laws, precluding new mining
claims within designated wilderness
after that date (see Hendee et al. 1990,
p. 508). A large number of mountain
yellow-legged frog locations occur
within wilderness areas managed by the
USFS and NPS and, therefore, are
afforded protection from direct loss or
degradation of habitat by some human
activities (such as development,
commercial timber harvest, road
construction, and some fire management
actions). Livestock grazing and fish
stocking both occur within designated
wilderness areas on lands within the
National Forest System.
National Forest Management Act of
1976
Under the National Forest
Management Act of 1976, as amended
(NFMA) (16 U.S.C. 1600 et seq.), the
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USFS is tasked with managing National
Forest lands based on multiple-use,
sustained-yield principles, and with
implementing land and resource
management plans (LRMP) on each
National Forest to provide for a
diversity of plant and animal
communities. The purpose of an LRMP
is to guide and set standards for all
natural resource management activities
for the life of the plan (10 to 15 years).
NFMA requires the USFS to incorporate
standards and guidelines into LRMPs.
The 1982 planning regulations for
implementing NFMA (47 FR 43026;
September 30, 1982), under which all
existing forest plans in the Sierra
Nevada were prepared until recently,
guided management of National Forests
and required that fish and wildlife
habitat on National Forest system lands
be managed to maintain viable
populations of existing native and
desired nonnative vertebrate species in
the planning area. A viable population
is defined as a population of a species
that continues to persist over the long
term with sufficient distribution to be
resilient and adaptable to stressors and
likely future environments. In order to
insure that viable populations would be
maintained, the 1982 planning
regulations directed that habitat must be
provided to support, at least, a
minimum number of reproductive
individuals and that habitat must be
well-distributed so that those
individuals could interact with others in
the planning area.
On April 9, 2012, the USFS published
a final rule (77 FR 21162) amending 36
CFR 219 to adopt new National Forest
System land management regulations
that guide the development,
amendment, and revision of LRMPs for
all Forest System lands. These revised
regulations, which became effective on
May 9, 2012, replaced the 1982
planning rule. The 2012 planning rule
requires that the USFS maintain viable
populations of species of conservation
concern at the discretion of regional
foresters. This rule could thereby result
in removal of the limited protections
that are currently in place for mountain
yellow-legged frogs under the Sierra
Nevada Forest Plan Amendment
(SNFPA), as described below.
Sierra Nevada Forest Plan Amendment
In 2001, a record of decision was
signed by the USFS for the Sierra
Nevada Forest Plan Amendment
(SNFPA), based on the final
environmental impact statement for the
SNFPA effort and prepared under the
1982 NFMA planning regulations. The
Record of Decision amends the USFS
Pacific Southwest Regional Guide, the
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Intermountain Regional Guide, and the
LRMPs for National Forests in the Sierra
Nevada and Modoc Plateau. This
document affects land management on
all National Forests throughout the
range of the mountain yellow-legged
frog complex. The SNFPA addresses
and gives management direction on
issues pertaining to old forest
ecosystems; aquatic, riparian, and
meadow ecosystems; fire and fuels;
noxious weeds; and lower west-side
hardwood ecosystems of the Sierra
Nevada. In January 2004, the USFS
amended the SNFPA, based on the final
supplemental environmental impact
statement, following a review of fire and
fuels treatments, compatibility with the
National Fire Plan, compatibility with
the Herger-Feinstein Quincy Library
Group Forest Recovery Pilot Project, and
effects of the SNFPA on grazing,
recreation, and local communities
(USDA 2004, pp. 26–30).
Relevant to the mountain yellowlegged frog complex, the Record of
Decision for SNFPA aims to protect and
restore aquatic, riparian, and meadow
ecosystems, and to provide for the
viability of associated native species
through implementation of an aquatic
management strategy. The aquatic
management strategy is a general
framework with broad policy direction.
Implementation of this strategy was
intended to take place at the landscape
and project levels. Nine goals are
associated with the aquatic management
strategy:
(1) The maintenance and restoration
of water quality to comply with the
Clean Water Act (CWA) and the Safe
Drinking Water Act;
(2) The maintenance and restoration
of habitat to support viable populations
of native and desired nonnative
riparian-dependent species, and to
reduce negative impacts of nonnative
species on native populations;
(3) The maintenance and restoration
of species diversity in riparian areas,
wetlands, and meadows to provide
desired habitats and ecological
functions;
(4) The maintenance and restoration
of the distribution and function of biotic
communities and biological diversity in
special aquatic habitats (such as springs,
seeps, vernal pools, fens, bogs, and
marshes);
(5) The maintenance and restoration
of spatial and temporal connectivity for
aquatic and riparian species within and
between watersheds to provide
physically, chemically, and biologically
unobstructed movement for their
survival, migration, and reproduction;
(6) The maintenance and restoration
of hydrologic connectivity between
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floodplains, channels, and water tables
to distribute flood flows and to sustain
diverse habitats;
(7) The maintenance and restoration
of watershed conditions as measured by
favorable infiltration characteristics of
soils and diverse vegetation cover to
absorb and filter precipitation, and to
sustain favorable conditions of
streamflows;
(8) The maintenance and restoration
of instream flows sufficient to sustain
desired conditions of riparian, aquatic,
wetland, and meadow habitats, and to
keep sediment regimes within the
natural range of variability; and
(9) The maintenance and restoration
of the physical structure and condition
of streambanks and shorelines to
minimize erosion and sustain desired
habitat diversity.
If these goals of the aquatic
management strategy are pursued and
met, threats to the mountain yellowlegged frog complex resulting from
habitat alterations could be reduced.
However, the aquatic management
strategy is a generalized approach that
does not contain specific
implementation timeframes or
objectives, and it does not provide
direct protections for the mountain
yellow-legged frog. Additionally, as
described above, the April 9, 2012, final
rule (77 FR 21162) that amended 36 CFR
219 to adopt new National Forest
System land management planning
regulations could result in removal of
the limited protections that are
currently in place for mountain yellowlegged frogs under the SNFPA.
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National Park Service Organic Act
The statute establishing the National
Park Service, commonly referred to as
the National Park Service Organic Act
(39 Stat. 535; 16 U.S.C. 1, 2, 3, and 4),
states that the NPS will administer areas
under their jurisdiction ‘‘. . . by such
means and measures as conform to the
fundamental purpose of said parks,
monuments, and reservations, which
purpose 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.’’ Park managers must take
action to ensure that ongoing NPS
activities do not cause impairment. In
cases of doubt as to the impact of
activities on park natural resource, the
Park Service is to decide in favor of
protecting the natural resources.
Sequoia, Kings Canyon, and Yosemite
National Parks began phasing out fish
stocking by the State in 1969 and
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terminated this practice entirely in 1991
(Knapp 1996, p. 9).
Federal Power Act
The Federal Power Act of 1920, as
amended (FPA) (16 U.S.C. 791 et seq.)
was enacted to regulate non-federal
hydroelectric projects to support the
development of rivers for energy
generation and other beneficial uses.
The FPA provides for cooperation
between the Federal Energy Regulatory
Commission (Commission) and other
Federal agencies in licensing and
relicensing power projects. The FPA
mandates that each license includes
conditions to protect, mitigate, and
enhance fish and wildlife and their
habitat affected by the project. However,
the FPA also requires that the
Commission give equal consideration to
competing priorities, such as power and
development, energy conservation,
protection of recreational opportunities,
and preservation of other aspects of
environmental quality. Further, the FPA
does not mandate protections of habitat
or enhancements for fish and wildlife
species, but provides a mechanism for
resource agency recommendations that
are incorporated into a license at the
discretion of the Commission.
Additionally, the FPA provides for the
issuance of a license for the duration of
up to 50 years, and the FPA contains no
provision for modification of the project
for the benefit of species, such as
mountain yellow-legged frogs, before a
current license expires.
Although most reservoirs and water
diversions are located at lower
elevations than those at which extant
mountain yellow-legged frog
populations occur, numerous extant
populations occur within watersheds
that feed into developed and managed
aquatic systems (such as reservoirs and
water diversions) operated for the
purpose of power generation and
regulated by the FPA and may be
considered during project relicensing.
State
California Endangered Species Act
This section has been updated from
the information presented in the
proposed rule, and discussion of
CDFW’s current fish-stocking practices
has been moved to the Factor A
discussion of Habitat Modification Due
to Introduction of Trout to Historically
Fishless Areas.
The California Endangered Species
Act (CESA) (California Fish and Game
Code, section 2080 et seq.) prohibits the
unauthorized take of State-listed
endangered or threatened species. CESA
requires State agencies to consult with
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CDFW on activities that may affect a
State-listed species, and mitigate for any
adverse impacts to the species or its
habitat. Pursuant to CESA, it is unlawful
to import or export, take, possess,
purchase, or sell any species or part or
product of any species listed as
endangered or threatened. The State
may authorize permits for scientific,
educational, or management purposes,
and allow take that is incidental to
otherwise lawful activities. On April 1,
2013, the Sierra Nevada yellow-legged
frog was listed as a threatened species
and the mountain yellow-legged frog
(Statewide) was listed as an endangered
species under CESA (CDFW 2013, p. 1).
While the listing of the Sierra Nevada
yellow-legged frog and the mountain
yellow-legged frog under CESA provide
some protections to these species, as
State regulation prohibits the
unauthorized take of State-listed
species, the definition of take under
CESA does not include habitat
modification or degradation.
Additionally, the majority of the lands
occupied by these species are federally
managed lands, so there is limited
jurisdiction in which to regulate land
management activities that may affect
these species.
Overall, existing Federal and State
laws and regulatory mechanisms
currently offer some level of protection
for the mountain yellow-legged frog
complex. While not the intent of the
Wilderness Act, the mountain yellowlegged frogs receive ancillary protection
from the Wilderness Act due to its
prohibitions on development, road
construction, and timber harvest, and
associated standards and guidelines that
limit visitor and packstock group sizes
and use. With the exception of the
National Park Service Organic Act, the
existing regulatory mechanisms have
not been effective in reducing threats to
mountain yellow-legged frogs and their
habitat from fish stocking and the
continuing presence of nonnative fish.
Nor have these mechanisms been
effective in protecting populations from
infection by diseases, although Forest
Service standards and guidelines have
likely reduced threats associated with
grazing, timber harvest, and recreation
use. Although State regulations under
CESA provide some protection against
take of the mountain yellow-legged
frogs, the definition of take under CESA
does not include habitat modification or
degradation.
Factor E. Other Natural or Manmade
Factors Affecting Its Continued
Existence
The mountain yellow-legged frog is
sensitive to environmental change or
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degradation because it has an aquatic
and terrestrial life history and highly
permeable skin that increases exposure
of individuals to substances in the
water, air, and terrestrial substrates
(Blaustein and Wake 1990, p. 203;
Bradford and Gordon 1992. p. 9;
Blaustein and Wake 1995, p. 52;
Stebbins and Cohen 1995, pp. 227–228).
Several natural or anthropogenically
influenced changes, including
contaminant deposition, acid
precipitation, increases in ambient
ultraviolet radiation, and climate
change, have been implicated as
contributing to amphibian declines
(Corn 1994, pp. 62–63; Alford and
Richards 1999, pp. 2–7). There are also
documented incidences of direct
mortality of, or the potential for direct
disturbance to, individuals from some
activities already discussed; in severe
instances, these actions may have
population-level consequences. As
presented in the proposed rule (78 FR
24472, April 25, 2013), contaminants,
acid precipitation, and ambient
ultraviolet radiation are not known to
pose a threat (current or historical) to
the mountain yellow-legged frog and,
therefore, are not discussed further.
Please refer to the proposed listing rule
for the Sierra Nevada yellow-legged
frog, the northern DPS of the mountain
yellow-legged frog, and the Yosemite
toad (78 FR 24472, April 25, 2013) for
a detailed discussion of contaminants,
acid precipitation, and ambient
ultraviolet radiation.
Climate Change
Our analysis under the Act includes
consideration of ongoing and projected
changes in climate. The terms ‘‘climate’’
and ‘‘climate change’’ are defined by the
Intergovernmental Panel on Climate
Change (IPCC). The term ‘‘climate’’
refers to the mean and variability of
different types of weather conditions
over time, with 30 years being a typical
period for such measurements, although
shorter or longer periods also may be
used (IPCC 2007a, p. 1450; IPCC 2013a,
Annex III). The term ‘‘climate change’’
thus refers to a change in the mean or
variability of one or more measures of
climate (for example, temperature or
precipitation) that persists for an
extended period, typically decades or
longer, whether the change is due to
natural variability, human activity, or
both (IPCC 2007a, p. 1450; IPCC 2013a,
Annex III). A recent compilation of
climate change and its effects is
available from reports of the
Intergovernmental Panel on Climate
Change (IPCC) (IPCC 2013b, entire).
Global climate projections are
informative and, in some cases, the only
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or the best scientific information
available for us to use. However,
projected changes in climate and related
impacts can vary substantially across
and within different regions of the
world (for example, IPCC 2007a, pp. 8–
12). Therefore, we use downscaled
projections when they are available and
have been developed through
appropriate scientific procedures,
because such projections provide higher
resolution information that is more
relevant to the spatial scales used for
analyses of a given species (see Glick et
al. 2011, pp. 58–61, for a discussion of
downscaling). With regard to our
analysis for the Sierra Nevada of
California (and western United States),
downscaled projections are available,
yet even downscaled climate models
contain some uncertainty.
Variability exists in outputs from
different climate models, and
uncertainty regarding future GHG
emissions is also a factor in modeling
(PRBO 2011, p. 3). A general pattern
that holds for many predictive models
indicates northern areas of the United
States will become wetter, and southern
areas (particularly the Southwest) will
become drier. These models also predict
that extreme events, such as heavier
storms, heat waves, and regional
droughts, may become more frequent
(Glick et al. 2011, p. 7). Moreover, it is
generally expected that the duration and
intensity of droughts will increase in the
future (Glick et al. 2011, p. 45; PRBO
2011, p. 21).
The last century has included some of
the most variable climate reversals
documented, at both the annual and
near-decadal scales, including a high
˜
frequency of El Nino (associated with
˜
more severe winters) and La Nina
(associated with milder winters) events
(reflecting drought periods of 5 to 8
years alternating with wet periods)
(USDA 2001b, p. 33). Scientists have
confirmed a longer duration climate
cycle termed the Pacific Decadal
Oscillation (PDO), which operates on
cycles between 2 to 3 decades, and
generally is characterized by warm and
dry (PDO positive) followed by cool and
wet cycles (PDO negative) (Mantua et al.
1997, pp. 1069–1079; Zhang et al. 1997,
pp. 1004–1018). Snowpack is seen to
follow this pattern—heavier in the PDO
negative phase in California, and lighter
in the positive phase (Mantua et al.
1997, p. 14; Cayan et al. 1998, p. 3148;
McCabe and Dettinger 2002, p. 24).
For the Sierra Nevada ecoregion,
climate models predict that mean
annual temperatures will increase by 1.8
to 2.4 °C (3.2 to 4.3 °F) by 2070,
including warmer winters with earlier
spring snowmelt and higher summer
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temperatures. However, it is expected
that temperature and climate variability
will vary based on topographic diversity
(for example, wind intensity will
determine east versus west slope
variability) (PRBO 2011, p. 18). Mean
annual rainfall is projected to decrease
from 9.2–33.9 cm (3.6–13.3 in) by 2070;
however, projections have high
uncertainty and one study predicts the
opposite effect (PRBO 2011, p. 18).
Given the varied outputs from differing
modeling assumptions, and the
influence of complex topography on
microclimate patterns, it is difficult to
draw general conclusions about the
effects of climate change on
precipitation patterns in the Sierra
Nevada (PRBO 2011, p. 18). Snowpack
is, by all projections, going to decrease
dramatically (following the temperature
rise and more precipitation falling as
rain) (Kadir et al. 2013, pp. 76–80).
Higher winter streamflows, earlier
runoff, and reduced spring and summer
streamflows are projected, with
increasing severity in the southern
Sierra Nevada (PRBO 2011, pp. 20–22);
(Kadir et al. 2013, pp. 71–75).
Snow-dominated elevations of 2,000–
2,800 m (6,560–9,190 ft) will be the
most sensitive to temperature increases,
and a warming of 5 °C (9 °F) is projected
to shift center timing (the measure when
half a stream’s annual flow has passed
a given point in time) to more than 45
days earlier in the year as compared to
the 1961–1990 baseline (PRBO 2011, p.
23). Lakes, ponds, and other standing
waters fed by snowmelt or streams are
likely to dry out or be more ephemeral
during the non-winter months (Lacan et
al. 2008, pp. 216–222; PRBO 2011, p.
24). This pattern could influence ground
water transport, and springs may be
similarly depleted, leading to lower lake
levels.
Blaustein et al. (2010, pp. 285–300)
provide an exhaustive review of
potential direct and indirect and
habitat-related effects of climate change
to amphibian species, with
documentation of effects in a number of
species where such effects have been
studied. Altitudinal range shifts with
changes in climate have been reported
in some regions. They note that
temperature can influence the
concentration of dissolved oxygen in
aquatic habitats, with warmer water
generally having lower concentrations
of dissolved oxygen, and that water
balance heavily influences amphibian
physiology and behavior. They predict
that projected changes in temperature
and precipitation are likely to increase
habitat loss and alteration for those
species living in sensitive habitats, such
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as ephemeral ponds and alpine habitats
(Blaustein et al. 2010, pp. 285–287).
Because environmental cues such as
temperature and precipitation are
clearly linked to onset of reproduction
in many species, climate change will
likely affect the timing of reproduction
in many species, potentially with
different sexes responding differently to
climate change. For example, males of
two newt species (Triturus spp.) showed
a greater degree of change in arrival date
at breeding ponds (Blaustein et al. 2010,
p. 288). Lower concentrations of
dissolved oxygen in aquatic habitats
may negatively affect developing
embryos and larvae, in part because
increases in temperature increase the
oxygen consumption rate in
amphibians. Reduced oxygen
concentrations have also been shown to
result in accelerated hatching in ranid
frogs, but at a smaller size, while larval
development and behavior may also be
affected and may be mediated by larval
density and food availability (Blaustein
et al. 2010, pp. 288–289).
Increased temperatures can reduce
time to metamorphosis, which can
increase chances of survival where
ponds dry, but also result in
metamorphosis at a smaller size,
suggesting a likely trade-off between
development and growth, which may be
exacerbated by climate change and have
fitness consequences for adults
(Blaustein et al. 2010, pp. 289–290).
Changes in terrestrial habitat, such as
changed soil moisture and vegetation,
can also directly affect adult and
juvenile amphibians, especially those
adapted to moist forest floors and cool,
highly oxygenated water that
characterizes montane regions. Climate
change may also interact with other
stressors that may be acting on a
particular species, such as disease and
contaminants (Blaustein et al. 2010, pp.
290–299).
A recent paper (Kadir et al. 2013,
entire) provides specific information on
the effects of climate change in the
Sierra Nevada. The report found that
glaciers in the Sierra Nevada have
decreased in area over the past century,
and glacier shrinkage results in earlier
peak water runoff and drier summer
conditions. Another result from the
report is that the lower edge of the
conifer-dominated forests in the Sierra
Nevada has been retreating upslope over
the past 60 years. Regarding wildfire,
since 1950, annual acreage burned in
wildfires statewide has been increasing
in California, and in the western United
States, large wildfires have become
more frequent, increasing in tandem
with rising spring and summer
temperatures. Finally, the report found
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that today’s subalpine forests in the
Sierra Nevada are much denser—that is,
comprise more small-diameter trees—
than they were over 70 years ago.
During this time period, warmer
temperatures, earlier snowmelt, and
more rain than snow occurred in this
region. Many of these changes in the
Sierra Nevada of California due to
climate are likely to influence mountain
yellow-legged frogs because both
mountain yellow-legged frog species in
the Sierra Nevada are highly vulnerable
to climate change because changing
hydrology and habitat in the Sierra
Nevada will likely have impacts on
remaining populations (Viers et al.
2013, pp. 55, 56).
Vulnerability of species to climate
change is a function of three factors:
Sensitivity of a species or its habitat to
climate change, exposure of individuals
to such physical changes in the
environment, and their capacity to
adapt to those changes (Glick et al.
2011, pp. 19–22). Critical sensitivity
elements broadly applicable across
organizational levels (from species
through habitats to ecosystems) are
associated with physical variables, such
as hydrology (timing, magnitude, and
volume of waterflows), fire regime
(frequency, extent, and severity of fires),
and wind (Glick et al. 2011, pp. 39–40).
Species-level sensitivities generally
include physiological factors, such as
changes in temperature, moisture, or pH
as they influence individuals; these also
include dependence on sensitive
habitats, ecological linkages to other
species, and changes in phenology
(timing of key life-history events) (Glick
et al. 2011, pp. 40–41).
Exposure to environmental stressors
renders species vulnerable to climate
change impacts, either through direct
mechanisms (for example, physical
temperature extremes or changes in
solar radiation), or indirectly through
impacts upon habitat (hydrology; fire
regime; or abundance and distribution
of prey, competitors, or predator
species). A species’ capacity to adapt to
climate change is increased by
behavioral plasticity (the ability to
modify behavior to mitigate the impacts
of the stressor), dispersal ability (the
ability to relocate to meet shifting
conditions), and evolutionary potential
(for example, shorter lived species with
multiple generations have more capacity
to adapt through evolution) (Glick et al.
2011, pp. 48–49).
The International Union for
Conservation of Nature describes five
categories of life-history traits that
render species more vulnerable to
climate change (Foden et al. 2008 in
Glick et al. 2011, p. 33): (1) Specialized
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habitat or microhabitat requirements, (2)
narrow environmental tolerances or
thresholds that are likely to be exceeded
under climate change, (3) dependence
on specific triggers or cues that are
likely to be disrupted (for example,
rainfall or temperature cues for
breeding, migration, or hibernation), (4)
dependence on interactions between
species that are likely to be disrupted,
and (5) inability or poor ability to
disperse quickly or to colonize more
suitable range. We apply these criteria
in this final rule to assess the
vulnerability of mountain yellow-legged
frogs to climate change.
At high elevations, where most extant
populations occur, mountain yellowlegged frogs depend on high mountain
lakes where both adult and larval frogs
overwinter under ice for up to 9 months
of the year. Overwintering under ice
poses physiological problems for the
frogs, most notably the depletion of
oxygen in the water during the winter
(Bradford 1983, p. 1171). Bradford
(1983, pp. 1174–1182) has found, based
on lab and field results, that tadpoles
are more resistant to low dissolved
oxygen levels than adult frogs; after two
drought years that were followed by a
severe winter, all frogs in 21 of 26 study
lakes were lost (with the exception of
one 2.1-m (6.9-ft) deep lake that
contained only one individual), while
tadpoles survived in all but one of the
shallowest lakes. Losses were
apparently due to oxygen depletion in a
year when there was exceptional
precipitation, ice depths that were
thicker than usual, and lake thawing
was 5 to 6 weeks later than the previous
year. The survival of adults in
substantial numbers was significantly
correlated with lake depth and confined
to lakes deeper than 4 m (13.1 ft).
Bradford (1983, pp. 1174–1179) found
that mean oxygen concentration in lakes
was directly related to maximum lake
depth, with dissolved oxygen levels
declining throughout the winter. He also
found that a thickened ice layer on a
lake causes the lake to become
effectively more shallow, leading to an
increased rate of oxygen depletion
(Bradford 1983, p. 1178). Studies of
winterkill of fish due to oxygen
depletion also show that oxygen
depletion is inversely related to lake
depth and occurs most rapidly in
shallow lakes relative to deeper lakes
(See review in Bradford 1983, p. 1179).
Bradford (1983, p. 1179) considered the
possibility that winterkill of the frogs
was due to freezing, but dismissed the
potential because some of the lakes
where winterkill occurred were deeper
than the probable maximum ice depth
in that year. Because the deeper lakes
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that once supported frog populations
now harbor introduced trout
populations and are generally no longer
available as refugia for frogs, the
shallower lakes where frogs currently
occur may be more vulnerable to
weather extremes in a climate with
increased variability, including drought
years and years with exceptional severe
cold winters. Such episodic stressors
may have been infrequent in the past,
but appear to be increasing, and they are
important to long-lived species with
small populations.
In summer, reduced snowpack and
enhanced evapotranspiration following
higher temperatures can dry out ponds
that otherwise would have sustained
rearing tadpoles (Lacan et al. 2008, p.
220), and may also reduce fecundity
(egg production) (Lacan et al. 2008, p.
222). Lacan et al. (2008, p. 211)
observed that most frog breeding
occurred in the smaller, fishless lakes of
Kings Canyon National Park that are
shallow and prone to summer drying.
Thus, climate change will likely reduce
available breeding habitat for mountain
yellow-legged frogs and lead to greater
frequency of stranding and death of
tadpoles as such lakes dry out earlier in
the year (Corn 2005, p. 64; Lacan et al.
2008, p. 222).
Earlier snowmelt is expected to cue
breeding earlier in the year. The
advance of this primary signal for
breeding phenology in montane and
boreal habitats (Corn 2005, p. 61) may
have both positive and negative effects.
Additional time for growth and
development may render larger
individuals more fit to overwinter;
however, earlier breeding may also
expose young tadpoles (or eggs) to
killing frosts in more variable
conditions of early spring (Corn 2005, p.
60).
Whether mountain yellow-legged
frogs depend on other species that may
be affected either positively or
negatively by climate change is unclear.
Climate change may alter invertebrate
communities (PRBO 2011 p. 24). In one
study, an experimental increase in
stream temperature was shown to
decrease density and biomass of
invertebrates (Hogg and Williams 1996,
p. 401). Thus, climate change might
have a negative impact on the mountain
yellow-legged frog prey base.
Indirect effects from climate change
may lead to greater risk to mountain
yellow-legged frog population
persistence. For example, fire intensity
and magnitude are projected to increase
(PRBO 2011, pp. 24–25), and, therefore,
the contribution and influence of this
stressor upon frog habitat and
populations will increase. Climate
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change may alter lake productivity
through changes in water chemistry, the
extent and timing of mixing, and
nutrient inputs from increased fires, all
of which may influence community
dynamics and composition (Melack et
al. 1997, p. 971; Parker et al. 2008, p.
12927). These changes may not all be
negative; for example, water chemistry
and nutrient inputs, along with warmer
summer temperatures, could increase
net primary productivity in high
mountain lakes to enhance frog food
sources, although changes in net
primary productivity may also
negatively affect invertebrate prey
species endemic to oligotrophic lakes
(low nutrient, low productivity).
Carey (1993, p. 359) has suggested
that, where environmental changes
cause sufficient stress to cause
immunological suppression, cold body
temperatures that montane amphibians
experience over winter could play a
synergistic role in reducing further
immunological responses to disease.
Thus, such conditions might make
mountain yellow-legged frogs more
susceptible to disease. Additionally,
Blaustein et al. (2001, p. 1808) have
suggested that climate change could also
affect the distribution of pathogens and
their vectors, exposing amphibians to
new pathogens. Climate change
(warming) has been hypothesized as a
driver for the range shift of Bd (Pounds
et al. 2006, p. 161; Bosch et al. 2007, p.
253). However, other work has indicated
that survival and transmission of Bd is
more likely facilitated by cooler and
wetter conditions (Corn 2005, p. 63).
Fisher et al. (2009, p. 299) present a
review of information available to date
and evaluate the competing hypotheses
regarding Bd dynamics, and they
present some cases that suggest a
changing climate can change the host–
pathogen dynamic to a more virulent
state.
The key risk factor for climate change
impacts on mountain yellow-legged
frogs is likely the combined effect of
reduced water levels in high mountain
lakes and ponds and the relative
inability of individuals to disperse and
colonize across longer distances in order
to occupy more favorable habitat
conditions (if they exist). Although such
adaptive range shifts have been
observed in some plant and animal
species, they have not been reported in
amphibians. The changes observed in
amphibians to date have been more
associated with changes in timing of
breeding (phenology) (Corn 2005, p. 60).
This limited adaptive capacity for
mountain yellow-legged frogs is a
function of high site fidelity and the
extensive habitat fragmentation due to
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the introduction of fishes in many of the
more productive and persistent high
mountain lake habitats and streams that
constitute critical dispersal corridors
throughout much of the frogs’ range (see
Factor C discussion above).
An increase in the frequency,
intensity, and duration of droughts
caused by climate change may have
compounding effects on populations of
mountain yellow-legged frogs already in
decline. In situations where other
stressors (such as introduced fish) have
resulted in the isolation of mountain
yellow-legged frogs in marginal habitats,
localized mountain yellow-legged frog
population crashes or extirpations
resulting from drought may exacerbate
their isolation and preclude natural
recolonization (Bradford et al. 1993, p.
887; Drost and Fellers 1996, p. 424;
Lacan et al. 2008, p. 222). Viers et al.
(2013, pp. 55, 56) have used a variety of
risk metrics to determine that both
mountain yellow-legged frog species in
the Sierra Nevada are highly vulnerable
to climate change, and that changing
hydrology and habitat in the Sierra
Nevada will likely have drastic impacts
on remaining populations. Climate
change represents a substantial future
threat to the persistence of mountain
yellow-legged frog populations.
Direct and Indirect Mortality
Other risk factors include direct and
indirect mortality as an unintentional
consequence of activities within
mountain yellow-legged frog habitat.
Mortality due to trampling by grazing
livestock has been noted in a limited
number of situations, with expected
mortality risk thought to be greatest if
livestock concentrate in prime breeding
habitat early in the season when adults
are breeding and egg masses are present
(Brown et al. 2009, p. 59). Brown et al.
(2009, p. 59) note that standards in the
SNFPA are intended to mitigate this
risk. Recreational uses also have the
potential to result in direct or indirect
mortality of mountain yellow-legged
frog individuals at all life stages. The
Forest Service has identified activities,
including recreational activities that
occur in the frogs’ breeding sites as
being risk factors for the frogs, while
noting that recreation use is a risk that
USFS management can change (USDA
2001a, pp. 213–214). Brown et al. (2009,
pp. 65–66) note that tadpoles and
juveniles, in particular, may be injured
or killed by trampling, crushing, etc., by
hikers, bikers, anglers, pets, packstock,
or off-highway vehicles, although the
number of documented situations
appears limited. Recreational activities,
such as hiking and camping, are
associated primarily with physical site
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alteration (changes to soil and
vegetation conditions), and such effects
are found to be highly localized. For
example, estimates in a heavily-used
portion of the Eagle Cap Wilderness in
Oregon indicated that no more than 2
percent of the area had been altered by
recreational use (Cole and Landres 1996,
p. 170). However, where impacts of
recreational use are highly localized,
species impacts due to trampling have
been identified, especially for rare plant
species (Cole and Landres 1996, p. 170).
Fire management activities (i.e. fuels
reduction and prescribed fire) lead to
some direct mortality and have the
potential to disrupt behavior. Please
refer to the proposed listing rule for the
Sierra Nevada yellow-legged frog and
the northern DPS of the mountain
yellow-legged frog under the Act (16
U.S.C. 1531 et seq.) for information
about effects of fire retardants on
mountain yellow-legged frogs. Roads
create the potential for direct mortality
of amphibians by vehicle strikes
(deMaynadier and Hunter 2000, p. 56)
and the possible introduction of
contaminants into new areas; however,
most extant populations are not located
near roads. Collectively, direct mortality
risks to mountain yellow-legged frogs
are likely of sporadic significance. They
may be important on occasion on a sitespecific basis, but are likely of low
prevalence across the range of the
species.
Small Population Size
In many localities, remaining
populations for both the Sierra Nevada
yellow-legged frog and the mountain
yellow-legged frog are small (CDFW,
unpubl. data). Brown et al. (2011, p. 24)
reported that about 90 percent of
watersheds have fewer than 10 adults
and 80 percent have fewer than 10
subadults and 100 tadpoles. Remnant
populations in the northern portion of
the range for the Sierra Nevada yellowlegged frog (from Lake Tahoe north) and
the southern portion of the populations
of the northern DPS of the mountain
yellow-legged frog (south of Kings
Canyon National Park) currently also
exhibit very low abundances (CDFW,
unpubl. data).
Compared to large populations, small
populations are more vulnerable to
extirpation from environmental,
demographic, and genetic stochasticity
(random natural occurrences), and
unforeseen (natural or unnatural)
catastrophes (Shaffer 1981, p. 131).
Environmental stochasticity refers to
annual variation in birth and death rates
in response to weather, disease,
competition, predation, or other factors
external to the population (Shaffer 1981,
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p. 131). Small populations may be less
able to respond to natural
´
environmental changes (Kery et al.
2000, p. 28), such as a prolonged
drought or even a significant natural
predation event. Periods of prolonged
drought are more likely to have a
significant effect on mountain yellowlegged frogs because drought conditions
occur on a landscape scale and all life
stages are dependent on habitat with
suitable perennial water. Demographic
stochasticity is random variability in
survival or reproduction among
individuals within a population (Shaffer
1981, p. 131) and could increase the risk
of extirpation of the smaller remaining
populations. Genetic stochasticity
results from changes in gene frequencies
due to the founder effect (loss of genetic
variation that occurs when a new
population is established by a small
number of individuals) (Reiger 1968, p.
163); random fixation (the complete loss
of one of two alleles in a population, the
other allele reaching a frequency of 100
percent) (Reiger 1968, p. 371); or
inbreeding depression (loss of fitness or
vigor due to mating among relatives)
´
(Soule 1980, p. 96). Additionally, small
populations generally have an increased
chance of genetic drift (random changes
in gene frequencies from generation to
generation that can lead to a loss of
variation) and inbreeding (Ellstrand and
Elam 1993, p. 225).
Allee effects (Dennis 1989, pp. 481–
538) occur when a population loses its
positive stock-recruitment relationship
(when population is in decline). In a
declining population, an extinction
threshold or ‘‘Allee threshold’’ (Berec et
al. 2006, pp. 185–191) may be crossed,
where adults in the population either
cease to breed or the population
becomes so compromised that breeding
does not contribute to population
growth. Allee effects typically fall into
three broad categories (Courchamp et al.
1999, pp. 405–410): lack of facilitation
(including low mate detection and loss
of breeding cues), demographic
stochasticity, and loss of heterozygosity
(a measure of genetic variability).
Environmental stochasticity amplifies
Allee effects (Dennis 1989, pp. 481–538;
Dennis 2002, pp, 389–401). The Allee
effects of demographic stochasticity and
loss of heterozygosity are likely as
mountain yellow-legged frog
populations continue to diminish.
The extinction risk for a species
represented by few small populations is
magnified when those populations are
isolated from one another. This is
especially true for species whose
populations normally function in a
metapopulation structure, whereby
dispersal or migration of individuals to
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new or formerly occupied areas is
necessary. Connectivity between these
populations is essential to increase the
number of reproductively active
individuals in a population; mitigate the
genetic, demographic, and
environmental effects of small
population size; and recolonize
extirpated areas. Additionally, fewer
populations by itself increases the risk
of extinction.
The combination of low numbers with
the other extant stressors of disease, fish
persistence, and potential for climate
extremes could have adverse
consequences for the mountain yellowlegged frog complex as populations
approach the Allee threshold. Small
population size is currently a significant
threat to most populations of mountain
yellow-legged frogs across the range of
the species.
Cumulative Impacts of Extant Threats
Stressors may act additively or
synergistically. An additive effect would
mean that an accumulation of otherwise
low threat factors acting in combination
may collectively result in individual
losses that are meaningful at the
population level. A synergistic effect is
one where the interaction of one or
more stressors together leads to effects
greater than the sum of those individual
factors combined. Further, the
cumulative effect of multiple added
stressors can erode population viability
over successive generations and act as a
chronic strain on the viability of a
species, resulting in a progressive loss of
populations over time. Such interactive
effects from compounded stressors
thereby act synergistically to curtail the
viability of frog metapopulations and
increase the risks of extinction.
It is difficult to predict the precise
impact of the cumulative threat
represented by the relatively novel Bd
epidemic across a landscape already
fragmented by fish stocking. The
singular threat of the Bd epidemic wave
in the uninfected populations of the
mountain yellow-legged frog complex in
the southern Sierra Nevada could
extirpate those populations as the
pathogen spreads. A compounding
effect of disease-caused extirpation is
that recolonization may never occur
because streams connecting extirpated
sites to extant populations now contain
introduced fishes, which act as barriers
to frog movement within
metapopulations. This situation isolates
the remaining populations of mountain
yellow-legged frogs from one another
(Bradford 1991, p. 176; Bradford et al.
1993, p. 887). It is logical to presume
that the small, fragmented populations
left in the recent wake of Bd spread
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through the majority of the range of the
Sierra Nevada yellow-legged frog may
experience further extirpations as
surviving adults eventually die, and
recruitment into the breeding pool from
the Bd-positive subadult class is
significantly reduced. These impacts
may be exacerbated by the present and
growing threat of climate change,
although this effect may take years to
materialize.
In summary, based on the best
available scientific and commercial
information, we consider other natural
and manmade factors to be substantial
ongoing threats to the Sierra Nevada
yellow-legged frog and the northern DPS
of the mountain yellow-legged frog.
These include high, prevalent risk
associated with climate change and
small population sizes, and the
associated risk from the additive or
synergistic effects of these two stressors
interacting with other acknowledged
threats, including habitat fragmentation
and degradation (see Factor A), disease
and predation (see Factor C), or other
threats currently present but with low
relative contribution in isolation.
Determination for the Sierra Nevada
Yellow-Legged Frog
We have carefully assessed the best
scientific and commercial information
available regarding the past, present,
and future threats to the Sierra Nevada
yellow-legged frog. The best available
information for the Sierra Nevada
yellow-legged frog shows that the
geographic extent of the species’ range
has declined, with local populationlevel changes first noticed in the early
1900s (Grinnell and Storer 1924, p. 664)
although they were still abundant at
many sites in the Sierra Nevada until
the 1960s (Zweifel 1955, pp. 237–238).
Population losses continued between
the 1960s and 1990s (Bradford et al.
1993, p. 883) and have continued in
recent decades. Now fewer, increasingly
isolated populations maintain viable
recruitment (entry of post-metamorphic
frogs into the breeding population).
Coupled with the observation that
remnant populations are also
numerically smaller (in some cases
consisting of few individuals), this
reduction in occupancy and population
density across the landscape suggests
significant losses in metapopulation
viability and high attendant risk to the
overall population of the species. The
impacts of the declines on population
resilience are two-fold: (1) The
geographic extent and number of
populations are reduced across the
landscape, resulting in fewer and more
isolated populations (the species is less
able to withstand population stressors
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and unfavorable conditions exist for
genetic exchange or dispersal to
unoccupied areas (habitat
fragmentation)); and (2) species
abundance (in any given population) is
reduced, making local extirpations
much more likely (decreased population
viability). Knapp et al. (2007b, pp. 1–2)
estimated a 10 percent decline per year
in the number of remaining mountain
yellow-legged frog populations and
argued for the listing of the species as
endangered based on this observed rate
of population loss.
Threats that face the Sierra Nevada
mountain yellow-legged frog, discussed
above under Factors A, C, D, and E,
increase the risk of the species’
extinction, given the isolation of
remaining populations. The best
available science indicates that the
introduction of fishes to the frog’s
habitat to support recreational angling is
one of the primary causes of the decline
of the Sierra Nevada yellow-legged frog
and poses a current and continuing
threat to the species (Factor A). Water
bodies throughout this range have been
intensively stocked with introduced fish
(principally trout). It is a threat of
significant influence, and although
fewer lakes are stocked currently than
were stocked prior to 2001, it remains
prevalent today because fish persist in
many high-elevation habitats even
where stocking has ceased. Further, the
introduction of fish has generally
restricted remaining Sierra Nevada
yellow-legged frog populations to more
marginal habitats, thereby increasing the
likelihood of localized extinctions.
Recolonization in these situations is
difficult for a highly aquatic species
with high site fidelity and unfavorable
dispersal conditions.
Historical livestock grazing activities
may also have modified the habitat of
the Sierra Nevada yellow-legged frog
throughout much of its range (Factor A).
Grazing pressure has been significantly
reduced from historical levels, but is
expected to have legacy effects on
mountain yellow-legged frog habitat
where prior downcutting and
headcutting of streams have resulted in
reduced water tables and would benefit
from restoration. Current grazing that
complies with forest standards and
guidelines is not expected to cause
habitat-related effects to the species in
almost all cases, but in limited cases
may continue to contribute to some
localized degradation and loss of
suitable habitat. The habitat-related
effects of recreation, packstock grazing,
dams and water diversions, roads,
timber harvests, and fire management
activities on the Sierra Nevada yellowlegged frog (Factor A) may have
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contributed to historical losses when
protections and use limits that are
currently afforded by USFS and NPS
standards and guidelines did not exist.
Currently, Federal land management
agencies with jurisdiction within the
current range of the Sierra Nevada
yellow-legged frog have developed
management standards and guidelines
that limit habitat damage due to these
activities, although in localized areas
habitat-related changes may continue to
affect individual populations.
Competitive exclusion and predation
by fish have eliminated or reduced
mountain yellow-legged frog
populations in stocked habitats, and left
remnant populations isolated, while
bullfrogs are expected to have negative
effects where they occur (Factor C). It is
important to recognize that, throughout
the vast majority of its range, Sierra
Nevada yellow-legged frogs did not coevolve with any species of fish, as they
predominantly occur in water bodies
above natural fish barriers.
Consequently, the species has not
evolved defenses against fish predation.
Sierra Nevada yellow-legged frogs are
vulnerable to multiple pathogens (see
Factor C) whose effects range from low
levels of infection within persistent
populations to disease-induced
extirpation of entire populations. The
Bd epidemic has caused extirpations of
Sierra Nevada yellow-legged frog
populations throughout its range and
caused associated significant declines in
numbers of individuals. Though Bd was
only recently discovered to affect the
Sierra Nevada yellow-legged frog, it
appears to infect populations at much
higher rates than other pathogens. The
imminence of this risk to populations in
currently uninfected habitats is
immediate and the potential effects
severe. The already-realized effects to
the survival of sensitive amphibian life
stages in Bd-positive areas are welldocumented. Although some
populations survive the initial Bd wave,
survival rates of metamorphs and
population viability are markedly
reduced relative to historical (pre-Bd)
norms.
These threats described above are
likely to be exacerbated by widespread
changes associated with climate change
and by current small population sizes in
many locations (see Factor E), while
instances of direct and indirect
mortality are expected to have
population-level effects only in
relatively uncommon, localized
situations. On a rangewide basis, the
existing regulatory mechanisms (Factor
D) have not been effective in protecting
populations from declines due to fish
stocking and continuing presence of fish
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and to disease, although standards and
guidelines developed by the USFS and
the NPS have largely limited threats due
to livestock and packstock grazing,
recreation, and timber use.
The main and interactive effects of
these various risk factors have acted to
reduce Sierra Nevada yellow-legged frog
populations to small fractions of their
historical habitat and reduce population
abundances significantly throughout
most of its current range. Remaining
areas that have yet to be impacted by Bd
are at immediate and severe risk.
Given the life history of this species,
dispersal, recolonization, and genetic
exchange are largely precluded by the
fragmentation of habitat common
throughout its current range as a result
of fish introductions. Frogs that may
disperse are susceptible to hostile
conditions in many circumstances. In
essence, Sierra Nevada yellow-legged
frogs have been marginalized by
historical fish introductions.
Populations have recently been
decimated by Bd, and the cumulative
effect of other stressors (such as
anticipated reduction of required
aquatic breeding habitats with climate
change and more extreme weather)
upon a fragmented landscape make
adaptation and recovery a highly
improbable scenario without active
intervention. The cumulative risk from
these stressors to the persistence of the
Sierra Nevada yellow-legged frog
throughout its range is significant.
The Act defines an endangered
species as any species that is ‘‘in danger
of extinction throughout all or a
significant portion of its range’’ and a
threatened species as any species ‘‘that
is likely to become endangered
throughout all or a significant portion of
its range within the foreseeable future.’’
We find that the Sierra Nevada yellowlegged frog is presently in danger of
extinction throughout its entire range,
based on the immediacy, severity, and
scope of the threats described above.
Specifically, these include habitat
degradation and fragmentation under
Factor A, predation and disease under
Factor C, and climate change and the
interaction of these various stressors
cumulatively impacting small remnant
populations under Factor E. There has
been a rangewide reduction in
abundance and geographic extent of
surviving populations of the Sierra
Nevada yellow-legged frog following
decades of fish stocking, habitat
fragmentation, and, most recently, a
disease epidemic. Surviving
populations are smaller and more
isolated, and recruitment in Bd-positive
populations is much reduced relative to
historical norms. This combination of
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population stressors makes species
persistence precarious throughout the
current range in the Sierra Nevada.
We have carefully assessed the best
scientific and commercial information
available regarding the past, present,
and future threats to the species, and
have determined that the Sierra Nevada
yellow-legged frog meets the definition
of endangered under the Act, rather
than threatened. This is because
significant threats are occurring now
and will occur in the future, at a high
magnitude and across the species’ entire
range, making the species in danger of
extinction at the present time. The rate
of population decline remains high in
the wake of Bd epidemics, and the
remaining Sierra Nevada yellow-legged
frog populations are at high, imminent
risk. Population declines are expected to
continue as maturing tadpoles succumb
to Bd infection, and fragmented
populations at very low abundances
will face significant obstacles to
recovery. Therefore, on the basis of the
best available scientific and commercial
information, and the threats posed to
these species under the listing factors
above, we are listing the Sierra Nevada
yellow-legged frog as endangered in
accordance with sections 3(6) and
4(a)(1) of the Act.
Under the Act and our implementing
regulations, a species may warrant
listing if it is endangered or threatened
throughout all or a significant portion of
its range. The Sierra Nevada yellowlegged frog is restricted in its range, and
the threats occur throughout the
remaining occupied habitat. Therefore,
we assessed the status of this species
throughout its entire range. The threats
to the survival of the species occur
throughout the species’ range and are
not restricted to any particular
significant portion of that range.
Accordingly, our assessment and final
determination applies to the species
throughout its entire range.
Final Determination for the Northern
DPS of the Mountain Yellow-Legged
Frog
We have carefully assessed the best
scientific and commercial information
available regarding the past, present,
and future threats to the northern DPS
of the mountain yellow-legged frog. The
best available information for the
northern DPS of the mountain yellowlegged frog shows that the geographic
extent of the species’ range has
declined, with local population-level
changes first noticed in the early 1900s
(Grinnell and Storer 1924, p. 664),
although they were still abundant at
many sites in the Sierra Nevada until
the 1960s (Zweifel 1955, pp. 237–238).
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24283
Population losses continued between
the 1960s and 1990s (Bradford et al.
1993, p. 883) and have continued in
recent decades. Now fewer, increasingly
isolated populations maintain viable
recruitment (entry of post-metamorphic
frogs into the breeding population).
Coupled with the observation that
remnant populations are also
numerically smaller (in some cases
consisting of a few individuals), this
reduction in occupancy and population
density across the landscape suggests
significant losses in metapopulation
viability and high attendant risk to the
overall population of the species. The
impacts of the declines on population
resilience are two-fold: (1) The
geographic extent and number of
populations are reduced across the
landscape, resulting in fewer and more
isolated populations (the species is less
able to withstand population stressors
and unfavorable conditions exist for
genetic exchange or dispersal to
unoccupied areas (habitat
fragmentation)); and (2) species
abundance (in any given population) is
reduced, making local extirpations
much more likely (decreased population
viability). Knapp et al. (2007b, pp. 1–2)
estimated a 10 percent decline per year
in the number of remaining mountain
yellow-legged frog populations and
argued for the listing of the species as
endangered based on this observed rate
of population loss.
Threats that face the northern DPS of
the mountain yellow-legged frog,
discussed above under Factors A, C, D,
and E, increase the risk of the species’
extinction, given the isolation of
remaining populations. The best
available science indicates that the
introduction of fishes to the frog’s
habitat to support recreational angling is
one of the primary causes of the decline
of the northern DPS of the mountain
yellow-legged frog and poses a current
and continuing threat to the species
(Factor A). Water bodies throughout this
range have been intensively stocked
with introduced fish (principally trout).
It is a threat of significant influence, and
although fewer lakes are stocked
currently than were stocked prior to
2001, it remains prevalent today
because fish persist in many highelevation habitats even where stocking
has ceased. Recolonization in these
situations is difficult for a highly
aquatic species with high site fidelity
and unfavorable dispersal conditions.
Climate change is likely to exacerbate
these other threats and further threaten
population resilience.
Historical livestock grazing activities
may also have modified the habitat of
the northern DPS of the mountain
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yellow-legged frog throughout much of
its range (Factor A). Grazing pressure
has been significantly reduced from
historical levels, but is expected to have
legacy effects to mountain yellow-legged
frog habitat where prior downcutting
and headcutting of streams have
resulted in reduced water tables that
still need restoration to correct. Current
grazing that complies with forest
standards and guidelines is not
expected to cause habitat-related effects
to the species in almost all cases, but in
limited cases may continue to
contribute to some localized
degradation and loss of suitable habitat.
The habitat-related effects of recreation,
packstock grazing, dams and water
diversions, roads, timber harvests, and
fire management activities on the
northern DPS of the mountain yellowlegged frog (Factor A) may have
contributed to historical losses when
protections and use limits that are
currently afforded by USFS and NPS
standards and guidelines did not exist.
Currently, Federal agencies with
jurisdiction within the current range of
the northern DPS of the mountain
yellow-legged frog have developed
management standards and guidelines
that limit habitat damage due to these
activities, although in localized areas
habitat-related changes may continue to
affect individual populations.
Competitive exclusion and predation
by fish have eliminated or reduced
mountain yellow-legged frog
populations in stocked habitats, and left
remnant populations isolated, while
bullfrogs are expected to have negative
effects where they occur (Factor C). It is
important to recognize that throughout
the vast majority of its range, the
northern DPS of the mountain yellowlegged frogs did not co-evolve with any
species of fish, as this species
predominantly occurs in water bodies
above natural fish barriers.
Consequently, the species has not
evolved defenses against fish predation.
Mountain yellow-legged frogs are
vulnerable to multiple pathogens (see
Factor C) whose effects range from low
levels of infection within persistent
populations to disease-induced
extirpation of entire populations. The
Bd epidemic has caused rangewide
extirpations of populations of the
northern DPS of the mountain yellowlegged frog and associated significant
declines in numbers of individuals.
Though Bd was only recently
discovered to affect the mountain
yellow-legged frog, it appears to infect
populations at much higher rates than
other pathogens. The imminence of this
risk to currently uninfected habitats is
immediate, and the potential effects
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severe. The already-realized effects to
the survival of sensitive amphibian life
stages in Bd-positive areas are welldocumented. Although some
populations survive the initial Bd wave,
survival rates of metamorphs and
population viability are markedly
reduced relative to historical (pre-Bd)
norms.
These threats are likely to be
exacerbated by widespread changes
associated with climate change and by
current small population sizes in many
locations (see Factor E), while instances
of direct and indirect mortality are
expected to have population-level
effects only in relatively uncommon,
localized situations. Rangewide, the
existing regulatory mechanisms (Factor
D) have not been effective in protecting
populations from declines due to fish
stocking and continuing presence of fish
and to disease, although standards and
guidelines developed by the USFS and
the NPS have largely limited threats due
to livestock and packstock grazing,
recreation, and timber use.
The main and interactive effects of
these various risk factors have acted to
reduce the northern DPS of the
mountain yellow-legged frog to a small
fraction of its historical range and
reduce population abundances
significantly throughout most of its
current range. Populations of this
species in remaining areas in the
southern Sierra Nevada that have yet to
be impacted by Bd are at immediate and
severe risk.
Given the life history of this species,
dispersal, recolonization, and genetic
exchange are largely precluded by the
fragmentation of habitat common
throughout its current range as a result
of fish introductions. Frogs that may
disperse are susceptible to hostile
conditions in many circumstances. In
essence, mountain yellow-legged frogs
have been marginalized by historical
fish introductions. Populations have
recently been decimated by Bd, and the
accumulation of other stressors (such as
anticipated reduction of required
aquatic breeding habitats with climate
change and more extreme weather)
upon a fragmented landscape make
adaptation and recovery a highly
improbable scenario without active
intervention. The cumulative risk from
these stressors to the persistence of the
mountain yellow-legged frog throughout
its range is significant.
The Act defines an endangered
species as any species that is ‘‘in danger
of extinction throughout all or a
significant portion of its range’’ and a
threatened species as any species ‘‘that
is likely to become endangered
throughout all or a significant portion of
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its range within the foreseeable future.’’
We find that the northern DPS of the
mountain yellow-legged frog is
presently in danger of extinction
throughout its entire range, based on the
immediacy, severity, and scope of the
threats described above. Specifically,
these include habitat degradation and
fragmentation under Factor A, predation
and disease under Factor C, and climate
change and the interaction of these
various stressors cumulatively
impacting small remnant populations
under Factor E. There has been a
rangewide reduction in abundance and
geographic extent of surviving
populations of the northern DPS of the
mountain yellow-legged frog following
decades of fish stocking, habitat
fragmentation, and, most recently, a
disease epidemic. Surviving
populations are smaller and more
isolated, and recruitment in Bd-positive
populations is much reduced relative to
historical norms. This combination of
population stressors makes species
persistence precarious throughout the
current range in the Sierra Nevada.
We have carefully assessed the best
scientific and commercial information
available regarding the past, present,
and future threats to the species, and
have determined that the northern DPS
of the mountain yellow-legged frog,
meets the definition of endangered
under the Act, rather than threatened.
This is because significant threats are
occurring now and will occur in the
future, at a high magnitude and across
the DPS’ entire range, making the
northern DPS of the mountain yellowlegged frog in danger of extinction at the
present time. The rate of population
decline remains high in the wake of Bd
epidemics, and northern DPS of the
mountain yellow-legged frog areas are at
high, imminent risk. The recent rates of
decline for these populations are even
higher than declines in the populations
of the Sierra Nevada yellow-legged frog,
and as Bd infects remaining core areas,
population viability will be significantly
reduced, and extirpations or significant
population declines are expected.
Population declines are expected to
continue as maturing tadpoles succumb
to Bd infection, and fragmented
populations at very low abundances
will face significant obstacles to
recovery. Therefore, on the basis of the
best available scientific and commercial
information, and the threats posed to
these species discussed under the listing
factors above, we are listing the
northern DPS of the mountain yellowlegged frog as endangered in accordance
with sections 3(6) and 4(a)(1) of the Act.
Under the Act and our implementing
regulations, a species may warrant
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listing if it is endangered or threatened
throughout all or a significant portion of
its range. The northern DPS of the
mountain yellow-legged frog addressed
in this final listing rule is restricted in
its range, and the threats occur
throughout the remaining occupied
habitat. Therefore, we assessed the
status of this DPS throughout its entire
range in the Sierra Nevada of California.
The threats to the survival of this DPS
occur throughout its range in the
southern Sierra Nevada and are not
restricted to any particular significant
portion of that range. Accordingly, our
assessment and final determination
applies to the DPS throughout its entire
range.
Summary of Biological Status and
Threats Affecting the Yosemite Toad
Background
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Taxonomy and Species Description
Please refer to the proposed listing
rule for the Yosemite toad under the Act
(16 U.S.C. 1531 et seq.) for additional
species information, including detailed
information on taxonomy. In this
section of the final rule, it is our intent
to discuss only those topics directly
relevant to the listing of the Yosemite
toad (Anaxyrus canorus) as threatened.
Habitat and Life History
Breeding habitat—Yosemite toads are
associated with wet meadows due to
their breeding ecology. Camp (1916, pp.
59–62) found Yosemite toads in wet
meadow habitats and at lake shores
located among lodgepole (Pinus
contorta) at the lower elevations to
whitebark (P. albicaulis) pines at the
higher elevations. Mullally (1953, pp.
182–183) found adult toads common on
the margins of high-elevation lakes,
streams, and pools wherever the
meadow vegetation was thicker or more
luxuriant than usual or where there
were patches of low willows (Salix
spp.). Liang (2010, p. 81) observed
Yosemite toads most frequently
associated with (in order of preference):
wet meadows, alpine-dwarf scrub, red
fir (Abies magnifica), water, lodgepole
pine, and subalpine conifer habitats.
Yosemite toads were found as often at
large as at small sites (Liang 2010, p.
19), suggesting that this species is
capable of successfully utilizing small
habitat patches. Liang also found that
population persistence was greater at
higher elevations, with an affinity for
relatively flat sites with a southwesterly
aspect (Liang 2010, p. 20; see also
Mullally 1953, p. 182). These areas
receive higher solar radiation and are
capable of sustaining hydric (wet),
seasonally ponded, and mesic (moist)
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breeding and rearing habitat. The
Yosemite toad is more common in areas
with less variation in mean annual
temperature, or more temperate sites
with less climate variation (Liang 2010,
pp. 21–22).
Adults are thought to be long-lived,
and this factor allows for persistence in
variable conditions and more marginal
habitats where only periodic good years
allow high reproductive success (USFS
et al. 2009, p. 27). Females have been
documented to reach 15 years of age,
and males as many as 12 years (Kagarise
Sherman and Morton 1993, p. 195);
however, the average longevity of the
Yosemite toad in the wild is not known.
Jennings and Hayes (1994, p. 52)
indicated that females begin breeding at
ages 4 to 6 years, while males begin
breeding at ages 3 to 5 years.
Adults appear to have high sitefidelity; Liang (2010, pp. 99, 100) found
that the majority of individuals
identified in multiple years were
located in the same meadow pools,
although individuals will move between
breeding areas (Liang 2010, p. 52; Liang
2013, p. 561). Breeding habitat includes
shallow, warm-water areas in wet
meadows, such as shallow ponds and
flooded vegetation, ponds, lake edges,
and slow-flowing streams (Karlstrom
1962, pp. 8–12; Brown 2013,
unpaginated). Tadpoles have also been
observed in shallow areas of lakes
(Mullally 1953, pp. 182–183).
Adult Yosemite toads are most often
observed near water, but only
occasionally in water (Mullally and
Cunningham 1956b, pp. 57–67). Moist
upland areas such as seeps and
springheads are important summer
nonbreeding habitats for adult toads
(Martin 2002, pp. 1–3). The majority of
their life is spent in the upland habitats
proximate to their breeding meadows.
They use rodent burrows for
overwintering and probably for
temporary refuge during the summer
(Jennings and Hayes 1994, pp. 50–53),
and they spend most of their time in
burrows (Liang 2010, p. 95). They also
use spaces under surface objects,
including logs and rocks, for temporary
refuge (Stebbins 1951, pp. 245–248;
Karlstrom 1962, pp. 9–10). Males and
females also likely inhabit different
areas and habitats when not breeding,
and females tend to move farther from
breeding ponds than males (USFS et al.
2009, p. 28).
Males exit burrows first, and spend
more time in breeding pools than
females, who do not breed every year
(Kagarise Sherman and Morton, 1993, p.
196). Data suggest that higher lipid
storage in females, which enhances
overwinter survival, also precludes the
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energetic expense of breeding every year
(Morton 1981, p. 237). The Yosemite
toad is a prolific breeder, laying many
eggs immediately at snowmelt. This is
accomplished in a short period of time,
coinciding with water levels in meadow
habitats and ephemeral pools they use
for breeding. Female toads lay
approximately 700–2,000 eggs in two
strings (one from each ovary) (USFS et
al. 2009, p. 21). Females may split their
egg clutches within the same pool, or
even between different pools, and may
lay eggs communally with other toads
(USFS et al. 2009, p. 22).
Eggs hatch within 3–15 days,
depending on ambient water
temperatures (Kagarise Sherman 1980,
pp. 46–47; Jennings and Hayes 1994, p.
52). Tadpoles typically metamorphose
around 40–50 days after fertilization,
and are not known to overwinter
(Jennings and Hayes 1994. p. 52).
Tadpoles are black in color, tend to
congregate together (Brattstrom 1962,
pp. 38–46) in warm shallow waters
during the day (Cunningham 1963, pp.
60–61), and then retreat to deeper
waters at night (Mullaly 1953, p. 182).
Rearing through metamorphosis takes
approximately 5–7 weeks after eggs are
laid (USFS et al. 2009, p. 25). Toads
need shallow, warm surface water that
persists through the period during
which they metamorphose; shorter
hydroperiods in that habitat can reduce
reproductive success (Brown 2013,
unpaginated).
Reproductive success is dependent on
the persistence of tadpole rearing sites
and conditions for breeding, egg
deposition, hatching, and rearing to
metamorphosis (USFS et al. 2009, p.
23). Given their association with
shallow, ephemeral habitats, Yosemite
toads are susceptible to droughts and
weather extremes. Abiotic factors
leading to mortality (such as freezing or
desiccation) appear to be more
significant during the early life stages of
toads, while biotic factors (such as
predation) are probably more prominent
factors during later life stages (USFS et
al. 2009, p. 30). However, since adult
toads lead a much more inconspicuous
lifestyle, direct observation of adult
mortality is difficult and it is usually
not possible to determine causes of
adult mortality.
Yosemite toads can move farther than
1 km (0.63 mi) from their breeding
meadows (average movement is 275 m
(902 ft)), and they utilize terrestrial
environments extensively (Liang 2010,
p. 85). The average distance traveled by
females is twice as far as males, and
home ranges for females are 1.5 times
greater than those for males (Liang 2010,
p. 94). Movement into the upland
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terrestrial environment following
breeding does not follow a predictable
path, and toads tend to traverse longer
distances at night, perhaps to minimize
evaporative water loss (Liang 2010, p.
98). Martin (2008, p. 123) tracked adult
toads during the active season and
found that on average toads traveled a
total linear distance of 494 m (1,620 ft)
within the season, with minimum travel
distance of 78 m (256 ft) and maximum
of 1.76 km (1.09 mi).
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Historical Range and Distribution
The known historical range of the
Yosemite toad in the Sierra Nevada
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extended from the Blue Lakes region
north of Ebbetts Pass (Alpine County) to
south of the Evolution Lake area (Fresno
County) (Karlstrom 1962, p. 3; Stebbins
1985, p. 72; see also Knapp 2013,
unpaginated; Brown 2013,
unpaginated). Yosemite toad habitat
historically spanned elevations from
1,460 to 3,630 m (4,790 to 11,910 ft)
(Stebbins 1985, p. 72; Stephens 2001, p.
12).
similar to the historical range defined
above (USFS et al. 2009, p. 41).
However, within that range, toad
habitats have been degraded and may be
decreasing in area as a result of conifer
encroachment and historical livestock
grazing (see Factor A below). The vast
majority of the Yosemite toad’s range is
within federally managed land. Figure
2, Estimated Range of Yosemite Toad,
displays a range map for the species.
Current Range and Distribution
The current range of the Yosemite
toad, at least in terms of overall
geographic extent, remains largely
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Population Estimates and Status
Baseline data on the number and size
of historical Yosemite toad populations
are limited, and historic records are
largely based on accounts from field
notes, or pieced together through
museum collections, thereby providing
limited information on historical
populations. Systematic survey
information across the range of the
species on National Forest System
Lands largely follows the designation of
the Yosemite toad as a candidate species
under the Act. In addition, surveys for
the Yosemite toad have been conducted
within Yosemite, Kings Canyon, and
Sequoia National Parks (Knapp 2013,
unpaginated). From these recent
inventories, Yosemite toads have been
found at 469 localities collectively on
six National Forests (USFS et al. 2009,
p. 40; see also Brown and Olsen 2013,
pp. 675–691), at 179 breeding sites that
were surveyed between 1992 and 2010
in Yosemite National Park (Berlow et al.
2013, p. 3), and detected at 18 localities
in Kings Canyon National Park (NPS
2011, geospatial data). Although we did
not cite to the information from the
National Parks in the proposed rule, we
had the geospatial occupancy data that
is currently included in Berlow et al.
2013, and we utilized that data in our
analysis for the proposed listing (see
comments 6 and 7 below, and their
respective responses). The number of
localities identified in these surveys
reflects more occupied sites than were
known before such extensive surveys
were conducted, and indicates that the
species is still widespread throughout
its range. These inventories were
typically conducted to determine toad
presence or absence (they were not
censuses), and do not explicitly
compare historic sites to recent surveys.
Moreover, single-visit surveys of toads
are unreliable as indices of abundance
because timing is so critical to the
presence of detectable life stages and
not all potential breeding habitats
within the range of the species were
surveyed (USFS et al. 2009, p. 41; Liang
2010, p. 10; Brown and Olsen 2013, p.
685). Given these considerations,
conclusions about population trends,
abundance, or extirpation rates are not
possible from these datasets overall.
One pair of studies allows us to
compare current distribution with
historic distributions and indicates that
large reductions have occurred. In 1915
and 1919, Grinnell and Storer (1924, pp.
657–660) surveyed for vertebrates at 40
sites along a 143-km (89-mi) west-to-east
transect across the Sierra Nevada,
through Yosemite National Park, and
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found Yosemite toads at 13 of those
sites. In 1992, Drost and Fellers (1996,
pp. 414–425) conducted more thorough
surveys, specifically for amphibians, at
38 of the Grinnell and Storer sites plus
additional nearby sites. Drost and
Fellers (1996, pp. 418) found that
Yosemite toads were absent from 6 of 13
sites where they had been found in the
original Grinnell and Storer (1924)
survey. Moreover, at the sites where
they were present, Yosemite toads most
often occurred in very low numbers
relative to general abundance reported
in the historical record (Grinnell and
Storer 1924, pp. 657–660). Therefore, by
the early 1990s, the species was either
undetectable or had declined in
numbers at 9 of 13 (69 percent) of the
Grinnell and Storer sites (Drost and
Fellers 1996, p. 418).
Another study comparing historic and
current occurrences also found a large
decline in Yosemite toad distribution. In
1990, David Martin surveyed 75 sites
throughout the range of the Yosemite
toad for which there were historical
records of the species’ presence. This
study found that 47 percent of
historically occupied sites showed no
evidence of any life stage of the species
(Stebbins and Cohen 1995, pp. 213–
215). This result suggests a range-wide
decline to about one half of historical
sites, based on occupancy alone.
A third study comparing historic and
recent surveys indicates declines in
Yosemite toad distribution. Jennings
and Hayes (1994, pp. 50–53) reviewed
the current status of Yosemite toads
using museum records of historic and
recent sightings, published data, and
unpublished data and field notes from
biologists working with the species.
They estimated a loss of over 50 percent
of former Yosemite toad locations
throughout the range of the species
(based on 144 specific sites).
The only long-term, site-specific
population study for Yosemite toads
documented a dramatic decline over 2
decades of monitoring. Kagarise
Sherman and Morton (1993, pp. 186–
198) studied Yosemite toads at Tioga
Pass Meadow (Mono County, California)
from 1971 through 1991 (with the most
intensive monitoring through 1982).
They documented a decline in the
average number of males entering the
breeding pools from 258 to 28 during
the mid-1970s through 1982. During the
same time period, the number of
females varied between 45 and 100, but
there was no apparent trend in number
observed. During the 1980s, it appeared
that males continued to decline, females
also declined, and breeding activity
became sporadic. By 1991, they found
only one male and two egg masses.
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Sadinski (2004, p. 40) revisited the
survey locations annually from 1995
and 2001 and found a maximum of two
males and two egg masses, suggesting
the toads in Tioga Meadows had not
recovered from their decline. In the
study of Yosemite toads at nearby Dana
Meadows, Sadinski (2004, pp. 39–42)
documented few adults within the
habitats surveyed, finding substantial
mortality in embryos that he associated
with effects of ice, water mold, and
flatworms. Sadinski (2004, pp. 38–42)
also found high larval mortality when
breeding sites dried before larvae could
reach metamorphosis. Sadinski (2004)
stated that the proximity of the Kagarise
Sherman and Morton (1993) study sites
at Tioga Meadows and his sites in Dana
Meadows practically ensured that
animals from both sites were part of the
same metapopulation. Sadinski
surmised that perhaps much of that
metapopulation experienced events at
breeding sites similar to those that
Kagarise Sherman and Morton (1993)
observed (Sadinski 2004, pp. 39–40). He
further opined that, if each of his
substantial sites had previously
supported hundreds of breeding adults
in the 1970s, the overall population of
Yosemite toads had declined
dramatically throughout the area since
that time.
Kagarise Sherman and Morton (1993,
pp. 186–198) also conducted occasional
surveys of six other populations in the
eastern Sierra Nevada. Five of these
populations showed long-term declines
that were evident beginning between
1978 through 1981, while the sixth
population held relatively steady until
the final survey in 1990, at which time
it dropped. In 1991, E.L. Karlstrom
revisited the site where he had studied
a breeding population of Yosemite toads
from 1954 to 1958 (just south of Tioga
Pass Meadow within Yosemite National
Park), and found no evidence of toads
or signs of breeding (Kagarise Sherman
and Morton 1993, p. 190).
The most reliable information about
Yosemite toad population status and
trends is the USFS SNAMPH. This
study, conducted on National Forest
System Lands, is designed to provide
statistical comparisons across 5-year
monitoring cycles with 134 watersheds
(Brown et al. 2011, pp. 3–4). This
approach allows researchers to assess
trends for the entire range of the toad,
rather than at limited survey sites (C.
Brown 2012, pers. comm., see also
Brown and Olsen 2013). The results of
this assessment indicate the species has
declined from historical levels, with
Yosemite toads occurring in
approximately 13 percent of watersheds
where they existed prior to 1990. This
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study also found that breeding was
occurring in approximately 84 percent
of the watersheds that were occupied in
the period 1990–2001, suggesting that
the number of locations where breeding
occurs has continued to decline.
Additionally, the study found that
breeding currently occurs in an
estimated 22 percent of watersheds
within the current estimated range of
the species (Brown et al. 2012, p. 115).
Moreover, overall abundances in the
intensively monitored watersheds were
very low (fewer than 20 males per
meadow per year) relative to other
historically reported abundances of the
species (Brown et al. 2011, p. 4). Brown
et al. (2011, p. 35) suggest that
populations are now very small across
the range of the species. During their
monitoring over the past decade, they
found only 18 percent of occupied
survey watersheds range-wide had
‘‘large’’ populations (more than 1,000
tadpoles or 100 of any other lifestage
detected at the time of survey). While
not all surveys were conducted at the
peak of tadpole presence and adults are
not reliably found outside of the
breeding season, Brown et al. (2012)
surveyed many sites at appropriate
times and rarely found the large
numbers of tadpoles or metamorphs that
would be expected if population sizes
were similar to those reported
historically. The researchers interpret
these data, in combination with
documented local population declines
from other studies (see above), to
support the hypothesis that population
declines have occurred range-wide
(Brown et al. 2012, p. 11).
Summary of Changes From the
Proposed Rule for the Yosemite Toad
Based on peer review and Federal,
State, and public comments (see
comments in the Summary of
Comments and Recommendations
section, below), we clarified information
for the Yosemite toad to better
characterize our knowledge of the
species’ habitat requirements.
Specifically, we reorganized and
clarified the habitat details (Habitat and
Life History), southern extent of the
species’ range (Historic Range and
Distribution), and species surveys
(USFS and NPS). We also added
information on occupancy in National
Parks that was inadvertently omitted
from the proposed rule (Population
Estimates and Status).
In the Summary of Factors Affecting
the Species section, under Factor A, we
made small changes to the discussion
about meadow loss and degradation in
order to improve clarity. In the
Livestock Use (Grazing) Effects to
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Meadow Habitat section, we reorganized
the information and separated the
effects of historic livestock grazing from
the effects due to current grazing levels,
and we added additional references
received from the USFS. In the Roads
and Timber Harvest Effects to Meadow
Habitat section, we clarified the extent
to which these activities overlap with
the Yosemite toad’s range and
distinguished the effects of past
activities from the effects of current
activities. We added information on
road locations and on USFS Forest
standards and guidelines that currently
limit the effects of these activities on
riparian areas. In this final rule, we
found that roads and timber harvest
activities are not current and ongoing
threats to the species. However, there
may be localized effects where legacy
effects of past road building or timber
harvest continue to modify wet
meadows or where activities occur in
close proximity to extant Yosemite toad
populations.
In the Fire Management section, we
added information to clarify that
Yosemite toads primarily occur in
higher elevation areas where fire
suppression activities are rarely
conducted. This finding suggests that
fire suppression has had little effect on
forest encroachment into meadow
habitats in most areas where the species
occurs. In the Recreation and Packstock
Effects to Meadow Habitat section, we
added additional information on USFS
and NPS restoration activities to protect
meadows, off-highway vehicle effects,
packstock use, and agency monitoring
and protection activities to limit effects
due to packstock use. We revised our
conclusion to clarify that, in general, we
do not consider habitat-related changes
associated with current levels of hiking,
backpacking, or packstock use to pose a
risk to Yosemite toad populations.
Recreation may have habitat-related
effects to toads in localized areas where
use adjacent to occupied meadows is
exceptionally heavy, or where heavy or
motorized use results in changes to
meadow hydrology. Accordingly,
rangewide, recreation is a threat of low
prevalence. In the section on Dams and
Water Diversions, we added information
to clarify that almost all reservoirs are
located below the range of the Yosemite
toad. We include small changes in the
Climate Change section to improve
clarity or add information from
references provided during peer review.
In Factor B, we added information
provided during the comment period,
which documented the sale of one
Yosemite toad from a pet store in
Southern California (store now closed).
We also added information on
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protections provided by agency-required
research permits. In Factor C, based on
peer review comments, we added
information on a Bd study on Yosemite
toads. We removed the discussion of
contaminants under Factor E, and we
refer readers to the proposed rule
affirming that the best available
information indicates that contaminants
do not pose a current or continuing
threat to the Yosemite toad. We also
added new information in the Other
Sources of Direct and Indirect Mortality
section as a result of information
provided during peer review. Although
we have not changed the determination,
we have made a few small changes in
the wording of the determination for the
Yosemite toad to reflect the above
changes.
Summary of Factors Affecting the
Species
Section 4 of the Act (16 U.S.C. 1533),
and its implementing regulations at 50
CFR part 424, set forth the procedures
for adding species to the Federal Lists
of Endangered and Threatened Wildlife
and Plants. Under section 4(a)(1) of the
Act, we may list a species 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; and (E)
other natural or manmade factors
affecting its continued existence. Listing
actions may be warranted based on any
of the above threat factors, singly or in
combination. Each of these factors is
discussed below, and changes from the
proposed rule (78 FR 24472, April 25,
2013) are reflected in these discussions.
Factor A. The Present or Threatened
Destruction, Modification, or
Curtailment of Its Habitat or Range
The habitat comprising the current
range of the Yosemite toad is generally
characterized by low levels of physical
disturbance (there is little to no current
development pressure). However, these
areas are also generally more sensitive
to perturbation and take longer to
recover from disturbances due to
reduced growing seasons and harsher
environmental conditions. Since
Yosemite toads rely heavily on shallow,
ephemeral water, they may be more
sensitive to minor changes in their
habitat. Loss or alteration of suitable
breeding habitat can reduce
reproductive success, which may have a
profound impact when population
numbers are small. Past management
and development activity has played a
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role in the degradation of meadow
habitats within the Sierra Nevada.
Human activities within these habitats
include grazing, timber harvest, fuels
management, recreation, and water
development.
Meadow Habitat Loss and Degradation
Some of the habitat effects associated
with grazing activities that were
described for the mountain yellowlegged frogs (see the Summary of
Factors Affecting the Species section for
those species, above) also apply to
Yosemite toads. However, there are
differences based on the Yosemite toad’s
reliance on very shallow, ephemeral
water in meadow and pool habitats
versus the deeper lakes and streams
frequented by mountain yellow-legged
frogs. Because Yosemite toads rely on
very shallow, ephemeral water, they
may be sensitive to even minor changes
in their habitat, particularly to
hydrology (Brown 2013, unpaginated).
Meadow habitat quality in the Western
United States, and specifically the
Sierra Nevada, has been degraded by
past activities, such as overgrazing, tree
encroachment, fire suppression, and
road building, over the last century
(Stillwater Sciences 2008, pp. 1–53;
Halpern et al. 2010, pp. 717–732; Vale
1987, pp. 1–18; Ratliff 1985, pp. i–48).
These past activities have contributed to
erosion and stream incision in areas of
the Sierra Nevada, leading to meadow
dewatering and encroachment by
invasive vegetation (Menke et al. 1996,
pp. 25–28; Lindquist and Wilcox 2000,
p. 2).
Given the reliance of the Yosemite
toad on these meadow and pool habitats
for breeding, rearing, and adult survival,
it is logical to conclude that the various
stressors have had an indirect effect on
the viability of Yosemite toad
populations via degradation of their
habitat. Loss of connectivity of habitats
leads to further isolation and population
fragmentation. Because of physiological
constraints, the tendency to move only
short distances, and high site fidelity,
amphibians may be unable to recolonize
unoccupied sites following local
extinctions if the distance between sites
is too great, although recolonization can
occur over time (Blaustein et al. 1994a,
p. 8).
Since the existence of meadows is
largely dependent on their hydrologic
setting, most meadow degradation is
due fundamentally to hydrologic
alterations (Stillwater Sciences 2008, p.
13). There are many drivers of
hydrologic alterations in meadow
ecosystems. In some locations, historic
water development and ongoing water
management activities have physically
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changed the underlying hydrologic
system. Diversion and irrigation ditches
formed a vast network that altered local
and regional stream hydrology, although
these manmade systems are generally
below the range of the Yosemite toad.
Timber harvest and associated road
construction further altered erosion and
sediment delivery patterns in rivers and
meadow streams. Fire suppression and
an increase in the frequency of large
wildfires due to excessive fuel buildup
have introduced additional disturbance
pressures to the meadows of the Sierra
Nevada (Stillwater Sciences 2008, p.
13). Many meadows now have downcut
stream courses, compacted soils, altered
plant community compositions, and
diminished wildlife and aquatic habitats
(SNEP 1996, pp. 120–121).
Land uses causing channel erosion are
a threat to Sierra Nevada meadows.
These threats include erosive activities
within the watershed upslope of the
meadow, along with impacts from land
use directly in the meadows themselves.
Compaction of meadow soils by roads or
intensive trampling (for example,
overgrazing) can reduce infiltration,
accelerate surface run-off, and thereby
lead to channel incision (Menke et al.
1996, pp. 25–28). Mining, overgrazing,
timber harvesting, and railroad and road
construction and maintenance have
contributed to watershed degradation,
resulting in accelerated erosion,
sedimentation in streams and reservoirs,
meadow dewatering, and degraded
terrestrial and aquatic habitats (Linquist
2000, p. 2). Deep incision has been
documented in several meadows in the
Sierra Nevada. One example is Halstead
Meadow in Sequoia National Park,
where headcutting exceeds 10 feet in
many areas and is resulting in widening
channels, erosion in additional
meadows, and a lowered water table
(Cooper and Wolf 2006, p. 1).
The hydrologic effects of stream
incision on the groundwater system may
significantly impact groundwater
storage, affecting late summer soil
moisture and facilitating vegetation
change (Bergmann 2004, pp. 24–31). For
example, in the northern Sierra Nevada,
logging, overgrazing, and road/railroad
construction have caused stream
incision, resulting in dewatering of
riparian meadow sediments and a
succession from native wet meadow
vegetation to sagebrush and dryland
grasses (Loheide and Gorelick 2007, p.
2). A woody shrub (Artemisia
rothrockii) is invading meadows as
channel incision causes shallow-waterdependent herbs to die back, allowing
shrub seedlings to establish in disturbed
areas during wet years (Darrouzet-Nardi
et al. 2006, p. 31).
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Mountain meadows in the western
United States and Sierra Nevada have
also been progressively colonized by
trees (Thompson 2007, p. 3; Vale 1987,
p. 6), with an apparent pattern of
encroachment during two distinct
periods in the late 1800s and mid-1900s
(Halpern et al. 2010, p. 717). This trend
has been attributed to a number of
factors, including climate, changes in
fire regime, and cessation of sheep
grazing (Halpern et al. 2010, pp. 717–
718; Vale 1987, pp. 10–13), but analyses
are limited to correlational comparisons
and research results are mixed, so the
fundamental contribution of each
potential driver remains uncertain. We
discuss the contribution of these factors
to habitat loss and degradation for the
Yosemite toad below.
Livestock Use (Grazing) Effects to
Meadow Habitat
The combined effect of legacy
conditions from historically excessive
grazing use and current livestock
grazing activities have the potential to
impact habitat in the range of the
Yosemite toad. The following
subsections discuss the effects of
excessive historical grazing, current
extent of grazing, and current grazing
management practices.
Overgrazing has been associated with
accelerated erosion and gullying of
meadows (Kattelmann and Embury
1996, pp. 13, 18), which leads to
siltation and more rapid succession of
meadows. Grazing can cause erosion by
disturbing the ground, damaging and
reducing vegetative cover, and
destroying peat layers in meadows,
which lowers the groundwater table and
summer flows (Armour et al. 1994, pp.
9–12; Martin 2002, pp. 1–3; Kauffman
and Krueger 1984, pp. 431–434).
Downcut channels, no longer connected
to the historic, wide floodplains of the
meadow, instead are confined within
narrow, incised channels. Downstream,
formerly perennial (year-round) streams
often become intermittent or dry due to
loss of water storage capacity in the
meadow aquifers that formerly
sustained them (Lindquist et al. 1997,
pp. 7–8).
Heavy grazing can alter vegetative
species composition and contribute to
lodgepole pine (Pinus contorta)
invasion (Ratliff 1985, pp. 33–36).
Lowering of the water table facilitates
encroachment of conifers into meadows.
Gully formation and lowering of water
tables, changes in the composition of
herbaceous vegetation, increases in the
density of forested stands, and the
expansion of trees into areas that
formerly were treeless have been
documented in California wilderness
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the Effects of Excessive Historical
Grazing section in Factor A analysis for
the Sierra Nevada and mountain yellowlegged frogs, above).
Livestock grazing in the Sierra Nevada
has been widespread for so long that, in
most places, no ungrazed areas are
available to illustrate the natural
condition of the habitat (Kattelmann
and Embury 1996, pp. 16–18). Dull
(1999, p. 899) conducted stratigraphic
pollen analysis (identification of pollen
Effects of Historical Livestock Grazing
in sedimentary layers) in mountain
Grazing of livestock in Sierra Nevada
meadows of the Kern Plateau, and found
meadows and riparian areas (rivers,
significant vegetation changes
streams, and adjacent upland areas that
attributable to sheep and cattle grazing
directly affect them) began in the midby 1900 (though fire regime change was
1700s with the European settlement of
also implicated; see below). This
California (Menke et al. 1996, p. 7).
degradation is widespread across the
Following the gold rush of the midSierra Nevada. Cooper and Wolf 2006
1800s, grazing increased to a level
(p. 1) reports that 50 to 80 percent of
exceeding the carrying capacity of the
grazed meadows now dominated by dry
available range, causing significant
meadow plants were formerly wet
impacts to meadow and riparian
meadows (Cooper and Wolf 2006, p. 1).
ecosystems (Meehan and Platts 1978, p.
Due to the long history (Menke et al.
275; Menke et al. 1996, p. 7). By the turn 1996, Ch. 22, pp. 1–52) of livestock and
of the 20th century, high Sierra Nevada
packstock grazing in the Sierra Nevada
meadows were converted to summer
and the lack of historical Yosemite toad
rangelands for grazing cattle, sheep,
population size estimates, it is
horses, goats, and pigs, although the
impossible to establish a reliable
alpine areas were mainly grazed by
quantitative estimate for the historical
sheep (Beesley 1996, pp. 7–8; Menke et
significance and contribution of grazing
al. 1996, p. 14). Stocking rates of both
on Yosemite toad populations.
cattle and sheep in Sierra meadows in
However, because of the documented
the late 19th and early 20th centuries
negative effects of livestock on Yosemite
were very heavy (Kosco and Bartolome
toad habitat, and the documented direct
1981, pp. 248–250), and grazing
mortality caused by livestock, the
severely degraded many meadows
decline of some populations of
(Ratliff 1985, pp. 26–31; Menke et al.
Yosemite toad has been attributed to the
1996, p. 14). Grazing impacts occurred
effects of livestock grazing (Jennings and
across the entire range of the Yosemite
Hayes 1994, pp. 50–53; Jennings 1996,
toad, as cattle and sheep were driven
pp. 921–944). Because Yosemite toad
virtually everywhere in the Sierra
breeding habitat is generally in very
Nevada where forage was available
shallow waters within meadows, the
(Kinney 1996, pp. 37–42; Menke et al.
breeding habitat is thought to be more
1996, p. 14).
vulnerable to changes in hydrology
Grazing within the National Forests
caused by grazing because the small
has continued into recent times, with
shallow pools are more easily impacted
reduction in activity (motivated by
(Knapp 2002c, p. 1; Martin 2002, pp. 1–
resource concerns, conflicts with other
3; USFS et al. 2009, pp. 22, 59–62;
uses, and deteriorating range
Brown 2013, unpaginated). U.S.
conditions) beginning in the 1920s. A
Geological Survey records indicate that
brief wartime increase in the 1940s
Yosemite, Sequoia, and Kings Canyon
followed, before grazing continued to be have no meadows within the parks that
scaled back beginning in the 1950s
are documented to have degraded
through the early 1970s. However,
hydrology (see NPS 2013, p. 7);
despite these reductions, grazing still
conditions in the parks may be related
exceeded sustainable capacity in many
to the early elimination of most grazing
areas (Menke et al. 1996, p. 9; UC 1996a, on national parklands in the Sierra
p. 115). Historical evidence indicates
Nevada.
that heavy livestock use in the Sierra
Effects of Current Livestock Grazing
Nevada has resulted in widespread
Currently, approximately 33 percent
damage to rangelands and riparian
of the estimated range of the Yosemite
systems due to sod destruction in
toad is within active USFS grazing
meadows, vegetation destruction, and
gully erosion (see review in Brown et al. allotments (USFS 2008, geospatial data).
While stocking rates have been reduced
2009, pp. 56–58 and in USFS et al.
2009, p. 57). (For additional information or eliminated in most areas, legacy
effects including eroded channels, soil
on historical grazing regimes, refer to
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areas and National Parks (Cole and
Landres 1996, p. 171). This invasion has
been attributed to sheep grazing, though
the phenomenon has been observed on
both ungrazed meadows and on
meadows grazed continually since about
1900 (Ratliff 1985, p. 35), suggesting
that other drivers may be involved (see
‘‘Effects of Fire Suppression on Meadow
Habitats’’ and ‘‘Climate Effects to
Meadow Habitat’’ below).
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erosion, and stream entrenchment that
resulted in lowered water tables, drier
meadows, and tree encroachment could
still be observed in some Sierran
meadows, especially in National Forests
where grazing was more intense (Vankat
and Major 1978, pp. 386–397). Meadow
conditions in the Sierra Nevada have
improved over time, but local problems
could still be found as of 1985 (Ratliff
1985, pp. ii–iii) and numerous examples
of head-cutting and stream incision are
available within the range of the toad
(Knapp 2013, unpaginated). (For
additional information, see sections
above pertaining to effects of grazing on
the mountain yellow-legged frogs.)
The influence of grazing on toad
populations in recent history is
uncertain, despite more available data
on land use and Yosemite toad
occurrence. In 2005, the USFS, in
collaboration with other researchers,
began a 5-year study with multiple
components to assess the effects of
grazing on Yosemite toads (Allen-Diaz
et al. 2010, pp. 1–45; Roche et al. 2012a,
pp. 56–65; Roche et al. 2012b, pp. 1–11;
McIlroy et al.. 2013, pp. 1–11).
Specifically, the goals of the research
were to assess: (1) Whether livestock
grazing under SNFPA Riparian
Standards and Guidelines has a
measurable effect on Yosemite toad
populations and (2) effects of livestock
grazing on key habitat components that
affect survival and recruitment of
Yosemite toad populations. SNFPA
standards and guidelines limit livestock
utilization of grass and grass-like plants
to a maximum of 40 percent (or a
minimum 4-inch stubble height) (USDA
2004, p. 56). These companion studies
did not detect an effect from grazing
activity on young-of-year toad density or
breeding pool occupancy, water quality,
or cover (Allen-Diaz et al. 2010, p. 1;
Roche et al. 2012a, p. 56; Roche et al.
2012b, p. 1–1; McIlroy et al.. 2013, p. 1).
It is important to note that the results
of these studies did not present a direct
measurement of toad survival (for
example, mark—recapture analysis of
population trends), and the design was
limited in numbers of years and
treatment replicates. It is plausible that,
for longer lived species with irregular
female breeding activity over the time
course of this particular study,
statistical power was not sufficient to
discern a treatment effect. Further, a
time lag could occur between effect and
discernible impacts, and significant
confounding variability in known
drivers such as interannual variation in
climate.
Additionally, the experimental design
in the studies tested the hypothesis that
forest management guidelines (at 40
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percent use threshold) were impacting
toad populations, and this limited some
analyses and experimental design to
sites with lower treatment intensities.
Researchers reported annual utilization
by cattle ranging from 10–48 percent,
while individual meadow use ranged
from 0–76 percent (the SNFPA
allowable use is capped at 40 percent)
(Allen-Diaz et al. 2010, p. 5). As a result
of the study design, the Allen-Diaz
study does not provide sufficient
information on the impacts of grazing
on Yosemite toads above the prescribed
management guidelines. In general, it is
not clear to what extent brief episodes
of intense use (such as in cattle
gathering areas) have as negative
impacts on toads, or over what
percentage of the grazed meadow
landscape such heavier usage may
occur.
The researchers observed significant
variation in young-of-year occupancy in
pools between meadows and years, and
within meadows over years (Allen-Diaz
et al. 2010, p. 7). This variability would
likely mask treatment effects, unless the
grazing variable was a dominant factor
driving site occupancy, and the
magnitude of the effect was quite severe.
Further, in an addendum to the initial
report, Lind et al. (2011b, pp. 12–14)
report statistically significant negative
(inverse) relationships for tadpole
density and grazing intensity (tadpole
densities decreased when percent use
exceeded between 30 and 40 percent).
This result supports the hypothesis that
grazing at intensities approaching and
above the 40 percent threshold can
negatively affect Yosemite toad
populations.
Allen-Diaz et al. (2010, p. 2) and
Roche et al. (2012b, pp. 6–7) found that
toad occupancy is strongly driven by
meadow wetness (hydrology) and
suggested attention should focus on
contemporary factors directly impacting
meadow wetness, such as climate, fire
regime changes, and conifer
encroachment (see Factor A above). The
researchers also stated that meadow use
by cattle during the grazing season is
driven by selection of plant
communities found in drier meadows
(Allen-Diaz et al. 2010, p. 2). This
suggests that the apparent differences in
preference could provide for some
segregation of toad and livestock use in
meadow habitats, so that at least direct
mortality threats may be mitigated by
behavioral isolation. Based on the
limitations of the study as described
above, we find the initial results from
Allen-Diaz et al. (2010, pp. 1–45) to be
inconclusive to discern the impacts of
grazing on Yosemite toad populations
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where grazing and toads co-occur in
meadows.
The available grazing studies focus on
breeding habitat (wet meadows) and do
not consider impacts to upland habitats.
The USFS grazing guidelines for
protection of meadow habitats of the
Yosemite toad include fencing breeding
meadows, but they do not necessarily
protect upland habitat. Martin (2008)
surveyed 11 meadow sites located along
a stream channel in or near low growing
willows both before and after cattle
grazed the entire meadow, and Martin
found that Yosemite toads could no
longer be located along the stream
channel after the vegetation was grazed.
However, both adults and subadults
could be found in dense willow thickets
or in parts of the meadow that were less
heavily grazed (Martin 2008, p. 298).
Grazing can also degrade or destroy
moist upland areas used as nonbreeding
habitat by Yosemite toads (Martin 2008,
p. 159), especially when nearby
meadow and riparian areas have been
fenced to exclude livestock. Livestock
may also collapse rodent burrows used
by Yosemite toads as cover and
hibernation sites (Martin 2008, p. 159)
or disturb toads and disrupt their
behavior. Martin (2008, pp. 305–306)
observed that grazing significantly
reduced vegetation height at grazed
meadow foraging sites, and since these
areas are not protected by current
grazing guidelines, deduced that cattle
grazing is having a negative effect on
terrestrial life stage survivorship in
Yosemite toads. This problem was
exacerbated as fenced areas effectively
shifted grazing activity to upland areas
actively used by terrestrial life stages of
the Yosemite toad (Martin 2008, p. 306).
Although we lack definitive data to
assess the link between Yosemite toad
population dynamics and habitat
degradation by livestock grazing
activity, in light of the documented
impacts to meadow habitats (including
effects on local hydrology) from grazing
activity in general, we consider this
threat prevalent with moderate impacts
to the Yosemite toad and a potential
limiting factor in population recovery
rangewide. In addition, given the
potential for negative impacts from
heavy use, and the vulnerability of toad
habitat should grazing management
practices change with new management
plans, we expect this threat to continue
into the future.
Roads and Timber Harvest Effects to
Meadow Habitat
Road construction and use, along with
timber harvest activity, may impact
Yosemite toad habitat via fragmentation,
ground disturbance, and soil
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compaction or erosion (Helms and
Tappeiner 1996, pp. 439–476). Roads
may alter both the physical environment
and the chemical environment; roads
may present barriers to movement and
may alter hydrologic and geomorphic
processes that shape aquatic systems,
while vehicle emissions and road-runoff
are expected to contain chemicals that
may be toxic (USFS et al. 2009, pp. 71–
73). Timber harvests and past
development of roads could potentially
also lead to increased rates of siltation,
contributing to the loss of breeding
habitats for the Yosemite toad.
Prior to the formation of National
Parks and National Forests, timber
harvest was widespread and
unregulated in the Sierra Nevada;
however, most cutting occurred below
the current elevation range of the
Yosemite toad (University of California
at Davis (UCD) UC 1996b, pp. 17–45;
USFS et al. 2009, p. 77). Between 1900
and 1950, most timber harvest occurred
in old-growth forests on private land
(UC 1996b, pp. 17–45). During this
period, forest plans often lacked
standards to protect riparian areas and
associated meadows, leading to harvest
activities that included cutting to edges
of riparian areas and forest road
construction that often crossed streams,
associated aquatic habitat, and
meadows, and resulted in head-cutting,
lowered water tables, and loss of
riparian habitats; legacies of these past
activities remain today (USFS et al.
2009, p. 77). Currently on National
Forests, timber harvest and related
vegetation management activities
overlap with Yosemite toads primarily
in the lower elevation portions of the
species’ range; the red fir and lodgepole
forests that generally surround highelevation meadows that are Yosemite
toad habitat do not have commercial
value (USFS et al. 2009, pp. 76, 77).
Forest standards and guidelines
currently provide protections for
riparian areas, such as buffers for timber
and vegetation management activities.
The majority of forest roads in
National Forests of the Sierra Nevada
were built between 1950 and 1990, to
support major increases in timber
harvest on National Forests, (USDA
2001a, p. 443), suggesting that many
forest roads occur at elevations below
the current range of the Yosemite toad.
Relatively few public roads, including
trans-Sierran State Highways 4 (Ebbetts
Pass), 88 (Carson Pass), 108 (Sonora
Pass), and 120 (Tioga Pass), cross the
high elevations of the Sierra Nevada
within the range of the Yosemite toad
(USFS et al. 2009, p. 71), although
smaller public roads are present in some
high-elevation areas. One percent of
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Yosemite toad populations occur on
private lands where urbanization and
corresponding construction of new
roads may be more likely (USFS et al.
2009, p. 71); however, we are not aware
of any proposals for new road
construction at this time.
We expect that the majority of timber
harvest, road development, and
associated management impacts (see
‘‘Effects of Fire Suppression on Meadow
Habitats’’ below) to Yosemite toad
habitat took place during the expansion
period in the latter half of the 20th
century. Using a model, Liang et al.
(2010, p. 16) found that Yosemite toads
were more likely to occur in areas closer
to timber activity, although the high
correlation between elevation and the
distance to harvest activity in model
results definitive conclusions regarding
cause and effect. However, they noted
that, because timber harvest activities
may maintain breeding sites by opening
the forest canopy and potentially
preventing encroachment of trees into
sites, breeding animals might benefit
from timber activity (Liang et al. 2010,
p. 16). Limited information from timber
sale areas where low-elevation
populations occur indicates that such
activities may negatively affect upland
habitat use if burrow sites are crushed
(USFS 2013, p. 6). Although grounddisturbance due to timber harvest
activities has the potential to have
population-level effects on Yosemite
toad habitat, especially where habitat is
limited, currently the best available
information does not indicate that the
current level of timber harvest occurring
within watersheds currently inhabited
by the Yosemite toad is adversely
affecting habitat (USFS et al. 2009, p.
77). Therefore the best available
scientific and commercial information
does not indicate that ongoing road
construction and maintenance or timber
harvest are significant threats to the
Yosemite toad. There may be localized
effects of these activities in areas where
legacy effects continue to result in
modified wet meadow habitat
conditions, or where current harvest
and road activities occur in close
proximity to extant Yosemite toad
populations.
Effects of Fire Suppression on Meadow
Habitats
Fire management refers to activities
over the past century to combat forest
fires. Historically, both lightning-caused
fires and fires ignited by American
Indians were regularly observed in
western forests (Parsons and Botti 1996,
p. 29), and in the latter 19th century, the
active use of fire to eliminate tree
canopy in favor of forage plants
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continued by sheepherders (Kilgore and
Taylor 1979, p. 139). Beginning in the
20th century, land management in the
Sierra Nevada shifted to focus on fire
suppression as a guiding policy (UC
2007, p. 10).
Long-term fire suppression has
influenced forest structure and altered
ecosystem dynamics in the Sierra
Nevada. In general, the time between
fires is now much longer than it was
historically, and live and dead fuels are
more abundant and continuous (USDA
2001a, p. 35). Much of the habitat for
the Yosemite toad occurs in highelevation meadows within wilderness
and backcountry areas where vegetation
is sparse and fire suppression activities
are rarely conducted (USFS et al. 2009,
p. 55), suggesting that fire suppression
has played a limited role in such
locations. At high elevations,
encroachment of lodgepole pine at
meadow edges has been attributed to
cessation of sheep grazing or legacy
effects of high-intensity grazing that
reduced water tables, as opposed to fire
suppression activities (Vankat and
Major 1978, pp. 392–395). At lower
elevations, it is not clear how habitat
changes attributed to fire suppression
have affected Yosemite toad
populations. However, Liang et al.
(2010, p. 16) observed that toads were
less likely to occur in areas where the
fire regime was significantly altered
from historical conditions, and
suggested that the toads are affected by
some unknown or unmeasured factors
related to fire management.
Evidence indicates that fire plays a
significant role in the evolution and
maintenance of lower elevation forested
meadows of the Sierra Nevada. Under
natural conditions, conifers are
excluded from meadows by fire and
saturated soils. Small fires thin and/or
destroy encroaching conifers, while
large fires are believed to determine the
meadow—forest boundary (Vankat and
Major 1978, p. 394; Parsons and
DeBenedetti 1979, pp. 29–31). Fire is
thought to be important in maintaining
open aquatic and riparian habitats for
amphibians in some systems (Russel et
al. 1999, pp. 374–384), and fire
suppression may have thereby
contributed to conifer encroachment on
meadows (Chang 1996, pp. 1071–1099;
NPS 2002, p. 1). However, fire
suppression effects are thought to vary
with ecosystem fire regime; variableinterval fires are characteristic of the
upper montane red fir forests (Chang
1996, pp. 107, 1072) that are the setting
for Yosemite toad habitat at the lower
elevations of its range, while longinterval fires are characteristic of the
subalpine lodgepole pine forests (Chang
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1996, p. 1072) that are the setting for
Yosemite toad habitats at higher
elevations. The effects of fire
suppression on forest structure is
thought to be far less important in the
longer interval forest types (Chang 1996,
p. 1072).
While no studies have confirmed a
link between fire suppression and
rangewide population decline of the
Yosemite toad, circumstantial evidence
to date suggests that historic fire
suppression may be a factor underlying
meadow encroachment at lower
elevations. The effect of fire
suppression, therefore, is thought to be
largely restricted to lower elevations
within the Yosemite toad’s range; fire
suppression activities are rarely
conducted where much of the habitat
for the Yosemite toad occurs (USFS et
al. 2009, pp. 51–54). Based on the best
available information, we find it likely
that habitat modification due to reduced
fire frequency is a moderate threat to
Yosemite toad in those lower-elevation
areas where fire suppression has
resulted in conifer encroachment into
meadows.
Recreation and Packstock Effects to
Meadow Habitat
Recreational activities take place
throughout the Sierra Nevada, and they
can have significant negative impacts on
wildlife and their habitats (USDA
2001a, pp. 221, 453–500). Recreation
can cause considerable impact to
vegetation and soils in western U.S.
Wilderness Areas and National Parks
even with light use, with recovery
occurring only after considerable
periods of non-use (USFS et al. 2009, p.
66). Heavy foot traffic in riparian areas
tramples vegetation, compacts soils, and
can physically damage streambanks.
Trails (foot, horse, bicycle, or offhighway motor vehicle) can compact the
soil, displace vegetation, and increase
erosion, thereby potentially lowering
the water table (Kondolph et al. 1996,
pp. 1009–1026). However, the National
Park Service considers current hiking
and backpacking activities to be a
negligible risk factor for the Yosemite
toad within the Parks. The Parks have
also worked to improve impacted
meadows by reconstructing poorly
designed trails that have degraded
meadow hydrology, also identifying
additional Yosemite toad meadows to
prioritize additional restoration
activities (NPS 2013, p. 9). Similar
activities have been implemented on
National Forests; for example, the Inyo
National Forest has re-routed several
trails to avoid the toad’s breeding
habitat (USFS 2013, p. 5).
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Although much Yosemite toad habitat
is located in wilderness or other
backcountry areas removed from
motorized access, the USFS has noted
locations where proximity of roads or
off-highway vehicle routes to Yosemite
toad breeding habitat has resulted in
observed impacts to Yosemite breeding
habitat. Off-highway vehicles are often
the first vehicles to pass through roads
blocked by winter snows, occasionally
driving off the road to pass remaining
obstacles (USFS et al. 2009, p. 63).
Records of such off-highway vehicle
travel in breeding meadows and ponds
(USFS 2013, pp. 6, 7) suggests that such
activities have the potential to
negatively affect these habitats, although
the population-level effects to Yosemite
toads are thought to be limited.
Packstock use has similar effects to
those discussed for livestock grazing (for
additional information on current
packstock use levels and management
protections, see the Packstock Use
section under the mountain yellowlegged frogs, above), although this risk
factor is potentially more problematic as
this land use typically takes place in
more remote and higher-elevation areas
occupied by Yosemite toads, and
packstock tend to graze in many of the
same locations that the toads prefer
(USFS et al. 2009, p. 65). Currently,
there are very few studies on the effects
of packstock grazing on amphibians,
especially in the Sierra Nevada.
However, in Yosemite, Sequoia, and
Kings Canyon National Parks, packstock
use is monitored annually to prevent
long-term impacts. Additionally, the
NPS (2013, p. 9) has indicated that,
except for a few specific areas,
packstock use and Yosemite toads
typically do not overlap within the
Parks. Many areas are closed to
packstock use entirely or limited to day
use due to inadequate trail access or to
protect sensitive areas. Long-term use
data indicate that packstock use is
declining, with no evidence to suggest
that it will increase in the future (NPS
2013, pp. 6, 7). Where permitted,
current guidelines in the National Parks
limit trips to 20–25 animals, regulated
under conditional use permits (Brooks
2012, pers. comm.). Similar standards
and guidelines limit packstock group
size and use within the National Forests
(USFS 2013, pp. 3–5).
Habitat-related effects of recreational
activities on the Yosemite toad may
have population-level impacts in
localized areas and under site-specific
conditions, for example, where foot
traffic adjacent to occupied meadows is
exceptionally heavy and results in
meadow damage, where legacy effects of
high recreation use have resulted in
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continuing meadow damage, or where
off-highway vehicle use results in
changes in meadow hydrology.
However, in general, we do not consider
habitat-related changes associated with
current levels of hiking or backpacking
to pose a population-level risk to
Yosemite toads. Therefore, at this time
we consider recreational activities to be
a low prevalence threat across the range
of the Yosemite toad.
Dams and Water Diversions Effects to
Meadow Habitat
Past construction of dams, diversion,
and irrigation ditches resulted in a vast
man-made network that altered local
and regional stream hydrology in the
Sierra Nevada (SNEP 1996, p. 120),
although, with the exception of several
dozen small impoundments and
diversions, almost all of these are
located below the range of the Yosemite
toad (USFS et al. 2009, pp. 76, 77).
However, in the past a small number of
reservoirs were constructed within the
historic range of the Yosemite toad,
most notably Upper and Lower Blue
Lakes, Edison, Florence, Huntington,
Courtright, and Wishon Reservoirs.
Construction of several high-elevation
reservoirs (for example, Edison and
Florence) is thought to have inundated
shallow-water breeding habitat for the
toad (USFS et al. 2009, pp. 76, 77).
Where reservoirs are used for
hydroelectric power, water-level
declines caused by drawdown of
reservoirs can lead to the mortality of
eggs and tadpoles by stranding and
desiccation, although, with the
exception of Blue Lakes, Yosemite toads
are currently not known from the above
reservoirs (USFS et al. 2009, pp. 78, 79).
Past construction of these reservoirs
likely contributed to the decline of the
Yosemite toad in the area where they
were built. Increasing effects from
climate change, or new water supply
development in response to such effects,
may exacerbate this risk in the future if
new reservoirs are constructed within
areas occupied by the toad. However,
we are not aware of any proposals to
construct additional reservoirs within
the Yosemite toads range. We expect
that continuing reservoir operations
may have continued habitat-related
effects to toad populations in these
developed areas, but less so in the
current extent of the Yosemite toad’s
(remnant) range. Therefore, we consider
this threat to be of low prevalence to the
Yosemite toad across its range.
Climate Effects to Meadow Habitat
Different studies indicate that
multiple drivers are behind the
phenomenon of conifer encroachment
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into meadows. The first factor affecting
the rate of conifer encroachment into
meadow habitats, fire suppression, was
discussed above. Climate variability is
another factor affecting the rate of
conifer encroachment on meadow
habitats. A study by Franklin et al.
(1971, p. 215) concluded that fire had
little influence on meadow maintenance
in their study area, while another study
concluded that climate change is a more
likely explanation for encroachment of
trees into the adjacent meadow at their
site, rather than fire suppression or
changes in grazing intensity (Dyer and
Moffett, 1999, p. 444).
Climatic variability is strongly
correlated with tree encroachment into
dry subalpine meadows (Jakubos and
Romme 1993, p. 382). In the Sierra
Nevada, most lodgepole pine seedlings
become established during years of low
snowpack when meadow soil moisture
is reduced (Wood 1975, p. 129). The
length of the snow-free period may be
the most critical variable in tree
invasion of subalpine meadows
(Franklin et al. 1971, p. 222), with the
establishment of a good seed crop,
followed by an early snowmelt,
resulting in significant tree
establishment. It is apparent that
periods of low snowpack and early melt
may in fact be necessary for seedling
establishment (Ratliff, 1985, p. 35).
Millar et al. (2004, p. 181) reported that
increased temperature, coupled with
reduced moisture availability in relation
to large-scale temporal shifts in climate,
facilitated the invasion of 10 subalpine
meadows studied in the Sierra Nevada.
Our analyses under the Act include
consideration of ongoing and projected
changes in climate. The terms ‘‘climate’’
and ‘‘climate change’’ are defined by the
Intergovernmental Panel on Climate
Change (IPCC). ‘‘Climate’’ refers to the
mean and variability of different types
of weather conditions over time, with 30
years being a typical period for such
measurements, although shorter or
longer periods also may be used (IPCC
2007, p. 1450; IPCC 2013a, Annex III).
The term ‘‘climate change’’ thus refers
to a change in the mean or variability of
one or more measures of climate (for
example, temperature or precipitation)
that persists for an extended period,
typically decades or longer, whether the
change is due to natural variability,
human activity, or both (IPCC 2007, p.
1450; IPCC 2013a, Annex III). A recent
compilation of climate change and its
effects is available from reports of the
Intergovernmental Panel on Climate
Change (IPCC) (IPCC 2013b, entire).
Various types of changes in climate can
have direct or indirect effects on
species. These effects may be positive,
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neutral, or negative, and they may
change over time, depending on the
species and other relevant
considerations, such as the effects of
interactions of climate with other
variables (for example, habitat
fragmentation) (IPCC 2007, pp. 8–14,
18–19). In our analyses, we use our
expert judgment to weigh relevant
information, including uncertainty, in
our consideration of various aspects of
climate change.
For the Sierra Nevada ecoregion,
climate models predict that mean
annual temperatures will increase by 1.8
to 2.4 °C (3.2 to 4.3 °F) by 2070,
including warmer winters with earlier
spring snowmelt and higher summer
temperatures (PRBO 2011, p. 18).
Additionally, mean annual rainfall is
projected to decrease from the current
average by some 9.2–33.9 cm (3.6–13.3
in) by 2070 (PRBO 2011, p. 18).
However, projections have high
uncertainty, and one study predicts the
opposite effect (PRBO 2011, p. 18).
Snowpack is, by all projections, going to
decrease dramatically (following the
temperature rise and increase in
precipitation falling as rain) (PRBO
2011, p. 19); (Kadir et al. 2013, pp. 76–
80). Higher winter stream flows, earlier
runoff, and reduced spring and summer
stream flows are projected, with
increasing severity in the southern
Sierra Nevada (PRBO 2011, pp. 20–22);
(Kadir et al. 2013, pp. 71–75).
Snow-dominated elevations from
2,000–2,800 m (6,560–9,190 ft) will be
the most sensitive to temperature
increases (PRBO 2011, p. 23). Meadows
fed by snowmelt may dry out or be more
ephemeral during the non-winter
months (PRBO 2011, p. 24). This pattern
could influence groundwater transport,
and springs may be similarly depleted,
leading to lower water levels in
available breeding habitat and decreased
area and hydroperiod (i.e., duration of
water retention) of suitable habitat for
rearing tadpoles of Yosemite toads.
Changes in water transport may promote
channel incision and result in a shift to
non-meadow conditions (Viers et al.
2013, p. 31).
Blaustein et al. (2010, pp. 285–300)
provide an exhaustive review of
potential direct and indirect and
habitat-related effects of climate change
to amphibian species, with
documentation of effects in a number of
species where such effects have been
studied. Altitudinal range shifts with
changes in climate have been reported
in some regions. They note that
temperature can influence the
concentration of dissolved oxygen in
aquatic habitats, with warmer water
generally having lower concentrations
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of dissolved oxygen, and that water
balance heavily influences amphibian
physiology and behavior. They predict
that projected changes in temperature
and precipitation are likely to increase
habitat loss and alteration for those
species living in sensitive habitats, such
as ephemeral ponds and alpine habitats
(Blaustein et al. 2010, pp. 285–287).
Because environmental cues such as
temperature and precipitation are
clearly linked to onset of reproduction
in many species, climate change will
likely affect the timing of reproduction
in many species, potentially with
different sexes responding differently to
climate change. For example, males of
two newt species (Triturus spp.) showed
a greater degree of change in arrival date
at breeding ponds (Blaustein et al. 2010,
p. 288). Lower concentrations of
dissolved oxygen in aquatic habitats
may negatively affect developing
embryos and larvae, in part because
increases in temperature increase the
oxygen consumption rate in
amphibians. Reduced oxygen
concentrations have also been shown to
result in accelerated hatching in ranid
frogs, but at a smaller size, while larval
development and behavior may also be
affected and may be mediated by larval
density and food availability (Blaustein
et al. 2010, pp. 288–289).
Increased temperatures can reduce
time to metamorphosis, which can
increase chances of survival where
ponds dry, but also result in
metamorphosis at a smaller size,
suggesting a likely trade-off between
development and growth, which may be
exacerbated by climate change and have
fitness consequences for adults
(Blaustein et al. 2010, pp. 289–290).
Changes in terrestrial habitat, such as
changed soil moisture and vegetation,
can also directly affect adult and
juvenile amphibians, especially those
adapted to moist forest floors and cool,
highly oxygenated water that
characterizes montane regions. Climate
change may also interact with other
stressors that may be acting on a
particular species, such as disease and
contaminants (Blaustein et al. 2010, pp.
290–299).
A recent paper (Kadir et al. 2013,
entire) provides specific information on
the effects of climate change in the
Sierra Nevada. The report found that
glaciers in the Sierra Nevada have
decreased in area over the past century,
and glacier shrinkage results in earlier
peak water runoff and drier summer
conditions. Another result from the
report is that the lower edge of the
conifer-dominated forests in the Sierra
Nevada has been retreating upslope over
the past 60 years. Regarding wildfire,
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since 1950, annual acreage burned in
wildfires statewide has been increasing
in California, and in the western United
States, large wildfires have become
more frequent, increasing in tandem
with rising spring and summer
temperatures. Finally, the report found
that today’s subalpine forests in the
Sierra Nevada are much denser—that is,
comprise more small-diameter trees—
than they were over 70 years ago.
During this time period, warmer
temperatures, earlier snowmelt, and
more rain than snow occurred in this
region. Many of these changes in the
Sierra Nevada of California due to
climate are likely to influence Yosemite
toads because they are highly vulnerable
to climate change because changing
hydrology and habitat in the Sierra
Nevada will likely have impacts on
remaining populations (Viers et al.
2013, pp. 55, 56).
Historically, drought is thought to
have contributed to the decline of the
Yosemite toad (Kagarise Sherman and
Morton 1993, p. 186; Jennings and
Hayes 1994, pp. 50–53). Extended and
more severe droughts pose an ongoing,
rangewide risk to the species and are
expected to increase with predicted
climate changes (PRBO 2011, p. 18).
Such changes may reduce both the
amount of suitable breeding habitat and
the length of time that suitable water is
available in that habitat (Brown 2013,
unpaginated).
Davidson et al. (2002, p. 1598)
analyzed geographic decline patterns for
the Yosemite toad. They compared
known areas of extirpation against a
hypothesized model for climate change
that would predict greater numbers of
extirpations at lower altitudes, and in
more southern latitudes. The
researchers did not observe a pattern in
the available historic data to support the
climate change hypothesis as a driver of
historic population losses, although
they acknowledge that climate change
may be a contributor in more complex
or subtle ways. Additionally, this study
was limited by small sample size, and
it is possible that climate change effects
on the Yosemite toad (a long-lived
species) may not become evident for
many years (USFS et al. 2009, p. 48).
Finally, Davidson et al. (2002, p. 1598)
did find an increase in occupancy with
elevation (greater densities of
populations at altitude), and this
observation is consistent with a pattern
that would fit a response to climate
change (USFS et al. 2009, p. 48).
However, this observation would also be
consistent if the features of these
particular habitats (such as at higher
elevation) were more suited to the
special ecological requirements of the
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toad, or if other stressors acting on
populations at lower elevations were
responsible for the declines. We,
therefore, find these results
inconclusive.
Most recently, modeled vulnerability
assessments for Sierra Nevada montane
meadow systems have utilized life
history and habitat requirements to
gauge vulnerability of amphibian
species to climate change. This
assessment indicates that vulnerability
to hydro-climatic changes will likely be
very high for the Yosemite toad, and
that continued or worsening stream
channelization in montane meadows
from flashy storms may worsen effects
by further reductions in the water table
(Viers et al. 2013, p. 56).
The breeding ecology and life history
of the Yosemite toad are that of a habitat
specialist, as it utilizes pool and
meadow habitats during the onset of
snowmelt and carefully times its
reproduction to fit available conditions
within ephemeral breeding sites. The
most striking documented declines in
Yosemite toad populations in the
historical record are correlated with
extreme climate episodes (drought)
(Kagarise Sherman and Morton 1993,
pp. 186–198). Given these observations,
it is likely that climate change (see also
discussion in mountain yellow-legged
frog’s Summary of Factors Affecting the
Species, under Factor E) poses a
significant risk to the Yosemite toad
now and in the future. It is quite
possible that these impacts are
occurring currently, and have occurred
over the last few decades. However, it
is difficult in short time intervals to
discern the degree of effect from climate
change within the variability of natural
climate cycles.
In summary, based on the best
available scientific and commercial
information, we consider the threats of
destruction, modification, and
curtailment of the species’ habitat and
range to be significant ongoing threats to
the Yosemite toad. The legacy effects of
past land uses have altered meadow
communities through the mechanism of
stream incision by permanently
reducing habitat quantity and quality
unless active and costly restoration is
implemented. Climate change is a
current threat of high magnitude.
Threats considered of moderate
magnitude include livestock grazing and
fire management regime. Threats
considered currently low magnitude
include roads and timber harvest, dams
and water diversions, and recreational
land uses.
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Factor B. Overutilization for
Commercial, Recreational, Scientific, or
Educational Purposes
We do not have any scientific or
commercial information to indicate that
overutilization for commercial,
recreational, or scientific purposes
poses a threat to the Yosemite toad.
There is currently no known
commercial market for Yosemite toads,
although one pet store in Los Angeles
that is no longer in business had
previously sold at least one Yosemite
toad (USFS et al. 2009, pp. 65–66); and
there is also no documented recreational
or educational use for Yosemite toads.
Scientific research may cause some
stress to Yosemite toads through
disturbance and disruption of behavior,
handling, and injuries associated with
marking individuals. This activity has
resulted in the known death of
individuals through accidental
trampling (Green and Kagarise Sherman
2001, pp. 92–103), irradiation from
radioactive tags (Karlstrom 1957, pp.
187–195), and collection for museum
specimens (Jennings and Hayes 1994,
pp. 50–53). We expect that requirements
for Federal (USFS and NPS) and State
(CDFW) research and special use
permits, and University ethics
requirements provide some protections
for wildlife-research subjects and limit
negative effects to individuals.
Therefore, we do not currently consider
ongoing and future scientific research to
be a threat to the Yosemite toad. We also
anticipate that further research into the
genetics and life history of the Yosemite
toad and broader methodological
censuses will provide a net conservation
benefit to this under-studied species.
Based on the best available scientific
and commercial information, we do not
consider overutilization for commercial,
recreational, scientific, or educational
purposes to be a threat to the Yosemite
toad.
Factor C. Disease or Predation
Predation
Prior to the trout stocking of high
Sierra Nevada lakes, which began over
a century ago, fish were entirely absent
from most of this region (Bradford 1989,
pp. 775–778). Observations regarding
the effects of introduced fishes on the
Yosemite toad are mixed. However, resurveys of historical Yosemite toad sites
have shown that the species has
disappeared from several lakes where
they formerly bred, and these areas are
now occupied by fish (Stebbins and
Cohen 1995, pp. 213–215; Martin 2002,
p. 1).
Drost and Fellers (1994, pp. 414–425)
suggested that Yosemite toads are less
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vulnerable to fish predation than frogs
because they breed primarily in
ephemeral waters that do not support
fish. Further, Jennings and Hayes (1994,
pp. 50–53) stated that the palatability of
Yosemite toad tadpoles to fish predators
is unknown, but often assumed to be
low based on the unpalatability of
western toads (Drost and Fellers 1994,
pp. 414–425; Kiesecker et al. 1996, pp.
1237–1245), to which Yosemite toads
are closely related. Grasso (2005, p. 1)
observed brook trout swimming near,
but the trout ignored Yosemite toad
tadpoles, suggesting that tadpoles are
unpalatable. The study also found that
subadult Yosemite toads were not
consumed by brook trout (Grasso 2005,
p. 1), although the sublethal effects of
trout ‘‘sampling’’ (mouthing and
ejecting tadpoles) and the palatability of
subadults to other trout species are
unknown. Martin (2002, p. 1) observed
brook trout preying on Yosemite toad
tadpoles, and also saw them ‘‘pick at’’
Yosemite toad eggs (which later became
infected with fungus). In addition,
metamorphosed western toads have
been observed in golden trout stomach
contents (Knapp 2002c, p. 1).
Nevertheless, Grasso et al. (2010, p. 457)
concluded that early life stages of the
Yosemite toad likely possess chemical
defenses that provide sufficient
protection from native trout predation.
The observed predation of Yosemite
toad tadpoles by trout (Martin 1992, p.
1) indicates that introduced fishes may
pose a predation risk to the species in
some situations, which may be
accentuated during drought years. At a
site where Yosemite toads normally
breed in small meadow ponds, they
have been observed to successfully
switch breeding activities to stream
habitat containing fish during years of
low water (Strand 2002, p. 1). Thus,
drought conditions may increase the
toads’ exposure to predatory fish, and
place them in habitats where they
compete with fish for invertebrate prey.
Additionally, although the number of
lake breeding sites used by Yosemite
toads is small relative to the number of
ephemeral sites, lake sites may be
especially important because they are
more likely to be habitable during years
with low water (Knapp 2002c, p. 1).
Overall, the data and available
literature suggest that direct mortality
from fish predation is likely not an
important factor driving Yosemite toad
population dynamics. This does not
discount other indirect impacts, such as
the possibility that fish may be effective
disease vectors (see below). Yosemite
toad use of more ephemeral breeding
habitats (which are less habitable to fish
species as they cannot tolerate drying or
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freezing) minimizes the interaction of
fish and toad tadpoles. Further, where
fish and toads co-occur, it is possible
that food depletion (outcompetition) by
fish negatively affects Yosemite toads
(USFS et al. 2009, p. 58).
Other predators may also have an
effect on Yosemite toad populations.
Kagarise Sherman and Morton (1993, p.
194) reported evidence of toad
predation by common ravens (Corvus
corax) and concluded this activity was
responsible for the elimination of toads
from one site. These researchers also
confirmed, as reported in other studies,
predation on Yosemite toad by Clark’s
nutcrackers (Nucifraga columbiana).
The significance of avian predation may
increase if the abundance of common
ravens within the current range of the
Yosemite toad increases as it has in
nearby regions (Camp et al. 1993, p.
138; Boarman et al. 1995, p. 1; Kelly et
al. 2002, p. 202). However, the degree to
which avian predation may be affecting
Yosemite toad populations has not been
quantified.
Disease
Although not all vectors have been
confirmed in the Sierra Nevada,
introduced fishes, humans, pets,
livestock, packstock, vehicles, and wild
animals may all act to facilitate disease
transmission between amphibian
populations. Infection of both fish and
amphibians by a common disease has
been documented with viral (Mao et al.
1999, pp. 45–52) and fungal pathogens
in the western United States (Blaustein
et al. 1994b, pp. 251–254). Mass die-offs
of amphibians in the western United
States and around the world have been
attributed to Bd fungal infections of
metamorphs and adults (Carey et al.
1999, pp. 1–14), Saprolegnia fungal
infections of eggs (Blaustein et al.
1994b, pp. 251–254), ranavirus
infections, and bacterial infections
(Carey et al. 1999, pp. 1–14).
Various diseases are confirmed to be
lethal to Yosemite toads (Green and
Kagarise Sherman 2001, pp. 92–103),
and recent research has elucidated the
potential role of Bd infection as a threat
to Yosemite toad populations (Dodge
and Vredenburg 2012, p. 1). These
various diseases and infections, in
concert with other factors, have likely
contributed to the decline of the
Yosemite toad (Kagarise Sherman and
Morton 1993, pp. 193–194) and may
continue to pose a risk to the species
(Dodge and Vredenburg 2012, p. 1).
Die-offs in Yosemite toad populations
have been documented in the literature,
and an interaction with diseases in
these events has been confirmed.
However, no single cause has been
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validated by field studies. Tissue
samples from dead or dying adult
Yosemite toads and healthy tadpoles
were collected during a die-off at Tioga
Pass Meadow and Saddlebag Lake and
analyzed for disease (Green and
Kagarise Sherman 2001, pp. 92–103).
Six infections were found in the adults,
including infection with Bd, bacillary
bacterial septicemia (red-leg disease),
Dermosporidium (a fungus), myxozoa
spp. (parasitic cnidarians), Rhabdias
spp. (parasitic roundworms), and
several species of trematode (parasitic
flatworms). Despite positive detections,
no single infectious disease was found
in more than 25 percent of individuals,
and some dead toads showed no signs
of infection to explain their death.
Further, no evidence of infection was
found in tadpoles. A meta-analysis of
red-leg disease also revealed that the
disease is a secondary infection that
may be associated with a suite of
different pathogens, and so actual
causes of decline in these instances
were ambiguous (Kagarise Sherman and
Morton 1993, p. 194). The authors
concluded that the die-off was caused
by suppression of the immune system
caused by an undiagnosed viral
infection or chemical contamination
that made the toads susceptible to the
variety of diagnosed infections.
Saprolegnia ferax, a species of water
mold that commonly infects fish in
hatcheries, caused a massive lethal
infection of eggs of western toads at a
site in Oregon (Blaustein et al. 1994b, p.
252). It is unclear whether this event
was caused by the introduction of the
fungal pathogen via fish stocking, or if
the fungus was already present and the
eggs’ ability to resist infection was
inhibited by some unknown
environmental factor (Blaustein et al.
1994b, p. 253). Subsequent laboratory
experiments have shown that the fungus
could be passed from hatchery fish to
western toads (Kiesecker et al. 2001, pp.
1064–1070). Fungal growth on Yosemite
toad eggs has been observed in the field,
but the fungus was not identified and it
was unclear whether the fungus was the
source of the egg mortality (Kagarise
Sherman 1980, p. 46). Field studies
conducted in Yosemite National Park
found that an undetermined species of
water mold infected only the egg masses
that contained dead embryos of
Yosemite toads (Sadinski 2004, pp. 33–
34). The researchers also observed that
the water mold became established on
egg masses only after embryo death, and
subsequently spread, causing the
mortality of additional embryos of
Yosemite toads.
Sadinski (2004, p. 35) discovered that
mortality of Yosemite toad embryos may
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be attributed to an unidentified species
of a free-living flatworm (Turbellaria
spp.). In Yosemite National Park, these
worms were observed to penetrate
Yosemite toad egg masses and feed
directly on the embryos. In some
locations, Turbellaria spp. reached such
large densities that they consumed all
the embryos within a Yosemite toad egg
mass. Predation also facilitated the
colonization and spread of water mold
on egg masses, leading to further
embryo mortality. Further studies
would be needed to determine which
species of Turbellaria feeds on Yosemite
toad eggs, and the extent of this impact
on Yosemite toad populations.
Until recently, the contribution of Bd
infection to Yosemite toad population
declines was relatively unknown.
Although the toad is hypothetically
susceptible due to co-occurrence with
the mountain yellow-legged frog, the
spread and growth of Bd in the warmer
pool habitats, occupied for a much
shorter time relative to the frog, is
suspected to render individuals less
prone to epidemic outbreaks (USFS et
al. 2009, p. 50). Fellers et al. (2011, p.
391) documented the occurrence of Bd
infection in Yosemite National Park
toads over at least a couple of decades,
and they note population persistence in
spite of the continued presence of the
pathogen. In a survey of 196 museum
specimens, Dodge and Vredenburg
(2012, p. 1) report the first presence of
Bd infection in Yosemite toads
beginning in 1961, with the pathogen
becoming highly prevalent during the
recorded declines of the late 1970s,
before it peaked in the 1990s at 85
percent positive incidence. In live
specimen sampling, Dodge and
Vredenburg (2012, p. 1) collected 1,266
swabs of Yosemite toads between 2006
and 2011, and found Bd infection
intensities at 17–26 percent (with
juvenile toads most affected). The
studies detected a pattern indicative of
the historic emergence of Bd, which
coincided with the documented decline
in Yosemite toad (Dodge 2013, p. 1). As
such, results from these studies support
the hypothesis that Bd infection and
chytridiomycosis have played an
important role in Yosemite toad
population dynamics over the period of
their recent recorded decline.
Carey (1993, pp. 355–361) developed
a model to explain the disappearance of
boreal toads (Bufo boreas boreas) in the
Rocky Mountains, suggesting immune
system suppression from extreme winter
stress (‘‘winter stress syndrome’’) could
have contributed to the decline in that
species. This model may also fit
Yosemite toad die-offs observed by
Kagarise Sherman and Morton (1993,
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pp. 186–198), given the close
relationship between the two toads, and
their occupation of similar habitats.
However, an analysis of immune system
suppression and the potential role of
winter stress relative to Yosemite toad
population trends is not available at this
time. Yet, the decline pattern observed
in the Carey study is mirrored by the
pattern in the Yosemite toad (heavy
mortality exhibited in males first)
(Knapp 2012, pers. comm.). This
observation, in concert with the recent
results from museum swabs (Dodge and
Vredenburg 2012, p. 1), provides a
correlative link to the timing of the
recorded Yosemite toad declines and Bd
infection intensities.
Although disease as a threat factor to
the Yosemite toad is relatively less
documented, Bd infection causes mass
mortalities in the closely related boreal
toad (Carey et al. 2006, p. 19) and there
is evidence related to Bd’s role in
historical die-offs in Yosemite toads.
Much of the historic research
documenting Yosemite toad declines
predated our awareness of Bd as a major
amphibian pathogen. Additionally, the
life history of the Yosemite toad, as a
rapid breeder during early snowmelt,
limits the opportunities to observe
population crashes in the context of
varied environmental stressors.
Currently available evidence indicates
that Bd was likely a significant factor
contributing to the recent historical
declines observed in Yosemite toad
populations (Dodge and Vredenburg
2012, p. 1). Although infection
intensities are currently lower than
some peak historic measurements, this
threat remains a potential factor that
may continue to reduce survival
through metamorphosis, and therefore
recruitment to the breeding population
(Knapp 2012, pers. comm.).
Additionally, the interaction of disease
and other stressors, such as climate
extremes, is not well understood in the
Yosemite toad. Research does suggest
that the combination of these threats
represents a factor in the historical
decline of the species (Kagarise
Sherman and Morton 1993, p. 186).
In summary, based on the best
available scientific and commercial
information, we do not consider
predation to be a threat to the species.
We consider disease to be a threat to the
Yosemite toad that has a moderate,
ongoing effect on populations of the
species rangewide. The threat most
specifically includes the amphibian
pathogen, Bd. Although definitive
empirical data quantifying the
contribution of disease to Yosemite toad
population declines are not currently
available, population declines that were
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concurrent with the prevalence and
spread of Bd across the Sierra Nevada
support the assertion that disease has
played a role in the observed trend.
Further, Bd infection, even at lower
intensities, may interact with climate
extremes and continue to depress
recruitment of yearling and subadult
Yosemite toads to breeding Yosemite
toad populations. We suspect this threat
was historically significant, that it is
currently having a moderate influence
on toad populations, and we expect it to
be a future concern.
Factor D. The Inadequacy of Existing
Regulatory Mechanisms
In determining whether the
inadequacy of regulatory mechanisms
constitutes a threat to the Yosemite
toad, we analyzed the existing Federal
and State laws and regulations that may
address the threats to the species or
contain relevant protective measures.
Regulatory mechanisms are typically
nondiscretionary and enforceable, and
may preclude the need for listing if such
mechanisms are judged to adequately
address the threat(s) to the species such
that listing is not warranted. Conversely,
threats on the landscape are not
addressed by existing regulatory
mechanisms where the existing
mechanisms are not adequate (or not
adequately implemented or enforced).
We discussed the applicable State and
Federal laws and regulations, including
the Wilderness Act, NFMA above (see
Factor D discussion for mountain
yellow-legged frogs). In general, the
same administrative policies and
statutes are in effect for the Yosemite
toad. This section additionally
addresses regulatory mechanisms with a
specific emphasis on the Yosemite toad.
Taylor Grazing Act of 1934
In response to overgrazing of available
rangelands by livestock from the 1800s
to the 1930s, Congress passed the Taylor
Grazing Act in 1934 (43 U.S.C. 315 et
seq.). This action was an effort to stop
the damage to the remaining public
lands as a result of overgrazing and soil
depletion, to provide coordination for
grazing on public lands, and to attempt
to stabilize the livestock industry
(Meehan and Platts 1978, p. 275; Public
Lands Council et al. v. Babbitt Secretary
of the Interior et al. (167 F. 3d 1287)).
Passage of the Taylor Grazing Act
resulted in reduced grazing in some
areas, including the high Sierra Nevada.
However, localized use remained high,
precluding regeneration of many
meadow areas (Beesley 1996, p. 14;
Menke et al. 1996, p. 14; Public Lands
Council et al. v. Babbitt Secretary of the
Interior et al. (167 F. 3d 1287)).
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Existing Federal and State laws and
regulatory mechanisms currently offer
some level of protection for the
Yosemite toad. Specifically, these
include the Wilderness Act, the NFMA,
the SNFPA, and the FPA (see Factor D
discussion for mountain yellow-legged
frog complex). Based on the best
available scientific and commercial
information, we do not consider the
inadequacy of existing regulatory
mechanisms to be a threat to the
Yosemite toad.
Factor E. Other Natural or Manmade
Factors Affecting Its Continued
Existence
The Yosemite toad is sensitive to
environmental change or degradation
due to its life history, biology, and
existence in ephemeral habitats
characterized by climate extremes and
low productivity. It is also sensitive to
anthropogenically influenced factors.
For example, contaminants, acid
precipitation, ambient ultraviolet
radiation, and climate change have been
implicated as contributing to amphibian
declines (Corn 1994, pp. 62–63; Alford
and Richards 1999, pp. 2–7). However,
as with the case with the mountain
yellow-legged frog complex,
contaminants, acid precipitation, and
ambient ultraviolet radiation are not
known to pose a threat (current or
historical) to Yosemite toad and,
therefore, are not discussed further.
Please refer to the proposed listing rule
for the Sierra Nevada yellow-legged
frog, the northern DPS of the mountain
yellow-legged frog, and the Yosemite
toad (78 FR 24472, April 25, 2013) for
a detailed discussion of contaminants,
acid precipitation, and ambient
ultraviolet radiation. The following
discussion will focus on potential threat
factors specifically studied in the
Yosemite toad, based on the unique life
history, population status,
demographics, or biological factors
specific to Yosemite toad populations.
Climate Change Effects on Individuals
As discussed above in Factor A,
climate change can result in detrimental
impacts to Yosemite toad habitat.
Climate variability could also negatively
impact populations through alteration of
the frequency, duration, and magnitude
of either droughts or severe winters
(USFS et al. 2009, p. 47). Yosemite toads
breed and their tadpoles develop in
shallow meadow and ephemeral
habitats, where mortality from
desiccation and freezing can be very
high, often causing complete loss of an
annual cohort (USFS et al. 2009, p. 10).
Kagarise Sherman and Morton (1993,
pp. 192–193) documented in a long-
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term population study that Yosemite
toad hatching success and survival were
subject to a balance between the
snowpack water contribution to
breeding pools and the periodicity and
character of breeding season storms and
post-breeding climate (whether it is cold
or warm). When it is too cold, eggs and
tadpoles are lost to freezing. This
situation poses a risk as earlier
snowmelt is expected to cue breeding
earlier in the year, exposing young
tadpoles (or eggs) to killing frosts in
more variable conditions of early spring
(Corn 2005, p. 60). When it is too dry,
tadpoles are lost to pool desiccation.
Alterations in the annual and seasonal
hydrologic cycles that influence water
volume and persistence in Yosemite
toad breeding areas can thereby impact
breeding success. The threat of climate
change on individuals is significant, and
is of high prevalence now and into the
future.
Other Sources of Direct and Indirect
Mortality
Direct and indirect mortality of
Yosemite toads has occurred as a result
of livestock grazing. Mortality risk from
livestock trampling is expected to be the
greatest for non-larval stages where
livestock concentrate in Yosemite toad
habitat when toad densities are highest;
early in the season when breeding
adults are aggregated and egg masses are
laid; and at metamorphosis when
juveniles are metamorphosing in mass
along aquatic margins. However,
because cattle typically are not present
during the breeding season, the risk of
trampling is expected to be greatest for
metamorphs (USFS et al. 2009, p. 59).
Cattle have been observed to trample
Yosemite toad metamorphs and
subadult toads, and these life stages can
fall into deep hoofprints and die (Martin
2008, p. 158). Specifically, Martin
(2008, p. 158) witnessed some 60
subadult and metamorph toad deaths
during the movement of 25 cattle across
a stream channel bordered by willows
within a meadow complex. Adult
Yosemite toads trampled to death by
cattle have also been observed (Martin
2002, pp. 1–3). This risk factor is likely
of sporadic significance, and is of
greatest concern where active grazing
allotments coincide with breeding
meadows. However, it is difficult to
determine the degree of this impact
without quantitative data.
Trampling and collapse of rodent
burrows by recreationists, pets, and
vehicles could lead to direct mortality of
terrestrial life stages of the Yosemite
toad. Recreational activity may also
disturb toads and disrupt their behavior
(Karlstrom 1962, pp. 3–34). Recreational
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anglers may be a source of introduced
pathogens and parasites, and they have
been observed using toads and tadpoles
as bait (USFS et al. 2009, p. 66).
However, Kagarise Sherman and Morton
(1993, p. 196) did not find a relationship
between the distance from the nearest
road and the declines in their study
populations, suggesting that human
activity was not the cause of decline in
that situation. Recreational activity may
be of conservation concern, and this
threat may increase with greater activity
in mountain meadows. However,
current available information does not
indicate that recreational activity is a
significant stressor for Yosemite toads.
Fire management practices over the
last century have created the potential
for severe fires in the Sierra Nevada.
Wildfires do pose a potential direct
mortality threat to Yosemite toads,
although amphibians in general are
thought to retreat to moist or
subterranean refuges and thereby suffer
low mortality during natural fires
(Russel et al. 1999, pp. 374–384). In the
closely related boreal toad (Bufo
boreas), Hossak and Corn (2007, p.
1409) documented a positive response
(increase in occupied breeding sites and
population size) following a wildfire,
with returns to near pre-fire occupancy
levels after 4 to 5 years (Hossack et al.
2012, p. 224), suggesting that habitatrelated changes associated with
wildfires may provide at least shortterm benefits to Yosemite toad
populations. However, data on the
direct and indirect effects of fire on
Yosemite toads are lacking.
USFS et al. (2009, p. 74) suggested
that the negative effects of roads that
have been documented in other
amphibians, in concert with the
substantial road network across a
portion of the Yosemite toad’s range,
indicate this risk factor may be
potentially significant to the species.
Roads may facilitate direct mortality of
amphibians through vehicle strikes
(DeMaynadier and Hunter 2000, pp. 56–
65), and timber harvest activities
(including fuels management and
vegetation restoration activities) have
been documented to result in the direct
mortality of Yosemite toads (USFS 2013,
p. 94). Levels of timber harvest and road
construction have declined substantially
since implementation of the California
Spotted Owl Sierran Province Interim
Guidelines in 1993, and some existing
roads have been decommissioned or are
scheduled to be decommissioned
(USDA 2001a, p. 445). Therefore, the
risks posed by new roads and timber
harvests have declined, but those
already existing still may pose risks to
the species and its habitat.
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Toads could potentially be trampled
or crushed by activities implemented to
reduce fire danger. USFS et al. (2009, p.
53) report that the Forest Service has
initiated a fuels reduction program in
order to reduce the extent and intensity
of wildfires. However, most of these
projects will occur in the Wildland
Urban Interface, which is below the
elevational range of the Yosemite toad
and generally near human
developments. However, in the future
some fuels projects may occur in limited
areas around facilities, such as resorts,
pack stations, or summer homes, within
the lowest portion of the Yosemite toad
range.
Collectively, direct mortality from
land uses within the Yosemite toad
range may have impacts to the toad.
However, we are aware of no studies
that have quantified or estimated the
prevalence of this particular threat to be
able to assess its impact to Yosemite
toad populations. At the current time,
direct and indirect mortality from roads
are not considered to be a significant
factor affecting the Yosemite toad
rangewide.
Small Population Size
Although it is believed that the range
of the Yosemite toad has not
significantly contracted, the majority of
populations across this area have been
extirpated, and this loss has been
significant relative to the historical
condition (multitudes of populations
within many watersheds across their
geographic range) (see ‘‘Population
Estimates and Status’’ above). Further,
growing evidence suggest that the
populations that remain are small,
numbering fewer than 20 males in most
cases (Kagrise Sherman and Morton
1993, p. 190; Sadinski 2004, p. 40;
Brown et al. 2012, p. 125). This
situation renders these remnant
populations susceptible to risks
inherent to small populations (see
Factor E discussion, ‘‘Small Population
Size,’’ for mountain yellow-legged frogs,
above) including inbreeding depression
and genetic drift, along with a higher
probability of extirpation from
unpredictable events such as severe
storms or extended droughts.
Traill et al. (2009, p. 32) argued for a
benchmark viable population size of
5,000 adult individuals (and 500 to
prevent inbreeding) for a broad range of
taxa, although this type of blanket figure
has been disputed as an approach to
conservation (Flather et al. 2011, pp.
307–308). Another estimate, specific to
amphibians, is that populations of at
least 100 individuals are less
susceptible to demographic stochasticity
(Schad 2007, p. 10). Amphibian species
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with highly fluctuating population size,
high frequencies of local extinctions,
and living in changeable environments
may be especially susceptible to
curtailment of dispersal and restriction
of habitat (Green 2003, p. 331). These
conditions are all likely applicable to
the Yosemite toad.
Therefore, based on the best available
commercial and scientific information,
we conclude that small population size
is a prevalent and significant threat to
the species viability of the Yosemite
toad across its range, especially in
concert with other extant stressors (such
as climate change).
Cumulative Impacts of Extant Threats
Interactive effects or cumulative
impacts from multiple additive stressors
acting upon Yosemite toad populations
over time are indicated by the
documented declines in populations
and abundance across the range of the
species. Although no single causative
factor linked to population declines in
Yosemite toads has been confirmed in
the literature (excepting perhaps
extreme climate conditions such as
droughts) (Kagarise Sherman and
Morton 1993, p. 186; Jennings and
Hayes 1994, pp. 50–53), there has been
a decline in population abundance and
numbers of extant populations
inhabiting the landscape (Brown et al.
2012, pp. 115–131; Kagarise Sherman
and Morton 1993, pp. 186–198). This
pattern of decline suggests a factor or
combination of factors common
throughout the range of the toad. The
available literature (Kagarise Sherman
and Morton 1993, pp. 186–198; Jennings
and Hayes 1994, pp. 50–53; USFS et al.
2009, pp. 1–133; Martin 2008, pp. i–
393) supports the contention that a
combination of factors has interacted
and is responsible for the decline
observed in Yosemite toad populations
over the past few decades.
Disease has been documented in
Yosemite toad populations, and recent
data documenting historic trends in Bd
infection intensity are compelling
(Dodge and Vredenburg 2012, p. 1), but
disease has not been definitively tied to
the observed rangewide decline. There
is considerable evidence that various
stressors, mediated via impacts to
meadow hydrology following upslope
land management practices over the last
century, have detrimentally affected the
quantity and quality of breeding
meadows. Many of these stressors, such
as grazing, have been more significant in
the past than under current management
standards. However, legacy effects
remain, and meadows tend not to
recover without active intervention once
excessive stream incision in their
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watershed is set in motion (Vankat and
Major 1978, pp. 386–397). Certain
stressors may be of concern, such as
recreational impacts and avian
predation upon terrestrial life stages of
toads, although we do not have
sufficient data to document the
magnitude of these particular stressors.
Given the evidence supporting the
role of climate in reducing populations
and potentially leading to the
extirpation of many of the populations
studied through the 1970s and into the
early 1990s (Kagarise Sherman and
Morton 1993, pp. 186–198), this factor
is likely either a primary driver, or at
least a significant contributing factor in
the declines that have been observed.
Climate models predict increasing
drought intensity and changes to the
hydroperiod based on reduced
snowpack, along with greater climate
variability in the future (PRBO 2011, pp.
18–25). These changes will likely
exacerbate stress to the habitat specialist
Yosemite toad through a pronounced
impact on its ephemeral aquatic habitat,
and also through an increase in the
frequency of freezing and drying events
that kill Yosemite toad eggs and
tadpoles. These changes and the
resultant impacts likely will effectively
reduce breeding success of remnant
populations already at low abundance
and still in decline. If an interaction
such as winter stress and disease (Carey
1993, pp. 355–362) is the underlying
mechanism for Yosemite toad declines,
then the enhanced influence of climate
change as a stressor may tip the balance
further towards higher incidence and
increased virulence of disease, which
would also lead to greater population
declines and extirpations.
Determination for Yosemite Toad
Section 4 of the Act (16 U.S.C. 1533),
and its implementing regulations at 50
CFR part 424, set forth the procedures
for adding species to the Federal Lists
of Endangered and Threatened Wildlife
and Plants. Under section 4(a)(1) of the
Act, we may list a species based on (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 (E)
Other natural or manmade factors
affecting its continued existence. Listing
actions may be warranted based on any
of the above threat factors, singly or in
combination.
We have carefully assessed the best
scientific and commercial information
available regarding the past, present,
and future threats to the Yosemite toad.
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The Yosemite toad is the most narrowly
distributed Sierra Nevada endemic,
pond-breeding amphibian (Shaffer et al.
2000, p. 246). Although it apparently
still persists throughout a large portion
of its historical range, it has been
reduced to an estimated 13 percent of
historical watersheds. (The proposed
rule indicated that the toad was reduced
to an estimated 12 percent of its range,
peer review corrected this number to 13
percent (Brown 2013, unpaginated). In
addition, while the best available data
do not provide information on whether
populations are currently stable, or
whether there is a persistent decline,
remnant populations are predominantly
small.
Yosemite toad populations are subject
to threats from habitat degradation
associated with land uses that
negatively influence meadow
hydrology, fostering meadow
dewatering, and conifer and other
invasive plant encroachment. These
activities include the legacy effects of
historic grazing activities, the fire
management regime of the past century,
historic timber management activities,
and associated road construction. The
impacts from these threats are
cumulatively of moderate magnitude,
and their legacy impacts on meadow
habitats act as a constraint upon extant
populations now and are expected to
hinder persistence and recovery into the
future. Diseases are threats of
conservation concern that have likely
also had an effect on populations
leading to historical population decline,
and these threats are operating currently
and will continue to do so into the
future, likely with impacts of moderatemagnitude effects on Yosemite toad
populations.
The individual, interactive, and
cumulative effects of these various risk
factors have acted to reduce the
geographic extent and abundance of this
species throughout its habitat in the
Sierra Nevada. The combined effect of
these stressors acting upon small
remnant populations of Yosemite toads
is of significant conservation concern.
The Yosemite toad has a life history and
ecology that make it sensitive to drought
and anticipated weather extremes
associated with climate change. Climate
change is expected to become
increasingly significant to the Yosemite
toad and its habitat in the future
throughout its range. Therefore, climate
change represents a threat that has a
high magnitude of impact as an indirect
stressor via habitat loss and degradation,
and as a direct stressor via enhanced
risk of climate extremes to all life stages
of Yosemite toads.
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The Act defines an endangered
species as any species that is ‘‘in danger
of extinction throughout all or a
significant portion of its range’’ and a
threatened species as any species ‘‘that
is likely to become endangered
throughout all or a significant portion of
its range within the foreseeable future.’’
We find that the Yosemite toad is likely
to become endangered throughout all or
a significant portion of its range within
the foreseeable future, based on the
immediacy, severity, and scope of the
threats described above. These include
habitat loss associated with degradation
of meadow hydrology following stream
incision consequent to the cumulative
effects of historic land management
activities, notably livestock grazing, and
also the anticipated hydrologic effects
upon habitat from climate change under
listing Factor A. Additionally, we find
that disease under listing Factor C was
likely a contributor to the recent historic
decline of the Yosemite toad, and may
remain an important factor limiting
recruitment in remnant populations. We
also find that the Yosemite toad is likely
to become endangered through the
direct effects of climate change
impacting small remnant populations
under Factor E, likely compounded with
the cumulative effect of other threat
factors (such as disease).
We have carefully assessed the best
scientific and commercial information
available regarding the past, present,
and future threats to the species, and
have determined that the Yosemite toad
meets the definition of threatened under
the Act, rather than endangered. This
determination is because the impacts
from the threats are occurring now at
high and moderate magnitudes, but are
all likely to become of high magnitude
in the foreseeable future across the
species’ entire range, making the species
likely to become in danger of extinction.
While population decline has been
widespread, the rate of decline is not so
severe to indicate extinction is
imminent, but this rate could increase
as stressors such as climate change
impact small remnant populations.
Further, the geographic extent of the
species remains rather widespread
throughout its historic range, conferring
some measure of ecological and
geographic redundancy. Therefore, on
the basis of the best available scientific
and commercial information, we
finalize listing the Yosemite toad as
threatened in accordance with sections
3(20) and 4(a)(1) of the Act.
The term ‘‘threatened species’’ means
any species (or subspecies or, for
vertebrates, distinct population
segments) that is likely to become an
endangered species within the
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foreseeable future throughout all or a
significant portion of its range. The Act
does not define the term ‘‘foreseeable
future’’ but it likely describes the extent
to which the Service could reasonably
rely on predictions about the future in
making determinations about the future
conservation status of the species. In
considering the foreseeable future as it
relates to the status of the Yosemite
toad, we considered the historical data
to identify any relevant existing trends
that might allow for reliable prediction
of the future (in the form of
extrapolating the trends). We also
considered how current stressors are
affecting the species and whether we
could reliably predict any future trends
in those stressors that might affect the
species recognizing that our ability to
make reliable predictions for the future
is limited by the quantity and quality of
available data. Thus the foreseeable
future includes the species’ response to
these stressors and any trends.
Under the Act and our implementing
regulations, a species may warrant
listing if it is endangered or threatened
throughout all or a significant portion of
its range. The Yosemite toad is highly
restricted in its range, and the threats
occur throughout its range. Therefore,
we assessed the status of the species
throughout its entire range. The threats
to the survival of the species occur
throughout the species’ range and are
not restricted to any particular
significant portion of that range, nor are
they concentrated in a specific portion
of the range. Accordingly, our
assessment and final determination
applies to the species throughout its
entire range.
Summary of Comments
In the proposed rule published on
April 25, 2013 (78 FR 24472), we
requested that all interested parties
submit written comments on the
proposal by June 24, 2013. Given the
large number of requests that we
received to extend the public comment
period, we reopened the comment
period on July 19, 2013 (78 FR 43122),
requesting written comments on the
proposal by November 18, 2013, and
again reopened the comment period on
January 10, 2014 (79 FR 1805), with the
close of comment period on March 11,
1014. We also contacted appropriate
Federal and State agencies, scientific
experts and organizations, and other
interested parties and invited them to
comment on the proposal. Newspaper
notices inviting general public comment
were published in the Sacramento Bee
and Bakersfield Californian. We
received multiple requests for a public
hearing. We held two public hearings on
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January 30, 2014, in Sacramento,
California. We also held two public
informational meetings, one in
Bridgeport, California, on January 8,
2014, and the other in Fresno,
California, on January 13, 2014. We also
participated in several public forums,
one sponsored by Congressman
McClintock and two sponsored by
Congressman LaMalfa. All substantive
information provided during comment
periods has either been incorporated
directly into this final determination or
addressed below.
Peer Reviewer Comments
In accordance with our peer review
policy published on July 1, 1994 (59 FR
34270), we solicited expert opinion
from five knowledgeable individuals
with scientific expertise that included
familiarity with the Sierra Nevada
yellow-legged frog, the northern DPS of
the mountain yellow-legged frog, the
Yosemite toad, and the habitat and
biological needs of, and threats to each
species. We received responses from
four of the peer reviewers.
We reviewed all comments received
from the peer reviewers for substantive
issues and new information regarding
the listing of the Sierra Nevada yellowlegged frog, the northern DPS of the
mountain yellow-legged frog, and the
Yosemite toad. The peer reviewers
generally concurred with our methods
and conclusions and provided
additional information, clarifications,
and suggestions to improve the final
rule. However, one of the four peer
reviewers suggested the rationale for
listing Yosemite toad was poorly
supported. Peer reviewer comments are
addressed in the following summary
and incorporated into the final rule.
(1) Comment: Two peer reviewers
recommended that we refer to Rana
muscosa as the southern mountain
yellow-legged frog in order to reduce
reader confusion in the final rule.
Our Response: We have clarified the
common names we are using in this
final rule for each yellow-legged frog
species (see Background and Taxonomy
sections in this final rule). While
Crother et al. (2008, p. 11) accepted the
common name of southern mountain
yellow-legged frog for Rana muscosa,
the use of this common name may
create additional confusion as the reader
may interpret the name to imply the
yellow-legged frogs in southern
California that are already listed as the
southern DPS, rather than the R.
muscosa in the Sierra Nevada.
Therefore, we continue to refer to the
northern DPS of Rana muscosa as the
northern DPS of the mountain yellowlegged frog, as we did in the proposed
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rule, to minimize confusion for the
public.
(2) Comment: Two peer reviewers
suggested that we utilize a rangewide
analysis for listing Rana muscosa and
thereby combine the northern and
southern DPSs of the mountain yellowlegged frog into one listed entity.
Clarifying discussions with one peer
reviewer suggested that we not complete
a rangewide analysis, but rather keep
the DPSs separate (Knapp, pers. comm.).
Our Response: Given the geographic
isolation, different habitat requirements,
differences in threats, and different
management needs between Rana
muscosa in the Sierra Nevada compared
with southern California, we have
decided to retain the DPS analysis in the
proposed rule and to maintain the
northern and southern DPSs of
mountain yellow-legged frog as separate
listed entities. Within the Sierra
Nevada, R. muscosa is predominantly
found within high-elevation lake
habitats that freeze during the winter
months, while in southern California,
Rana muscosa populations occupy
stream habitats that are not typically
subject to winter freezing. The
differences in the habitats utilized by
the northern and southern DPSs of the
mountain yellow-legged frog and the
differences in the threats to each
population segment indicate that
management actions needed to recover
the northern California and southern
California populations will also be
different and are most expediently
addressed separately by DPS (see
Distinct Vertebrate Population Segment
Analysis in this final rule).
The factors that are threats to the
species also differ between the two
DPSs. We have identified fish stocking
and presence of fish as a threat for both
the northern and southern DPSs.
However, the other threats we identified
for the northern DPS are primarily
habitat degradation, disease, and
climate change, whereas the main
threats for the southern DPS consist of
recreational activities, roads, and
wildfire. While there is some overlap in
the threats identified for the two DPSs,
the threats that are important to the
species status vary substantially
between the Sierra Nevada and southern
California.
The differences between the northern
and southern DPSs of the mountain
yellow-legged frog in both habitat use
and the factors affecting the species
results in differences in the actions and
activities that would be needed to
conserve the species in each of the two
DPSs. Conservation planning, including
identifying actions and setting priorities
for recovery, will be more effective and
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better suited to meet the species’ needs
if two separate DPSs are retained.
(3) Comment: One peer reviewer
indicated that the frogs within the
Spanish and Bean Creek areas of Plumas
County (low-elevation areas within the
northern portion of the Sierra Nevada)
in which Wengert (2008) conducted
telemetry studies of frog movement
distances, may actually be foothill
yellow-legged frog (Rana boylii) rather
than Sierra Nevada yellow-legged frogs
(Rana sierrae) (see Habitat and Life
History section in Background for the
mountain yellow-legged frogs of this
final rule).
Our Response: We acknowledge and
understand some of the challenges in
correctly identifying the species in areas
where the ranges of Sierra Nevada and
foothill yellow-legged frogs overlap.
Recent genetic analysis of samples
collected from frogs in Spanish and
Bean Creeks has identified the frogs
occurring in Bean Creek as both Sierra
Nevada and foothill yellow-legged frogs
(Lind et al. 2011a, pp. 281–282), while
Spanish Creek frogs were identified as
foothill yellow-legged frog (Poorten et
al. 2013, p. 4). However, given the small
sample size, Poorten et al. (2013, p. 4)
suggested that followup investigation
was needed to determine whether Sierra
Nevada yellow-legged frogs also occur
in Spanish Creek.
While it is not clear whether Wengert
(2008) studied Sierra Nevada or foothill
yellow-legged frogs, given the streambased ecological setting of the study, we
expect that the movement distances
recorded are applicable to the Sierra
Nevada yellow-legged frog within a
stream-based system, as the ecology is
comparable between the two sister taxa
in regard to stream systems.
Additionally, a study conducted by
Fellers et al. (2013, p. 159) documented
Sierra Nevada yellow-legged frog
movement distances up to 1,032 m in a
29-day period, suggesting the seasonlong movement distance documented by
Wengert (2008, p. 20) is applicable.
(4) Comment: One peer reviewer
provided comment that our proposed
rule did not include more-recent
literature on the effects of airborne
contaminants on the mountain yellow
legged frog, including Bradford et al.
2011, which measured contaminant
concentrations at multiple sites in the
southern Sierra Nevada and compared
their distribution with population
declines of mountain yellow-legged
frogs, finding no association between
the two. The peer reviewer further
recommended that we state that frogs
are sensitive to contaminants, but
measured contaminant concentrations
in multiple media indicate very low
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exposures to contaminants from upwind
sources.
Our Response: In our proposed rule,
we included a discussion of
environmental factors that affect the
mountain yellow-legged frog complex,
including contaminants. Based on our
analysis in the proposed rule, we did
not identify this environmental factor as
a threat to the species. Upon our review
of additional literature, including a
study focused specifically on the
mountain yellow-legged frog complex,
our initial discussion remains valid,
which indicated that the potential threat
posed by contaminants is not a factor in
the listing of this species. We refer to
the proposed rule for the discussion of
the effects of contaminants on the
mountain yellow-legged frog.
(5) Comment: One peer reviewer
suggested that recent genetic studies
(Shaffer et al. 2000, Stevens 2001, and
Goebel et al. 2009) do not support our
conclusion that Yosemite toad is a valid
species.
Our Response: When conducting our
review of the Yosemite toad as a listable
entity under the Act, we incorporated
the results of the studies mentioned by
the peer reviewer. In addition to the
previously included literature on the
genetics of Yosemite toad, we have
included in this final rule results from
Switzer et al. (2009), which provide
genetic data supporting the Yosemite
toad as a valid species. While we
acknowledge that the evolutionary
history of the Yosemite toad is
complicated and not fully understood,
given our conclusions after reviewing
the taxonomy of the species, and given
that the scientific community as a whole
continues to recognize the Yosemite
toad as a valid species, we continue to
recognize Yosemite toad as a valid
species (for further discussion, see
Taxonomy section above).
(6) Comment: One peer reviewer
provided information regarding the
number of localities of Yosemite toad
within two National Parks, and
suggested that, had we included these
locations, the analysis may have had a
different outcome.
Our Response: When we conducted
our analysis for the proposed rule to
determine whether the Yosemite toad
warrants listing under the Act, we
utilized the best available scientific and
commercial information. Part of that
information included the geospatial data
for Yosemite toad locations within both
Yosemite and Sequoia National Parks.
These data were subsequently used for
the proposed critical habitat
designation. While we did have (and
used) the information on Yosemite toad
locations within the National Parks in
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our analysis, we did not cite to this
information into the text of the
proposed rule. This was updated with
the data included in Berlow et al.
(2013), as well as information received
from Sequoia National Park staff.
Regardless, we utilized the geospatial
data in the proposed rule, determining
that the information suggests that the
Yosemite toad has disappeared from
approximately 47–69 percent of
formerly occupied sites (Berlow et al.
2013, p. 2). In addition, at many of the
remaining sites, Yosemite toads exist in
very low numbers, indicating that many
remaining populations are vulnerable to
extirpation. Our use of the data from
both National Forests and National
Parks led us to our proposed status
determination, which is affirmed here.
(7) Comment: One peer reviewer
stated that there is scant evidence
available to argue that there has been a
decline in abundance of the Yosemite
toad and that the difficulty in accurately
quantifying toad abundance, coupled
with the fact that the proposed rule did
not include locality data from the
National Parks, has weakened the
argument for our determination.
Our Response: While we agree that no
studies have documented a rangewide
decline in population abundances in
Yosemite toads, and we do not have
sufficient data to conduct a robust trend
analysis or detect negative population
growth rates, we relied on published
literature for our determination. At a
minimum, the published literature
provides anecdotally documented
declines in numbers of individual
Yosemite toads at the respective study
sites. The best available information
shows that the Yosemite toad
populations have declined, and that the
remnant populations comprise low
numbers of individual adult toads. For
our analysis, we did utilize the data on
toad locations in the National Parks (see
our response to comment 6) and
included it as part of our analysis on the
estimated loss of historically occupied
sites (47–69 percent of formerly
occupied sites (Berlow et al. 2013, p. 2)).
We mainly focused our analysis on the
potential drivers of population stability
and identified the predominate threats
to the species as the continuing effects
of degradation of meadow hydrology
associated with historical land
management practices and the effects of
climate change and anthropogenic
stressors acting on the small remnant
populations. (For complete discussion
see Summary of Factors Affecting the
Species section above.)
(8) Comment: One peer reviewer
stated that there are scientific
uncertainties regarding the long-term
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population trends and threats to
Yosemite toad and that these
uncertainties should be explicitly
described.
Our response: As required by the Act,
we based our proposed rule and this
final rule on the best available scientific
and commercial data. While there are
some uncertainties in the information,
we clearly articulated these
uncertainties when conducting our
analysis for the rule. (See Population
Estimate and Status and Meadow
Habitat Loss and Degradation sections
for examples.)
Federal Agency Comments
(9) Comment: The Forest Service
suggested that the rule does not
represent the best available scientific
and commercial information in
proposing a determination.
Our Response: In conducting our
analysis, we rely on the best available
scientific and commercial information,
as required by the Act. On occasion, we
are not aware of certain information that
is available at the time we issue a
proposed rule or new information
becomes available around the time of
publication, which is part of the reason
we request public comment, as well as
peer review. That portion of the process
helps to inform our final decision by
soliciting input and seeking additional
available information. As a result of this
process, we have received new scientific
and commercial information that we
have reviewed and incorporated into
this final rule.
(10) Comment: The USFS noted that
the proposed rule did not identify
mining activities as a threat to the
mountain yellow-legged frog.
Our Response: We acknowledge that
there is some overlap between current
mining activities and areas occupied by
the mountain yellow-legged frogs,
particularly in the northern part of the
range; however, we do not have
information to assess the impact that
mining has on the species in those areas
where mining occurs, and how it acts as
either an historical or current threat to
the species. Within designated
wilderness, new mining claims have
been prohibited since January 1, 1984.
Additionally, while suction dredge
mining may have the potential to alter
microhabitat uses by the species, the
current moratorium on this practice
removes this potential threat. However,
we acknowledge that this situation may
change in the future.
(11) Comment: The USFS suggested
that the uncertainties we presented
under Factor D as it relates to their
Forest Plan revision process and
protections for mountain yellow-legged
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frog are not applicable and that the
protections under the SNFPA will
continue as a result of consultation with
the Service.
Our Response: We did not identify
Factor D as a threat to the mountain
yellow-legged frog, and we incorporated
an analysis of the protection that the
current Forest Plans offer the species.
While there is some uncertainty as to
whether these protections will remain
in the revised Forest Plans, the USFS is
not required to consult with the Service
on the Sierra Nevada yellow-legged frog
and northern DPS of the mountain
yellow-legged frog in the absence of the
protections afforded under the Act. As
such, we must evaluate the adequacy of
existing regulatory mechanisms from
the baseline of the species not being
federally listed under the Act.
(12) Comment: The USFS suggested
the final rule include a discussion of the
impacts of bullfrog predation on the
mountain yellow-legged frog.
Our Response: We have limited
information on the presence of bullfrogs
in the Sierra Nevada, but we have
included a section on the potential
threat of American bullfrogs where they
are known to occur in the Lake Tahoe
Basin (see discussion under Factor C for
mountain yellow-legged frogs).
(13) Comment: The USFS and several
other commenters suggested that the
information presented as it relates to the
impacts of grazing on Yosemite toad
was inaccurate. Specifically, they
suggested that we did not include the
results of peer-reviewed journal articles
in our analysis of the impacts posed by
livestock grazing.
Our Response: At the time of the
proposed rule, we were aware of the
peer-reviewed literature related to the
impacts of livestock grazing on
Yosemite toad, and inadvertently
omitted the literature from the rule. We
have reviewed and included the
relevant articles in this final rule.
Additionally, while we did not
incorporate all of the specifics of the
journal articles, we did incorporate the
results of a 5-year study that
investigated the impacts of cattle
grazing on Yosemite toad in our
analysis, as they were presented in
Allen Diaz et al. 2010, and subsequently
in the Lind et al. (2011b, addendum).
(14) Comment: The USFS and several
other commenters suggested that our
reliance on a single non-peer-reviewed
study to assess the impacts of cattle
grazing on Yosemite toads, through
direct mortality or the modification of
their habitat, was inappropriate.
Additionally, they suggested we
discounted the peer-reviewed published
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journal articles related to the impacts of
cattle grazing on Yosemite toad.
Our Response: In conducting our
analysis, we rely on the best available
scientific and commercial information,
as required by the Act. This information
does not need to be specifically
published in a scientific journal. The
Martin (2008) study that is being
referred to by the commenters is a
doctoral dissertation that was, in fact,
reviewed prior to release. We relied on
the information presented by Martin in
assessing the potential for direct
mortality of Yosemite toad that is
attributed to livestock. We also relied on
Martin for the potential impacts of
livestock grazing on overwintering and
upland areas utilized by Yosemite toad,
as the peer-reviewed publications that
the commenters referred to were based
on a study that only assessed grazing
effects on breeding. As such, the best
available scientific and commercial
information includes Martin (2008). In
our proposed rule, we evaluated the
information that ran contrary to Martin
(2008), and we have subsequently
incorporated the information presented
in the peer-reviewed journal articles in
this final rule. Please also see response
to comment #13.
(15) Comment: The USFS commented
that chytrid fungus, fish stocking, and
climate change pose the greatest threats
to the mountain yellow-legged frogs,
and that threats from authorized
management activities are insignificant
threats to the species.
Our Response: We have concluded in
this final rule that, in general,
authorized activities on public lands
managed by the USFS and the NPS are
not significant threats to the mountain
yellow-legged frogs, but we also
recognize that there may be limited sitespecific conditions where authorized
activities could have population-level
effects, especially where populations are
small or habitat areas are limited (see
Summary of Factors Affecting the
Species in this final rule).
(16) Comment: The USFS noted that
recent publications indicate that
livestock grazing that meets current
USFS standards and guidelines is less of
a threat to the Yosemite toad than was
described in the proposed rule.
Our Response: We have revised our
discussion of grazing in this final rule
to clarify the conditions under which
we consider current grazing activities to
pose habitat-related threats to the
Yosemite toad (see Summary of Changes
and Factor A discussion for the
Yosemite toad).
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Comments From States
(17) Comment: The California
Department of Fish and Wildlife
(CDFW) originally commented that the
threats presented in the proposed rule
suggested that a determination of
threatened status would be more
appropriate than endangered for the
Sierra Nevada yellow-legged frog.
However, CDFW reconsidered this
suggestion after discussions with
Service staff and submitted a followup
comment letter that agrees with the
Service determination and supports
listing the Sierra Nevada yellow-legged
frog as endangered.
Our Response: We find that an
endangered status for the Sierra Nevada
yellow-legged frog is an appropriate
determination and appreciate CDFW’s
reconsideration of their initial
comments.
(18) Comment: CDFW commented
that they remain concerned that listing
the species as endangered could hinder
timely implementation of the
Department’s recovery and restoration
efforts for the species pursuant to its
State-listing under CESA. CDFW notes
that they have a responsibility to
continue activities and expand efforts
that will contribute to the recovery of
the Sierra Nevada yellow-legged frog
and hope that such efforts can be
fostered through the 1991 Cooperative
Agreement between the California
Department of Fish and Game and the
U.S. Fish and Wildlife Service. They
also comment that, in his June 13, 2012,
memo to the Service’s Regional
Directors, the Director of the Fish and
Wildlife Service acknowledged the
Federal-State collaborative nature of
conservation activities for listed species.
Our Response: We note that, for
research activities that aid in the
recovery of the species, and that may
result in take, a permit issued under
section 10a(1)A of the Act is the
appropriate mechanism. However, our
regulations at 50 CFR 17.21 state that
any qualified employee or agent who is
designated by CDFW for such purposes,
may, when acting in the course of his
official duties, take endangered wildlife
species covered by a Cooperative
Agreement (developed pursuant to
Section 6 of the Act) between the
Service and the State provided such
take is not reasonably anticipated to
result in: (1) The death or permanent
disabling of the specimen; (2) the
removal of the specimen from the State
of California; (3) the introduction of the
specimen or any of its progeny into an
area beyond the historical range of the
species; or (4) the holding of the
specimen in captivity for a period of
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more than 45 days. Take that does not
meet these four conditions would
require a section 10(a)(1)(A) permit. We
acknowledge and appreciate the
important role that CDFW will play in
the recovery of the Sierra Nevada
yellow-legged frog, and look forward to
continuing collaborative conservation
actions with CDFW for this and other
listed species in California.
(19) Comment: CDFW agreed that we
should retain the northern DPS and the
southern DPS designations for the
mountain yellow-legged frog (Rana
muscosa). They provided updates to our
discussion of take related to State-listing
of the mountain yellow-legged frog
complex.
Our Response: We appreciate the
support, and we have retained the two
DPSs in the final determination (see
Distinct Vertebrate Population Segment
Analysis). We have also revised our
discussion of CESA to provide the
updated information on take related to
State-listing of the mountain yellowlegged frog complex (see Factor D for
mountain yellow-legged frog).
(20) Comment: CDFW provided
comments on our discussion of the
following threats to the mountain
yellow-legged frog complex:
Recreational activities, past trout
stocking versus continued trout
stocking, and pesticide detection in the
Sierra Nevada. They commented that
the evidence presented in the
Recreation section did not support the
conclusion, urging us to readdress the
section and remove claims unsupported
by appropriate citations, and noted that
recreation effects to the environment
were supported, but no evidence
indicates that such activities affect the
frog populations. In the Recreation
section, they also noted several errors
and inaccuracies in citing other authors.
CDFW provided extensive comments on
our discussion of dams and water
diversions, commenting that they were
of the opinion that dams and diversion
posed a threat of low significance to the
continued existence of the mountain
yellow-legged frogs and suggesting that
the section required significant
amendments to accurately capture the
degree of potential impacts. They noted
that most dams were constructed below
the range of extant frog populations, and
that some information was misapplied
from research on lower-elevation
amphibian species, such as the foothill
yellow-legged frog, which resulted in
overstatement of the potential impact of
dams and water diversions on the
mountain yellow-legged frog complex.
They provided numerous smaller
specific comments on text within the
section.
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Our Response: We thank the CDFW
for the additional information provided
to strengthen our analysis. We have
addressed these comments through
changes to the Fish Stocking,
Recreation, and Dams and Water
Diversions sections for the Sierra
Nevada and mountain yellow-legged
frogs in this final rule. We re-checked
references and revised the sections
noted to state more clearly the potential
effects of these activities, to rely on
appropriate citations, and to refine our
conclusions in agreement with CDFW’s
comments. Please see Factor A in
Summary of Factors Affecting the
Species for updated information.
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Public Comments
(21) Comment: Several commenters
suggested that the Service does not have
the authority or jurisdiction to designate
the Sierra Nevada yellow-legged frog
and the northern DPS of the mountain
yellow-legged frog as endangered nor
the Yosemite toad as threatened.
Our Response: The authority for the
Service to issue this rulemaking comes
from the Endangered Species Act of
1973 (16 U.S.C. 1531 et seq.), as
amended, through the 108th Congress.
The Service is designated as the lead
Federal agency for implementing the
Act for terrestrial and freshwater
species. Authority to implement the Act
does not require Federal jurisdiction or
land ownership
(22) Comment: Multiple commenters
indicated that existing Federal and State
legislation and regulations, such as the
Wilderness Act, CESA, and CDFW
regulations, provide sufficient
protection for these amphibians, and
thereby eliminate the need for listing
the species.
Our Response: We agree that existing
Federal and State legislation and
regulations, such as the Wilderness Act,
CESA, and CDFW regulations provide
some protection for the Sierra Nevada
yellow-legged frog, the northern DPS of
the mountain yellow-legged frog, and
the Yosemite toad. However, while
existing legislation and regulations
provide some level of protection for the
Sierra Nevada yellow-legged frog, the
northern DPS of the mountain yellowlegged frog, and the Yosemite toad, they
do not require that Federal agencies
ensure that actions that they fund,
authorize, or carry out will not likely
jeopardize the species’ continued
existence (for further information see
discussions under Factor D). Therefore,
we have determined that the Sierra
Nevada yellow-legged frog and the
northern DPS of the mountain yellowlegged frog are endangered and that the
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Yosemite toad is threatened under the
Act.
(23) Comment: Several commenters
suggest that it is necessary for the
Service to conduct an analysis of the
impacts that listing a species may have
on local economies prior to issuance of
a final rule.
Our Response: Under the Act, the
Service is not required to conduct an
analysis regarding the economic impact
of listing endangered or threatened
species. However, the Act does require
that the Service consider the economic
impacts of a designation of critical
habitat. A draft of this analysis is
available to the public on https://
www.regulations.gov (79 FR 1805).
(24) Comment: Several commenters
suggested that the decline of the Sierra
Nevada yellow-legged frog, northern
DPS of the mountain yellow-legged frog,
and the Yosemite toad is a natural
evolutionary process, and that the
presence of environmental stressors is a
normal driver of evolution and/or
extinction.
Our Response: Under the Act, we are
required to use the best available
scientific and commercial information
to assess the factors affecting a species
in order to make a status determination.
The Act requires the Service to consider
all threats and impacts that may be
responsible for declines as potential
listing factors. The evidence presented
suggests that the threats to the species
are both natural and manmade (see
Factor E—Other Natural or Manmade
Factors Affecting the Species), but that
they are primarily the result of
anthropogenic influences (see Summary
of Factors Affecting the Species in this
final rule). Thus, the threats associated
with the declines of these species are
not part of a natural evolutionary
process.
(25) Comment: Several commenters
were concerned about the effects of
listing on mining and associated
activities conducted under the General
Mining Law of 1872. They suggested
that the listing of these species will
remove 5 million acres from mining and
other productive uses of the land. One
commenter was concerned that there
would be no assurances that
development of a mining claim will
result in the ability to mine it.
Our Response: In the proposed rule,
we identified unauthorized discharge of
chemicals or fill material into any water
upon which the Sierra Nevada yellowlegged frog, the northern DPS of the
mountain yellow-legged frog, and the
Yosemite toad are known to occur as a
potential threat to these species. On
National Forests outside of designated
wilderness, new mining may occur
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pursuant to the Mining Law of 1872 (30
U.S.C. 21 et seq.), which was enacted to
promote exploration and development
of domestic mineral resources, as well
as the settlement of the western United
States. It permits U.S. citizens and
businesses to prospect hardrock
(locatable) minerals and, if a valuable
deposit is found, file a claim giving
them the right to use the land for mining
activities and sell the minerals
extracted, without having to pay the
Federal Government any holding fees or
royalties (GAO 1989, p. 2). Gold and
other minerals are frequently mined as
locatable minerals, and, as such, mining
is subject to the Mining Law of 1872.
However, Federal wilderness areas were
closed to new mining claims at the
beginning of 1984 (see Factor D under
mountain yellow-legged frogs above),
thereby precluding the filing of new
mining claims in those areas designated
as Federal wilderness (a large part of the
area in which the species occur).
Authorization of mining under the
Mining Law of 1872 is a discretionary
agency action pursuant to section 7 of
the Act. Therefore, Federal agencies
with jurisdiction over land where
mining occurs will review mining and
other actions that they fund, authorize,
or carry out to determine if listed
species may be affected in accordance
with section 7 of the Act.
(26) Comment: Numerous
commenters suggested that the listing of
the Sierra Nevada yellow-legged frog,
the northern DPS of the mountain
yellow-legged frog, and the Yosemite
toad are being misused to restrict or
prohibit access for fishing, hiking,
camping, and other recreational uses,
and implement land use restrictions,
management requirements, and personal
liabilities on the public that are not
prudent, clearly defined, or necessary.
Our Response: The listing of the
Sierra Nevada yellow-legged frog, the
northern DPS of the mountain yellowlegged frog, and the Yosemite toad does
not prevent access to any land, whether
private, tribal, State, or Federal. The
listing of a species does not affect land
ownership or establish a refuge,
wilderness, reserve, or other
conservation area. A listing does not
allow the government or public to
access private lands without the
permission of the landowner. It does not
require implementation of restoration,
recovery, or enhancement measures by
non-Federal landowners. Federal
agencies will review actions that they
fund, authorize, or carry out to
determine if any of these three
amphibians, and other listed species as
appropriate, may be affected by the
Federal action. The Federal agency will
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consult with the Service, in accordance
with Section 7 of the Act (see also
response to comment 25).
(27) Comment: Several commenters
suggested that listing the Sierra Nevada
yellow-legged frog and the northern DPS
of the mountain yellow-legged frog
under the Act is not necessary given
that a majority of the range of these
species is within wilderness areas
afforded protection under the
Wilderness Act and by the protections
afforded under CESA.
Our Response: We agree that existing
Federal and State legislation and
regulations, such as the Wilderness Act
and CESA, provide some protection for
the Sierra Nevada mountain yellowlegged frog, the northern DPS of the
mountain yellow-legged frog, and the
Yosemite toad. However, we identified
the main threats to the two frog species
as habitat degradation and
fragmentation, predation and disease,
climate change, and the interactions of
these stressors on small populations.
Neither the Wilderness Act nor the
State’s listing status under CESA
ameliorates these threats to levels that
would preclude the need to list the
species under the Act. (See discussion
under Factor D).
(28) Comment: One commenter
suggested that habitat and range of the
mountain yellow-legged frog is not
threatened with destruction or
modification based on a large portion
being located in wilderness, and the
proposed rule stating ‘‘physical habitat
destruction does not appear to be the
primary factor associated with the
decline of the mountain yellow-legged
frogs.’’
Our Response: While we agree that
the loss, destruction, or conversion of
physical habitat is not a primary factor
in the decline of the mountain yellowlegged frogs, we discuss both the
biological modification of habitat due to
changes in predator communities, prey
communities, and in nutrient levels,
and due to the habitat fragmentation
associated with the presence of
introduced fish. Although the presence
of introduced fish does not result in
conversion or loss of the physical
attributes of habitat (for example,
removal or filling of lakes, ponds, etc.),
fish presence does effectively preclude
the use of the habitat by the mountain
yellow-legged frog (see our discussion
under Factor A). While a large portion
of the range of the mountain yellowlegged frog is within federally
designated wilderness, or on National
Parks, we identified the main threats to
the species as habitat degradation and
fragmentation, predation and disease,
climate change, and the interactions of
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these stressors on small populations.
Neither the Wilderness Act nor the
protections afforded within National
Parks ameliorates these threats to levels
that would preclude the need to list the
species under the Act (see discussion
under Factor D).
(29) Comment: One commenter stated
that we failed to consider the
effectiveness of restoration activities
being conducted by CDFW as part of
their High Mountain Lakes Project and
plans for Yosemite and Sequoia and
Kings National Parks that are intended
to implement restoration actions.
Our Response: We are aware of the
activities, including the High Mountain
Lakes Project (see Factor A discussions
above in this final rule), being
conducted by CDFW, USFS, NPS, and
researchers aimed at restoring habitat
for the mountain yellow-legged frog.
While efforts of interested parties have
resulted in the restoration of habitat for
these species, the restored habitat
represents a small portion of the range
of the species, and has occurred only in
localized areas. As such, these activities,
while beneficial and important for the
recovery of the species, do not
significantly counter the threats of
introduced predators, disease, or
climate change. Additionally, we are
aware of planning efforts by Yosemite
and Sequoia and Kings National Parks,
partially implemented, and we are
aware that these restoration plans have
not been finalized.
(30) Comment: One commenter
provided information suggesting
livestock are responsible for the
transportation of Bd in the environment.
Our Response: While livestock may
provide a vector for the transmission of
amphibian disease within the Sierra
Nevada, there are numerous other
mechanisms of transport, including
wildlife, as well as anthropogenic
vectors. Since the importance of
differing disease vectors related to Bd is
poorly understood, we did not include
a discussion of disease transport
associated with livestock grazing in this
rule (see Factor C for discussion of
disease).
(31) Comment: One commenter
provided information to suggest that
activities associated with illicit
cultivation of marijuana on National
Forest System lands should be
identified as a potential threat to the
mountain yellow-legged frog.
Our Response: We agree that aspects
associated with illegal cultivation of
marijuana on National Forest System
lands may pose a risk to the mountain
yellow-legged frogs, such as dewatering
of habitats and contamination from
pesticides and fertilizers. There is
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potential overlap with this illegal
activity and areas occupied by mountain
yellow-legged frogs; however, not
enough information is available at this
point to assess the impact that illegal
cultivation of marijuana has on the
species.
(32) Comment: Several commenters
suggest that there is insufficient
evidence to make a listing
determination for the mountain yellowlegged frog in accordance with the Act.
Our Response: As we have presented
in both the proposed rule and this final
rule, a substantial compilation of
scientific and commercial information is
available to support listing both the
Sierra Nevada yellow-legged frog and
the northern DPS of the mountain
yellow-legged frog under the Act. We
have presented evidence that there has
been a curtailment in range and
numbers attributed to habitat
degradation and fragmentation under
Factor A, predation and disease under
Factor C, and climate change and the
interaction of these various stressors
cumulatively impacting small remnant
populations under Factor E (see
Determination for the Sierra Nevada
Yellow-legged Frog and Determination
for the Northern DPS of the Mountain
Yellow-legged Frog sections above for a
synopsis and see the Summary of
Factors Affecting the Species for a
detailed analysis).
(33) Comment: Numerous
commenters purported that the greatest
threat to the mountain yellow-legged
frog is Bd, and since listing the species
will not alleviate the threat, the species
should not be listed. Additionally, it
was suggested that these species should
be reared in captivity until the threat of
Bd is resolved.
Our Response: We agree that Bd is
one of the primary contributing factors
in the current decline of these species;
however, it is not the only factor
responsible for their decline or the only
one forming the basis of our
determination. All Factors are
considered when making a listing
determination (see the Summary of
Factors Affecting the Species for a
detailed discussion). We have also
identified habitat fragmentation and
predation attributed to the introduction
of fish and climate change as threats to
the species. We are required to evaluate
all the threats affecting a species,
including disease under Factor C.
With respect to the prospect of
captive breeding, we acknowledge that
this activity is one of the suite of tools
that can be utilized for the conservation
of the species. Captive breeding is
currently being conducted for the
southern DPS of the mountain yellow-
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legged frog, and we are currently
working with various facilities to
explore this option. Additionally, when
a species is listed as either endangered
or threatened, the Act provides many
tools to advance the conservation of
listed species; available tools including
recovery planning under section 4 of the
Act, interagency cooperation and
consultation under section 7 of the Act,
and grants to the States under section 6
of the Act. All of these mechanisms
assist in the conservation of the species.
(34) Comment: Several commenters
provided information to suggest that
livestock grazing is not detrimental to
amphibian species and that the
proposed rule did not adequately
capture the neutral or beneficial effects
of livestock grazing on amphibian
species.
Our Response: We have revised our
discussion of grazing in this final rule
to clarify the conditions under which
we consider current grazing activities to
pose habitat-related threats (see Factor
A above). In addition, research with a
related ranid frog of western montane
environments, (the Columbia spotted
frog, Rana luteiventris) has indicated
that livestock grazing may reduce
vegetation levels in riparian and wet
meadow habitat, but does not have
short-term effects on the frog
populations, although they caution that
the length of the study may not capture
potential long-term effects (Adams et al.
2009, pp. 132, 137). However, George et
al. (2011, pp. 216, 232) in a review of
the effectiveness of management actions
on riparian areas, noted that continuous
grazing often results in heavy grazing
use of riparian areas, even if an area is
lightly stocked, because livestock are
attracted to the areas from adjacent
uplands. They note substantial literature
that documents that livestock grazing
could damage riparian areas, and the
resulting move, beginning in the 1980s,
in Federal and State resource agencies
to apply conservation practices to
protecting and improving riparian
habitats (George et al. 2011, p. 217).
They note that studies provide sufficient
evidence that riparian grazing
management that maintains or enhances
key vegetation attributes will enhance
stream channel and riparian soil
stability, although variable biotic and
abiotic conditions can have site-specific
effects on results (George et al. 2011, pp.
217–227).
In our proposed rule, we focused on
livestock grazing as a potential listing
factor, and while there are potentially
some current, localized effects to the
Sierra Nevada yellow-legged frog, the
northern DPS of the mountain yellowlegged frog, and the Yosemite toad, we
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consider the majority of the impacts
associated with livestock grazing are the
legacy effects of historically high
grazing intensities.
(35) Comment: One commenter stated
that the discussion of the effects of
global climate change in the proposed
rule for the Sierra Nevada yellow-legged
frog, northern DPS of the mountain
yellow-legged frog, and Yosemite toad
was not appropriate. The commenter
believed that the Service ‘‘pushes’’ the
climate models, both spatially and
temporally, beyond what the commenter
considered to be reliable, and ignores
their uncertainty. In addition, the
commenter claims that no credible
models can project potential climate
change in the Sierra Nevada. The
commenter stated the Act is not an
appropriate mechanism to regulate
global climate change and greenhouse
gases. Finally, the commenter suggested
if the Service does list the three
amphibians, that they be designated as
threatened species with a section 4(d)
rule that excludes lawful greenhouse
gases from the prohibitions of the Act.
Our Response: We used the best
available scientific and commercial
information available as it pertains to
climate change. In addition to the peerreviewed scientific journal articles and
reports that were utilized in our analysis
and cited in the proposed rule, recently
published studies have presented data
and conclusions that increase the level
of confidence that global climate change
is the result of anthropogenic actions
(summarized in Blaustein et al. 2010
and discussed above). A recent paper
(Kadir et al. 2013) provides specific
information on the effects of climate
change in the Sierra Nevada and is
discussed above. While the Service is
concerned about the effects of global
climate change on listed species,
wildlife, and their habitats, to date, we
have not used the Act to regulate
greenhouse gases. We evaluated the
suggestion that the three amphibians be
listed as threatened species with a
section 4(d) rule excluding prohibitions
or restrictions on greenhouse gases.
However, our determination is that the
Sierra Nevada yellow-legged frog and
the northern DPS of the mountain
yellow-legged frog meet the definition of
endangered, the Yosemite toad meets
the definition of threatened, and a
section 4(d) rule for greenhouse gases is
not appropriate.
(36) Comment: One commenter
suggested that the discussion of genetics
for the mountain yellow-legged frog
does not support the taxonomy of the
Sierra Nevada yellow-legged frog and
the northern DPS of the mountain
yellow-legged frog as separate species.
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The commenter further suggested the
text of the rule specifying two major
genetic lineages and four groups does
not support listing of the frogs as
separate genetic groups.
Our Response: Vredenburg et al.
(2007, p. 317) did not rely solely on
DNA evidence in the recognition of two
distinct species of mountain yellowlegged frog in the Sierra Nevada, but
instead used a combination of DNA
evidence, morphological information,
and acoustic studies. The taxonomy of
the mountain yellow-legged frogs as two
distinct species in the Sierras has been
widely accepted in the scientific
community and by species experts. We
are not listing a subspecies but rather
two separate, recognized species, the
Sierra Nevada yellow-legged frog and
the northern DPS of the mountain
yellow-legged frog.
(37) Comment: Several commenters
suggested that activities such as timber
harvest, road construction, recreation,
and livestock grazing are in decline in
the Sierras compared with historical
levels and should not be included as
potential threats to the Sierra Nevada
yellow-legged frog, the northern DPS of
the mountain yellow-legged frog, or the
Yosemite toad.
Our Response: In conducting our
analysis of the factors affecting the
species, we did include timber harvest,
road construction, recreation, and
livestock grazing, as potential threats to
the species, but acknowledge that the
major impact on the species was the
result of the legacy effects of historical
practices, and that these activities
currently pose a lower intensity,
localized threat. We have attempted to
clarify the distinction in this final rule
(see Factor A discussions above).
(38) Comment: Numerous
commenters stated that listing the
mountain yellow-legged frogs and the
Yosemite toad would prevent fuelsreduction activities, leading to fires and
loss of habitat.
Our Response: In this final rule under
Factor A for the mountain yellow-legged
frogs and Yosemite toad, we address
potential habitat changes that may be
related to timber harvest activities,
including harvests for fuels reduction
purposes. We found that most
populations of the three species occur at
high elevations above areas where
timber harvests are likely. At lower
elevations, forest standards and
guidelines would be expected to limit
potential threats to the species in most
cases, although limited site-specific
situations might result in habitat effects
with population consequences. We also
found that changed fire regimes have, in
some of the same lower elevation areas,
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led to an increased potential for highintensity fires, which could alter habitat
and, therefore, pose relatively localized
population-level effects to the species.
For the Yosemite toad, we found that
although ground-disturbance due to
timber harvest activities has the
potential to have population-level
effects at lower elevations, especially
where habitat is limited, currently the
best available information indicates
toads might achieve long-term benefits
from activities that reduce
encroachment of trees into breeding
sites. Therefore, we expect that fuelsreduction activities in lower elevation
areas will be generally beneficial to
these species.
(39) Comment: A number of
commenters suggested that, given the
results of more-recent studies that were
not included in the proposed rule,
livestock grazing should be removed as
a threat to the Yosemite toad (See also
comment 13 from the USFS).
Our Response: In our proposed rule,
we addressed the potential impacts of
grazing on Yosemite toad based on
Allen-Diaz et al. (2010). The morerecent studies referenced (such as Roche
et al. 2012a and 2012b, and McIlroy et
al. 2013) are different publications but
are based on the results of the
companion studies whose initial report,
and subsequent addendum, we
referenced as Allen-Diaz et al. (2010)
and Lind et al. (2011b). The study
conducted determined that livestock
grazing in accordance with the USFS’s
standards and guidelines does not affect
Yosemite toad breeding success. While
appropriately managed levels of grazing
do not impact breeding success, these
grazing standards are not always met.
Additionally, the main impact of
grazing on Yosemite toad is due to the
legacy effects of historical grazing
intensities on Yosemite toad habitat.
Given the limitations of the study (see
discussion under Factor A) and the
documentation that these standards are
not always met, livestock grazing may
continue to pose a localized threat to the
species.
(40) Comment: One commenter
provided several comments suggesting
that livestock grazing is not a threat to
Yosemite toad in light of the results of
a current study, the documentation of
Yosemite toads existing in areas that
have been subject to grazing for
centuries, and because the population
declines cited in our proposed rule
occurred in an area not subject to
grazing.
Our Response: See response to
comments 13, 14, and 39. In our
proposed rule, we identified the impacts
of livestock grazing primarily from an
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historical context as a potential
contributor to meadow degradation.
There is a great deal of information,
while not specific to Yosemite toad, on
the negative impacts of high-intensity
grazing regimes on ecosystem dynamics.
Grazing under current Forest Service
standards does not appear to impact
Yosemite toad breeding, however when
inappropriate levels of grazing do occur,
grazing may still present a localized
impact on Yosemite toads via direct
mortality or through practices that
prevent the hydrologic recovery of
historically wet meadow systems. While
the documented declines of Yosemite
toad have occurred in areas that are not
currently subject to livestock grazing,
historical grazing occurred throughout
the Sierra Nevada. We did not implicate
livestock grazing in the decline in
population sizes, rather as a potential
historical driver in meadow degradation
rangewide. We have clarified this
distinction in the final rule (see Factor
A discussion and Summary of Factors
Affecting the Species for the Yosemite
toad).
(41) Comment: One commenter
suggested that livestock grazing
continues to provide a threat to the
Sierra Nevada yellow-legged frog and
Yosemite toad and provided
information documenting habitat
degradation attributed to current
livestock grazing and utilization above
the standards of the SNFPA.
Our Response: As we have presented
in the proposed and final rules, the
impact of livestock grazing on these
species is primarily one of historical
significance, with the potential for
future localized impacts to the species
and/or their habitat. Based on the
information provided regarding habitat
conditions and potential impacts to
habitat, we have maintained our
position that current livestock grazing
poses a localized impact to the
mountain yellow-legged frogs and a
prevalent threat with moderate impacts
to the Yosemite toad.
(42) Comment: One party commented
that we have not demonstrated that the
Sierra Nevada population of the
mountain yellow-legged frog is a DPS.
They indicate that we have not shown
that the population is significant to the
taxon as a whole because we have not
shown whether other populations of the
species could persist in the highelevation Sierra Nevada portion of the
species’ range or discussed how the
Sierra Nevada populations are adapted
to the area. In addition, they indicate
that we failed to show that extirpation
of the northern population would result
in a significant gap in the range of the
species, and we did not show that the
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24307
populations had markedly different
genetics characteristics.
Our Response: The commenters
correctly noted that, to recognize a
population of a species as a DPS, we
must establish that the population is (1)
discrete from the remainder of the
populations to which the species
belongs, and (2) if determined to be
discrete, it is also found to be significant
to the species to which it belongs.
However, the commenters incorrectly
conclude that the population must meet
all three criteria for significance. We
find the northern population of the
mountain yellow-legged frog to be
discrete from the southern population
because it is separated from the
southern frogs by a 225-km (140-mi)
barrier of unsuitable habitat. The
primary basis for our finding that the
northern population is significant to the
species as a whole is that loss of the
northern population would mean the
loss of the species from a large portion
of its range and reduce the species to
small isolated occurrences in southern
California. The population also meets
two additional criteria for significance:
(1) Evidence of the persistence of the
discrete population segment in an
ecological setting unusual or unique for
the taxon, and (2) evidence that the
discrete population segment differs
markedly from the remainder of the
species in its genetic characteristics. We
have revised the language in our DPS
analysis to clarify the basis for the
determination (see Distinct Vertebrate
Population Segment Analysis).
(43) Comment: Numerous
commenters commented that we were
required to complete a NEPA analysis of
the proposed listing.
Our Response: We have determined
that environmental assessments and
environmental impact statements, as
defined under the authority of the
National Environmental Policy Act
(NEPA; 42 U.S.C. 4321 et seq.), need not
be prepared in connection with listing
a species as an endangered or
threatened species under the
Endangered Species Act. We published
a notice outlining our reasons for this
determination in the Federal Register
on October 25, 1983 (48 FR 49244) (see
Required Determinations section of this
rule).
(44) Comment: One commenter asked
that, if we determine that the three
amphibian species under consideration
are endangered or threatened under the
Act, then we enter into a cooperative
agreement with the State of California
under section 6 of the Act.
Our Response: We have been
operating under such a cooperative
agreement with the California
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Department of Fish and Game (now
Department of Fish and Wildlife
(CDFW)) since 1991. https://
www.dfg.ca.gov/wildlife/nongame/
publications/docs/CDFGCooperative
AgreementWithUSFWS.pdf
(45) Comment: One commenter stated
that if the three amphibians considered
are listed as threatened or endangered,
then research should continue into the
causes of population decline.
Our Response: We expect research on
these issues to continue into the future.
Once the three amphibians are listed as
threatened or endangered species under
the Act, additional funding for research
and other conservation programs for
those species will become available
through grants established under section
6 of the Act. Such grants are provided
to State agencies with which we have
established cooperative agreements.
(46) Comment: One commenter
indicated that because of a County
resolution, we must coordinate with the
board of supervisors of that County
prior to publishing a final rule.
Our Response: We provide all
interested parties an equal opportunity
to submit comments or information
prior to publication of a final rule, and
we give equal consideration to all such
information and comments, regardless
of source. Our requirements for
‘‘coordination,’’ however, are
established by the Act, by other Federal
statutes such as the Administrative
Procedure Act, and by executive order.
(47) Comment: Several commenters
asked for additional time to provide
comments. One commenter added that
we provided little public outreach.
Our Response: As discussed in the
first paragraph of the Summary of
Comments and Recommendations
section (above), we provided two
additional public comment periods for a
total of 240 days (approximately 8
months) of public comment. We also
hosted two public hearings and two
public informational meetings at various
locations within the range of the species
under consideration. We also attended
two additional public meetings hosted
by Congressmen representing districts
within the range of the species. We
contacted and sought input from
appropriate Federal and State agencies,
scientific experts and organizations, and
other interested parties. We also
published notices in the newspapers
with the largest readerships within both
the northern and southern portions of
the ranges of the species. Additional
public comment periods or outreach
were not feasible given limitations
imposed by available funds and
requirements imposed by the Act
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regarding available time in which to
publish a final rule.
(48) Comment: One commenter noted
that the Act authorizes the Secretary to
extend the time available for publication
of a final rule by up to 6 months if
‘‘there is substantial disagreement
regarding the sufficiency or accuracy of
the available data.’’ The commenter
stated that such substantial
disagreement does exist and so
requested that the available time be
extended by 6 months. Specifically, the
commenter indicated that the available
data are not sufficient to support listing
after taking into account various Federal
and State statutes and programs
currently benefiting the three species.
Such statutes and programs include the
Wilderness Act, the Sierra Nevada
Forest Plan, the Clean Water Act, the
California Endangered Species Act, and
the discontinuation of fish stocking by
CDFW in much of the range of the two
frogs.
Our Response: While we agree that
these efforts aid in the conservation of
the three amphibians, we do not
consider substantial disagreement to
exist regarding our conclusion that the
Sierra Nevada yellow-legged frog and
the northern DPS of the mountain
yellow-legged frog meet the definition of
‘‘endangered species’’ under the Act.
We considered the existing Federal and
State statutes and programs in our
determination. The data documenting
population declines and extirpations
associated with Bd and the presence of
introduced fish are sufficient for the
Service to determine that the two
species are ‘‘in danger of extinction
throughout all or a significant portion of
[their] range[s].’’ Data also show that the
Yosemite toad is vulnerable to habitat
changes and climate change, and thus
merits listing as a threatened species,
which is defined as ‘‘likely to become
an endangered species within the
foreseeable future within all or a
significant portion of its range.’’
Available Conservation Measures
Conservation measures provided to
species listed as endangered or
threatened under the Act include
recognition, recovery actions,
requirements for Federal protection, and
prohibitions against certain practices.
Recognition through listing results in
public awareness, and conservation by
Federal, State, Tribal, and local
agencies, private organizations, and
individuals. The Act encourages
cooperation with the States and requires
that recovery actions be carried out for
all listed species. The protection
required by Federal agencies and the
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prohibitions against certain activities
are discussed, in part, below.
The primary purpose of the Act is the
conservation of endangered and
threatened species and the ecosystems
upon which they depend. The ultimate
goal of such conservation efforts is the
recovery of these listed species, so that
they no longer need the protective
measures of the Act. Subsection 4(f) of
the Act requires the Service to develop
and implement recovery plans for the
conservation of endangered and
threatened species. The recovery
planning process involves the
identification of actions that are
necessary to halt or reverse the species’
decline by addressing the threats to its
survival and recovery. The goal of this
process is to restore listed species to a
point where they are secure, selfsustaining, and functioning components
of their ecosystems.
Recovery planning includes the
development of a recovery outline
shortly after a species is listed and
preparation of a draft and final recovery
plan. The recovery outline guides the
immediate implementation of urgent
recovery actions and describes the
process to be used to develop a recovery
plan. Revisions of the plan may be done
to address continuing or new threats to
the species, as new substantive
information becomes available. The
recovery plan identifies site-specific
management actions that set a trigger for
review of the five factors that control
whether a species remains endangered
or may be downlisted or delisted, and
methods for monitoring recovery
progress. Recovery plans also establish
a framework for agencies to coordinate
their recovery efforts and provide
estimates of the cost of implementing
recovery tasks. Recovery teams
(composed of species experts, Federal
and State agencies, nongovernmental
organizations, and stakeholders) are
often established to develop recovery
plans. When completed, the recovery
outline, draft recovery plan, and the
final recovery plan will be available on
our Web site (https://www.fws.gov/
endangered), or from our Sacramento
Fish and Wildlife Office (see FOR
FURTHER INFORMATION CONTACT).
Implementation of recovery actions
generally requires the participation of a
broad range of partners, including other
Federal agencies, States, Tribal,
nongovernmental organizations,
businesses, and private landowners.
Examples of recovery actions include
habitat restoration (e.g., restoration of
native vegetation), research, captive
propagation and reintroduction, and
outreach and education. The recovery of
many listed species cannot be
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accomplished solely on Federal lands
because their range may occur primarily
or solely on non-Federal lands. To
achieve recovery of these species
requires cooperative conservation efforts
on private, State, and Tribal lands.
Following publication of this final
listing rule, funding for recovery actions
will be available from a variety of
sources, including Federal budgets,
State programs, and cost share grants for
non-Federal landowners, the academic
community, and nongovernmental
organizations. In addition, pursuant to
section 6 of the Act, the States of
California and Nevada would be eligible
for Federal funds to implement
management actions that promote the
protection or recovery of the Sierra
Nevada mountain yellow-legged frog,
Northern Distinct Population Segment
of the mountain yellow-legged frog, and
the Yosemite toad. Information on our
grant programs that are available to aid
species recovery can be found at: https://
www.fws.gov/grants.
Please let us know if you are
interested in participating in recovery
efforts for the Sierra Nevada yellowlegged frog, the northern DPS of the
mountain yellow-legged frog, or the
Yosemite toad. Additionally, we invite
you to submit any new information on
these species whenever it becomes
available and any information you may
have for recovery planning purposes
(see FOR FURTHER INFORMATION CONTACT).
Section 7(a) of the Act requires
Federal agencies to evaluate their
actions with respect to any species that
is listed as an endangered or threatened
species and with respect to its critical
habitat, if any is designated. Regulations
implementing this interagency
cooperation provision of the Act are
codified at 50 CFR part 402. Section
7(a)(2) of the Act requires Federal
agencies to ensure that any action
authorized, funded or carried out by
such agency is not likely to jeopardize
the continued existence of the species or
destroy or adversely modify its critical
habitat. If a Federal action may affect a
listed species or its critical habitat, the
responsible Federal agency must enter
into consultation with the Service.
Federal agency actions within the
species’ habitat that may require
consultation, as described in the
preceding paragraph, include
management and any other landscapealtering activities on Federal lands
administered by the USFS, NPS, and
other Federal agencies as appropriate.
The Act and its implementing
regulations set forth a series of general
prohibitions and exceptions that apply
to all endangered and threatened
wildlife. The prohibitions of section
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9(a)(2) of the Act, codified at 50 CFR
17.21 for endangered wildlife, in part,
make it illegal for any person subject to
the jurisdiction of the United States to
take (includes harass, harm, pursue,
hunt, shoot, wound, kill, trap, capture,
or collect; or to attempt any of these),
import, export, ship in interstate
commerce in the course of commercial
activity, or sell or offer for sale in
interstate or foreign commerce any
listed species. Under the Lacey Act (18
U.S.C. 42–43; 16 U.S.C. 3371–3378), it
is also illegal to possess, sell, deliver,
carry, transport, or ship any such
wildlife that has been taken illegally.
Certain exceptions apply to agents of the
Service and State conservation agencies.
We may issue permits to carry out
otherwise prohibited activities
involving endangered and threatened
wildlife species under certain
circumstances. Regulations governing
permits are codified at 50 CFR 17.22 for
endangered species, and at 17.32 for
threatened species. With regard to
endangered wildlife, a permit must be
issued for the following purposes: for
scientific purposes, to enhance the
propagation or survival of the species,
and for incidental take in connection
with otherwise lawful activities.
It is our policy, as published in the
Federal Register on July 1, 1994 (59 FR
34272), to identify to the maximum
extent practicable at the time a species
is listed, those activities that would or
would not constitute a violation of
section 9 of the Act. The intent of this
policy is to increase public awareness of
the effect of a listing on proposed and
ongoing activities within the range of
listed species. The following activities
could potentially result in a violation of
section 9 of the Act; this list is not
comprehensive:
(1) Unauthorized collecting, handling,
possessing, selling, delivering, carrying,
or transporting of the species, including
import or export across State lines and
international boundaries, except for
properly documented antique
specimens of these taxa at least 100
years old, as defined by section 10(h)(1)
of the Act;
(2) Introduction of species that
compete with or prey upon the Sierra
Nevada yellow-legged frog, the northern
DPS of the mountain yellow-legged frog,
or the Yosemite toad;
(3) The unauthorized release of
biological control agents that attack any
life stage of these species;
(4) Unauthorized modification of the
mountain meadow habitats or
associated upland areas important for
the breeding, rearing, and survival of
these species; and
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(5) Unauthorized discharge of
chemicals or fill material into any
waters in which the Sierra Nevada
yellow-legged frog, the northern DPS of
the mountain yellow-legged frog, or the
Yosemite toad are known to occur.
Questions regarding whether specific
activities would constitute a violation of
section 9 of the Act should be directed
to the Sacramento Fish and Wildlife
Office (see FOR FURTHER INFORMATION
CONTACT).
Under section 4(d) of the ESA, the
Secretary has discretion to issue such
regulations as he deems necessary and
advisable to provide for the
conservation of threatened species. Our
implementing regulations (50 CFR
17.31) for threatened wildlife generally
incorporate the prohibitions of section 9
of the Act for endangered wildlife,
except when a ‘‘special rule’’
promulgated pursuant to section 4(d) of
the Act has been issued with respect to
a particular threatened species. In such
a case, the general prohibitions in 50
CFR 17.31 would not apply to that
species, and instead, the special rule
would define the specific take
prohibitions and exceptions that would
apply for that particular threatened
species, which we consider necessary
and advisable to conserve the species.
The Secretary also has the discretion to
prohibit by regulation with respect to a
threatened species any act prohibited by
section 9(a)(1) of the ESA. Exercising
this discretion, which has been
delegated to the Service by the
Secretary, the Service has developed
general prohibitions that are appropriate
for most threatened species in 50 CFR
17.31 and exceptions to those
prohibitions in 50 CFR 17.32. Since we
are not promulgating a special section
4(d) rule, all of the section 9
prohibitions, including the ‘‘take’’
prohibitions, will apply to the Yosemite
toad.
Required Determinations
National Environmental Policy Act (42
U.S.C. 4321 et seq.)
We have determined that
environmental assessments and
environmental impact statements, as
defined under the authority of the
National Environmental Policy Act
(NEPA; 42 U.S.C. 4321 et seq.), need not
be prepared in connection with listing
a species as an endangered or
threatened species under the
Endangered Species Act. We published
a notice outlining our reasons for this
determination in the Federal Register
on October 25, 1983 (48 FR 49244).
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Government-to-Government
Relationship With Tribes
In accordance with the President’s
memorandum of April 29, 1994
(Government-to-Government Relations
with Native American Tribal
Governments; 59 FR 22951), Executive
Order 13175 (Consultation and
Coordination With Indian Tribal
Governments), and the Department of
the Interior’s manual at 512 DM 2, we
readily acknowledge our responsibility
to communicate meaningfully with
recognized Federal Tribes on a
government-to-government basis. In
accordance with Secretarial Order 3206
of June 5, 1997 (American Indian Tribal
Rights, Federal-Tribal Trust
Responsibilities, and the Endangered
Species Act), we readily acknowledge
our responsibilities to work directly
with tribes in developing programs for
healthy ecosystems, to acknowledge that
tribal lands are not subject to the same
controls as Federal public lands, to
remain sensitive to Indian culture, and
to make information available to tribes.
PART 17—[AMENDED]
References Cited
A complete list of references cited in
this rulemaking is available on the
Internet at https://www.regulations.gov
and upon request from the Sacramento
Fish and Wildlife Office (see FOR
FURTHER INFORMATION CONTACT).
■
Authors
The primary authors of this final rule
are the staff members of the Sacramento
Fish and Wildlife Office.
List of Subjects in 50 CFR Part 17
Endangered and threatened species,
Exports, Imports, Reporting and
recordkeeping requirements,
Transportation.
Regulation Promulgation
Accordingly, we amend part 17,
subchapter B of chapter I, title 50 of the
Code of Federal Regulations, as follows:
Species
Vertebrate population where endangered or threatened
Historic range
Common name
Scientific name
*
AMPHIBIANS
*
*
*
1. The authority citation for part 17
continues to read as follows:
Authority: 16 U.S.C. 1361–1407; 1531–
1544; 4201–4245; unless otherwise noted.
2. Amend § 17.11(h), the List of
Endangered and Threatened Wildlife, by
revising the entry for ‘‘Frog, mountain
yellow-legged (southern California
DPS)’’ and adding entries for ‘‘Frog,
mountain yellow-legged (northern
California DPS)’’, ‘‘Frog, Sierra Nevada
yellow-legged’’, and ‘‘Toad, Yosemite’’
to the List of Endangered and
Threatened Wildlife in alphabetical
order under Amphibians to read as
follows:
■
§ 17.11 Endangered and threatened
wildlife.
*
*
*
(h) * * *
Status
*
When listed
*
*
Critical
habitat
*
Special
rules
*
*
Frog, mountain yellow-legged (northern California
DPS).
Frog, mountain yellow-legged (southern California
DPS).
*
Rana muscosa .......
*
U.S.A. (CA) .............
*
U.S.A., northern
California.
*
E
*
834
NA
NA
Rana muscosa .......
U.S.A. (CA) .............
U.S.A., southern
California.
E
728
17.95(d)
NA
*
Frog, Sierra Nevada
yellow-legged.
*
Rana sierrae ...........
*
U.S.A. (CA, NV) .....
*
Entire ......................
*
E
*
834
NA
*
Toad, Yosemite .......
*
Anaxyrus canorus ...
*
U.S.A. (CA) .............
*
Entire ......................
*
T
*
834
NA
*
*
*
*
*
*
*
*
*
*
Dated: April 21, 2014.
Daniel M. Ashe,
Director, U.S. Fish and Wildlife Service.
*
[FR Doc. 2014–09488 Filed 4–25–14; 1:30 pm]
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*
*
NA
*
NA
*
Agencies
[Federal Register Volume 79, Number 82 (Tuesday, April 29, 2014)]
[Rules and Regulations]
[Pages 24255-24310]
From the Federal Register Online via the Government Printing Office [www.gpo.gov]
[FR Doc No: 2014-09488]
[[Page 24255]]
Vol. 79
Tuesday,
No. 82
April 29, 2014
Part IV
Department of the Interior
-----------------------------------------------------------------------
Fish and Wildlife Service
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50 CFR Part 17
Endangered and Threatened Wildlife and Plants; Endangered Species
Status for Sierra Nevada Yellow-Legged Frog and Northern Distinct
Population Segment of the Mountain Yellow-Legged Frog, and Threatened
Species Status for Yosemite Toad; Final Rule
Federal Register / Vol. 79 , No. 82 / Tuesday, April 29, 2014 / Rules
and Regulations
[[Page 24256]]
-----------------------------------------------------------------------
DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS-R8-ES-2012-0100; 4500030113]
RIN 1018-AZ21
Endangered and Threatened Wildlife and Plants; Endangered Species
Status for Sierra Nevada Yellow-Legged Frog and Northern Distinct
Population Segment of the Mountain Yellow-Legged Frog, and Threatened
Species Status for Yosemite Toad
AGENCY: Fish and Wildlife Service, Interior.
ACTION: Final rule.
-----------------------------------------------------------------------
SUMMARY: We, the U.S. Fish and Wildlife Service (Service), determine
endangered species status under the Endangered Species Act of 1973
(Act), as amended, for the Sierra Nevada yellow-legged frog and the
northern distinct population segment (DPS) of the mountain yellow-
legged frog (mountain yellow-legged frog populations that occur north
of the Tehachapi Mountains), and determine threatened species status
under the Act for the Yosemite toad. The effect of this regulation will
be to add these species to the List of Endangered and Threatened
Wildlife.
DATES: This rule becomes effective June 30, 2014.
ADDRESSES: This final rule is available on the Internet at https://www.regulations.gov and at the Sacramento Fish and Wildlife Office.
Comments and materials we received, as well as supporting documentation
used in preparing this rule, are available for public inspection at
https://www.regulations.gov. All of the comments, materials, and
documentation that we considered in this rulemaking are available by
appointment, during normal business hours at: U.S. Fish and Wildlife
Service, Sacramento Fish and Wildlife Office, 2800 Cottage Way, Room W-
2605, Sacramento, CA 95825; 916-414-6600 (telephone); 916-414-6712
(facsimile).
FOR FURTHER INFORMATION CONTACT: Jennifer Norris, Field Supervisor,
U.S. Fish and Wildlife Service, Sacramento Fish and Wildlife Office,
2800 Cottage Way, Room W-2605, Sacramento, CA 95825; 916-414-6600
(telephone); 916-414-6712 (facsimile). Persons who use a
telecommunications device for the deaf (TDD) may call the Federal
Information Relay Service (FIRS) at 800-877-8339.
SUPPLEMENTARY INFORMATION:
Executive Summary
Why we need to publish a rule. Under the Endangered Species Act, a
species may warrant protection through listing if it is endangered or
threatened throughout all or a significant portion of its range.
Listing a species as an endangered or threatened species can be only
completed by issuing a rule.
This rule will finalize the listing of the Sierra Nevada yellow-
legged frog (Rana sierrae) as an endangered species, the northern DPS
of the mountain yellow-legged frog (Rana muscosa) as an endangered
species, and the Yosemite toad (Anaxyrus canorus) as a threatened
species.
The basis for our action. Under the Endangered Species Act, we can
determine that a species is an endangered or threatened species based
on any of 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 (E) Other natural or manmade factors
affecting its continued existence.
We have determined that both the Sierra Nevada yellow-legged frog
and the northern DPS of the mountain yellow-legged frog are presently
in danger of extinction throughout their entire ranges, based on the
immediacy, severity, and scope of the threats to their continued
existence. These include habitat degradation and fragmentation,
predation and disease, climate change, inadequate regulatory
protections, and the interaction of these various stressors impacting
small remnant populations. A rangewide reduction in abundance and
geographic extent of surviving populations of frogs has occurred
following decades of fish stocking, habitat fragmentation, and most
recently a disease epidemic. Surviving populations are smaller and more
isolated, and recruitment in diseased populations is much reduced
relative to historic norms. This combination of population stressors
makes persistence of these species precarious throughout the currently
occupied range in the Sierra Nevada.
We have also determined that the Yosemite toad is likely to become
endangered throughout its range within the foreseeable future, based on
the immediacy, severity, and scope of the threats to its continued
existence. These include habitat loss associated with degradation of
meadow hydrology following stream incision consequent to the cumulative
effects of historical land management activities, notably livestock
grazing, and also the anticipated hydrologic effects upon habitat from
climate change. We also find that the Yosemite toad is likely to become
endangered through the direct effects of climate change impacting small
remnant populations, likely compounded with the cumulative effect of
other threat factors (such as disease).
Peer review and public comment. We sought comments from independent
specialists to ensure that our designations are based on scientifically
sound data, assumptions, and analyses. We invited these peer reviewers
to comment on our listing proposal. We also considered all comments and
information received during the comment period.
Previous Federal Actions
Please refer to the proposed listing rule for the Sierra Nevada
yellow-legged frog, the northern DPS of the mountain yellow-legged
frog, and the Yosemite toad (78 FR 24472, April 25, 2013) for a
detailed description of previous Federal actions concerning these
species.
We will also be finalizing critical habitat designations for the
Sierra Nevada yellow-legged frog, the northern DPS of the mountain
yellow-legged, and the Yosemite toad under the Act in the near future.
Summary of Biological Status and Threats for the Sierra Nevada Yellow-
Legged Frog and the Northern DPS of the Mountain Yellow-Legged Frog
Background
Please refer to the proposed listing rule for the Sierra Nevada
yellow-legged frog and the northern DPS of the mountain yellow-legged
frog under the Act (16 U.S.C. 1531 et seq.) for additional species
information. In the proposed rule, we described two separate species of
yellow-legged frogs, Rana sierrae and Rana muscosa, that resulted from
the recent taxonomic split (see Taxonomy section below) of the
previously known Rana muscosa, which we referred to in our proposed
rule as the mountain yellow-legged frog ``species complex.'' For
clarity and in order to maintain consistency with our previous
treatment of the southern DPS of the mountain yellow legged frog in
southern California (67 FR 44382, July 2, 2002) as well as with our
proposed rule, and for the purposes of this document, we retain the
common name of mountain yellow-legged frog for Rana muscosa, as opposed
to the new common name, southern mountain yellow-legged frog, as
published by
[[Page 24257]]
Crother et al. (2008, p. 11). We also note that the California
Department of Fish and Game (CDFG) was recently renamed the California
Department of Fish and Wildlife (CDFW). We refer to the California
Department of Fish and Wildlife in all cases when discussing the agency
in the text. Where citations are from CDFG documents, we include CDFW
in parentheses for clarification.
Taxonomy
Please refer to the proposed listing rule for the Sierra Nevada
yellow-legged frog and the northern DPS of the mountain yellow-legged
frog under the Act (16 U.S.C. 1531 et seq.) for detailed species
information on taxonomy (78 FR 24472, April 25, 2013).
Vredenburg et al. (2007, p. 371) determined that Rana sierrae
occurs in the Sierra Nevada north of the South Fork Kings River
watershed, along the east slope of the Sierra Nevada south into Inyo
County at the southern extent of its range, and in the Glass Mountains
just south of Mono Lake; and that R. muscosa occurs in the southern
portion of the Sierra Nevada within and south of the South Fork Kings
River watershed to the west of the Sierra Nevada crest (along with
those populations inhabiting southern California) (Vredenburg et al.
2007, pp. 370-371). The Monarch Divide separates these species in the
western Sierra Nevada, while they are separated by the Cirque Crest to
the east (Knapp 2013, unpaginated).
For purposes of this rule, we recognize the species differentiation
as presented in Vredenburg et al. (2007, p. 371) and adopted by the
official societies mentioned above (Crother et al. 2008, p. 11), and in
this final rule we refer to Rana sierrae as the Sierra Nevada yellow-
legged frog, and we refer to the Sierra Nevada populations of R.
muscosa as the northern DPS of the mountain yellow-legged frog. In
California and Nevada, the Sierra Nevada yellow-legged frogs occupy the
western Sierra Nevada north of the Monarch Divide (in Fresno County)
and the eastern slope of the Sierra Nevada (east of the crest) from
Inyo County through Mono County (including the Glass Mountains), to
areas north of Lake Tahoe. The northern DPS of the mountain yellow-
legged frog occurs only in California in the western Sierra Nevada and
extends from south of the Monarch Divide in Fresno County through
portions of the Kern River drainage. Figure 1 shows the approximate
species boundaries within their historical ranges as determined by
Knapp (unpubl. data).
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Many studies cited in the rest of this document include articles
and reports that were published prior to the official species
reclassification, where the researchers may reference either one or
both species. Where possible and appropriate, information will be
referenced specifically (either as Sierra Nevada yellow-legged frog or
the northern DPS of the mountain yellow-legged frog) to reflect the
split of the species. Where information applies to both species, the
two species will be referred to collectively as mountain yellow-legged
frog or mountain yellow-legged frog species complex.
Species Description
Please refer to the proposed listing rule for the Sierra Nevada
yellow-legged frog and the northern DPS of the mountain yellow-legged
frog under the Act (16 U.S.C. 1531 et seq.) for additional information
about species descriptions (78 FR 24472, April 25, 2013). The body
lengths (snout to vent) of the mountain yellow-legged frogs range from
40 to 80 millimeters (mm) (1.5 to 3.25 inches (in)) (Jennings and Hayes
1994, p. 74). Females average slightly larger than males, and males
have a swollen, darkened thumb base (Wright and Wright 1949, pp. 424-
430; Stebbins 1951, pp. 330-335; Zweifel 1955, p. 235; Zweifel 1968, p.
65.1). Dorsal (upper) coloration in adults is variable, exhibiting a
mix of brown and yellow, but also can be grey, red, or green-brown, and
is usually patterned with dark spots (Jennings and Hayes 1994, p. 74;
Stebbins 2003, p. 233). These spots may be large (6 mm (0.25 in)) and
few, smaller and more numerous, or a mixture of both (Zweifel 1955, p.
230). Irregular lichen- or moss-like patches (to which the name muscosa
refers) may also be present on the dorsal surface (Zweifel 1955, pp.
230, 235; Stebbins 2003, p. 233).
[[Page 24259]]
The belly and undersurfaces of the hind limbs are yellow or orange,
and this pigmentation may extend forward from the abdomen to the
forelimbs (Wright and Wright 1949, pp. 424-429; Stebbins 2003, p. 233).
Mountain yellow-legged frogs may produce a distinctive mink or garlic-
like odor when disturbed (Wright and Wright 1949, p. 432; Stebbins
2003, p. 233). Although these species lack vocal sacs, they can
vocalize in or out of water, producing what has been described as a
faint clicking sound (Zweifel 1955, p. 234; Ziesmer 1997, pp. 46-47;
Stebbins 2003, p. 233). Mountain yellow-legged frogs have smoother
skin, generally with heavier spotting and mottling dorsally, darker toe
tips (Zweifel 1955, p. 234), and more opaque ventral coloration
(Stebbins 2003, p. 233) than the foothill yellow-legged frog.
The Sierra Nevada yellow-legged frog and the northern DPS of the
mountain yellow-legged frog are similar morphologically and
behaviorally (hence their shared taxonomic designation until recently).
However, these two species can be distinguished from each other
physically by the ratio of the lower leg (fibulotibia) length to snout
vent length. The northern DPS of the mountain yellow-legged frog has
longer limbs (Vredenburg et al. 2007, p. 368). Typically, this ratio is
greater than or equal to 0.55 in the northern DPS of the mountain
yellow-legged frog and less than 0.55 in the Sierra Nevada yellow-
legged frog.
Mountain yellow-legged frogs deposit their eggs in globular clumps,
which are often somewhat flattened and roughly 2.5 to 5 centimeters
(cm) (1 to 2 in) in diameter (Stebbins 2003, p. 444). When eggs are
close to hatching, egg mass volume averages 198 cubic cm (78 cubic in)
(Pope 1999, p. 30). Eggs have three firm, jelly-like, transparent
envelopes surrounding a grey-tan or black vitelline (egg yolk) capsule
(Wright and Wright 1949, pp. 431-433). Clutch size varies from 15 to
350 eggs per egg mass (Livezey and Wright 1945, p. 703; Vredenburg et
al. 2005, p. 565). Egg development is temperature dependent. In
laboratory breeding experiments, egg hatching time ranged from 18 to 21
days at temperatures of 5 to 13.5 degrees Celsius ([deg]C) (41 to 56
degrees Fahrenheit ([deg]F)) (Zweifel 1955, pp. 262-264). Field
observations show similar results (Pope 1999, p. 31).
The tadpoles of mountain yellow-legged frogs generally are mottled
brown on the dorsal side with a faintly yellow venter (underside)
(Zweifel 1955, p. 231; Stebbins 2003, p. 460). Total tadpole length
reaches 72 mm (2.8 in), the body is flattened, and the tail musculature
is wide (about 2.5 cm (1 in) or more) before tapering into a rounded
tip (Wright and Wright 1949, p. 431). The mouth has a maximum of eight
labial (lip) tooth rows (two to four upper and four lower) (Stebbins
2003, p. 460). Tadpoles may take more than 1 year (Wright and Wright
1949, p. 431), and often require 2 to 4 years, to reach metamorphosis
(transformation from tadpoles to frogs) (Cory 1962b, p. 515; Bradford
1983, pp. 1171, 1182; Bradford et al. 1993, p. 883; Knapp and Matthews
2000, p. 435), depending on local climate conditions and site-specific
variables.
The time required to reach reproductive maturity in mountain
yellow-legged frogs is thought to vary between 3 and 4 years post
metamorphosis (Zweifel 1955, p. 254). This information, in combination
with the extended amount of time as a tadpole before metamorphosis,
means that it may take 5 to 8 years for mountain yellow-legged frogs to
begin reproducing. While the typical lifespan of mountain yellow-legged
frogs is largely unknown, Matthews and Miaud (2007, p. 991) estimated
that the total lifespan (including tadpole and adult life stages)
ranges up to 14 years, with other documented estimates of up to 16
years of age for the Sierra Nevada yellow-legged frog (Fellers et al.
2013, p. 155), suggesting that mountain yellow-legged frogs are long-
lived amphibians.
Habitat and Life History
Mountain yellow-legged frogs currently exist in montane regions of
the Sierra Nevada of California. Throughout their range, these species
historically inhabited lakes, ponds, marshes, meadows, and streams at
elevations typically ranging from 1,370 to 3,660 meters (m) (4,500 to
12,000 feet (ft)) ((CDFG (CDFW)) 2011, pp. A-1-A-5), but can occur as
low as 1,067 m (3,500 ft) in the northern portions of their range (USFS
2011, geospatial data; USFS 2013, p. 4). Mountain yellow-legged frogs
are highly aquatic; they are generally not found more than 1 m (3.3 ft)
from water (Stebbins 1951, p. 340; Mullally and Cunningham 1956a, p.
191; Bradford et al. 1993, p. 886). Mullally and Cunningham (1956a, p.
191) found adults sitting on rocks along the shoreline, where there was
little or no vegetation. Although mountain yellow-legged frogs may use
a variety of shoreline habitats, both tadpoles and adults are observed
less frequently at shorelines that drop abruptly to a depth of 60 cm (2
ft) than at open shorelines that gently slope up to shallow waters of
only 5 to 8 cm (2 to 3 in) in depth (Mullally and Cunningham 1956a, p.
191; Jennings and Hayes 1994, p. 77).
At lower elevations within their historical range, these species
have been associated with rocky streambeds and wet meadows surrounded
by coniferous forest (Zweifel 1955, p. 237; Zeiner et al. 1988, p. 88),
although, in general, little is known about the ecology of mountain
yellow-legged frogs in Sierra Nevada stream habitats (Brown 2013,
unpaginated). Zweifel (1955, p. 237) found that streams utilized by
adults varied from streams having high gradients and numerous pools,
rapids, and small waterfalls, to streams with low gradients and slow
flows, marshy edges, and sod banks, while aquatic substrates varied
from bedrock to fine sand, rubble (rock fragments), and boulders.
Jennings and Hayes (1994, p. 77) have indicated that mountain yellow-
legged frogs appear absent from the smallest creeks, and suggest that
it is probably because these creeks have insufficient depth for
adequate refuge and overwintering habitat. However, Brown (2013,
unpaginated) reports that the frogs are found in small creeks, although
she notes that the extent to which these are remnant populations now
excluded from preferred habitat is not known. In the northern portion
of the Sierra Nevada yellow-legged frog range, the remnant populations
primarily occur in stream habitats.
At higher elevations, these species occupy lakes, ponds, tarns
(small steep-banked mountain lakes or pools, generally of glacial
origin), and streams (Zweifel 1955, p. 237; Mullally and Cunningham
1956a, p. 191). Mountain yellow-legged frogs in the Sierra Nevada are
most abundant in high-elevation lakes and slow-moving portions of
streams (Zweifel 1955, p. 237; Mullally and Cunningham 1956a, p. 191).
The borders of alpine (above the tree line) lakes and mountain meadow
streams used by mountain yellow-legged frogs are frequently grassy or
muddy, although many are bordered by exposed glaciated bedrock. Zweifel
(1955, pp. 237-238) suggested that alpine lakeshores differ from the
sandy or rocky shores inhabited by mountain yellow-legged frogs in
lower elevation streams.
Adult mountain yellow-legged frogs breed in a variety of habitats
including the shallows of stillwater habitat (lakes or ponds) and
flowing inlet streams (Zweifel 1955, p. 243; Pope 1999, p. 30). Adults
emerge from overwintering sites immediately following snowmelt, and
will even move over ice to reach breeding sites (Pope 1999, pp. 46-47;
Vredenburg et al. 2005, p. 565). Mountain yellow-legged frogs deposit
[[Page 24260]]
their eggs underwater in clusters, which they attach to rocks, gravel,
or vegetation, or which they deposit under banks (Wright and Wright
1949, p. 431; Stebbins 1951, p. 341; Zweifel 1955, p. 243; Pope 1999,
p. 30).
Lake depth is an important attribute defining habitat suitability
for mountain yellow-legged frogs. At high elevations, both frogs and
tadpoles overwinter under ice in lakes and streams. As tadpoles must
overwinter multiple years before metamorphosis, successful breeding
sites are located in (or connected to) lakes and ponds that do not dry
out in the summer, and also are deep enough that they do not completely
freeze or become oxygen-depleted (anoxic) in winter. Both adults and
tadpole mountain yellow-legged frogs overwinter for up to 9 months in
the bottoms of lakes that are at least 1.7 m (5.6 ft) deep; however,
overwinter survival may be greater in lakes that are at least 2.5 m
(8.2 ft) deep (Bradford 1983, p. 1179; Vredenburg et al. 2005, p. 565).
Bradford (1983, pp. 1173, 1178-1179) found that, in years with
exceptional precipitation (61 percent above average) and greater than
normal ice-depths, mountain yellow-legged frog die-offs sometimes
result from oxygen depletion during winter in lakes less than 4 m (13
ft) in depth, finding that in ice-covered lakes, oxygen depletion
occurs most rapidly in shallow lakes relative to deeper lakes. However,
tadpoles may survive for months in nearly anoxic conditions when
shallow lakes are frozen to the bottom. More recent work reported
populations of mountain yellow-legged frogs overwintering in lakes less
than 1.5 m (5 ft) deep that were assumed to have frozen to the bottom,
and yet healthy frogs emerged the following July (Matthews and Pope
1999, pp. 622-623; Pope 1999, pp. 42-43). Matthews and Pope 1999, p.
619) used radio telemetry to find that, when lakes had begun to freeze
over, the frogs were utilizing rock crevices, holes, and ledges near
shore, where water depths ranged from 0.2 m (0.7 ft) to 1.5 m (5 ft).
Vredenburg et al. (2005, p. 565) noted that such behavior may be a
response to presence of introduced fish. Matthews and Pope (1999, p.
622) suggested that the granite surrounding these overwintering
habitats probably insulates mountain yellow-legged frogs from extreme
winter temperatures, and that they can survive, provided there is an
adequate supply of oxygen.
Mountain yellow-legged frog tadpoles maintain a relatively high
body temperature by selecting warmer microhabitats (Bradford 1984, p.
973). During winter, tadpoles remain in warmer water below the
thermocline (the transition layer between thermally stratified water).
After spring overturn (thaw and thermal mixing of the water), they
behaviorally modulate their body temperature by moving to shallow,
near-shore water when warmer days raise surface water temperatures.
During the late afternoon and evening, mountain yellow-legged frogs
retreat to offshore waters that are less subject to night cooling
(Bradford 1984, p. 974).
Available evidence suggests that adult mountain yellow-legged frogs
display strong site fidelity and return to the same overwintering and
summer habitats from year to year (Pope 1999, p. 45; Matthews and
Preisler 2010, p. 252). Matthews and Pope (1999, pp. 618-623) observed
that the frogs' movement patterns and habitat associations shifted
seasonally. Frogs were well-distributed in most lakes, ponds, and
creeks during August, but moved to only a few lakes by October.
Matthews and Pope (1999, pp. 618-623) established home-range areas for
10 frogs and found that frogs remained through August in the lake or
creek where they'd been captured, with movement confined to areas
ranging from 19.4 to 1,028 square meters (m\2\) (23.20 to 1,229 square
yards (y\2\)). In September, movements increased, with home-ranges
varying from 53 to 9,807 m\2\ in size (63.4 to 11,729 y\2\); six of
nine frogs tagged in September moved from that lake by the end of the
month, suggesting a pattern in which adult mountain yellow-legged frogs
move among overwintering, breeding, and feeding sites during the year,
with narrow distributions in early spring and late fall due to
restricted overwintering habitat (Pope and Matthews 2001, p. 791).
Although terrestrial movements of more than two or three hops from
water were previously undocumented, overland movements exceeding 66 m
(217 ft) were observed in 17 percent of tagged frogs, demonstrating
that mountain yellow-legged frogs move overland as well as along
aquatic pathways (Pope and Matthews 2001, p. 791). Pope and Matthews
(2001, p. 791) also recorded a movement distance of over 1 km
(including a minimum of 420 m (0.26 miles) overland movement and
movement through a stream course). The farthest reported distance of a
mountain yellow-legged frog from water is 400 m (1,300 ft) (Vredenburg
2002, p. 4).
Within stream systems, Sierra Nevada yellow-legged frogs have been
documented to move 1,032 m (3,385 ft) over a 29-day period (Fellers et
al. 2013, p. 159). Wengert (2008, p. 18) conducted a telemetry study
that documented single-season movement distances for Sierra Nevada
yellow-legged frog of up to 3.3 kilometers (km) (2.05 miles (mi)) along
streams. Along stream habitats, adults have been observed greater than
22 m (71 ft) from the water during the overwintering period (Wengert
2008, p. 20). Additionally, during the duration of the study, Wengert
(2008, p. 13) found that 14 percent of the documented frog locations
occurred greater than 0.2 m (0.66 ft) from the stream edge. While
recent information suggests that the frogs in the Wengert study may
have actually been foothill yellow-legged frog (Rana boylii) (Poorten
et al., 2013, p. 4), we expect that the movement distances recorded are
applicable to the Sierra Nevada yellow-legged frog within a stream-
based system, as the ecology is comparable between the two sister taxa
in regard to stream systems.
Almost no data exist on the dispersal of juvenile mountain yellow-
legged frogs away from breeding sites; however, juveniles that may be
dispersing have been observed in small intermittent streams (Bradford
1991, p. 176). Regionally, mountain yellow-legged frogs are thought to
exhibit a metapopulation structure (Bradford et al. 1993, p. 886; Drost
and Fellers 1996, p. 424). Metapopulations are spatially separated
population subunits within migratory distance of one another such that
individuals may interbreed among subunits and populations may become
reestablished if they are extirpated (Hanski and Simberloff 1997, p.
6).
Historical Range and Distribution
Mountain yellow-legged frogs were historically abundant and
ubiquitous across many of the higher elevations within the Sierra
Nevada. Grinnell and Storer (1924, p. 664) reported the Sierra Nevada
yellow-legged frog to be the most common amphibian surveyed in the
Yosemite area. It is difficult to know the precise historical ranges of
the Sierra Nevada yellow-legged frog and the northern DPS of the
mountain yellow-legged frog, because projections must be inferred from
museum collections that do not reflect systematic surveys, and survey
information predating significant rangewide reduction is very limited.
However, projections of historical ranges are available using
predictive habitat modeling based on recent research (Knapp, unpubl.
data).
Historically, the range of the Sierra Nevada yellow-legged frog
extended in California from north of the Feather River, in Butte and
Plumas Counties, south to the Monarch Divide on the west side of the
Sierra Nevada crest in Fresno County. East of the Sierra Nevada crest
in California, the historical
[[Page 24261]]
range of the Sierra Nevada yellow-legged frog extends from areas north
of Lake Tahoe, through Mono County (including the Glass Mountains) to
Inyo County. Historical records indicate that the Sierra Nevada yellow-
legged frog also occurred at locations within the Carson Range of
Nevada, including Mount Rose in Washoe County, and also occurred in the
vicinity of Lake Tahoe in Douglas County, Nevada (Linsdale 1940, pp.
208-210; Zweifel 1955, p. 231; Jennings 1984, p. 52; Knapp 2013,
unpaginated).
Historically, the northern DPS of the mountain yellow-legged frog
ranged from the Monarch Divide in Fresno County as far southward as
Breckenridge Mountain, in Kern County (Vredenburg et al. 2007, p. 371).
The historical ranges of the two frog species within the mountain
yellow-legged complex, therefore, meet each other roughly along the
Monarch Divide to the north, and along the crest of the Sierra Nevada
to the east. Because we have determined that the historic range of R.
muscosa is entirely within the State of California, in this final rule
we correct the listing for the southern DPS of the mountain yellow-
legged frog to remove Nevada from its historic range.
Current Range and Distribution
Since the time of the mountain yellow-legged frog observations of
Grinnell and Storer (1924, pp. 664-665), a number of researchers have
reported disappearances of these species from a large fraction of their
historical ranges in the Sierra Nevada (Hayes and Jennings 1986, p.
490; Bradford 1989, p. 775; Bradford et al. 1994, pp. 323-327; Jennings
and Hayes 1994, p. 78; Jennings 1995, p. 133; Stebbins and Cohen 1995,
pp. 225-226; Drost and Fellers 1996, p. 414; Jennings 1996, pp. 934-
935; Knapp and Matthews 2000, p. 428; Vredenburg et al. 2005, p. 564).
The current distributions of the Sierra Nevada yellow-legged frog
and the northern DPS of the mountain yellow-legged frog are restricted
primarily to publicly managed lands at high elevations, including
streams, lakes, ponds, and meadow wetlands located within National
Forests and National Parks. National Forests with extant (surviving)
populations of mountain yellow-legged frogs include the Plumas National
Forest, Tahoe National Forest, Humboldt-Toiyabe National Forest, Lake
Tahoe Basin Management Unit, Eldorado National Forest, Stanislaus
National Forest, Sierra National Forest, Sequoia National Forest, and
Inyo National Forest. National Parks with extant populations of
mountain yellow-legged frogs include Yosemite National Park, Kings
Canyon National Park, and Sequoia National Park.
The most pronounced declines within the mountain yellow-legged frog
complex have occurred north of Lake Tahoe in the northernmost 125-km
(78-mi) portion of the range (Sierra Nevada yellow-legged frog) and
south of Kings Canyon National Park in Tulare County (the northern DPS
of the mountain yellow-legged frog). In the southernmost 50-km (31-mi)
portion of the range, only a few populations of the northern DPS of the
mountain yellow-legged frog remain (Fellers 1994, p. 5; Jennings and
Hayes 1994, pp. 74-78); except for a few small populations in the Kern
River drainage, the northern DPS of the mountain yellow-legged frog is
entirely extirpated from all of Sequoia National Park (Knapp 2013,
unpaginated). As of 2000, mountain yellow-legged frog populations were
known to have persisted in greater density in the National Parks of the
Sierra Nevada as compared to the surrounding U.S. Forest Service (USFS)
lands, and the populations that did occur in the National Parks
generally exhibited higher abundances than those on USFS lands
(Bradford et al. 1994, p. 323; Knapp and Matthews 2000, p. 430).
Population Estimates and Status
Monitoring efforts and research studies have documented substantial
declines of mountain yellow-legged frog populations in the Sierra
Nevada. The number of extant populations has declined greatly over the
last few decades. Remaining populations are patchily scattered
throughout the historical range (Jennings and Hayes 1994, pp. 74-78;
Jennings 1995, p. 133; Jennings 1996, p. 936). In the northernmost
portion of the range (Butte and Plumas Counties), only a few Sierra
Nevada yellow-legged frog populations have been documented since 1970
(Jennings and Hayes 1994, pp. 74-78; CDFG (CDFW) et al., unpubl. data).
Declines of both species have also been noted in the central and
southern Sierra Nevada (Drost and Fellers 1996, p. 420; Knapp and
Matthews 2001, pp. 433-437; Knapp 2013, unpaginated). In the southern
Sierra Nevada (Sierra, Sequoia, and Inyo National Forests; and Kings
Canyon and Yosemite National Parks), modest to relatively large
populations (for example, breeding populations of approximately 40 to
more than 200 adults) of mountain yellow-legged frogs do remain;
however, in recent years some large populations have been extirpated in
this area (Bradford 1991, p. 176; Bradford et al. 1994, pp. 325-326;
Knapp 2002a, p. 10, Wake and Vredenburg 2009, pp. 11467-11470).
Davidson et al. (2002, p. 1591) reviewed 255 previously documented
mountain yellow-legged frog locations (based on Jennings and Hayes
1994, pp. 74-78) throughout the historical range and concluded that 83
percent of these sites no longer support frog populations. Vredenburg
et al. (2007, pp. 369-371) compared recent survey records (1995-2004)
with museum records from 1899-1994 and reported that 92.5 percent of
historical Sierra Nevada yellow-legged frog populations and 92.3
percent of populations of the northern DPS of mountain yellow-legged
frog are now extirpated.
CDFW (CDFG (CDFW) 2011, pp. 17-20) used historical localities from
museum records covering the same time interval (1899-1994), but updated
recent locality information with additional survey data (1995-2010) to
significantly increase proportional coverage from the Vredenburg et al.
(2007) study. These more recent surveys failed to detect any extant
frog populations (within 1 km (0.63 mi), a metric used to capture
interbreeding individuals within metapopulations) at 220 of 318
historical Sierra Nevada yellow-legged frog localities and 94 of 109
historical northern DPS of the mountain yellow-legged frog localities
(in the Sierran portion of their range). This calculates to an
estimated loss of 69 percent of Sierra Nevada yellow-legged frog
metapopulations and 86 percent of northern DPS of the mountain yellow-
legged frog metapopulations from historical occurrences.
In addition to comparisons based on individual localities, CDFW
(CDFG 2011, pp. 20-25) compared historical and recent population status
at the watershed scale. This is a rough index of the geographic extent
of the species through their respective ranges. Within the Sierra
Nevada, 44 percent of watersheds historically utilized by Sierra Nevada
yellow-legged frogs, and 59 percent of watersheds historically utilized
by northern DPS mountain yellow-legged frogs, no longer support extant
populations. However, this watershed-level survey methodology is not a
good indicator of population changes because a watershed is counted as
recently occupied if a single individual (at any life stage) is
observed within the entire watershed even though several individual
populations may have been lost (CDFG (CDFW) 2011b, p. 20). Therefore,
these surveys likely underestimate population declines. Many watersheds
support only a single extant metapopulation, which occupies one to
several adjacent water bodies
[[Page 24262]]
(CDFG (CDFW) 2011, p. 20). Remaining populations are generally very
small.
Rangewide, declines of mountain yellow-legged frog populations were
estimated at around one-half of historical populations by the end of
the 1980s (Bradford et al. 1994, p. 323). Between 1988 and 1991,
Bradford et al. (1994a, pp. 323-327) resurveyed sites known
historically (1955 through 1979 surveys) to support mountain yellow-
legged frogs. They did not detect frogs at 27 historical sites on the
Kaweah River, and they detected frogs at 52 percent of historical sites
within Sequoia and Kings Canyon National Parks and 12.5 percent of
historical sites outside of Sequoia and Kings Canyon National Parks.
Because this work was completed before the taxonomic division of
mountain yellow-legged frogs, we have not differentiated between the
two species here. When both species are combined, this resurvey effort
detected mountain yellow-legged frogs at 19.4 percent of historical
sites (Bradford et al. 1994, pp. 324-325).
Available information discussed below indicates that the rates of
population decline have not abated, and they have likely accelerated
during the 1990s into the 2000s. Drost and Fellers (1996, p. 417)
repeated Grinnell and Storer's early 20th century surveys in Yosemite
National Park, and reported frog presence at 2 of 14 historical sites
where what is now known as Sierra Nevada yellow-legged frogs occurred.
The two positive sightings consisted of a single tadpole at one site
and a single adult female at another. They identified 17 additional
sites with suitable mountain yellow-legged frog habitat, and in those
surveys, they detected 3 additional populations. In 2002, Knapp (2002a,
p. 10) resurveyed 302 water bodies known to be occupied by mountain
yellow-legged frogs between 1995 and 1997, and 744 sites where frogs
were not previously detected. Knapp found frogs at 59 percent of the
previously occupied sites, whereas 8 percent of previously unoccupied
sites were colonized. These data suggest an extirpation rate five to
six times higher than the colonization rate within this study area. The
documented extirpations appeared to occur non-randomly across the
landscape, were typically spatially clumped, and involved the
disappearance of all or nearly all of the mountain yellow-legged frog
populations in a watershed (Knapp 2002a, p. 9). CDFW (CDFG 2011, p. 20)
assessed data from sites where multiple surveys were completed after
1995 (at least 5 years apart). They found that the Sierra Nevada
yellow-legged frog was not detected at 45 percent of sites where they
previously had been confirmed, while the mountain yellow-legged frog
(rangewide, including southern California) was no longer detectable at
81 percent of historically occupied sites.
The USFS has been conducting a rangewide, long-term monitoring
program for the Sierra Nevada yellow-legged frog and the northern DPS
of the mountain yellow-legged frog on National Forest lands in the
Sierra Nevada, known as the Sierra Nevada Amphibian Monitoring Program
(SNAMPH). This monitoring effort provides unbiased estimates by using
an integrated unequal probability design, and it provides numbers for
robust statistical comparisons across 5-year monitoring cycles spanning
208 watersheds (Brown et al. 2011, pp. 3-4). The results of this
assessment indicate that the species have declined in both distribution
and abundance. Based on surveys conducted from 2002 through 2009,
breeding activity was found in about half (48 percent) of the
watersheds where the species were found in the decade prior to SNAMPH
monitoring (1990 and 2001) (Brown et al. 2011, p. 4). Breeding was
found in 3 percent of watersheds where species had been found prior to
1990. Rangewide, breeding was found in 4 percent of watersheds.
Moreover, relative abundances were low; an estimated 9 percent of
populations were large (numbering more than 100 frogs or 500 tadpoles);
about 90 percent of the watersheds had fewer than 10 adults, while 80
percent had fewer than 10 subadults and 100 tadpoles (Brown et al.
2011, p. 24).
To summarize population trends over the available historical
record, estimates range from losses between 69 to 93 percent of Sierra
Nevada yellow-legged frog populations and 86 to 92 percent of the
northern DPS of the mountain yellow-legged frog. Rangewide reduction
has diminished the number of watersheds that support mountain yellow-
legged frogs somewhere between the conservative estimates of 44 percent
in the case of Sierra Nevada yellow-legged frogs and at least 59
percent in the case of the northern DPS of the mountain yellow-legged
frogs, to as high as 97 percent of watersheds for the mountain yellow-
legged frog complex across the Sierra Nevada. Remaining populations are
much smaller than historical norms, and the density of populations per
watershed has declined substantially; as a result, many watersheds
currently support single metapopulations at low abundances.
Distinct Vertebrate Population Segment Analysis
Under the Act, we must consider for listing any species,
subspecies, or, for vertebrates, any DPS of these taxa if there is
sufficient information to indicate that such action may be warranted.
To implement the measures prescribed by the Act, we, along with the
National Marine Fisheries Service (National Oceanic and Atmospheric
Administration-Fisheries), developed a joint policy that addresses the
recognition of DPSs for potential listing actions (61 FR 4722). The
policy allows for a more refined application of the Act that better
reflects the biological needs of the taxon being considered and avoids
the inclusion of entities that do not require the Act's protective
measures.
Under our DPS policy, three elements are considered in a decision
regarding the status of a possible DPS as endangered or threatened
under the Act. The elements are: (1) 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. In other words,
if we determine that a population segment of a vertebrate species being
considered for listing is both discrete and significant, we would
conclude that it represents a DPS, and thus a ``species'' under section
3(16) of the Act, whereupon we would evaluate the level of threat to
the DPS based on the five listing factors established under section
4(a)(1) of the Act to determine whether listing the DPS as an
``endangered species'' or a ``threatened species'' is warranted.
Please refer to the proposed listing rule for detailed information
about the distinct vertebrate population segment analysis for the
northern DPS of the mountain yellow-legged frog (78 FR 24472, April 25,
2013). We previously confirmed the status of the southern California
population of the mountain yellow-legged frog as a DPS at the time that
it was listed as endangered under the Act (67 FR 44382, pp. 44384-
44385). We summarize below the analysis for discreteness and
significance for the northern California population of the mountain
yellow-legged frog (in the Sierra Nevada); this summary includes
changes from the proposed rule to address comments received from the
public (78 FR 24472, April 25, 2013).
[[Page 24263]]
Discreteness
Under our DPS Policy, a population segment of a vertebrate species
may be considered discrete if it satisfies either 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 may provide evidence of this separation); or (2) it is
delimited by international governmental boundaries within which
significant differences in control of exploitation, management of
habitat, conservation, status, or regulatory mechanisms exist.
The analysis of the northern population segment of the mountain
yellow-legged frog (Rana muscosa) (in the Sierra Nevada) is based on
the marked separation from other populations. The range of the mountain
yellow-legged frog is divided by a natural geographic barrier, the
Tehachapi Mountains, which physically isolates the populations in the
southern Sierra Nevada from those in the mountains of southern
California. The distance of the geographic separation is about 225 km
(140 mi). The geographic separation of the Sierra Nevada and southern
California frogs was recognized in the earliest description of the
species by Camp (1917), who treated frogs from the two areas as
separate subspecies within the R. boylii group (see more on
classification of the mountain yellow-legged frogs in Taxonomy). There
is no contiguous habitat that provides connectivity between the two
populations that is sufficient for the migration, growth, rearing, or
reproduction of dispersing frogs. Genetic differences well-supported in
the scientific literature also provide evidence of this separation (see
Taxonomy). Therefore, we find that the northern population segment of
the mountain yellow-legged frog (Rana muscosa) (in the Sierra Nevada)
is discrete from the remainder of the species.
Significance
Under our DPS Policy, once we have determined that a population
segment is discrete, we consider its biological and ecological
significance to the larger taxon to which it belongs. Our DPS policy
provides several potential considerations that may demonstrate the
significance of a population segment to the remainder of its taxon,
including: (1) Evidence of the persistence of the discrete population
segment in an ecological setting unusual or unique for the taxon, (2)
evidence that loss of the discrete population segment would result in a
significant gap in the range of the taxon, (3) evidence that the
population segment represents the only surviving natural occurrence of
a taxon that may be more abundant elsewhere as an introduced population
outside its historic range, or (4) evidence that the discrete
population segment differs markedly from the remainder of the species
in its genetic characteristics.
We have found substantial evidence that three of the four
significance criteria are met by the discrete northern population
segment of the mountain yellow-legged frog that occurs in the Sierra
Nevada. These include its persistence in an ecological setting that is
unique for the taxon, evidence that its loss would result in a
significant gap in the range of the taxon, and its genetic uniqueness
(reflecting significant reproductive isolation over time). To establish
the significance of the discrete northern population segment, we rely
on the effect that the loss of this population segment would have on
the range of the taxon, and supplement that with evidence that the
population segment persists in an ecological setting unusual or unique
for the taxon and also differs from other population segments in its
genetic characteristics. There are no introduced populations of the
northern DPS of the mountain yellow-legged frog outside of the species'
historical range.
Evidence indicates that loss of the northern population segment of
the mountain yellow-legged frog (in the Sierra Nevada) would result in
a significant gap in the range of the taxon. The Sierran mountain
yellow-legged frogs comprise the entire distribution of the species in
approximately the northern half of the species' range, and loss of the
distinct population segment in the northern portion of the range could
have significant conservation implications for the species.
Furthermore, loss of the northern population segment of the mountain
yellow-legged frog (in the Sierra Nevada) would reduce the species to
the remaining small, isolated sites in the streams of southern
California (USFWS, Jul 2012, pp. 11-12). Loss of the northern
population segment of the mountain yellow-legged frog would leave an
area of the southern Sierra Nevada over 150 km (93 mi) in length
without any ranid (frogs in the genus Ranidae) frogs, which were once
abundant and widespread in the higher elevation Sierra Nevada (Cory
1962b, p. 515; Fellers 1994, p. 5). The potential loss of the northern
population segment of the mountain yellow-legged frog would constitute
a significant gap in the range of the species.
One of the most striking differences between the northern
population segment and the southern population segment of the mountain
yellow-legged frogs is the difference in the ecological setting in
which they each persist. Zweifel (1955, pp. 237-241) observed that the
frogs in southern California are typically found in steep-gradient
streams in the chaparral belt at low elevations (370 m (1,220 ft)),
even though they may range into small meadow streams at higher
elevations up to 2,290 m (7,560 ft). In contrast, frogs from the
northern population segment of mountain yellow-legged frogs are most
abundant in high-elevation lakes and slow-moving portions of streams
where winter conditions are extreme. David Bradford's (1989) southern
Sierra Nevada study of mountain yellow-legged frogs, for example, was
conducted in Sequoia and Kings Canyon National Parks at high elevations
between 2,910 and 3,430 m (9,600 to 11,319 ft). The rugged canyons of
the arid mountain ranges of southern California, where waters seldom
freeze, bear little resemblance to the alpine lakes and streams of the
Sierra Nevada where adult frogs and tadpoles must overwinter at the
bottoms of ice and snow-covered lakes for up to 9 months of the year.
The significantly different ecological settings between mountain
yellow-legged frogs in southern California and those in the northern
population segment (in the Sierra Nevada) distinguish these populations
from each other.
Finally, the northern population segment of the mountain yellow-
legged frog is biologically significant based on genetic differences.
Vredenburg et al. (2007, p. 361) identified that two of three distinct
genetic clades (groups of distinct lineage) constitute the northern
range of the mountain yellow-legged frog found in the Sierra Nevada,
with the remaining clade represented by the endangered southern
California DPS of the mountain yellow-legged frog. Macey et al. (2001,
p. 141) estimated the genetic divergence between the northern
population of mountain yellow-legged frogs (in the Sierra Nevada) and
the southern population of mountain yellow-legged frogs (in southern
California) to have occurred 1.4 million years before present (mybp),
thereby indicating functional isolation.
The loss of the northern population of the mountain yellow-legged
frog would result in a significant gap in the range of the mountain
yellow-legged frog species. The differences between the ecological
settings for the southern
[[Page 24264]]
population of mountain yellow-legged frogs (steep-gradient streams that
seldom freeze) and the northern population of mountain yellow-legged
frogs (high-elevation lakes and slow-moving portions of streams where
frogs overwinter under ice and snow for up to 75 percent of the year)
are significant. Additionally, the genetic distinction between these
two populations reflects isolation for over a million years. Therefore
based on the information discussed above, we find that northern
population of the mountain yellow-legged frog (in the Sierra Nevada
mountains) meets the significance criteria under our Policy Regarding
the Recognition of Distinct Vertebrate Population Segments (61 FR
4722).
DPS Conclusion
Based on the best scientific and commercial data available on
distribution as well as ecological setting and genetic characteristics
of the species, we have determined that the northern population segment
of the mountain yellow-legged frog (in the Sierra Nevada) is both
discrete and significant per our DPS policy. Therefore, we conclude
that the northern discrete population segment of the mountain yellow-
legged frog is a DPS, and thus a ``species'' under section 3(16) of the
Act. Our determination of biological and ecological significance is
appropriate because the population segment has a geographical
distribution that is biologically meaningful.
Summary of Changes From the Proposed Rule for the Sierra Nevada Yellow-
Legged Frog and the Northern DPS of the Mountain Yellow-Legged Frog
Based on peer review, Federal and State, and public comments (see
comments in the Summary of Comments and Recommendations section below),
we have clarified information in the sections provided for the Sierra
Nevada yellow-legged frog and the northern DPS of the mountain yellow-
legged frog to better characterize our knowledge of the species'
habitat requirements, correcting some information based on peer review
(vocalizations (Species Description), species ranges (Taxonomy and
Historic and Current Ranges and Distribution sections), current
distribution in Sequoia National Park (Historic and Current Ranges and
Distribution), and clarifying the basis for our determination of
significance for the northern population of the mountain yellow-legged
frog in response to public comments (Distinct Vertebrate Population
Segment)), occasionally adding additional information where needed. In
the Summary of Factors Affecting the Species section, we have re-
ordered threats in Factor A so that the primary activity that has
modified the habitat of the mountain yellow-legged frog complex is
addressed first, while activities with potential only for localized
effects are addressed later. Based on peer review, and Federal, State,
county, and public comments, we have added information where needed and
clarified our findings on the role of current activities, such as
grazing, recreation, packstock use, etc., in species declines. We
reviewed the analysis of dams and diversions that we presented in the
proposed rule and determined that most large reservoir facilities are
below the current range of the mountain yellow-legged frogs. We revised
the dams and water diversions threat magnitude from moderate prevalent
in the proposed rule to minor localized where such structures occur in
this final rule.
In the proposed rule, we stated that grazing presented a minor
prevalent threat. We reworded this final rule to more accurately
reflect the contribution of legacy effects of past grazing levels to
this threat assessment. We found that current livestock grazing that
complies with forest standards and guidelines is not expected to
negatively affect mountain yellow-legged frog populations in most
cases, although limited exceptions could occur (where extant habitat is
limited and legacy effects to meadows still require restoration, where
habitat is limited such as in stream riparian zones or small meadows,
or where grazing standards are exceeded). Rangewide, livestock grazing
is not a substantial threat to the species.
In response to information provided during the public comment
period, we added a discussion of mining activities in the Factor A
discussion. In this final rule, we determine that, while most mining
activities take place below the extant ranges of the species, where
some types of mining activities occur, localized habitat-related
effects may result.
We added new information available on packstock grazing, retaining
our finding that packstock grazing is only likely to be a threat to
mountain yellow-legged frogs in limited situations. We also added more
information on roads and timber harvests, and we clarified that these
activities primarily do not occur where there are extant populations
(except where frogs occur in the northern or lower elevation portions
of the range), and that USFS standards are generally designed to limit
potential effects of such activities. We clarified the threat magnitude
for roads and timber harvest from minor prevalence rangewide to not a
threat to extant populations across much of the species' ranges
(although they may pose important habitat-related effects to the
species in localized areas). We reviewed information provided by the
U.S. Forest Service (USFS), the National Park Service (NPS), CDFW, and
others on recreation activities, and we changed our conclusion on the
recreation threat magnitude from low significance to the species
overall to not considered a threat to populations over much of their
range. However, we recognize that there may be localized effects,
especially outside of backcountry areas where use is high or where
motorized and mechanical use occurs in extant frog habitat.
We added a brief discussion of bullfrogs (Lithobates catesbeiana)
under Factor C for mountain yellow-legged frogs noting that bullfrog
predation and competition is expected to have population-level effects
to mountain yellow-legged frog populations in those low elevation
areas, or in the Lake Tahoe Basin, where the two species may co-occur.
We slightly revised our characterization of the recent population
declines of the mountain yellow-legged frogs due to Batrachochytrium
dendrobatidis (Bd), identifying the fungus as one of the primary
drivers of recent declines, and adding information provided by peer
reviewers and agencies. We also added information to our discussion
under Factor D, including information about the National Park Service
Organic Act, information on the provision in the Wilderness Act about
withdrawing minerals, and information on the status of the Sierra
Nevada yellow-legged frog and the mountain yellow-legged frog under the
California Endangered Species Act (CESA). We also moved discussion of
current CDFW fisheries management to the ``Habitat Modification Due to
Introduction of Trout to Historically Fishless Areas'' section under
Factor A.
We removed the discussion of contaminants under Factor E and refer
readers to the proposed rule. Although we received additional
information that clarified some text and provided additional references
regarding contaminants, the clarifications supported our conclusions in
the proposed rule that the best available information indicates that
contaminants do not pose a current or continuing threat to the species.
We also added additional information either available in our files, or
provided by commenters,
[[Page 24265]]
to clarify and support our finding on the threat of climate change. We
revised the explanation in the determinations for each species to
reflect the above changes.
Summary of Factors Affecting the Species
Section 4 of the Act (16 U.S.C. 1533), and its implementing
regulations at 50 CFR part 424, set forth the procedures for adding
species to the Federal Lists of Endangered and Threatened Wildlife and
Plants. Under section 4(a)(1) of the Act, we may list a species 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; and (E) other natural or manmade
factors affecting its continued existence. Listing actions may be
warranted based on any of the above threat factors, singly or in
combination. Each of these factors is discussed below, and changes from
the proposed rule (78 FR 24472, April 25, 2013) are reflected in these
discussions. The following analysis is applicable to both the Sierra
Nevada yellow-legged frog (Rana sierrae) and the northern distinct
population segment of the mountain yellow-legged frog (Rana muscosa).
Factor A. The Present or Threatened Destruction, Modification, or
Curtailment of Its Habitat or Range
A number of hypotheses, including habitat modification (including
loss of vegetation, loss of wetlands, habitat modification for urban
development, and degradation of upland habitats) have been proposed for
recent global amphibian declines (Bradford et al. 1993, p. 883; Corn
1994, p. 62; Alford and Richards 1999, p. 134). However, physical
habitat modification has not been associated with the rangewide decline
of mountain yellow-legged frogs. Mountain yellow-legged frogs occur
primarily at high elevations in the Sierra Nevada, which have not had
the types or extent of large-scale habitat conversion and physical
disturbance that have occurred at lower elevations (Knapp and Matthews
2000, p. 429). Thus, direct habitat destruction or modification
associated with intensive human activities has not been implicated in
the decline of this species (Davidson et al. 2002, p. 1597).
However, other human activities may have played a role in the
modification of mountain yellow-legged frog habitat. We have identified
the following habitat-related activities as potentially relevant to the
conservation status of the mountain yellow-legged frog complex: Fish
introductions (see also Factor C, below), dams and water diversions,
livestock grazing, timber management, road construction and
maintenance, packstock use, recreational activities, and fire
management activities. Such activities may have degraded habitat in
ways that have reduced its capacity to sustain viable populations and
may have fragmented and isolated mountain yellow-legged frog
populations from each other.
Habitat Modification Due to Introduction of Trout to Historically
Fishless Areas
One habitat feature that is documented to have a significant
detrimental impact to mountain yellow-legged frog populations is the
presence of introduced trout resulting from stocking programs for the
creation and maintenance of a recreational fishery. To further angling
success and opportunity, trout stocking programs in the Sierra Nevada
started in the late 19th century (Bahls 1992, p. 185; Pister 2001, p.
280). This anthropogenic activity has community-level effects and is
one of the primary threats to mountain yellow-legged frog habitat and
species viability.
Prior to extensive trout planting programs, almost all streams and
lakes in the Sierra Nevada at elevations above 1,800 m (6,000 ft) were
fishless. Several native fish species occur naturally in aquatic
habitats below this elevation in the Sierra Nevada (Knapp 1996, pp. 12-
14; Moyle et al. 1996, p. 354; Moyle 2002, p. 25), but natural barriers
prevented fish from colonizing the higher-elevation waters of the
Sierra Nevada watershed (Moyle et al. 1996, p. 354). The upper reaches
of the Kern River, where native fish such as the Little Kern golden
trout (Oncorhynchus mykiss whitei) and California golden trout (O. m.
aguabonita) evolved, represent the only major exception to the 1,800-m
(6,000-ft) elevation limit for fishes within the range of the mountain
yellow-legged frog in the Sierra Nevada (Moyle 2002, p. 25).
Additionally, prior to extensive planting, native Paiute cutthroat (O.
clarki seleneris) and Lahontan cutthroat (O. c. henshawi) were limited
in their distribution to several rivers, streams, and limited large
lakes in the eastern Sierra Nevada (Knapp 1996, p. 369; Moyle 1996 et
al., pp. 954-958), indicating some overlap with the range of the Sierra
Nevada yellow-legged frog.
Some of the first practitioners of trout stocking in the Sierra
Nevada were the Sierra Club, local sportsmen's clubs, private citizens,
and the U.S. military (Knapp 1996, p. 8; Pister 2001, p. 280). As more
hatcheries were built and the management of the trout fishery became
better organized, fish planting continued for the purpose of increased
angler opportunities and success (Pister 2001, p. 281). After World War
II, the method of transporting trout to high-elevation areas changed
from packstock to aircraft, which allowed stocking in more remote lakes
and in greater numbers. With the advent of aerial stocking, trout
planting expanded to new areas, with higher efficiency.
Brook trout (Salvelinus fontinalis), brown trout (Salmo trutta),
rainbow trout (Oncorhynchus mykiss), and other trout species
assemblages have been planted in most streams and lakes of the Sierra
Nevada (Knapp 1996, p. 8; Moyle 2002, p. 25). Since the advent of
aerial stocking, backcountry areas not accessible by truck are stocked
by air (Pert 2002, pers. comm.), which limits stocking to lakes.
National Forests in the Sierra Nevada have a higher proportion of lakes
with fish occupancy than do National Parks (Knapp 1996, p. 3),
primarily because the National Park Service (NPS) began phasing out
fish stocking within their jurisdictional boundaries in 1969, with
limited stocking occurring until it was terminated altogether in Sierra
Nevada National Parks in 1991 (Knapp 1996, p. 9). California Department
of Fish and Wildlife (CDFW) continues to stock trout in National Forest
water bodies, but in 2001 reduced the number of stocked water bodies to
reduce impacts to native amphibians (ICF Jones & Stokes 2010, pp. ES-1-
ES-16). Current stocking decisions are based on criteria outlined in
the Environmental Impact Report for the Hatchery and Stocking Program
(ICF Jones & Stokes 2010, Appendix K).
Fish stocking as a practice has been widespread throughout the
range of both species of mountain yellow-legged frogs. Knapp and
Matthews (2000, p. 428) indicated that 65 percent of the water bodies
that were 1 ha (2.5 ac) or larger in National Forests they studied were
stocked with fish on a regular basis. Over 90 percent of the total
water body surface area in the John Muir Wilderness was occupied by
nonnative trout (Knapp and Matthews 2000, p. 434).
Another detrimental feature of fish stocking is that, in the Sierra
Nevada, fish often persist in water bodies even after stocking ceases.
Thirty-five to 50 percent of lakes larger than 1 ha (2.5 ac) within
Sierra Nevada National Parks are occupied by nonnative fish, which is
[[Page 24266]]
only a 29 to 44 percent decrease in fish occupancy since fish stocking
was terminated around 2 decades before the estimate was made (Knapp
1996, p. 1). Though data on fish occupancy in streams are lacking
throughout the Sierra Nevada, Knapp (1996, pp. 9-11) estimated that 60
percent of the streams in Yosemite National Park were still occupied by
introduced trout because trout readily move out of lakes to colonize
both inlet and outlet streams. The presence of trout in these once
fishless waters has modified the habitat at a landscape scale.
Thus, the frog's habitat has been modified due to the introduction
of a nonnative predator that both competes for limited food resources
and directly preys on mountain yellow-legged frog tadpoles and adults
(see Factor C below). Presence of nonnative trout in naturally fishless
ecosystems has had profound effects on the structure and composition of
faunal assemblages, severely reducing not only amphibians, but also
zooplankton and large invertebrate species (see Knapp 1996, p. 6;
Bradford et al. 1998, p. 2489; Finlay and Vredenburg 2007, pp. 2194-
2197). Within the frog's historical range, past trout introductions and
the continuing presence of fish in most lakes resulted in the
elimination of frogs from most waters that were suitable for fish.
Across the range of these species in the Sierra Nevada, the presence of
fish in most of the deeper lakes has altered the aquatic habitat that
mountain yellow-legged frogs rely on for overwintering and breeding,
and has also reduced connectivity among frog populations. Fish now
populate the deeper lakes and connecting streams and largely separate
and increase the distance between the current sites inhabited by the
highly-aquatic frogs (the connectivity of occupied sites in present
versus former fishless conditions differs by approximately 10-fold)
(Bradford et al. 1993, pp. 884-887; Knapp 1996, pp. 373-379). Where
reservoirs harbor introduced fish, successful reproduction of mountain
yellow-legged frogs may be reduced if there are no shallow side
channels or separate pools (Jennings 1996, p. 939). Most reservoirs do
not overlap significantly with the current extant range of the species
(CDFW 2013, p. 3) (see Dams and Water Diversions below); however, a
number of reservoirs were constructed in the mid-1900s at mid-
elevations within lower edges of the species' historic range (for
example, Sierra Nevada yellow-legged frogs were taken from Bear River
Reservoir (Eldorado National Forest), Union Reservoir (Stanislaus
National Forest), and several others). With the exception of one 1999
record from Faggs Reservoir on the Plumas National Forest, all of
several dozen records of the species from reservoirs are pre-1975, and
at least half pre-date the water development projects at those
locations (Brown et al. 2009, p. 78). All of these reservoirs now
harbor introduced fish species, and at least two also harbor bullfrogs,
suggesting that subsequent introductions may have played a role in past
declines in those areas (see Brown et al. 2009, p. 78).
The body of scientific research has demonstrated that introduced
trout have negatively impacted mountain yellow-legged frogs over much
of the Sierra Nevada (Grinnell and Storer 1924, p. 664; Bradford 1989,
pp. 775-778; Bradford et al. 1993, pp. 882-888; Knapp 1994, p. 3; Drost
and Fellers 1996, p. 422; Knapp 1996, pp. 13-15; Bradford et al. 1998,
pp. 2482, 2489; Knapp and Matthews 2000, p. 428; Knapp et al. 2001, p.
401). Fish stocking programs have negative ecological implications
because fish eat aquatic fauna, including amphibians and invertebrates
(Bahls 1992, p. 191; Erman 1996, p. 992; Jennings 1996, p. 939; Knapp
1996, pp. 373-379; Matthews et al. 2001, pp. 1135-1136; Pilliod and
Peterson 2001, p. 329; Schindler et al. 2001, p. 309; Moyle 2002, p.
58; Epanchin et al. 2010, p. 2406). Finlay and Vredenburg (2007, p.
2187) documented that the same benthic (bottom-dwelling) invertebrate
resource base sustains the growth of both frogs and trout, suggesting
that competition with trout for prey is an important factor that may
contribute to the decline of the mountain yellow-legged frog.
Introductions of salmonids to fishless lakes have also been associated
with alteration of nutrient cycles and primary productivity in mountain
lakes, including those in the Sierra Nevada (Schindler et al. 2001, pp.
308, 313-319).
Knapp and Matthews (2000, p. 428) surveyed more than 1,700 water
bodies, and concluded that a strong negative correlation exists between
introduced trout and mountain yellow-legged frogs (Knapp and Matthews
2000, p. 435). Consistent with this finding are the results of an
analysis of the distribution of mountain yellow-legged frog tadpoles,
which indicate that the presence and abundance of this life stage are
reduced dramatically in fish-stocked lakes (Knapp et al. 2001, p. 408).
Knapp (2005a, pp. 265-279) also compared the distribution of nonnative
trout with the distributions of several amphibian and reptile species
in 2,239 lakes and ponds in Yosemite National Park, and found that
mountain yellow-legged frogs were five times less likely to be detected
in waters where trout were present. Even though stocking within the
National Park ceased in 1991, more than 50 percent of water bodies
deeper than 4 m (13 ft) and 75 percent deeper than 16 m (52 ft) still
contained trout populations in 2000-2002 (Knapp 2005a, p. 270). Both
trout and mountain yellow-legged frogs utilize deeper water bodies.
Based on the results from Knapp (2005a), the reduced detection of frogs
in trout-occupied waters indicates that trout are excluding mountain
yellow-legged frogs from some of the best aquatic habitat.
Several aspects of the mountain yellow-legged frog's life history
are thought to exacerbate its vulnerability to extirpation by trout
(Bradford 1989, pp. 777-778; Bradford et al. 1993, pp. 886-888; Knapp
1996, p. 14; Knapp and Matthews 2000, p. 435). Mountain yellow-legged
frogs are highly aquatic and are found primarily in lakes, most of
which now contain trout (Knapp 1996, p. 14). In comparison to other
Sierran frogs, mountain yellow-legged frog tadpoles generally need at
least 2 years to reach metamorphosis, which restricts breeding to
waters that are deep enough to avoid depletion of oxygen when ice-
covered (Knapp 1996, p.14). Overwintering adults must also avoid oxygen
depletion when the water is covered by ice, generally limiting
overwintering to deeper waters that do not become anoxic (Mullally and
Cunningham 1956a, p. 194; Bradford 1983, p. 1179; Knapp and Matthews
2000, pp. 435-436). At high elevations, both tadpoles and adults
overwinter under ice for up to 9 months (Bradford 1983, p. 1171). These
habitat requirements appear to restrict successful breeding and
overwintering to the deeper water bodies where the chances of summer
drying and winter freezing are reduced, the same water bodies that are
most suitable for fishes; fishes also need deeper water bodies where
the chances of summer drying and winter freezing are reduced (Bradford
1983, pp. 1172-1179; Knapp 1996, p. 14; Knapp and Matthews 2000, pp.
429, 435-436). Past fish-stocking practices targeted the deeper lakes,
so the percentage of water bodies containing fish has increased with
water depth, resulting in elimination of mountain yellow-legged frogs
from once suitable habitats in which they were historically most
common, and thereby generally isolating populations to the shallower,
marginal habitats that do not have fish (Bradford 1983, pp. 1172-1179;
Bradford et al. 1993, pp. 884, 886-
[[Page 24267]]
887; Knapp and Matthews 2000, pp. 435-436).
Mountain yellow-legged frogs and trout (native and nonnative) do
co-occur at some sites, but these co-occurrences are generally thought
to represent mountain yellow-legged frog ``sink'' populations (areas
with negative population growth rates in the absence of immigration)
(Bradford et al. 1998, p. 2489; Knapp and Matthews 2000, p. 436).
Mountain yellow-legged frogs have also been extirpated at some fishless
bodies of water (Bradford 1991, p. 176; Drost and Fellers 1996, p.
422). A possible explanation is the isolation and fragmentation of
remaining populations due to introduced fishes in the streams that once
provided mountain yellow-legged frogs with dispersal and recolonization
routes; these remote populations are now non-functional as
metapopulations (Bradford 1991, p. 176; Bradford et al. 1993, p. 887).
Based on a survey of 95 basins within Sequoia and Kings Canyon National
Parks, Bradford et al. (1993, pp. 885-886) estimated that the
introduction of fishes into the study area resulted in an approximately
10-fold increase in habitat fragmentation between populations of
mountain yellow-legged frogs. Knapp and Matthews (2000, p. 436) believe
that this fragmentation has further isolated mountain yellow-legged
frogs within the already marginal habitat left unused by fishes.
Fragmentation of mountain yellow-legged frog habitat renders
populations more vulnerable to extirpation from random events (such as
disease) (Wilcox 1980, pp. 114-115; Bradford et al. 1993, p. 887;
Hanski and Simberloff 1997, p. 21; Knapp and Matthews 2000, p. 436).
Isolated population locations may have higher extinction rates because
trout prevent successful recolonization and dispersal to and from these
sites (Bradford et al. 1993, p. 887; Blaustein et al. 1994a, p. 7;
Knapp and Matthews 2000, p. 436). If the distance between sites is too
great, amphibians may not readily recolonize unoccupied sites following
local extinctions because of physiological constraints, the tendency to
move only short distances, and high site fidelity. Finally, frogs that
do attempt recolonization may emigrate into fish-occupied habitat and
perish, rendering sites with such metapopulation dynamics less able to
sustain frog populations.
In 2001, CDFW revised fish stocking practices and implemented an
informal policy on fish stocking in the range of the Sierra Nevada
yellow-legged frog and northern DPS of the mountain yellow-legged frog.
This policy directs that: (1) Fish will not be stocked in lakes with
known populations of mountain yellow-legged frogs, nor in lakes that
have not yet been surveyed for mountain yellow-legged frog presence;
(2) waters will be stocked only with a fisheries management
justification; and (3) the number of stocked lakes will be reduced over
time. In 2001, the number of lakes stocked with fish within the range
of the mountain yellow-legged frog in the Sierra Nevada was reduced by
75 percent (Milliron 2002, pp. 6-7; Pert et al. 2002, pers. comm.).
Current CDFW guidelines stipulate that water bodies within the same
basin and 2 km (1.25 mi) from a known mountain yellow-legged frog
population will not be stocked with fish unless stocking is justified
through a management plan that considers all the aquatic resources in
the basin, or unless there is heavy angler use and no opportunity to
improve the mountain yellow-legged frog habitat (Milliron 2002a, p. 5).
The Hatchery and Stocking Program Environmental Impact Report/
Environmental Impact Statement, finalized in 2010 (ICF Jones & Stokes
2010, Appendix K), outlines a decision approach to mitigate fish
stocking effects on Sierra amphibians that prohibits fish stocking in
lakes with confirmed presence of a limited number of designated
species, including the mountain yellow-legged frogs (see ICF Jones &
Stokes 2010, Appendix E) using recognized survey protocols. Large
reservoirs generally continue to be stocked to provide a put-and-take
fishery for recreational angling.
As part of the High Mountain Lakes Project, CDFW is in the process
of developing management plans for basins within the range of the
Sierra Nevada yellow-legged frog and the northern DPS of mountain
yellow-legged frog (CDFG (CDFW) 2001, p. 1; Lockhart 2011, pers.
comm.). CDFW states that objectives of the basin plans specific to the
mountain yellow-legged frog include management in a manner that
maintains or restores native biodiversity and habitat quality, supports
viable populations of native species, and provides for recreational
opportunities that consider historical use patterns (CDFG (CDFW) 2001,
p. 3). They state that, under this approach, lakes that support
mountain yellow-legged populations in breeding, foraging, or dispersal,
and/or present opportunities to restore or expand habitat, are managed
for the conservation of the species. Lakes that do not support mountain
yellow-legged frogs are not viable restoration opportunities, and lakes
that support trout populations are managed primarily for recreational
angling (CDFG (CDFW) 2001, p. 3). They further note that lakes managed
for recreational angling may be stocked if CDFW determines that
stocking the lake will achieve a desirable fisheries management
objective and is not otherwise precluded by stocking decision
guidelines and agreements (for stocking decision documents, see CDFW
2013, pp. 1, 2).
Since the mid-1990s, various parties, including researchers, CDFW,
NPS, and the USFS, have implemented a variety of projects to actively
restore habitat for the mountain yellow-legged frog via the removal of
nonnative trout (USFS 2011, pp. 128-130; NPS 2013, pp. 3-5).
Although fish stocking has been curtailed within many occupied
basins, the impacts to frog populations persist due to the presence of
self-sustaining fish populations in some of the best habitat that
normally would have sustained mountain yellow-legged frogs. The
fragmentation that persists across the range of these frog species
renders them more vulnerable to other population stressors, and
recovery is slow, if not impossible, without costly and physically
difficult direct human intervention (such as physical and chemical
trout removal) (see Knapp et al. 2007a, pp. 11-19). While most of the
impacts occurred historically, the impact upon the biogeographic
(population/metapopulation) integrity of the species will be long-
lasting. Currently, habitat degradation and fragmentation by fish is
considered a highly significant and prevalent threat to persistence and
recovery of the species.
Dams and Water Diversions
While a majority of dams and water diversions within the Sierra
Nevada are located at lower elevations (USFS 2011, p. 83), some large
reservoirs have been constructed within the historic range of the
mountain yellow-legged frog complex. These large reservoirs include,
but are not limited to Huntington Lake, Florence Lake, Lake Thomas A.
Edison, Saddlebag Lake, Cherry Lake, Hetch Hetchy, Upper and Lower Blue
Lakes, Lake Aloha, Silver Lake, Hell Hole Reservoir, French Meadow
Reservoir, Lake Spaulding, Alpine Lake, Loon Lake, and Ice House
Reservoir. A number of these occur at elevations below the current
range of the species, indicating that the network of large water and
power projects found at lower elevations does not overlap significantly
with the current accepted distribution of the mountain yellow-legged
frogs in the Sierra Nevada (CDFW 2013, p. 3).
Kondolf et al. (1996, p. 1014) report that dams can have direct
effects to
[[Page 24268]]
riparian habitat through permanent removal of habitat to construct
roads, penstocks, powerhouses, canals, and dams. Impacts of reservoirs
include flooding of riparian vegetation and impediments to
establishment of new shoreline vegetation by fluctuating water levels.
Dams can alter the temperature and sediment load of the rivers they
impound (Cole and Landres 1996, p. 175). Dams, water diversions, and
their associated structures can also alter the natural flow regime with
unseasonal and fluctuating releases of water (Kondolf et al. 1996, p.
1014). We expect most such effects to occur in stream systems below the
extant range of the mountain yellow-legged frogs, although it is
possible that stream localities at the northern extent of the range or
at low elevations may be affected (see also CDFW 2013, pp. 2-4).
The extent of past impacts to mountain yellow-legged frog
populations from habitat loss or modification due to reservoir projects
has not been quantified. CDFW (2013, p. 3) has noted that there are
locations where the habitat inundated as the result of dam construction
(for example, Lake Aloha in the Desolation Wilderness) may have been of
higher quality for mountain yellow-legged frogs than the created
impoundment. Reservoirs can provide habitat for introduced predators,
including fish, bullfrogs, and crayfish, and in some cases, the past
construction of reservoirs has facilitated the spread of nonnative fish
(CDFW 2013, pp. 3, 4). In such cases, reservoirs may function as
barriers to movement of mountain yellow-legged frogs. However, CDFW
reported observing mountain yellow-legged frogs dispersing through
fishless reservoirs (CDFW 2013, p. 4). (For a complete discussion of
the impacts of fish stocking see Habitat Modification Due to
Introduction of Trout to Historically Fishless Areas above and the
discussion under Factor C.).
Most of the dams constructed within the historic range of the
mountain yellow-legged frogs are small streamflow-maintenance dams
(CDFW 2013, p. 13) at the outflows of high-elevation lakes. These small
dams may create additional habitat for the species and can act as
barriers to fish migration from downstream tributaries into fishless
habitats, although they do not impede frog movement (CDFW 2013, p. 3).
CDFW staff (2013, p. 13) have observed that extant frog populations may
have persisted where such dams have helped to preserve a fishless
environment behind the dam.
Based on comments from CDFW and others and the provision of
additional information, we have reviewed the analysis of dams and
diversions that we presented in the proposed rule. We find that most
large facilities are below the current range of the mountain yellow-
legged frogs and have revised our finding. In the proposed rule, we
stated that dams and diversions presented a moderate, prevalent threat
to persistence and recovery of the species. In this final rule, we find
that dams and water diversions present a minor, localized threat to
persistence and recovery of the species where structures occur.
Livestock Use (Grazing)
The combined effect of legacy conditions from historically
excessive grazing use and current livestock grazing activities has the
potential to impact habitat in the range of the mountain yellow-legged
frog. The following subsections discuss the effects of excessive
historical grazing, current extent of grazing, and current grazing
management practices. As discussed below, grazing has the potential to
reduce the suitability of habitat for mountain yellow-legged frogs by
reducing its capability to sustain frogs and facilitate dispersal and
migration, especially in stream areas.
Grazing of livestock in riparian areas impacts the function of the
aquatic system in multiple ways, including soil compaction, which
increases runoff and decreases water availability to plants; vegetation
removal, which promotes increased soil temperatures and evaporation
rates at the soil surface; and direct physical damage to the vegetation
(Kauffman and Krueger 1984, pp. 433-434; Cole and Landres 1996, pp.
171-172; Knapp and Matthews 1996, pp. 816-817). Streamside vegetation
protects and stabilizes streambanks by binding soils to resist erosion
and trap sediment (Kauffman et al. 1983, p. 683; Chaney et al. 1990, p.
2). Grazing within mountain yellow-legged frog habitat has been
observed to remove vegetative cover, potentially exposing frogs to
predation and increased desiccation (Knapp 1993b, p. 1; Jennings 1996,
p. 539), and to lead to erosion which may silt in ponds and thereby
reduce the water depth needed for overwinter survival (Knapp 1993b, p.
1). However, an appropriately managed grazing regime (including timing
and intensity) can enhance primary riparian vegetation attributes that
are strongly correlated to stream channel and riparian soil stability
conditions necessary to maintain a functioning riparian system (George
et al. 2011, p. 227). Although, where highly degraded conditions such
as downcut channels exist, grazing management alone may not be
sufficient to restore former riparian conditions (George et al. 2011,
p. 227).
Aquatic habitat can also be degraded by grazing. Mass erosion from
trampling and hoof slide causes streambank collapse and an accelerated
rate of soil transport to streams (Meehan and Platts 1978, p. 274).
Accelerated rates of erosion lead to elevated instream sediment loads
and depositions, and changes in stream-channel morphology (Meehan and
Platts 1978, pp. 275-276; Kauffman and Krueger 1984, p. 432). Livestock
grazing may lead to diminished perennial streamflows (Armour et al.
1994, p. 10). Livestock can increase nutrient-loading in water bodies
due to urination and defecation in or near the water, and can cause
elevated bacteria levels in areas where cattle are concentrated (Meehan
and Platts 1978, p. 276; Stephenson and Street 1978, p. 156; Kauffman
and Krueger 1984, p. 432). With increased grazing intensity, these
adverse effects to the aquatic ecosystem increase proportionately
(Meehan and Platts 1978, p. 275; Clary and Kinney 2000, p. 294).
Observational data indicate that livestock can negatively impact
mountain yellow-legged frogs by altering riparian habitat (Knapp 1993a,
p. 1; 1993b, p. 1; 1994, p. 3; Jennings 1996, p. 938; Carlson 2002,
pers. comm.; Knapp 2002a, p. 29). Livestock tend to concentrate along
streams and wet areas where there is water and herbaceous vegetation;
grazing impacts are, therefore, most pronounced in these habitats
(Meehan and Platts 1978, p. 274; U.S. Government Accounting Office
(GAO) 1988, pp. 10-11; Fleischner 1994, p. 635; Menke et al. 1996, p.
17). This concentration of livestock contributes to the destabilization
of streambanks, causing undercuts and bank failures (Kauffman et al.
1983, p. 684; Marlow and Pogacnik 1985, pp. 282-283; Knapp and Matthews
1996, p. 816; Moyle 2002, p. 55). Grazing activity can contribute to
the downcutting of streambeds and lower the water table. The degree of
erosion caused by livestock grazing can vary with slope gradient,
aspect, soil condition, vegetation density, and accessibility to
livestock, with soil disturbance greater in areas overused by livestock
(Meehan and Platts 1978, pp. 275-276; Kauffman et al. 1983, p. 685;
Kauffman and Krueger 1984, p. 432; Bohn and Buckhouse 1985, p. 378; GAO
1988, p. 11; Armour et al. 1994, pp. 9-11; Moyle 2002, p. 55).
Livestock grazing may impact other wetland systems, including ponds
that can serve as mountain yellow-legged
[[Page 24269]]
frog habitat. Grazing can modify shoreline habitats by removing
overhanging banks that provide shelter, and grazing contributes to the
siltation of breeding ponds. Bradford (1983, p. 1179) and Pope (1999,
pp. 43-44) have documented the importance of deep lakes to overwinter
survival of these species. We expect that pond siltation due to grazing
may reduce the depth of breeding ponds and cover underwater crevices in
some circumstances where grazing is heavy and where soils are highly
erodable, thereby making the ponds less suitable, or unsuitable, as
overwintering habitat for tadpoles and adult mountain yellow-legged
frogs.
Effects of Excessive Historical Grazing
In general, historical livestock grazing within the range of the
mountain yellow-legged frog was at a high (although undocumented),
unregulated and unsustainable level until the establishment of National
Parks (beginning in 1890) and National Forests (beginning in 1905) (UC
1996a, p. 114; Menke et al. 1996, p. 14). Historical evidence indicates
that heavy livestock use in the Sierra Nevada has resulted in
widespread damage to rangelands and riparian systems due to sod
destruction in meadows, vegetation destruction, and gully erosion (see
review in Brown et al. 2009, pp. 56-58). Within the newly established
National Parks, grazing by cattle and sheep was eliminated, although
grazing by packstock, such as horses and mules, continued. Within the
National Forests, the amount of livestock grazing was gradually
reduced, and the types of animals shifted away from sheep and toward
cattle and packstock, with cattle becoming the dominant livestock.
During World Wars I and II, increased livestock use occurred on
National Forests in the west, causing overuse in the periods 1914-1920
and 1939-1946. Between 1950 and 1970 livestock numbers were permanently
reduced due to allotment closures and uneconomical operations, with
increased emphasis on resource protection and riparian enhancement.
Further reductions in livestock use began again in the 1990s, due in
part to USFS reductions in permitted livestock numbers, seasons of use,
implementation of rest-rotation grazing systems, and to responses to
drought (Menke et al. 1996, pp. 7, 8). Between 1981 and 1998, livestock
numbers on National Forests in the Sierra Nevada decreased from 163,000
to approximately 97,000 head, concurrent with Forest Service
implementation of standards and guidelines for grazing and other
resource management (USFS 2001, pp. 399-416).
Effects of Current Grazing
Yosemite, Sequoia, and Kings Canyon National Parks remain closed to
livestock grazing. On USFS-administered lands that overlap the
historical ranges of the mountain yellow-legged frog in the Sierra
Nevada, there are currently 161 active Rangeland Management Unit
Allotments for livestock grazing. However, based on frog surveys
performed since 2005, only 27 of these allotments have extant mountain
yellow-legged frog populations, while some allotments that were located
in sensitive areas have been closed (USFS 2008, unpubl. data; CDFW
(CDFG) unpubl. data). As of 2009, USFS data indicated that grazing
occurs on about 65 percent of National Forest lands within the range of
the mountain yellow-legged frog; that livestock numbers remain greatly
reduced from historical levels; and that numerous watershed restoration
projects have been implemented, although grazing may still impact many
meadows above mid-elevation and restoration efforts are far from
complete (Brown et al. 2009, pp. 56, 57). However, Brown et al. (2009,
p. 56) report that livestock grazing is more likely to occur in certain
habitat types used by mountain yellow-legged frogs than others,
indicating that populations found in meadows, stream riparian zones,
and lakes in meadows are more likely to encounter habitat effects of
grazing than populations found in the deeper alpine lakes that the
species more likely inhabit (Brown et al. 2009, p. 56).
USFS standards and guidelines in forest land and resource
management plans have been implemented to protect water quality,
sensitive species, vegetation, and stream morphology. Further, USFS
standards have been implemented in remaining allotments to protect
aquatic habitats (see discussion of the aquatic management strategy
under Factor D for examples). USFS data from long-term meadow
monitoring collected from 1999 to 2006 indicate that most meadows
appear to be in an intermediate quality condition class, with seeming
limited change in condition class over the first 6 years of monitoring.
In addition, USFS grazing standards and guidelines are based on current
science and are designed to improve or maintain range ecological
conditions, and standards for managing habitat for threatened,
endangered, and sensitive species have also been incorporated (Brown et
al. 2009, pp. 56-58). The seasonal turn-out dates (dates at which
livestock are permitted to move onto USFS allotments) are set yearly
based on factors such as elevation, annual precipitation, soil
moisture, and forage plant phenology, and meadow readiness dates are
also set for montane meadows. However, animals turned out to graze on
low-elevation range (until higher elevation meadows are ready) may
reach upper portions of allotments before the meadows have reached
range readiness (Brown et al. 2009, p. 58).
Menke et al. (1996) have reported that grazing livestock in numbers
that are consistent with grazing capacity and use of sustainable
methods led to better range management in the Sierra Nevada over the 20
years prior to development of the report. They also noted that moderate
livestock grazing has the potential to increase native species
diversity in wet and mesic meadows by allowing native plant cover to
increase on site. Brown et al. (2009, p. 58) expect proper livestock
management, such as proper timing, intensity, and duration, to result
in a trend towards increased riparian species and a trend towards
restored wet and mesic meadows on National Forests. To date, the
scientific and commercial information available to us does not include
descriptive or cause-effect research that establishes a causal link
between habitat effects of livestock grazing and mountain yellow-legged
frog populations; however, anecdotal information of specific habitat
effects suggests that, in specific locations, the current grazing
levels may have population-level effects (see Knapp 1993b, p. 1; Brown
et al. 2009, p. 56). In addition, where low-elevation populations occur
in meadows, additional conservation measures may be required for
recovery (USFS 2013, p. 5).
In summary, the legacy effects to habitat from historical grazing
levels, such as increased erosion, stream downcutting and headcutting,
lowered water tables, and increased siltation, are a threat to mountain
yellow-legged frogs in those areas where such conditions still occur
and may need active restoration. In the proposed rule, we stated that
grazing presented a minor prevalent threat. Based on USFS and public
comments, we have reevaluated our analysis of grazing to clarify
effects of past versus current grazing. We have reworded the finding to
more accurately reflect the contribution of legacy effects of past
grazing levels to this threat assessment, as follows: Current livestock
grazing activities may present an ongoing, localized threat to
individual populations in locations where the populations occur in
stream
[[Page 24270]]
riparian zones and in small waters within meadow systems, where active
grazing co-occurs with extant frog populations. Livestock grazing that
complies with forest standards and guidelines is not expected to
negatively affect mountain yellow-legged frog populations in most
cases, although limited exceptions could occur, especially where extant
habitat is limited. In addition, mountain yellow-legged frogs may be
negatively affected where grazing standards are exceeded. Rangewide,
current livestock grazing is not a substantial threat to the species.
Mining
Several types of mining activities have occurred, or may currently
occur, on National Forests, including aggregate mining (the extraction
of materials from streams or stream terraces for use in construction),
hardrock mining (the extraction of minerals by drilling or digging into
solid rock), hydraulic mining (a historical practice using pressurized
water to erode hillsides, outlawed in 1884), placer mining (mining in
sand or gravel, or on the surface, without resorting to mechanically
assisted means or explosives), and suction-dredge mining (the
extraction of gold from riverine materials, in which water, sediment,
and rocks are vacuumed from portions of streams and rivers, sorted to
obtain gold, and the spoils redeposited in the stream (see review in
Brown et al. 2009, pp. 62-64).
Aggregate mining can alter sediment transport in streams, altering
and incising stream channels, and can cause downstream deposition of
sediment, altering or eliminating habitat. Aggregate mining typically
occurs in large riverine channels that are downstream of much of the
range of the mountain yellow-legged frog complex (see review in Brown
et al. 2009, pp. 62-64). However, Brown et al. (2009, pp. 62-64) note
that effects of aggregate mining may occur in some portions of the
Feather River system where such operations occur within the historic
range of the Sierra Nevada yellow-legged frog, and potentially in
localized areas within the range of both species, where the USFS
maintains small quarries for road work. They note that, although
effects of aggregate mining on mountain yellow-legged frogs are
unstudied, impacts are probably slight.
Hardrock mining can be a source of pollution where potentially
toxic metals are solubilized by waters that are slightly acidic. Past
mining activities have resulted in the existence of many shaft or
tunnel mines on the forest in the Sierra Nevada, although most are
thought to occur below the range of the species. Most operations that
are thought to have the potential to impact the mountain yellow-legged
frogs occur in the lower elevation portions of the Sierra Nevada
yellow-legged frog range on the Plumas National Forest and in the
ranges of both species on the Inyo National Forest (see review in Brown
et al. 2009, pp. 62-64).
Hydraulic mining has exposed previously concealed rocks that can
increase pollutants such as acid, cadmium, mercury, and asbestos, and
its effect on water pollution may still be apparent on the Feather
River. However, most of the area that was mined in this way is below
the elevation where Sierra Nevada yellow-legged frogs are present, so
effects are likely highly localized (see review in Brown et al. 2009,
pp. 63, 64). Although placer mining was dominant historically, today
it's almost exclusively recreational and is not expected to have
habitat-related effects.
Brown et al. (2009, p. 64) report that suction-dredge mining is
also primarily recreational noting that, because nozzles are currently
restricted to 6 inches or smaller, CDFW (CDFG, 1994) expects disturbed
areas to recover quickly (although CDFW notes that such dredging may
increase suspended sediments, change stream geomorphology, and bury or
suffocate larvae). Suction dredge mining occurs primarily in the
foothills of the Sierra Nevada, thus presenting a risk primarily to
mountain yellow-legged frog populations at the lower elevations of the
species' range. Suction dredging is highly regulated by the CDFW, and
in the past, many streams have been seasonally or permanently closed
(see review in Brown et al. 2009, p. 64). Currently CDFW has imposed a
moratorium on suction dredging.
The high-elevation areas where most Sierra Nevada yellow-legged
frogs and mountain yellow-legged frogs occur are within designated
wilderness, where mechanical uses are prohibited by the Wilderness Act.
Designated wilderness was withdrawn for new mining claims on January 1,
1984, although a limited number of active mines that predated the
withdrawal still occur within wilderness (see Wilderness Act under
Factor D, below). Therefore, we expect that mining activities may pose
local habitat-related impacts to the species at specific localities
where mining occurs.
Packstock Use
Similar to cattle, horses and mules may significantly overgraze,
trample, or pollute riparian and aquatic habitat if too many are
concentrated in riparian areas too often or for too long. Commercial
packstock trips are permitted in National Forests and National Parks
within the Sierra Nevada, often providing transport services into
wilderness areas through the use of horses or mules. Use of packstock
in the Sierra Nevada increased after World War II as road access,
leisure time, and disposable income increased (Menke et al. 1996, p.
919). Packstock grazing is the only grazing currently permitted in the
National Parks of the Sierra Nevada. Since the mid-1970s, National
Forests and National Parks have generally implemented regulations to
manage visitor use and group sizes, including measures to reduce
packstock impacts to vegetation and soils in order to protect
wilderness resources. For example, Sequoia and Kings Canyon National
Parks have the backcountry area with the longest history of research
and management of packstock impacts (Hendee et al. 1990, p. 461).
Hendee et al. (1990, p. 461) report that the extensive and long-term
monitoring for Sequoia, Kings Canyon, and Yosemite National Parks makes
it possible to quantify impacts of packstock use, showing that the vast
majority of Sierra Nevada yellow-legged frog and mountain yellow-legged
frog populations in the Parks show no to negligible impacts from
packstock use (National Park Service 2013, p. 3). In the Sixty-Lakes
Basin of Kings Canyon National Park, packstock use is regulated in wet
meadows to protect mountain yellow-legged frog breeding habitat in bogs
and along lake shores from trampling and associated degradation
(Vredenburg 2002, p. 11; Werner 2002, p. 2; National Park Service 2013,
p. 3). Packstock use is also regulated in designated wilderness in
National Forests within the Sierra Nevada.
Packstock use is likely a threat of low significance to mountain
yellow-legged frogs at the current time, except on a limited, site-
specific basis. As California's human population increases, the impact
of recreational activities, including packstock use and riding on the
National Forests in the Sierra Nevada, are projected to increase (USDA
2001a, pp. 473-474). However, on the Inyo National Forest, current
commercial packstock use is approximately 27 percent of the level of
use in the 1980s reflecting a decline in the public's need and demand
for packstock trips. From 2001 to 2005, commercial packstock outfitters
within the Golden Trout and South Sierra Wilderness Areas averaged 28
percent of their current authorized use (USFS
[[Page 24271]]
2006, p. 3-18). Similarly, long-term permitting data for
administrative, commercial, and recreational packstock use in the three
National Parks indicates that packstock use is declining in the Parks,
providing no evidence to suggest that packstock use will increase in
the future in the Parks (National Park Service 2013, pp. 3, 4). Habitat
changes due to packstock grazing may pose a risk to some remnant
populations of frogs and, in certain circumstances, a hindrance to
recovery of populations in heavily used areas.
Roads and Timber Harvest
Activities that alter the terrestrial environment (such as road
construction and timber harvest) may impact amphibian populations in
the Sierra Nevada (Jennings 1996, p. 938) at locations where these
activities occur. Historically, road construction and timber harvest
may have acted to reduce the species' range prior to the more recent
detailed studies and systematic monitoring that have quantified and
documented species losses. Prior to the formation of National Parks in
1890 and National Forests in 1905, timber harvest was widespread and
unregulated, but primarily took place at elevations on the western
slope of the Sierra Nevada below the range of the mountain yellow-
legged frog (University of California (UC) 1996b, pp. 24-25). Between
1900 and 1950, the majority of timber harvest occurred in old-growth
forests on private land (UC 1996b, p. 25). Between 1950 and the early
1990s, timber harvest on National Forests increased, and the majority
of timber harvest-associated impacts on mountain yellow-legged frogs
may therefore have taken place during this period in lower elevation
locations where timber harvest and species occurrences overlapped.
Currently, these activities are expected to occur outside National
Parks or National Forest wilderness areas, with limited exceptions.
Timber harvest activities (including vegetation management and
fuels management) remove vegetation and cause ground disturbance and
compaction, making the ground more susceptible to erosion (Helms and
Tappeiner 1996, p. 446). This erosion can increase siltation downstream
and potentially damage mountain yellow-legged frog breeding habitat.
Timber harvest may alter the annual hydrograph (timing and volume of
surface flows) in areas where harvests occur. The majority of erosion
caused by timber harvests is from logging roads (Helms and Tappeiner
1996, p. 447). A recent monitoring effort, which was conducted by the
USFS in stream habitats in the northern part of the Sierra Nevada
yellow-legged frog's range, attempted to assess the impact of
vegetation management activities, which would include activities
similar to timber harvest, on mountain yellow-legged frog populations
(Foote et al. 2013, p. 2). However, given the timing of project
implementation, the results were limited to the impacts of these
management activities on mountain yellow-legged frog habitat. The
results of the monitoring suggest these activities did not
significantly impact perennial stream habitat for the mountain yellow-
legged frog, although there were instances of habitat degradation
attributed to sedimentation resulting from road decommissioning and
culvert replacement (Foote et al. 2013, p. 32).
Roadways have the potential to affect riparian habitat by altering
the physical and chemical environment, including alteration of surface-
water run-off, with potential changes to hydrology in high-mountain
lake and stream systems (Brown et al. 2009, pp. 71-72). Roads,
including those associated with timber harvests, have also been found
to contribute to habitat fragmentation and limit amphibian movement,
thus having a negative effect on amphibian species richness. Therefore,
road construction could fragment mountain yellow-legged frog habitat if
a road bisects habitat consisting of water bodies in close proximity.
In the prairies and forests of Minnesota, Lehtinen et al. (1999, pp. 8-
9) found that increased road density reduced amphibian species
richness. DeMaynadier and Hunter (2000, p. 56) found similar results in
a study of eight amphibian species in Maine, although results varied
with road type and width. Results showed that anuran (true frogs, the
group of frogs that includes the mountain yellow-legged frogs) habitat
use and movement were not affected even by a wide, heavily used logging
road (deMaynadier and Hunter 2000, p. 56); this finding suggests that
forest roads may not fragment populations where such roads occur.
Currently, most of the mountain yellow-legged frog populations
occur in National Parks or designated wilderness areas where timber is
not harvested (Bradford et al. 1994, p. 323; Drost and Fellers 1996, p.
421; Knapp and Matthews 2000, p. 430) and where motorized access (and
roads) does not occur. Mountain yellow-legged frog populations outside
of these areas are most often located above the timberline, so timber
harvest activity is not expected to affect the majority of extant
mountain yellow-legged frog populations. There is a higher potential
overlap of timber harvest activities with the species in the northern
and lower elevation portions of the species' ranges where the frogs
occur in streams and meadows in forested environments; in these areas,
populations are very small and fragmented (Brown 2013, unpaginated).
Likewise, at lower elevations of the Sierra Nevada, forest roads and
logging roads are more common (Brown et al. 2009, p. 71). Habitat
effects associated with roads are most likely to occur where existing
roadways occur (for example, see Knapp 1993b, unpaginated). Although
additional roads may be constructed within the range of the mountain
yellow-legged frogs, we are not aware of any proposals to build new
roads at this time.
In riparian areas, the USFS generally maintains standards and
guidelines for land management activities, such as timber harvests,
that are designed to maintain the hydrologic, geomorphic, and ecologic
processes that directly affect streams, stream processes, and aquatic
habitats, and which can limit potential effects of such activities
(Foote et al. 2013, pp. 4, 32). In general, we expect the standards to
be effective in preventing habitat-related effects to these species.
Additionally, neither timber harvests nor roads have been implicated as
important contributors to the decline of this species (Jennings 1996,
pp. 921-941), although habitat alterations due to these activities may,
in site-specific, localized cases, have population-level effects to
mountain yellow-legged frogs. We expect that such cases would be more
likely at lower elevations or in the more northern portion of the
species' range where limited extant populations occur in close
proximity to timber harvest, or where populations occur in drainages
adjacent to roadways. In the proposed rule, we stated that roads and
timber harvest likely present minor prevalent threats to the mountain
yellow-legged frogs factored across the range of the species. We are
clarifying that language, noting that they may pose important habitat-
related effects to the species in localized areas, but are not likely
threats across most of the species' ranges.
Fire and Fire Management Activities
Mountain yellow-legged frogs are generally found at high elevations
in wilderness areas and National Parks where vegetation is sparse and
where fire may have historically played a limited role in the
ecosystem. However, at lower elevations and in the northern portion of
the range, mountain yellow-legged frogs occur in stream or lake
environments within areas that are
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forested to various extents. In some areas within the current range of
the mountain yellow-legged frog, long-term fire suppression has changed
the forest structure and created conditions that increase fire severity
and intensity (McKelvey et al. 1996, pp. 1934-1935). Excessive erosion
and siltation of mountain yellow-legged frog habitats following
wildfire is a concern where shallow, lower elevation aquatic areas
occur below forested stands. However, prescribed fire has been used by
land managers to achieve various silvicultural objectives, including
fuel load reduction. In some systems, fire is thought to be important
in maintaining open aquatic and riparian habitats for amphibians
(Russell et al. 1999, p. 378), although severe and intense wildfires
may reduce amphibian survival, as the moist and permeable skin of
amphibians increases their susceptibility to heat and desiccation
(Russell et al. 1999, p. 374). Amphibians may avoid direct mortality
from fire by retreating to wet habitats or sheltering in subterranean
burrows.
The effects of past fire and fire management activities on
historical populations of mountain yellow-legged frogs are not known.
Neither the direct nor indirect effects of prescribed fire or wildfire
on the mountain yellow-legged frog have been studied. Hossack et al.
(2012, pp. 221, 226), in a study of the effects of six stand-replacing
fires on three amphibians that breed in temporary ponds in low-
elevation dense coniferous forests or in high-elevation open, subalpine
forests in Glacier National Park, found that effects of wildfire on
amphibians may not be evident for several years post-fire with time-
lagged declines. The decline in populations was presumably due to the
proximity of high-severity fires to important breeding habitats, which
resulted in low recruitment of juveniles into the breeding population.
They cautioned, however, that amphibian responses to fire are context
specific and cannot be generalized too broadly; they found no change in
occupancy after wildfire at high elevations where wetlands were in
sparse forest or open meadows where there was less change in canopy
cover and insolation after wildfire. Where fire has occurred in the
steep canyons of southern California where the southern DPS of the
mountain yellow-legged frog occurs, the character of the habitat has
been significantly altered, leading to erosive scouring and flooding of
creeks after surface vegetation is denuded (North 2012, pers. comm.).
North (2012, pers. comm.) reported that at least one population of the
federally endangered southern DPS of the mountain yellow-legged frog,
which occurs in streams, declined substantially after fire on the East
Fork City Creek (San Bernardino Mountains) in 2003 and, by 2012, was
approaching extirpation. Although most populations of mountain yellow-
legged frogs are in alpine habitat that differs from the habitat in
southern California, when they occur in lower-elevation stream
habitats, they could be similarly affected by large wildfires. When a
large fire does occur in occupied habitat, mountain yellow-legged frogs
can be susceptible to both direct mortality (leading to significantly
reduced population sizes) and indirect effects (habitat alteration and
reduced breeding habitat). It is possible that fire has caused
localized extirpations in the past. However, because these species
generally occupy high-elevation habitat, we have determined that fire
is not a significant threat to the mountain yellow-legged frog complex
over much of its current range, although where the species occur at
lower elevations or in the most northerly portion of their ranges,
fire-related changes to habitat may have population-level effects to
the species.
Recreation
Recreational activities that include hiking, camping, and
backpacking take place throughout the Sierra Nevada, whereas off-road
vehicle (ORV) use takes place in areas outside of designated
wilderness. These activities can have significant negative impacts on
many plant and animal species and their habitats (U.S. Department of
Agriculture (USDA) 2001a, pp. 483-493). Extant populations of the
mountain yellow-legged frog complex are primarily located at high
elevations in sub-alpine and alpine habitat within designated
wilderness. High-elevation wilderness areas are ecosystems that are
subject to intense solar exposure; extremes in temperatures,
precipitation levels, and wind; short growing seasons; and shallow,
nutrient-poor soil. Such habitats are typically not resilient to
disturbance (Schoenherr 1992, p. 167; Cole and Landres 1996, p. 170).
In easily accessible areas, heavy foot traffic in riparian areas
can trample vegetation, compact soils, and physically damage stream
banks (Kondolf et al. 1996, pp. 1014, 1019). Human foot, horse,
bicycle, or off-highway motor vehicle trails can replace riparian
habitat with compacted soil (Kondolph et al. 1996, pp. 1014, 1017,
1019), lower the water table, and cause increased erosion where such
activities occur. Bahls (1992, p. 190) reported that the recreational
activity of anglers at high mountain lakes can be locally intense in
western wilderness areas, with most regions reporting a level of use
greater than the fragile lakeshore environments can withstand. Heavy
recreation use has been associated with changes in the basic ecology of
lakes. In the 1970s, Silverman and Erman (1979) found that the most
heavily used back-country lakes in their study had less nitrate and
more iron and aquatic plants than other lakes. These researchers
suggested that erosion at trails and campsites, improper waste
disposal, destruction of vegetation, and campsites might cause an
increase in elements that formerly limited plant growth (Hendee et al.
1990, pp. 435, 436). The NPS considers hiking and backpacking to be a
negligible risk for the mountain yellow-legged frogs within the Parks,
noting that, while hiking and backpacking occur adjacent to many
populations, evidence indicates that risk to habitat is slight to none.
For example, monitoring of a high-use trail that allows thousands of
hikers annually to come into close contact with several populations of
mountain yellow-legged frogs, whose habitat is immediately adjacent to
the trail, shows that the populations have grown substantially over the
last decade (NPS 2013, p. 6). In one location where high hiking levels
may be having an impact due to access via an adjacent road, Yosemite
National Park personnel have restricted access (NPS 2013, p. 6).
Although recreation was noted in 1998 as the fastest growing use of
National Forests (USFS 2001a, p. 453), to our knowledge, no studies to
date have identified a correlation between such recreation-related
impacts to habitat and effects to populations of the mountain yellow-
legged frog complex.
Because of demand for wilderness recreational experiences and
concern about wilderness resource conditions, wilderness land
management now includes standards for wilderness conditions,
implementing permit systems and group-size limits for visitors and
packstock, prohibitions on camping and packstock use close to water,
and other visitor management techniques to reduce impacts to habitat,
including riparian habitat (Cole 2001, pp. 4-5). These wilderness land
management techniques are currently being used in National Forest
Wilderness areas in the Sierra Nevada and in backcountry areas of
Yosemite, Sequoia, and Kings Canyon National Parks. In the proposed
rule, we stated that current recreation activities were considered a
threat of low significance to the species' habitat overall. Based on
[[Page 24273]]
comments from the National Park Service, the USFS, CDFW, and the
public, we have reevaluated the previous analysis and have revised our
finding. Therefore, current habitat effects of recreational activities
are not considered to have population-level effects to mountain yellow-
legged frogs over much of their respective ranges, although there may
be localized effects especially outside of backcountry areas where use
levels are not limited, or where motorized use occurs in extant frog
habitat.
In summary, based on the best available scientific and commercial
information, we consider the modification of habitat and curtailment of
the species' ranges to be a significant and ongoing threat to the
Sierra Nevada yellow-legged frog and northern DPS of the mountain
yellow-legged frog. Habitat fragmentation and degradation (loss of
habitat through competitive exclusion) from stocking and the continued
presence of introduced trout across the majority of the species' range
is a threat of high prevalence. This threat is a significant limiting
factor to persistence and recovery of the species rangewide. Threats of
low prevalence (threats that may be important limiting factors in some
areas, but not across a large part of the mountain yellow-legged frog
complex's range) include dams and water diversions, grazing, packstock
use, timber harvest and roads, recreation, and fire management
activities.
Factor B. Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes
No commercial market for mountain yellow-legged frogs exists, nor
any documented recreational or educational uses for these species.
Scientific research may cause stress to mountain yellow-legged frogs
through disturbance, including disruption of the species' behavior,
handling of individual frogs, and injuries associated with marking and
tracking individuals. However, this is a relatively minor nuisance and
not likely a negative impact to the survival and reproduction of
individuals or the viability of the populations.
Based on the best available scientific and commercial information,
we do not consider overutilization for commercial, recreational,
scientific, or educational purposes to be a threat to the mountain
yellow-legged frog complex now or in the future.
Factor C. Disease or Predation
Predation
Researchers have observed predation of mountain yellow-legged frogs
by the mountain garter snake (Thamnophis elegans elegans), Brewer's
blackbird (Euphagus cyanocephalus), Clark's nutcracker (Nucifraga
columbiana), coyote (Canis latrans), and black bear (Ursus americanus)
(Mullally and Cunningham 1956a, p. 193; Bradford 1991, pp. 176-177;
Jennings et al. 1992, p. 505; Feldman and Wilkinson 2000, p. 102;
Vredenburg et al. 2005, p. 565). However, none of these has been
implicated as a driver of population dynamics, and we expect that such
predation events do not generally have population-level impacts except
where so few individuals remain that such predation is associated with
loss of a population (Bradford 1991, pp 174-177; Jennings 1996, p.
938).
The American bullfrog (Lithobates catesbeiana) is native to the
United States east of the Rocky Mountains, but was introduced to
California about a century ago. The American bullfrog has become common
in California in most permanent lakes and ponds below 1,829 m (6,000
ft) and is implicated in the declines of a number of native frog
species (Jennings 1996, p. 931). Mountain yellow-legged frogs are
thought to be particularly vulnerable to bullfrogs and introduced
crayfish, potentially because the frogs did not evolve with a predator
(Jennings 1996, p. 939). In addition, research indicates that bullfrogs
may outcompete other species of amphibians where fish are present
because bullfrogs are both unpalatable to fish and are naturally
vulnerable to invertebrate predators such as dragonfly (Anisoptera)
nymphs, which fish preferentially consume. Bullfrogs may co-occur with
mountain yellow-legged frogs at lower elevations. On the Plumas
National Forest, sites created as a result of restoration activities
have been invaded by bullfrogs (Brown et al. 2009, pp. 48, 49).
Bullfrogs also occur in the Lake Tahoe Basin (USFS 2000, pp. 530, G-12)
in the vicinity of Fallen Leaf Lake. Bullfrog predation and competition
is expected to have population-level effects where bullfrog populations
occupy the same areas as extant mountain yellow-legged frog
populations.
The most prominent predator of mountain yellow-legged frogs is
introduced trout, whose significance is well-established because it has
been repeatedly observed that the frogs rarely coexist with fish, and
it is known that introduced trout can and do prey on all frog life
stages except for eggs (Grinnell and Storer 1924, p. 664; Mullally and
Cunningham 1956a, p. 190; Cory 1962a, p. 401; 1963, p. 172; Bradford
1989, pp. 775-778; Bradford and Gordon 1992, p. 65; Bradford et al.
1993, pp. 882-888; 1994a, p. 326; Drost and Fellers 1996, p. 422;
Jennings 1996, p. 940; Knapp 1996, p. 14; Knapp and Matthews 2000, p.
428; Knapp et al. 2001, p. 401; Vredenburg 2004, p. 7649; Knapp 2013,
unpaginated). Knapp (1996, pp. 1-44) estimated that 63 percent of lakes
larger than 1 ha (2.5 ac) in the Sierra Nevada contain one or more
nonnative trout species, and that greater than 60 percent of streams
contain nonnative trout. In some areas, trout-occupied waters comprise
greater than 90 percent of total water body surface area (Knapp and
Matthews 2000, p. 434).
The multiple-year tadpole stage of the mountain yellow-legged frog
requires submersion in the aquatic habitat year-round until
metamorphosis. Moreover, all life stages are highly aquatic, increasing
the frog's susceptibility to predation by trout (where they co-occur)
throughout its lifespan. Overwinter mortality due to predation is
especially significant because, when water bodies ice over in winter,
adults and tadpoles move from shallow margins of lakes and ponds into
deeper unfrozen water where they are more vulnerable to predation; fish
encounters in such areas increase, while refuge is less available.
The predation of mountain yellow-legged frogs by fishes observed in
the early 20th century by Grinnell and Storer and the documented
population declines of the 1970s (Bradford 1991, pp. 174-177; Bradford
et al. 1994, pp. 323-327; Stebbins and Cohen 1995, pp. 226-227) were
not the beginning of the mountain yellow-legged frog's decline, but
rather the continuation of a long decline that started soon after fish
introductions to the Sierra Nevada began in the mid-1800s (Knapp and
Matthews 2000, p. 436). Metapopulation theory (Hanski 1997, pp. 85-86)
predicts this type of time lag from habitat modification to population
extinction (Knapp and Matthews 2000, p. 436). In 2004, Vredenburg
(2004, p. 7647) concluded that introduced trout are effective predators
on mountain yellow-legged frog tadpoles and suggested that the
introduction of trout is the most likely reason for the decline of the
mountain yellow-legged frog complex. This threat due to predation by
introduced trout is a significant, prevalent (rangewide) risk to
mountain yellow-legged frogs, and it will persist into the future in
those locations where fish are present. The effect of introduced
bullfrogs is expected to be a substantial continuing threat in those
locations
[[Page 24274]]
where bullfrogs are known to occur presently, but may present more of a
future threat if bullfrogs expand their elevational range as a result
of climate change.
Disease
Over roughly the last 2 decades, pathogens have been associated
with amphibian population declines, mass die-offs, and even extinctions
worldwide (Bradford 1991, pp. 174-177; Blaustein et al. 1994b, pp. 251-
254; Alford and Richards 1999, pp. 506; Muths et al. 2003, p. 357;
Weldon et al. 2004, p. 2100; Rachowicz et al. 2005, p. 1446; Fisher et
al. 2009, p. 292). One pathogen strongly associated with dramatic
declines on all continents that harbor amphibians (all continents
except Antarctica) is the chytrid fungus, Batrachochytrium
dendrobatidis (Bd) (Rachowicz et al. 2005, p. 1442). This chytrid
fungus has now been reported in amphibian species worldwide (Fellers et
al. 2001, p. 945; Rachowicz et al. 2005, p. 1442). Early doubt that
this particular pathogen was responsible for worldwide die-offs has
largely been overcome by the weight of evidence documenting the
appearance, spread, and detrimental effects to affected populations
(Vredenburg et al. 2010, p. 9689). The correlation of notable recent
amphibian declines with reports of outbreaks of fatal chytridiomycosis
(the disease caused by Bd) in montane areas has led to a general
association between high altitude, cooler climates, and population
extirpations associated with Bd (Fisher et al. 2009, p. 298).
Bd affects the mouth parts and epidermal (skin) tissue of tadpoles
and metamorphosed frogs (Fellers et al. 2001, pp. 950-951). The fungus
can reproduce asexually, and can generally withstand adverse conditions
such as freezing or drought (Briggs et al. 2002, p. 38). It also may
reproduce sexually, leading to thick-walled sporangia that would be
capable of long-term survival (for distant transport and persistence in
sites even after all susceptible host animal populations are
extirpated) (Morgan et al. 2007, p. 13849). Adult frogs can acquire
this fungus from tadpoles, and it can also be transmitted between
tadpoles (Rachowicz and Vredenburg 2004, p. 80).
In California, chytridiomycosis has been detected in many amphibian
species, including mountain yellow-legged frogs (Briggs et al. 2002, p.
38; Knapp 2002b, p. 1). The earliest documented case in the mountain
yellow-legged frog complex was in 1998, at Yosemite National Park
(Fellers et al. 2001, p. 945); however, more recent literature shows Bd
occurring in mountain yellow-legged frogs as early as 1975 (Ouellet et
al. (2005, p. 1436; Vredenberg et al. 2010, p. 9689). It is unclear how
Bd was originally transmitted to the frogs (Briggs et al. 2002, p. 39).
Visual examination of 43 tadpole specimens collected between 1955 and
1976 revealed no evidence of Bd infection, yet 14 of 36 specimens
preserved between 1993 and 1999 did have abnormalities attributable to
Bd (Fellers et al. 2001, p. 947). The earliest recorded case of Bd in
mountain yellow-legged frogs is from 1975, and Bd was also identified
on two adult Yosemite toads among over 50 dead, dying, or healthy
Yosemite toads collected during a die-off in 1976 (Green and Kagarise
Sherman 2001, p. 92), although it was not thought to be the cause of
the die-off in the population. Given these records, it is possible that
this pathogen has affected all three amphibian species covered in this
final rule since at least the mid-1970s. Mountain yellow-legged frogs
may be especially vulnerable to Bd infections because all life stages
share the same aquatic habitat nearly year round, facilitating the
transmission of this fungus among individuals at different life stages
(Fellers et al. 2001, p. 951).
During the epidemic phase of chytrid infection into unexposed
populations, rapid die-offs of adult and subadult lifestages are
observed (Vredenburg et al. 2010, p. 9691), with metamorphs being
extremely sensitive to Bd infection (Kilpatrick et al. 2009, p. 113;
Vredenburg et al. 2010, p. 9691; see also Vredenburg 2013,
unpaginated). Field and laboratory experiments indicate that Bd
infection is generally lethal to mountain yellow-legged frogs (Knapp
2005b; Rachowicz 2005, pers. comm.), and is likely responsible for
declines in sites that were occupied as recently as 2002, but where
frogs were absent by 2005 (Knapp 2005b). Rachowicz et al. (2006, p.
1671) monitored several infected and uninfected populations in Sequoia
and Kings Canyon National Parks over multiple years, documenting
dramatic declines and extirpations in only the infected populations.
Rapid die-offs of mountain yellow-legged frogs from chytridiomycosis
have been observed in more than 50 water bodies in the southern Sierra
Nevada in recent years (Briggs et al. 2005, p. 3151). Studies of the
microscopic structure of tissue and other evidence suggests Bd caused
many of the recent extinctions in the Sierra National Forest's John
Muir Wilderness Area and in Kings Canyon National Park, where 41
percent of the populations went extinct between 1995 and 2002 (Knapp
2002a, p. 10).
In several areas where detailed studies of the effects of Bd on the
mountain yellow-legged frog are ongoing, substantial declines have been
observed following the course of the disease infection and spread.
Survey results from 2000 in Yosemite and Sequoia and Kings Canyon
National Parks indicated that 17 percent of frog populations in
Yosemite and 27 percent of the mountain yellow-legged frog populations
sampled across both Sequoia and Kings Canyon National Parks showed
evidence of Bd infection, although the proportion of infected frogs at
each site varied greatly and disease incidence varied within each Park
(Briggs et al. 2002, p. 40) (In the proposed rule, these two figures
were averaged across all three parks; these numbers reflect the text
presented in Briggs et al. 2002). In both 2003 and 2004, 19 percent of
the populations that were sampled in Sequoia and Kings Canyon National
Parks were infected with Bd (Rachowicz 2005, pp. 2-3). By 2005, 91
percent of assayed populations in Yosemite National Park showed
evidence of Bd infection (Knapp 2005b, pp. 1-2), and the number of
occupied sites in Sequoia and Kings Canyon National Parks had decreased
by 47 percent from those known to be occupied 3 to 8 years previously
(Knapp 2005b, pers. comm). Currently, it is believed that all
populations in Yosemite Park are infected with Bd (Knapp et al. 2011,
p. 9).
The effects of Bd on host populations of the mountain yellow-legged
frog are variable, ranging from extirpation to persistence with a low
level of infection (Briggs et al. 2002, pp. 40-41). When Bd infection
first occurs in a population, the most common outcome is epidemic
spread of the disease and population extirpation (Briggs et al. 2010,
p. 9699). Die-offs are characterized by rapid onset of high-level Bd
infections, followed by death due to chytridiomycosis. Although most
populations that are newly exposed to Bd are driven to extirpation
following the arrival of Bd, some populations that experience Bd-caused
population crashes are not extirpated, and some may even recover
despite ongoing chytridiomycosis (Briggs et al. 2010, pp. 9695-9696).
However, it is apparent that even at sites exhibiting population
persistence with Bd, high mortality of metamorphosing frogs persists,
and this phenomenon may explain the lower abundances observed in such
populations (Briggs et al. 2010, p. 9699).
Vredenburg et al. (2010a, pp. 2-4) studied frog populations before,
during, and after the infection and spread of Bd in three study basins
constituting 13, 33,
[[Page 24275]]
and 42 frog populations, respectively, then comprising the most intact
metapopulations remaining for these species throughout their range. The
spread of Bd averaged 688 m/year (yr) (2,257 ft/yr), reaching all areas
of the smaller basin in 1 year, and taking 3 to 5 years to completely
infect the larger basins, progressing like a wave across the landscape.
The researchers documented die-offs following the spread of Bd, with
decreased population growth rates evident within the first year of
infection. Basinwide, metapopulations crashed from 1,680 to 22
individuals (northern DPS of the mountain yellow-legged frog) in
Milestone Basin, with 9 of 13 populations extirpated; from 2,193 to 47
individuals (northern DPS of the mountain yellow-legged frog) in Sixty
Lakes Basin, with 27 of 33 populations extirpated; and from 5,588 to
436 individuals (Sierra Nevada yellow-legged frog) in Barrett Lakes
Basin, with 33 of 42 populations extirpated. The evidence is clear that
Bd can and does decimate newly infected frog populations. Moreover,
this rangewide population threat is acting upon a landscape already
impacted by habitat modification and degradation by introduced fishes
(see Factor A discussion, above). As a result, remnant populations in
fishless lakes are now affected by Bd.
Vredenburg et al. (2010a, p. 3) projected that, at current
extinction rates, and given the disease dynamics of Bd (infected
tadpoles succumb to chytridiomycosis at metamorphosis), most if not
all, extant populations within the recently infected basins they
studied would go extinct within the next 3 years. Available data (CDFW,
unpubl. data; Knapp 2005b; Rachowicz 2005, pers. comm.; Rachowicz et
al. 2006, p. 1671) indicate that Bd is now widespread throughout the
Sierra Nevada and, although it has not infected all populations at this
time, it is a serious and substantial threat rangewide to the mountain
yellow-legged frog complex.
Other diseases have also been reported as adversely affecting
amphibian species, and these may be present within the range of the
mountain yellow-legged frog. Bradford (1991, pp. 174-177) reported an
outbreak of red-leg disease in Kings Canyon National Park, and
suggested this was a result of overcrowding within a mountain yellow-
legged frog population. Red-leg disease is caused by the bacterial
pathogen Aeromonas hydrophila, along with other pathogens. Red-leg
disease is opportunistic and successfully attacks immune-suppressed
individuals, and this pathogen appears to be highly contagious,
affecting the epidermis and digestive tract of otherwise healthy
amphibians (Shotts 1984, pp. 51-52; Carey 1993, p. 358; Carey and
Bryant 1995, pp. 14-15). Although it has been correlated with decline
of a frog population in at least one case, red-leg disease is not
thought to be a significant contributor to observed frog population
declines rangewide, based on the available literature.
Saprolegnia is a globally distributed fungus that commonly attacks
all life stages of fishes (especially hatchery-reared fishes), and has
recently been documented to attack and kill egg masses of western toads
(Bufo boreas) (Blaustein et al. 1994b, p. 252). This pathogen may be
introduced through fish stocking, or it may already be established in
the aquatic ecosystem. Fishes and migrating or dispersing amphibians
may be vectors for this fungus (Blaustein et al. 1994b, p. 253;
Kiesecker et al. 2001, p. 1068). Saprolegnia has been reported in the
southern DPS of the mountain yellow-legged frog (North 2012, pers.
comm.); however, its occurrence within the Sierran range of the
mountain yellow-legged frog complex and associated influence on
population dynamics (if any) are unknown.
Other pathogens of concern for amphibian species include
ranaviruses (Family Iridoviridae). Mao et al. (1999, pp. 49-50)
isolated identical iridoviruses from co-occurring populations of the
threespine stickleback (Gasterosteus aculeatus) and the red-legged frog
(Rana aurora), indicating that infection by a given virus is not
limited to a single species, and that iridoviruses can infect animals
of different taxonomic classes. This suggests that virus-hosting trout
introduced into mountain yellow-legged frog habitat may be a vector for
amphibian viruses. However, definitive mechanisms for the transmission
to the mountain yellow-legged frog remain unknown. No viruses were
detected in the mountain yellow-legged frogs that Fellers et al. (2001,
p. 950) analyzed for Bd. In Kings Canyon National Park, Knapp (2002a,
p. 20) found mountain yellow-legged frogs showing symptoms attributed
to a ranavirus (Knapp 2013, unpaginated). To date, ranaviruses remain a
concern for the mountain yellow-legged frog complex, but the available
information does not indicate they are negatively affecting
populations.
It is unknown whether amphibian pathogens in the high Sierra Nevada
have always coexisted with amphibian populations or if the presence of
such pathogens is a recent phenomenon. However, it has been suggested
that the susceptibility of amphibians to pathogens may have recently
increased in response to anthropogenic environmental disruption (Carey
1993, pp. 355-360; Blaustein et al. 1994b, p. 253; Carey et al. 1999,
p. 7). This hypothesis suggests that environmental changes may be
indirectly responsible for certain amphibian die-offs due to immune
system suppression of tadpoles or post-metamorphic amphibians (Carey
1993, p. 358; Blaustein et al. 1994b, p. 253; Carey et al. 1999, pp. 7-
8). Pathogens such as Aeromonas hydrophila, which are present in fresh
water and in healthy organisms, may become more of a threat,
potentially causing localized amphibian population die-offs when the
immune systems of individuals within the host population are suppressed
(Carey 1993, p. 358; Carey and Bryant 1995, p. 14).
The contribution of Bd as an environmental stressor and limiting
factor on mountain yellow-legged frog population dynamics is currently
extremely high, and it poses a significant current and continuing
threat to remnant uninfected populations in the southern Sierra Nevada.
Its effects are most dramatic following the epidemic stage as it
spreads across newly infected habitats; massive die-off events follow
the spread of the fungus, and it is likely that survival of mountain
yellow-legged frogs through the metamorphosis stage is substantially
reduced even years after the initial epidemic (Rachowicz et al. 2006,
pp. 1679-1680). The relative impact from other diseases and the
interaction of other stressors and disease on the immune systems of
mountain yellow-legged frogs remains poorly documented to date.
In summary, based on the best available scientific and commercial
information, we consider the threats of predation and disease to be
significant, ongoing threats to the Sierra Nevada yellow-legged frog
and the northern DPS of the mountain yellow-legged frog. These threats
include predation by bullfrogs and introduced fishes, and amphibian
pathogens (most specifically, the chytrid fungus), two primary driving
forces leading to population declines in the mountain yellow-legged
frog complex. These are highly prevalent threats, and they are
predominant limiting factors hindering population viability and
precluding recovery across the ranges of the mountain yellow-legged
frog complex.
[[Page 24276]]
Factor D. The Inadequacy of Existing Regulatory Mechanisms
In determining whether the inadequacy of regulatory mechanisms
constitutes a threat to the mountain yellow-legged frog complex, we
analyzed the existing Federal and State laws and regulations that may
address the threats to these species or contain relevant protective
measures. Regulatory mechanisms are typically nondiscretionary and
enforceable, and may preclude the need for listing if such mechanisms
are judged to adequately address the threat(s) to the species such that
listing is not warranted. Conversely, threats on the landscape are not
ameliorated where existing regulatory mechanisms are not adequate (or
when existing mechanisms are not adequately implemented or enforced).
Federal Wilderness Act
The Wilderness Act of 1964 (16 U.S.C. 1131 et seq.) established a
National Wilderness Preservation System made up of federally owned
areas designated by Congress as ``wilderness'' for the purpose of
preserving and protecting designated areas in their natural condition.
The Wilderness Act states the use of these areas with limited exception
are subject to the following restrictions: (1) New or temporary roads
cannot be built; (2) motor vehicles, motorized equipment, or motorboats
cannot be used; (3) aircraft cannot land; (4) no form of mechanical
transport can occur; and (5) no structure or installation may be built.
In addition, a special provision within the Wilderness Act stipulated
that, except for valid existing rights, effective January 1, 1984, the
minerals within designated wilderness areas would be withdrawn from all
forms of appropriation under mining laws, precluding new mining claims
within designated wilderness after that date (see Hendee et al. 1990,
p. 508). A large number of mountain yellow-legged frog locations occur
within wilderness areas managed by the USFS and NPS and, therefore, are
afforded protection from direct loss or degradation of habitat by some
human activities (such as development, commercial timber harvest, road
construction, and some fire management actions). Livestock grazing and
fish stocking both occur within designated wilderness areas on lands
within the National Forest System.
National Forest Management Act of 1976
Under the National Forest Management Act of 1976, as amended (NFMA)
(16 U.S.C. 1600 et seq.), the USFS is tasked with managing National
Forest lands based on multiple-use, sustained-yield principles, and
with implementing land and resource management plans (LRMP) on each
National Forest to provide for a diversity of plant and animal
communities. The purpose of an LRMP is to guide and set standards for
all natural resource management activities for the life of the plan (10
to 15 years). NFMA requires the USFS to incorporate standards and
guidelines into LRMPs. The 1982 planning regulations for implementing
NFMA (47 FR 43026; September 30, 1982), under which all existing forest
plans in the Sierra Nevada were prepared until recently, guided
management of National Forests and required that fish and wildlife
habitat on National Forest system lands be managed to maintain viable
populations of existing native and desired nonnative vertebrate species
in the planning area. A viable population is defined as a population of
a species that continues to persist over the long term with sufficient
distribution to be resilient and adaptable to stressors and likely
future environments. In order to insure that viable populations would
be maintained, the 1982 planning regulations directed that habitat must
be provided to support, at least, a minimum number of reproductive
individuals and that habitat must be well-distributed so that those
individuals could interact with others in the planning area.
On April 9, 2012, the USFS published a final rule (77 FR 21162)
amending 36 CFR 219 to adopt new National Forest System land management
regulations that guide the development, amendment, and revision of
LRMPs for all Forest System lands. These revised regulations, which
became effective on May 9, 2012, replaced the 1982 planning rule. The
2012 planning rule requires that the USFS maintain viable populations
of species of conservation concern at the discretion of regional
foresters. This rule could thereby result in removal of the limited
protections that are currently in place for mountain yellow-legged
frogs under the Sierra Nevada Forest Plan Amendment (SNFPA), as
described below.
Sierra Nevada Forest Plan Amendment
In 2001, a record of decision was signed by the USFS for the Sierra
Nevada Forest Plan Amendment (SNFPA), based on the final environmental
impact statement for the SNFPA effort and prepared under the 1982 NFMA
planning regulations. The Record of Decision amends the USFS Pacific
Southwest Regional Guide, the Intermountain Regional Guide, and the
LRMPs for National Forests in the Sierra Nevada and Modoc Plateau. This
document affects land management on all National Forests throughout the
range of the mountain yellow-legged frog complex. The SNFPA addresses
and gives management direction on issues pertaining to old forest
ecosystems; aquatic, riparian, and meadow ecosystems; fire and fuels;
noxious weeds; and lower west-side hardwood ecosystems of the Sierra
Nevada. In January 2004, the USFS amended the SNFPA, based on the final
supplemental environmental impact statement, following a review of fire
and fuels treatments, compatibility with the National Fire Plan,
compatibility with the Herger-Feinstein Quincy Library Group Forest
Recovery Pilot Project, and effects of the SNFPA on grazing,
recreation, and local communities (USDA 2004, pp. 26-30).
Relevant to the mountain yellow-legged frog complex, the Record of
Decision for SNFPA aims to protect and restore aquatic, riparian, and
meadow ecosystems, and to provide for the viability of associated
native species through implementation of an aquatic management
strategy. The aquatic management strategy is a general framework with
broad policy direction. Implementation of this strategy was intended to
take place at the landscape and project levels. Nine goals are
associated with the aquatic management strategy:
(1) The maintenance and restoration of water quality to comply with
the Clean Water Act (CWA) and the Safe Drinking Water Act;
(2) The maintenance and restoration of habitat to support viable
populations of native and desired nonnative riparian-dependent species,
and to reduce negative impacts of nonnative species on native
populations;
(3) The maintenance and restoration of species diversity in
riparian areas, wetlands, and meadows to provide desired habitats and
ecological functions;
(4) The maintenance and restoration of the distribution and
function of biotic communities and biological diversity in special
aquatic habitats (such as springs, seeps, vernal pools, fens, bogs, and
marshes);
(5) The maintenance and restoration of spatial and temporal
connectivity for aquatic and riparian species within and between
watersheds to provide physically, chemically, and biologically
unobstructed movement for their survival, migration, and reproduction;
(6) The maintenance and restoration of hydrologic connectivity
between
[[Page 24277]]
floodplains, channels, and water tables to distribute flood flows and
to sustain diverse habitats;
(7) The maintenance and restoration of watershed conditions as
measured by favorable infiltration characteristics of soils and diverse
vegetation cover to absorb and filter precipitation, and to sustain
favorable conditions of streamflows;
(8) The maintenance and restoration of instream flows sufficient to
sustain desired conditions of riparian, aquatic, wetland, and meadow
habitats, and to keep sediment regimes within the natural range of
variability; and
(9) The maintenance and restoration of the physical structure and
condition of streambanks and shorelines to minimize erosion and sustain
desired habitat diversity.
If these goals of the aquatic management strategy are pursued and
met, threats to the mountain yellow-legged frog complex resulting from
habitat alterations could be reduced. However, the aquatic management
strategy is a generalized approach that does not contain specific
implementation timeframes or objectives, and it does not provide direct
protections for the mountain yellow-legged frog. Additionally, as
described above, the April 9, 2012, final rule (77 FR 21162) that
amended 36 CFR 219 to adopt new National Forest System land management
planning regulations could result in removal of the limited protections
that are currently in place for mountain yellow-legged frogs under the
SNFPA.
National Park Service Organic Act
The statute establishing the National Park Service, commonly
referred to as the National Park Service Organic Act (39 Stat. 535; 16
U.S.C. 1, 2, 3, and 4), states that the NPS will administer areas under
their jurisdiction ``. . . by such means and measures as conform to the
fundamental purpose of said parks, monuments, and reservations, which
purpose 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.'' Park managers must take action to
ensure that ongoing NPS activities do not cause impairment. In cases of
doubt as to the impact of activities on park natural resource, the Park
Service is to decide in favor of protecting the natural resources.
Sequoia, Kings Canyon, and Yosemite National Parks began phasing out
fish stocking by the State in 1969 and terminated this practice
entirely in 1991 (Knapp 1996, p. 9).
Federal Power Act
The Federal Power Act of 1920, as amended (FPA) (16 U.S.C. 791 et
seq.) was enacted to regulate non-federal hydroelectric projects to
support the development of rivers for energy generation and other
beneficial uses. The FPA provides for cooperation between the Federal
Energy Regulatory Commission (Commission) and other Federal agencies in
licensing and relicensing power projects. The FPA mandates that each
license includes conditions to protect, mitigate, and enhance fish and
wildlife and their habitat affected by the project. However, the FPA
also requires that the Commission give equal consideration to competing
priorities, such as power and development, energy conservation,
protection of recreational opportunities, and preservation of other
aspects of environmental quality. Further, the FPA does not mandate
protections of habitat or enhancements for fish and wildlife species,
but provides a mechanism for resource agency recommendations that are
incorporated into a license at the discretion of the Commission.
Additionally, the FPA provides for the issuance of a license for the
duration of up to 50 years, and the FPA contains no provision for
modification of the project for the benefit of species, such as
mountain yellow-legged frogs, before a current license expires.
Although most reservoirs and water diversions are located at lower
elevations than those at which extant mountain yellow-legged frog
populations occur, numerous extant populations occur within watersheds
that feed into developed and managed aquatic systems (such as
reservoirs and water diversions) operated for the purpose of power
generation and regulated by the FPA and may be considered during
project relicensing.
State
California Endangered Species Act
This section has been updated from the information presented in the
proposed rule, and discussion of CDFW's current fish-stocking practices
has been moved to the Factor A discussion of Habitat Modification Due
to Introduction of Trout to Historically Fishless Areas.
The California Endangered Species Act (CESA) (California Fish and
Game Code, section 2080 et seq.) prohibits the unauthorized take of
State-listed endangered or threatened species. CESA requires State
agencies to consult with CDFW on activities that may affect a State-
listed species, and mitigate for any adverse impacts to the species or
its habitat. Pursuant to CESA, it is unlawful to import or export,
take, possess, purchase, or sell any species or part or product of any
species listed as endangered or threatened. The State may authorize
permits for scientific, educational, or management purposes, and allow
take that is incidental to otherwise lawful activities. On April 1,
2013, the Sierra Nevada yellow-legged frog was listed as a threatened
species and the mountain yellow-legged frog (Statewide) was listed as
an endangered species under CESA (CDFW 2013, p. 1).
While the listing of the Sierra Nevada yellow-legged frog and the
mountain yellow-legged frog under CESA provide some protections to
these species, as State regulation prohibits the unauthorized take of
State-listed species, the definition of take under CESA does not
include habitat modification or degradation. Additionally, the majority
of the lands occupied by these species are federally managed lands, so
there is limited jurisdiction in which to regulate land management
activities that may affect these species.
Overall, existing Federal and State laws and regulatory mechanisms
currently offer some level of protection for the mountain yellow-legged
frog complex. While not the intent of the Wilderness Act, the mountain
yellow-legged frogs receive ancillary protection from the Wilderness
Act due to its prohibitions on development, road construction, and
timber harvest, and associated standards and guidelines that limit
visitor and packstock group sizes and use. With the exception of the
National Park Service Organic Act, the existing regulatory mechanisms
have not been effective in reducing threats to mountain yellow-legged
frogs and their habitat from fish stocking and the continuing presence
of nonnative fish. Nor have these mechanisms been effective in
protecting populations from infection by diseases, although Forest
Service standards and guidelines have likely reduced threats associated
with grazing, timber harvest, and recreation use. Although State
regulations under CESA provide some protection against take of the
mountain yellow-legged frogs, the definition of take under CESA does
not include habitat modification or degradation.
Factor E. Other Natural or Manmade Factors Affecting Its Continued
Existence
The mountain yellow-legged frog is sensitive to environmental
change or
[[Page 24278]]
degradation because it has an aquatic and terrestrial life history and
highly permeable skin that increases exposure of individuals to
substances in the water, air, and terrestrial substrates (Blaustein and
Wake 1990, p. 203; Bradford and Gordon 1992. p. 9; Blaustein and Wake
1995, p. 52; Stebbins and Cohen 1995, pp. 227-228). Several natural or
anthropogenically influenced changes, including contaminant deposition,
acid precipitation, increases in ambient ultraviolet radiation, and
climate change, have been implicated as contributing to amphibian
declines (Corn 1994, pp. 62-63; Alford and Richards 1999, pp. 2-7).
There are also documented incidences of direct mortality of, or the
potential for direct disturbance to, individuals from some activities
already discussed; in severe instances, these actions may have
population-level consequences. As presented in the proposed rule (78 FR
24472, April 25, 2013), contaminants, acid precipitation, and ambient
ultraviolet radiation are not known to pose a threat (current or
historical) to the mountain yellow-legged frog and, therefore, are not
discussed further. Please refer to the proposed listing rule for the
Sierra Nevada yellow-legged frog, the northern DPS of the mountain
yellow-legged frog, and the Yosemite toad (78 FR 24472, April 25, 2013)
for a detailed discussion of contaminants, acid precipitation, and
ambient ultraviolet radiation.
Climate Change
Our analysis under the Act includes consideration of ongoing and
projected changes in climate. The terms ``climate'' and ``climate
change'' are defined by the Intergovernmental Panel on Climate Change
(IPCC). The term ``climate'' refers to the mean and variability of
different types of weather conditions over time, with 30 years being a
typical period for such measurements, although shorter or longer
periods also may be used (IPCC 2007a, p. 1450; IPCC 2013a, Annex III).
The term ``climate change'' thus refers to a change in the mean or
variability of one or more measures of climate (for example,
temperature or precipitation) that persists for an extended period,
typically decades or longer, whether the change is due to natural
variability, human activity, or both (IPCC 2007a, p. 1450; IPCC 2013a,
Annex III). A recent compilation of climate change and its effects is
available from reports of the Intergovernmental Panel on Climate Change
(IPCC) (IPCC 2013b, entire).
Global climate projections are informative and, in some cases, the
only or the best scientific information available for us to use.
However, projected changes in climate and related impacts can vary
substantially across and within different regions of the world (for
example, IPCC 2007a, pp. 8-12). Therefore, we use downscaled
projections when they are available and have been developed through
appropriate scientific procedures, because such projections provide
higher resolution information that is more relevant to the spatial
scales used for analyses of a given species (see Glick et al. 2011, pp.
58-61, for a discussion of downscaling). With regard to our analysis
for the Sierra Nevada of California (and western United States),
downscaled projections are available, yet even downscaled climate
models contain some uncertainty.
Variability exists in outputs from different climate models, and
uncertainty regarding future GHG emissions is also a factor in modeling
(PRBO 2011, p. 3). A general pattern that holds for many predictive
models indicates northern areas of the United States will become
wetter, and southern areas (particularly the Southwest) will become
drier. These models also predict that extreme events, such as heavier
storms, heat waves, and regional droughts, may become more frequent
(Glick et al. 2011, p. 7). Moreover, it is generally expected that the
duration and intensity of droughts will increase in the future (Glick
et al. 2011, p. 45; PRBO 2011, p. 21).
The last century has included some of the most variable climate
reversals documented, at both the annual and near-decadal scales,
including a high frequency of El Ni[ntilde]o (associated with more
severe winters) and La Ni[ntilde]a (associated with milder winters)
events (reflecting drought periods of 5 to 8 years alternating with wet
periods) (USDA 2001b, p. 33). Scientists have confirmed a longer
duration climate cycle termed the Pacific Decadal Oscillation (PDO),
which operates on cycles between 2 to 3 decades, and generally is
characterized by warm and dry (PDO positive) followed by cool and wet
cycles (PDO negative) (Mantua et al. 1997, pp. 1069-1079; Zhang et al.
1997, pp. 1004-1018). Snowpack is seen to follow this pattern--heavier
in the PDO negative phase in California, and lighter in the positive
phase (Mantua et al. 1997, p. 14; Cayan et al. 1998, p. 3148; McCabe
and Dettinger 2002, p. 24).
For the Sierra Nevada ecoregion, climate models predict that mean
annual temperatures will increase by 1.8 to 2.4 [deg]C (3.2 to
4.3[emsp14][deg]F) by 2070, including warmer winters with earlier
spring snowmelt and higher summer temperatures. However, it is expected
that temperature and climate variability will vary based on topographic
diversity (for example, wind intensity will determine east versus west
slope variability) (PRBO 2011, p. 18). Mean annual rainfall is
projected to decrease from 9.2-33.9 cm (3.6-13.3 in) by 2070; however,
projections have high uncertainty and one study predicts the opposite
effect (PRBO 2011, p. 18). Given the varied outputs from differing
modeling assumptions, and the influence of complex topography on
microclimate patterns, it is difficult to draw general conclusions
about the effects of climate change on precipitation patterns in the
Sierra Nevada (PRBO 2011, p. 18). Snowpack is, by all projections,
going to decrease dramatically (following the temperature rise and more
precipitation falling as rain) (Kadir et al. 2013, pp. 76-80). Higher
winter streamflows, earlier runoff, and reduced spring and summer
streamflows are projected, with increasing severity in the southern
Sierra Nevada (PRBO 2011, pp. 20-22); (Kadir et al. 2013, pp. 71-75).
Snow-dominated elevations of 2,000-2,800 m (6,560-9,190 ft) will be
the most sensitive to temperature increases, and a warming of 5 [deg]C
(9[emsp14][deg]F) is projected to shift center timing (the measure when
half a stream's annual flow has passed a given point in time) to more
than 45 days earlier in the year as compared to the 1961-1990 baseline
(PRBO 2011, p. 23). Lakes, ponds, and other standing waters fed by
snowmelt or streams are likely to dry out or be more ephemeral during
the non-winter months (Lacan et al. 2008, pp. 216-222; PRBO 2011, p.
24). This pattern could influence ground water transport, and springs
may be similarly depleted, leading to lower lake levels.
Blaustein et al. (2010, pp. 285-300) provide an exhaustive review
of potential direct and indirect and habitat-related effects of climate
change to amphibian species, with documentation of effects in a number
of species where such effects have been studied. Altitudinal range
shifts with changes in climate have been reported in some regions. They
note that temperature can influence the concentration of dissolved
oxygen in aquatic habitats, with warmer water generally having lower
concentrations of dissolved oxygen, and that water balance heavily
influences amphibian physiology and behavior. They predict that
projected changes in temperature and precipitation are likely to
increase habitat loss and alteration for those species living in
sensitive habitats, such
[[Page 24279]]
as ephemeral ponds and alpine habitats (Blaustein et al. 2010, pp. 285-
287).
Because environmental cues such as temperature and precipitation
are clearly linked to onset of reproduction in many species, climate
change will likely affect the timing of reproduction in many species,
potentially with different sexes responding differently to climate
change. For example, males of two newt species (Triturus spp.) showed a
greater degree of change in arrival date at breeding ponds (Blaustein
et al. 2010, p. 288). Lower concentrations of dissolved oxygen in
aquatic habitats may negatively affect developing embryos and larvae,
in part because increases in temperature increase the oxygen
consumption rate in amphibians. Reduced oxygen concentrations have also
been shown to result in accelerated hatching in ranid frogs, but at a
smaller size, while larval development and behavior may also be
affected and may be mediated by larval density and food availability
(Blaustein et al. 2010, pp. 288-289).
Increased temperatures can reduce time to metamorphosis, which can
increase chances of survival where ponds dry, but also result in
metamorphosis at a smaller size, suggesting a likely trade-off between
development and growth, which may be exacerbated by climate change and
have fitness consequences for adults (Blaustein et al. 2010, pp. 289-
290). Changes in terrestrial habitat, such as changed soil moisture and
vegetation, can also directly affect adult and juvenile amphibians,
especially those adapted to moist forest floors and cool, highly
oxygenated water that characterizes montane regions. Climate change may
also interact with other stressors that may be acting on a particular
species, such as disease and contaminants (Blaustein et al. 2010, pp.
290-299).
A recent paper (Kadir et al. 2013, entire) provides specific
information on the effects of climate change in the Sierra Nevada. The
report found that glaciers in the Sierra Nevada have decreased in area
over the past century, and glacier shrinkage results in earlier peak
water runoff and drier summer conditions. Another result from the
report is that the lower edge of the conifer-dominated forests in the
Sierra Nevada has been retreating upslope over the past 60 years.
Regarding wildfire, since 1950, annual acreage burned in wildfires
statewide has been increasing in California, and in the western United
States, large wildfires have become more frequent, increasing in tandem
with rising spring and summer temperatures. Finally, the report found
that today's subalpine forests in the Sierra Nevada are much denser--
that is, comprise more small-diameter trees--than they were over 70
years ago. During this time period, warmer temperatures, earlier
snowmelt, and more rain than snow occurred in this region. Many of
these changes in the Sierra Nevada of California due to climate are
likely to influence mountain yellow-legged frogs because both mountain
yellow-legged frog species in the Sierra Nevada are highly vulnerable
to climate change because changing hydrology and habitat in the Sierra
Nevada will likely have impacts on remaining populations (Viers et al.
2013, pp. 55, 56).
Vulnerability of species to climate change is a function of three
factors: Sensitivity of a species or its habitat to climate change,
exposure of individuals to such physical changes in the environment,
and their capacity to adapt to those changes (Glick et al. 2011, pp.
19-22). Critical sensitivity elements broadly applicable across
organizational levels (from species through habitats to ecosystems) are
associated with physical variables, such as hydrology (timing,
magnitude, and volume of waterflows), fire regime (frequency, extent,
and severity of fires), and wind (Glick et al. 2011, pp. 39-40).
Species-level sensitivities generally include physiological factors,
such as changes in temperature, moisture, or pH as they influence
individuals; these also include dependence on sensitive habitats,
ecological linkages to other species, and changes in phenology (timing
of key life-history events) (Glick et al. 2011, pp. 40-41).
Exposure to environmental stressors renders species vulnerable to
climate change impacts, either through direct mechanisms (for example,
physical temperature extremes or changes in solar radiation), or
indirectly through impacts upon habitat (hydrology; fire regime; or
abundance and distribution of prey, competitors, or predator species).
A species' capacity to adapt to climate change is increased by
behavioral plasticity (the ability to modify behavior to mitigate the
impacts of the stressor), dispersal ability (the ability to relocate to
meet shifting conditions), and evolutionary potential (for example,
shorter lived species with multiple generations have more capacity to
adapt through evolution) (Glick et al. 2011, pp. 48-49).
The International Union for Conservation of Nature describes five
categories of life-history traits that render species more vulnerable
to climate change (Foden et al. 2008 in Glick et al. 2011, p. 33): (1)
Specialized habitat or microhabitat requirements, (2) narrow
environmental tolerances or thresholds that are likely to be exceeded
under climate change, (3) dependence on specific triggers or cues that
are likely to be disrupted (for example, rainfall or temperature cues
for breeding, migration, or hibernation), (4) dependence on
interactions between species that are likely to be disrupted, and (5)
inability or poor ability to disperse quickly or to colonize more
suitable range. We apply these criteria in this final rule to assess
the vulnerability of mountain yellow-legged frogs to climate change.
At high elevations, where most extant populations occur, mountain
yellow-legged frogs depend on high mountain lakes where both adult and
larval frogs overwinter under ice for up to 9 months of the year.
Overwintering under ice poses physiological problems for the frogs,
most notably the depletion of oxygen in the water during the winter
(Bradford 1983, p. 1171). Bradford (1983, pp. 1174-1182) has found,
based on lab and field results, that tadpoles are more resistant to low
dissolved oxygen levels than adult frogs; after two drought years that
were followed by a severe winter, all frogs in 21 of 26 study lakes
were lost (with the exception of one 2.1-m (6.9-ft) deep lake that
contained only one individual), while tadpoles survived in all but one
of the shallowest lakes. Losses were apparently due to oxygen depletion
in a year when there was exceptional precipitation, ice depths that
were thicker than usual, and lake thawing was 5 to 6 weeks later than
the previous year. The survival of adults in substantial numbers was
significantly correlated with lake depth and confined to lakes deeper
than 4 m (13.1 ft).
Bradford (1983, pp. 1174-1179) found that mean oxygen concentration
in lakes was directly related to maximum lake depth, with dissolved
oxygen levels declining throughout the winter. He also found that a
thickened ice layer on a lake causes the lake to become effectively
more shallow, leading to an increased rate of oxygen depletion
(Bradford 1983, p. 1178). Studies of winterkill of fish due to oxygen
depletion also show that oxygen depletion is inversely related to lake
depth and occurs most rapidly in shallow lakes relative to deeper lakes
(See review in Bradford 1983, p. 1179). Bradford (1983, p. 1179)
considered the possibility that winterkill of the frogs was due to
freezing, but dismissed the potential because some of the lakes where
winterkill occurred were deeper than the probable maximum ice depth in
that year. Because the deeper lakes
[[Page 24280]]
that once supported frog populations now harbor introduced trout
populations and are generally no longer available as refugia for frogs,
the shallower lakes where frogs currently occur may be more vulnerable
to weather extremes in a climate with increased variability, including
drought years and years with exceptional severe cold winters. Such
episodic stressors may have been infrequent in the past, but appear to
be increasing, and they are important to long-lived species with small
populations.
In summer, reduced snowpack and enhanced evapotranspiration
following higher temperatures can dry out ponds that otherwise would
have sustained rearing tadpoles (Lacan et al. 2008, p. 220), and may
also reduce fecundity (egg production) (Lacan et al. 2008, p. 222).
Lacan et al. (2008, p. 211) observed that most frog breeding occurred
in the smaller, fishless lakes of Kings Canyon National Park that are
shallow and prone to summer drying. Thus, climate change will likely
reduce available breeding habitat for mountain yellow-legged frogs and
lead to greater frequency of stranding and death of tadpoles as such
lakes dry out earlier in the year (Corn 2005, p. 64; Lacan et al. 2008,
p. 222).
Earlier snowmelt is expected to cue breeding earlier in the year.
The advance of this primary signal for breeding phenology in montane
and boreal habitats (Corn 2005, p. 61) may have both positive and
negative effects. Additional time for growth and development may render
larger individuals more fit to overwinter; however, earlier breeding
may also expose young tadpoles (or eggs) to killing frosts in more
variable conditions of early spring (Corn 2005, p. 60).
Whether mountain yellow-legged frogs depend on other species that
may be affected either positively or negatively by climate change is
unclear. Climate change may alter invertebrate communities (PRBO 2011
p. 24). In one study, an experimental increase in stream temperature
was shown to decrease density and biomass of invertebrates (Hogg and
Williams 1996, p. 401). Thus, climate change might have a negative
impact on the mountain yellow-legged frog prey base.
Indirect effects from climate change may lead to greater risk to
mountain yellow-legged frog population persistence. For example, fire
intensity and magnitude are projected to increase (PRBO 2011, pp. 24-
25), and, therefore, the contribution and influence of this stressor
upon frog habitat and populations will increase. Climate change may
alter lake productivity through changes in water chemistry, the extent
and timing of mixing, and nutrient inputs from increased fires, all of
which may influence community dynamics and composition (Melack et al.
1997, p. 971; Parker et al. 2008, p. 12927). These changes may not all
be negative; for example, water chemistry and nutrient inputs, along
with warmer summer temperatures, could increase net primary
productivity in high mountain lakes to enhance frog food sources,
although changes in net primary productivity may also negatively affect
invertebrate prey species endemic to oligotrophic lakes (low nutrient,
low productivity).
Carey (1993, p. 359) has suggested that, where environmental
changes cause sufficient stress to cause immunological suppression,
cold body temperatures that montane amphibians experience over winter
could play a synergistic role in reducing further immunological
responses to disease. Thus, such conditions might make mountain yellow-
legged frogs more susceptible to disease. Additionally, Blaustein et
al. (2001, p. 1808) have suggested that climate change could also
affect the distribution of pathogens and their vectors, exposing
amphibians to new pathogens. Climate change (warming) has been
hypothesized as a driver for the range shift of Bd (Pounds et al. 2006,
p. 161; Bosch et al. 2007, p. 253). However, other work has indicated
that survival and transmission of Bd is more likely facilitated by
cooler and wetter conditions (Corn 2005, p. 63). Fisher et al. (2009,
p. 299) present a review of information available to date and evaluate
the competing hypotheses regarding Bd dynamics, and they present some
cases that suggest a changing climate can change the host-pathogen
dynamic to a more virulent state.
The key risk factor for climate change impacts on mountain yellow-
legged frogs is likely the combined effect of reduced water levels in
high mountain lakes and ponds and the relative inability of individuals
to disperse and colonize across longer distances in order to occupy
more favorable habitat conditions (if they exist). Although such
adaptive range shifts have been observed in some plant and animal
species, they have not been reported in amphibians. The changes
observed in amphibians to date have been more associated with changes
in timing of breeding (phenology) (Corn 2005, p. 60). This limited
adaptive capacity for mountain yellow-legged frogs is a function of
high site fidelity and the extensive habitat fragmentation due to the
introduction of fishes in many of the more productive and persistent
high mountain lake habitats and streams that constitute critical
dispersal corridors throughout much of the frogs' range (see Factor C
discussion above).
An increase in the frequency, intensity, and duration of droughts
caused by climate change may have compounding effects on populations of
mountain yellow-legged frogs already in decline. In situations where
other stressors (such as introduced fish) have resulted in the
isolation of mountain yellow-legged frogs in marginal habitats,
localized mountain yellow-legged frog population crashes or
extirpations resulting from drought may exacerbate their isolation and
preclude natural recolonization (Bradford et al. 1993, p. 887; Drost
and Fellers 1996, p. 424; Lacan et al. 2008, p. 222). Viers et al.
(2013, pp. 55, 56) have used a variety of risk metrics to determine
that both mountain yellow-legged frog species in the Sierra Nevada are
highly vulnerable to climate change, and that changing hydrology and
habitat in the Sierra Nevada will likely have drastic impacts on
remaining populations. Climate change represents a substantial future
threat to the persistence of mountain yellow-legged frog populations.
Direct and Indirect Mortality
Other risk factors include direct and indirect mortality as an
unintentional consequence of activities within mountain yellow-legged
frog habitat. Mortality due to trampling by grazing livestock has been
noted in a limited number of situations, with expected mortality risk
thought to be greatest if livestock concentrate in prime breeding
habitat early in the season when adults are breeding and egg masses are
present (Brown et al. 2009, p. 59). Brown et al. (2009, p. 59) note
that standards in the SNFPA are intended to mitigate this risk.
Recreational uses also have the potential to result in direct or
indirect mortality of mountain yellow-legged frog individuals at all
life stages. The Forest Service has identified activities, including
recreational activities that occur in the frogs' breeding sites as
being risk factors for the frogs, while noting that recreation use is a
risk that USFS management can change (USDA 2001a, pp. 213-214). Brown
et al. (2009, pp. 65-66) note that tadpoles and juveniles, in
particular, may be injured or killed by trampling, crushing, etc., by
hikers, bikers, anglers, pets, packstock, or off-highway vehicles,
although the number of documented situations appears limited.
Recreational activities, such as hiking and camping, are associated
primarily with physical site
[[Page 24281]]
alteration (changes to soil and vegetation conditions), and such
effects are found to be highly localized. For example, estimates in a
heavily-used portion of the Eagle Cap Wilderness in Oregon indicated
that no more than 2 percent of the area had been altered by
recreational use (Cole and Landres 1996, p. 170). However, where
impacts of recreational use are highly localized, species impacts due
to trampling have been identified, especially for rare plant species
(Cole and Landres 1996, p. 170). Fire management activities (i.e. fuels
reduction and prescribed fire) lead to some direct mortality and have
the potential to disrupt behavior. Please refer to the proposed listing
rule for the Sierra Nevada yellow-legged frog and the northern DPS of
the mountain yellow-legged frog under the Act (16 U.S.C. 1531 et seq.)
for information about effects of fire retardants on mountain yellow-
legged frogs. Roads create the potential for direct mortality of
amphibians by vehicle strikes (deMaynadier and Hunter 2000, p. 56) and
the possible introduction of contaminants into new areas; however, most
extant populations are not located near roads. Collectively, direct
mortality risks to mountain yellow-legged frogs are likely of sporadic
significance. They may be important on occasion on a site-specific
basis, but are likely of low prevalence across the range of the
species.
Small Population Size
In many localities, remaining populations for both the Sierra
Nevada yellow-legged frog and the mountain yellow-legged frog are small
(CDFW, unpubl. data). Brown et al. (2011, p. 24) reported that about 90
percent of watersheds have fewer than 10 adults and 80 percent have
fewer than 10 subadults and 100 tadpoles. Remnant populations in the
northern portion of the range for the Sierra Nevada yellow-legged frog
(from Lake Tahoe north) and the southern portion of the populations of
the northern DPS of the mountain yellow-legged frog (south of Kings
Canyon National Park) currently also exhibit very low abundances (CDFW,
unpubl. data).
Compared to large populations, small populations are more
vulnerable to extirpation from environmental, demographic, and genetic
stochasticity (random natural occurrences), and unforeseen (natural or
unnatural) catastrophes (Shaffer 1981, p. 131).
Environmental stochasticity refers to annual variation in birth and
death rates in response to weather, disease, competition, predation, or
other factors external to the population (Shaffer 1981, p. 131). Small
populations may be less able to respond to natural environmental
changes (K[eacute]ry et al. 2000, p. 28), such as a prolonged drought
or even a significant natural predation event. Periods of prolonged
drought are more likely to have a significant effect on mountain
yellow-legged frogs because drought conditions occur on a landscape
scale and all life stages are dependent on habitat with suitable
perennial water. Demographic stochasticity is random variability in
survival or reproduction among individuals within a population (Shaffer
1981, p. 131) and could increase the risk of extirpation of the smaller
remaining populations. Genetic stochasticity results from changes in
gene frequencies due to the founder effect (loss of genetic variation
that occurs when a new population is established by a small number of
individuals) (Reiger 1968, p. 163); random fixation (the complete loss
of one of two alleles in a population, the other allele reaching a
frequency of 100 percent) (Reiger 1968, p. 371); or inbreeding
depression (loss of fitness or vigor due to mating among relatives)
(Soul[eacute] 1980, p. 96). Additionally, small populations generally
have an increased chance of genetic drift (random changes in gene
frequencies from generation to generation that can lead to a loss of
variation) and inbreeding (Ellstrand and Elam 1993, p. 225).
Allee effects (Dennis 1989, pp. 481-538) occur when a population
loses its positive stock-recruitment relationship (when population is
in decline). In a declining population, an extinction threshold or
``Allee threshold'' (Berec et al. 2006, pp. 185-191) may be crossed,
where adults in the population either cease to breed or the population
becomes so compromised that breeding does not contribute to population
growth. Allee effects typically fall into three broad categories
(Courchamp et al. 1999, pp. 405-410): lack of facilitation (including
low mate detection and loss of breeding cues), demographic
stochasticity, and loss of heterozygosity (a measure of genetic
variability). Environmental stochasticity amplifies Allee effects
(Dennis 1989, pp. 481-538; Dennis 2002, pp, 389-401). The Allee effects
of demographic stochasticity and loss of heterozygosity are likely as
mountain yellow-legged frog populations continue to diminish.
The extinction risk for a species represented by few small
populations is magnified when those populations are isolated from one
another. This is especially true for species whose populations normally
function in a metapopulation structure, whereby dispersal or migration
of individuals to new or formerly occupied areas is necessary.
Connectivity between these populations is essential to increase the
number of reproductively active individuals in a population; mitigate
the genetic, demographic, and environmental effects of small population
size; and recolonize extirpated areas. Additionally, fewer populations
by itself increases the risk of extinction.
The combination of low numbers with the other extant stressors of
disease, fish persistence, and potential for climate extremes could
have adverse consequences for the mountain yellow-legged frog complex
as populations approach the Allee threshold. Small population size is
currently a significant threat to most populations of mountain yellow-
legged frogs across the range of the species.
Cumulative Impacts of Extant Threats
Stressors may act additively or synergistically. An additive effect
would mean that an accumulation of otherwise low threat factors acting
in combination may collectively result in individual losses that are
meaningful at the population level. A synergistic effect is one where
the interaction of one or more stressors together leads to effects
greater than the sum of those individual factors combined. Further, the
cumulative effect of multiple added stressors can erode population
viability over successive generations and act as a chronic strain on
the viability of a species, resulting in a progressive loss of
populations over time. Such interactive effects from compounded
stressors thereby act synergistically to curtail the viability of frog
metapopulations and increase the risks of extinction.
It is difficult to predict the precise impact of the cumulative
threat represented by the relatively novel Bd epidemic across a
landscape already fragmented by fish stocking. The singular threat of
the Bd epidemic wave in the uninfected populations of the mountain
yellow-legged frog complex in the southern Sierra Nevada could
extirpate those populations as the pathogen spreads. A compounding
effect of disease-caused extirpation is that recolonization may never
occur because streams connecting extirpated sites to extant populations
now contain introduced fishes, which act as barriers to frog movement
within metapopulations. This situation isolates the remaining
populations of mountain yellow-legged frogs from one another (Bradford
1991, p. 176; Bradford et al. 1993, p. 887). It is logical to presume
that the small, fragmented populations left in the recent wake of Bd
spread
[[Page 24282]]
through the majority of the range of the Sierra Nevada yellow-legged
frog may experience further extirpations as surviving adults eventually
die, and recruitment into the breeding pool from the Bd-positive
subadult class is significantly reduced. These impacts may be
exacerbated by the present and growing threat of climate change,
although this effect may take years to materialize.
In summary, based on the best available scientific and commercial
information, we consider other natural and manmade factors to be
substantial ongoing threats to the Sierra Nevada yellow-legged frog and
the northern DPS of the mountain yellow-legged frog. These include
high, prevalent risk associated with climate change and small
population sizes, and the associated risk from the additive or
synergistic effects of these two stressors interacting with other
acknowledged threats, including habitat fragmentation and degradation
(see Factor A), disease and predation (see Factor C), or other threats
currently present but with low relative contribution in isolation.
Determination for the Sierra Nevada Yellow-Legged Frog
We have carefully assessed the best scientific and commercial
information available regarding the past, present, and future threats
to the Sierra Nevada yellow-legged frog. The best available information
for the Sierra Nevada yellow-legged frog shows that the geographic
extent of the species' range has declined, with local population-level
changes first noticed in the early 1900s (Grinnell and Storer 1924, p.
664) although they were still abundant at many sites in the Sierra
Nevada until the 1960s (Zweifel 1955, pp. 237-238). Population losses
continued between the 1960s and 1990s (Bradford et al. 1993, p. 883)
and have continued in recent decades. Now fewer, increasingly isolated
populations maintain viable recruitment (entry of post-metamorphic
frogs into the breeding population). Coupled with the observation that
remnant populations are also numerically smaller (in some cases
consisting of few individuals), this reduction in occupancy and
population density across the landscape suggests significant losses in
metapopulation viability and high attendant risk to the overall
population of the species. The impacts of the declines on population
resilience are two-fold: (1) The geographic extent and number of
populations are reduced across the landscape, resulting in fewer and
more isolated populations (the species is less able to withstand
population stressors and unfavorable conditions exist for genetic
exchange or dispersal to unoccupied areas (habitat fragmentation)); and
(2) species abundance (in any given population) is reduced, making
local extirpations much more likely (decreased population viability).
Knapp et al. (2007b, pp. 1-2) estimated a 10 percent decline per year
in the number of remaining mountain yellow-legged frog populations and
argued for the listing of the species as endangered based on this
observed rate of population loss.
Threats that face the Sierra Nevada mountain yellow-legged frog,
discussed above under Factors A, C, D, and E, increase the risk of the
species' extinction, given the isolation of remaining populations. The
best available science indicates that the introduction of fishes to the
frog's habitat to support recreational angling is one of the primary
causes of the decline of the Sierra Nevada yellow-legged frog and poses
a current and continuing threat to the species (Factor A). Water bodies
throughout this range have been intensively stocked with introduced
fish (principally trout). It is a threat of significant influence, and
although fewer lakes are stocked currently than were stocked prior to
2001, it remains prevalent today because fish persist in many high-
elevation habitats even where stocking has ceased. Further, the
introduction of fish has generally restricted remaining Sierra Nevada
yellow-legged frog populations to more marginal habitats, thereby
increasing the likelihood of localized extinctions. Recolonization in
these situations is difficult for a highly aquatic species with high
site fidelity and unfavorable dispersal conditions.
Historical livestock grazing activities may also have modified the
habitat of the Sierra Nevada yellow-legged frog throughout much of its
range (Factor A). Grazing pressure has been significantly reduced from
historical levels, but is expected to have legacy effects on mountain
yellow-legged frog habitat where prior downcutting and headcutting of
streams have resulted in reduced water tables and would benefit from
restoration. Current grazing that complies with forest standards and
guidelines is not expected to cause habitat-related effects to the
species in almost all cases, but in limited cases may continue to
contribute to some localized degradation and loss of suitable habitat.
The habitat-related effects of recreation, packstock grazing, dams and
water diversions, roads, timber harvests, and fire management
activities on the Sierra Nevada yellow-legged frog (Factor A) may have
contributed to historical losses when protections and use limits that
are currently afforded by USFS and NPS standards and guidelines did not
exist. Currently, Federal land management agencies with jurisdiction
within the current range of the Sierra Nevada yellow-legged frog have
developed management standards and guidelines that limit habitat damage
due to these activities, although in localized areas habitat-related
changes may continue to affect individual populations.
Competitive exclusion and predation by fish have eliminated or
reduced mountain yellow-legged frog populations in stocked habitats,
and left remnant populations isolated, while bullfrogs are expected to
have negative effects where they occur (Factor C). It is important to
recognize that, throughout the vast majority of its range, Sierra
Nevada yellow-legged frogs did not co-evolve with any species of fish,
as they predominantly occur in water bodies above natural fish
barriers. Consequently, the species has not evolved defenses against
fish predation.
Sierra Nevada yellow-legged frogs are vulnerable to multiple
pathogens (see Factor C) whose effects range from low levels of
infection within persistent populations to disease-induced extirpation
of entire populations. The Bd epidemic has caused extirpations of
Sierra Nevada yellow-legged frog populations throughout its range and
caused associated significant declines in numbers of individuals.
Though Bd was only recently discovered to affect the Sierra Nevada
yellow-legged frog, it appears to infect populations at much higher
rates than other pathogens. The imminence of this risk to populations
in currently uninfected habitats is immediate and the potential effects
severe. The already-realized effects to the survival of sensitive
amphibian life stages in Bd-positive areas are well-documented.
Although some populations survive the initial Bd wave, survival rates
of metamorphs and population viability are markedly reduced relative to
historical (pre-Bd) norms.
These threats described above are likely to be exacerbated by
widespread changes associated with climate change and by current small
population sizes in many locations (see Factor E), while instances of
direct and indirect mortality are expected to have population-level
effects only in relatively uncommon, localized situations. On a
rangewide basis, the existing regulatory mechanisms (Factor D) have not
been effective in protecting populations from declines due to fish
stocking and continuing presence of fish
[[Page 24283]]
and to disease, although standards and guidelines developed by the USFS
and the NPS have largely limited threats due to livestock and packstock
grazing, recreation, and timber use.
The main and interactive effects of these various risk factors have
acted to reduce Sierra Nevada yellow-legged frog populations to small
fractions of their historical habitat and reduce population abundances
significantly throughout most of its current range. Remaining areas
that have yet to be impacted by Bd are at immediate and severe risk.
Given the life history of this species, dispersal, recolonization,
and genetic exchange are largely precluded by the fragmentation of
habitat common throughout its current range as a result of fish
introductions. Frogs that may disperse are susceptible to hostile
conditions in many circumstances. In essence, Sierra Nevada yellow-
legged frogs have been marginalized by historical fish introductions.
Populations have recently been decimated by Bd, and the cumulative
effect of other stressors (such as anticipated reduction of required
aquatic breeding habitats with climate change and more extreme weather)
upon a fragmented landscape make adaptation and recovery a highly
improbable scenario without active intervention. The cumulative risk
from these stressors to the persistence of the Sierra Nevada yellow-
legged frog throughout its range is significant.
The Act defines an endangered species as any species that is ``in
danger of extinction throughout all or a significant portion of its
range'' and a threatened species as any species ``that is likely to
become endangered throughout all or a significant portion of its range
within the foreseeable future.'' We find that the Sierra Nevada yellow-
legged frog is presently in danger of extinction throughout its entire
range, based on the immediacy, severity, and scope of the threats
described above. Specifically, these include habitat degradation and
fragmentation under Factor A, predation and disease under Factor C, and
climate change and the interaction of these various stressors
cumulatively impacting small remnant populations under Factor E. There
has been a rangewide reduction in abundance and geographic extent of
surviving populations of the Sierra Nevada yellow-legged frog following
decades of fish stocking, habitat fragmentation, and, most recently, a
disease epidemic. Surviving populations are smaller and more isolated,
and recruitment in Bd-positive populations is much reduced relative to
historical norms. This combination of population stressors makes
species persistence precarious throughout the current range in the
Sierra Nevada.
We have carefully assessed the best scientific and commercial
information available regarding the past, present, and future threats
to the species, and have determined that the Sierra Nevada yellow-
legged frog meets the definition of endangered under the Act, rather
than threatened. This is because significant threats are occurring now
and will occur in the future, at a high magnitude and across the
species' entire range, making the species in danger of extinction at
the present time. The rate of population decline remains high in the
wake of Bd epidemics, and the remaining Sierra Nevada yellow-legged
frog populations are at high, imminent risk. Population declines are
expected to continue as maturing tadpoles succumb to Bd infection, and
fragmented populations at very low abundances will face significant
obstacles to recovery. Therefore, on the basis of the best available
scientific and commercial information, and the threats posed to these
species under the listing factors above, we are listing the Sierra
Nevada yellow-legged frog as endangered in accordance with sections
3(6) and 4(a)(1) of the Act.
Under the Act and our implementing regulations, a species may
warrant listing if it is endangered or threatened throughout all or a
significant portion of its range. The Sierra Nevada yellow-legged frog
is restricted in its range, and the threats occur throughout the
remaining occupied habitat. Therefore, we assessed the status of this
species throughout its entire range. The threats to the survival of the
species occur throughout the species' range and are not restricted to
any particular significant portion of that range. Accordingly, our
assessment and final determination applies to the species throughout
its entire range.
Final Determination for the Northern DPS of the Mountain Yellow-Legged
Frog
We have carefully assessed the best scientific and commercial
information available regarding the past, present, and future threats
to the northern DPS of the mountain yellow-legged frog. The best
available information for the northern DPS of the mountain yellow-
legged frog shows that the geographic extent of the species' range has
declined, with local population-level changes first noticed in the
early 1900s (Grinnell and Storer 1924, p. 664), although they were
still abundant at many sites in the Sierra Nevada until the 1960s
(Zweifel 1955, pp. 237-238). Population losses continued between the
1960s and 1990s (Bradford et al. 1993, p. 883) and have continued in
recent decades. Now fewer, increasingly isolated populations maintain
viable recruitment (entry of post-metamorphic frogs into the breeding
population). Coupled with the observation that remnant populations are
also numerically smaller (in some cases consisting of a few
individuals), this reduction in occupancy and population density across
the landscape suggests significant losses in metapopulation viability
and high attendant risk to the overall population of the species. The
impacts of the declines on population resilience are two-fold: (1) The
geographic extent and number of populations are reduced across the
landscape, resulting in fewer and more isolated populations (the
species is less able to withstand population stressors and unfavorable
conditions exist for genetic exchange or dispersal to unoccupied areas
(habitat fragmentation)); and (2) species abundance (in any given
population) is reduced, making local extirpations much more likely
(decreased population viability). Knapp et al. (2007b, pp. 1-2)
estimated a 10 percent decline per year in the number of remaining
mountain yellow-legged frog populations and argued for the listing of
the species as endangered based on this observed rate of population
loss.
Threats that face the northern DPS of the mountain yellow-legged
frog, discussed above under Factors A, C, D, and E, increase the risk
of the species' extinction, given the isolation of remaining
populations. The best available science indicates that the introduction
of fishes to the frog's habitat to support recreational angling is one
of the primary causes of the decline of the northern DPS of the
mountain yellow-legged frog and poses a current and continuing threat
to the species (Factor A). Water bodies throughout this range have been
intensively stocked with introduced fish (principally trout). It is a
threat of significant influence, and although fewer lakes are stocked
currently than were stocked prior to 2001, it remains prevalent today
because fish persist in many high-elevation habitats even where
stocking has ceased. Recolonization in these situations is difficult
for a highly aquatic species with high site fidelity and unfavorable
dispersal conditions. Climate change is likely to exacerbate these
other threats and further threaten population resilience.
Historical livestock grazing activities may also have modified the
habitat of the northern DPS of the mountain
[[Page 24284]]
yellow-legged frog throughout much of its range (Factor A). Grazing
pressure has been significantly reduced from historical levels, but is
expected to have legacy effects to mountain yellow-legged frog habitat
where prior downcutting and headcutting of streams have resulted in
reduced water tables that still need restoration to correct. Current
grazing that complies with forest standards and guidelines is not
expected to cause habitat-related effects to the species in almost all
cases, but in limited cases may continue to contribute to some
localized degradation and loss of suitable habitat. The habitat-related
effects of recreation, packstock grazing, dams and water diversions,
roads, timber harvests, and fire management activities on the northern
DPS of the mountain yellow-legged frog (Factor A) may have contributed
to historical losses when protections and use limits that are currently
afforded by USFS and NPS standards and guidelines did not exist.
Currently, Federal agencies with jurisdiction within the current range
of the northern DPS of the mountain yellow-legged frog have developed
management standards and guidelines that limit habitat damage due to
these activities, although in localized areas habitat-related changes
may continue to affect individual populations.
Competitive exclusion and predation by fish have eliminated or
reduced mountain yellow-legged frog populations in stocked habitats,
and left remnant populations isolated, while bullfrogs are expected to
have negative effects where they occur (Factor C). It is important to
recognize that throughout the vast majority of its range, the northern
DPS of the mountain yellow-legged frogs did not co-evolve with any
species of fish, as this species predominantly occurs in water bodies
above natural fish barriers. Consequently, the species has not evolved
defenses against fish predation.
Mountain yellow-legged frogs are vulnerable to multiple pathogens
(see Factor C) whose effects range from low levels of infection within
persistent populations to disease-induced extirpation of entire
populations. The Bd epidemic has caused rangewide extirpations of
populations of the northern DPS of the mountain yellow-legged frog and
associated significant declines in numbers of individuals. Though Bd
was only recently discovered to affect the mountain yellow-legged frog,
it appears to infect populations at much higher rates than other
pathogens. The imminence of this risk to currently uninfected habitats
is immediate, and the potential effects severe. The already-realized
effects to the survival of sensitive amphibian life stages in Bd-
positive areas are well-documented. Although some populations survive
the initial Bd wave, survival rates of metamorphs and population
viability are markedly reduced relative to historical (pre-Bd) norms.
These threats are likely to be exacerbated by widespread changes
associated with climate change and by current small population sizes in
many locations (see Factor E), while instances of direct and indirect
mortality are expected to have population-level effects only in
relatively uncommon, localized situations. Rangewide, the existing
regulatory mechanisms (Factor D) have not been effective in protecting
populations from declines due to fish stocking and continuing presence
of fish and to disease, although standards and guidelines developed by
the USFS and the NPS have largely limited threats due to livestock and
packstock grazing, recreation, and timber use.
The main and interactive effects of these various risk factors have
acted to reduce the northern DPS of the mountain yellow-legged frog to
a small fraction of its historical range and reduce population
abundances significantly throughout most of its current range.
Populations of this species in remaining areas in the southern Sierra
Nevada that have yet to be impacted by Bd are at immediate and severe
risk.
Given the life history of this species, dispersal, recolonization,
and genetic exchange are largely precluded by the fragmentation of
habitat common throughout its current range as a result of fish
introductions. Frogs that may disperse are susceptible to hostile
conditions in many circumstances. In essence, mountain yellow-legged
frogs have been marginalized by historical fish introductions.
Populations have recently been decimated by Bd, and the accumulation of
other stressors (such as anticipated reduction of required aquatic
breeding habitats with climate change and more extreme weather) upon a
fragmented landscape make adaptation and recovery a highly improbable
scenario without active intervention. The cumulative risk from these
stressors to the persistence of the mountain yellow-legged frog
throughout its range is significant.
The Act defines an endangered species as any species that is ``in
danger of extinction throughout all or a significant portion of its
range'' and a threatened species as any species ``that is likely to
become endangered throughout all or a significant portion of its range
within the foreseeable future.'' We find that the northern DPS of the
mountain yellow-legged frog is presently in danger of extinction
throughout its entire range, based on the immediacy, severity, and
scope of the threats described above. Specifically, these include
habitat degradation and fragmentation under Factor A, predation and
disease under Factor C, and climate change and the interaction of these
various stressors cumulatively impacting small remnant populations
under Factor E. There has been a rangewide reduction in abundance and
geographic extent of surviving populations of the northern DPS of the
mountain yellow-legged frog following decades of fish stocking, habitat
fragmentation, and, most recently, a disease epidemic. Surviving
populations are smaller and more isolated, and recruitment in Bd-
positive populations is much reduced relative to historical norms. This
combination of population stressors makes species persistence
precarious throughout the current range in the Sierra Nevada.
We have carefully assessed the best scientific and commercial
information available regarding the past, present, and future threats
to the species, and have determined that the northern DPS of the
mountain yellow-legged frog, meets the definition of endangered under
the Act, rather than threatened. This is because significant threats
are occurring now and will occur in the future, at a high magnitude and
across the DPS' entire range, making the northern DPS of the mountain
yellow-legged frog in danger of extinction at the present time. The
rate of population decline remains high in the wake of Bd epidemics,
and northern DPS of the mountain yellow-legged frog areas are at high,
imminent risk. The recent rates of decline for these populations are
even higher than declines in the populations of the Sierra Nevada
yellow-legged frog, and as Bd infects remaining core areas, population
viability will be significantly reduced, and extirpations or
significant population declines are expected. Population declines are
expected to continue as maturing tadpoles succumb to Bd infection, and
fragmented populations at very low abundances will face significant
obstacles to recovery. Therefore, on the basis of the best available
scientific and commercial information, and the threats posed to these
species discussed under the listing factors above, we are listing the
northern DPS of the mountain yellow-legged frog as endangered in
accordance with sections 3(6) and 4(a)(1) of the Act.
Under the Act and our implementing regulations, a species may
warrant
[[Page 24285]]
listing if it is endangered or threatened throughout all or a
significant portion of its range. The northern DPS of the mountain
yellow-legged frog addressed in this final listing rule is restricted
in its range, and the threats occur throughout the remaining occupied
habitat. Therefore, we assessed the status of this DPS throughout its
entire range in the Sierra Nevada of California. The threats to the
survival of this DPS occur throughout its range in the southern Sierra
Nevada and are not restricted to any particular significant portion of
that range. Accordingly, our assessment and final determination applies
to the DPS throughout its entire range.
Summary of Biological Status and Threats Affecting the Yosemite Toad
Background
Taxonomy and Species Description
Please refer to the proposed listing rule for the Yosemite toad
under the Act (16 U.S.C. 1531 et seq.) for additional species
information, including detailed information on taxonomy. In this
section of the final rule, it is our intent to discuss only those
topics directly relevant to the listing of the Yosemite toad (Anaxyrus
canorus) as threatened.
Habitat and Life History
Breeding habitat--Yosemite toads are associated with wet meadows
due to their breeding ecology. Camp (1916, pp. 59-62) found Yosemite
toads in wet meadow habitats and at lake shores located among lodgepole
(Pinus contorta) at the lower elevations to whitebark (P. albicaulis)
pines at the higher elevations. Mullally (1953, pp. 182-183) found
adult toads common on the margins of high-elevation lakes, streams, and
pools wherever the meadow vegetation was thicker or more luxuriant than
usual or where there were patches of low willows (Salix spp.). Liang
(2010, p. 81) observed Yosemite toads most frequently associated with
(in order of preference): wet meadows, alpine-dwarf scrub, red fir
(Abies magnifica), water, lodgepole pine, and subalpine conifer
habitats.
Yosemite toads were found as often at large as at small sites
(Liang 2010, p. 19), suggesting that this species is capable of
successfully utilizing small habitat patches. Liang also found that
population persistence was greater at higher elevations, with an
affinity for relatively flat sites with a southwesterly aspect (Liang
2010, p. 20; see also Mullally 1953, p. 182). These areas receive
higher solar radiation and are capable of sustaining hydric (wet),
seasonally ponded, and mesic (moist) breeding and rearing habitat. The
Yosemite toad is more common in areas with less variation in mean
annual temperature, or more temperate sites with less climate variation
(Liang 2010, pp. 21-22).
Adults are thought to be long-lived, and this factor allows for
persistence in variable conditions and more marginal habitats where
only periodic good years allow high reproductive success (USFS et al.
2009, p. 27). Females have been documented to reach 15 years of age,
and males as many as 12 years (Kagarise Sherman and Morton 1993, p.
195); however, the average longevity of the Yosemite toad in the wild
is not known. Jennings and Hayes (1994, p. 52) indicated that females
begin breeding at ages 4 to 6 years, while males begin breeding at ages
3 to 5 years.
Adults appear to have high site-fidelity; Liang (2010, pp. 99, 100)
found that the majority of individuals identified in multiple years
were located in the same meadow pools, although individuals will move
between breeding areas (Liang 2010, p. 52; Liang 2013, p. 561).
Breeding habitat includes shallow, warm-water areas in wet meadows,
such as shallow ponds and flooded vegetation, ponds, lake edges, and
slow-flowing streams (Karlstrom 1962, pp. 8-12; Brown 2013,
unpaginated). Tadpoles have also been observed in shallow areas of
lakes (Mullally 1953, pp. 182-183).
Adult Yosemite toads are most often observed near water, but only
occasionally in water (Mullally and Cunningham 1956b, pp. 57-67). Moist
upland areas such as seeps and springheads are important summer
nonbreeding habitats for adult toads (Martin 2002, pp. 1-3). The
majority of their life is spent in the upland habitats proximate to
their breeding meadows. They use rodent burrows for overwintering and
probably for temporary refuge during the summer (Jennings and Hayes
1994, pp. 50-53), and they spend most of their time in burrows (Liang
2010, p. 95). They also use spaces under surface objects, including
logs and rocks, for temporary refuge (Stebbins 1951, pp. 245-248;
Karlstrom 1962, pp. 9-10). Males and females also likely inhabit
different areas and habitats when not breeding, and females tend to
move farther from breeding ponds than males (USFS et al. 2009, p. 28).
Males exit burrows first, and spend more time in breeding pools
than females, who do not breed every year (Kagarise Sherman and Morton,
1993, p. 196). Data suggest that higher lipid storage in females, which
enhances overwinter survival, also precludes the energetic expense of
breeding every year (Morton 1981, p. 237). The Yosemite toad is a
prolific breeder, laying many eggs immediately at snowmelt. This is
accomplished in a short period of time, coinciding with water levels in
meadow habitats and ephemeral pools they use for breeding. Female toads
lay approximately 700-2,000 eggs in two strings (one from each ovary)
(USFS et al. 2009, p. 21). Females may split their egg clutches within
the same pool, or even between different pools, and may lay eggs
communally with other toads (USFS et al. 2009, p. 22).
Eggs hatch within 3-15 days, depending on ambient water
temperatures (Kagarise Sherman 1980, pp. 46-47; Jennings and Hayes
1994, p. 52). Tadpoles typically metamorphose around 40-50 days after
fertilization, and are not known to overwinter (Jennings and Hayes
1994. p. 52). Tadpoles are black in color, tend to congregate together
(Brattstrom 1962, pp. 38-46) in warm shallow waters during the day
(Cunningham 1963, pp. 60-61), and then retreat to deeper waters at
night (Mullaly 1953, p. 182). Rearing through metamorphosis takes
approximately 5-7 weeks after eggs are laid (USFS et al. 2009, p. 25).
Toads need shallow, warm surface water that persists through the period
during which they metamorphose; shorter hydroperiods in that habitat
can reduce reproductive success (Brown 2013, unpaginated).
Reproductive success is dependent on the persistence of tadpole
rearing sites and conditions for breeding, egg deposition, hatching,
and rearing to metamorphosis (USFS et al. 2009, p. 23). Given their
association with shallow, ephemeral habitats, Yosemite toads are
susceptible to droughts and weather extremes. Abiotic factors leading
to mortality (such as freezing or desiccation) appear to be more
significant during the early life stages of toads, while biotic factors
(such as predation) are probably more prominent factors during later
life stages (USFS et al. 2009, p. 30). However, since adult toads lead
a much more inconspicuous lifestyle, direct observation of adult
mortality is difficult and it is usually not possible to determine
causes of adult mortality.
Yosemite toads can move farther than 1 km (0.63 mi) from their
breeding meadows (average movement is 275 m (902 ft)), and they utilize
terrestrial environments extensively (Liang 2010, p. 85). The average
distance traveled by females is twice as far as males, and home ranges
for females are 1.5 times greater than those for males (Liang 2010, p.
94). Movement into the upland
[[Page 24286]]
terrestrial environment following breeding does not follow a
predictable path, and toads tend to traverse longer distances at night,
perhaps to minimize evaporative water loss (Liang 2010, p. 98). Martin
(2008, p. 123) tracked adult toads during the active season and found
that on average toads traveled a total linear distance of 494 m (1,620
ft) within the season, with minimum travel distance of 78 m (256 ft)
and maximum of 1.76 km (1.09 mi).
Historical Range and Distribution
The known historical range of the Yosemite toad in the Sierra
Nevada extended from the Blue Lakes region north of Ebbetts Pass
(Alpine County) to south of the Evolution Lake area (Fresno County)
(Karlstrom 1962, p. 3; Stebbins 1985, p. 72; see also Knapp 2013,
unpaginated; Brown 2013, unpaginated). Yosemite toad habitat
historically spanned elevations from 1,460 to 3,630 m (4,790 to 11,910
ft) (Stebbins 1985, p. 72; Stephens 2001, p. 12).
Current Range and Distribution
The current range of the Yosemite toad, at least in terms of
overall geographic extent, remains largely similar to the historical
range defined above (USFS et al. 2009, p. 41). However, within that
range, toad habitats have been degraded and may be decreasing in area
as a result of conifer encroachment and historical livestock grazing
(see Factor A below). The vast majority of the Yosemite toad's range is
within federally managed land. Figure 2, Estimated Range of Yosemite
Toad, displays a range map for the species.
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Population Estimates and Status
Baseline data on the number and size of historical Yosemite toad
populations are limited, and historic records are largely based on
accounts from field notes, or pieced together through museum
collections, thereby providing limited information on historical
populations. Systematic survey information across the range of the
species on National Forest System Lands largely follows the designation
of the Yosemite toad as a candidate species under the Act. In addition,
surveys for the Yosemite toad have been conducted within Yosemite,
Kings Canyon, and Sequoia National Parks (Knapp 2013, unpaginated).
From these recent inventories, Yosemite toads have been found at 469
localities collectively on six National Forests (USFS et al. 2009, p.
40; see also Brown and Olsen 2013, pp. 675-691), at 179 breeding sites
that were surveyed between 1992 and 2010 in Yosemite National Park
(Berlow et al. 2013, p. 3), and detected at 18 localities in Kings
Canyon National Park (NPS 2011, geospatial data). Although we did not
cite to the information from the National Parks in the proposed rule,
we had the geospatial occupancy data that is currently included in
Berlow et al. 2013, and we utilized that data in our analysis for the
proposed listing (see comments 6 and 7 below, and their respective
responses). The number of localities identified in these surveys
reflects more occupied sites than were known before such extensive
surveys were conducted, and indicates that the species is still
widespread throughout its range. These inventories were typically
conducted to determine toad presence or absence (they were not
censuses), and do not explicitly compare historic sites to recent
surveys. Moreover, single-visit surveys of toads are unreliable as
indices of abundance because timing is so critical to the presence of
detectable life stages and not all potential breeding habitats within
the range of the species were surveyed (USFS et al. 2009, p. 41; Liang
2010, p. 10; Brown and Olsen 2013, p. 685). Given these considerations,
conclusions about population trends, abundance, or extirpation rates
are not possible from these datasets overall.
One pair of studies allows us to compare current distribution with
historic distributions and indicates that large reductions have
occurred. In 1915 and 1919, Grinnell and Storer (1924, pp. 657-660)
surveyed for vertebrates at 40 sites along a 143-km (89-mi) west-to-
east transect across the Sierra Nevada, through Yosemite National Park,
and found Yosemite toads at 13 of those sites. In 1992, Drost and
Fellers (1996, pp. 414-425) conducted more thorough surveys,
specifically for amphibians, at 38 of the Grinnell and Storer sites
plus additional nearby sites. Drost and Fellers (1996, pp. 418) found
that Yosemite toads were absent from 6 of 13 sites where they had been
found in the original Grinnell and Storer (1924) survey. Moreover, at
the sites where they were present, Yosemite toads most often occurred
in very low numbers relative to general abundance reported in the
historical record (Grinnell and Storer 1924, pp. 657-660). Therefore,
by the early 1990s, the species was either undetectable or had declined
in numbers at 9 of 13 (69 percent) of the Grinnell and Storer sites
(Drost and Fellers 1996, p. 418).
Another study comparing historic and current occurrences also found
a large decline in Yosemite toad distribution. In 1990, David Martin
surveyed 75 sites throughout the range of the Yosemite toad for which
there were historical records of the species' presence. This study
found that 47 percent of historically occupied sites showed no evidence
of any life stage of the species (Stebbins and Cohen 1995, pp. 213-
215). This result suggests a range-wide decline to about one half of
historical sites, based on occupancy alone.
A third study comparing historic and recent surveys indicates
declines in Yosemite toad distribution. Jennings and Hayes (1994, pp.
50-53) reviewed the current status of Yosemite toads using museum
records of historic and recent sightings, published data, and
unpublished data and field notes from biologists working with the
species. They estimated a loss of over 50 percent of former Yosemite
toad locations throughout the range of the species (based on 144
specific sites).
The only long-term, site-specific population study for Yosemite
toads documented a dramatic decline over 2 decades of monitoring.
Kagarise Sherman and Morton (1993, pp. 186-198) studied Yosemite toads
at Tioga Pass Meadow (Mono County, California) from 1971 through 1991
(with the most intensive monitoring through 1982). They documented a
decline in the average number of males entering the breeding pools from
258 to 28 during the mid-1970s through 1982. During the same time
period, the number of females varied between 45 and 100, but there was
no apparent trend in number observed. During the 1980s, it appeared
that males continued to decline, females also declined, and breeding
activity became sporadic. By 1991, they found only one male and two egg
masses. Sadinski (2004, p. 40) revisited the survey locations annually
from 1995 and 2001 and found a maximum of two males and two egg masses,
suggesting the toads in Tioga Meadows had not recovered from their
decline. In the study of Yosemite toads at nearby Dana Meadows,
Sadinski (2004, pp. 39-42) documented few adults within the habitats
surveyed, finding substantial mortality in embryos that he associated
with effects of ice, water mold, and flatworms. Sadinski (2004, pp. 38-
42) also found high larval mortality when breeding sites dried before
larvae could reach metamorphosis. Sadinski (2004) stated that the
proximity of the Kagarise Sherman and Morton (1993) study sites at
Tioga Meadows and his sites in Dana Meadows practically ensured that
animals from both sites were part of the same metapopulation. Sadinski
surmised that perhaps much of that metapopulation experienced events at
breeding sites similar to those that Kagarise Sherman and Morton (1993)
observed (Sadinski 2004, pp. 39-40). He further opined that, if each of
his substantial sites had previously supported hundreds of breeding
adults in the 1970s, the overall population of Yosemite toads had
declined dramatically throughout the area since that time.
Kagarise Sherman and Morton (1993, pp. 186-198) also conducted
occasional surveys of six other populations in the eastern Sierra
Nevada. Five of these populations showed long-term declines that were
evident beginning between 1978 through 1981, while the sixth population
held relatively steady until the final survey in 1990, at which time it
dropped. In 1991, E.L. Karlstrom revisited the site where he had
studied a breeding population of Yosemite toads from 1954 to 1958 (just
south of Tioga Pass Meadow within Yosemite National Park), and found no
evidence of toads or signs of breeding (Kagarise Sherman and Morton
1993, p. 190).
The most reliable information about Yosemite toad population status
and trends is the USFS SNAMPH. This study, conducted on National Forest
System Lands, is designed to provide statistical comparisons across 5-
year monitoring cycles with 134 watersheds (Brown et al. 2011, pp. 3-
4). This approach allows researchers to assess trends for the entire
range of the toad, rather than at limited survey sites (C. Brown 2012,
pers. comm., see also Brown and Olsen 2013). The results of this
assessment indicate the species has declined from historical levels,
with Yosemite toads occurring in approximately 13 percent of watersheds
where they existed prior to 1990. This
[[Page 24288]]
study also found that breeding was occurring in approximately 84
percent of the watersheds that were occupied in the period 1990-2001,
suggesting that the number of locations where breeding occurs has
continued to decline. Additionally, the study found that breeding
currently occurs in an estimated 22 percent of watersheds within the
current estimated range of the species (Brown et al. 2012, p. 115).
Moreover, overall abundances in the intensively monitored
watersheds were very low (fewer than 20 males per meadow per year)
relative to other historically reported abundances of the species
(Brown et al. 2011, p. 4). Brown et al. (2011, p. 35) suggest that
populations are now very small across the range of the species. During
their monitoring over the past decade, they found only 18 percent of
occupied survey watersheds range-wide had ``large'' populations (more
than 1,000 tadpoles or 100 of any other lifestage detected at the time
of survey). While not all surveys were conducted at the peak of tadpole
presence and adults are not reliably found outside of the breeding
season, Brown et al. (2012) surveyed many sites at appropriate times
and rarely found the large numbers of tadpoles or metamorphs that would
be expected if population sizes were similar to those reported
historically. The researchers interpret these data, in combination with
documented local population declines from other studies (see above), to
support the hypothesis that population declines have occurred range-
wide (Brown et al. 2012, p. 11).
Summary of Changes From the Proposed Rule for the Yosemite Toad
Based on peer review and Federal, State, and public comments (see
comments in the Summary of Comments and Recommendations section,
below), we clarified information for the Yosemite toad to better
characterize our knowledge of the species' habitat requirements.
Specifically, we reorganized and clarified the habitat details (Habitat
and Life History), southern extent of the species' range (Historic
Range and Distribution), and species surveys (USFS and NPS). We also
added information on occupancy in National Parks that was inadvertently
omitted from the proposed rule (Population Estimates and Status).
In the Summary of Factors Affecting the Species section, under
Factor A, we made small changes to the discussion about meadow loss and
degradation in order to improve clarity. In the Livestock Use (Grazing)
Effects to Meadow Habitat section, we reorganized the information and
separated the effects of historic livestock grazing from the effects
due to current grazing levels, and we added additional references
received from the USFS. In the Roads and Timber Harvest Effects to
Meadow Habitat section, we clarified the extent to which these
activities overlap with the Yosemite toad's range and distinguished the
effects of past activities from the effects of current activities. We
added information on road locations and on USFS Forest standards and
guidelines that currently limit the effects of these activities on
riparian areas. In this final rule, we found that roads and timber
harvest activities are not current and ongoing threats to the species.
However, there may be localized effects where legacy effects of past
road building or timber harvest continue to modify wet meadows or where
activities occur in close proximity to extant Yosemite toad
populations.
In the Fire Management section, we added information to clarify
that Yosemite toads primarily occur in higher elevation areas where
fire suppression activities are rarely conducted. This finding suggests
that fire suppression has had little effect on forest encroachment into
meadow habitats in most areas where the species occurs. In the
Recreation and Packstock Effects to Meadow Habitat section, we added
additional information on USFS and NPS restoration activities to
protect meadows, off-highway vehicle effects, packstock use, and agency
monitoring and protection activities to limit effects due to packstock
use. We revised our conclusion to clarify that, in general, we do not
consider habitat-related changes associated with current levels of
hiking, backpacking, or packstock use to pose a risk to Yosemite toad
populations. Recreation may have habitat-related effects to toads in
localized areas where use adjacent to occupied meadows is exceptionally
heavy, or where heavy or motorized use results in changes to meadow
hydrology. Accordingly, rangewide, recreation is a threat of low
prevalence. In the section on Dams and Water Diversions, we added
information to clarify that almost all reservoirs are located below the
range of the Yosemite toad. We include small changes in the Climate
Change section to improve clarity or add information from references
provided during peer review.
In Factor B, we added information provided during the comment
period, which documented the sale of one Yosemite toad from a pet store
in Southern California (store now closed). We also added information on
protections provided by agency-required research permits. In Factor C,
based on peer review comments, we added information on a Bd study on
Yosemite toads. We removed the discussion of contaminants under Factor
E, and we refer readers to the proposed rule affirming that the best
available information indicates that contaminants do not pose a current
or continuing threat to the Yosemite toad. We also added new
information in the Other Sources of Direct and Indirect Mortality
section as a result of information provided during peer review.
Although we have not changed the determination, we have made a few
small changes in the wording of the determination for the Yosemite toad
to reflect the above changes.
Summary of Factors Affecting the Species
Section 4 of the Act (16 U.S.C. 1533), and its implementing
regulations at 50 CFR part 424, set forth the procedures for adding
species to the Federal Lists of Endangered and Threatened Wildlife and
Plants. Under section 4(a)(1) of the Act, we may list a species 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; and (E) other natural or manmade
factors affecting its continued existence. Listing actions may be
warranted based on any of the above threat factors, singly or in
combination. Each of these factors is discussed below, and changes from
the proposed rule (78 FR 24472, April 25, 2013) are reflected in these
discussions.
Factor A. The Present or Threatened Destruction, Modification, or
Curtailment of Its Habitat or Range
The habitat comprising the current range of the Yosemite toad is
generally characterized by low levels of physical disturbance (there is
little to no current development pressure). However, these areas are
also generally more sensitive to perturbation and take longer to
recover from disturbances due to reduced growing seasons and harsher
environmental conditions. Since Yosemite toads rely heavily on shallow,
ephemeral water, they may be more sensitive to minor changes in their
habitat. Loss or alteration of suitable breeding habitat can reduce
reproductive success, which may have a profound impact when population
numbers are small. Past management and development activity has played
a
[[Page 24289]]
role in the degradation of meadow habitats within the Sierra Nevada.
Human activities within these habitats include grazing, timber harvest,
fuels management, recreation, and water development.
Meadow Habitat Loss and Degradation
Some of the habitat effects associated with grazing activities that
were described for the mountain yellow-legged frogs (see the Summary of
Factors Affecting the Species section for those species, above) also
apply to Yosemite toads. However, there are differences based on the
Yosemite toad's reliance on very shallow, ephemeral water in meadow and
pool habitats versus the deeper lakes and streams frequented by
mountain yellow-legged frogs. Because Yosemite toads rely on very
shallow, ephemeral water, they may be sensitive to even minor changes
in their habitat, particularly to hydrology (Brown 2013, unpaginated).
Meadow habitat quality in the Western United States, and specifically
the Sierra Nevada, has been degraded by past activities, such as
overgrazing, tree encroachment, fire suppression, and road building,
over the last century (Stillwater Sciences 2008, pp. 1-53; Halpern et
al. 2010, pp. 717-732; Vale 1987, pp. 1-18; Ratliff 1985, pp. i-48).
These past activities have contributed to erosion and stream incision
in areas of the Sierra Nevada, leading to meadow dewatering and
encroachment by invasive vegetation (Menke et al. 1996, pp. 25-28;
Lindquist and Wilcox 2000, p. 2).
Given the reliance of the Yosemite toad on these meadow and pool
habitats for breeding, rearing, and adult survival, it is logical to
conclude that the various stressors have had an indirect effect on the
viability of Yosemite toad populations via degradation of their
habitat. Loss of connectivity of habitats leads to further isolation
and population fragmentation. Because of physiological constraints, the
tendency to move only short distances, and high site fidelity,
amphibians may be unable to recolonize unoccupied sites following local
extinctions if the distance between sites is too great, although
recolonization can occur over time (Blaustein et al. 1994a, p. 8).
Since the existence of meadows is largely dependent on their
hydrologic setting, most meadow degradation is due fundamentally to
hydrologic alterations (Stillwater Sciences 2008, p. 13). There are
many drivers of hydrologic alterations in meadow ecosystems. In some
locations, historic water development and ongoing water management
activities have physically changed the underlying hydrologic system.
Diversion and irrigation ditches formed a vast network that altered
local and regional stream hydrology, although these manmade systems are
generally below the range of the Yosemite toad. Timber harvest and
associated road construction further altered erosion and sediment
delivery patterns in rivers and meadow streams. Fire suppression and an
increase in the frequency of large wildfires due to excessive fuel
buildup have introduced additional disturbance pressures to the meadows
of the Sierra Nevada (Stillwater Sciences 2008, p. 13). Many meadows
now have downcut stream courses, compacted soils, altered plant
community compositions, and diminished wildlife and aquatic habitats
(SNEP 1996, pp. 120-121).
Land uses causing channel erosion are a threat to Sierra Nevada
meadows. These threats include erosive activities within the watershed
upslope of the meadow, along with impacts from land use directly in the
meadows themselves. Compaction of meadow soils by roads or intensive
trampling (for example, overgrazing) can reduce infiltration,
accelerate surface run-off, and thereby lead to channel incision (Menke
et al. 1996, pp. 25-28). Mining, overgrazing, timber harvesting, and
railroad and road construction and maintenance have contributed to
watershed degradation, resulting in accelerated erosion, sedimentation
in streams and reservoirs, meadow dewatering, and degraded terrestrial
and aquatic habitats (Linquist 2000, p. 2). Deep incision has been
documented in several meadows in the Sierra Nevada. One example is
Halstead Meadow in Sequoia National Park, where headcutting exceeds 10
feet in many areas and is resulting in widening channels, erosion in
additional meadows, and a lowered water table (Cooper and Wolf 2006, p.
1).
The hydrologic effects of stream incision on the groundwater system
may significantly impact groundwater storage, affecting late summer
soil moisture and facilitating vegetation change (Bergmann 2004, pp.
24-31). For example, in the northern Sierra Nevada, logging,
overgrazing, and road/railroad construction have caused stream
incision, resulting in dewatering of riparian meadow sediments and a
succession from native wet meadow vegetation to sagebrush and dryland
grasses (Loheide and Gorelick 2007, p. 2). A woody shrub (Artemisia
rothrockii) is invading meadows as channel incision causes shallow-
water-dependent herbs to die back, allowing shrub seedlings to
establish in disturbed areas during wet years (Darrouzet-Nardi et al.
2006, p. 31).
Mountain meadows in the western United States and Sierra Nevada
have also been progressively colonized by trees (Thompson 2007, p. 3;
Vale 1987, p. 6), with an apparent pattern of encroachment during two
distinct periods in the late 1800s and mid-1900s (Halpern et al. 2010,
p. 717). This trend has been attributed to a number of factors,
including climate, changes in fire regime, and cessation of sheep
grazing (Halpern et al. 2010, pp. 717-718; Vale 1987, pp. 10-13), but
analyses are limited to correlational comparisons and research results
are mixed, so the fundamental contribution of each potential driver
remains uncertain. We discuss the contribution of these factors to
habitat loss and degradation for the Yosemite toad below.
Livestock Use (Grazing) Effects to Meadow Habitat
The combined effect of legacy conditions from historically
excessive grazing use and current livestock grazing activities have the
potential to impact habitat in the range of the Yosemite toad. The
following subsections discuss the effects of excessive historical
grazing, current extent of grazing, and current grazing management
practices.
Overgrazing has been associated with accelerated erosion and
gullying of meadows (Kattelmann and Embury 1996, pp. 13, 18), which
leads to siltation and more rapid succession of meadows. Grazing can
cause erosion by disturbing the ground, damaging and reducing
vegetative cover, and destroying peat layers in meadows, which lowers
the groundwater table and summer flows (Armour et al. 1994, pp. 9-12;
Martin 2002, pp. 1-3; Kauffman and Krueger 1984, pp. 431-434). Downcut
channels, no longer connected to the historic, wide floodplains of the
meadow, instead are confined within narrow, incised channels.
Downstream, formerly perennial (year-round) streams often become
intermittent or dry due to loss of water storage capacity in the meadow
aquifers that formerly sustained them (Lindquist et al. 1997, pp. 7-8).
Heavy grazing can alter vegetative species composition and
contribute to lodgepole pine (Pinus contorta) invasion (Ratliff 1985,
pp. 33-36). Lowering of the water table facilitates encroachment of
conifers into meadows. Gully formation and lowering of water tables,
changes in the composition of herbaceous vegetation, increases in the
density of forested stands, and the expansion of trees into areas that
formerly were treeless have been documented in California wilderness
[[Page 24290]]
areas and National Parks (Cole and Landres 1996, p. 171). This invasion
has been attributed to sheep grazing, though the phenomenon has been
observed on both ungrazed meadows and on meadows grazed continually
since about 1900 (Ratliff 1985, p. 35), suggesting that other drivers
may be involved (see ``Effects of Fire Suppression on Meadow Habitats''
and ``Climate Effects to Meadow Habitat'' below).
Effects of Historical Livestock Grazing
Grazing of livestock in Sierra Nevada meadows and riparian areas
(rivers, streams, and adjacent upland areas that directly affect them)
began in the mid-1700s with the European settlement of California
(Menke et al. 1996, p. 7). Following the gold rush of the mid-1800s,
grazing increased to a level exceeding the carrying capacity of the
available range, causing significant impacts to meadow and riparian
ecosystems (Meehan and Platts 1978, p. 275; Menke et al. 1996, p. 7).
By the turn of the 20th century, high Sierra Nevada meadows were
converted to summer rangelands for grazing cattle, sheep, horses,
goats, and pigs, although the alpine areas were mainly grazed by sheep
(Beesley 1996, pp. 7-8; Menke et al. 1996, p. 14). Stocking rates of
both cattle and sheep in Sierra meadows in the late 19th and early 20th
centuries were very heavy (Kosco and Bartolome 1981, pp. 248-250), and
grazing severely degraded many meadows (Ratliff 1985, pp. 26-31; Menke
et al. 1996, p. 14). Grazing impacts occurred across the entire range
of the Yosemite toad, as cattle and sheep were driven virtually
everywhere in the Sierra Nevada where forage was available (Kinney
1996, pp. 37-42; Menke et al. 1996, p. 14).
Grazing within the National Forests has continued into recent
times, with reduction in activity (motivated by resource concerns,
conflicts with other uses, and deteriorating range conditions)
beginning in the 1920s. A brief wartime increase in the 1940s followed,
before grazing continued to be scaled back beginning in the 1950s
through the early 1970s. However, despite these reductions, grazing
still exceeded sustainable capacity in many areas (Menke et al. 1996,
p. 9; UC 1996a, p. 115). Historical evidence indicates that heavy
livestock use in the Sierra Nevada has resulted in widespread damage to
rangelands and riparian systems due to sod destruction in meadows,
vegetation destruction, and gully erosion (see review in Brown et al.
2009, pp. 56-58 and in USFS et al. 2009, p. 57). (For additional
information on historical grazing regimes, refer to the Effects of
Excessive Historical Grazing section in Factor A analysis for the
Sierra Nevada and mountain yellow-legged frogs, above).
Livestock grazing in the Sierra Nevada has been widespread for so
long that, in most places, no ungrazed areas are available to
illustrate the natural condition of the habitat (Kattelmann and Embury
1996, pp. 16-18). Dull (1999, p. 899) conducted stratigraphic pollen
analysis (identification of pollen in sedimentary layers) in mountain
meadows of the Kern Plateau, and found significant vegetation changes
attributable to sheep and cattle grazing by 1900 (though fire regime
change was also implicated; see below). This degradation is widespread
across the Sierra Nevada. Cooper and Wolf 2006 (p. 1) reports that 50
to 80 percent of grazed meadows now dominated by dry meadow plants were
formerly wet meadows (Cooper and Wolf 2006, p. 1).
Due to the long history (Menke et al. 1996, Ch. 22, pp. 1-52) of
livestock and packstock grazing in the Sierra Nevada and the lack of
historical Yosemite toad population size estimates, it is impossible to
establish a reliable quantitative estimate for the historical
significance and contribution of grazing on Yosemite toad populations.
However, because of the documented negative effects of livestock on
Yosemite toad habitat, and the documented direct mortality caused by
livestock, the decline of some populations of Yosemite toad has been
attributed to the effects of livestock grazing (Jennings and Hayes
1994, pp. 50-53; Jennings 1996, pp. 921-944). Because Yosemite toad
breeding habitat is generally in very shallow waters within meadows,
the breeding habitat is thought to be more vulnerable to changes in
hydrology caused by grazing because the small shallow pools are more
easily impacted (Knapp 2002c, p. 1; Martin 2002, pp. 1-3; USFS et al.
2009, pp. 22, 59-62; Brown 2013, unpaginated). U.S. Geological Survey
records indicate that Yosemite, Sequoia, and Kings Canyon have no
meadows within the parks that are documented to have degraded hydrology
(see NPS 2013, p. 7); conditions in the parks may be related to the
early elimination of most grazing on national parklands in the Sierra
Nevada.
Effects of Current Livestock Grazing
Currently, approximately 33 percent of the estimated range of the
Yosemite toad is within active USFS grazing allotments (USFS 2008,
geospatial data). While stocking rates have been reduced or eliminated
in most areas, legacy effects including eroded channels, soil erosion,
and stream entrenchment that resulted in lowered water tables, drier
meadows, and tree encroachment could still be observed in some Sierran
meadows, especially in National Forests where grazing was more intense
(Vankat and Major 1978, pp. 386-397). Meadow conditions in the Sierra
Nevada have improved over time, but local problems could still be found
as of 1985 (Ratliff 1985, pp. ii-iii) and numerous examples of head-
cutting and stream incision are available within the range of the toad
(Knapp 2013, unpaginated). (For additional information, see sections
above pertaining to effects of grazing on the mountain yellow-legged
frogs.)
The influence of grazing on toad populations in recent history is
uncertain, despite more available data on land use and Yosemite toad
occurrence. In 2005, the USFS, in collaboration with other researchers,
began a 5-year study with multiple components to assess the effects of
grazing on Yosemite toads (Allen-Diaz et al. 2010, pp. 1-45; Roche et
al. 2012a, pp. 56-65; Roche et al. 2012b, pp. 1-11; McIlroy et al..
2013, pp. 1-11). Specifically, the goals of the research were to
assess: (1) Whether livestock grazing under SNFPA Riparian Standards
and Guidelines has a measurable effect on Yosemite toad populations and
(2) effects of livestock grazing on key habitat components that affect
survival and recruitment of Yosemite toad populations. SNFPA standards
and guidelines limit livestock utilization of grass and grass-like
plants to a maximum of 40 percent (or a minimum 4-inch stubble height)
(USDA 2004, p. 56). These companion studies did not detect an effect
from grazing activity on young-of-year toad density or breeding pool
occupancy, water quality, or cover (Allen-Diaz et al. 2010, p. 1; Roche
et al. 2012a, p. 56; Roche et al. 2012b, p. 1-1; McIlroy et al.. 2013,
p. 1).
It is important to note that the results of these studies did not
present a direct measurement of toad survival (for example, mark--
recapture analysis of population trends), and the design was limited in
numbers of years and treatment replicates. It is plausible that, for
longer lived species with irregular female breeding activity over the
time course of this particular study, statistical power was not
sufficient to discern a treatment effect. Further, a time lag could
occur between effect and discernible impacts, and significant
confounding variability in known drivers such as interannual variation
in climate.
Additionally, the experimental design in the studies tested the
hypothesis that forest management guidelines (at 40
[[Page 24291]]
percent use threshold) were impacting toad populations, and this
limited some analyses and experimental design to sites with lower
treatment intensities. Researchers reported annual utilization by
cattle ranging from 10-48 percent, while individual meadow use ranged
from 0-76 percent (the SNFPA allowable use is capped at 40 percent)
(Allen-Diaz et al. 2010, p. 5). As a result of the study design, the
Allen-Diaz study does not provide sufficient information on the impacts
of grazing on Yosemite toads above the prescribed management
guidelines. In general, it is not clear to what extent brief episodes
of intense use (such as in cattle gathering areas) have as negative
impacts on toads, or over what percentage of the grazed meadow
landscape such heavier usage may occur.
The researchers observed significant variation in young-of-year
occupancy in pools between meadows and years, and within meadows over
years (Allen-Diaz et al. 2010, p. 7). This variability would likely
mask treatment effects, unless the grazing variable was a dominant
factor driving site occupancy, and the magnitude of the effect was
quite severe. Further, in an addendum to the initial report, Lind et
al. (2011b, pp. 12-14) report statistically significant negative
(inverse) relationships for tadpole density and grazing intensity
(tadpole densities decreased when percent use exceeded between 30 and
40 percent). This result supports the hypothesis that grazing at
intensities approaching and above the 40 percent threshold can
negatively affect Yosemite toad populations.
Allen-Diaz et al. (2010, p. 2) and Roche et al. (2012b, pp. 6-7)
found that toad occupancy is strongly driven by meadow wetness
(hydrology) and suggested attention should focus on contemporary
factors directly impacting meadow wetness, such as climate, fire regime
changes, and conifer encroachment (see Factor A above). The researchers
also stated that meadow use by cattle during the grazing season is
driven by selection of plant communities found in drier meadows (Allen-
Diaz et al. 2010, p. 2). This suggests that the apparent differences in
preference could provide for some segregation of toad and livestock use
in meadow habitats, so that at least direct mortality threats may be
mitigated by behavioral isolation. Based on the limitations of the
study as described above, we find the initial results from Allen-Diaz
et al. (2010, pp. 1-45) to be inconclusive to discern the impacts of
grazing on Yosemite toad populations where grazing and toads co-occur
in meadows.
The available grazing studies focus on breeding habitat (wet
meadows) and do not consider impacts to upland habitats. The USFS
grazing guidelines for protection of meadow habitats of the Yosemite
toad include fencing breeding meadows, but they do not necessarily
protect upland habitat. Martin (2008) surveyed 11 meadow sites located
along a stream channel in or near low growing willows both before and
after cattle grazed the entire meadow, and Martin found that Yosemite
toads could no longer be located along the stream channel after the
vegetation was grazed. However, both adults and subadults could be
found in dense willow thickets or in parts of the meadow that were less
heavily grazed (Martin 2008, p. 298). Grazing can also degrade or
destroy moist upland areas used as nonbreeding habitat by Yosemite
toads (Martin 2008, p. 159), especially when nearby meadow and riparian
areas have been fenced to exclude livestock. Livestock may also
collapse rodent burrows used by Yosemite toads as cover and hibernation
sites (Martin 2008, p. 159) or disturb toads and disrupt their
behavior. Martin (2008, pp. 305-306) observed that grazing
significantly reduced vegetation height at grazed meadow foraging
sites, and since these areas are not protected by current grazing
guidelines, deduced that cattle grazing is having a negative effect on
terrestrial life stage survivorship in Yosemite toads. This problem was
exacerbated as fenced areas effectively shifted grazing activity to
upland areas actively used by terrestrial life stages of the Yosemite
toad (Martin 2008, p. 306).
Although we lack definitive data to assess the link between
Yosemite toad population dynamics and habitat degradation by livestock
grazing activity, in light of the documented impacts to meadow habitats
(including effects on local hydrology) from grazing activity in
general, we consider this threat prevalent with moderate impacts to the
Yosemite toad and a potential limiting factor in population recovery
rangewide. In addition, given the potential for negative impacts from
heavy use, and the vulnerability of toad habitat should grazing
management practices change with new management plans, we expect this
threat to continue into the future.
Roads and Timber Harvest Effects to Meadow Habitat
Road construction and use, along with timber harvest activity, may
impact Yosemite toad habitat via fragmentation, ground disturbance, and
soil compaction or erosion (Helms and Tappeiner 1996, pp. 439-476).
Roads may alter both the physical environment and the chemical
environment; roads may present barriers to movement and may alter
hydrologic and geomorphic processes that shape aquatic systems, while
vehicle emissions and road-runoff are expected to contain chemicals
that may be toxic (USFS et al. 2009, pp. 71-73). Timber harvests and
past development of roads could potentially also lead to increased
rates of siltation, contributing to the loss of breeding habitats for
the Yosemite toad.
Prior to the formation of National Parks and National Forests,
timber harvest was widespread and unregulated in the Sierra Nevada;
however, most cutting occurred below the current elevation range of the
Yosemite toad (University of California at Davis (UCD) UC 1996b, pp.
17-45; USFS et al. 2009, p. 77). Between 1900 and 1950, most timber
harvest occurred in old-growth forests on private land (UC 1996b, pp.
17-45). During this period, forest plans often lacked standards to
protect riparian areas and associated meadows, leading to harvest
activities that included cutting to edges of riparian areas and forest
road construction that often crossed streams, associated aquatic
habitat, and meadows, and resulted in head-cutting, lowered water
tables, and loss of riparian habitats; legacies of these past
activities remain today (USFS et al. 2009, p. 77). Currently on
National Forests, timber harvest and related vegetation management
activities overlap with Yosemite toads primarily in the lower elevation
portions of the species' range; the red fir and lodgepole forests that
generally surround high-elevation meadows that are Yosemite toad
habitat do not have commercial value (USFS et al. 2009, pp. 76, 77).
Forest standards and guidelines currently provide protections for
riparian areas, such as buffers for timber and vegetation management
activities.
The majority of forest roads in National Forests of the Sierra
Nevada were built between 1950 and 1990, to support major increases in
timber harvest on National Forests, (USDA 2001a, p. 443), suggesting
that many forest roads occur at elevations below the current range of
the Yosemite toad. Relatively few public roads, including trans-Sierran
State Highways 4 (Ebbetts Pass), 88 (Carson Pass), 108 (Sonora Pass),
and 120 (Tioga Pass), cross the high elevations of the Sierra Nevada
within the range of the Yosemite toad (USFS et al. 2009, p. 71),
although smaller public roads are present in some high-elevation areas.
One percent of
[[Page 24292]]
Yosemite toad populations occur on private lands where urbanization and
corresponding construction of new roads may be more likely (USFS et al.
2009, p. 71); however, we are not aware of any proposals for new road
construction at this time.
We expect that the majority of timber harvest, road development,
and associated management impacts (see ``Effects of Fire Suppression on
Meadow Habitats'' below) to Yosemite toad habitat took place during the
expansion period in the latter half of the 20th century. Using a model,
Liang et al. (2010, p. 16) found that Yosemite toads were more likely
to occur in areas closer to timber activity, although the high
correlation between elevation and the distance to harvest activity in
model results definitive conclusions regarding cause and effect.
However, they noted that, because timber harvest activities may
maintain breeding sites by opening the forest canopy and potentially
preventing encroachment of trees into sites, breeding animals might
benefit from timber activity (Liang et al. 2010, p. 16). Limited
information from timber sale areas where low-elevation populations
occur indicates that such activities may negatively affect upland
habitat use if burrow sites are crushed (USFS 2013, p. 6). Although
ground-disturbance due to timber harvest activities has the potential
to have population-level effects on Yosemite toad habitat, especially
where habitat is limited, currently the best available information does
not indicate that the current level of timber harvest occurring within
watersheds currently inhabited by the Yosemite toad is adversely
affecting habitat (USFS et al. 2009, p. 77). Therefore the best
available scientific and commercial information does not indicate that
ongoing road construction and maintenance or timber harvest are
significant threats to the Yosemite toad. There may be localized
effects of these activities in areas where legacy effects continue to
result in modified wet meadow habitat conditions, or where current
harvest and road activities occur in close proximity to extant Yosemite
toad populations.
Effects of Fire Suppression on Meadow Habitats
Fire management refers to activities over the past century to
combat forest fires. Historically, both lightning-caused fires and
fires ignited by American Indians were regularly observed in western
forests (Parsons and Botti 1996, p. 29), and in the latter 19th
century, the active use of fire to eliminate tree canopy in favor of
forage plants continued by sheepherders (Kilgore and Taylor 1979, p.
139). Beginning in the 20th century, land management in the Sierra
Nevada shifted to focus on fire suppression as a guiding policy (UC
2007, p. 10).
Long-term fire suppression has influenced forest structure and
altered ecosystem dynamics in the Sierra Nevada. In general, the time
between fires is now much longer than it was historically, and live and
dead fuels are more abundant and continuous (USDA 2001a, p. 35). Much
of the habitat for the Yosemite toad occurs in high-elevation meadows
within wilderness and backcountry areas where vegetation is sparse and
fire suppression activities are rarely conducted (USFS et al. 2009, p.
55), suggesting that fire suppression has played a limited role in such
locations. At high elevations, encroachment of lodgepole pine at meadow
edges has been attributed to cessation of sheep grazing or legacy
effects of high-intensity grazing that reduced water tables, as opposed
to fire suppression activities (Vankat and Major 1978, pp. 392-395). At
lower elevations, it is not clear how habitat changes attributed to
fire suppression have affected Yosemite toad populations. However,
Liang et al. (2010, p. 16) observed that toads were less likely to
occur in areas where the fire regime was significantly altered from
historical conditions, and suggested that the toads are affected by
some unknown or unmeasured factors related to fire management.
Evidence indicates that fire plays a significant role in the
evolution and maintenance of lower elevation forested meadows of the
Sierra Nevada. Under natural conditions, conifers are excluded from
meadows by fire and saturated soils. Small fires thin and/or destroy
encroaching conifers, while large fires are believed to determine the
meadow--forest boundary (Vankat and Major 1978, p. 394; Parsons and
DeBenedetti 1979, pp. 29-31). Fire is thought to be important in
maintaining open aquatic and riparian habitats for amphibians in some
systems (Russel et al. 1999, pp. 374-384), and fire suppression may
have thereby contributed to conifer encroachment on meadows (Chang
1996, pp. 1071-1099; NPS 2002, p. 1). However, fire suppression effects
are thought to vary with ecosystem fire regime; variable-interval fires
are characteristic of the upper montane red fir forests (Chang 1996,
pp. 107, 1072) that are the setting for Yosemite toad habitat at the
lower elevations of its range, while long-interval fires are
characteristic of the subalpine lodgepole pine forests (Chang 1996, p.
1072) that are the setting for Yosemite toad habitats at higher
elevations. The effects of fire suppression on forest structure is
thought to be far less important in the longer interval forest types
(Chang 1996, p. 1072).
While no studies have confirmed a link between fire suppression and
rangewide population decline of the Yosemite toad, circumstantial
evidence to date suggests that historic fire suppression may be a
factor underlying meadow encroachment at lower elevations. The effect
of fire suppression, therefore, is thought to be largely restricted to
lower elevations within the Yosemite toad's range; fire suppression
activities are rarely conducted where much of the habitat for the
Yosemite toad occurs (USFS et al. 2009, pp. 51-54). Based on the best
available information, we find it likely that habitat modification due
to reduced fire frequency is a moderate threat to Yosemite toad in
those lower-elevation areas where fire suppression has resulted in
conifer encroachment into meadows.
Recreation and Packstock Effects to Meadow Habitat
Recreational activities take place throughout the Sierra Nevada,
and they can have significant negative impacts on wildlife and their
habitats (USDA 2001a, pp. 221, 453-500). Recreation can cause
considerable impact to vegetation and soils in western U.S. Wilderness
Areas and National Parks even with light use, with recovery occurring
only after considerable periods of non-use (USFS et al. 2009, p. 66).
Heavy foot traffic in riparian areas tramples vegetation, compacts
soils, and can physically damage streambanks. Trails (foot, horse,
bicycle, or off-highway motor vehicle) can compact the soil, displace
vegetation, and increase erosion, thereby potentially lowering the
water table (Kondolph et al. 1996, pp. 1009-1026). However, the
National Park Service considers current hiking and backpacking
activities to be a negligible risk factor for the Yosemite toad within
the Parks. The Parks have also worked to improve impacted meadows by
reconstructing poorly designed trails that have degraded meadow
hydrology, also identifying additional Yosemite toad meadows to
prioritize additional restoration activities (NPS 2013, p. 9). Similar
activities have been implemented on National Forests; for example, the
Inyo National Forest has re-routed several trails to avoid the toad's
breeding habitat (USFS 2013, p. 5).
[[Page 24293]]
Although much Yosemite toad habitat is located in wilderness or
other backcountry areas removed from motorized access, the USFS has
noted locations where proximity of roads or off-highway vehicle routes
to Yosemite toad breeding habitat has resulted in observed impacts to
Yosemite breeding habitat. Off-highway vehicles are often the first
vehicles to pass through roads blocked by winter snows, occasionally
driving off the road to pass remaining obstacles (USFS et al. 2009, p.
63). Records of such off-highway vehicle travel in breeding meadows and
ponds (USFS 2013, pp. 6, 7) suggests that such activities have the
potential to negatively affect these habitats, although the population-
level effects to Yosemite toads are thought to be limited.
Packstock use has similar effects to those discussed for livestock
grazing (for additional information on current packstock use levels and
management protections, see the Packstock Use section under the
mountain yellow-legged frogs, above), although this risk factor is
potentially more problematic as this land use typically takes place in
more remote and higher-elevation areas occupied by Yosemite toads, and
packstock tend to graze in many of the same locations that the toads
prefer (USFS et al. 2009, p. 65). Currently, there are very few studies
on the effects of packstock grazing on amphibians, especially in the
Sierra Nevada. However, in Yosemite, Sequoia, and Kings Canyon National
Parks, packstock use is monitored annually to prevent long-term
impacts. Additionally, the NPS (2013, p. 9) has indicated that, except
for a few specific areas, packstock use and Yosemite toads typically do
not overlap within the Parks. Many areas are closed to packstock use
entirely or limited to day use due to inadequate trail access or to
protect sensitive areas. Long-term use data indicate that packstock use
is declining, with no evidence to suggest that it will increase in the
future (NPS 2013, pp. 6, 7). Where permitted, current guidelines in the
National Parks limit trips to 20-25 animals, regulated under
conditional use permits (Brooks 2012, pers. comm.). Similar standards
and guidelines limit packstock group size and use within the National
Forests (USFS 2013, pp. 3-5).
Habitat-related effects of recreational activities on the Yosemite
toad may have population-level impacts in localized areas and under
site-specific conditions, for example, where foot traffic adjacent to
occupied meadows is exceptionally heavy and results in meadow damage,
where legacy effects of high recreation use have resulted in continuing
meadow damage, or where off-highway vehicle use results in changes in
meadow hydrology. However, in general, we do not consider habitat-
related changes associated with current levels of hiking or backpacking
to pose a population-level risk to Yosemite toads. Therefore, at this
time we consider recreational activities to be a low prevalence threat
across the range of the Yosemite toad.
Dams and Water Diversions Effects to Meadow Habitat
Past construction of dams, diversion, and irrigation ditches
resulted in a vast man-made network that altered local and regional
stream hydrology in the Sierra Nevada (SNEP 1996, p. 120), although,
with the exception of several dozen small impoundments and diversions,
almost all of these are located below the range of the Yosemite toad
(USFS et al. 2009, pp. 76, 77). However, in the past a small number of
reservoirs were constructed within the historic range of the Yosemite
toad, most notably Upper and Lower Blue Lakes, Edison, Florence,
Huntington, Courtright, and Wishon Reservoirs. Construction of several
high-elevation reservoirs (for example, Edison and Florence) is thought
to have inundated shallow-water breeding habitat for the toad (USFS et
al. 2009, pp. 76, 77). Where reservoirs are used for hydroelectric
power, water-level declines caused by drawdown of reservoirs can lead
to the mortality of eggs and tadpoles by stranding and desiccation,
although, with the exception of Blue Lakes, Yosemite toads are
currently not known from the above reservoirs (USFS et al. 2009, pp.
78, 79).
Past construction of these reservoirs likely contributed to the
decline of the Yosemite toad in the area where they were built.
Increasing effects from climate change, or new water supply development
in response to such effects, may exacerbate this risk in the future if
new reservoirs are constructed within areas occupied by the toad.
However, we are not aware of any proposals to construct additional
reservoirs within the Yosemite toads range. We expect that continuing
reservoir operations may have continued habitat-related effects to toad
populations in these developed areas, but less so in the current extent
of the Yosemite toad's (remnant) range. Therefore, we consider this
threat to be of low prevalence to the Yosemite toad across its range.
Climate Effects to Meadow Habitat
Different studies indicate that multiple drivers are behind the
phenomenon of conifer encroachment into meadows. The first factor
affecting the rate of conifer encroachment into meadow habitats, fire
suppression, was discussed above. Climate variability is another factor
affecting the rate of conifer encroachment on meadow habitats. A study
by Franklin et al. (1971, p. 215) concluded that fire had little
influence on meadow maintenance in their study area, while another
study concluded that climate change is a more likely explanation for
encroachment of trees into the adjacent meadow at their site, rather
than fire suppression or changes in grazing intensity (Dyer and
Moffett, 1999, p. 444).
Climatic variability is strongly correlated with tree encroachment
into dry subalpine meadows (Jakubos and Romme 1993, p. 382). In the
Sierra Nevada, most lodgepole pine seedlings become established during
years of low snowpack when meadow soil moisture is reduced (Wood 1975,
p. 129). The length of the snow-free period may be the most critical
variable in tree invasion of subalpine meadows (Franklin et al. 1971,
p. 222), with the establishment of a good seed crop, followed by an
early snowmelt, resulting in significant tree establishment. It is
apparent that periods of low snowpack and early melt may in fact be
necessary for seedling establishment (Ratliff, 1985, p. 35). Millar et
al. (2004, p. 181) reported that increased temperature, coupled with
reduced moisture availability in relation to large-scale temporal
shifts in climate, facilitated the invasion of 10 subalpine meadows
studied in the Sierra Nevada.
Our analyses under the Act include consideration of ongoing and
projected changes in climate. The terms ``climate'' and ``climate
change'' are defined by the Intergovernmental Panel on Climate Change
(IPCC). ``Climate'' refers to the mean and variability of different
types of weather conditions over time, with 30 years being a typical
period for such measurements, although shorter or longer periods also
may be used (IPCC 2007, p. 1450; IPCC 2013a, Annex III). The term
``climate change'' thus refers to a change in the mean or variability
of one or more measures of climate (for example, temperature or
precipitation) that persists for an extended period, typically decades
or longer, whether the change is due to natural variability, human
activity, or both (IPCC 2007, p. 1450; IPCC 2013a, Annex III). A recent
compilation of climate change and its effects is available from reports
of the Intergovernmental Panel on Climate Change (IPCC) (IPCC 2013b,
entire). Various types of changes in climate can have direct or
indirect effects on species. These effects may be positive,
[[Page 24294]]
neutral, or negative, and they may change over time, depending on the
species and other relevant considerations, such as the effects of
interactions of climate with other variables (for example, habitat
fragmentation) (IPCC 2007, pp. 8-14, 18-19). In our analyses, we use
our expert judgment to weigh relevant information, including
uncertainty, in our consideration of various aspects of climate change.
For the Sierra Nevada ecoregion, climate models predict that mean
annual temperatures will increase by 1.8 to 2.4 [deg]C (3.2 to
4.3[emsp14][deg]F) by 2070, including warmer winters with earlier
spring snowmelt and higher summer temperatures (PRBO 2011, p. 18).
Additionally, mean annual rainfall is projected to decrease from the
current average by some 9.2-33.9 cm (3.6-13.3 in) by 2070 (PRBO 2011,
p. 18). However, projections have high uncertainty, and one study
predicts the opposite effect (PRBO 2011, p. 18). Snowpack is, by all
projections, going to decrease dramatically (following the temperature
rise and increase in precipitation falling as rain) (PRBO 2011, p. 19);
(Kadir et al. 2013, pp. 76-80). Higher winter stream flows, earlier
runoff, and reduced spring and summer stream flows are projected, with
increasing severity in the southern Sierra Nevada (PRBO 2011, pp. 20-
22); (Kadir et al. 2013, pp. 71-75).
Snow-dominated elevations from 2,000-2,800 m (6,560-9,190 ft) will
be the most sensitive to temperature increases (PRBO 2011, p. 23).
Meadows fed by snowmelt may dry out or be more ephemeral during the
non-winter months (PRBO 2011, p. 24). This pattern could influence
groundwater transport, and springs may be similarly depleted, leading
to lower water levels in available breeding habitat and decreased area
and hydroperiod (i.e., duration of water retention) of suitable habitat
for rearing tadpoles of Yosemite toads. Changes in water transport may
promote channel incision and result in a shift to non-meadow conditions
(Viers et al. 2013, p. 31).
Blaustein et al. (2010, pp. 285-300) provide an exhaustive review
of potential direct and indirect and habitat-related effects of climate
change to amphibian species, with documentation of effects in a number
of species where such effects have been studied. Altitudinal range
shifts with changes in climate have been reported in some regions. They
note that temperature can influence the concentration of dissolved
oxygen in aquatic habitats, with warmer water generally having lower
concentrations of dissolved oxygen, and that water balance heavily
influences amphibian physiology and behavior. They predict that
projected changes in temperature and precipitation are likely to
increase habitat loss and alteration for those species living in
sensitive habitats, such as ephemeral ponds and alpine habitats
(Blaustein et al. 2010, pp. 285-287).
Because environmental cues such as temperature and precipitation
are clearly linked to onset of reproduction in many species, climate
change will likely affect the timing of reproduction in many species,
potentially with different sexes responding differently to climate
change. For example, males of two newt species (Triturus spp.) showed a
greater degree of change in arrival date at breeding ponds (Blaustein
et al. 2010, p. 288). Lower concentrations of dissolved oxygen in
aquatic habitats may negatively affect developing embryos and larvae,
in part because increases in temperature increase the oxygen
consumption rate in amphibians. Reduced oxygen concentrations have also
been shown to result in accelerated hatching in ranid frogs, but at a
smaller size, while larval development and behavior may also be
affected and may be mediated by larval density and food availability
(Blaustein et al. 2010, pp. 288-289).
Increased temperatures can reduce time to metamorphosis, which can
increase chances of survival where ponds dry, but also result in
metamorphosis at a smaller size, suggesting a likely trade-off between
development and growth, which may be exacerbated by climate change and
have fitness consequences for adults (Blaustein et al. 2010, pp. 289-
290). Changes in terrestrial habitat, such as changed soil moisture and
vegetation, can also directly affect adult and juvenile amphibians,
especially those adapted to moist forest floors and cool, highly
oxygenated water that characterizes montane regions. Climate change may
also interact with other stressors that may be acting on a particular
species, such as disease and contaminants (Blaustein et al. 2010, pp.
290-299).
A recent paper (Kadir et al. 2013, entire) provides specific
information on the effects of climate change in the Sierra Nevada. The
report found that glaciers in the Sierra Nevada have decreased in area
over the past century, and glacier shrinkage results in earlier peak
water runoff and drier summer conditions. Another result from the
report is that the lower edge of the conifer-dominated forests in the
Sierra Nevada has been retreating upslope over the past 60 years.
Regarding wildfire, since 1950, annual acreage burned in wildfires
statewide has been increasing in California, and in the western United
States, large wildfires have become more frequent, increasing in tandem
with rising spring and summer temperatures. Finally, the report found
that today's subalpine forests in the Sierra Nevada are much denser--
that is, comprise more small-diameter trees--than they were over 70
years ago. During this time period, warmer temperatures, earlier
snowmelt, and more rain than snow occurred in this region. Many of
these changes in the Sierra Nevada of California due to climate are
likely to influence Yosemite toads because they are highly vulnerable
to climate change because changing hydrology and habitat in the Sierra
Nevada will likely have impacts on remaining populations (Viers et al.
2013, pp. 55, 56).
Historically, drought is thought to have contributed to the decline
of the Yosemite toad (Kagarise Sherman and Morton 1993, p. 186;
Jennings and Hayes 1994, pp. 50-53). Extended and more severe droughts
pose an ongoing, rangewide risk to the species and are expected to
increase with predicted climate changes (PRBO 2011, p. 18). Such
changes may reduce both the amount of suitable breeding habitat and the
length of time that suitable water is available in that habitat (Brown
2013, unpaginated).
Davidson et al. (2002, p. 1598) analyzed geographic decline
patterns for the Yosemite toad. They compared known areas of
extirpation against a hypothesized model for climate change that would
predict greater numbers of extirpations at lower altitudes, and in more
southern latitudes. The researchers did not observe a pattern in the
available historic data to support the climate change hypothesis as a
driver of historic population losses, although they acknowledge that
climate change may be a contributor in more complex or subtle ways.
Additionally, this study was limited by small sample size, and it is
possible that climate change effects on the Yosemite toad (a long-lived
species) may not become evident for many years (USFS et al. 2009, p.
48). Finally, Davidson et al. (2002, p. 1598) did find an increase in
occupancy with elevation (greater densities of populations at
altitude), and this observation is consistent with a pattern that would
fit a response to climate change (USFS et al. 2009, p. 48). However,
this observation would also be consistent if the features of these
particular habitats (such as at higher elevation) were more suited to
the special ecological requirements of the
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toad, or if other stressors acting on populations at lower elevations
were responsible for the declines. We, therefore, find these results
inconclusive.
Most recently, modeled vulnerability assessments for Sierra Nevada
montane meadow systems have utilized life history and habitat
requirements to gauge vulnerability of amphibian species to climate
change. This assessment indicates that vulnerability to hydro-climatic
changes will likely be very high for the Yosemite toad, and that
continued or worsening stream channelization in montane meadows from
flashy storms may worsen effects by further reductions in the water
table (Viers et al. 2013, p. 56).
The breeding ecology and life history of the Yosemite toad are that
of a habitat specialist, as it utilizes pool and meadow habitats during
the onset of snowmelt and carefully times its reproduction to fit
available conditions within ephemeral breeding sites. The most striking
documented declines in Yosemite toad populations in the historical
record are correlated with extreme climate episodes (drought) (Kagarise
Sherman and Morton 1993, pp. 186-198). Given these observations, it is
likely that climate change (see also discussion in mountain yellow-
legged frog's Summary of Factors Affecting the Species, under Factor E)
poses a significant risk to the Yosemite toad now and in the future. It
is quite possible that these impacts are occurring currently, and have
occurred over the last few decades. However, it is difficult in short
time intervals to discern the degree of effect from climate change
within the variability of natural climate cycles.
In summary, based on the best available scientific and commercial
information, we consider the threats of destruction, modification, and
curtailment of the species' habitat and range to be significant ongoing
threats to the Yosemite toad. The legacy effects of past land uses have
altered meadow communities through the mechanism of stream incision by
permanently reducing habitat quantity and quality unless active and
costly restoration is implemented. Climate change is a current threat
of high magnitude. Threats considered of moderate magnitude include
livestock grazing and fire management regime. Threats considered
currently low magnitude include roads and timber harvest, dams and
water diversions, and recreational land uses.
Factor B. Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes
We do not have any scientific or commercial information to indicate
that overutilization for commercial, recreational, or scientific
purposes poses a threat to the Yosemite toad. There is currently no
known commercial market for Yosemite toads, although one pet store in
Los Angeles that is no longer in business had previously sold at least
one Yosemite toad (USFS et al. 2009, pp. 65-66); and there is also no
documented recreational or educational use for Yosemite toads.
Scientific research may cause some stress to Yosemite toads through
disturbance and disruption of behavior, handling, and injuries
associated with marking individuals. This activity has resulted in the
known death of individuals through accidental trampling (Green and
Kagarise Sherman 2001, pp. 92-103), irradiation from radioactive tags
(Karlstrom 1957, pp. 187-195), and collection for museum specimens
(Jennings and Hayes 1994, pp. 50-53). We expect that requirements for
Federal (USFS and NPS) and State (CDFW) research and special use
permits, and University ethics requirements provide some protections
for wildlife-research subjects and limit negative effects to
individuals. Therefore, we do not currently consider ongoing and future
scientific research to be a threat to the Yosemite toad. We also
anticipate that further research into the genetics and life history of
the Yosemite toad and broader methodological censuses will provide a
net conservation benefit to this under-studied species.
Based on the best available scientific and commercial information,
we do not consider overutilization for commercial, recreational,
scientific, or educational purposes to be a threat to the Yosemite
toad.
Factor C. Disease or Predation
Predation
Prior to the trout stocking of high Sierra Nevada lakes, which
began over a century ago, fish were entirely absent from most of this
region (Bradford 1989, pp. 775-778). Observations regarding the effects
of introduced fishes on the Yosemite toad are mixed. However, re-
surveys of historical Yosemite toad sites have shown that the species
has disappeared from several lakes where they formerly bred, and these
areas are now occupied by fish (Stebbins and Cohen 1995, pp. 213-215;
Martin 2002, p. 1).
Drost and Fellers (1994, pp. 414-425) suggested that Yosemite toads
are less vulnerable to fish predation than frogs because they breed
primarily in ephemeral waters that do not support fish. Further,
Jennings and Hayes (1994, pp. 50-53) stated that the palatability of
Yosemite toad tadpoles to fish predators is unknown, but often assumed
to be low based on the unpalatability of western toads (Drost and
Fellers 1994, pp. 414-425; Kiesecker et al. 1996, pp. 1237-1245), to
which Yosemite toads are closely related. Grasso (2005, p. 1) observed
brook trout swimming near, but the trout ignored Yosemite toad
tadpoles, suggesting that tadpoles are unpalatable. The study also
found that subadult Yosemite toads were not consumed by brook trout
(Grasso 2005, p. 1), although the sublethal effects of trout
``sampling'' (mouthing and ejecting tadpoles) and the palatability of
subadults to other trout species are unknown. Martin (2002, p. 1)
observed brook trout preying on Yosemite toad tadpoles, and also saw
them ``pick at'' Yosemite toad eggs (which later became infected with
fungus). In addition, metamorphosed western toads have been observed in
golden trout stomach contents (Knapp 2002c, p. 1). Nevertheless, Grasso
et al. (2010, p. 457) concluded that early life stages of the Yosemite
toad likely possess chemical defenses that provide sufficient
protection from native trout predation.
The observed predation of Yosemite toad tadpoles by trout (Martin
1992, p. 1) indicates that introduced fishes may pose a predation risk
to the species in some situations, which may be accentuated during
drought years. At a site where Yosemite toads normally breed in small
meadow ponds, they have been observed to successfully switch breeding
activities to stream habitat containing fish during years of low water
(Strand 2002, p. 1). Thus, drought conditions may increase the toads'
exposure to predatory fish, and place them in habitats where they
compete with fish for invertebrate prey. Additionally, although the
number of lake breeding sites used by Yosemite toads is small relative
to the number of ephemeral sites, lake sites may be especially
important because they are more likely to be habitable during years
with low water (Knapp 2002c, p. 1).
Overall, the data and available literature suggest that direct
mortality from fish predation is likely not an important factor driving
Yosemite toad population dynamics. This does not discount other
indirect impacts, such as the possibility that fish may be effective
disease vectors (see below). Yosemite toad use of more ephemeral
breeding habitats (which are less habitable to fish species as they
cannot tolerate drying or
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freezing) minimizes the interaction of fish and toad tadpoles. Further,
where fish and toads co-occur, it is possible that food depletion
(outcompetition) by fish negatively affects Yosemite toads (USFS et al.
2009, p. 58).
Other predators may also have an effect on Yosemite toad
populations. Kagarise Sherman and Morton (1993, p. 194) reported
evidence of toad predation by common ravens (Corvus corax) and
concluded this activity was responsible for the elimination of toads
from one site. These researchers also confirmed, as reported in other
studies, predation on Yosemite toad by Clark's nutcrackers (Nucifraga
columbiana). The significance of avian predation may increase if the
abundance of common ravens within the current range of the Yosemite
toad increases as it has in nearby regions (Camp et al. 1993, p. 138;
Boarman et al. 1995, p. 1; Kelly et al. 2002, p. 202). However, the
degree to which avian predation may be affecting Yosemite toad
populations has not been quantified.
Disease
Although not all vectors have been confirmed in the Sierra Nevada,
introduced fishes, humans, pets, livestock, packstock, vehicles, and
wild animals may all act to facilitate disease transmission between
amphibian populations. Infection of both fish and amphibians by a
common disease has been documented with viral (Mao et al. 1999, pp. 45-
52) and fungal pathogens in the western United States (Blaustein et al.
1994b, pp. 251-254). Mass die-offs of amphibians in the western United
States and around the world have been attributed to Bd fungal
infections of metamorphs and adults (Carey et al. 1999, pp. 1-14),
Saprolegnia fungal infections of eggs (Blaustein et al. 1994b, pp. 251-
254), ranavirus infections, and bacterial infections (Carey et al.
1999, pp. 1-14).
Various diseases are confirmed to be lethal to Yosemite toads
(Green and Kagarise Sherman 2001, pp. 92-103), and recent research has
elucidated the potential role of Bd infection as a threat to Yosemite
toad populations (Dodge and Vredenburg 2012, p. 1). These various
diseases and infections, in concert with other factors, have likely
contributed to the decline of the Yosemite toad (Kagarise Sherman and
Morton 1993, pp. 193-194) and may continue to pose a risk to the
species (Dodge and Vredenburg 2012, p. 1).
Die-offs in Yosemite toad populations have been documented in the
literature, and an interaction with diseases in these events has been
confirmed. However, no single cause has been validated by field
studies. Tissue samples from dead or dying adult Yosemite toads and
healthy tadpoles were collected during a die-off at Tioga Pass Meadow
and Saddlebag Lake and analyzed for disease (Green and Kagarise Sherman
2001, pp. 92-103). Six infections were found in the adults, including
infection with Bd, bacillary bacterial septicemia (red-leg disease),
Dermosporidium (a fungus), myxozoa spp. (parasitic cnidarians),
Rhabdias spp. (parasitic roundworms), and several species of trematode
(parasitic flatworms). Despite positive detections, no single
infectious disease was found in more than 25 percent of individuals,
and some dead toads showed no signs of infection to explain their
death. Further, no evidence of infection was found in tadpoles. A meta-
analysis of red-leg disease also revealed that the disease is a
secondary infection that may be associated with a suite of different
pathogens, and so actual causes of decline in these instances were
ambiguous (Kagarise Sherman and Morton 1993, p. 194). The authors
concluded that the die-off was caused by suppression of the immune
system caused by an undiagnosed viral infection or chemical
contamination that made the toads susceptible to the variety of
diagnosed infections.
Saprolegnia ferax, a species of water mold that commonly infects
fish in hatcheries, caused a massive lethal infection of eggs of
western toads at a site in Oregon (Blaustein et al. 1994b, p. 252). It
is unclear whether this event was caused by the introduction of the
fungal pathogen via fish stocking, or if the fungus was already present
and the eggs' ability to resist infection was inhibited by some unknown
environmental factor (Blaustein et al. 1994b, p. 253). Subsequent
laboratory experiments have shown that the fungus could be passed from
hatchery fish to western toads (Kiesecker et al. 2001, pp. 1064-1070).
Fungal growth on Yosemite toad eggs has been observed in the field, but
the fungus was not identified and it was unclear whether the fungus was
the source of the egg mortality (Kagarise Sherman 1980, p. 46). Field
studies conducted in Yosemite National Park found that an undetermined
species of water mold infected only the egg masses that contained dead
embryos of Yosemite toads (Sadinski 2004, pp. 33-34). The researchers
also observed that the water mold became established on egg masses only
after embryo death, and subsequently spread, causing the mortality of
additional embryos of Yosemite toads.
Sadinski (2004, p. 35) discovered that mortality of Yosemite toad
embryos may be attributed to an unidentified species of a free-living
flatworm (Turbellaria spp.). In Yosemite National Park, these worms
were observed to penetrate Yosemite toad egg masses and feed directly
on the embryos. In some locations, Turbellaria spp. reached such large
densities that they consumed all the embryos within a Yosemite toad egg
mass. Predation also facilitated the colonization and spread of water
mold on egg masses, leading to further embryo mortality. Further
studies would be needed to determine which species of Turbellaria feeds
on Yosemite toad eggs, and the extent of this impact on Yosemite toad
populations.
Until recently, the contribution of Bd infection to Yosemite toad
population declines was relatively unknown. Although the toad is
hypothetically susceptible due to co-occurrence with the mountain
yellow-legged frog, the spread and growth of Bd in the warmer pool
habitats, occupied for a much shorter time relative to the frog, is
suspected to render individuals less prone to epidemic outbreaks (USFS
et al. 2009, p. 50). Fellers et al. (2011, p. 391) documented the
occurrence of Bd infection in Yosemite National Park toads over at
least a couple of decades, and they note population persistence in
spite of the continued presence of the pathogen. In a survey of 196
museum specimens, Dodge and Vredenburg (2012, p. 1) report the first
presence of Bd infection in Yosemite toads beginning in 1961, with the
pathogen becoming highly prevalent during the recorded declines of the
late 1970s, before it peaked in the 1990s at 85 percent positive
incidence. In live specimen sampling, Dodge and Vredenburg (2012, p. 1)
collected 1,266 swabs of Yosemite toads between 2006 and 2011, and
found Bd infection intensities at 17-26 percent (with juvenile toads
most affected). The studies detected a pattern indicative of the
historic emergence of Bd, which coincided with the documented decline
in Yosemite toad (Dodge 2013, p. 1). As such, results from these
studies support the hypothesis that Bd infection and chytridiomycosis
have played an important role in Yosemite toad population dynamics over
the period of their recent recorded decline.
Carey (1993, pp. 355-361) developed a model to explain the
disappearance of boreal toads (Bufo boreas boreas) in the Rocky
Mountains, suggesting immune system suppression from extreme winter
stress (``winter stress syndrome'') could have contributed to the
decline in that species. This model may also fit Yosemite toad die-offs
observed by Kagarise Sherman and Morton (1993,
[[Page 24297]]
pp. 186-198), given the close relationship between the two toads, and
their occupation of similar habitats. However, an analysis of immune
system suppression and the potential role of winter stress relative to
Yosemite toad population trends is not available at this time. Yet, the
decline pattern observed in the Carey study is mirrored by the pattern
in the Yosemite toad (heavy mortality exhibited in males first) (Knapp
2012, pers. comm.). This observation, in concert with the recent
results from museum swabs (Dodge and Vredenburg 2012, p. 1), provides a
correlative link to the timing of the recorded Yosemite toad declines
and Bd infection intensities.
Although disease as a threat factor to the Yosemite toad is
relatively less documented, Bd infection causes mass mortalities in the
closely related boreal toad (Carey et al. 2006, p. 19) and there is
evidence related to Bd's role in historical die-offs in Yosemite toads.
Much of the historic research documenting Yosemite toad declines
predated our awareness of Bd as a major amphibian pathogen.
Additionally, the life history of the Yosemite toad, as a rapid breeder
during early snowmelt, limits the opportunities to observe population
crashes in the context of varied environmental stressors. Currently
available evidence indicates that Bd was likely a significant factor
contributing to the recent historical declines observed in Yosemite
toad populations (Dodge and Vredenburg 2012, p. 1). Although infection
intensities are currently lower than some peak historic measurements,
this threat remains a potential factor that may continue to reduce
survival through metamorphosis, and therefore recruitment to the
breeding population (Knapp 2012, pers. comm.). Additionally, the
interaction of disease and other stressors, such as climate extremes,
is not well understood in the Yosemite toad. Research does suggest that
the combination of these threats represents a factor in the historical
decline of the species (Kagarise Sherman and Morton 1993, p. 186).
In summary, based on the best available scientific and commercial
information, we do not consider predation to be a threat to the
species. We consider disease to be a threat to the Yosemite toad that
has a moderate, ongoing effect on populations of the species rangewide.
The threat most specifically includes the amphibian pathogen, Bd.
Although definitive empirical data quantifying the contribution of
disease to Yosemite toad population declines are not currently
available, population declines that were concurrent with the prevalence
and spread of Bd across the Sierra Nevada support the assertion that
disease has played a role in the observed trend. Further, Bd infection,
even at lower intensities, may interact with climate extremes and
continue to depress recruitment of yearling and subadult Yosemite toads
to breeding Yosemite toad populations. We suspect this threat was
historically significant, that it is currently having a moderate
influence on toad populations, and we expect it to be a future concern.
Factor D. The Inadequacy of Existing Regulatory Mechanisms
In determining whether the inadequacy of regulatory mechanisms
constitutes a threat to the Yosemite toad, we analyzed the existing
Federal and State laws and regulations that may address the threats to
the species or contain relevant protective measures. Regulatory
mechanisms are typically nondiscretionary and enforceable, and may
preclude the need for listing if such mechanisms are judged to
adequately address the threat(s) to the species such that listing is
not warranted. Conversely, threats on the landscape are not addressed
by existing regulatory mechanisms where the existing mechanisms are not
adequate (or not adequately implemented or enforced).
We discussed the applicable State and Federal laws and regulations,
including the Wilderness Act, NFMA above (see Factor D discussion for
mountain yellow-legged frogs). In general, the same administrative
policies and statutes are in effect for the Yosemite toad. This section
additionally addresses regulatory mechanisms with a specific emphasis
on the Yosemite toad.
Taylor Grazing Act of 1934
In response to overgrazing of available rangelands by livestock
from the 1800s to the 1930s, Congress passed the Taylor Grazing Act in
1934 (43 U.S.C. 315 et seq.). This action was an effort to stop the
damage to the remaining public lands as a result of overgrazing and
soil depletion, to provide coordination for grazing on public lands,
and to attempt to stabilize the livestock industry (Meehan and Platts
1978, p. 275; Public Lands Council et al. v. Babbitt Secretary of the
Interior et al. (167 F. 3d 1287)). Passage of the Taylor Grazing Act
resulted in reduced grazing in some areas, including the high Sierra
Nevada. However, localized use remained high, precluding regeneration
of many meadow areas (Beesley 1996, p. 14; Menke et al. 1996, p. 14;
Public Lands Council et al. v. Babbitt Secretary of the Interior et al.
(167 F. 3d 1287)).
Existing Federal and State laws and regulatory mechanisms currently
offer some level of protection for the Yosemite toad. Specifically,
these include the Wilderness Act, the NFMA, the SNFPA, and the FPA (see
Factor D discussion for mountain yellow-legged frog complex). Based on
the best available scientific and commercial information, we do not
consider the inadequacy of existing regulatory mechanisms to be a
threat to the Yosemite toad.
Factor E. Other Natural or Manmade Factors Affecting Its Continued
Existence
The Yosemite toad is sensitive to environmental change or
degradation due to its life history, biology, and existence in
ephemeral habitats characterized by climate extremes and low
productivity. It is also sensitive to anthropogenically influenced
factors. For example, contaminants, acid precipitation, ambient
ultraviolet radiation, and climate change have been implicated as
contributing to amphibian declines (Corn 1994, pp. 62-63; Alford and
Richards 1999, pp. 2-7). However, as with the case with the mountain
yellow-legged frog complex, contaminants, acid precipitation, and
ambient ultraviolet radiation are not known to pose a threat (current
or historical) to Yosemite toad and, therefore, are not discussed
further. Please refer to the proposed listing rule for the Sierra
Nevada yellow-legged frog, the northern DPS of the mountain yellow-
legged frog, and the Yosemite toad (78 FR 24472, April 25, 2013) for a
detailed discussion of contaminants, acid precipitation, and ambient
ultraviolet radiation. The following discussion will focus on potential
threat factors specifically studied in the Yosemite toad, based on the
unique life history, population status, demographics, or biological
factors specific to Yosemite toad populations.
Climate Change Effects on Individuals
As discussed above in Factor A, climate change can result in
detrimental impacts to Yosemite toad habitat. Climate variability could
also negatively impact populations through alteration of the frequency,
duration, and magnitude of either droughts or severe winters (USFS et
al. 2009, p. 47). Yosemite toads breed and their tadpoles develop in
shallow meadow and ephemeral habitats, where mortality from desiccation
and freezing can be very high, often causing complete loss of an annual
cohort (USFS et al. 2009, p. 10). Kagarise Sherman and Morton (1993,
pp. 192-193) documented in a long-
[[Page 24298]]
term population study that Yosemite toad hatching success and survival
were subject to a balance between the snowpack water contribution to
breeding pools and the periodicity and character of breeding season
storms and post-breeding climate (whether it is cold or warm). When it
is too cold, eggs and tadpoles are lost to freezing. This situation
poses a risk as earlier snowmelt is expected to cue breeding earlier in
the year, exposing young tadpoles (or eggs) to killing frosts in more
variable conditions of early spring (Corn 2005, p. 60). When it is too
dry, tadpoles are lost to pool desiccation. Alterations in the annual
and seasonal hydrologic cycles that influence water volume and
persistence in Yosemite toad breeding areas can thereby impact breeding
success. The threat of climate change on individuals is significant,
and is of high prevalence now and into the future.
Other Sources of Direct and Indirect Mortality
Direct and indirect mortality of Yosemite toads has occurred as a
result of livestock grazing. Mortality risk from livestock trampling is
expected to be the greatest for non-larval stages where livestock
concentrate in Yosemite toad habitat when toad densities are highest;
early in the season when breeding adults are aggregated and egg masses
are laid; and at metamorphosis when juveniles are metamorphosing in
mass along aquatic margins. However, because cattle typically are not
present during the breeding season, the risk of trampling is expected
to be greatest for metamorphs (USFS et al. 2009, p. 59). Cattle have
been observed to trample Yosemite toad metamorphs and subadult toads,
and these life stages can fall into deep hoofprints and die (Martin
2008, p. 158). Specifically, Martin (2008, p. 158) witnessed some 60
subadult and metamorph toad deaths during the movement of 25 cattle
across a stream channel bordered by willows within a meadow complex.
Adult Yosemite toads trampled to death by cattle have also been
observed (Martin 2002, pp. 1-3). This risk factor is likely of sporadic
significance, and is of greatest concern where active grazing
allotments coincide with breeding meadows. However, it is difficult to
determine the degree of this impact without quantitative data.
Trampling and collapse of rodent burrows by recreationists, pets,
and vehicles could lead to direct mortality of terrestrial life stages
of the Yosemite toad. Recreational activity may also disturb toads and
disrupt their behavior (Karlstrom 1962, pp. 3-34). Recreational anglers
may be a source of introduced pathogens and parasites, and they have
been observed using toads and tadpoles as bait (USFS et al. 2009, p.
66). However, Kagarise Sherman and Morton (1993, p. 196) did not find a
relationship between the distance from the nearest road and the
declines in their study populations, suggesting that human activity was
not the cause of decline in that situation. Recreational activity may
be of conservation concern, and this threat may increase with greater
activity in mountain meadows. However, current available information
does not indicate that recreational activity is a significant stressor
for Yosemite toads.
Fire management practices over the last century have created the
potential for severe fires in the Sierra Nevada. Wildfires do pose a
potential direct mortality threat to Yosemite toads, although
amphibians in general are thought to retreat to moist or subterranean
refuges and thereby suffer low mortality during natural fires (Russel
et al. 1999, pp. 374-384). In the closely related boreal toad (Bufo
boreas), Hossak and Corn (2007, p. 1409) documented a positive response
(increase in occupied breeding sites and population size) following a
wildfire, with returns to near pre-fire occupancy levels after 4 to 5
years (Hossack et al. 2012, p. 224), suggesting that habitat-related
changes associated with wildfires may provide at least short-term
benefits to Yosemite toad populations. However, data on the direct and
indirect effects of fire on Yosemite toads are lacking.
USFS et al. (2009, p. 74) suggested that the negative effects of
roads that have been documented in other amphibians, in concert with
the substantial road network across a portion of the Yosemite toad's
range, indicate this risk factor may be potentially significant to the
species. Roads may facilitate direct mortality of amphibians through
vehicle strikes (DeMaynadier and Hunter 2000, pp. 56-65), and timber
harvest activities (including fuels management and vegetation
restoration activities) have been documented to result in the direct
mortality of Yosemite toads (USFS 2013, p. 94). Levels of timber
harvest and road construction have declined substantially since
implementation of the California Spotted Owl Sierran Province Interim
Guidelines in 1993, and some existing roads have been decommissioned or
are scheduled to be decommissioned (USDA 2001a, p. 445). Therefore, the
risks posed by new roads and timber harvests have declined, but those
already existing still may pose risks to the species and its habitat.
Toads could potentially be trampled or crushed by activities
implemented to reduce fire danger. USFS et al. (2009, p. 53) report
that the Forest Service has initiated a fuels reduction program in
order to reduce the extent and intensity of wildfires. However, most of
these projects will occur in the Wildland Urban Interface, which is
below the elevational range of the Yosemite toad and generally near
human developments. However, in the future some fuels projects may
occur in limited areas around facilities, such as resorts, pack
stations, or summer homes, within the lowest portion of the Yosemite
toad range.
Collectively, direct mortality from land uses within the Yosemite
toad range may have impacts to the toad. However, we are aware of no
studies that have quantified or estimated the prevalence of this
particular threat to be able to assess its impact to Yosemite toad
populations. At the current time, direct and indirect mortality from
roads are not considered to be a significant factor affecting the
Yosemite toad rangewide.
Small Population Size
Although it is believed that the range of the Yosemite toad has not
significantly contracted, the majority of populations across this area
have been extirpated, and this loss has been significant relative to
the historical condition (multitudes of populations within many
watersheds across their geographic range) (see ``Population Estimates
and Status'' above). Further, growing evidence suggest that the
populations that remain are small, numbering fewer than 20 males in
most cases (Kagrise Sherman and Morton 1993, p. 190; Sadinski 2004, p.
40; Brown et al. 2012, p. 125). This situation renders these remnant
populations susceptible to risks inherent to small populations (see
Factor E discussion, ``Small Population Size,'' for mountain yellow-
legged frogs, above) including inbreeding depression and genetic drift,
along with a higher probability of extirpation from unpredictable
events such as severe storms or extended droughts.
Traill et al. (2009, p. 32) argued for a benchmark viable
population size of 5,000 adult individuals (and 500 to prevent
inbreeding) for a broad range of taxa, although this type of blanket
figure has been disputed as an approach to conservation (Flather et al.
2011, pp. 307-308). Another estimate, specific to amphibians, is that
populations of at least 100 individuals are less susceptible to
demographic stochasticity (Schad 2007, p. 10). Amphibian species
[[Page 24299]]
with highly fluctuating population size, high frequencies of local
extinctions, and living in changeable environments may be especially
susceptible to curtailment of dispersal and restriction of habitat
(Green 2003, p. 331). These conditions are all likely applicable to the
Yosemite toad.
Therefore, based on the best available commercial and scientific
information, we conclude that small population size is a prevalent and
significant threat to the species viability of the Yosemite toad across
its range, especially in concert with other extant stressors (such as
climate change).
Cumulative Impacts of Extant Threats
Interactive effects or cumulative impacts from multiple additive
stressors acting upon Yosemite toad populations over time are indicated
by the documented declines in populations and abundance across the
range of the species. Although no single causative factor linked to
population declines in Yosemite toads has been confirmed in the
literature (excepting perhaps extreme climate conditions such as
droughts) (Kagarise Sherman and Morton 1993, p. 186; Jennings and Hayes
1994, pp. 50-53), there has been a decline in population abundance and
numbers of extant populations inhabiting the landscape (Brown et al.
2012, pp. 115-131; Kagarise Sherman and Morton 1993, pp. 186-198). This
pattern of decline suggests a factor or combination of factors common
throughout the range of the toad. The available literature (Kagarise
Sherman and Morton 1993, pp. 186-198; Jennings and Hayes 1994, pp. 50-
53; USFS et al. 2009, pp. 1-133; Martin 2008, pp. i-393) supports the
contention that a combination of factors has interacted and is
responsible for the decline observed in Yosemite toad populations over
the past few decades.
Disease has been documented in Yosemite toad populations, and
recent data documenting historic trends in Bd infection intensity are
compelling (Dodge and Vredenburg 2012, p. 1), but disease has not been
definitively tied to the observed rangewide decline. There is
considerable evidence that various stressors, mediated via impacts to
meadow hydrology following upslope land management practices over the
last century, have detrimentally affected the quantity and quality of
breeding meadows. Many of these stressors, such as grazing, have been
more significant in the past than under current management standards.
However, legacy effects remain, and meadows tend not to recover without
active intervention once excessive stream incision in their watershed
is set in motion (Vankat and Major 1978, pp. 386-397). Certain
stressors may be of concern, such as recreational impacts and avian
predation upon terrestrial life stages of toads, although we do not
have sufficient data to document the magnitude of these particular
stressors.
Given the evidence supporting the role of climate in reducing
populations and potentially leading to the extirpation of many of the
populations studied through the 1970s and into the early 1990s
(Kagarise Sherman and Morton 1993, pp. 186-198), this factor is likely
either a primary driver, or at least a significant contributing factor
in the declines that have been observed. Climate models predict
increasing drought intensity and changes to the hydroperiod based on
reduced snowpack, along with greater climate variability in the future
(PRBO 2011, pp. 18-25). These changes will likely exacerbate stress to
the habitat specialist Yosemite toad through a pronounced impact on its
ephemeral aquatic habitat, and also through an increase in the
frequency of freezing and drying events that kill Yosemite toad eggs
and tadpoles. These changes and the resultant impacts likely will
effectively reduce breeding success of remnant populations already at
low abundance and still in decline. If an interaction such as winter
stress and disease (Carey 1993, pp. 355-362) is the underlying
mechanism for Yosemite toad declines, then the enhanced influence of
climate change as a stressor may tip the balance further towards higher
incidence and increased virulence of disease, which would also lead to
greater population declines and extirpations.
Determination for Yosemite Toad
Section 4 of the Act (16 U.S.C. 1533), and its implementing
regulations at 50 CFR part 424, set forth the procedures for adding
species to the Federal Lists of Endangered and Threatened Wildlife and
Plants. Under section 4(a)(1) of the Act, we may list a species based
on (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 (E) Other natural or manmade factors affecting its
continued existence. Listing actions may be warranted based on any of
the above threat factors, singly or in combination.
We have carefully assessed the best scientific and commercial
information available regarding the past, present, and future threats
to the Yosemite toad. The Yosemite toad is the most narrowly
distributed Sierra Nevada endemic, pond-breeding amphibian (Shaffer et
al. 2000, p. 246). Although it apparently still persists throughout a
large portion of its historical range, it has been reduced to an
estimated 13 percent of historical watersheds. (The proposed rule
indicated that the toad was reduced to an estimated 12 percent of its
range, peer review corrected this number to 13 percent (Brown 2013,
unpaginated). In addition, while the best available data do not provide
information on whether populations are currently stable, or whether
there is a persistent decline, remnant populations are predominantly
small.
Yosemite toad populations are subject to threats from habitat
degradation associated with land uses that negatively influence meadow
hydrology, fostering meadow dewatering, and conifer and other invasive
plant encroachment. These activities include the legacy effects of
historic grazing activities, the fire management regime of the past
century, historic timber management activities, and associated road
construction. The impacts from these threats are cumulatively of
moderate magnitude, and their legacy impacts on meadow habitats act as
a constraint upon extant populations now and are expected to hinder
persistence and recovery into the future. Diseases are threats of
conservation concern that have likely also had an effect on populations
leading to historical population decline, and these threats are
operating currently and will continue to do so into the future, likely
with impacts of moderate-magnitude effects on Yosemite toad
populations.
The individual, interactive, and cumulative effects of these
various risk factors have acted to reduce the geographic extent and
abundance of this species throughout its habitat in the Sierra Nevada.
The combined effect of these stressors acting upon small remnant
populations of Yosemite toads is of significant conservation concern.
The Yosemite toad has a life history and ecology that make it sensitive
to drought and anticipated weather extremes associated with climate
change. Climate change is expected to become increasingly significant
to the Yosemite toad and its habitat in the future throughout its
range. Therefore, climate change represents a threat that has a high
magnitude of impact as an indirect stressor via habitat loss and
degradation, and as a direct stressor via enhanced risk of climate
extremes to all life stages of Yosemite toads.
[[Page 24300]]
The Act defines an endangered species as any species that is ``in
danger of extinction throughout all or a significant portion of its
range'' and a threatened species as any species ``that is likely to
become endangered throughout all or a significant portion of its range
within the foreseeable future.'' We find that the Yosemite toad is
likely to become endangered throughout all or a significant portion of
its range within the foreseeable future, based on the immediacy,
severity, and scope of the threats described above. These include
habitat loss associated with degradation of meadow hydrology following
stream incision consequent to the cumulative effects of historic land
management activities, notably livestock grazing, and also the
anticipated hydrologic effects upon habitat from climate change under
listing Factor A. Additionally, we find that disease under listing
Factor C was likely a contributor to the recent historic decline of the
Yosemite toad, and may remain an important factor limiting recruitment
in remnant populations. We also find that the Yosemite toad is likely
to become endangered through the direct effects of climate change
impacting small remnant populations under Factor E, likely compounded
with the cumulative effect of other threat factors (such as disease).
We have carefully assessed the best scientific and commercial
information available regarding the past, present, and future threats
to the species, and have determined that the Yosemite toad meets the
definition of threatened under the Act, rather than endangered. This
determination is because the impacts from the threats are occurring now
at high and moderate magnitudes, but are all likely to become of high
magnitude in the foreseeable future across the species' entire range,
making the species likely to become in danger of extinction. While
population decline has been widespread, the rate of decline is not so
severe to indicate extinction is imminent, but this rate could increase
as stressors such as climate change impact small remnant populations.
Further, the geographic extent of the species remains rather widespread
throughout its historic range, conferring some measure of ecological
and geographic redundancy. Therefore, on the basis of the best
available scientific and commercial information, we finalize listing
the Yosemite toad as threatened in accordance with sections 3(20) and
4(a)(1) of the Act.
The term ``threatened species'' means any species (or subspecies
or, for vertebrates, distinct population segments) that is likely to
become an endangered species within the foreseeable future throughout
all or a significant portion of its range. The Act does not define the
term ``foreseeable future'' but it likely describes the extent to which
the Service could reasonably rely on predictions about the future in
making determinations about the future conservation status of the
species. In considering the foreseeable future as it relates to the
status of the Yosemite toad, we considered the historical data to
identify any relevant existing trends that might allow for reliable
prediction of the future (in the form of extrapolating the trends). We
also considered how current stressors are affecting the species and
whether we could reliably predict any future trends in those stressors
that might affect the species recognizing that our ability to make
reliable predictions for the future is limited by the quantity and
quality of available data. Thus the foreseeable future includes the
species' response to these stressors and any trends.
Under the Act and our implementing regulations, a species may
warrant listing if it is endangered or threatened throughout all or a
significant portion of its range. The Yosemite toad is highly
restricted in its range, and the threats occur throughout its range.
Therefore, we assessed the status of the species throughout its entire
range. The threats to the survival of the species occur throughout the
species' range and are not restricted to any particular significant
portion of that range, nor are they concentrated in a specific portion
of the range. Accordingly, our assessment and final determination
applies to the species throughout its entire range.
Summary of Comments
In the proposed rule published on April 25, 2013 (78 FR 24472), we
requested that all interested parties submit written comments on the
proposal by June 24, 2013. Given the large number of requests that we
received to extend the public comment period, we reopened the comment
period on July 19, 2013 (78 FR 43122), requesting written comments on
the proposal by November 18, 2013, and again reopened the comment
period on January 10, 2014 (79 FR 1805), with the close of comment
period on March 11, 1014. We also contacted appropriate Federal and
State agencies, scientific experts and organizations, and other
interested parties and invited them to comment on the proposal.
Newspaper notices inviting general public comment were published in the
Sacramento Bee and Bakersfield Californian. We received multiple
requests for a public hearing. We held two public hearings on January
30, 2014, in Sacramento, California. We also held two public
informational meetings, one in Bridgeport, California, on January 8,
2014, and the other in Fresno, California, on January 13, 2014. We also
participated in several public forums, one sponsored by Congressman
McClintock and two sponsored by Congressman LaMalfa. All substantive
information provided during comment periods has either been
incorporated directly into this final determination or addressed below.
Peer Reviewer Comments
In accordance with our peer review policy published on July 1, 1994
(59 FR 34270), we solicited expert opinion from five knowledgeable
individuals with scientific expertise that included familiarity with
the Sierra Nevada yellow-legged frog, the northern DPS of the mountain
yellow-legged frog, the Yosemite toad, and the habitat and biological
needs of, and threats to each species. We received responses from four
of the peer reviewers.
We reviewed all comments received from the peer reviewers for
substantive issues and new information regarding the listing of the
Sierra Nevada yellow-legged frog, the northern DPS of the mountain
yellow-legged frog, and the Yosemite toad. The peer reviewers generally
concurred with our methods and conclusions and provided additional
information, clarifications, and suggestions to improve the final rule.
However, one of the four peer reviewers suggested the rationale for
listing Yosemite toad was poorly supported. Peer reviewer comments are
addressed in the following summary and incorporated into the final
rule.
(1) Comment: Two peer reviewers recommended that we refer to Rana
muscosa as the southern mountain yellow-legged frog in order to reduce
reader confusion in the final rule.
Our Response: We have clarified the common names we are using in
this final rule for each yellow-legged frog species (see Background and
Taxonomy sections in this final rule). While Crother et al. (2008, p.
11) accepted the common name of southern mountain yellow-legged frog
for Rana muscosa, the use of this common name may create additional
confusion as the reader may interpret the name to imply the yellow-
legged frogs in southern California that are already listed as the
southern DPS, rather than the R. muscosa in the Sierra Nevada.
Therefore, we continue to refer to the northern DPS of Rana muscosa as
the northern DPS of the mountain yellow-legged frog, as we did in the
proposed
[[Page 24301]]
rule, to minimize confusion for the public.
(2) Comment: Two peer reviewers suggested that we utilize a
rangewide analysis for listing Rana muscosa and thereby combine the
northern and southern DPSs of the mountain yellow-legged frog into one
listed entity. Clarifying discussions with one peer reviewer suggested
that we not complete a rangewide analysis, but rather keep the DPSs
separate (Knapp, pers. comm.).
Our Response: Given the geographic isolation, different habitat
requirements, differences in threats, and different management needs
between Rana muscosa in the Sierra Nevada compared with southern
California, we have decided to retain the DPS analysis in the proposed
rule and to maintain the northern and southern DPSs of mountain yellow-
legged frog as separate listed entities. Within the Sierra Nevada, R.
muscosa is predominantly found within high-elevation lake habitats that
freeze during the winter months, while in southern California, Rana
muscosa populations occupy stream habitats that are not typically
subject to winter freezing. The differences in the habitats utilized by
the northern and southern DPSs of the mountain yellow-legged frog and
the differences in the threats to each population segment indicate that
management actions needed to recover the northern California and
southern California populations will also be different and are most
expediently addressed separately by DPS (see Distinct Vertebrate
Population Segment Analysis in this final rule).
The factors that are threats to the species also differ between the
two DPSs. We have identified fish stocking and presence of fish as a
threat for both the northern and southern DPSs. However, the other
threats we identified for the northern DPS are primarily habitat
degradation, disease, and climate change, whereas the main threats for
the southern DPS consist of recreational activities, roads, and
wildfire. While there is some overlap in the threats identified for the
two DPSs, the threats that are important to the species status vary
substantially between the Sierra Nevada and southern California.
The differences between the northern and southern DPSs of the
mountain yellow-legged frog in both habitat use and the factors
affecting the species results in differences in the actions and
activities that would be needed to conserve the species in each of the
two DPSs. Conservation planning, including identifying actions and
setting priorities for recovery, will be more effective and better
suited to meet the species' needs if two separate DPSs are retained.
(3) Comment: One peer reviewer indicated that the frogs within the
Spanish and Bean Creek areas of Plumas County (low-elevation areas
within the northern portion of the Sierra Nevada) in which Wengert
(2008) conducted telemetry studies of frog movement distances, may
actually be foothill yellow-legged frog (Rana boylii) rather than
Sierra Nevada yellow-legged frogs (Rana sierrae) (see Habitat and Life
History section in Background for the mountain yellow-legged frogs of
this final rule).
Our Response: We acknowledge and understand some of the challenges
in correctly identifying the species in areas where the ranges of
Sierra Nevada and foothill yellow-legged frogs overlap. Recent genetic
analysis of samples collected from frogs in Spanish and Bean Creeks has
identified the frogs occurring in Bean Creek as both Sierra Nevada and
foothill yellow-legged frogs (Lind et al. 2011a, pp. 281-282), while
Spanish Creek frogs were identified as foothill yellow-legged frog
(Poorten et al. 2013, p. 4). However, given the small sample size,
Poorten et al. (2013, p. 4) suggested that followup investigation was
needed to determine whether Sierra Nevada yellow-legged frogs also
occur in Spanish Creek.
While it is not clear whether Wengert (2008) studied Sierra Nevada
or foothill yellow-legged frogs, given the stream-based ecological
setting of the study, we expect that the movement distances recorded
are applicable to the Sierra Nevada yellow-legged frog within a stream-
based system, as the ecology is comparable between the two sister taxa
in regard to stream systems. Additionally, a study conducted by Fellers
et al. (2013, p. 159) documented Sierra Nevada yellow-legged frog
movement distances up to 1,032 m in a 29-day period, suggesting the
season-long movement distance documented by Wengert (2008, p. 20) is
applicable.
(4) Comment: One peer reviewer provided comment that our proposed
rule did not include more-recent literature on the effects of airborne
contaminants on the mountain yellow legged frog, including Bradford et
al. 2011, which measured contaminant concentrations at multiple sites
in the southern Sierra Nevada and compared their distribution with
population declines of mountain yellow-legged frogs, finding no
association between the two. The peer reviewer further recommended that
we state that frogs are sensitive to contaminants, but measured
contaminant concentrations in multiple media indicate very low
exposures to contaminants from upwind sources.
Our Response: In our proposed rule, we included a discussion of
environmental factors that affect the mountain yellow-legged frog
complex, including contaminants. Based on our analysis in the proposed
rule, we did not identify this environmental factor as a threat to the
species. Upon our review of additional literature, including a study
focused specifically on the mountain yellow-legged frog complex, our
initial discussion remains valid, which indicated that the potential
threat posed by contaminants is not a factor in the listing of this
species. We refer to the proposed rule for the discussion of the
effects of contaminants on the mountain yellow-legged frog.
(5) Comment: One peer reviewer suggested that recent genetic
studies (Shaffer et al. 2000, Stevens 2001, and Goebel et al. 2009) do
not support our conclusion that Yosemite toad is a valid species.
Our Response: When conducting our review of the Yosemite toad as a
listable entity under the Act, we incorporated the results of the
studies mentioned by the peer reviewer. In addition to the previously
included literature on the genetics of Yosemite toad, we have included
in this final rule results from Switzer et al. (2009), which provide
genetic data supporting the Yosemite toad as a valid species. While we
acknowledge that the evolutionary history of the Yosemite toad is
complicated and not fully understood, given our conclusions after
reviewing the taxonomy of the species, and given that the scientific
community as a whole continues to recognize the Yosemite toad as a
valid species, we continue to recognize Yosemite toad as a valid
species (for further discussion, see Taxonomy section above).
(6) Comment: One peer reviewer provided information regarding the
number of localities of Yosemite toad within two National Parks, and
suggested that, had we included these locations, the analysis may have
had a different outcome.
Our Response: When we conducted our analysis for the proposed rule
to determine whether the Yosemite toad warrants listing under the Act,
we utilized the best available scientific and commercial information.
Part of that information included the geospatial data for Yosemite toad
locations within both Yosemite and Sequoia National Parks. These data
were subsequently used for the proposed critical habitat designation.
While we did have (and used) the information on Yosemite toad locations
within the National Parks in
[[Page 24302]]
our analysis, we did not cite to this information into the text of the
proposed rule. This was updated with the data included in Berlow et al.
(2013), as well as information received from Sequoia National Park
staff. Regardless, we utilized the geospatial data in the proposed
rule, determining that the information suggests that the Yosemite toad
has disappeared from approximately 47-69 percent of formerly occupied
sites (Berlow et al. 2013, p. 2). In addition, at many of the remaining
sites, Yosemite toads exist in very low numbers, indicating that many
remaining populations are vulnerable to extirpation. Our use of the
data from both National Forests and National Parks led us to our
proposed status determination, which is affirmed here.
(7) Comment: One peer reviewer stated that there is scant evidence
available to argue that there has been a decline in abundance of the
Yosemite toad and that the difficulty in accurately quantifying toad
abundance, coupled with the fact that the proposed rule did not include
locality data from the National Parks, has weakened the argument for
our determination.
Our Response: While we agree that no studies have documented a
rangewide decline in population abundances in Yosemite toads, and we do
not have sufficient data to conduct a robust trend analysis or detect
negative population growth rates, we relied on published literature for
our determination. At a minimum, the published literature provides
anecdotally documented declines in numbers of individual Yosemite toads
at the respective study sites. The best available information shows
that the Yosemite toad populations have declined, and that the remnant
populations comprise low numbers of individual adult toads. For our
analysis, we did utilize the data on toad locations in the National
Parks (see our response to comment 6) and included it as part of our
analysis on the estimated loss of historically occupied sites (47-69
percent of formerly occupied sites (Berlow et al. 2013, p. 2)). We
mainly focused our analysis on the potential drivers of population
stability and identified the predominate threats to the species as the
continuing effects of degradation of meadow hydrology associated with
historical land management practices and the effects of climate change
and anthropogenic stressors acting on the small remnant populations.
(For complete discussion see Summary of Factors Affecting the Species
section above.)
(8) Comment: One peer reviewer stated that there are scientific
uncertainties regarding the long-term population trends and threats to
Yosemite toad and that these uncertainties should be explicitly
described.
Our response: As required by the Act, we based our proposed rule
and this final rule on the best available scientific and commercial
data. While there are some uncertainties in the information, we clearly
articulated these uncertainties when conducting our analysis for the
rule. (See Population Estimate and Status and Meadow Habitat Loss and
Degradation sections for examples.)
Federal Agency Comments
(9) Comment: The Forest Service suggested that the rule does not
represent the best available scientific and commercial information in
proposing a determination.
Our Response: In conducting our analysis, we rely on the best
available scientific and commercial information, as required by the
Act. On occasion, we are not aware of certain information that is
available at the time we issue a proposed rule or new information
becomes available around the time of publication, which is part of the
reason we request public comment, as well as peer review. That portion
of the process helps to inform our final decision by soliciting input
and seeking additional available information. As a result of this
process, we have received new scientific and commercial information
that we have reviewed and incorporated into this final rule.
(10) Comment: The USFS noted that the proposed rule did not
identify mining activities as a threat to the mountain yellow-legged
frog.
Our Response: We acknowledge that there is some overlap between
current mining activities and areas occupied by the mountain yellow-
legged frogs, particularly in the northern part of the range; however,
we do not have information to assess the impact that mining has on the
species in those areas where mining occurs, and how it acts as either
an historical or current threat to the species. Within designated
wilderness, new mining claims have been prohibited since January 1,
1984. Additionally, while suction dredge mining may have the potential
to alter microhabitat uses by the species, the current moratorium on
this practice removes this potential threat. However, we acknowledge
that this situation may change in the future.
(11) Comment: The USFS suggested that the uncertainties we
presented under Factor D as it relates to their Forest Plan revision
process and protections for mountain yellow-legged frog are not
applicable and that the protections under the SNFPA will continue as a
result of consultation with the Service.
Our Response: We did not identify Factor D as a threat to the
mountain yellow-legged frog, and we incorporated an analysis of the
protection that the current Forest Plans offer the species. While there
is some uncertainty as to whether these protections will remain in the
revised Forest Plans, the USFS is not required to consult with the
Service on the Sierra Nevada yellow-legged frog and northern DPS of the
mountain yellow-legged frog in the absence of the protections afforded
under the Act. As such, we must evaluate the adequacy of existing
regulatory mechanisms from the baseline of the species not being
federally listed under the Act.
(12) Comment: The USFS suggested the final rule include a
discussion of the impacts of bullfrog predation on the mountain yellow-
legged frog.
Our Response: We have limited information on the presence of
bullfrogs in the Sierra Nevada, but we have included a section on the
potential threat of American bullfrogs where they are known to occur in
the Lake Tahoe Basin (see discussion under Factor C for mountain
yellow-legged frogs).
(13) Comment: The USFS and several other commenters suggested that
the information presented as it relates to the impacts of grazing on
Yosemite toad was inaccurate. Specifically, they suggested that we did
not include the results of peer-reviewed journal articles in our
analysis of the impacts posed by livestock grazing.
Our Response: At the time of the proposed rule, we were aware of
the peer-reviewed literature related to the impacts of livestock
grazing on Yosemite toad, and inadvertently omitted the literature from
the rule. We have reviewed and included the relevant articles in this
final rule. Additionally, while we did not incorporate all of the
specifics of the journal articles, we did incorporate the results of a
5-year study that investigated the impacts of cattle grazing on
Yosemite toad in our analysis, as they were presented in Allen Diaz et
al. 2010, and subsequently in the Lind et al. (2011b, addendum).
(14) Comment: The USFS and several other commenters suggested that
our reliance on a single non-peer-reviewed study to assess the impacts
of cattle grazing on Yosemite toads, through direct mortality or the
modification of their habitat, was inappropriate. Additionally, they
suggested we discounted the peer-reviewed published
[[Page 24303]]
journal articles related to the impacts of cattle grazing on Yosemite
toad.
Our Response: In conducting our analysis, we rely on the best
available scientific and commercial information, as required by the
Act. This information does not need to be specifically published in a
scientific journal. The Martin (2008) study that is being referred to
by the commenters is a doctoral dissertation that was, in fact,
reviewed prior to release. We relied on the information presented by
Martin in assessing the potential for direct mortality of Yosemite toad
that is attributed to livestock. We also relied on Martin for the
potential impacts of livestock grazing on overwintering and upland
areas utilized by Yosemite toad, as the peer-reviewed publications that
the commenters referred to were based on a study that only assessed
grazing effects on breeding. As such, the best available scientific and
commercial information includes Martin (2008). In our proposed rule, we
evaluated the information that ran contrary to Martin (2008), and we
have subsequently incorporated the information presented in the peer-
reviewed journal articles in this final rule. Please also see response
to comment 13.
(15) Comment: The USFS commented that chytrid fungus, fish
stocking, and climate change pose the greatest threats to the mountain
yellow-legged frogs, and that threats from authorized management
activities are insignificant threats to the species.
Our Response: We have concluded in this final rule that, in
general, authorized activities on public lands managed by the USFS and
the NPS are not significant threats to the mountain yellow-legged
frogs, but we also recognize that there may be limited site-specific
conditions where authorized activities could have population-level
effects, especially where populations are small or habitat areas are
limited (see Summary of Factors Affecting the Species in this final
rule).
(16) Comment: The USFS noted that recent publications indicate that
livestock grazing that meets current USFS standards and guidelines is
less of a threat to the Yosemite toad than was described in the
proposed rule.
Our Response: We have revised our discussion of grazing in this
final rule to clarify the conditions under which we consider current
grazing activities to pose habitat-related threats to the Yosemite toad
(see Summary of Changes and Factor A discussion for the Yosemite toad).
Comments From States
(17) Comment: The California Department of Fish and Wildlife (CDFW)
originally commented that the threats presented in the proposed rule
suggested that a determination of threatened status would be more
appropriate than endangered for the Sierra Nevada yellow-legged frog.
However, CDFW reconsidered this suggestion after discussions with
Service staff and submitted a followup comment letter that agrees with
the Service determination and supports listing the Sierra Nevada
yellow-legged frog as endangered.
Our Response: We find that an endangered status for the Sierra
Nevada yellow-legged frog is an appropriate determination and
appreciate CDFW's reconsideration of their initial comments.
(18) Comment: CDFW commented that they remain concerned that
listing the species as endangered could hinder timely implementation of
the Department's recovery and restoration efforts for the species
pursuant to its State-listing under CESA. CDFW notes that they have a
responsibility to continue activities and expand efforts that will
contribute to the recovery of the Sierra Nevada yellow-legged frog and
hope that such efforts can be fostered through the 1991 Cooperative
Agreement between the California Department of Fish and Game and the
U.S. Fish and Wildlife Service. They also comment that, in his June 13,
2012, memo to the Service's Regional Directors, the Director of the
Fish and Wildlife Service acknowledged the Federal-State collaborative
nature of conservation activities for listed species.
Our Response: We note that, for research activities that aid in the
recovery of the species, and that may result in take, a permit issued
under section 10a(1)A of the Act is the appropriate mechanism. However,
our regulations at 50 CFR 17.21 state that any qualified employee or
agent who is designated by CDFW for such purposes, may, when acting in
the course of his official duties, take endangered wildlife species
covered by a Cooperative Agreement (developed pursuant to Section 6 of
the Act) between the Service and the State provided such take is not
reasonably anticipated to result in: (1) The death or permanent
disabling of the specimen; (2) the removal of the specimen from the
State of California; (3) the introduction of the specimen or any of its
progeny into an area beyond the historical range of the species; or (4)
the holding of the specimen in captivity for a period of more than 45
days. Take that does not meet these four conditions would require a
section 10(a)(1)(A) permit. We acknowledge and appreciate the important
role that CDFW will play in the recovery of the Sierra Nevada yellow-
legged frog, and look forward to continuing collaborative conservation
actions with CDFW for this and other listed species in California.
(19) Comment: CDFW agreed that we should retain the northern DPS
and the southern DPS designations for the mountain yellow-legged frog
(Rana muscosa). They provided updates to our discussion of take related
to State-listing of the mountain yellow-legged frog complex.
Our Response: We appreciate the support, and we have retained the
two DPSs in the final determination (see Distinct Vertebrate Population
Segment Analysis). We have also revised our discussion of CESA to
provide the updated information on take related to State-listing of the
mountain yellow-legged frog complex (see Factor D for mountain yellow-
legged frog).
(20) Comment: CDFW provided comments on our discussion of the
following threats to the mountain yellow-legged frog complex:
Recreational activities, past trout stocking versus continued trout
stocking, and pesticide detection in the Sierra Nevada. They commented
that the evidence presented in the Recreation section did not support
the conclusion, urging us to readdress the section and remove claims
unsupported by appropriate citations, and noted that recreation effects
to the environment were supported, but no evidence indicates that such
activities affect the frog populations. In the Recreation section, they
also noted several errors and inaccuracies in citing other authors.
CDFW provided extensive comments on our discussion of dams and water
diversions, commenting that they were of the opinion that dams and
diversion posed a threat of low significance to the continued existence
of the mountain yellow-legged frogs and suggesting that the section
required significant amendments to accurately capture the degree of
potential impacts. They noted that most dams were constructed below the
range of extant frog populations, and that some information was
misapplied from research on lower-elevation amphibian species, such as
the foothill yellow-legged frog, which resulted in overstatement of the
potential impact of dams and water diversions on the mountain yellow-
legged frog complex. They provided numerous smaller specific comments
on text within the section.
[[Page 24304]]
Our Response: We thank the CDFW for the additional information
provided to strengthen our analysis. We have addressed these comments
through changes to the Fish Stocking, Recreation, and Dams and Water
Diversions sections for the Sierra Nevada and mountain yellow-legged
frogs in this final rule. We re-checked references and revised the
sections noted to state more clearly the potential effects of these
activities, to rely on appropriate citations, and to refine our
conclusions in agreement with CDFW's comments. Please see Factor A in
Summary of Factors Affecting the Species for updated information.
Public Comments
(21) Comment: Several commenters suggested that the Service does
not have the authority or jurisdiction to designate the Sierra Nevada
yellow-legged frog and the northern DPS of the mountain yellow-legged
frog as endangered nor the Yosemite toad as threatened.
Our Response: The authority for the Service to issue this
rulemaking comes from the Endangered Species Act of 1973 (16 U.S.C.
1531 et seq.), as amended, through the 108th Congress. The Service is
designated as the lead Federal agency for implementing the Act for
terrestrial and freshwater species. Authority to implement the Act does
not require Federal jurisdiction or land ownership
(22) Comment: Multiple commenters indicated that existing Federal
and State legislation and regulations, such as the Wilderness Act,
CESA, and CDFW regulations, provide sufficient protection for these
amphibians, and thereby eliminate the need for listing the species.
Our Response: We agree that existing Federal and State legislation
and regulations, such as the Wilderness Act, CESA, and CDFW regulations
provide some protection for the Sierra Nevada yellow-legged frog, the
northern DPS of the mountain yellow-legged frog, and the Yosemite toad.
However, while existing legislation and regulations provide some level
of protection for the Sierra Nevada yellow-legged frog, the northern
DPS of the mountain yellow-legged frog, and the Yosemite toad, they do
not require that Federal agencies ensure that actions that they fund,
authorize, or carry out will not likely jeopardize the species'
continued existence (for further information see discussions under
Factor D). Therefore, we have determined that the Sierra Nevada yellow-
legged frog and the northern DPS of the mountain yellow-legged frog are
endangered and that the Yosemite toad is threatened under the Act.
(23) Comment: Several commenters suggest that it is necessary for
the Service to conduct an analysis of the impacts that listing a
species may have on local economies prior to issuance of a final rule.
Our Response: Under the Act, the Service is not required to conduct
an analysis regarding the economic impact of listing endangered or
threatened species. However, the Act does require that the Service
consider the economic impacts of a designation of critical habitat. A
draft of this analysis is available to the public on https://www.regulations.gov (79 FR 1805).
(24) Comment: Several commenters suggested that the decline of the
Sierra Nevada yellow-legged frog, northern DPS of the mountain yellow-
legged frog, and the Yosemite toad is a natural evolutionary process,
and that the presence of environmental stressors is a normal driver of
evolution and/or extinction.
Our Response: Under the Act, we are required to use the best
available scientific and commercial information to assess the factors
affecting a species in order to make a status determination. The Act
requires the Service to consider all threats and impacts that may be
responsible for declines as potential listing factors. The evidence
presented suggests that the threats to the species are both natural and
manmade (see Factor E--Other Natural or Manmade Factors Affecting the
Species), but that they are primarily the result of anthropogenic
influences (see Summary of Factors Affecting the Species in this final
rule). Thus, the threats associated with the declines of these species
are not part of a natural evolutionary process.
(25) Comment: Several commenters were concerned about the effects
of listing on mining and associated activities conducted under the
General Mining Law of 1872. They suggested that the listing of these
species will remove 5 million acres from mining and other productive
uses of the land. One commenter was concerned that there would be no
assurances that development of a mining claim will result in the
ability to mine it.
Our Response: In the proposed rule, we identified unauthorized
discharge of chemicals or fill material into any water upon which the
Sierra Nevada yellow-legged frog, the northern DPS of the mountain
yellow-legged frog, and the Yosemite toad are known to occur as a
potential threat to these species. On National Forests outside of
designated wilderness, new mining may occur pursuant to the Mining Law
of 1872 (30 U.S.C. 21 et seq.), which was enacted to promote
exploration and development of domestic mineral resources, as well as
the settlement of the western United States. It permits U.S. citizens
and businesses to prospect hardrock (locatable) minerals and, if a
valuable deposit is found, file a claim giving them the right to use
the land for mining activities and sell the minerals extracted, without
having to pay the Federal Government any holding fees or royalties (GAO
1989, p. 2). Gold and other minerals are frequently mined as locatable
minerals, and, as such, mining is subject to the Mining Law of 1872.
However, Federal wilderness areas were closed to new mining claims at
the beginning of 1984 (see Factor D under mountain yellow-legged frogs
above), thereby precluding the filing of new mining claims in those
areas designated as Federal wilderness (a large part of the area in
which the species occur). Authorization of mining under the Mining Law
of 1872 is a discretionary agency action pursuant to section 7 of the
Act. Therefore, Federal agencies with jurisdiction over land where
mining occurs will review mining and other actions that they fund,
authorize, or carry out to determine if listed species may be affected
in accordance with section 7 of the Act.
(26) Comment: Numerous commenters suggested that the listing of the
Sierra Nevada yellow-legged frog, the northern DPS of the mountain
yellow-legged frog, and the Yosemite toad are being misused to restrict
or prohibit access for fishing, hiking, camping, and other recreational
uses, and implement land use restrictions, management requirements, and
personal liabilities on the public that are not prudent, clearly
defined, or necessary.
Our Response: The listing of the Sierra Nevada yellow-legged frog,
the northern DPS of the mountain yellow-legged frog, and the Yosemite
toad does not prevent access to any land, whether private, tribal,
State, or Federal. The listing of a species does not affect land
ownership or establish a refuge, wilderness, reserve, or other
conservation area. A listing does not allow the government or public to
access private lands without the permission of the landowner. It does
not require implementation of restoration, recovery, or enhancement
measures by non-Federal landowners. Federal agencies will review
actions that they fund, authorize, or carry out to determine if any of
these three amphibians, and other listed species as appropriate, may be
affected by the Federal action. The Federal agency will
[[Page 24305]]
consult with the Service, in accordance with Section 7 of the Act (see
also response to comment 25).
(27) Comment: Several commenters suggested that listing the Sierra
Nevada yellow-legged frog and the northern DPS of the mountain yellow-
legged frog under the Act is not necessary given that a majority of the
range of these species is within wilderness areas afforded protection
under the Wilderness Act and by the protections afforded under CESA.
Our Response: We agree that existing Federal and State legislation
and regulations, such as the Wilderness Act and CESA, provide some
protection for the Sierra Nevada mountain yellow-legged frog, the
northern DPS of the mountain yellow-legged frog, and the Yosemite toad.
However, we identified the main threats to the two frog species as
habitat degradation and fragmentation, predation and disease, climate
change, and the interactions of these stressors on small populations.
Neither the Wilderness Act nor the State's listing status under CESA
ameliorates these threats to levels that would preclude the need to
list the species under the Act. (See discussion under Factor D).
(28) Comment: One commenter suggested that habitat and range of the
mountain yellow-legged frog is not threatened with destruction or
modification based on a large portion being located in wilderness, and
the proposed rule stating ``physical habitat destruction does not
appear to be the primary factor associated with the decline of the
mountain yellow-legged frogs.''
Our Response: While we agree that the loss, destruction, or
conversion of physical habitat is not a primary factor in the decline
of the mountain yellow-legged frogs, we discuss both the biological
modification of habitat due to changes in predator communities, prey
communities, and in nutrient levels, and due to the habitat
fragmentation associated with the presence of introduced fish. Although
the presence of introduced fish does not result in conversion or loss
of the physical attributes of habitat (for example, removal or filling
of lakes, ponds, etc.), fish presence does effectively preclude the use
of the habitat by the mountain yellow-legged frog (see our discussion
under Factor A). While a large portion of the range of the mountain
yellow-legged frog is within federally designated wilderness, or on
National Parks, we identified the main threats to the species as
habitat degradation and fragmentation, predation and disease, climate
change, and the interactions of these stressors on small populations.
Neither the Wilderness Act nor the protections afforded within National
Parks ameliorates these threats to levels that would preclude the need
to list the species under the Act (see discussion under Factor D).
(29) Comment: One commenter stated that we failed to consider the
effectiveness of restoration activities being conducted by CDFW as part
of their High Mountain Lakes Project and plans for Yosemite and Sequoia
and Kings National Parks that are intended to implement restoration
actions.
Our Response: We are aware of the activities, including the High
Mountain Lakes Project (see Factor A discussions above in this final
rule), being conducted by CDFW, USFS, NPS, and researchers aimed at
restoring habitat for the mountain yellow-legged frog. While efforts of
interested parties have resulted in the restoration of habitat for
these species, the restored habitat represents a small portion of the
range of the species, and has occurred only in localized areas. As
such, these activities, while beneficial and important for the recovery
of the species, do not significantly counter the threats of introduced
predators, disease, or climate change. Additionally, we are aware of
planning efforts by Yosemite and Sequoia and Kings National Parks,
partially implemented, and we are aware that these restoration plans
have not been finalized.
(30) Comment: One commenter provided information suggesting
livestock are responsible for the transportation of Bd in the
environment.
Our Response: While livestock may provide a vector for the
transmission of amphibian disease within the Sierra Nevada, there are
numerous other mechanisms of transport, including wildlife, as well as
anthropogenic vectors. Since the importance of differing disease
vectors related to Bd is poorly understood, we did not include a
discussion of disease transport associated with livestock grazing in
this rule (see Factor C for discussion of disease).
(31) Comment: One commenter provided information to suggest that
activities associated with illicit cultivation of marijuana on National
Forest System lands should be identified as a potential threat to the
mountain yellow-legged frog.
Our Response: We agree that aspects associated with illegal
cultivation of marijuana on National Forest System lands may pose a
risk to the mountain yellow-legged frogs, such as dewatering of
habitats and contamination from pesticides and fertilizers. There is
potential overlap with this illegal activity and areas occupied by
mountain yellow-legged frogs; however, not enough information is
available at this point to assess the impact that illegal cultivation
of marijuana has on the species.
(32) Comment: Several commenters suggest that there is insufficient
evidence to make a listing determination for the mountain yellow-legged
frog in accordance with the Act.
Our Response: As we have presented in both the proposed rule and
this final rule, a substantial compilation of scientific and commercial
information is available to support listing both the Sierra Nevada
yellow-legged frog and the northern DPS of the mountain yellow-legged
frog under the Act. We have presented evidence that there has been a
curtailment in range and numbers attributed to habitat degradation and
fragmentation under Factor A, predation and disease under Factor C, and
climate change and the interaction of these various stressors
cumulatively impacting small remnant populations under Factor E (see
Determination for the Sierra Nevada Yellow-legged Frog and
Determination for the Northern DPS of the Mountain Yellow-legged Frog
sections above for a synopsis and see the Summary of Factors Affecting
the Species for a detailed analysis).
(33) Comment: Numerous commenters purported that the greatest
threat to the mountain yellow-legged frog is Bd, and since listing the
species will not alleviate the threat, the species should not be
listed. Additionally, it was suggested that these species should be
reared in captivity until the threat of Bd is resolved.
Our Response: We agree that Bd is one of the primary contributing
factors in the current decline of these species; however, it is not the
only factor responsible for their decline or the only one forming the
basis of our determination. All Factors are considered when making a
listing determination (see the Summary of Factors Affecting the Species
for a detailed discussion). We have also identified habitat
fragmentation and predation attributed to the introduction of fish and
climate change as threats to the species. We are required to evaluate
all the threats affecting a species, including disease under Factor C.
With respect to the prospect of captive breeding, we acknowledge
that this activity is one of the suite of tools that can be utilized
for the conservation of the species. Captive breeding is currently
being conducted for the southern DPS of the mountain yellow-
[[Page 24306]]
legged frog, and we are currently working with various facilities to
explore this option. Additionally, when a species is listed as either
endangered or threatened, the Act provides many tools to advance the
conservation of listed species; available tools including recovery
planning under section 4 of the Act, interagency cooperation and
consultation under section 7 of the Act, and grants to the States under
section 6 of the Act. All of these mechanisms assist in the
conservation of the species.
(34) Comment: Several commenters provided information to suggest
that livestock grazing is not detrimental to amphibian species and that
the proposed rule did not adequately capture the neutral or beneficial
effects of livestock grazing on amphibian species.
Our Response: We have revised our discussion of grazing in this
final rule to clarify the conditions under which we consider current
grazing activities to pose habitat-related threats (see Factor A
above). In addition, research with a related ranid frog of western
montane environments, (the Columbia spotted frog, Rana luteiventris)
has indicated that livestock grazing may reduce vegetation levels in
riparian and wet meadow habitat, but does not have short-term effects
on the frog populations, although they caution that the length of the
study may not capture potential long-term effects (Adams et al. 2009,
pp. 132, 137). However, George et al. (2011, pp. 216, 232) in a review
of the effectiveness of management actions on riparian areas, noted
that continuous grazing often results in heavy grazing use of riparian
areas, even if an area is lightly stocked, because livestock are
attracted to the areas from adjacent uplands. They note substantial
literature that documents that livestock grazing could damage riparian
areas, and the resulting move, beginning in the 1980s, in Federal and
State resource agencies to apply conservation practices to protecting
and improving riparian habitats (George et al. 2011, p. 217). They note
that studies provide sufficient evidence that riparian grazing
management that maintains or enhances key vegetation attributes will
enhance stream channel and riparian soil stability, although variable
biotic and abiotic conditions can have site-specific effects on results
(George et al. 2011, pp. 217-227).
In our proposed rule, we focused on livestock grazing as a
potential listing factor, and while there are potentially some current,
localized effects to the Sierra Nevada yellow-legged frog, the northern
DPS of the mountain yellow-legged frog, and the Yosemite toad, we
consider the majority of the impacts associated with livestock grazing
are the legacy effects of historically high grazing intensities.
(35) Comment: One commenter stated that the discussion of the
effects of global climate change in the proposed rule for the Sierra
Nevada yellow-legged frog, northern DPS of the mountain yellow-legged
frog, and Yosemite toad was not appropriate. The commenter believed
that the Service ``pushes'' the climate models, both spatially and
temporally, beyond what the commenter considered to be reliable, and
ignores their uncertainty. In addition, the commenter claims that no
credible models can project potential climate change in the Sierra
Nevada. The commenter stated the Act is not an appropriate mechanism to
regulate global climate change and greenhouse gases. Finally, the
commenter suggested if the Service does list the three amphibians, that
they be designated as threatened species with a section 4(d) rule that
excludes lawful greenhouse gases from the prohibitions of the Act.
Our Response: We used the best available scientific and commercial
information available as it pertains to climate change. In addition to
the peer-reviewed scientific journal articles and reports that were
utilized in our analysis and cited in the proposed rule, recently
published studies have presented data and conclusions that increase the
level of confidence that global climate change is the result of
anthropogenic actions (summarized in Blaustein et al. 2010 and
discussed above). A recent paper (Kadir et al. 2013) provides specific
information on the effects of climate change in the Sierra Nevada and
is discussed above. While the Service is concerned about the effects of
global climate change on listed species, wildlife, and their habitats,
to date, we have not used the Act to regulate greenhouse gases. We
evaluated the suggestion that the three amphibians be listed as
threatened species with a section 4(d) rule excluding prohibitions or
restrictions on greenhouse gases. However, our determination is that
the Sierra Nevada yellow-legged frog and the northern DPS of the
mountain yellow-legged frog meet the definition of endangered, the
Yosemite toad meets the definition of threatened, and a section 4(d)
rule for greenhouse gases is not appropriate.
(36) Comment: One commenter suggested that the discussion of
genetics for the mountain yellow-legged frog does not support the
taxonomy of the Sierra Nevada yellow-legged frog and the northern DPS
of the mountain yellow-legged frog as separate species. The commenter
further suggested the text of the rule specifying two major genetic
lineages and four groups does not support listing of the frogs as
separate genetic groups.
Our Response: Vredenburg et al. (2007, p. 317) did not rely solely
on DNA evidence in the recognition of two distinct species of mountain
yellow-legged frog in the Sierra Nevada, but instead used a combination
of DNA evidence, morphological information, and acoustic studies. The
taxonomy of the mountain yellow-legged frogs as two distinct species in
the Sierras has been widely accepted in the scientific community and by
species experts. We are not listing a subspecies but rather two
separate, recognized species, the Sierra Nevada yellow-legged frog and
the northern DPS of the mountain yellow-legged frog.
(37) Comment: Several commenters suggested that activities such as
timber harvest, road construction, recreation, and livestock grazing
are in decline in the Sierras compared with historical levels and
should not be included as potential threats to the Sierra Nevada
yellow-legged frog, the northern DPS of the mountain yellow-legged
frog, or the Yosemite toad.
Our Response: In conducting our analysis of the factors affecting
the species, we did include timber harvest, road construction,
recreation, and livestock grazing, as potential threats to the species,
but acknowledge that the major impact on the species was the result of
the legacy effects of historical practices, and that these activities
currently pose a lower intensity, localized threat. We have attempted
to clarify the distinction in this final rule (see Factor A discussions
above).
(38) Comment: Numerous commenters stated that listing the mountain
yellow-legged frogs and the Yosemite toad would prevent fuels-reduction
activities, leading to fires and loss of habitat.
Our Response: In this final rule under Factor A for the mountain
yellow-legged frogs and Yosemite toad, we address potential habitat
changes that may be related to timber harvest activities, including
harvests for fuels reduction purposes. We found that most populations
of the three species occur at high elevations above areas where timber
harvests are likely. At lower elevations, forest standards and
guidelines would be expected to limit potential threats to the species
in most cases, although limited site-specific situations might result
in habitat effects with population consequences. We also found that
changed fire regimes have, in some of the same lower elevation areas,
[[Page 24307]]
led to an increased potential for high-intensity fires, which could
alter habitat and, therefore, pose relatively localized population-
level effects to the species. For the Yosemite toad, we found that
although ground-disturbance due to timber harvest activities has the
potential to have population-level effects at lower elevations,
especially where habitat is limited, currently the best available
information indicates toads might achieve long-term benefits from
activities that reduce encroachment of trees into breeding sites.
Therefore, we expect that fuels-reduction activities in lower elevation
areas will be generally beneficial to these species.
(39) Comment: A number of commenters suggested that, given the
results of more-recent studies that were not included in the proposed
rule, livestock grazing should be removed as a threat to the Yosemite
toad (See also comment 13 from the USFS).
Our Response: In our proposed rule, we addressed the potential
impacts of grazing on Yosemite toad based on Allen-Diaz et al. (2010).
The more-recent studies referenced (such as Roche et al. 2012a and
2012b, and McIlroy et al. 2013) are different publications but are
based on the results of the companion studies whose initial report, and
subsequent addendum, we referenced as Allen-Diaz et al. (2010) and Lind
et al. (2011b). The study conducted determined that livestock grazing
in accordance with the USFS's standards and guidelines does not affect
Yosemite toad breeding success. While appropriately managed levels of
grazing do not impact breeding success, these grazing standards are not
always met. Additionally, the main impact of grazing on Yosemite toad
is due to the legacy effects of historical grazing intensities on
Yosemite toad habitat. Given the limitations of the study (see
discussion under Factor A) and the documentation that these standards
are not always met, livestock grazing may continue to pose a localized
threat to the species.
(40) Comment: One commenter provided several comments suggesting
that livestock grazing is not a threat to Yosemite toad in light of the
results of a current study, the documentation of Yosemite toads
existing in areas that have been subject to grazing for centuries, and
because the population declines cited in our proposed rule occurred in
an area not subject to grazing.
Our Response: See response to comments 13, 14, and 39. In our
proposed rule, we identified the impacts of livestock grazing primarily
from an historical context as a potential contributor to meadow
degradation. There is a great deal of information, while not specific
to Yosemite toad, on the negative impacts of high-intensity grazing
regimes on ecosystem dynamics. Grazing under current Forest Service
standards does not appear to impact Yosemite toad breeding, however
when inappropriate levels of grazing do occur, grazing may still
present a localized impact on Yosemite toads via direct mortality or
through practices that prevent the hydrologic recovery of historically
wet meadow systems. While the documented declines of Yosemite toad have
occurred in areas that are not currently subject to livestock grazing,
historical grazing occurred throughout the Sierra Nevada. We did not
implicate livestock grazing in the decline in population sizes, rather
as a potential historical driver in meadow degradation rangewide. We
have clarified this distinction in the final rule (see Factor A
discussion and Summary of Factors Affecting the Species for the
Yosemite toad).
(41) Comment: One commenter suggested that livestock grazing
continues to provide a threat to the Sierra Nevada yellow-legged frog
and Yosemite toad and provided information documenting habitat
degradation attributed to current livestock grazing and utilization
above the standards of the SNFPA.
Our Response: As we have presented in the proposed and final rules,
the impact of livestock grazing on these species is primarily one of
historical significance, with the potential for future localized
impacts to the species and/or their habitat. Based on the information
provided regarding habitat conditions and potential impacts to habitat,
we have maintained our position that current livestock grazing poses a
localized impact to the mountain yellow-legged frogs and a prevalent
threat with moderate impacts to the Yosemite toad.
(42) Comment: One party commented that we have not demonstrated
that the Sierra Nevada population of the mountain yellow-legged frog is
a DPS. They indicate that we have not shown that the population is
significant to the taxon as a whole because we have not shown whether
other populations of the species could persist in the high-elevation
Sierra Nevada portion of the species' range or discussed how the Sierra
Nevada populations are adapted to the area. In addition, they indicate
that we failed to show that extirpation of the northern population
would result in a significant gap in the range of the species, and we
did not show that the populations had markedly different genetics
characteristics.
Our Response: The commenters correctly noted that, to recognize a
population of a species as a DPS, we must establish that the population
is (1) discrete from the remainder of the populations to which the
species belongs, and (2) if determined to be discrete, it is also found
to be significant to the species to which it belongs. However, the
commenters incorrectly conclude that the population must meet all three
criteria for significance. We find the northern population of the
mountain yellow-legged frog to be discrete from the southern population
because it is separated from the southern frogs by a 225-km (140-mi)
barrier of unsuitable habitat. The primary basis for our finding that
the northern population is significant to the species as a whole is
that loss of the northern population would mean the loss of the species
from a large portion of its range and reduce the species to small
isolated occurrences in southern California. The population also meets
two additional criteria for significance: (1) Evidence of the
persistence of the discrete population segment in an ecological setting
unusual or unique for the taxon, and (2) evidence that the discrete
population segment differs markedly from the remainder of the species
in its genetic characteristics. We have revised the language in our DPS
analysis to clarify the basis for the determination (see Distinct
Vertebrate Population Segment Analysis).
(43) Comment: Numerous commenters commented that we were required
to complete a NEPA analysis of the proposed listing.
Our Response: We have determined that environmental assessments and
environmental impact statements, as defined under the authority of the
National Environmental Policy Act (NEPA; 42 U.S.C. 4321 et seq.), need
not be prepared in connection with listing a species as an endangered
or threatened species under the Endangered Species Act. We published a
notice outlining our reasons for this determination in the Federal
Register on October 25, 1983 (48 FR 49244) (see Required Determinations
section of this rule).
(44) Comment: One commenter asked that, if we determine that the
three amphibian species under consideration are endangered or
threatened under the Act, then we enter into a cooperative agreement
with the State of California under section 6 of the Act.
Our Response: We have been operating under such a cooperative
agreement with the California
[[Page 24308]]
Department of Fish and Game (now Department of Fish and Wildlife
(CDFW)) since 1991. https://www.dfg.ca.gov/wildlife/nongame/publications/docs/CDFGCooperativeAgreementWithUSFWS.pdf
(45) Comment: One commenter stated that if the three amphibians
considered are listed as threatened or endangered, then research should
continue into the causes of population decline.
Our Response: We expect research on these issues to continue into
the future. Once the three amphibians are listed as threatened or
endangered species under the Act, additional funding for research and
other conservation programs for those species will become available
through grants established under section 6 of the Act. Such grants are
provided to State agencies with which we have established cooperative
agreements.
(46) Comment: One commenter indicated that because of a County
resolution, we must coordinate with the board of supervisors of that
County prior to publishing a final rule.
Our Response: We provide all interested parties an equal
opportunity to submit comments or information prior to publication of a
final rule, and we give equal consideration to all such information and
comments, regardless of source. Our requirements for ``coordination,''
however, are established by the Act, by other Federal statutes such as
the Administrative Procedure Act, and by executive order.
(47) Comment: Several commenters asked for additional time to
provide comments. One commenter added that we provided little public
outreach.
Our Response: As discussed in the first paragraph of the Summary of
Comments and Recommendations section (above), we provided two
additional public comment periods for a total of 240 days
(approximately 8 months) of public comment. We also hosted two public
hearings and two public informational meetings at various locations
within the range of the species under consideration. We also attended
two additional public meetings hosted by Congressmen representing
districts within the range of the species. We contacted and sought
input from appropriate Federal and State agencies, scientific experts
and organizations, and other interested parties. We also published
notices in the newspapers with the largest readerships within both the
northern and southern portions of the ranges of the species. Additional
public comment periods or outreach were not feasible given limitations
imposed by available funds and requirements imposed by the Act
regarding available time in which to publish a final rule.
(48) Comment: One commenter noted that the Act authorizes the
Secretary to extend the time available for publication of a final rule
by up to 6 months if ``there is substantial disagreement regarding the
sufficiency or accuracy of the available data.'' The commenter stated
that such substantial disagreement does exist and so requested that the
available time be extended by 6 months. Specifically, the commenter
indicated that the available data are not sufficient to support listing
after taking into account various Federal and State statutes and
programs currently benefiting the three species. Such statutes and
programs include the Wilderness Act, the Sierra Nevada Forest Plan, the
Clean Water Act, the California Endangered Species Act, and the
discontinuation of fish stocking by CDFW in much of the range of the
two frogs.
Our Response: While we agree that these efforts aid in the
conservation of the three amphibians, we do not consider substantial
disagreement to exist regarding our conclusion that the Sierra Nevada
yellow-legged frog and the northern DPS of the mountain yellow-legged
frog meet the definition of ``endangered species'' under the Act. We
considered the existing Federal and State statutes and programs in our
determination. The data documenting population declines and
extirpations associated with Bd and the presence of introduced fish are
sufficient for the Service to determine that the two species are ``in
danger of extinction throughout all or a significant portion of [their]
range[s].'' Data also show that the Yosemite toad is vulnerable to
habitat changes and climate change, and thus merits listing as a
threatened species, which is defined as ``likely to become an
endangered species within the foreseeable future within all or a
significant portion of its range.''
Available Conservation Measures
Conservation measures provided to species listed as endangered or
threatened under the Act include recognition, recovery actions,
requirements for Federal protection, and prohibitions against certain
practices. Recognition through listing results in public awareness, and
conservation by Federal, State, Tribal, and local agencies, private
organizations, and individuals. The Act encourages cooperation with the
States and requires that recovery actions be carried out for all listed
species. The protection required by Federal agencies and the
prohibitions against certain activities are discussed, in part, below.
The primary purpose of the Act is the conservation of endangered
and threatened species and the ecosystems upon which they depend. The
ultimate goal of such conservation efforts is the recovery of these
listed species, so that they no longer need the protective measures of
the Act. Subsection 4(f) of the Act requires the Service to develop and
implement recovery plans for the conservation of endangered and
threatened species. The recovery planning process involves the
identification of actions that are necessary to halt or reverse the
species' decline by addressing the threats to its survival and
recovery. The goal of this process is to restore listed species to a
point where they are secure, self-sustaining, and functioning
components of their ecosystems.
Recovery planning includes the development of a recovery outline
shortly after a species is listed and preparation of a draft and final
recovery plan. The recovery outline guides the immediate implementation
of urgent recovery actions and describes the process to be used to
develop a recovery plan. Revisions of the plan may be done to address
continuing or new threats to the species, as new substantive
information becomes available. The recovery plan identifies site-
specific management actions that set a trigger for review of the five
factors that control whether a species remains endangered or may be
downlisted or delisted, and methods for monitoring recovery progress.
Recovery plans also establish a framework for agencies to coordinate
their recovery efforts and provide estimates of the cost of
implementing recovery tasks. Recovery teams (composed of species
experts, Federal and State agencies, nongovernmental organizations, and
stakeholders) are often established to develop recovery plans. When
completed, the recovery outline, draft recovery plan, and the final
recovery plan will be available on our Web site (https://www.fws.gov/endangered), or from our Sacramento Fish and Wildlife Office (see FOR
FURTHER INFORMATION CONTACT).
Implementation of recovery actions generally requires the
participation of a broad range of partners, including other Federal
agencies, States, Tribal, nongovernmental organizations, businesses,
and private landowners. Examples of recovery actions include habitat
restoration (e.g., restoration of native vegetation), research, captive
propagation and reintroduction, and outreach and education. The
recovery of many listed species cannot be
[[Page 24309]]
accomplished solely on Federal lands because their range may occur
primarily or solely on non-Federal lands. To achieve recovery of these
species requires cooperative conservation efforts on private, State,
and Tribal lands.
Following publication of this final listing rule, funding for
recovery actions will be available from a variety of sources, including
Federal budgets, State programs, and cost share grants for non-Federal
landowners, the academic community, and nongovernmental organizations.
In addition, pursuant to section 6 of the Act, the States of California
and Nevada would be eligible for Federal funds to implement management
actions that promote the protection or recovery of the Sierra Nevada
mountain yellow-legged frog, Northern Distinct Population Segment of
the mountain yellow-legged frog, and the Yosemite toad. Information on
our grant programs that are available to aid species recovery can be
found at: https://www.fws.gov/grants.
Please let us know if you are interested in participating in
recovery efforts for the Sierra Nevada yellow-legged frog, the northern
DPS of the mountain yellow-legged frog, or the Yosemite toad.
Additionally, we invite you to submit any new information on these
species whenever it becomes available and any information you may have
for recovery planning purposes (see FOR FURTHER INFORMATION CONTACT).
Section 7(a) of the Act requires Federal agencies to evaluate their
actions with respect to any species that is listed as an endangered or
threatened species and with respect to its critical habitat, if any is
designated. Regulations implementing this interagency cooperation
provision of the Act are codified at 50 CFR part 402. Section 7(a)(2)
of the Act requires Federal agencies to ensure that any action
authorized, funded or carried out by such agency is not likely to
jeopardize the continued existence of the species or destroy or
adversely modify its critical habitat. If a Federal action may affect a
listed species or its critical habitat, the responsible Federal agency
must enter into consultation with the Service.
Federal agency actions within the species' habitat that may require
consultation, as described in the preceding paragraph, include
management and any other landscape-altering activities on Federal lands
administered by the USFS, NPS, and other Federal agencies as
appropriate.
The Act and its implementing regulations set forth a series of
general prohibitions and exceptions that apply to all endangered and
threatened wildlife. The prohibitions of section 9(a)(2) of the Act,
codified at 50 CFR 17.21 for endangered wildlife, in part, make it
illegal for any person subject to the jurisdiction of the United States
to take (includes harass, harm, pursue, hunt, shoot, wound, kill, trap,
capture, or collect; or to attempt any of these), import, export, ship
in interstate commerce in the course of commercial activity, or sell or
offer for sale in interstate or foreign commerce any listed species.
Under the Lacey Act (18 U.S.C. 42-43; 16 U.S.C. 3371-3378), it is also
illegal to possess, sell, deliver, carry, transport, or ship any such
wildlife that has been taken illegally. Certain exceptions apply to
agents of the Service and State conservation agencies.
We may issue permits to carry out otherwise prohibited activities
involving endangered and threatened wildlife species under certain
circumstances. Regulations governing permits are codified at 50 CFR
17.22 for endangered species, and at 17.32 for threatened species. With
regard to endangered wildlife, a permit must be issued for the
following purposes: for scientific purposes, to enhance the propagation
or survival of the species, and for incidental take in connection with
otherwise lawful activities.
It is our policy, as published in the Federal Register on July 1,
1994 (59 FR 34272), to identify to the maximum extent practicable at
the time a species is listed, those activities that would or would not
constitute a violation of section 9 of the Act. The intent of this
policy is to increase public awareness of the effect of a listing on
proposed and ongoing activities within the range of listed species. The
following activities could potentially result in a violation of section
9 of the Act; this list is not comprehensive:
(1) Unauthorized collecting, handling, possessing, selling,
delivering, carrying, or transporting of the species, including import
or export across State lines and international boundaries, except for
properly documented antique specimens of these taxa at least 100 years
old, as defined by section 10(h)(1) of the Act;
(2) Introduction of species that compete with or prey upon the
Sierra Nevada yellow-legged frog, the northern DPS of the mountain
yellow-legged frog, or the Yosemite toad;
(3) The unauthorized release of biological control agents that
attack any life stage of these species;
(4) Unauthorized modification of the mountain meadow habitats or
associated upland areas important for the breeding, rearing, and
survival of these species; and
(5) Unauthorized discharge of chemicals or fill material into any
waters in which the Sierra Nevada yellow-legged frog, the northern DPS
of the mountain yellow-legged frog, or the Yosemite toad are known to
occur.
Questions regarding whether specific activities would constitute a
violation of section 9 of the Act should be directed to the Sacramento
Fish and Wildlife Office (see FOR FURTHER INFORMATION CONTACT).
Under section 4(d) of the ESA, the Secretary has discretion to
issue such regulations as he deems necessary and advisable to provide
for the conservation of threatened species. Our implementing
regulations (50 CFR 17.31) for threatened wildlife generally
incorporate the prohibitions of section 9 of the Act for endangered
wildlife, except when a ``special rule'' promulgated pursuant to
section 4(d) of the Act has been issued with respect to a particular
threatened species. In such a case, the general prohibitions in 50 CFR
17.31 would not apply to that species, and instead, the special rule
would define the specific take prohibitions and exceptions that would
apply for that particular threatened species, which we consider
necessary and advisable to conserve the species. The Secretary also has
the discretion to prohibit by regulation with respect to a threatened
species any act prohibited by section 9(a)(1) of the ESA. Exercising
this discretion, which has been delegated to the Service by the
Secretary, the Service has developed general prohibitions that are
appropriate for most threatened species in 50 CFR 17.31 and exceptions
to those prohibitions in 50 CFR 17.32. Since we are not promulgating a
special section 4(d) rule, all of the section 9 prohibitions, including
the ``take'' prohibitions, will apply to the Yosemite toad.
Required Determinations
National Environmental Policy Act (42 U.S.C. 4321 et seq.)
We have determined that environmental assessments and environmental
impact statements, as defined under the authority of the National
Environmental Policy Act (NEPA; 42 U.S.C. 4321 et seq.), need not be
prepared in connection with listing a species as an endangered or
threatened species under the Endangered Species Act. We published a
notice outlining our reasons for this determination in the Federal
Register on October 25, 1983 (48 FR 49244).
[[Page 24310]]
Government-to-Government Relationship With Tribes
In accordance with the President's memorandum of April 29, 1994
(Government-to-Government Relations with Native American Tribal
Governments; 59 FR 22951), Executive Order 13175 (Consultation and
Coordination With Indian Tribal Governments), and the Department of the
Interior's manual at 512 DM 2, we readily acknowledge our
responsibility to communicate meaningfully with recognized Federal
Tribes on a government-to-government basis. In accordance with
Secretarial Order 3206 of June 5, 1997 (American Indian Tribal Rights,
Federal-Tribal Trust Responsibilities, and the Endangered Species Act),
we readily acknowledge our responsibilities to work directly with
tribes in developing programs for healthy ecosystems, to acknowledge
that tribal lands are not subject to the same controls as Federal
public lands, to remain sensitive to Indian culture, and to make
information available to tribes.
References Cited
A complete list of references cited in this rulemaking is available
on the Internet at https://www.regulations.gov and upon request from the
Sacramento Fish and Wildlife Office (see FOR FURTHER INFORMATION
CONTACT).
Authors
The primary authors of this final rule are the staff members of the
Sacramento Fish and Wildlife Office.
List of Subjects in 50 CFR Part 17
Endangered and threatened species, Exports, Imports, Reporting and
recordkeeping requirements, Transportation.
Regulation Promulgation
Accordingly, we amend part 17, subchapter B of chapter I, title 50
of the Code of Federal Regulations, as follows:
PART 17--[AMENDED]
0
1. The authority citation for part 17 continues to read as follows:
Authority: 16 U.S.C. 1361-1407; 1531-1544; 4201-4245; unless
otherwise noted.
0
2. Amend Sec. 17.11(h), the List of Endangered and Threatened
Wildlife, by revising the entry for ``Frog, mountain yellow-legged
(southern California DPS)'' and adding entries for ``Frog, mountain
yellow-legged (northern California DPS)'', ``Frog, Sierra Nevada
yellow-legged'', and ``Toad, Yosemite'' to the List of Endangered and
Threatened Wildlife in alphabetical order under Amphibians to read as
follows:
Sec. 17.11 Endangered and threatened wildlife.
* * * * *
(h) * * *
--------------------------------------------------------------------------------------------------------------------------------------------------------
Species Vertebrate
-------------------------------------------------------- population where Critical Special
Historic range endangered or Status When listed habitat rules
Common name Scientific name threatened
--------------------------------------------------------------------------------------------------------------------------------------------------------
* * * * * * *
Amphibians
* * * * * * *
Frog, mountain yellow-legged Rana muscosa........ U.S.A. (CA)........ U.S.A., northern E 834 NA NA
(northern California DPS). California.
Frog, mountain yellow-legged Rana muscosa........ U.S.A. (CA)........ U.S.A., southern E 728 17.95(d) NA
(southern California DPS). California.
* * * * * * *
Frog, Sierra Nevada yellow-legged Rana sierrae........ U.S.A. (CA, NV).... Entire............. E 834 NA NA
* * * * * * *
Toad, Yosemite................... Anaxyrus canorus.... U.S.A. (CA)........ Entire............. T 834 NA NA
* * * * * * *
--------------------------------------------------------------------------------------------------------------------------------------------------------
* * * * *
Dated: April 21, 2014.
Daniel M. Ashe,
Director, U.S. Fish and Wildlife Service.
[FR Doc. 2014-09488 Filed 4-25-14; 1:30 pm]
BILLING CODE 4310-55-P