Endangered and Threatened Wildlife and Plants; Finding for the Gopher Tortoise Eastern and Western Distinct Population Segments, 61834-61868 [2022-21659]

Agencies

• Fish and Wildlife Service
• DEPARTMENT OF THE INTERIOR

[Federal Register Volume 87, Number 196 (Wednesday, October 12, 2022)]
[Proposed Rules]
[Pages 61834-61868]
From the Federal Register Online via the Government Publishing Office [www.gpo.gov]
[FR Doc No: 2022-21659]



[[Page 61833]]

Vol. 87

Wednesday,

No. 196

October 12, 2022

Part III





 Department of the Interior





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Fish and Wildlife Service





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50 CFR Part 17





Endangered and Threatened Wildlife and Plants; Finding for the Gopher 
Tortoise Eastern and Western Distinct Population Segments; Proposed 
Rule

Federal Register / Vol. 87, No. 196 / Wednesday, October 12, 2022 / 
Proposed Rules

[[Page 61834]]


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DEPARTMENT OF THE INTERIOR

Fish and Wildlife Service

50 CFR Part 17

[Docket No. FWS-R4-ES-2009-0029; FF09E21000 FXES1111090FEDR 223]


Endangered and Threatened Wildlife and Plants; Finding for the 
Gopher Tortoise Eastern and Western Distinct Population Segments

AGENCY: Fish and Wildlife Service, Interior.

ACTION: Notification of findings.

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SUMMARY: We, the U.S. Fish and Wildlife Service (Service), announce 
findings on the status of the gopher tortoise (Gopherus polyphemus) 
rangewide and in the eastern (east of the Mobile and Tombigbee Rivers) 
and western (west of the Mobile and Tombigbee Rivers) portions of the 
range under the Endangered Species Act of 1973, as amended (Act). After 
a review of the best available scientific and commercial information, 
we find that listing the gopher tortoise as an endangered or a 
threatened species rangewide is not warranted. We find that the gopher 
tortoise in the eastern portion of its range and the gopher tortoise in 
the western portion of its range meet the criteria of separate distinct 
population segments (DPS), as defined by our Policy Regarding the 
Recognition of Distinct Vertebrate Population Segments Under the 
Endangered Species Act. We determine the Eastern DPS of the gopher 
tortoise is not warranted for listing at this time. Further, we confirm 
that the Western DPS of the gopher tortoise meets the definition of a 
threatened species. Additionally, this notice serves as our completed 
5-year review of the Western DPS of the gopher tortoise. We ask the 
public to submit to us any new information that becomes available 
concerning the threats to the gopher tortoise or its habitat at any 
time.

DATES: The finding announced in this document was made on October 12, 
2022.

ADDRESSES: This finding is available on the internet at https://www.regulations.gov at Docket No. FWS-R4-ES-2009-0029. Supporting 
information that we developed for this finding including the species 
status assessment report, peer review, and future condition modeling, 
are found in the decision file available at https://www.regulations.gov 
at Docket No. FWS-R4-ES-2009-0029 and on the Service's website at 
https://www.fws.gov/office/florida-ecological-services/library, and is 
available for public inspection, by appointment, during normal business 
hours at the U.S. Fish and Wildlife Service, Florida Field Office, 7915 
Baymeadows Way, Suite 200, Jacksonville, FL 32256. Please submit any 
new information or materials concerning this finding to the above 
address.

FOR FURTHER INFORMATION CONTACT: Lourdes Mena, Division Manager, 
Florida Classification and Recovery, U.S. Fish and Wildlife Service, 
Florida Ecological Services Field Office, 7915 Baymeadows Way, Suite 
200, Jacksonville, FL 32256-7517; telephone 904-731-3134; or James 
Austin, Acting Field Supervisor, Mississippi Ecological Services Field 
Office, 6578 Dogwood View Parkway, Jackson, MS 39213; telephone 601-
321-1129. Individuals in the United States who are deaf, deafblind, 
hard of hearing, or have a speech disability may dial 711 (TTY, TTDD, 
or TeleBraille) to access telecommunications relay services. 
Individuals outside the United States should use the relay services 
offered within their country to make international calls to the point-
of-contact in the United States.

SUPPLEMENTARY INFORMATION:

Previous Federal Actions

    On July 7, 1987, the gopher tortoise (Gopherus polyphemus) was 
listed under the Act as a threatened species (52 FR 25376) in the 
western portion of its range, from the Tombigbee and Mobile Rivers in 
Alabama west to southeastern Louisiana on the lower Gulf Coastal Plain. 
On January 18, 2006, we received a petition dated January 13, 2006, 
from Save Our Big Scrub, Inc. and Wild South requesting that the 
population of the gopher tortoise in the eastern portion of its range 
be listed as a threatened species under the Act and critical habitat be 
designated. On September 9, 2009, we published a 90-day finding (74 FR 
46401) that the petition contained substantial information indicating 
that listing may be warranted for the eastern population of the gopher 
tortoise. On July 27, 2011, we published a 12-month finding (76 FR 
45130) on the petition to list the gopher tortoise in the eastern 
portion of its range, and, in that finding, we evaluated the status of 
the gopher tortoise in the western portion of its range. We reaffirmed 
that the gopher tortoise warranted listing as a threatened species in 
the western portion of its range. We found the gopher tortoise in the 
eastern portion of its range was warranted for listing but precluded by 
higher priority listing actions (warranted but precluded finding).
    The species was placed on the candidate list (our list of species 
that have been found to warrant listing, but which are precluded by 
higher priority listing actions) and received a listing priority number 
of 8 based on the magnitude and immediacy of the threats. The eastern 
population of gopher tortoise was included in subsequent annual 
candidate notices of review (CNORs) (76 FR 66370, October 26, 2011; 77 
FR 69994, November 21, 2012; 78 FR 70104, November 22, 2013; 79 FR 
72450, December 5, 2014; 80 FR 80584, December 24, 2015; 81 FR 87246, 
December 2, 2016; 84 FR 54732, October 10, 2019; 85 FR 73164, November 
16, 2020; 87 FR 26152, May 3, 2022).
    On April 1, 2021, the Center for Biological Diversity (CBD) filed a 
complaint alleging our ``warranted but precluded'' finding for the 
eastern population of the gopher tortoise violated the Act because we 
were not making ``expeditious progress'' in adding qualified species to 
the lists of endangered or threatened species and because we had not 
shown that the immediate proposal of the eastern population of the 
gopher tortoise was precluded by higher priority actions consistent 
with 16 U.S.C. 1533(b)(3)(B)(iii). On April 26, 2022, the Service 
entered into a court-approved settlement agreement with CBD requiring 
the Service to submit either a warranted or a not warranted finding for 
the eastern population of gopher tortoise to the Federal Register by 
September 30, 2022.
    On June 20, 2019, we initiated a 5-year review for the western 
population of the gopher tortoise (84 FR 28850), and this document 
completes our status review under section 4(c)(2) of the Act. See 
https://ecos.fws.gov/ecp/species/C044 for the species profile for the 
gopher tortoise.

Supporting Documents

    A species status assessment (SSA) team prepared an SSA report for 
the gopher tortoise. The SSA team was composed of Service biologists, 
in consultation with other species experts. The SSA report represents 
compilations of the best scientific and commercial data available 
concerning the status of the species, including the impacts of past, 
present, and future factors (both negative and beneficial) affecting 
the species. In accordance with our joint policy on peer review 
published in the Federal Register on July 1, 1994 (59 FR 34270), and 
our August 22, 2016, memorandum updating and clarifying the role of 
peer review of listing actions

[[Page 61835]]

under the Act, we sought the expert opinions of seven appropriate 
specialists regarding the gopher tortoise SSA. We received responses 
from two peer reviewers. We coordinated with the nine Tribal nations in 
the range of the species early in the SSA process for the gopher 
tortoise, including the Catawba Nation, the Jena Band of Choctaw 
Indians, the Tunica-Biloxi Indian Tribe, the Miccosukee Tribe of 
Indians, the Seminole Tribe of Florida, the Chitimacha Tribe of 
Louisiana, the Coushatta Tribe of Louisiana, the Mississippi Band of 
Choctaw Indians, and the Poarch Band of Creek Indians. We sent the 
draft SSA report for review to 10 Tribes (with the addition of the 
Cherokee Nation).

Background

Species Information

    In this section, we present an overview of the biological 
information for gopher tortoise. A more thorough review of the 
taxonomy, species description, life history, species needs, and ecology 
of the gopher tortoise is presented in detail in the SSA report 
(Service 2022, pp. 24-45).
Taxonomy and Species Description
    The gopher tortoise is the only tortoise (family Testudinidae) east 
of the Mississippi River; one of six species in the genus Gopherus in 
North America (Ernst and Lovich 2009, p. 581; Edwards et al. 2016, p. 
131). The scientific name, Gopherus polyphemus, has remained unchanged 
since it was first described by F.M. Daudin in 1802. There is no 
taxonomic distinction between the gopher tortoise in the western and 
eastern portions of its range or at any level of geographic 
subdivision. However, genetic differences do occur in populations 
across the range of the species. Genetic variation across the range is 
best explained by the geographic features of the Apalachicola-
Chattahoochee River system and the Mobile and Tombigbee Rivers in 
Alabama (Osentoski and Lamb 1995, p. 709; Clostio et al. 2012, pp. 613-
625; Ennen et al. 2012, pp. 110-122; Gaillard et al. 2017, p. 497) (see 
Genetics section below for more information).
    The gopher tortoise is larger than any other terrestrial turtle in 
the Southeast and is characterized by a domed, dark brown to grayish-
black carapace (upper shell) and a yellowish plastron (lower shell). 
Adult gopher tortoises are typically 10 to 12 inches (in) (25.4 to 30.5 
centimeters (cm)) long and weigh 9 to 13 pounds (4 to 6 kilograms) 
(Ernst et al. 1994, p. 466; Ashton and Ashton 2008, p. 17; Bramble and 
Hutchison 2014, p. 4). Hatchlings are up to 2 in (5 cm) in length, with 
a somewhat soft, yellow-orange shell. When young, female gopher 
tortoises may be smaller than males, but, as adults, female tortoises 
are generally larger than males. Females have a flat plastron, while 
that of males is more concave. Male gopher tortoises can also be 
distinguished by a larger gland under the chin and a longer throat 
projection. As a fossorial species, its hind feet are elephantine or 
stumpy, and the forelimbs are shovel-like, with claws used for digging.
Range and Distribution
    The gopher tortoise occurs in the Southeastern Atlantic and Gulf 
Coastal Plains from southern South Carolina, west through Georgia, the 
Florida panhandle, Alabama, and Mississippi to eastern Louisiana, and 
south through peninsular Florida (Auffenberg and Franz 1982, p. 95). 
The current range of the gopher tortoise generally aligns with the 
species' historical range and the historical range of the longleaf pine 
ecosystem (Auffenberg and Franz 1982, pp. 99-120). The eastern portion 
of the gopher tortoise's range includes Alabama (east of the Tombigbee 
and Mobile Rivers), Florida, Georgia, and southern South Carolina. The 
western portion of the range includes areas west of the Tombigbee River 
in Alabama, Mississippi, and Louisiana.
    The gopher tortoise is more widespread and abundant in the eastern 
portion of its range, particularly in central and north Florida and 
eastern and southern Georgia. These areas in Florida and Georgia make 
up the core of the species' distribution (Tuberville et al. 2009, p. 
12). The best available information indicates gopher tortoises occur on 
approximately 844,812 acres (ac) (341,883 hectares (ha)) across the 
species' range (areal extent of populations as delineated for our 
analysis below in Analysis Unit and Population Delineation). An 
additional 16,338,932 ac (6,612,131 ha) of potential habitat has been 
identified by a species-specific habitat suitability model (Crawford et 
al. 2020, entire; Service 2022, pp. 122-126). For the SSA assessment, 
potential habitat is described as suitable habitat with unknown gopher 
tortoise presence outside delineated local gopher tortoise populations 
but within the species' current range. Rangewide, approximately 80 
percent of potential gopher tortoise habitat occurs in private 
ownership, with the remainder owned or managed by local, State, 
Federal, or private conservation entities (Wear and Greis 2013, p. 103; 
Natural Resources Conservation Service (NRCS) 2018, p. 2).
Life History
    The gopher tortoise's life history is characterized by a late age 
of reproductive maturity, low reproductive output (fecundity), and long 
lifespan, which make this long-lived species more vulnerable to 
demographic perturbations and slower to rebound from impacts to 
populations (Lohoefener and Lohmeier 1984, p. 2; Service 2013, p. 21; 
Tuberville et al. 2014, p. 1151). Gopher tortoises reach reproductive 
maturity between 9 and 20 years of age, although reproductive maturity 
is determined by size rather than age. Growth rates and sizes at sexual 
maturity vary among populations and habitat quality (Landers et al. 
1982, pp. 104-105; Mushinsky et al. 1994, pp. 123-125).
    Gopher tortoises generally breed from May through October; however, 
the gopher tortoise populations in south Florida have an extended 
reproductive season (Landers et al. 1980, p. 355; McRae et al. 1981, 
pp. 172-173; Taylor 1982, entire; Diemer 1992a, pp. 282-283; Ott-
Eubanks et al. 2003, p. 317; Moore et al. 2009, p. 391). The warmer 
weather in south Florida is associated with year-round courtship 
behavior, greater site productivity, and larger clutches leading to 
production of young over a much longer time period than populations 
farther north (Ashton et al. 2007, p. 359; Moore et al. 2009, p. 391). 
Female gopher tortoises usually lay eggs from mid-May through mid-July, 
and incubation lasts 80-110 days (Diemer 1986, p. 127). Rangewide, 
average clutch size varies from 4-8 eggs per clutch, with clutches in 
the western portion of the range averaging lower with 4.8-5.6 eggs per 
clutch (Seigel and Hurley 1993, p. 6; Seigel and Smith 1996, pp. 10-11; 
Tuma 1996, pp. 22-23; Epperson and Heise 2003, pp. 318-321; Ashton et 
al. 2007, p. 357). Sex determination is temperature dependent for 
gopher tortoises, with lower temperatures producing more males and 
higher temperatures producing more females. The pivotal temperature for 
a 1:1 sex ratio has been observed to be 29.3 degrees Celsius ([deg]C) 
(84.7 degrees Fahrenheit ([deg]F)) (DeMuth 2001, pp. 1612-1613). The 
lifespan of gopher tortoises is generally estimated at 50-80 years.
    The gopher tortoise's diet reflects that of a generalist herbivore 
(e.g., eating mainly grasses, plants, fallen flowers, fruits, and 
leaves) and may also include insects and carrion (Auffenberg and 
Iverson 1979, p. 558; Landers 1980, p. 9; Garner and Landers 1981, p. 
123;

[[Page 61836]]

Wright 1982, p. 25; Macdonald and Mushinsky 1988, pp. 349-351; Birkhead 
et al. 2005, pp. 146, 155; Mushinsky et al. 2006, p. 480; Richardson 
and Stiling 2019, pp. 387-388). Gopher tortoises prefer grassy, open-
canopy microhabitats, and their population density directly relates to 
the density and diversity of herbaceous biomass and a lack of canopy 
(Auffenberg and Iverson 1979, p. 558; Landers and Speake 1980, p. 522; 
Wright 1982, p. 22; Stewart et al. 1993, p. 79; Breininger et al. 1994, 
p. 63; Boglioli et al. 2000, p. 703; Ashton and Ashton 2008, p. 78).
Habitat
    Gopher tortoise habitat comprises well-drained, sandy soils (needed 
for burrowing, sheltering, and nest construction/breeding), with an 
open canopy, sparsely vegetated midstory, and abundant herbaceous 
groundcover (for feeding). Soil characteristics are an important 
component of gopher tortoise habitat and affect burrow density and 
extent. The soils in the eastern portion of the range are characterized 
by a higher sand content, although the percentage of sand varies by 
habitat type (i.e., coastal soils often contain more sand than more 
inland upland soils) (Auffenberg and Franz 1982, pp. 98-105, 113-118, 
120-121). In the western portion of the range, soils are loamy and 
contain more clay, and xeric (dry) conditions are less common west of 
the Florida panhandle (Lohoefener and Lohmeier 1981, p. 240; Auffenberg 
and Franz 1982, pp. 114-115; Mann 1995, pp. 10-11; Craul et al. 2005, 
pp. 11-13). Higher clay content in soils may contribute to lower 
abundance and density of tortoises (Means 1982, p. 524; Wright 1982, p. 
21; Ultsch and Anderson 1986, p. 790; Estes and Mann 1996, p. 24; Smith 
et al. 1997, p. 599; Jones and Dorr 2004, p. 461).
    Historically, gopher tortoise's habitats were open pine forests, 
savannahs, and xeric grasslands. Today, upland natural vegetative 
communities, including longleaf pine (Pinus palustris) and other open 
pine systems, sandhill, xeric (dry) oak (Quercus spp.) uplands, xeric 
hammock, xeric Florida scrub, and maritime scrub coastal habitats, most 
often provide the conditions necessary (e.g., open canopy and abundant 
herbaceous groundcover) to support gopher tortoises (Auffenberg and 
Franz 1982, p. 99; Diemer 1986, p. 126; Diemer 1987, pp. 73-74; 
Breininger et al. 1994, p. 60). In addition to the upland natural 
communities, some ruderal (disturbed) habitat may also provide the open 
canopy or sunny conditions and herbaceous groundcover needed by gopher 
tortoises (Auffenberg and Franz 1982, p. 99; Howell et al. 2020, p. 1). 
An open canopy allows sunlight to reach the forest floor to stimulate 
the growth and development of herbaceous groundcover and provide warmth 
for basking and egg incubation (Landers 1980, pp. 6, 8; Landers and 
Speake 1980, p. 522; Lohoefener and Lohmeier 1981, entire; Auffenberg 
and Franz 1982, pp. 98-99, 104-107, 111, 120; Boglioli et al. 2000, p. 
703; Rostal and Jones 2002, p. 485; Jones and Dorr 2004, p. 461; 
McDearman 2006, p. 2; McIntyre et al. 2019, p. 287). When canopies 
become too dense in an area, gopher tortoises move into ruderal 
habitats such as roadsides with more herbaceous ground cover, lower 
tree cover, and sun exposure (Garner and Landers 1981, p. 122; McCoy et 
al. 1993, p. 38; Baskaran et al. 2006, p. 346). Ruderal habitats may 
also include utility rights-of-way, edges, fencerows, pasturelands, and 
planted pine stands.
    Historically, open-canopied southern pine forests were maintained 
by frequent, lightning-generated fires. Currently, a variety of land 
management practices including prescribed fire, grazing, mowing, roller 
chopping, timber harvesting, and selective herbicide application, are 
used in the restoration, enhancement, and maintenance of gopher 
tortoise habitats (Cox et al. 2004, p. 10; Ashton and Ashton 2008, p. 
78; Georgia Department of Natural Resources (GDNR) 2014, unpaginated; 
Rautsaw et al. 2018, p. 141). These habitat management activities 
implemented singularly or in combination (e.g., roller chopping 
followed by prescribed fire) are used to restore and maintain the open 
canopy, sparsely vegetated midstory, and abundant herbaceous 
groundcover conditions needed by gopher tortoises.
    Gopher tortoise burrows are central to normal feeding, breeding, 
and sheltering activity. Gopher tortoises can excavate many burrows 
over their lifetime and often use several each year. Burrows typically 
extend 15 to 25 feet (ft; 4.6 to 7.6 meters (m)), can be up to 12 ft 
(3.7 m) deep, and provide shelter from predators, winter cold, fire, 
and summer heat (Hansen 1963, p. 359; Landers 1980, p. 6; Wright 1982, 
p. 50; Diemer 1986, p. 127; Boglioli 2000, p. 699). Tortoises spend 
most of their time within burrows and emerge during the day to bask, 
feed, and reproduce (Service 2022, p. 28). During the cool weather 
dormant season, gopher tortoises throughout most of the range shelter 
within their burrows, become torpid, do not eat, and rarely emerge, 
except on warm days to bask in sunlight at the burrow entrance (Service 
2013, p. 21).
    As a keystone species (which is a species that has a 
disproportionately large effect on its natural environment relative to 
its abundance), gopher tortoise burrow systems provide benefits to the 
landscape and return leached nutrients to the soil surface; increase 
habitat heterogeneity; shelter seeds from fires; and provide resources 
and refugia for other species (Auffenberg and Weaver 1969, p. 191; 
Landers 1980, pp. 2, 515; Kaczor and Hartnett 1990, pp. 107-108). An 
estimated 60 vertebrates and 302 invertebrates, including the 
threatened Eastern indigo snake, the gopher mouse, the six-lined 
roadrunner, the gopher frog, the cave cricket, and casual visitants, 
such as the tiger beetle, skunk, opossum, and rattlesnakes, share 
tortoise burrows (Jackson and Milstrey 1989, p. 87).
Genetics
    Genetic flow in gopher tortoise populations is known to be 
influenced by distance, geographic features, and human influence by 
transporting tortoises across the range. Several studies show genetic 
assemblages across the geographic range, but these studies have not 
been entirely congruent in their delineations of western and eastern 
genetic assemblages (Osentoski and Lamb 1995, p. 713; Clostio et al. 
2012, pp. 617-620; Ennen et al. 2012, pp. 113-120; Gaillard et al. 
2017, pp. 501-503). Recent microsatellite analysis suggests there are 
five main genetic groups delineated by the Tombigbee and Mobile Rivers, 
Apalachicola and Chattahoochee Rivers, and the transitional areas 
between several physiographic province sections of the Coastal Plains 
(i.e., Eastern Gulf, Sea Island, and Floridian) (figure 1) (Gaillard et 
al. 2017, pp. 505-507).

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[GRAPHIC] [TIFF OMITTED] TP12OC22.000

    The last decade of genetic research has shown that genetic 
diversity exists among individuals in a population, among populations, 
and across the range (Ennen et al. 2010, entire; Clostio et al. 2012, 
entire; Gaillard et al. 2017, entire). The most recent rangewide 
genetic analysis confirmed that the edges (periphery) of the range have 
lower levels of genetic diversity relative to the core but also showed 
genetic mixing at the borders between units (Gaillard et al. 2017, p. 
507). Evidence of tortoises with ancestry from different genetic sites 
is most likely due to the decades of tortoises being moved by humans as 
part of various formal and informal translocation and population 
augmentation efforts as well as non-conservation, human-mediated 
movements (see Translocation and Headstarting, below) (Gaillard et al. 
2017, pp. 504-505). In addition, contemporary gene flow is asymmetric 
across the gopher tortoise range as a result of recent migrations 
affecting changes in genetic diversity. For example, gene flow is 
higher from the Central to Western genetic regions and from the Florida 
panhandle to the East Georgia genetic region, while the Florida 
panhandle area has low genetic flow with the West Georgia genetic 
region (Gaillard et al. 2017, pp. 504-509). In general, migration rates 
between genetic regions were low, with the highest proportion of 
movements and genetic exchange from within the same genetic unit 
(Gaillard et al. 2017, pp. 505-506).
Home Range and Movement
    As mentioned previously, gopher tortoises often use several burrows 
per year. The burrows of a gopher tortoise represent the general 
boundaries of a home range, which is the area used for feeding, 
breeding, and sheltering (McRae et al. 1981, p. 176). Gopher tortoise 
home ranges tend to vary in size depending on habitat quality, with 
larger areas in lower quality habitat (Auffenberg and Iverson 1979, pp. 
559-561; Castellon et al. 2012, p. 159; Guyer et al. 2012, p. 130). 
Home ranges are larger in the western portion of the gopher tortoise 
range than those typically observed for tortoises in Alabama (east of 
the Tombigbee and Mobile Rivers), Georgia, South Carolina, and Florida, 
and this variation is most likely due to habitat quality differences 
(Lohoefener and Lohmeier 1984, pp. 1-25; Epperson and Heise 2003, p. 
315; Tuberville et al. 2005, p. 356; Richter et al. 2011, p. 408). 
Males typically have larger home ranges and tend to travel farther 
distances than females; this is primarily for breeding opportunities 
and related to burrow density and social hierarchical behaviors (McRae 
et al. 1981, p. 175; Guyer et al. 2012, pp. 129-

[[Page 61838]]

132; Castellon et al. 2018, pp. 11-12). For example, average home 
ranges in Mississippi, Alabama, Florida, and Georgia have varied from 
0.1 to 39.8 ac (0.04 to 16.1 hectares ha) (McRae et al. 1981, pp. 175-
176; Diemer 1992b, pp. 160-161; Tuma 1996, pp. 28-43; Ott-Eubanks et 
al. 2003, pp. 315-316; Guyer et al. 2012, pp. 128-129; Castellon et al. 
2018, p. 17).
    Just as gopher tortoise home ranges are larger in lower quality 
habitat, gopher tortoise movements also increase as herbaceous biomass 
and habitat quality decrease and tortoises must search farther for 
adequate resources (Auffenberg and Iverson 1979, p. 558; Auffenberg and 
Franz 1982, p. 121; Castellon et al. 2018, p. 18). As distances 
increase between gopher tortoise burrows, isolation among gopher 
tortoises also increases due to the decreasing rate of visitation and 
breeding by males to females (Boglioli et al. 2003, p. 848; Guyer et 
al. 2012, p. 131). Most breeding populations have been found to consist 
of burrows no greater than about 549 ft (167 m) apart, although males 
may move up to 1,640 ft (500 m) for mating opportunities (Guyer and 
Johnson 2002, pp. 6-8; Ott-Eubanks et al. 2003, p. 320; Guyer et al. 
2012, p. 131).
Population Dynamics
    At the landscape scale, the gopher tortoise requires large swaths 
of interconnected, high-quality habitat patches to support healthy 
populations. Large swaths of high-quality habitat provide habitat 
connectivity for gopher tortoise life-history needs of dispersal 
(immigration and emigration), breeding, and foraging. Interconnected, 
high-quality habitat that supports gopher tortoise requirements 
influences population dynamics and demographics through the carrying 
capacity of the area and opportunities for genetic exchange.
    As long-lived animals, gopher tortoises naturally experience 
delayed sexual maturity, low reproductive rates, high mortality at 
young ages and small size-classes, and relatively low adult mortality. 
Factors affecting population growth, decline, and dynamics include the 
number or proportion of annually breeding and egg-laying females 
(breeding population size), clutch size, nest depredation rates, egg 
hatching success, mortality (hatchling-yearling, juvenile-subadult, 
adult), the age or size at first reproduction, age- or stage-class 
population structure, maximum age of reproduction, and immigration and 
emigration rates.
    Gopher tortoise population dynamics are sensitive to demographic 
changes in adult, hatchling, and juvenile survival (Gibbons 1987, 
entire; Congdon et al. 1993, entire; Heppell 1998, entire; Epperson and 
Heise 2001, entire; Miller 2001, entire; Wester 2005, entire; McDearman 
2006, p. 7). Hatchling survivorship is the most critical life history 
stage due to the high mortality in this life stage (Tuberville et al. 
2009, p. 33). For example, a simulated 5 percent decrease in hatchling 
mortality shifted the population growth rate from slowly declining (1.5 
percent decrease) to slowly increasing (1.1 percent increase) and 
eliminated the probability of extinction within 200 years (Tuberville 
et al. 2009, p. 33). Changes in other vital parameters, including age 
of first reproduction and average clutch size, also affect population 
growth, although generally not to the extent of hatchling and juvenile 
mortality (McDearman 2006, pp. 7, 20).
    Demographic factors have been evaluated in population viability 
analysis (PVA) studies to estimate the probabilities of gopher tortoise 
population extinction over time and the important factors affecting the 
species' viability (Cox et al. 1987, pp. 24-34; Lohoefener and Lohmeier 
1984, entire; Cox 1989, p. 10; Epperson and Heise 2001, pp. 37-39; 
Miller 2001, entire; Wester 2005, pp. 16-20; McDearman 2006, entire; 
Tuberville et al. 2009, entire; Folt et al. 2022, entire). The number 
of gopher tortoises required for a population to remain on the 
landscape for 200 years varies from 50 to 200 individuals depending on 
habitat and management conditions (Cox et al. 1987, pp. 27-29; Cox et 
al. 1994, p. 29). Although populations as small as 50 tortoises have 
exhibited positive growth rates and are projected to remain on the 
landscape in the future in some PVA models, the inclusion of threats 
such as upper respiratory tract disease (URTD) or fire ant (Conomyrma 
spp., Solenopsis invicta) predation led to population decline and 
eventual extirpation of these smaller populations in these models 
(Miller 2001, pp. 13, 26-27; McDearman 2006, pp. 6-7). In models that 
resulted in projected gopher tortoise population declines of 1 to 3 
percent per year, the factors that affected gopher tortoise population 
growth rates included the geographic location of the population and 
habitat quality (Tuberville et al. 2009, pp. 17-22). Populations of at 
least 100 gopher tortoises were found to be reasonably resilient to 
variations in habitat quality; however, larger populations of at least 
250 tortoises were needed to remain on the landscape in lower quality 
habitat (Tuberville et al. 2009, p. 19).
    A minimum viable population (MVP) in terms of acceptable benchmarks 
for the purpose of conservation and recovery efforts of gopher tortoise 
has been established by the Gopher Tortoise Council (GTC; GTC 2013, 
entire). Viability, as defined in the MVP, is valuable for conservation 
planning purposes and differs from the definition of viability used in 
the SSA (Service 2022, p. 20). The GTC adopted the definition of a 
viable tortoise population as consisting of at least 250 adult 
tortoises, at a density of at least 0.4 tortoises per ha, with an even 
sex ratio, and evidence of all age classes present, on a property with 
at least 247 ac (100 ha) of high-quality habitat managed for the 
benefit of the gopher tortoise (GTC 2013, pp. 2-3). Within our SSA 
report and this document, we use the GTC's definition of a ``viable 
population.'' A primary support population was defined as consisting of 
50-250 adult gopher tortoises. Primary support populations may improve 
viability through habitat restoration, natural recruitment increases, 
or population augmentation. A secondary support population was defined 
as fewer than 50 tortoises that have more constraints to reach 
sufficient viability, but are important for education, community 
interest, and augmentation, and can maintain sufficient viability to 
remain on the landscape in the long term with rigorous habitat 
management and/or connectivity with other populations (GTC 2014, p. 4). 
It should be noted that smaller support populations may remain on the 
landscape for a long period of time under high-quality habitat 
conditions but are more vulnerable to stochastic events than 
populations that meet the MVP threshold (Miller 2001, p. 28; GTC 2014, 
p. 4; Folt et al. 2021, entire). We rely on these defined population 
benchmarks in our assessment of gopher tortoise viability, as described 
below in Current Condition.

Regulatory and Analytical Framework

Regulatory Framework

    Section 4 of the Act (16 U.S.C. 1533) and the implementing 
regulations in title 50 of the Code of Federal Regulations set forth 
the procedures for determining whether a species is an endangered 
species or a threatened species, issuing protective regulations for 
threatened species, and designating critical habitat for threatened and 
endangered species. In 2019, jointly with the National Marine Fisheries 
Service, the Service issued final rules that revised the regulations in 
50 CFR parts 17 and 424 regarding how we add, remove, and reclassify 
threatened and endangered species and the criteria for designating 
listed species' critical

[[Page 61839]]

habitat (84 FR 45020 and 84 FR 44752; August 27, 2019). At the same 
time the Service also issued final regulations that, for species listed 
as threatened species after September 26, 2019, eliminated the 
Service's general protective regulations automatically applying to 
threatened species the prohibitions that section 9 of the Act applies 
to endangered species (collectively, the 2019 regulations).
    However, on July 5, 2022, the U.S. District Court for the Northern 
District of California vacated the 2019 regulations (Center for 
Biological Diversity v. Haaland, No. 4:19-cv-05206-JST, Doc. 168 (N.D. 
Cal. July 5, 2022) (CBD v. Haaland)), reinstating the regulations that 
were in effect before the effective date of the 2019 regulations as the 
law governing species classification and critical-habitat decisions. 
Accordingly, in developing the analysis contained in this finding, we 
applied the pre-2019 regulations, which may be reviewed in the 2018 
edition of the Code of Federal Regulations at 50 CFR 424.11(d). Those 
pre-2019 regulations did not include provisions clarifying the meaning 
of ``foreseeable future,'' so we applied a 2009 Department of the 
Interior Solicitor's opinion (M-37021, ``The Meaning of `Foreseeable 
Future' in Section 3(2) of the Endangered Species Act'' (Jan. 16, 2009) 
(M-37021). Because of the ongoing litigation regarding the court's 
vacatur of the 2019 regulations, and the resulting uncertainty 
surrounding the legal status of the regulations, we also undertook an 
analysis of whether the finding would be different if we were to apply 
the 2019 regulations. That analysis, which we described in a separate 
memo in the decisional file and posted on https://www.regulations.gov, 
concluded that we would have reached the same finding if we had applied 
the 2019 regulations because, based on the modeling and scenarios 
evaluated, we considered our ability to make reliable predictions in 
the future and the uncertainty in how and to what degree the gopher 
tortoise could respond to those risk factors in this timeframe. We 
determined that this timeframe represents a period of time for which we 
can reliably predict both the threats to the species and the species' 
response to those threats under the 2019 regulations. We also find this 
determination to be ``rooted in the best available data that allow 
predictions into the future'' and extend as far as those predictions 
are ``sufficiently reliable to provide a reasonable degree of 
confidence in the prediction, in light of the conservation purposes of 
the Act'' in accordance with the 2009 Solicitor's Opinion.
    On September 21, 2022, the U.S. Circuit Court of Appeals for the 
Ninth Circuit stayed the district court's July 5, 2022, order vacating 
the 2019 regulations until a pending motion for reconsideration before 
the district court is resolved (In re: Cattlemen's Ass'n, No. 22-
70194). The effect of the stay is that the 2019 regulations are 
currently the governing law. Because a court order requires us to 
submit this finding to the Federal Register by September 30, 2022, it 
is not feasible for us to revise the finding in response to the Ninth 
Circuit's decision. Instead, we hereby adopt the analysis in the 
separate memo that applied the 2019 regulations as our primary 
justification for the finding. However, due to the continued 
uncertainty resulting from the ongoing litigation, we also retain the 
analysis in this preamble that applies the pre-2019 regulations and we 
conclude that, for the reasons stated in our separate memo analyzing 
the 2019 regulations, this finding would have been the same if we had 
applied the pre-2019 regulations.
    The Act defines an ``endangered species'' as a species that is in 
danger of extinction throughout all or a significant portion of its 
range, and a ``threatened species'' as a species that is likely to 
become an endangered species within the foreseeable future throughout 
all or a significant portion of its range. The Act requires that we 
determine whether any species is an endangered species or a threatened 
species because of any of the following 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.
    These factors represent broad categories of natural or human-caused 
actions or conditions that could have an effect on a species' continued 
existence. In evaluating these actions and conditions, we look for 
those that may have a negative effect on individuals of the species, as 
well as other actions or conditions that may ameliorate any negative 
effects or may have positive effects.
    We use the term ``threat'' to refer in general to actions or 
conditions that are known to or are reasonably likely to negatively 
affect individuals of a species. The term ``threat'' includes actions 
or conditions that have a direct impact on individuals (direct 
impacts), as well as those that affect individuals through alteration 
of their habitat or required resources (stressors). The term ``threat'' 
may encompass--either together or separately--the source of the action 
or condition or the action or condition itself.
    However, the mere identification of any threat(s) does not 
necessarily mean that the species meets the statutory definition of an 
``endangered species'' or a ``threatened species.'' In determining 
whether a species meets either definition, we must evaluate all 
identified threats by considering the species' expected response and 
the effects of the threats--in light of those actions and conditions 
that will ameliorate the threats--on an individual, population, and 
species level. We evaluate each threat and its expected effects on the 
species, then analyze the cumulative effect of all of the threats on 
the species as a whole. We also consider the cumulative effect of the 
threats in light of those actions and conditions that will have 
positive effects on the species, such as any existing regulatory 
mechanisms or conservation efforts. The Secretary determines whether 
the species meets the definition of an ``endangered species'' or a 
``threatened species'' only after conducting this cumulative analysis 
and describing the expected effect on the species now and in the 
foreseeable future.
    The Act does not define the term ``foreseeable future,'' which 
appears in the statutory definition of ``threatened species.'' Because 
the decision in CBD v. Haaland vacated our 2019 regulations regarding 
the foreseeable future, we refer to a 2009 Department of the Interior 
Solicitor's opinion entitled ``The Meaning of `Foreseeable Future' in 
Section 3(20) of the Endangered Species Act'' (M-37021). That 
Solicitor's opinion states that the foreseeable future ``must be rooted 
in the best available data that allow predictions into the future'' and 
extends as far as those predictions are ``sufficiently reliable to 
provide a reasonable degree of confidence in the prediction, in light 
of the conservation purposes of the Act.'' Id. at 13.
    It is not always possible or necessary to define the foreseeable 
future as a particular number of years. Analysis of the foreseeable 
future uses the best scientific and commercial data available and 
should consider the timeframes applicable to the relevant threats and 
to the species' likely responses to those threats in view of its life-
history characteristics. Data that are typically relevant to assessing 
the species'

[[Page 61840]]

biological response include species-specific factors such as lifespan, 
reproductive rates or productivity, certain behaviors, and other 
demographic factors.

Analytical Framework

    The SSA report documents the results of our comprehensive 
biological review of the best scientific and commercial data regarding 
the status of the species, including an assessment of the potential 
threats to the species. The SSA report does not represent our decision 
on whether the species should be proposed for listing as an endangered 
or threatened species under the Act. However, it does provide the 
scientific basis that informs our regulatory decisions, which involve 
the further application of standards within the Act and its 
implementing regulations and policies. The following is a summary of 
the key results and conclusions from the SSA report; the full SSA 
report can be found at Docket FWS-R4-ES-2009-0029 on https://www.regulations.gov and at https://www.fws.gov/office/florida-ecological-services/library.
    To assess gopher tortoise viability, we used the three conservation 
biology principles of resiliency, redundancy, and representation 
(Shaffer and Stein 2000, pp. 306-310). Briefly, resiliency supports the 
ability of the species to withstand environmental and demographic 
stochasticity (for example, wet or dry, warm or cold years), redundancy 
supports the ability of the species to withstand catastrophic events 
(for example, droughts, large pollution events), and representation 
supports the ability of the species to adapt over time to long-term 
changes in the environment (for example, climate changes). In general, 
the more resilient and redundant a species is and the more 
representation it has, the more likely it is to sustain populations 
over time, even under changing environmental conditions. Using these 
principles, we identified the species' ecological requirements for 
survival and reproduction at the individual, population, and species 
levels, and described the beneficial and risk factors influencing the 
species' viability.
    The SSA process can be categorized into three sequential stages. 
During the first stage, we evaluated the individual species' life-
history needs. The next stage involved an assessment of the historical 
and current condition of the species' demographics and habitat 
characteristics, including an explanation of how the species arrived at 
its current condition. The final stage of the SSA involved making 
predictions about the species' responses to positive and negative 
environmental and anthropogenic influences. Throughout all of these 
stages, we used the best available information to characterize 
viability as the ability of a species to sustain populations in the 
wild over time. We use this information to inform our regulatory 
decision.

Summary of Biological Status and Threats

    In this discussion, we review the biological condition of the 
species and its resources, and the threats that influence the species' 
current and future condition, in order to assess the species' overall 
viability and the risks to that viability. The following discussions 
include evaluations of the following threats and associated sources 
influencing the gopher tortoise and its habitat: (1) Habitat loss, 
degradation, and fragmentation, (2) climate change, and (3) 
insufficient and/or incompatible habitat management. Other factors 
influencing gopher tortoise viability include road mortality, disease, 
harvest and rattlesnake roundups, predation, nonnative invasive 
species, and conservation measures, including relocation, 
translocation, and headstarting programs. Conservation of habitat 
through land acquisition and conservation actions on public and private 
lands and the retention of private forest lands reduces the severity of 
some of these threats by providing protection of habitat across the 
landscape, maintaining connectivity between habitat patches, and 
increasing the opportunity for beneficial habitat management actions. 
In this section, we describe the threats that influence the species' 
current and future conditions and conservation measures that may 
mitigate those threats. Additional information may be found in the SSA 
report (Service 2022, pp. 46-102).

Habitat Loss, Degradation, and Fragmentation

    Habitat loss, degradation, and fragmentation have affected the 
gopher tortoise and its habitat. The gopher tortoise was historically 
associated with fire-dependent longleaf pine ecosystems. Longleaf pine 
communities declined to less than 3 million ac (1.2 million ha) by the 
20th century from a historical estimate of 92 million ac (37 million 
ha) due to forest clearing and conversion for agriculture, conversion 
from longleaf to other pine species, and development (Frost 1993, p. 
20; Ware et al. 1993, p. 447; Landers et al. 1995, p. 39). As a result 
of fire suppression and exclusion in many areas, approximately two to 
three percent of longleaf pine ecosystems remain in relatively natural 
condition (Frost 1993, p. 17; Simberloff 1993, p. 3; Jose et al. 2007, 
p. ix; Jensen et al. 2008, p. 16; Oswalt et al. 2012, p. 7). Although 
historically associated with longleaf pine communities, the species 
currently occurs in open canopy stands of several southern pine 
species.
    Currently, habitat loss, degradation, and fragmentation caused by a 
variety of sources across the species' range continue to negatively 
affect gopher tortoise viability. Urbanization and development, major 
road construction, incompatible and/or insufficient habitat management, 
and certain types of agriculture negatively impact the gopher tortoise 
and its habitat (Auffenberg and Franz 1982, pp. 105, 112; Lohoefener 
and Lohmeier 1984, pp. 2-6; Diemer 1986, p. 128; Diemer 1987, pp. 74-
75; Hermann et al. 2002, pp. 294-295; Enge et al. 2006, p. 4). While 
large-scale development of solar farms may impact the gopher tortoise 
and its habitat in connection with other threats, we have determined 
that solar energy development is not a key factor influencing the 
species' viability at this time (Ong et al. 2013, p. iv; Service 2022, 
p. 52). Invasive species introduced as a result of habitat 
fragmentation or urbanization can influence gopher tortoises either 
through predation or alterations to habitat structure and function 
(Mann 1995, p. 24; Lippincott 1997, pp. 48-65; Basiotis 2007, p. 24; 
Engeman et al. 2009, p. 84; Engeman et al. 2011, p. 607; Dziadzio et 
al. 2016, p. 531; Bartoszek et al. 2018, pp. 353-354). Climate change 
has the potential to negatively impact habitat through the loss of 
habitat due to sea level rise, limitations on number of suitable burn 
days due to changes in temperature, precipitation, increased flooding 
due to predicted increases in the severity of hurricanes, and human 
migration from inundated coastal areas to inland areas, with subsequent 
impacts to gopher tortoises (Ruppert et al. 2008, p. 127; Castellon et 
al. 2018, pp. 11-14; Hayhoe et al. 2018, entire; Kupfer et al. 2020, 
entire). Although habitat management and climate change influence 
gopher tortoise habitat and contribute to habitat loss, fragmentation, 
and degradation, they are discussed as separate factors, below. In this 
section, we discuss below the primary sources (Urbanization and 
Development, Road Effects and Mortality) for habitat loss, 
fragmentation, and degradation.
Urbanization and Development
    At a landscape scale, the gopher tortoise needs large swaths of 
interconnected, high-quality habitat patches to support viable 
populations.

[[Page 61841]]

Within these large swaths of high-quality habitat on the landscape, 
gopher tortoises require habitat connectivity for dispersal 
(immigration and emigration), breeding, and foraging. Urbanization and 
development of the landscape fragments and replaces natural areas with 
artificial structures, impervious surfaces, and lawns and gardens 
containing nonnative plant species; this activity impacts gopher 
tortoise populations that rely on a mosaic of interconnected uplands 
(Sutherland 2009, p. 35). Development and urbanization can also impact 
gopher tortoise populations on conservation lands (lands in public or 
private ownership managed for conservation under a management plan) by 
disrupting habitat connectivity across the landscape and disrupting 
habitat management activities on conservation lands, particularly 
through the reduction of prescribed fire activities. Urbanization and 
development impacts to individuals, populations, and habitats have been 
documented, although not specifically quantified in terms of survival, 
recruitment, and health of gopher tortoises prior to our SSA. Our 
modeling for the future condition analysis in the SSA includes 
urbanization projected by the SLEUTH model as part of the threats 
scenarios as described in Future Condition (Service 2022, pp. 144-175; 
Folt et al. 2022, entire).
    Human population growth is a primary driver of urbanization and 
subsequent habitat fragmentation that is impacting gopher tortoises 
rangewide. Rangewide, Alabama, Florida, Georgia, Louisiana, and South 
Carolina have experienced population growth from 3 to 15 percent since 
2010, while Mississippi has experienced a 6 percent decrease in human 
population. Population growth from 2 to 13.4 percent is projected to 
occur in each State rangewide from 2020 to 2030 (Blanchard 2007, p. 7; 
FEDR 2021, unpaginated; Culverhouse College of Business 2021, 
unpaginated; Georgia Census 2021, unpaginated; Population Projections 
2005, unpaginated; U.S. Census Bureau 2021, unpaginated). As the human 
population continues to grow in the Southeast, development is expected 
to increase demand for forest resources and lead to habitat 
fragmentation and degradation of forests through the conversion of 
high-quality gopher tortoise habitat to lands in forest production that 
may not be managed in a way compatible with gopher tortoise needs. 
Forest loss and fragmentation reduce the ecological function and 
connectivity essential for the dispersal of gopher tortoises across the 
landscape (Guyer et al. 2012, p. 131; Jones and Dorr 2004, p. 461).
    Gopher tortoises can occur in residential areas despite the fact 
that these areas are typically of lower habitat quality. However, 
conversion of gopher tortoise habitat to residential areas results in 
mortality of gopher tortoises when individuals are entombed in burrows 
during construction activities. In the western portion of the range 
where the species is federally listed, individual gopher tortoises are 
translocated from development sites to avoid mortality from land 
development activities. Since 2007, the Florida Fish and Wildlife 
Conservation Commission (FWC) requires developers to relocate tortoises 
out of harm's way, either onsite or at an approved recipient site (FWC 
2007, p. 10). Other States (Georgia, Alabama, and South Carolina) have 
some measure of legal protection for gopher tortoises, though gopher 
tortoise burrows are not protected uniformly across the range. When 
notified, these States work with developers to minimize impacts when 
tortoises occur on development sites.
    Human development of the landscape (i.e., urbanization) affects 
terrestrial wildlife communities in the Southeastern United States, 
including gopher tortoise populations that often rely on upland 
habitats that are popular sites for urban development or agriculture. 
Gopher tortoise populations on protected and managed lands are somewhat 
buffered from habitat loss as a result of urbanization, but landscape-
level connectivity is negatively affected. Urbanization and development 
have influenced the gopher tortoise and its habitat historically, and 
we expect these effects to continue in the future. This threat is 
present across the range of the species, although populations near 
already urbanized areas and areas of projected development are more 
affected. For example, in Florida, urban growth and development is 
identified as one of the primary threats to gopher tortoises 
(Auffenberg and Franz 1982, p. 112; Diemer 1986, p. 128; Diemer 1987, 
pp. 74-75; Enge et al. 2006, p. 4). Georgia is also anticipated to 
experience dramatic human population increases (Georgia Census 2021, 
unpaginated), leading to subsequent development and potential loss of 
gopher tortoise habitat.
Road Effects and Mortality
    Roads pose a barrier to gopher tortoise movement, fragment habitat, 
isolate areas of habitat, and increase mortality of gopher tortoises 
(Andrews and Gibbons 2005, p. 772; Hughson and Darby 2013, pp. 227-
228). Roads that bisect habitat pose a hazard to gopher tortoises by 
forcing individuals into unsuitable areas and onto highways (Diemer 
1987, p. 75; Mushinsky et al. 2006, p. 38). Roads occurring within or 
adjacent to tortoise habitat impact gopher tortoises, because tortoises 
are attracted to road shoulders where open-canopy, grassy areas are 
maintained (Steen and Gibbs 2004, entire; Steen et al. 2006, p. 271). 
Gopher tortoises appear to use roadsides independently of larger 
habitat patches, treating them as areas for residency as opposed to 
travel corridors among other habitat patches (Rautsaw et al. 2018, p. 
141). Gopher tortoise nests in roadsides are more susceptible to 
predators, such as raccoons (Procyon lotor), which are common in 
ecological edges and fragmented, suburban landscapes (Hoffman and 
Gottschang 1977, p. 633; Wilcove 1985, pp. 1213-1214). The installation 
of wildlife barrier fences along roadways has the potential to minimize 
gopher tortoise road mortality. While barrier fencing along roads may 
reduce road mortality, fencing may also further limit the movement of 
gopher tortoises.
    While road mortality occurs in gopher tortoise populations, the 
extent to which it affects populations or the species is not well 
documented. There are no current rangewide monitoring efforts for 
gopher tortoise road mortality. Florida is the only state that has a 
database for reporting sick, injured, or dead tortoises; of tortoises 
reported to the Florida FWC as sick, injured, or dead, 41 percent were 
found injured or dead on roads (CCA 2018, p. 95).
    As development and subsequent habitat loss and fragmentation 
occurs, gopher tortoises will disperse to find better quality habitat, 
putting individual gopher tortoises at risk of road mortality. Impacts 
to habitat and road mortality are expected to increase as road 
densities and traffic volumes increase and habitat patches become more 
isolated and more difficult to manage (Enge et al. 2006, p. 10). 
Highway mortality of gopher tortoises will be highest where there are 
improved roads adjacent to gopher tortoise populations. Increased 
traffic on new or expanded roads adjacent to a gopher tortoise 
population will expose individuals to direct mortality from vehicles 
and potentially to increased predation. In addition, gopher tortoises 
in the vicinity of urban areas will be particularly vulnerable 
(Mushinsky et al. 2006, p. 362), especially in areas with heavy traffic 
patterns or high speed limits. The threat posed by roads is ongoing and 
is expected to continue, particularly in peninsular Florida and

[[Page 61842]]

urban centers in coastal portions of Georgia, Alabama, and Mississippi, 
where human populations are likely to increase as seen in urban 
modeling projections using SLEUTH (Terando et al. 2014, entire).
Agricultural Lands
    Agricultural lands are an important component of land use 
activities in the gopher tortoise range. Agricultural lands on suitable 
soils are 6 times less likely to have burrows and contain 20 times 
fewer gopher tortoise burrows than open pine sites (Hermann et al. 
2002, pp. 294-295). Gopher tortoises do not use the poor-quality 
habitat in annually tilled fields that do not provide necessary forage 
(Auffenberg and Franz 1982, p. 105). However, adult tortoises will 
return to abandoned agricultural fields in a few years when the land is 
dominated by perennial herbaceous species and remain until succession 
results in closed canopy conditions that do not provide the species' 
requirements (Auffenberg and Franz 1982, pp. 105, 107-108). 
Accordingly, habitat that is normally suitable for gopher tortoise but 
that is cleared for agricultural activities is not suitable for gopher 
tortoise use while it is in production or until forage and soil 
conditions provide gopher tortoise requirements for feeding, nesting, 
and sheltering.
    Cropland (i.e., agriculture) in the gopher tortoise range is 
projected to decline by 19 percent from 1997 to 2060 (Wear and Greis 
2013, p. 45). Restoration of abandoned agricultural fields with 
appropriate soils into potential gopher tortoise habitat is feasible 
and has been accomplished through the U.S. Department of Agriculture 
Conservation Reserve Program (CRP). For example, in the eastern portion 
of the gopher tortoise range, over 10.5 million acres were reported as 
enrolled in CRP from 2000 to 2019 in counties with gopher tortoise 
occurrences (USDA 2020, unpaginated). Although not all of these lands 
are expected to support gopher tortoise or fall into potential habitat, 
we expect these restoration actions will improve gopher tortoise 
habitat. However, at this time, we cannot project the extent to which 
abandoned agricultural fields will be restored to a level of 
suitability necessary to support gopher tortoise populations.
Solar Farms
    As interest in renewable energy increases, the development of solar 
farms across the gopher tortoise's range in the Southeast is also 
increasing, particularly in Florida and South Carolina (EIA 2021, 
unpaginated). A primary concern regarding large-scale deployment of 
solar energy is the potentially significant land use requirements, 
habitat fragmentation, possible exclusion of gopher tortoises as a 
result of fencing, and the need to relocate tortoises from solar farm 
sites prior to construction (Ong et al. 2013, p. iv). Some solar 
utility developers and companies recognize the potential to impact the 
gopher tortoise and its habitats and work with conservation 
organizations to avoid and minimize impacts via strategic siting 
assessments (NASA Develop 2018, unpaginated). The best available 
science indicates it is not a key factor in species viability, although 
information quantifying the extent and magnitude of the impact of solar 
farms on the gopher tortoise is limited.

Climate Change

    The effects of changing climate conditions have influenced and are 
expected to continue to influence gopher tortoises and their habitat. 
In the Southeastern United States, the impacts of climate change are 
currently occurring in the form of sea level rise and extreme weather 
events (Carter et al. 2018, p. 749). Changes in temperatures are 
projected to result in more frequent drought, more extreme heat 
(increases in air and water temperatures), increased heavy 
precipitation events (e.g., flooding), more intense storms (e.g., 
frequency of major hurricanes increases), and rising sea level and 
accompanying storm surge (Intergovernmental Panel on Climate Change 
(IPCC) 2022, entire). Higher temperatures and an increase in the 
duration and frequency of droughts are projected to increase the 
occurrence of wildfires and reduce the effectiveness of prescribed 
fires (Carter et al. 2018, pp. 773-774).
    Predicted increases in temperature across the gopher tortoise's 
range due to climate change are expected to affect the species' life 
history characteristics and demography through skewed sex ratios, 
larger clutch sizes, increased hatchling success, and larger hatchling 
size (DeMuth 2001, p. 1614; Ashton et al. 2007, pp. 355-362; Hunter et 
al. 2021, pp. 215, 221-224). Although these life history and 
demographic effects may not initially appear to have negative impacts, 
we do not have available modeling to project the effects of these 
changes on gopher tortoise demography in terms of forage availability, 
carrying capacity of areas where the gopher tortoise occurs, or other 
life history and demographic changes. However, the gopher tortoise may 
ameliorate these effects by selection of cooler nest sites and altering 
timing of nesting to earlier in the season (Czaja et al. 2020, entire). 
Some populations of gopher tortoises already exhibit both of these 
behaviors (Ashton and Ashton 2008, entire; Moore et al. 2009, entire; 
Craft 2021, pp. 42-45).
    Frequency of severe hurricanes is predicted to increase in the 
future (IPCC 2022, entire; Carter et al. 2018, entire). Gopher tortoise 
burrows, particularly those in coastal ecosystems, will be impacted by 
flooding after a hurricane, causing abandonment, though the burrow may 
become usable again (Waddle et al. 2006, pp. 281-283; Castellon et al. 
2018, pp. 11-14; Falk 2018, entire). In addition, overwash of coastal 
dunes may result in ``salt burn'' and loss of coastal vegetation, 
temporarily reducing forage availability in coastal natural communities 
used by gopher tortoises.
    Predicted changes in rangewide temperature and precipitation due to 
climate change will reduce the number of days with suitable conditions 
for prescribed burns needed to manage gopher tortoise habitat in the 
future compared to current conditions (Kupfer et al. 2020, entire). 
This reduction in prescribed fire, combined with the effects of 
urbanization, will further restrict the ability to manage gopher 
tortoise habitat with prescribed fire. In addition to the constrained 
ability to implement prescribed fire in the future, modeling for the 
Southeastern United States projects an increased wildfire risk and a 
longer fire season, with at least a 30 percent increase in lightning-
ignited wildfire from 2011 to 2060 (Vose et al. 2018, p. 239).
    Sea level rise associated with climate change is expected to affect 
coastal populations of gopher tortoises through subsequent inundation 
and loss of habitat in coastal areas. As sea levels continue to rise, 
coastal water levels--from the mean to the extreme--are growing deeper 
and reaching farther inland along most U.S. coastlines (Sweet et al. 
2022, p. 28). Global mean sea level has risen 7 to 8 in (16 to 21 cm) 
since 1900, with about half of that rise occurring since 1993 (Hayhoe 
et al. 2018, p. 85). In areas of the Southeastern United States, tide 
gauge analysis reveals as much as 1 to 3 ft (0.30 to 0.91 m) of local 
relative sea level rise in the past 100 years (Carter et al. 2018, p. 
757). The future estimated amount that sea level will rise varies based 
on the responses of the climate system to warming and human-caused 
emissions (Hayhoe et al. 2018, p. 85). The amount of gopher tortoise 
habitat predicted to be lost within a given population due to

[[Page 61843]]

sea level rise depends on the location of the population and site-
specific characteristics. Populations affected by habitat loss and 
degradation due to saltwater inundation and vegetation changes are 
expected to experience reduced abundance and resiliency. In addition, 
impacts to gopher tortoises and their habitat are expected due to the 
relocation of people from flood-prone coastal areas to inland areas, 
including the relocation of millions of people to currently undeveloped 
interior natural areas (Stanton and Ackerman 2007, p. 15; Ruppert et 
al. 2008, p. 127).
    The effects of climate change are projected to impact the gopher 
tortoise and its habitat. These impacts will be direct through loss of 
individuals and indirect through the loss of habitat due to sea level 
rise, lack of habitat management due to reduction in burn days, 
increased flooding, and human migration from inundated coastal areas to 
inland areas (Ruppert et al. 2008, p. 127; Castellon et al. 2018, pp. 
11-14; Hayhoe et al. 2018, entire; Kupfer et al. 2020, entire). Despite 
the recognition of climate effects on ecosystem processes, there is 
some uncertainty about the timing of these effects for the Southeastern 
United States and how the gopher tortoise will respond to these 
changes. Factors associated with a changing climate may act as risk 
multipliers by increasing the risk and severity of other threats, as 
described in Synergistic and Cumulative Effects, below.

Habitat Management

    As mentioned previously, the gopher tortoise needs large swaths of 
interconnected, high-quality habitat patches with open canopy and 
abundant herbaceous groundcover to support viable populations, and a 
variety of land management practices are used in the restoration, 
enhancement, and maintenance of gopher tortoise habitats. Insufficient 
habitat management (e.g., no prescribed fire program) has been 
identified as a major threat to the gopher tortoise (Smith et al. 2006, 
pp. 326-327). High-quality gopher tortoise habitat will require 
prescribed fire only at regular intervals, while areas of degraded or 
low-quality gopher tortoise habitat will require more active habitat 
management (e.g., multiple habitat management tools including 
mechanical and chemical treatments in conjunction with the 
reintroduction of prescribed fire to restore natural conditions). 
However, not all habitat management activities are uniformly beneficial 
to the gopher tortoise. In general, management actions that minimize 
soil disturbance, protect burrows, and maintain a diversity of 
groundcover plants, to ensure that sufficient sunlight reaches the 
ground, are beneficial to the gopher tortoise. Conversely, actions that 
cause significant soil disturbances or result in the loss of diverse 
groundcover are detrimental to the species. A variety of habitat 
management methods are implemented rangewide at varying degrees across 
land ownership and use types (e.g., conservation land, commercial 
forestry, family-owned lands, etc.). Prescribed fire, selective use of 
herbicide, mechanical vegetation management (e.g., roller chopping and 
mowing), and timber harvest are valuable management techniques in the 
restoration, management, and maintenance of gopher tortoise habitat and 
are frequently used in combination to achieve habitat condition goals.
    The regular application of prescribed fire is important for the 
maintenance of habitat conditions required by the gopher tortoise. When 
applied at appropriate intervals, prescribed fire reduces shrub and 
hardwood encroachment, and stimulates growth of forage plants such as 
grasses, forbs, and legumes, particularly when applied during the 
growing season (Thaxton and Platt 2006, p. 1336; FWC 2007, p. 32; Iglay 
et al. 2014, pp. 39-40; Fill et al. 2017, pp. 156-157). In addition, a 
more open canopy and midstory created with the use of prescribed fire 
allows for proper incubation of eggs and thermal regulation (basking) 
of tortoises. Without habitat management including fire management, 
gopher tortoises may abandon an area of previously suitable habitat 
after as little as 20 years of fire exclusion (Ashton et al. 2008, p. 
528). In the future, reduced habitat management is expected to result 
in habitat degradation or loss, negatively impacting the gopher 
tortoise.
    Mechanical or chemical (herbicide) management techniques may be 
needed to reduce hardwood competition to levels where prescribed fire 
can be effective and are increasingly important for areas where 
prescribed fire use is not a viable option, such as habitat in 
urbanized areas (Ashton and Ashton 2008, p. 78; Miller and Chamberlain 
2008, pp. 776-777; Jones et al. 2009, p. 1168; Iglay et al. 2014, p. 
40; Platt et al. 2015, p. 913; Greene et al. 2020, p. 50). Habitat 
management using mechanical means can be effective in reducing shrub 
and tree density to promote conditions favorable to herbaceous 
vegetation. Mechanical treatments are used in habitat restoration, site 
preparation to promote pine seedling survival and growth, maintenance, 
and in other agricultural and forestry endeavors. Mechanical vegetation 
management examples include mulching/chipping, subsoiling, shearing, 
stumping, root raking into piles or windrows, roller chopping, discing, 
and bedding. Depending on management objectives and treatment type, 
mechanical site preparation may result in substantial soil disturbance 
affecting soil structure and chemistry and may increase invasive 
species on a site (Hobbs and Huenneke 1992, pp. 324-325; Jack and 
McIntyre 2017, p. 189). Heavy equipment used to manage gopher tortoise 
habitat may also cause impacts to gopher tortoise through crushing or 
damage to burrows (Landers and Buckner 1981, pp. 1-7; Greene et al. 
2020, p. 54). Some land managers incorporate best management practices 
for gopher tortoise habitat into their management plans, including a 
buffer distance around burrows to minimize disturbance and hazards 
(Smith et al. 2015, pp. 459-460).
    Mechanical vegetation management followed by herbicide application 
is used as a short-term option to maintain habitat in areas where fire 
use is restricted. Herbicide can reduce midstory vegetation growth 
resulting in more sunlight reaching the ground. Although mechanical 
vegetation management is effective in reducing the vertical structure 
and overgrowth in the mid- and overstories, mechanical treatments alone 
do not replicate the stimulation of plant growth, flowering, and seed 
release, and soil nutrient cycling provided by fire (Dean et al. 2015, 
pp. 55-56). Best conservation practices for mechanical and herbicide 
management practices in gopher tortoise habitat are available for 
landowners and managers and are increasingly implemented (FWC 2013, 
entire; Service 2013, entire; GDNR 2014, entire; Florida Department of 
Agriculture and Consumer Services (FDACS) 2014, entire; FDACS 2015, 
entire; Jack and McIntyre 2017, p. 200).
Forest (Timber) Management
    Management of forests, either public or private, influences habitat 
where gopher tortoises occur or habitat that may be suitable for gopher 
tortoises. Although specific forest or timber management techniques 
vary by site, management goals, and ownership, we summarize the 
influence of forest or timber management in general on gopher tortoise 
below. More details and information on this influence may be found in 
the SSA section 3.8.4 Timber Management (Service 2022, pp. 76-79).
    Not all forested lands provide appropriate conditions for gopher 
tortoises. However, forests on lands with suitable soils and compatible 
forest

[[Page 61844]]

management objectives in the gopher tortoise range can be managed in 
such a way as to provide the open canopy and the dense herbaceous 
groundcover conditions needed for gopher tortoise viability. Some types 
of timber and gopher tortoise habitat management include the reduction 
of hardwood competition. This activity results in reduced tree density 
and increased sunlight, promoting herbaceous forage proliferation and 
suitable conditions for gopher tortoise basking and egg incubation 
(NRCS 2020, entire). Several management practices associated with 
working forests, such as planting densities, rotation length, and time 
until first and subsequent thinning(s), have a direct influence on 
whether these lands provide and maintain habitat for the species. 
Gopher tortoises occur in production pine forests with suitable 
conditions, although at lower densities than reported in other cover 
types, and densities may be below the threshold necessary to sustain a 
viable population (Diemer-Berish et al. 2012, pp. 51-52; Wigley et al. 
2012, p. 42; Greene et al. 2019, p. 51). In pine forests managed for 
timber or pulp (typically slash or loblolly pine) where suitable 
conditions are not maintained, gopher tortoises more frequently 
abandoned burrows and emigrated from low-quality habitat conditions 
associated with closed canopy pine plantations (Diemer 1992a, p. 288; 
Aresco and Guyer 1999, p. 32). Most modern forests managed more 
intensely for traditional wood products (i.e., timber, pulp) 
incorporate management strategies to maintain open canopy conditions 
for much of the life of a commercial stand (Weatherford et al. 2020, p. 
4). For private lands, programs such as forest certifications (e.g., 
Sustainable Forestry Initiative (SFI) or Forest Stewardship Council) 
and the development of diversified markets for forest products have 
increased forest management practices that benefit gopher tortoises 
(Greene et al. 2019, p. 201; Greene et al. 2020, p. 55).
    Public lands managed for multiple use or conservation objectives 
that include timber production employ some of the same habitat 
management techniques and additionally may be guided by land management 
plans or forest plans. The Forest and Rangeland Renewable Resources 
Planning Act (16 U.S.C. 36), as amended by the National Forest 
Management Act of 1976 (16 U.S.C. 1600-1614), requires that each 
National Forest (NF) be managed under a forest plan that is revised 
every 10 years. Forest plans provide an integrated framework for 
analyzing and approving projects and programs, including conservation 
of listed species. Several National Forests (e.g., Ocala NF, Desoto NF, 
Conecuh NF, Apalachicola NF, etc.) occur within the current range of 
the gopher tortoise, providing important habitat conservation for the 
species. Identification and implementation of land management and 
conservation measures to benefit gopher tortoises vary among National 
Forests, but generally include habitat restoration and management 
objectives and maintaining buffers around gopher tortoise burrows 
during various forest management activities.
    However, not all public or private lands are managed to these 
standards, and incompatible practices and insufficient management 
continue to affect gopher tortoise habitat and influence gopher 
tortoise viability. Reductions in required groundcover forage may be 
caused by nearly complete groundcover weed control, high seedling 
stocking rates, or short timber rotations with a minimal proportion of 
the rotation being open canopied. In addition, exclusion of prescribed 
fire and dense hardwood midstory encroachment within open canopied 
forests degrade habitat through suppression of groundcover and loss of 
open areas for burrowing and movement.
    Historical declines of longleaf forests are well established, with 
estimates of 95 percent loss from the historical estimate of 88 million 
ac (35.6 million ha) (Oswalt et al. 2012, p. 13). However, the 
magnitude and extent of insufficient and incompatible forestry and 
timber management currently occurring on the landscape and impacting 
gopher tortoise populations and habitat has not been quantified. 
Rangewide, approximately 80 percent of potential gopher tortoise 
habitat occurs in private ownership, with the remainder owned or 
managed by local, State, Federal, or private conservation entities 
(Wear and Greis 2013, p. 103; Natural Resources Conservation Service 
(NRCS) 2018, p. 2). Private landowners hold more than 86 percent of 
forests in the South and produce nearly all of the forest investment 
and timber harvesting in the region (Most of the potential gopher 
tortoise habitat is privately held, and much of this is in 
silviculture. Rangewide conservation and management efforts between 
private landowners and conservation agencies, such as best conservation 
practices for gopher tortoises developed by States and conservation 
incentive programs and partnerships, promote compatibility between 
timber and gopher tortoise management; these are further described in 
Conservation Efforts and Regulatory Mechanisms, below. We have included 
the best available information regarding gopher tortoises in timber 
production pine forests in our SSA; however, to date, systematic 
surveys in pine forests intensively managed for timber and pulp 
products across the range of the gopher tortoise have not been 
conducted.

Other Factors--Disease, Predation, Harvest and Roundups, Nonnative 
Invasive Species

Disease
    A number of diseases, including fungal, viral, bacterial, and 
parasitic diseases, have been documented in gopher tortoises (Ashton 
and Ashton 2008, pp. 39-41; Johnson et al. 2008, entire; Myers et al. 
2009, p. 582; Desiderio et al. 2021, entire). Upper Respiratory Tract 
Disease (URTD) resulting from two bacterial species (Mycoplasma 
agassizii and M. testudineum) has been documented throughout much of 
the tortoise's range (McLaughlin 1997, p. 6; Gates et al. 2002, entire; 
Rabatsky and Blihovde 2002, entire; Dziadzio et al. 2018, entire; 
Goessling et al. 2019, pp. 5-6). While large-scale die-offs due to URTD 
appear to be rare, correlations between exposure to Mycoplasma spp. and 
population declines are variable among populations (McCoy et al. 2007, 
p. 173). URTD has been linked to several large mortality events 
(defined as the loss of greater than 3 percent of adults in 1 year) in 
Florida with an estimated loss of 25-50 percent of the adult population 
in one event and 35 to 125 adults in other events (McLaughlin 1997, p. 
6; Gates et al. 2002, entire; Rabatsky and Blihovde 2002, entire; 
Dziadzio et al. 2018, entire). However, tortoises have natural 
antibodies to Mycoplasma spp., and these natural immune mechanisms may 
explain why die-offs are less prevalent rangewide than may be expected 
from the degree of seroprevalence in gopher tortoise populations 
(Hunter et al. 2008, p. 464; Gonynor and Yabsley 2009, pp. 1-2; 
Sandmeier et al. 2009, pp. 1261-1262). In addition, URTD may result in 
altered movement (e.g., increased dispersal) and behavior (e.g., 
changes to basking) among gopher tortoises (McGuire et al. 2014, pp. 
750-754; Goessling et al. 2017, p. 488). Tortoises dispersing long 
distances increase their likelihood of encountering a road (i.e., a 
barrier), potentially limiting spread of disease but increasing risk of 
road mortality. The magnitude of threat that URTD poses to gopher 
tortoise populations and tortoise demographics is currently

[[Page 61845]]

unknown, but the best available science indicates it is not a key 
factor in species viability (Karlin 2008, p. 145).
Predation
    Gopher tortoise nest predation varies annually and across sites, 
ranging from approximately 45 to 90 percent in a given year (Landers et 
al. 1980, p. 358; Wright 1982, p. 59; Marshall 1987, pp. 29-32). Gopher 
tortoises are most susceptible to predation within their first year of 
life, primarily within 30 days of hatching (Pike and Seigel 2006, p. 
128; Smith et al. 2013, pp. 4-5). Overall annual hatchling survival has 
been estimated to be approximately 13 percent (Perez-Heydrich et al. 
2012, p. 342). Raccoons (P. lotor) are the most frequently reported 
predator of nests and juvenile gopher tortoises (Landers et al. 1980, 
p. 358; Butler and Sowell 1996, p. 456). However, 25 species--12 
mammals, 5 birds, 6 reptiles, and 2 invertebrates--are known to be 
predators of eggs, emerging neonates, hatchlings, and older tortoises 
(Ashton and Ashton 2008, p. 27). Adult gopher tortoises are less likely 
to experience predation compared to hatchlings and eggs, but predation 
by canines (e.g., domestic dogs, coyotes, foxes) and humans has 
occurred (Causey and Cude 1978, pp. 94-95; Taylor 1982, p. 79; Hawkins 
and Burke 1989, p. 99, Mann 1995, p. 24). Some predation can be 
attributed to habitat fragmentation and edge effects, roads and 
infrastructure, increased availability of food for predators in 
proximity to human-inhabited areas, reduction or elimination of top 
canid carnivores, ecological perturbations allowing predator range 
expansion, and domestic animals associated with humans (Stiles and 
Jones 1998, p. 343; Crooks and Soule 1999, entire; Wetterer and Moore 
2005, pp. 352-353).
    As mentioned previously, the gopher tortoise is a long-lived 
species that naturally experiences high levels of mortality in early 
life stages. However, as urbanization increases in the future, we 
expect that higher levels of hatchling and juvenile mortality 
associated with increased predation near anthropogenic sites will have 
a negative impact on gopher tortoise recruitment in affected 
populations.
Harvest and Rattlesnake Roundups
    Historical harvest of gopher tortoises for consumption has 
influenced gopher tortoise populations in the past, particularly in 
portions of the Florida panhandle (Lohoefener and Lohmeier 1984, pp. 1-
30; Mann 1995, p. 18; Estes and Mann 1996, p. 21; Tuma and Sanford 
2014, pp. 145-146). Although this practice is now uncommon, localized 
harvest still occurs in some rural areas (Rostal et al. 2014, p. 146). 
Although loss of individuals may impact affected populations, we have 
determined that harvest is not a significant species-level threat to 
the gopher tortoise (Service 2022, p. 63).
    Historically, multiple rattlesnake roundups were held throughout 
the Southeast (Means 2009, p. 132). Snakes were collected by blowing 
fumes of noxious liquids (``gassing'') in gopher tortoise burrows to 
collect snakes for these roundups. Gassing of inhabited burrows 
negatively impacts the resident tortoise, though research that 
quantifies mortality associated with this practice is limited (Means 
2009, p. 139). The practice of gassing tortoise burrows is now 
prohibited across the species' range. Gopher tortoise mortality due to 
rattlesnake collection is primarily historical and is not likely a 
significant current influence on populations, as only one roundup still 
takes place in Alabama and the use of gasoline or other chemical or 
gaseous substances to drive snakes from burrows is now prohibited 
across the Southeast (Alabama Regulation 220-2-.11, Georgia codes 
sections 27-1-130 and 27-3-130, Florida Administrative Code 68A-
4.001(2), and Mississippi Code R 5-2.2 B). Therefore, harvest and take 
resulting from rattlesnake roundups are considered historical threats 
to the species, and the best available science indicates these are not 
current threats to the species.
Nonnative Invasive Species--Flora and Fauna
    The spread of nonnative invasive plant species alters and degrades 
gopher tortoise habitat by reducing forage quality and quantity and the 
availability of burrowing and nesting locations, and ultimately 
influences gopher tortoise viability. Some species postulated to impact 
tortoise habitat include kudzu (Pueraria montana), Chinese privet 
(Ligustrum sinense), Callery pear (Pyrus calleryana), natal grass 
(Melinis repens), and Japanese climbing fern (Lygodium japonicum), 
though quantified impacts of these species on tortoises are unknown. 
One species known to impact gopher tortoise use of habitat is 
cogongrass (Imperata cylindrica), a prolific invasive that occurs 
throughout much of the gopher tortoise's range. Unlike other invasive 
plant species in upland communities, cogongrass can rapidly spread 
following disturbances including prescribed fire (Yager et al. 2010, 
entire; Holzmueller and Jose 2011, pp. 436-437). It can quickly form a 
tall, dense ground cover with a dense rhizome layer and can outcompete 
native vegetation (Dozier et al. 1998, pp. 737-740; Mushinsky et al. 
2006, p. 360; Minogue et al. 2018, pp. 1-4). Widespread areas of dense 
cogongrass could result in habitat loss as gopher tortoises do not use 
these areas, nor do they consume cogongrass (Basiotis 2007, p. 21). 
Cogongrass can also decrease gopher tortoise habitat quality by 
reducing forage quality and quantity and the availability of burrowing 
and nesting locations (Lippincott 1997, pp. 48-65; Basiotis 2007, p. 
24).
    Nonnative invasive fauna can also negatively influence the gopher 
tortoise and its habitat. Throughout the gopher tortoise's range, the 
red imported fire ant (Solenopsis invicta) occurs in disturbed soil in 
upland habitats (Wetterer and Moore 2005, p. 352; Shearin 2011, pp. 22, 
30; USDA 2017, unpaginated). Fire ants are not able to breach gopher 
tortoise eggs, but the ants will depredate hatchlings (Mann 1995, p. 
24; Butler and Hull 1996, p. 17; Epperson and Heise 2003, p. 320; 
Diffie et al. 2010, p. 295; Dziadzio et al. 2016, pp. 531, 536). Fire 
ants are aggressive, and their stings can result in direct mortality 
and reduced survival by limiting growth, altering behavior, and 
changing foraging patterns of hatchlings (Wilcox and Giuliano 2014, pp. 
3-4; Dziadzio et al. 2016, pp. 532-533). In the western portion of the 
range, gopher tortoise conservation banks and other related sites must 
include fire ant monitoring and control as part of their management 
plan to reduce the effects of predation on tortoise eggs and hatchlings 
(74 FR 46401, September 9, 2009).
    The nine-banded armadillo (Dasypus novemcinctus), Argentine black 
and white tegu (Salvator merianae), Burmese python (Python bivittatus), 
and black spiny-tailed iguana (Ctenosaura similis) use gopher tortoise 
burrows and are known predators of tortoise eggs (Service 2022, pp. 68-
69). Frequent damage to gopher tortoise burrows by wild pigs (Sus 
scrofa), domestic dogs (Canis lupus familiaris), and possibly domestic 
cats (Felis catus) may impact some gopher tortoises as well.
    The current impact of these nonnative invasive floral and faunal 
species on gopher tortoise appears low at the species level. Although 
impacts to individuals and populations have been documented to occur, 
we did not find nonnative invasive species to be a key factor in gopher 
tortoise viability.

Conservation Efforts and Regulatory Mechanisms

    In this section, we describe key protections and conservation 
efforts

[[Page 61846]]

provided by various Federal and State entities, private landowners, and 
nongovernmental organizations. Additional information regarding 
conservation efforts and Federal and State protections may be found is 
the SSA report (Service 2022, pp. 79-102).
Federal and State Protections
    In addition to the protections provided to the gopher tortoise in 
the listed portion of the range under sections 7 and 10 of the Act, we 
implement conservation delivery tools and programs that aid in the 
conservation of listed and at-risk species, such as the gopher 
tortoise, on non-Federal lands. Cooperative conservation programs such 
as the Partners for Fish and Wildlife Program provide technical and 
financial assistance to private landowners and others for the 
conservation of wildlife and associated habitat. Between 2010 and 2019, 
under the Partners for Fish and Wildlife Program, approximately 65,000 
ac (26,305 ha) of restoration and enhancement activities were 
implemented in gopher tortoise habitat on private lands in Alabama, 
Florida, Georgia, and Mississippi (Service 2020, unpaginated).
    The Gopher Tortoise Conservation and Crediting Strategy (Strategy) 
is a conservation initiative designed to balance military mission 
activities and gopher tortoise conservation on Department of Defense 
(DoD) lands in the Southeast (Service 2017, entire); see below under 
Conservation Lands for further discussion about DoD lands. The Service-
approved Strategy establishes the framework for determining credit for 
DoD conservation actions and is intended to achieve a net conservation 
benefit to the species. It focuses on identification, prioritization, 
management, and protection of viable gopher tortoise populations and 
the best remaining habitat. It provides guidelines designed to result 
in an increase in the size and/or carrying capacity of populations 
while promoting the establishment of new populations through increased 
habitat connectivity or translocation of gopher tortoises (Service 
2017, entire).
    The U.S. Department of Agriculture Natural Resources Conservation 
Service (NRCS) offers technical and financial assistance to help 
agricultural producers voluntarily implement conservation activities 
and practices that benefit the gopher tortoise. The gopher tortoise is 
identified as a target species eligible for conservation funding in the 
national Working Lands for Wildlife partnership, which is a 
collaborative approach to conserving habitat on working lands. In 
addition, the NRCS works to restore longleaf pine across its historical 
range through the Longleaf Pine Initiative. Between 2012 and 2021, 
private landowners across the range of the species have received 
assistance to implement management practices that benefit gopher 
tortoises and gopher tortoise habitat on 943,740ac (381,918ha) through 
NRCS programs.
    Each State within the range of the gopher tortoise provides some 
measure of protection for the species. The States of Florida, Georgia, 
and South Carolina provide protection for the gopher tortoise through 
the requirement of land management plans for State lands. The gopher 
tortoise is protected by regulation as a non-game species in Alabama, 
is State-listed as threatened in Florida, Georgia, and Louisiana, and 
is State-listed as endangered in Mississippi and South Carolina. Gopher 
tortoise protections vary by State; however, laws within most States in 
the range focus on prohibitions against the take, possession, export/
sale, and killing of gopher tortoises. States in the gopher tortoise 
range also implement conservation programs in partnership with private 
landowners. For example, Florida's Landowner Assistance Program assists 
private landowners with plans to improve their wildlife habitat through 
the development of 10-year management plans on an estimated 44,000 ac 
(17,806 ha) of gopher tortoise habitat per year (FWC 2020b, p. 6). 
Florida has also developed the Gopher Tortoise Management and Gopher 
Tortoise Permitting Guidelines to guide gopher tortoise recovery 
efforts and regulatory actions (FWC 2007, revised 2012, entire; FWC 
2008, revised July 2020; entire). Florida regulations also require that 
construction or other activities that disturb gopher tortoise burrows 
must obtain a relocation permit and that the impacts be considered and 
mitigated.
Translocation and Headstarting
    Gopher tortoises have been considered one of the most translocated 
species in the Southeast, and translocation is commonly used as a 
conservation strategy to mitigate the loss of tortoises from land under 
development (Dodd and Seigel 1991, p. 340). Displaced tortoises are 
often translocated to suitable habitat to reestablish extirpated 
populations or augment existing populations (Griffith et al. 1989, p. 
477). Numerous studies have attempted to evaluate the success of gopher 
tortoise translocation and improve its efficacy. However, gopher 
tortoise life history characteristics (e.g., long-lived, slow-growing, 
and slow to reach maturity) make it difficult to determine if 
translocations result in sufficiently viable tortoise populations since 
the typical monitoring periods are shorter than the generation time for 
the species. Gopher tortoises disperse at a high rate in the year 
following translocation; however, soft-releases, or the temporary 
penning of gopher tortoises within a recipient area, are highly 
effective at limiting dispersal post-translocation (Tuberville et al. 
2005, pp. 353-354; Tuberville et al. 2008, pp. 2694-2695; Bauder et al. 
2014, pp. 1449-1450). Translocation is successful at removing tortoises 
from immediate danger due to development (Tuberville et al. 2005, p. 
356; Tuberville et al. 2008, p. 2695).
    Gopher tortoise relocation and translocation practices are being 
implemented and included as guidance across the range of the species 
(Service 2022, pp. 85-87). The primary goals for recipient sites are to 
prevent the loss of tortoises and retain the existing tortoises; and 
while habitat is lost on the development site, recipient sites can 
contribute to habitat conservation if sites receive long-term 
protection and subsequent habitat management. These sites can provide 
high conservation value by restocking tortoises to appropriately 
suitable lands where populations have previously been depleted. 
However, this practice could result in an overall net loss of habitat 
if not implemented in conjunction with acquisition and additional 
protection of habitat when needed. Additional information regarding 
specific translocation efforts in each State may be found in the SSA 
report (Service 2022, pp. 83-87).
    Headstarting, or the process of hatching and/or rearing juvenile 
turtles in captivity through their most vulnerable period, has shown 
success as a technique to boost depleted gopher tortoise populations 
(Holbrook et al. 2015, pp. 542-543; Tuberville et al. 2015, pp. 467-
468; Spencer et al. 2017, p. 1341; Quinn et al. 2018, p. 1552; 
Tuberville et al. 2021, p. 92). Headstarting has been explored as a 
management tool for the gopher tortoise with increasing recognition of 
its potential role, particularly when used in concert with other 
management actions (Spencer et al. 2017, entire; Quinn et al. 2018, pp. 
1552-1553). For example, the gopher tortoise headstarting program at 
Camp Shelby in Forrest County, Mississippi (funded by the Mississippi 
Army National Guard and in partnership with The Nature Conservancy) has 
been ongoing since 2013 and has shown initial success with headstarted 
juveniles surviving at a

[[Page 61847]]

much higher rate than their wild counterparts (70-80 percent versus 30 
percent for wild 2- to 3-year-old tortoises). Similar survival rates 
were noted in post-release monitoring of headstarted yearling gopher 
tortoises in Georgia and South Carolina (Tuberville et al. 2015, 
entire).
Other Conservation Mechanisms
    In the eastern portion of the range, the gopher tortoise is 
included in a candidate conservation agreement (CCA) (revised 2018) 
with State, nongovernmental and private organizations and in a 
candidate conservation agreement with assurances (CCAA) (2017) with 
Camp Blanding Joint Training in Florida. These Service-approved 
agreements outline management actions that landowners implement to 
benefit the gopher tortoise and its habitat across the candidate range. 
We developed the 2013 Rangewide Conservation Strategy for the Gopher 
Tortoise to guide conservation of the gopher tortoise by our partners, 
including States within gopher tortoise range, the Service, and other 
public and private entities to collect and share information on gopher 
tortoise threats, outline highest priority conservation actions, and 
identify organizations best suited to undertake those conservation 
actions (Service 2013, entire).
    In Florida, where the greatest number of tortoises have been 
identified, several additional conservation efforts are ongoing. The 
Forestry Wildlife Best Management Practices for State Imperiled Species 
and the Agriculture Wildlife Best Management Practices for State 
Imperiled Species were developed in 2014 and 2015, respectively, to 
enhance silviculture's contribution to the conservation of wildlife, 
provide guidance to landowners who chose to implement these voluntary 
practices, and reduce take of gopher tortoises (FDACS 2015, entire). By 
2021, landowners provided notice of intention to FWC to implement 
forestry best management practices (BMPs) on more than 3.7 million ac 
(1.5 million ha) and conservation practices on approximately 425,031 ac 
(172,004 ha) of agricultural lands in Florida (FWC 2020a, unpaginated; 
FWC 2021, p. 1). FWC also provides technical assistance to private and 
industry landowners to implement beneficial management and/or 
mitigation activities across 40 counties through other programs and 
agreements (FWC 2020b, p. 2; FWC 2021, p. 1).
    There are numerous other gopher tortoise conservation tools and 
guides, including several in the core of the species' range in Georgia. 
For example, the Best Conservation Practices for Gopher Tortoise 
Habitat on Working Forest Landscapes was developed to assist in best 
conservation practices for the creation and maintenance of gopher 
tortoise habitat in the candidate portion of the range (GDNR et al. 
2018, entire). Additionally, Forest Management Practices to Enhance 
Habitat for the Gopher Tortoise details the essentials of managing 
habitat for gopher tortoises, including prescribed fire, timber 
harvest, and selective herbicide use (GDNR 2014, unpaginated). Further, 
the Georgia Gopher Tortoise Initiative is an extension of the Georgia 
Department of Natural Resource's long-standing effort in conserving 
longleaf pine systems. The initiative is a collaborative effort between 
several public and private entities and is geared towards the 
protection, restoration, and long-term management of gopher tortoise 
habitat.
    Implemented rangewide, America's Longleaf Restoration Initiative is 
a collaborative effort involving multiple public and private partners 
actively supporting efforts to restore and conserve longleaf pine 
ecosystems with a goal to increase longleaf coverage on the landscape 
to 8.0 million ac (3.2 million ha) (ALRI 2021, unpaginated). Several 
local implementation teams work across the gopher tortoise range to 
help restore longleaf pine on habitat where gopher tortoises occur.
Conservation Lands
    The conservation of multiple large, contiguous tracts of habitat 
provides the connectivity and landscape heterogeneity requirements to 
support gopher tortoise viability. Gopher tortoise habitat occurs 
across a wide range of lands in public ownership with varying levels of 
management. An estimated 1.7 million ac (688,000 ha) of potential 
gopher tortoise habitat occurs on protected lands including lands in 
Federal, State, and local government, nongovernmental organization, and 
private ownership (e.g., conservation easements) throughout the 
species' range.
    Managing publicly owned lands in a way that benefits the gopher 
tortoise is an important mechanism for reducing the effects of habitat 
loss, fragmentation, and degradation on the species. Habitat management 
occurring on public conservation lands is often accomplished via 
natural resource planning instruments (e.g., land management plans, 
comprehensive conservation plans, resource management plans, etc.). 
Each State in the gopher tortoise's range has statutory authority to 
acquire land for conservation purposes. Since publication of the 12-
month finding (76 FR 45130, July 27, 2011), all States within the 
species' range have made concerted efforts to protect gopher tortoise 
habitat and potential gopher tortoise habitat via strategic land 
acquisition. Between 2011 and 2019, Alabama, Florida, Georgia, and 
South Carolina have reported fee-simple acquisition of approximately 
42,000 ac (16,996 ha) of potential gopher tortoise habitat with an 
additional approximately 78,000 ac (31,565 ha) acquired in conservation 
easements (CCA 2019, pp. 52-73). Federal entities including the U.S. 
Air Force, the U.S. Forest Service, and the Service recorded an 
additional 2,740 ac (1,109 ha) of potential gopher tortoise habitat 
acquired and approximately 24,000 ac (9,712 ha) of conservation 
easements acquired (CCA 2019, pp. 52-73).
    Several National Wildlife Refuges (NWRs) (e.g., Merritt Island NWR, 
Lake Wales Ridge NWR, Lower Suwannee NWR, St. Marks NWR) occur within 
the range of the gopher tortoise, providing important habitat 
conservation for the species. Management activities included in NWR 
Comprehensive Conservation Plans that influence gopher tortoises 
include habitat restoration activities such as prescribed fire, pine 
thinning, and other mechanical vegetation management for restoring 
desired vegetative conditions in pine and scrub systems, and tortoise 
management and monitoring actions based on priorities of the refuge and 
available resources.
    Rangewide, the gopher tortoise occurs on 31 DoD installations, with 
potential habitat on additional installations (DoD 2022, p. 4). Many of 
these installations specifically include gopher tortoise habitat and 
population management prescriptions and goals within their individual 
integrated natural resources management plans (INRMPs) prepared in 
conjunction with the Service. Most INRMPs also include land management 
for other upland species that benefit gopher tortoise habitat (and 
gopher tortoises) as well. Rangewide, approximately 830,000 ac (335,889 
ha) of potential gopher tortoise habitat occur on military 
installations. Limited information is currently available regarding the 
condition of this potential habitat and the extent to which these areas 
are occupied by gopher tortoises.
    National Forest (NF) plans provide an integrated framework for 
analyzing and approving projects and programs, including conservation 
of listed species. Several National Forests (e.g., Ocala NF, Desoto NF, 
Conecuh NF, Apalachicola NF, etc.) occur within the range of the gopher 
tortoise and provide important habitat conservation for the species. 
Identification and implementation of

[[Page 61848]]

land management and conservation measures to benefit gopher tortoises 
vary among NFs, but generally include habitat restoration and 
management objectives and maintaining buffers around gopher tortoise 
burrows during various forest management activities. For example, the 
Desoto NF recently completed a 10-year Collaborative Forest Landscape 
Restoration Program, during which actions to restore longleaf pine were 
implemented on 374,000 ac (151,352 ha) of NF lands. In addition, the 
Desoto NF has prioritized any management treatment that contributes to 
improvement of gopher tortoise, as set forth in their Mission, Vision, 
and Operational Strategy (USFS 2020, entire).
Private Lands Conservation Efforts
    Most forested land within the gopher tortoise range is privately 
owned. Privately owned lands account for approximately 80 percent of 
potential gopher tortoise habitat, of which approximately half are 
managed for forest production (NRCS 2018, p. 2; Greene et al. 2019, p. 
201). Across the gopher tortoise range, large working forests account 
for over 6 million ac (2.4 million ha) of forest land, representing a 
significant land use with the potential to influence gopher tortoise 
resiliency and viability (Weatherford et al. 2020, p. 3). While not all 
working forest lands include appropriate habitat conditions for gopher 
tortoises, approximately 2.78 million ac (1.12 million ha) of suitable 
soil types and 2.98 million ac (1.21 million ha) of open pine 
conditions are estimated to occur on private forest lands (NCASI 2021, 
p. 1). We included the best available data on gopher tortoise 
observations between 1977 and 2019 on private forest lands in our SSA 
(Weatherford et al. 2020, pp. 9-11; Service 2022, pp. 95-99). These 
observations occur on Member Company lands that are part of the 
National Council for Air and Stream Improvement and landowners may 
implement conservation measures including those outlined in the 
Sustainable Forestry Initiative guidelines.
    While working to meet a range of objectives, including timber 
production, many larger private working forests also accomplish 
conservation within a broad network of collaboration with Federal, 
State, and local government agencies, universities, and nongovernmental 
organizations. For example, forest landowners may create and maintain 
areas of open pine conditions, conduct gopher tortoise burrow surveys, 
conduct research, and implement BMPs that benefit the gopher tortoise. 
In addition, forest certification programs, such as the Sustainable 
Forestry Initiative (SFI) and Forest Stewardship Council, require 
participants to adhere to a set of principles including providing 
wildlife habitat to conserve biological diversity (Weatherford et al. 
2020, p. 11). Adhering to these principles likely provides a benefit to 
maintaining suitable gopher tortoise habitat in private working 
forests. An estimated 13.7 million ac (5.5 million ha) within the 
gopher tortoise's range are certified through SFI, although the 
proportion of certified acres that include gopher tortoise populations 
or their current habitat is unknown (SFI 2021, unpaginated). Other 
forest certifications, including the American Tree Farm System, are 
authorized by the Program for the Endorsement of Forest Certification, 
a third-party audited certification system.
    The largest forest landowner group in the United States is the 
family forest landowners, controlling approximately 87 percent of 
forest land in the South (Oswalt et al. 2014, p. 6). The American 
Forest Foundation works with smaller, family forest landowners and has 
partnered with the Service's Partners for Fish and Wildlife Program to 
develop habitat improvement plans as part of a 10-year agreement. Since 
2017, the partnership has implemented habitat management activities on 
more than 3,500 ac (1,416 ha) and identified 762 gopher tortoises, 
including 2 populations that meet the MVP criteria (AFF 2021, 
unpaginated).
    Additionally, The Longleaf Alliance works with private landowners 
and other partners across the range of the gopher tortoise to restore 
and maintain habitat as an essential part of their larger focus in 
restoring the longleaf pine ecosystem. Through The Longleaf Alliance, 
in 2019, landowners implemented more than 55,000 ac (22,258 ha) of 
prescribed fire within gopher tortoise habitat, in addition to longleaf 
pine plantings, groundcover restoration, and invasive plant management 
efforts (SERPPAS 2020, p. 17).
    Other private conservation efforts include several privately owned 
tracts of land managed as mitigation/conservation areas for gopher 
tortoises in both Mississippi and Alabama, which provide suitable 
habitat, protection, and habitat management. Four conservation areas in 
Alabama are managed through Service-approved habitat conservation 
plans, while the Mississippi conservation bank follows national 
mitigation banking guidelines for maintaining optimal habitat, 
including aggressive prescribed fire and longleaf restoration programs.

Synergistic and Cumulative Effects

    We note that, by using the SSA framework to guide our analysis of 
the scientific information documented in the SSA report, we have not 
only analyzed individual effects on the species, but we have also 
analyzed their potential cumulative effects. We incorporate the 
cumulative effects into our SSA analysis when we characterize the 
current and future condition of the species. To assess the current and 
future condition of the species, we undertake an iterative analysis 
that encompasses and incorporates the threats individually and then 
accumulates and evaluates the effects of all the factors that may be 
influencing the species, including threats and conservation efforts. 
Because the SSA framework considers not just the presence of the 
factors, but to what degree they collectively influence risk to the 
entire species, our assessment integrates the cumulative effects of the 
factors and replaces a standalone cumulative effects analysis.
    Several factors influencing gopher tortoise viability are 
synergistic and related. Urbanization and development results in 
habitat loss, fragmentation, and degradation through land use change 
and increased road infrastructure. The anthropogenic changes associated 
with urbanization and development also affect the gopher tortoise 
through the introduction of nonnative invasive species and predators. 
Climate change is expected to influence the gopher tortoise through 
several changes as described in Climate Change, above. Sea level rise 
is expected to result in an inland migration of the human population 
away from inundated areas, resulting in increased urbanization and 
developed inland areas that are currently undeveloped and potentially 
suitable upland habitat for gopher tortoise. In addition, changes in 
precipitation and temperature are expected to result in a decrease in 
the number of suitable burn days in gopher tortoise habitat, leading to 
reduced habitat management (another threat to gopher tortoise 
viability). Urbanization and development also limit the implementation 
of prescribed burns as a habitat management tool due to safety concerns 
and proximity to inhabited areas.
    Influences on the gopher tortoise that are not considered key 
factors influencing the species' status may exacerbate the effects of 
urbanization, climate change, and habitat management in affected gopher 
tortoise

[[Page 61849]]

populations. Conservation of habitat through land acquisition and 
conservation actions on public and private lands and the retention of 
private forest lands reduces the severity of some of these threats by 
providing protection of habitat across the landscape, maintaining 
connectivity between habitat patches, and increasing the opportunity 
for beneficial habitat management actions now and into the future.

Summary of Factors Influencing the Species

    The best available information regarding the gopher tortoise and 
its habitat indicates that habitat loss, degradation, and fragmentation 
(due to land use changes from urbanization), climate change, 
insufficient and/or incompatible habitat management, and conservation 
actions are the most significant factors influencing gopher tortoise 
viability. Urbanization results in a range of impacts that either 
remove, degrade, or fragment remaining habitat, or impact gopher 
tortoises directly through development. Urbanization brings road 
construction and expansion, which may cause direct mortality of gopher 
tortoises and fragment remaining habitats. In addition, this type of 
development may also create conditions that prove to be beneficial to 
invasive species, serve to increase predators, and establish inadequate 
conditions for fire management. Temperature increases associated with 
long-term climate change are likely to further constrain use of 
prescribed fire through a decrease in the number of suitable burn days.
    Habitat loss resulting from sea level rise associated with climate 
change is a risk for coastal populations of gopher tortoise. Habitat 
management through prescribed fire and other methods is important to 
maintaining suitable habitat conditions, and insufficient and/or 
incompatible habitat management now and in the future, especially based 
on projections in reduction of prescribed fire, impacts the viability 
of gopher tortoise populations. Conservation efforts to benefit the 
gopher tortoise and its habitat implemented by Federal, State, and 
private partners occur across the species' range and influence the 
gopher tortoise condition. These factors are considered to have 
population-level effects and were evaluated further in the current 
condition and future condition analysis.

Current Condition

    We describe the current condition of the gopher tortoise in terms 
of population resiliency and species redundancy and representation. The 
analysis of these conservation principles to understand the species' 
current viability is described in more detail in the gopher tortoise 
SSA report (Service 2022, pp. 103-143).
Data Sources
    To inform the gopher tortoise SSA, we requested, received, and 
reviewed a variety of data including information from State and Federal 
agencies, local governments, and private lands. Data received included 
two general types of information: spatially explicit data with location 
information (typically from conservation lands) and private lands data 
without location information. These data represent a subset of gopher 
tortoises likely to occur on the landscape due to the lack of a 
comprehensive private lands data set from systematic surveys. Data were 
collected using burrow surveys of various methodologies and included 
burrow surveys with and without burrow scoping, and line transect 
distance sampling (Buckland et al. 1993, entire; Thomas et al. 2010, 
entire); some burrow data were submitted with unknown methodology. 
Because data were provided by a variety of sources, contained disparate 
levels of data resolution, and were collected in various ways, we could 
not reliably determine abundance, density, habitat availability, or 
other metrics for all populations.
    All population data provided were integral to evaluating the 
current condition of the gopher tortoise, although different data types 
come with different assumptions and limitations. Data that come from 
standardized and systematic surveys result in spatially explicit burrow 
locations and subsequent population estimates. The use of these 
spatially explicit data allowed us to make more reliable estimates of 
population size; use spatial buffering to delineate populations based 
on species biology; tie site-specific habitat and management factors to 
locations of gopher tortoises; and estimate future parameters, such as 
estimated future abundance of gopher tortoise populations. Most 
spatially explicit data (e.g., burrow locations and subsequent 
population estimates) in our analyses came from assessments of 
populations on lands managed for the conservation of biodiversity or 
natural resources.
    A large percentage of potential gopher tortoise habitat occurs on 
lands in private ownership. To best assess the current and future 
condition of the gopher tortoise, including populations on private 
lands, we developed a landowner questionnaire and used responses to 
estimate population, habitat, and management factors at a county scale 
to ensure privacy for respondents (Service 2022, appendix A). The vast 
majority of the private lands data obtained for the SSA lack a spatial 
component because of issues associated with confidentiality of location 
data; however, this concern does not preclude the use and importance of 
these data in the SSA. Responses represent a small percentage of 
private lands that currently support gopher tortoises, as many private 
landowners express reluctance to share gopher tortoise occurrence data. 
We also included information from a subsequent Florida Forestry 
Association questionnaire in our analyses; however, no population 
estimates were available for these lands, and we were unable to 
estimate current resiliency for populations on these properties.
    Because data received from these questionnaires are not spatially 
explicit, there are limitations to the applicability of the data as it 
relates to delineation of populations, assessment of site-specific 
factors such as habitat quality and quantity and management regimes, 
and use of abundance data in projections of future scenarios. We 
include data from private landowners in the current condition analysis 
as county-level data and also categorize habitat condition based on 
landowner responses. The additional data we received on gopher tortoise 
populations on private lands when developing the SSA informed our 
current condition analysis of gopher tortoise viability and contributed 
to the understanding of species' viability.
    In this finding, we present results of the current and future 
condition analyses for delineated spatially explicit populations as 
described below for clarity and comparison purposes. However, the SSA 
report also presents results for current conditions for county-level 
data following the same analysis methodology (Service 2022, pp. 130-
142). We used spatially explicit data to inform the population model 
used to forecast future scenarios for the gopher tortoise, as described 
below. We did not use county-level data in our future analysis because 
most information in this category lacks abundance data and we could not 
apply spatially based modeling used in future analysis to the default 
county center point. We note that the data included in our current and 
future condition analyses represent a subset of gopher tortoises likely 
to occur on the landscape, as data from private lands were lacking 
(Service 2022, pp. 103-107). Thus, population estimates do not

[[Page 61850]]

represent an assessment of all populations of gopher tortoises, but 
rather represent information that was provided by partners through much 
of the species' range. Given we were able to use only a subset of 
populations that likely occur on the landscape, our future projections 
are likely an underestimate of gopher tortoises on the landscape.
Analysis Unit and Population Delineation
    To assess rangewide representation for gopher tortoise, we 
delineated five analysis units based on genetic differences (identified 
in Gaillard et al. 2017, entire), physiographic regions, and the input 
of species experts (figure 2). The Tombigbee and Mobile Rivers act as a 
boundary between Unit 1 (Western) and Unit 2 (Central) analysis units, 
and the Apalachicola-Chattahoochee Rivers act as a boundary between 
Unit 2 (Central) and Unit 3 (West Georgia) analysis units. Because of 
the high degree of admixture and lack of well-defined boundaries found 
within transitional zones of physiographic regions, we used other 
biogeographic barriers and expert input to delineate boundaries of the 
following units: Unit 3, Unit 4 (East Georgia), and Unit 5 (Florida) 
analysis units. We used U.S. Environmental Protection Agency Level IV 
ecoregions to delineate the boundaries between Units 3 and 4, and Units 
4 and 5 (EPA 2013, unpaginated). We used the Suwanee River to separate 
Units 3 and 5, as this river represents a significant barrier to 
dispersal, and gene flow between these two units is known to be low 
(Gaillard et al. 2017, p. 509). Additional details regarding the 
delineation of analysis units used to analyze the current and future 
condition of the gopher tortoise may be found in the SSA report 
(Service 2022, pp. 111-114).
BILLING CODE 4333-15-P

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[GRAPHIC] [TIFF OMITTED] TP12OC22.001

BILLING CODE 4333-15-C
    In order to analyze gopher tortoise population resiliency, we 
defined populations for the species as contiguous areas surrounding 
known gopher tortoise burrows with habitat conducive to survival, 
movement, and interbreeding among individuals within the area. Using 
survey data from across the range of the gopher tortoise, we delineated 
populations at two spatial scales: local populations and landscape 
populations, as defined below.
    Local populations are geographic aggregations of individuals that 
interact significantly with one another in social contexts making 
reproduction significantly greater between individuals within the 
aggregation than with individuals outside of the aggregation (sensu 
Smallwood 1999, pp. 103, 108). We operationally delineated local 
populations by identifying aggregations of individuals or burrows where 
individuals were clustered together within a 1,968-ft (600-m) buffer to 
the exclusion of other adjacent

[[Page 61852]]

individuals or burrows. Gopher tortoise habitat and demography vary 
across the range; therefore, the 1,968-ft (600-m) buffer represents an 
average and best estimate across geography and habitat variations based 
on a thorough literature search and species expert input (Diemer 1992b, 
p. 161; Guyer et al. 2012, pp. 122, 125, 132, Castellon et al. 2018, p. 
17; Service 2019, entire; Greene et al. 2020, pp. 52-53). We delineated 
656 local gopher tortoise populations with available spatially explicit 
data (table 1). We assumed that some areas were unsuitable for gopher 
tortoise movement or survival and considered those barriers to movement 
when delimiting local populations. These barriers included interstates, 
freeways, and expressways; major rivers and lakes; wetlands; and highly 
urbanized areas (USDOT 2016, unpaginated; ESRI imagery 2021, 
unpaginated).
    Landscape populations are a series of local populations that are 
connected by some form of movement; individuals within a landscape 
population are significantly more likely to interact with other 
individuals within the landscape population than individuals outside of 
the landscape population. Gopher tortoises have been shown to move more 
than 4,921 feet (1,500 m) throughout multiple years, with distances as 
large as 8,802-15,220 feet (2,683-4,639 m) (McRae et al. 1981, p. 172; 
Ott-Eubanks et al. 2003, p. 317; Diemer-Berish et al. 2012, p. 52; 
Guyer et al. 2012, entire; Castellon et al. 2018, entire). We 
operationally delineated landscape populations by identifying local 
populations connected by habitat within an 8,202-ft (2.5-km) buffer 
around each local population. To be most inclusive of local 
populations, we selected a landscape-population buffer consistent with 
the longer gopher tortoise movements observed (McRae et al. 1981, p, 
173; Diemer 1992b, p. 163; Bauder et al. 2014, pp. 1448-1449; Service 
2019, entire). We delineated 253 landscape populations with available 
spatial data (table 1).

 Table 1--Spatially Delineated Local and Landscape Populations of Gopher
                       Tortoises by State in 2021
------------------------------------------------------------------------
                                               Spatially delineated
                                                    populations
                                         -------------------------------
                                               Local         Landscape
------------------------------------------------------------------------
Florida.................................             316             161
Georgia.................................             151              63
Mississippi.............................              99               7
Alabama.................................              77              14
Louisiana...............................               7               5
South Carolina..........................               6               4
    Total:..............................             656           * 254
------------------------------------------------------------------------
* One delineated landscape population falls in both Georgia and Florida
  and is reflected in both States' landscape population total.

Resiliency
    Resiliency describes the ability of a species to withstand 
stochastic events and is associated with population size, growth rate, 
and habitat quality. Highly resilient populations are more likely to 
withstand disturbances such as random fluctuations in fecundity 
(demographic stochasticity), variation in mean annual temperature 
(environmental stochasticity), or the effects of anthropogenic 
activities, such as local development projects. Viability denotes a 
species' ability to sustain populations over a determined timeframe and 
is closely tied with population resiliency and species-level 
representation and redundancy. For gopher tortoise populations to have 
sufficient viability over the long term, they must have an adequate 
number of individuals (population size), be above a particular density 
(population density), and have sufficient genetic exchange between 
local populations to maintain genetic diversity. There must also be 
sufficient habitat that is beneficially managed for gopher tortoise in 
order to support individual and population needs. Population size and 
density are driven by a variety of underlying demographic parameters, 
including fecundity, sex ratio, and survival at various life history 
stages (egg, nest, hatchling, juvenile, and adult survival). Genetic 
diversity is primarily driven by rates of emigration and immigration 
between local populations.
    We relied on the MVP criteria adopted by the Gopher Tortoise 
Council for abundance, area of managed high-quality habitat, sex ratio, 
evidence of recruitment, variability in size and age classes, and no 
major constraints to gopher tortoise movement as described above (GTC 
2013, pp. 2-3). As previously mentioned, the best available data 
contain disparate levels of data resolution, thus we could not reliably 
determine abundance, density, or other metrics for all populations. 
Therefore, we used a burrow conversion factor for properties that 
provided burrow counts and locations, but did not have a corresponding 
abundance estimate. Although there is no single burrow conversion 
factor that would be appropriate for all populations across the range 
of the species, we selected the representative burrow conversion factor 
of 0.4 individuals per burrow to calculate an estimated current 
population size described in gopher tortoise literature (Guyer et al. 
2012, pp. 127, 129-131). The burrow-to-tortoise conversion factor 
allows the burrow count information to give an estimate of tortoises on 
the landscape, although we recognize that variance in burrow abundance 
is related to factors other than the number of tortoises (Burke 1989, 
p. entire; Breininger et al. 1991, pp. 319-320; McCoy and Mushinsky 
1992, pp. 402, 406).
    We used estimated abundance of adult gopher tortoises in a local 
population as a metric for categorical levels of resiliency: high 
(greater than or equal to 250), moderate (51 to 249), and low (fewer 
than 50). These resiliency levels align with the GTC working group's 
categories for viable (high resiliency), primary support (moderate 
resiliency), and secondary support (low resiliency) populations (GTC 
2014, p. 4).
    Current condition abundance estimates are based only on data from 
spatially delineated populations (i.e., do not contain county-level 
data or gopher tortoises that are present but not reported), and these 
estimates substantially underestimate the true number of gopher 
tortoises present across the species' range. Based on available data, 
there are an estimated 149,152 gopher tortoises from 656 spatially 
delineated local populations across the range of the species, with 
local populations categorized as follows: 360 in low condition, 169 in 
moderate condition, and 127 in high condition.

[[Page 61853]]

Resiliency of populations by analysis unit are described below and in 
table 2. Most gopher tortoises are found in the eastern portion of the 
range with Unit 5 (Florida) supporting 47 percent of the estimated 
rangewide population total, and Units 3 (West Georgia) and 4 (East 
Georgia) supporting 26 percent and 19 percent, respectively. Units 1 
(Western) and 2 (Central) support much smaller numbers of gopher 
tortoises, with 2 percent and 6 percent of the estimated rangewide 
population total, respectively, likely driven by differences in soils, 
as discussed earlier in Habitat.

           Table 2--Site-Specific Data Population Factors and Current Resiliency for Spatially Delineated Local Populations of Gopher Tortoise
--------------------------------------------------------------------------------------------------------------------------------------------------------
                                                 Landscape         Local
        Analysis unit             Burrows       populations     populations      Abundance                        Current resiliency
--------------------------------------------------------------------------------------------------------------------------------------------------------
1...........................           8,815              13             106           3,100  Low (94), Moderate (10), High (2).
--------------------------------------------------------------------------------------------------------------------------------------------------------
2...........................           5,809              30             106           8,642  Low (71), Moderate (27), High (8).
--------------------------------------------------------------------------------------------------------------------------------------------------------
3...........................          17,867              55             109          38,947  Low (42), Moderate (24), High (43).
--------------------------------------------------------------------------------------------------------------------------------------------------------
4...........................          20,216              46             124          28,408  Low (35), Moderate (58), High (31).
--------------------------------------------------------------------------------------------------------------------------------------------------------
5...........................          24,783             109             211          70,055  Low (118), Moderate (50), High (43).
--------------------------------------------------------------------------------------------------------------------------------------------------------
Rangewide...................          77,490             253             656         149,152  Low (360), Moderate (169), High (127).
--------------------------------------------------------------------------------------------------------------------------------------------------------

    We relied on gopher tortoise abundance to assess resiliency of 
populations as the abundance of individuals strongly reflects the 
condition of the habitat and implementation of beneficial management 
actions. We summarize our assessment of habitat condition and 
management actions below and provide more details regarding information 
used and analysis unit results in the SSA report (Service 2022, pp. 
122-130). The influence of habitat size, quality, and management on the 
resiliency and viability of gopher tortoise populations was also 
described in the MVP criteria (GTC 2013, p. 2).
    Habitat data were provided by a variety of sources and contain 
disparate levels of data resolution; thus, we could not reliably 
determine estimates of habitat within all populations across the range 
of the gopher tortoise. Estimates of habitat with known gopher tortoise 
occurrences (local populations) and potential habitat (outside local 
populations, but within the species' range) are derived from the 
species-specific Habitat Suitability Index (HSI) and suitable soils 
(Crawford et al. 2020, entire). Rangewide, we determined using the HSI 
that approximately 844,912 ac (341,923 ha) of suitable habitat occur 
within spatially explicit local populations with gopher tortoise 
occurrences and approximately 16,338,932 ac (6,612,131 ha) of potential 
habitat (suitable habitat with unknown gopher tortoise presence) occur 
outside delineated populations within the range of the species. 
Additionally, information from the landowner questionnaire was used to 
estimate the condition of potential habitat in each analysis unit with 
24 percent of the 447,340 ac (181,032 ha) characterized as low 
condition, 42 percent as moderate condition, and 34 percent as high 
condition (Service 2022, p. 126). Estimates of habitat were not used to 
assess resiliency of gopher tortoise populations; only abundance was 
used to assess resiliency. However, estimates of potential habitat and 
potential habitat quality on private lands give some information 
regarding the extent of habitat where gopher tortoises could occur 
compared to the extent of habitat where occurrences are known.
    To assess management of gopher tortoise habitat, we used several 
data sets available from multiple sources and at multiple spatial 
scales, and these data may include some overlap. Again, we did not use 
any management metrics in our resiliency assessment; only abundance was 
used to assess population resiliency. We determined an estimate of 
acres burned (prescribed fire and wildfire) using Tall Timbers 
Southeast fire history dataset, derived from the U.S. Geological Survey 
Burned Area (v2) Products (Hawbaker et al. 2020, entire) representing 
years 1994-2019 (Hawbaker et al. 2020, entire). Acres burned across all 
units have generally increased over time, with significantly more 
burning occurring in Unit 5 (Florida).
    We also used summary data for prescribed fire and other midstory 
maintenance activities available from America's Longleaf Restoration 
Initiative (ALRI) FY2019 annual report (ALRI 2019). Florida reported by 
far the most acres of habitat managed for longleaf by fire and other 
methods, with nearly 600,000 ac (242,811 ha) treated between October 
2018 and September 2019. Much of the management implemented by partners 
under the ALRI umbrella is likely to benefit gopher tortoise.
    Next, we summarized management practices as detailed in the gopher 
tortoise CCA 2021 annual report, which covers management actions 
implemented between October 2020 and September 2021. CCA management 
data have the advantages of being specific to sites known to support 
gopher tortoises and include both prescribed fire and other beneficial 
practices such as chemical and mechanical treatments and invasive 
species control. Unfortunately, the CCA data are limited to the eastern 
portion of the range, and thus do not include information for the 
western portion. Finally, we summarized the responses to the landowner 
questionnaire regarding acres of prescribed fire, burn frequency, and 
other management practices to benefit the gopher tortoise. Most 
prescribed burns occurred in Units 3 (West Georgia) and 5 (Florida); 
burn frequency is often on a 1- to 3-year cycle; and many landowners 
implement additional beneficial practices (Service 2022, pp. 129-130, 
133-139).
    We describe the results of our analysis of the abundance 
(resiliency), habitat, and management metrics for each analysis unit, 
below. Populations described are those delineated using spatially 
explicit data and may underestimate the number of gopher tortoises and 
populations on the landscape.

[[Page 61854]]

Analysis Unit 1 (Western)
    Based on available data, Unit 1 is composed of many small, 
disconnected populations and very few larger populations (106 local 
populations; 13 landscape populations), spread across private and 
public land. Abundance estimates indicate there are 94 low-, 10 
moderate-, and 2 high-resiliency local populations within this unit. 
Camp Shelby, a DoD property, is the stronghold of Unit 1 with a 
population estimate of 1,003 individual gopher tortoises. Based on 
responses to the landowner survey, 17 properties on private lands in 
the unit support gopher tortoise populations, with 7 properties 
reporting signs of reproduction.
    More than 103,000 ac (41,682 ha) of habitat occurs within gopher 
tortoise populations in Unit 1, with an additional 2 million ac 
(809,371 ha) of potential gopher tortoise habitat where gopher tortoise 
occurrence is unknown. The current estimates for prescribed fire 
implementation show that over 35,795 ac (14,485 ha) were burned within 
this unit in 2019, double the area burned since 1994. Over 90 percent 
of landowners who responded to the questionnaire report implementing 
prescribed fire on a 1- to 3-year rotation, with all respondents 
reporting implementation of additional beneficial practices for gopher 
tortoises.
Analysis Unit 2 (Central)
    Based on available data, Unit 2 has 106 local populations and 30 
landscape populations. Based on current abundance estimates, this unit 
is composed of 71 low-, 27 moderate-, and 8 high-resiliency local 
populations. The eight highly resilient populations are found on 
conservation lands including Fort Rucker, Conecuh NF, Apalachee 
Wildlife Management Area (WMA), Perdido WMA, Geneva State Forest, and 
an unnamed private property. Based on responses to the landowner 
survey, 32 properties on private lands in the unit support gopher 
tortoise populations with 17 properties reporting signs of 
reproduction.
    More than 68,000 ac (27,518 ha) of habitat occurs within gopher 
tortoise populations in Unit 2, with an additional 3.4 million ac (1.37 
million ha) of potential gopher tortoise habitat where gopher tortoise 
occurrence is unknown. The current estimates for prescribed fire 
implementation show that approximately 106,000 ac (42,896 ha) were 
burned in 2019, triple the area burned since 1994. Sixty percent of 
landowners who responded to the questionnaire report implementing 
prescribed fire on a 1- to 3-year rotation, with 72 percent of 
respondents reporting implementation of additional beneficial practices 
for gopher tortoises.
Analysis Unit 3 (West Georgia)
    Based on available data, Unit 3 has 109 local populations and 55 
landscape populations. Based on current abundance estimates, Unit 3 is 
composed of 42 low-, 24 moderate-, and 43 high-resiliency local 
populations. Of the 43 highly resilient populations, 7 populations have 
estimates exceeding 1,000 individuals, including Twin Rivers State 
Forest, Chattahoochee Fall Line WMA, River Bend WMA, Alapaha River WMA, 
Apalachicola NF, and the Jones Center at Ichauway. Based on responses 
to the landowner survey, 48 properties on private land in Unit 3 
support gopher tortoise populations with 21 properties reporting signs 
of reproduction.
    More than 220,000 ac (89,030 ha) of habitat occurs within gopher 
tortoise populations in Unit 3, with an additional 2.9 million ac (1.17 
million ha) of potential gopher tortoise habitat where gopher tortoise 
occurrence is unknown. The current estimates for prescribed fire 
implementation show that more than 194,000 ac (78,509 ha) were burned 
in 2019, almost a 10-fold increase since 1994. Sixty-seven percent of 
landowners who responded to the questionnaire report implementing 
prescribed fire on a 1- to 3-year rotation, with 44 percent of 
respondents reporting implementing additional beneficial practices for 
gopher tortoises.
Analysis Unit 4 (East Georgia)
    Based on available data, Unit 4 has 124 local populations and 46 
landscape populations. Based on current abundance estimates, Unit 4 is 
composed of 35 low-, 58 moderate-, and 31 high-resiliency local 
populations. Of the 31 highly resilient populations, 5 populations have 
estimates exceeding 1,000 individuals, including Ohoopee Dunes WMA, 
Ralph E. Simmons State Forest, Jennings State Forest, and Fort Stewart. 
Based on responses to the landowner survey, 22 properties on private 
land in the unit support gopher tortoise populations with 11 properties 
reporting signs of reproduction.
    More than 149,000 ac (60,298 ha) of habitat occurs within the 
gopher tortoise population in Unit 4, with an additional 2.7 million ac 
(1.09 million ha) of potential gopher tortoise habitat where gopher 
tortoise occurrence is unknown. The current estimates for prescribed 
fire implementation show that more than 161,000 ac (65,154 ha) were 
burned in 2019, over a 7 times increase since 1994. Fifty-three percent 
of landowners who responded to the questionnaire report implementing 
prescribed fire on a 1- to 3-year rotation, with 77 percent of 
respondents reporting implementing additional beneficial practices for 
gopher tortoises.
Analysis Unit 5 (Florida)
    Based on available data, Unit 5 has 211 spatially explicit local 
populations and 109 landscape populations. Based on current abundance 
estimates, Unit 5 is composed of 118 low-, 50 moderate-, and 43 high-
resiliency local populations. Of the 43 highly resilient populations, 
12 populations have estimates exceeding 1,000 individuals, including 
Camp Blanding and Goldhead Branch State Park, Ocala NF, Chassahowitzka 
WMA, Ichetucknee Springs State Park, Bell Ridge Wildlife and 
Environmental Area, Etoniah Creek State Forest, Halpata Tastanaki and 
Cross Florida Greenway, Lake Louisa State Park, Kissimmee Prairie 
Preserve State Park, Green Swamp West Unit WMA, Withlacoochee State 
Forest's Citrus Tract, and Perry Oldenburg Wildlife and Environmental 
Area and Withlachoochee State Forest's Croom Tract. Based on responses 
to the landowner survey, 48 properties on private land in the unit 
support gopher tortoise populations with 35 properties reporting signs 
of reproduction.
    More than 300,000 ac (121,405 ha) of habitat occurs within gopher 
tortoise populations in Unit 5, with an additional 5.3 million ac (2.14 
million ha) of potential gopher tortoise habitat where gopher tortoise 
occurrence is unknown. The current estimates for prescribed fire 
implementation show that more than 582,368 ac (235,675 ha) were burned 
in 2019, a nearly 14 times increase over time since 1994. Twenty-three 
percent of landowners who responded to the questionnaire report 
implementing prescribed fire on a 1- to 3-year rotation, with 83 
percent of respondents reporting implementing additional beneficial 
practices for gopher tortoises.
Representation and Redundancy
    We evaluated current representation by examining the genetic and 
environmental diversity within and among populations across the 
species' range (Gaillard et al. 2017, entire). We report redundancy for 
gopher tortoise as the number and resiliency of gopher tortoise 
populations and their distribution within and among analysis units. 
Current representation and redundancy have likely decreased relative to 
the historical condition of the species due to loss of open pine 
conditions and substantial reduction in

[[Page 61855]]

longleaf pine ecosystems in the species' range.
    The five delineated analysis units are based primarily on genetic 
variation in gopher tortoises across the range of the species. We 
expect this genetic variation to be generally indicative of the 
inherent adaptive capacity of the gopher tortoise as a species (Thurman 
et al. 2020, p. 522). In addition, the variety of environmental 
conditions across the species' range, particularly soil characteristics 
and associated life history characteristics differences between the 
western and eastern portions of the range, may be used as an indication 
of adaptive capacity for the gopher tortoise, allowing the species to 
withstand changing conditions (Thurman et al. 2020, p. 522). Gopher 
tortoise populations are distributed within and among analysis units 
across the species' range, contributing to potential adaptive capacity 
and current representation.
    Currently, multiple local and landscape populations occur in all 
five analysis units. Although the resiliency of these populations 
varies across the range, all analysis units contain populations in high 
and moderate resiliency. Rangewide, 45 percent of spatially explicit 
local populations exhibit moderate or high resiliency. These 
populations are distributed across the range of the species, 
contributing to future adaptive capacity (representation) and buffering 
against the potential of future catastrophic events (redundancy). 
Because the species is widely distributed across its range, it is 
highly unlikely any single event would put the species as a whole at 
risk, although the westernmost portions of the range are likely more 
vulnerable to such catastrophes given that a greater percentage of the 
populations present in this unit are of low resiliency compared to 
other analysis units.

Future Condition

Future Condition Modeling
    To assess future viability for the gopher tortoise, we developed an 
analytical framework that integrates projections from multiple models 
of future anthropogenic and climatic change to project future 
trajectories or trends of gopher tortoise populations and identify 
stressors with the greatest influence on future populations. The 
modeling framework estimates the change in population growth and number 
of populations while accounting for geographic variation in life 
history. The model links intrinsic factors (demographic vital rates) to 
four extrinsic anthropogenic factors that are expected to impact gopher 
tortoise population viability (climate warming, sea level rise, 
urbanization, and shifts in habitat management). We used published 
models describing extrinsic factors in the future to project gopher 
tortoise demographics under six future scenarios varying in threat 
magnitude and presence at three timesteps--40, 60, and 80 years in the 
future. A regression analysis of model outputs was used to identify 
threats that are predicted to have the greatest impact on gopher 
tortoise populations. We summarize the model framework below; 
additional information is available in the SSA report (Service 2022, 
pp. 144-159, appendix B; Folt et al. 2022, entire).
    We developed a population viability analysis (PVA) framework to 
predict population growth and extinction risk for the gopher tortoise. 
For the PVA, the demography of spatially explicit local gopher tortoise 
populations was brought into a multi-stage, female-only model with two 
discrete life stages: juveniles and adults. Recruitment into the adult 
stage by immigration was also modeled. Specific demographic parameters 
including recruitment, maturity age, survival, immigration, and initial 
population size were modeled based on values in gopher tortoise 
literature (Landers et al. 1980, p. 359; Mushinsky et al. 1994, p. 123; 
Rostal and Jones 2002, p. 7; Ott-Eubanks et al. 2003, p. 319; Ashton et 
al. 2007, p. 360; Guyer et al. 2012, p. 130; Perez-Heydrich et al. 
2012, p. 342; Smith et al. 2013, p. 355; Tuberville et al. 2014, p. 
1155; Meshaka Jr. et al. 2019, pp. 105-106; Howell et al. 2020, entire; 
Folt et al. 2021, pp. 624-625, 627; Hunter and Rostal 2021, p. 661; E. 
Hunter unpubl. data, 2021; J. Goessling 2021, p. 141). For the 
demographic parameters (e.g., recruitment, maturity age, survival) that 
vary substantially by temperature among populations, we determined the 
relationships between demographic rates and mean annual temperature 
(MAT) sourced from the WorldClim database (Hijmans 2020, entire).
    We initialized the model with estimates of population size from 
spatially delineated populations (as described in Current Condition). 
In the future condition analysis in the SSA, we did not model local 
populations with fewer than three adult individuals as part of the 
future condition analysis as these populations do not have sufficient 
viability to remain on the landscape during the timeframes modeled (40, 
60, and 80 years) (i.e., these populations have reached the quasi-
extinction threshold). The process of delineating spatially explicit 
local populations and landscape populations for the future condition 
model resulted in a dataset of 626 local populations that formed 244 
landscape populations with 70,600 individual (female) gopher tortoises 
that are included in our analysis of future conditions (Service 2022, 
p. 149).
    A recently published peer-reviewed model uses a very similar 
methodology to the future condition analysis in the SSA (Folt et al. 
2022, entire). The published model varied slightly from that in the SSA 
and did not model populations across the range with current abundance 
of fewer than eight individuals or fewer than three adult females. 
Populations with seven or fewer tortoises likely lack sufficient 
genetic diversity to support sufficient long-term viability (Chesser et 
al. 1980, entire; Frankham et al. 2011, p. 466; Folt et al. 2022, p. 
e02143). Both the recently published and the future condition analysis 
runs of the model assumed a 1:1 sex ratio and a 3:1 adult:juvenile 
ratio in populations and used the ratios to isolate and separate the 
female population into juvenile and adult components (Service 2022, p. 
149; Folt et al. 2021, p. 626; Folt 2022, p. e02143). The published 
iteration of the model resulted in the delineation of 457 local 
populations that formed 202 landscape populations (metapopulations) and 
approximated 70,500 female tortoises (Folt et al. 2022, p. e02143). The 
slight variation in the published model did not substantively change 
the considerations in our analyses of the gopher tortoise's future 
condition.
Influences on Gopher Tortoise Future Viability
    In coordination with scientists with expert knowledge in both 
gopher tortoise population biology and habitat management, we 
identified factors expected to influence gopher tortoise demographics 
in the future as described in Summary of Biological Status and Threats. 
We determined the key drivers of the gopher tortoise's future condition 
that we could incorporate into the model are climate warming, habitat 
management, urbanization, and sea level rise.
    Climate change is predicted to drive warming temperatures and 
seasonal shifts in precipitation across the Southeast (Carter et al. 
2018, entire). Of these two effects, warming temperatures may have the 
greater impact on gopher tortoises, because gopher tortoise demography 
is known to be sensitive to temperature gradients across the species' 
range. Specifically, maturity age and fecundity vary along a north-
south latitudinal gradient, where warmer, southern populations have 
faster growth rates, younger maturity ages, and

[[Page 61856]]

increased fecundity relative to cooler, northern populations (Ashton et 
al. 2007, p. 123; Meshaka Jr. et al. 2019, pp. 105-106). We modeled how 
climate warming may influence gopher tortoise demography by using the 
estimated linear relationships of mean annual temperature with maturity 
age and fecundity to predict how warming temperatures experienced by 
populations in the future will drive concurrent changes in demography.
    Although the gopher tortoise exhibits temperature-dependent sex 
determination, we did not include this effect in the model as gopher 
tortoises can modify nest site selection and timing of nesting, as 
discussed in chapter 3 of the SSA (Service 2022, p. 58). We also did 
not model any potential range expansion or contraction that could occur 
due to long-term climate change, because we are aware of no consensus 
or projection framework related to vegetative community changes and 
climate change projections; also, we expect any significant expansion 
or contraction of the gopher tortoise range is likely to occur late in 
or beyond our projection timeframe of 80 years.
    Climate change models predict favorable burn window conditions to 
shift over future decades, with favorable conditions for prescribed 
fire increasing in the winter but decreasing in the spring and summer 
(Kupfer et al. 2020, pp. 769-770). Overall, projections show that 
seasonal shifts in favorable burn window conditions will decrease 
overall opportunity for management with prescribed fire. We estimated 
how habitat management influences gopher tortoise populations by 
modeling use of fire as a management tool and linking the frequency of 
management to adult survival (Kupfer et al. 2020, entire; Service 2022, 
appendix B; Folt et al. 2022, pp. 4, 8-11). We modeled four changes in 
the burn window based on climate shifts projected by Representative 
Concentration Pathway (RCP) 4.5 and RCP 8.5: (1) decreased fire, (2) 
very decreased fire, (3) increased fire, and (4) status quo.
    Urbanization and development are expected to affect gopher tortoise 
populations in the future, even those on conservation lands, through 
reduced connectivity and effects to gene flow and population migration 
dynamics. Urbanization may also reduce the use of prescribed fire in an 
area and contribute to road mortality and the introduction of nonnative 
invasive species. We modeled effects of urbanization pressure on gopher 
tortoise populations by linking urbanization projections from the 
SLEUTH urbanization model to habitat management of local populations 
with prescribed fire and with baseline immigration rates of gopher 
tortoises across landscape populations (Terando et al. 2014, entire). 
We modeled three potential thresholds in urbanization: (1) Low 
urbanization where cells have a 95 percent or greater probability of 
being developed; (2) moderate urbanization where cells have a 50 
percent or greater probability of being developed; and (3) high 
urbanization where cells have a 20 percent or greater probability of 
being developed. Modeled cells with a high probability of urbanization 
are likely to be urbanized under any scenario (higher certainty), while 
areas with a lower probability of urbanization are likely to be 
urbanized in scenarios with increased impacts or greater effects. 
Inclusion of areas with a lower chance of development leads to an 
overall greater area expected to be developed.
    Sea level rise is expected to negatively affect gopher tortoise 
populations in low-lying coastal areas, such as coastal sand dune 
environments (Blonder et al. 2021, pp. 6-8). We modeled effects of sea 
level rise on gopher tortoises using three projections of sea level 
rise: The ``intermediate-high,'' ``high,'' and ``extreme'' projections 
correspond to projections from global emission scenarios RCP 6 and RCP 
8.5 (IPCC 2022, entire; NOAA 2020, entire). We projected the effects of 
sea level rise on the gopher tortoise in the future by modeling the 
height above sea level of local populations and through reduced 
connectivity between local populations.
Future Scenarios
    We developed six plausible scenarios of future climate warming, 
urbanization, habitat management, and sea level rise to simulate 
population growth and extinction risk for gopher tortoises for 40, 60, 
and 80 years into the future (table 3). Specifically, we created three 
scenarios with different levels of stressors (low stressors, medium 
stressors, and high stressors) that experienced habitat management 
consistent with contemporary target management goals. We then held the 
medium stressor values constant and developed three scenarios that 
varied in habitat management treatments, ranging from scenarios for the 
most habitat management to the least habitat management (table 3).
    Little information is available describing gopher tortoise 
immigration rates in wild populations. Given the uncertainty around 
this parameter, we included four additional scenarios with the medium 
stressor values and status quo habitat management to understand the 
effects of varying rates of immigration on the gopher tortoise future 
condition.

  Table 3--Threats, Habitat Management, and Immigration Values in the Nine Plausible Scenarios Used To Project
                           Future Population Growth and Abundance of Gopher Tortoises
----------------------------------------------------------------------------------------------------------------
                                                   Stressors
                              --------------------------------------------------                    Immigration
          Scenarios                Climate                                           Habitat         into the
                                   warming     Sea level rise   Probability of      management      population
                                  ([deg]C)           (m)         urbanization                        (percent)
----------------------------------------------------------------------------------------------------------------
Low stressors................             1.0            0.54  95 percent or     Status quo.....               1
                                                                greater.
Medium stressors.............             1.5            1.83  50 percent or     Status quo.....               1
                                                                greater.
High stressors...............             2.0            3.16  20 percent or     Status quo.....               1
                                                                greater.
Decreased management.........             1.5            1.83  50 percent or     Less fire......               1
                                                                greater.
Very decreased management....             1.5            1.83  50 percent or     Much less fire.               1
                                                                greater.
Improved management..........             1.5            1.83  50 percent or     More fire......               1
                                                                greater.
No immigration...............             1.5            1.83  50 percent or     Status quo.....               0
                                                                greater.
Intermediate immigration.....             1.5            1.83  50 percent or     Status quo.....               1
                                                                greater.
High immigration.............             1.5            1.83  50 percent or     Status quo.....               2
                                                                greater.
Very high immigration........             1.5            1.83  50 percent or     Status quo.....               4
                                                                greater.
----------------------------------------------------------------------------------------------------------------


[[Page 61857]]

    [The first three scenarios vary the levels of stressors (climate 
warming, sea level rise, and urbanization), while holding habitat 
management and immigration constant.
    The second three scenarios vary the levels of habitat management 
(through prescribed fire), while holding stressors and immigration 
constant.
    The last four scenarios vary only in the level of immigration into 
the population and hold stressors and habitat management constant.]
    To assess future resiliency, redundancy, and representation of the 
gopher tortoise, we used population projections to estimate changes in 
gopher tortoise populations in the future under each of the nine 
scenarios. We assessed the resiliency of future populations to changing 
environments by estimating persistence probability. Persistence 
probability is defined in this assessment as a measure of the risk of 
extinction and is expressed as the percent of current populations 
projected to occur on the landscape in a given future scenario. 
Although the SSA report uses the categories of ``extremely likely to 
persist,'' ``very likely to persist,'' ``more likely than not to 
persist,'' and ``unlikely to persist'' to characterize the future 
condition of gopher tortoise populations, these terms represent a 
portion of our analysis and are not fully representative of the status 
on the species. We will use the phrase ``remain on the landscape'' or 
``not extirpated'' in this finding to indicate the modeled future 
condition categories of gopher tortoise populations of ``extremely 
likely to persist,'' ``very likely to persist,'' and ``more likely than 
not to persist,'' and will indicate the timeframe to which that 
projection applies.
    We assessed redundancy by evaluating projected changes in the total 
number of individuals (abundance or resiliency), number of local 
populations, number of landscape populations, and their distribution 
across the landscape in the future. We summarized population trends by 
estimating population growth rate as increasing (greater than 1), 
stable (1), or decreasing (less than 1). We evaluated how 
representation is predicted to change in the future by examining how 
population growth of total population size (number of individual female 
gopher tortoises), number of local populations, and number of landscape 
populations will vary by the five population genetic groups of 
tortoises across the species' range.
    We report the rangewide model projections for each scenario at the 
three future time steps, summarize the results across all populations 
across the species' range, and describe differences among analysis 
units in Summary of Future Analysis, below. Details regarding future 
projections may also be found in the SSA report and the peer-reviewed 
model resulting from the SSA analyses (Service 2022, pp. 159-175; Folt 
et al. 2022, entire).

Summary of Future Analysis

    While declines in abundance and number of populations are 
predicted, overall projections suggest that extinction risk for the 
gopher tortoise is relatively low in the future. Population projections 
under six future scenarios (threats and management scenarios) predicted 
declines in the number of gopher tortoise individuals, local 
populations, and landscape populations at the 40-, 60-, and 80-year 
timesteps. Relative to current levels of total population size, 
projections for total population size suggested declines by 2060 (33-35 
percent declines), 2080 (30-34 percent declines), and 2100 (28-33 
percent declines). The declines reflect the projected loss of small 
gopher tortoise populations in the earlier timestep (40 years), while 
remaining larger populations remain on the landscape longer. The six 
scenarios varied little in the impact on the total number of 
individuals, local populations, and landscape populations within each 
timestep, but impacts increased in each successive timestep. In 
addition, the 95 percent confidence interval overlapped with 1.0 in all 
cases, indicating no difference in the scenarios.
    Among the future scenario projections, the number of local 
populations and landscape populations were predicted to decline in each 
projection interval (40-, 60-, and 80-year timesteps). Declines in 
local populations and landscape populations were 47-48 percent and 25-
27 percent declines among scenarios, respectively, at the 40-year 
timestep; 60-61 percent and 41-43 percent declines, respectively, at 
the 60-year timestep; and 68-70 percent and 53-57 percent declines, 
respectively, at the 80-year timestep. With these declines, mean 
projections among scenarios at the 80-year timestep indicate 47,202-
50,846 adult female gopher tortoises remain on the landscape in 188-198 
spatially explicit local populations across the range of the species.
    The number of individuals, local populations, and landscape 
populations varied by analysis unit. Abundance in Units 1, 3, and 5 was 
projected to decline overall (27-40 percent, 51-53 percent, and 42-48 
percent declines, respectively). Unit 4 was projected to experience a 
more modest decline (2-14 percent decrease in abundance), and Unit 2 
was projected to increase in abundance. However, declines in the number 
of local populations are projected for all units. The predicted 
declines in number of local populations are greatest in Units 1, 2, and 
5. More populations in Units 1 and 2 currently exhibit low resiliency, 
while Unit 5 contains the highest abundance and number of local 
populations across the range.
    Threats and habitat management scenarios did not strongly affect 
projections of gopher tortoise total population size (number of females 
in the total population), or the number of local and landscape 
populations. No single threat scenario (low, medium, or high stressors) 
or management scenario (more, less, or much less management) was 
sufficient to prevent population declines. However, model projections 
did change substantially based on the immigration rate in the scenario 
(very high, high, intermediate, or no immigration). For example, the 
total population size and the number of local and landscape populations 
projected to remain on the landscape in 2080 under the ``medium 
stressors'' scenario were reduced substantially when simulated with an 
immigration rate of 0. Conversely, higher values for immigration (2 and 
4 percent) produced projections with substantially increased total 
population size above initial starting population size and decreased 
declines in local and landscape populations. In addition to 
immigration, the initial total population size, areal extent of the 
population (ha (ac)), and predicted implementation of habitat 
management through prescribed fire positively affected the chance the 
population would remain on the landscape in the future. The declines in 
number of local populations occurred, in part, because many local 
populations (27.8 percent) had very few individuals to start with in 
the current conditions. Assuming a 3:1 adult to juvenile ratio and an 
even sex ratio, local populations with fewer than 8 individuals were 
functionally extirpated at the start of projections, given our quasi-
extinction probability (3 or fewer adult females).
    Our analysis simulated the fate of known populations largely on 
protected conservation lands that we expect will be managed for 
conservation in the future. Future condition projections based only on 
data from spatially delineated populations (i.e., do not contain 
county-level data or gopher tortoises that are present, but not 
reported) likely substantially

[[Page 61858]]

underestimate the true number of gopher tortoises present across the 
species' range. We expect populations on managed conservation lands to 
be characterized by greater demographic rates and lower extinction risk 
relative to populations that we were unable to model in our framework 
(populations with no spatially explicit data). To this end, we did not 
project the abundance of existing populations not included in our 
dataset or estimate the formation of new populations outside of 
conservation lands. While other tortoise populations exist outside of 
the ones we simulated with our projection model and new tortoise 
populations may form due to natural dispersal and colonization 
dynamics, they may occur on lands lacking long-term protection from 
development, and we did not project those populations into the future 
under assumptions of land management and protection for wildlife 
conservation. Similarly, we could not estimate the formation of new 
populations outside of the sites we projected, or the migration of 
entire populations to new areas, because we have no guarantee of land 
available for the formation or migration of populations.
    While the numbers of individuals, populations, and landscape 
populations were all expected to decline across each projection 
interval, overall projections suggest that extinction risk for the 
gopher tortoise is relatively low in the future. Of the individuals, 
local populations, and landscape populations modeled (a small subset of 
populations likely to occur across the landscape), mean projections 
among scenarios for 80 years in the future suggested the presence of 
47,202-50,846 individuals (females only) among 188-198 local 
populations within 106-114 landscape populations across most of the 
range of the species. The presence of relatively large numbers of 
individuals and populations suggests resiliency of the species in the 
face of change, and redundancy to buffer from future catastrophic 
events. The spatial distribution of populations predicted to occur on 
the landscape in the future are distributed evenly among genetic 
analysis units, which suggests adaptive capacity or representation in 
the future as well.
    Although we do not project any of the analysis units to be 
extirpated in any scenario, we do anticipate declines in species' 
representation and redundancy through the projected loss of total 
number of individuals and number of local and landscape populations. 
Gopher tortoise populations are projected to remain on the landscape in 
all scenarios and included timesteps in each analysis unit, providing 
genetic variability across the range and adaptive capacity for the 
species. We expect that future gopher tortoise redundancy will be 
somewhat reduced from current redundancy due to the loss of some local 
and landscape populations. For example, in Unit 1, approximately 16 
percent of current populations are expected to remain on the landscape 
at the 80-year timestep, under the medium stressor and less management 
scenario. Populations in this unit are more isolated, small, and 
fragmented compared to the remainder of the range.

Determination of Gopher Tortoise's Status

    Section 4 of the Act (16 U.S.C. 1533) and its implementing 
regulations (50 CFR part 424) set forth the procedures for determining 
whether a species meets the definition of ``endangered species'' or 
``threatened species.'' The Act defines an ``endangered species'' as a 
species that is in danger of extinction throughout all or a significant 
portion of its range, and a ``threatened species'' as a species that is 
likely to become an endangered species within the foreseeable future 
throughout all or a significant portion of its range. The Act requires 
that we determine whether a species meets the definition of 
``endangered species'' or ``threatened species'' because of any of the 
following 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.

Status Throughout All of Its Range

    After evaluating threats to the species and assessing the 
cumulative effect of the threats under the section 4(a)(1) factors, we 
determined that the species currently has sufficient resiliency, 
redundancy, and representation contributing to its overall viability 
across its range. The primary stressors affecting the gopher tortoise's 
biological status include habitat loss, degradation, and fragmentation 
due to land use changes from urbanization (Factor A), climate change 
(Factor E), and insufficient and/or incompatible habitat management 
(Factor E). Upper respiratory tract disease and other viral, bacterial, 
fungal, and parasitic infections (Factor C) affect individual gopher 
tortoises and can have localized effects, but these threats do not 
appear to have species-level impacts. Predation of eggs, hatchlings, 
and juvenile tortoises (Factor C) impacts some gopher tortoise 
populations. Overutilization for commercial or recreational purposes 
(harvest and rattlesnake roundups) (Factor B) of gopher tortoises was a 
historical threat and may affect individuals, but is not currently an 
impact to the species rangewide. The effects of nonnative invasive 
species (Factor E) on gopher tortoise habitat also negatively influence 
gopher tortoise viability. Conservation efforts and regulatory 
mechanisms are in place across the range of the species and are 
addressing some of the identified threats by restoring, enhancing, or 
providing gopher tortoise habitat, relocating tortoises, and augmenting 
populations through captive propagation.
    Urbanization results in a range of impacts that either remove or 
degrade/fragment remaining habitat, or can impact gopher tortoises 
directly through development. Urbanization brings road construction and 
expansion, which may cause direct mortality of gopher tortoises. In 
addition, this stressor creates conditions beneficial to nonnative 
invasive species and predators as well as conditions that limit fire 
management of gopher tortoise habitat. Temperature increases associated 
with long-term climate change are likely to further constrain use of 
prescribed fire through a decrease in the number of suitable burn days. 
Additionally, habitat loss resulting from sea level rise associated 
with climate change is a risk for coastal populations of gopher 
tortoise.
    A variety of conservation efforts to benefit the gopher tortoise 
and its habitat have been implemented by Federal and State agencies, 
nongovernmental organizations, private landowners, and partnerships 
across the range of the species. These conservation measures and 
existing regulatory mechanisms also influence gopher tortoise viability 
through the conservation and restoration of gopher tortoise habitat and 
prevention of habitat loss, particularly efforts implemented since our 
July 27, 2011, 12-month finding on the petition to list the eastern 
portion of the gopher tortoise range as threatened.
    While threats have acted on the species to reduce available habitat 
and species abundance, the gopher tortoise occurs in the six States 
comprising the historical and current range of the species. In 
addition, based on best available information, we estimate that more 
than 149,000 gopher tortoises occur in 656 spatially delineated local 
populations across the range of the species. Approximately 38 percent 
of

[[Page 61859]]

local populations exhibit high or moderate current resiliency, and the 
species is widely distributed across much of its range. In addition, 
the 360 gopher tortoise populations in low resiliency are widely 
distributed across the species' range. These low-resiliency populations 
often occur near other local populations (within a landscape 
population) and contribute to the resiliency of the landscape 
populations and the species' redundancy and representation. Despite the 
historical and current loss of habitat with the open pine conditions 
required by the gopher tortoise, sufficient quality and quantity of 
habitat remains to provide adequate resiliency to contribute to the 
viability of the species. Although the species-level redundancy has 
likely decreased from historical levels due to loss of habitat and the 
effects to the 3Rs, the gopher tortoise retains a sufficient number of 
populations with high or moderate resiliency that are distributed 
across the range to respond to catastrophic events. The five genetic 
groups delineated across the species' range provide adaptive capacity 
and sufficient species-level representation for the gopher tortoise. 
Thus, after assessing the best available information, we conclude that 
the gopher tortoise currently exhibits levels of resiliency, 
redundancy, and representation such that the species is not in danger 
of extinction throughout all of its range.
    Therefore, we proceed with determining whether the gopher tortoise 
is likely to become an endangered species within the foreseeable future 
throughout all of its range. We evaluated the future condition of the 
species based on projections under nine plausible scenarios. We 
evaluated the viability of the species under these scenarios over the 
foreseeable future and considered the condition of the species in 
relation to its resiliency, redundancy, and representation. We analyzed 
future conditions based on input from species experts, generation time 
for the species, and the confidence in predicting patterns of climate 
warming, sea level rise, urbanization, and habitat management, enabling 
us to reliably predict threats and the species' response over time. 
Using the best available information, we evaluated future conditions at 
40, 60, and 80 years in the future. These timesteps allowed us to 
project relevant threats to the species in view of its life-history 
characteristics, including lifespan and reproduction and recruitment. 
Within this timeframe, these projections are sufficiently reliable to 
provide a reasonable degree of confidence in the predictions. Details 
regarding the future condition analyses are available in the SSA report 
and associated future condition model (Folt et al. 2022; Service 2022, 
appendix B).
    In modeling the future condition of the species, we projected the 
number of individuals, local populations, and landscape populations, 
population growth, and the probability that populations will remain on 
the landscape (percent of current local populations extant on the 
landscape) under each scenario at timesteps 40, 60, and 80 years into 
the future as described in Future Condition, above. The projection 
outcomes did not differ significantly by different threat scenarios; 
however, immigration and management actions did affect model results. 
The threats included in future condition modeling are projected to 
result in a decline in the number of individuals, populations, and 
landscape populations across each projection interval. Of the 
individuals, local populations, and landscape populations modeled (a 
subset of populations likely to occur across the landscape), mean 
projections among scenarios for 80 years in the future suggested the 
presence of 47,202-50,846 individuals (adult females) among 188-198 
local populations within 106-114 landscape populations. We recognize 
this is likely an underestimation of the gopher tortoise's future 
condition since only existing populations on protected lands were 
modeled. In addition, any new populations in the future (formed or 
translocated) were not included in this future projection modeling. 
Many of the populations predicted not to remain on the landscape were 
currently small populations. Although the model projects declines in 
the future that include the loss of these smaller populations, the 
overall projections suggest that extinction risk for the gopher 
tortoise is low in the future.
    Although the threats to the species of habitat loss and 
fragmentation due to urbanization, climate change, sea level rise, and 
habitat management are expected to persist in the foreseeable future 
and the effects of these threats on this long-lived species will 
continue at some level, some threats have been reduced and will 
continue to be reduced through implemented and ongoing conservation 
actions and regulatory mechanisms, as discussed above under 
Conservation Efforts and Regulatory Mechanisms. Rangewide, the future 
condition of the species with relatively large numbers of individuals 
and populations suggests resiliency to withstand stochastic 
environmental and demographic change, and redundancy to buffer from 
future catastrophic events. The spatial distribution of populations 
predicted to remain extant in the future is distributed among genetic 
analysis units, which suggests sufficient genetic representation in the 
future as well.
    After evaluating threats to the species and assessing the 
cumulative effect of the threats under the section 4(a)(1) factors, we 
conclude that the risk factors acting on the gopher tortoise and its 
habitat, either singly or in combination, are not of sufficient 
imminence, scope, or magnitude to rise to the level to indicate that 
the species is in danger of extinction now (an endangered species), or 
likely to become endangered within the foreseeable future (a threatened 
species), throughout all of its range.

Status Throughout a Significant Portion of Its Range

    Under the Act and our implementing regulations, a species may 
warrant listing if it is in danger of extinction or likely to become so 
within the foreseeable future throughout all or a significant portion 
of its range. Having determined that the gopher tortoise is not in 
danger of extinction or likely to become so in the foreseeable future 
throughout all of its range, we now consider whether it may be in 
danger of extinction or likely to become so in the foreseeable future 
in a significant portion of its range--that is, whether there is any 
portion of the species' range for which it is true that both (1) the 
portion is significant; and (2) the species is in danger of extinction 
now or likely to become so in the foreseeable future in that portion. 
Depending on the case, it might be more efficient for us to address the 
``significance'' question or the ``status'' question first. We can 
choose to address either question first. Regardless of which question 
we address first, if we reach a negative answer with respect to the 
first question that we address, we do not need to evaluate the other 
question for that portion of the species' range.
    In undertaking this analysis for the gopher tortoise, we chose to 
address the status question first. We began by identifying any portions 
of the range where the biological status of the species may be 
different from its biological status elsewhere in its range. The range 
of a species can theoretically be divided into portions in an infinite 
number of ways, so we focus our analysis on portions of the species' 
range that contribute to the conservation of the species in a 
biologically meaningful way. For this purpose, we considered 
information pertaining to the geographic distribution of (a) 
individuals of the species, (b) the threats that the species faces, and 
(c) the

[[Page 61860]]

resiliency condition of populations. For the gopher tortoise, we 
considered whether the threats or their effects are occurring in any 
portion of the species' range such that the species is in danger of 
extinction now or likely to become so in the foreseeable future in that 
portion of the range.
    We examined the following past, ongoing, and future anticipated 
threats: habitat loss and fragmentation due to urbanization, climate 
warming, sea level rise, habitat management, disease, predation, and 
nonnative invasive species, including cumulative effects. The location 
and magnitude of some threats varies across the species' range and 
accordingly may impact the species differently in different portions. 
For example, sea level rise influences gopher tortoise viability 
primarily in coastal areas.
    Less habitat management to benefit gopher tortoise has been 
implemented in the western portion of the range (Units 1 and 2) 
compared to the remainder of the range; therefore, the effects of lack 
of habitat management influences gopher tortoise populations in the 
westernmost unit to a greater extent. Although threats to the gopher 
tortoise's viability differ spatially and in magnitude, we find that 
the overall level of threats is similar in populations or analysis 
units across the range of the species. These threats are certain to 
occur, and in those analysis units with fewer populations that exhibit 
predominantly low resiliency, these populations are facing the same 
level of threats. In those analysis units with populations that are 
overall less resilient compared to those in other units, we expect that 
a similar level of threats will have a disproportionate impact in these 
areas with lower resiliency populations. These low resiliency 
populations (or analysis units) will be impacted or have a stronger 
negative response to threats than moderate or high resiliency 
populations (or analysis units). We looked across the range of the 
gopher tortoise and identified three portions of the range where the 
biological status may be different than the rangewide status. The three 
areas we found to warrant further evaluation were the two westernmost 
analysis units corresponding to Unit 1 (Western; west of the Mobile and 
Tombigbee Rivers) and Unit 2 (Central; west of the Apalachicola and 
Chattahoochee Rivers and east of Unit 1) and Unit 5 (Florida).
    The impacts of habitat loss and fragmentation, climate change, and 
habitat management combined with other stressors are expected to reduce 
the viability of the populations to withstand stochastic and 
catastrophic events. Although most threats occur at a similar level 
throughout the range of the species, the threats of habitat management 
and sea level rise differ across the range.
    Sea level rise primarily affect populations along the coast in Unit 
5 (Florida). Although sea level rise is projected to affect coastal 
populations of gopher tortoise, the number of populations affected 
varies by location and elevation of the population, site-specific 
characteristics, and climate change scenario. Unit 5 currently has 43 
populations that exhibit high resiliency and 50 populations that 
exhibit moderate resiliency. Even though declines are predicted to be 
more significant in this unit than others, future condition modeling 
projects between 58 and 62 local populations and 37 to 43 landscape 
populations will remain on the landscape in Unit 5, including the very 
large populations (exceeding 1,000 individuals). The current and future 
condition analyses of gopher tortoise indicate sufficient resiliency, 
representation and redundancy in Unit 5. Given the species' current and 
future condition within this unit, we determined that the gopher 
tortoise in Unit 5 does not have a different status than the remainder 
of the range.
    The best available information indicates that less habitat 
management occurs in the western portion of the range (Units 1 and 2) 
compared to the remainder of the range. The populations in the western 
two units (particularly Unit 1) are characterized by ecological and 
physiological characteristics that lead to lower resiliency. 
Populations in Units 1 (Western) and 2 (Central) experience lower 
abundance, smaller clutch size, lower hatch rate, slower growth, and 
less extensive suitable habitat leading to lower resiliency for a 
higher proportion of populations in the two units. In Units 1 (Western) 
and 2 (Central), approximately 11 and 33 percent of populations exhibit 
moderate or high resiliency, respectively, compared to 45 percent 
rangewide. A higher proportion of populations in Units 1 (Western) and 
2 (Central) exhibit low resiliency, with 88 percent of populations in 
Unit 1 (Western) and 67 percent of populations in Unit 2 (Central) in 
low resiliency. Less habitat management beneficial to gopher tortoise 
occurs in Units 1 and 2, and the overall lower resiliency of 
populations in these units is lower. As a result of lower resiliency, 
the species' response is more pronounced, and the rangewide threats and 
lower levels of habitat management are having a greater impact than 
elsewhere in the range. Despite the lower current resiliency of 
populations in Units 1 (Western) and 2 (Central), the gopher tortoise 
is still widespread throughout this extensive geographic area and high 
and moderate resiliency populations also occur throughout the units. In 
addition, given the current population distribution across these units, 
it is not likely that a single catastrophic event would currently place 
the species from this portion of its range at risk of extinction.
    Modeling of future conditions projects declines in abundance and 
fewer extant local and landscape populations in Units 1 (Western) and 2 
(Central) compared to the rest of the range in the foreseeable future. 
For example, Unit 1 (Western) and Unit 2 (Central) are projected to 
have 15 and 14 local populations, respectively, on the landscape in 
2100 under the medium stressors and less habitat management scenario. 
These projected declines would significantly increase the risk of 
extirpation of Units 1 (Western) and 2 (Central) from a catastrophic or 
stochastic event. Although the species currently has sufficient 
resiliency and distribution to withstand a stochastic or catastrophic 
event, projected declines in resiliency or extirpation of populations 
will further reduce the species redundancy and representation in this 
portion of the range. Given the species' future condition within these 
units, we have identified Units 1 (Western) and 2 (Central) of the 
gopher tortoise as an area that has a different status than the 
remainder of the range.
    We then proceeded to the significance question, asking whether this 
portion of the range (i.e., Units 1 (Western) and 2 (Central)) is 
significant. The Service's most recent definition of ``significant'' 
within agency policy guidance has been invalidated by court order (see 
Desert Survivors v. U.S. Department of the Interior, 321 F. Supp. 3d 
1011, 1070-74 (N.D. Cal. 2018)). In undertaking this analysis for the 
gopher tortoise, we considered whether this portion of the species' 
range is significant based on its biological importance to the overall 
viability of the gopher tortoise. Therefore, for the purposes of this 
analysis, when considering whether this portion is significant, we 
considered whether the portion may (1) occur in a unique habitat or 
ecoregion for the species, (2) contain high-quality or high-value 
habitat relative to the remaining portions of the range, for the 
species' continued viability in light of the existing threats, (3) 
contain habitat that is essential to a specific life-history function 
for the species and that is not found in the other portions, or (4)

[[Page 61861]]

contain a large geographic portion of the suitable habitat relative to 
the remaining portions of the range for the species.
    We evaluated the available information about this portion of the 
species to assess its significance. The portion of the range that 
comprises Units 1 (Western) and 2 (Central) contains approximately 20 
percent of the suitable habitat currently occupied by the species, with 
approximately 103,582 ac (41,918 ha) in Unit 1 (Western) and 68,430 ac 
(27,692 ha) in Unit 2 (Central). Although these units contribute to the 
rangewide representation and redundancy of the gopher tortoise, Units 1 
(Western) and 2 (Central) do not constitute a large geographic area 
relative to the remaining portions of the range of the species. This 
portion does not contribute high-quality habitat or constitute high 
value habitat for gopher tortoise. The best available science indicates 
this portion generally contains lower quality or less extensive habitat 
for gopher tortoises than in the remainder of the range. In addition, 
this portion does not constitute an area of habitat that is essential 
to a specific life-history function for the species that is not found 
in the remainder of the range.
    Overall, we found no substantial information that would indicate 
this portion of the gopher tortoise's range is significant in terms of 
the above habitat considerations. As a result, we determined that the 
portion comprising Units 1 (Western) and 2 (Central) does not represent 
a significant portion of the gopher tortoise's range. Therefore, we 
conclude that the species is not in danger of extinction now or likely 
to become so in the foreseeable future in any significant portion of 
its range. This finding does not conflict with the courts' holdings in 
Desert Survivors v. Department of the Interior, 321 F. Supp. 3d 1011, 
1070-74 (N.D. Cal. 2018), and Center for Biological Diversity v. 
Jewell, 248 F. Supp. 3d 946, 959 (D. Ariz. 2017) because, in reaching 
this conclusion, we did not apply the aspects of the Final Policy's 
definition of ``significant'' that those court decisions held to be 
invalid.
    We have carefully assessed the best scientific and commercial 
information available regarding the current and future threats to the 
gopher tortoise. Because the species is neither in danger of extinction 
now nor likely to become so in the foreseeable future throughout all or 
any significant portion of its range, the gopher tortoise does not meet 
the definition of an endangered species or threatened species. 
Therefore, we find that listing the gopher tortoise as an endangered or 
threatened species rangewide under the Act is not warranted at this 
time.

Distinct Population Segment (DPS) Analysis

    Under the Act, we have the authority to consider for listing any 
species, subspecies, or, for vertebrates, any distinct population 
segment (DPS) of these taxa if there is sufficient information to 
indicate that such action may be warranted. The term ``species'' 
includes any subspecies of fish or wildlife or plants and any DPS of 
any species of vertebrate fish or wildlife that interbreeds when mature 
(16 U.S.C. 1532(16)). To guide the implementation of the DPS provisions 
of the Act, we and the National Marine Fisheries Service (National 
Oceanic and Atmospheric Administration--Fisheries), published the 
Policy Regarding the Recognition of Distinct Vertebrate Population 
Segments Under the Endangered Species Act (DPS Policy) in the Federal 
Register on February 7, 1996 (61 FR 4722). Under our DPS Policy, we use 
two elements to assess whether a population segment under consideration 
for listing may be recognized as a DPS: (1) The population segment's 
discreteness from the remainder of the species to which it belongs, and 
(2) the significance of the population segment to the species to which 
it belongs. If we determine that a population segment being considered 
for listing is a DPS, then the population segment's conservation status 
is evaluated based on the five listing factors established by the Act 
to determine if listing it as either endangered or threatened is 
warranted.
    Based on the information available regarding potential discreteness 
and significance for the species, we determined it was appropriate to 
review the status of the gopher tortoise by conducting a DPS analysis 
for the species. The western portion of the gopher tortoise range 
(Western) where the species is currently listed as threatened (52 FR 
25376, July 7, 1987)) consists of those populations of gopher tortoise 
found west of the Mobile and Tombigbee Rivers in Alabama, Louisiana, 
and Mississippi. The eastern portion of the range (Eastern), where the 
species was identified as a candidate in 2011, consists of those gopher 
tortoise populations east of the Mobile and Tombigbee Rivers in 
Alabama, Georgia, Florida, and South Carolina. Below, we evaluate the 
western and eastern portions of the gopher tortoise range as population 
segments to determine whether they meet the definition of a DPS under 
our DPS Policy.
BILLING CODE 4333-15-P

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[GRAPHIC] [TIFF OMITTED] TP12OC22.002

BILLING CODE 4333-15-C

Discreteness

    Under our DPS Policy, a population segment of a vertebrate taxon 
may be considered discrete if it satisfies either of the following 
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 
differences in control of exploitation, management of habitat, 
conservation status, or regulatory mechanisms exist

[[Page 61863]]

that are significant in light of section 4(a)(1)(D) of the Act. In 
determining whether the test for discreteness has been met under the 
DPS policy, we allow, but do not require genetic evidence to be used.

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. This 
consideration may include, but is not limited to: (1) Evidence of the 
persistence of the discrete population segment in an ecological setting 
that is unusual or unique for the taxon, (2) evidence that loss of the 
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 historical range, or 
(4) evidence that the discrete population segment differs markedly from 
other populations of the species in its genetic characteristics. Of 
particular note, as we explained in our draft (76 FR 76987, December 9, 
2011, p. 76998) and final (79 FR 37577, July 1, 2014, pp. 79 FR 37579, 
37585) Policy on Interpretation of the Phrase ``Significant Portion of 
Its Range'' in the Endangered Species Act's Definitions of ``Endangered 
Species'' and ``Threatened Species'' (SPR Policy), the definition of 
``significant'' for the purpose of significant portion of the range 
analysis differs from the definition of ``significant'' found in our 
DPS Policy and used for DPS analysis. Considering the potential results 
of using the same standard for significance under the DPS policy to 
define ``significant'' in the SPR Policy led us to conclude that the 
two provisions cannot use the same definitions for ``significant.'' 
Accordingly, the analysis for ``significant'' under the DPS Policy 
differs from the analysis of ``significant'' under the SPR provision. 
While the definition contained in the SPR Policy has been vacated, our 
consideration of ``significant'' in the ``significant portion of its 
range'' provision for this analysis is also different than the standard 
for significance under the DPS Policy for the same reasons.
    The DPS Policy requires that for a vertebrate population to meet 
the Act's definition of ``species,'' it must be discrete from other 
populations and must be significant to the taxon as a whole. The 
significance criterion under the DPS Policy is necessarily broad and 
could be met under a wider variety of circumstances even if it could 
not be met under the SPR Policy. For example, in this case, we 
determined (see section below) that the western and eastern population 
segments are ``significant'' for the purposes of DPS, and we did not, 
as discussed above, conclude that the western portion constituted a 
``significant'' portion of the gopher tortoise's range.

Discreteness of the Western and Eastern Population Segments of the 
Gopher Tortoise Range

    The western and eastern population segments of the gopher tortoise 
range are markedly separated from each other (other populations) 
geographically (physical) and genetically. The western and eastern 
population segments of the range are separated by the Mobile and 
Tombigbee Rivers. Thus, the western population segment includes all 
gopher tortoises occurring in southwestern Alabama, southern 
Mississippi, and southeastern Louisiana, and the eastern population 
segment includes all gopher tortoises occurring in the remainder of 
Alabama and all of Georgia, South Carolina, and Florida (figure 3). 
These rivers act as a physical impediment to crossing by gopher 
tortoises in either direction and represent a barrier to dispersal and 
gene flow. The rivers are wide and deep year-round, and human 
development (e.g., roads and towns) is adjacent to some areas of the 
rivers. Due to the physical separation of these two population segments 
by the Mobile and Tombigbee Rivers, gopher tortoises in these portions 
do not, and will likely never, naturally interact with individuals or 
populations in the other population segment.
    In terms of genetic separation, there is a phylogenetic break 
(difference in genetics) between the western and eastern population 
segments of the gopher tortoise's range (Ennen et al. 2012, pp. 113-
116). Several studies show genetic assemblages across the geographic 
range, but these studies are not entirely congruent in their 
delineations of western and eastern genetic assemblages (Osentoski and 
Lamb 1995, p. 713; Clostio et al. 2012, pp. 617-620; Ennen et al. 2012, 
pp. 113-120; Gaillard et al., 2017, pp. 501-503). No shared haplotypes 
on a mitochondrial gene were noted in populations found on opposite 
sides of the Mobile and Tombigbee Rivers (Clostio et al. 2012, pp. 619-
620). However, the phylogenetic break does not entirely correspond to a 
particular geographic barrier with some shared haplotypes found in both 
the western portions of the tortoise's range and the panhandle of 
Florida and Georgia populations in a similar study (Ennen et al. 2012, 
pp. 113-116). Recent microsatellite analysis suggests there are five 
main genetic groups in the taxon, delineated by the Tombigbee and 
Mobile Rivers, Apalachicola and Chattahoochee Rivers, and the 
transitional areas between several physiographic province sections of 
the Coastal Plains (i.e., Eastern Gulf, Sea Island, and Floridian) 
(Gaillard et al. 2017, pp. 505-507).
    Based on our review of the best available information, we conclude 
the western and eastern population segments of the gopher tortoise 
range are markedly separated from each other due to geographic 
(physical) and genetic separation. Therefore, we have determined that 
the western and eastern population segments of the gopher tortoise 
range each meet the condition of discreteness under our DPS policy.

Significance of the Western and Eastern Population Segments of the 
Gopher Tortoise Range

    We determine that the western and eastern discrete population 
segments are significant based, in part, upon evidence that loss of 
portions would result in a significant gap in the range of the taxon. 
The loss of either the western or eastern population segment would 
result in a substantial change in the overall range and distribution of 
the gopher tortoise. The loss of the western portion would shift the 
taxon's western range boundary eastward and result in the loss of 
species' presence west of the Mobile and Tombigbee Rivers, which are 
natural barriers to the eastern portion. A loss of the eastern portion 
of the range would result in a significant gap in the range by losing 
98 percent of the current estimated rangewide abundance (in spatially 
explicit populations), 88 percent of the geographic area of the range, 
and the core of the current species' distribution (Service 2022, pp. 
119-120).
    In addition, the western and eastern population segments differ 
markedly from each other in their genetic characteristics (unique 
haplotypes and pronounced nuclear differentiation), as described in 
Discreteness, above. The loss of the western population segment would 
result in a substantial reduction in the presence of these genetic 
characteristics in the species. The eastern population segment is 
genetically valuable to the taxon, because it contains the greatest 
genetic diversity and may contribute more to the overall adaptive 
capacity of the species. Therefore, we have determined that the western 
and eastern population segments differ markedly in the genetic

[[Page 61864]]

characteristics, and loss of this genetic diversity would likely impact 
the species' adaptive capacity.
    Given the evidence that the western and eastern population segments 
would result in a significant gap in the gopher tortoise's range if 
lost, and that these population segments differ markedly from each 
other based on their genetic characteristics, we consider the western 
and eastern population segments to be significant to the species as a 
whole. Thus, the western and eastern population segments of the gopher 
tortoise's range meet the criteria for significance under our DPS 
Policy.

DPS Conclusion for the Western and Eastern Portions

    Our DPS Policy directs us to evaluate the significance of a 
discrete population in the context of its biological and ecological 
significance to the remainder of the species to which it belongs. Under 
our DPS policy, the standard for discreteness does not require absolute 
separation because such separation can rarely be demonstrated for any 
population of organism. Based on an analysis of the best available 
scientific and commercial data, we conclude that the western and 
eastern portions of the gopher tortoise's range are discrete due to 
marked separation geographically, ecologically, and genetically from 
one another. Furthermore, we conclude that the western and eastern 
portions of the range are significant for the reasons described above, 
including that loss of either portion would result in a significant gap 
in the range of the taxon. Therefore, we conclude that the western and 
eastern portions of the gopher tortoise's range are both discrete and 
significant under our DPS policy, and, therefore, these populations are 
listable entities under the Act. We will subsequently refer to them as 
the Western DPS and the Eastern DPS.
    As mentioned above, we have determined the gopher tortoise in the 
western portion of its range, the current listed entity of gopher 
tortoise, meets the criteria of a DPS, but the best available 
information does not support any taxonomic change for the species. This 
document does not propose a revision of the defined entity. We will 
take regulatory action in the future to assign the correct nomenclature 
to the listed entity if we deem this action to be necessary for 
clarity.
    Based on our DPS Policy, if a population segment of a vertebrate 
species is both discrete and significant relative to the taxon as a 
whole (i.e., it is a distinct population segment), its evaluation for 
endangered or threatened status will be based on the Act's definition 
of those terms and a review of the factors enumerated in section 4(a) 
of the Act. Having found that the western and eastern portions of the 
gopher tortoise's range each meet the definition of a distinct 
population segment, we now evaluate the status of each DPS to determine 
whether it meets the definition of an endangered or threatened species 
under the Act.

Status Throughout All of the Western DPS's Range

    In the analysis above for the gopher tortoise as a whole, we have 
carefully assessed the best scientific and commercial information 
available regarding the past, present, and future threats to the 
Western DPS (i.e., Unit 1) of the species. We considered whether the 
Western DPS of the gopher tortoise is presently in danger of extinction 
throughout all of its range. As described above under Status Throughout 
a Significant Portion of its Range, the ongoing and future impacts of 
habitat loss and fragmentation, climate change, and habitat management 
in combination with secondary threats act to reduce the viability of 
the Western DPS. Other secondary, rangewide threats, including disease, 
predation, and nonnative invasive species, also have some effect on the 
Western DPS. However, the magnitude and impacts of these threats are 
exacerbated by population characteristics in this DPS.
    The local gopher tortoise populations in the Western DPS are 
generally smaller than in the Eastern DPS; in particular, the local 
populations have lower abundance, decreased reproduction, and decreased 
recruitment compared to the remainder of the range. However, 106 
spatially explicit local populations at varying levels of resiliency 
occur in the Western DPS and are distributed across the geographic area 
of the DPS. Approximately 87 percent of local populations in the 
Western DPS currently exhibit low resiliency, with 10 percent (12 
populations) in moderate or high resiliency. Populations in the Western 
DPS occur in habitat that is more fragmented than in the Eastern DPS 
with the De Soto National Forest in southern Mississippi as one of the 
few extensive reaches of suitable habitat.
    More than 103,000 ac (41,682 hectares) of habitat with gopher 
tortoise occurrences are currently known in the Western DPS with almost 
2 million ac (809,371 ha) of potential habitat where gopher tortoise 
occupancy is unknown. The best available information indicates that 
less habitat management occurs in the Western DPS compared to the 
Eastern DPS, although fire implementation has more than doubled since 
1994 (Service 2022, p. 130). Gopher tortoises are a long-lived species 
and populations in high (2) or moderate (10) resiliency currently occur 
in the Western DPS with reproduction and recruitment reported from 
populations on public and private lands. We expect individuals will 
remain on the landscape for several decades despite current and ongoing 
threats. Despite the lower current resiliency of populations in the 
Western DPS, the gopher tortoise is still widespread throughout this 
extensive geographic area. In addition, it is not likely that a single 
catastrophic event would result in the extirpation of the species from 
this portion, but loss of populations would reduce gopher tortoise 
representation and redundancy. We have determined that the Western DPS 
is not currently in danger of extinction throughout its range.
    We next analyzed whether the Western DPS is likely to become an 
endangered species within the foreseeable future throughout its range. 
In our consideration of foreseeable future, we evaluated how far into 
the future we could reliably predict the threats to this unit, as well 
as the gopher tortoise's response to those threats. Based on the 
modeling and scenarios evaluated, we considered our ability to make 
reliable predictions in the future and the uncertainty in how and to 
what degree the unit could respond to those risk factors in this 
timeframe. We determined a foreseeable future of 80 years for the 
Western DPS. We analyzed future conditions based on input from species 
experts, generation time for the species, and the confidence in 
predicting patterns of climate warming, sea level rise, urbanization, 
and habitat management, enabling us to reliably predict threats and the 
species' response over time. Details regarding the future condition 
analyses are available in the SSA report and associated future 
condition model (Folt et al. 2022, SSA 2022, appendix B).
    In future condition models, the populations in the Western DPS show 
low or no recruitment and population growth, leading to projected loss 
of populations, particularly small populations, in the foreseeable 
future. As described above, we developed nine plausible future 
scenarios to include varying levels of stressors and habitat management 
to project the future number of individuals, population growth rate, 
and number of local and landscape populations. The Western DPS is 
predicted to decline overall with reduced abundance and reductions in 
local and landscape populations. We included spatially explicit 
populations

[[Page 61865]]

with current population estimates of more than three tortoises in our 
analysis of future conditions. In the Western DPS, 102 spatially 
explicit local populations met this criteria and were modeled in our 
future condition analysis. In the moderate stressors and status quo 
habitat management scenario, 84 percent of modeled populations in the 
Western DPS are unlikely to remain on the landscape in 2100.
    For example, with the exception of one population, the model 
projects the remaining six spatially explicit populations in Louisiana 
were unlikely to remain on the landscape in 80 years in the future. 
Mississippi was projected to lose 77 percent of current local 
populations, but maintain 71 percent of its landscape populations 
(landscape populations will be composed of fewer local populations). 
Further, approximately 80 percent of spatially explicit local 
populations in the Western DPS are projected as unlikely to remain on 
the landscape in 80 years under the status quo threats, less management 
(prescribed fire), and immigration scenario. As mentioned above, less 
habitat management currently occurs in the Western DPS compared to the 
Eastern DPS. Therefore, we expect that status quo threats (medium 
stressors) and less habitat management are reasonable and a plausible 
mechanism to project future species' condition in the Western DPS. The 
low resiliency of these populations significantly increases the impact 
of current and ongoing threats to the populations in the Western DPS. 
In addition to reduced resiliency, the impact of a catastrophic or 
stochastic event would reduce representation and redundancy in the 
Western DPS within the foreseeable future.
    After assessing the best available information, we conclude that 
the Western DPS of gopher tortoise is likely to become endangered 
within the foreseeable future throughout the Western DPS.

Status Throughout a Significant Portion of the Western DPS's Range

    Under the Act and our implementing regulations, a species may 
warrant listing if it is in danger of extinction or likely to become so 
in the foreseeable future throughout all or a significant portion of 
its range. The court in Center for Biological Diversity v. Everson, 435 
F. Supp. 3d 69 (D.D.C. 2020) (Everson), vacated the aspect of the Final 
Policy on Interpretation of the Phrase ``Significant Portion of Its 
Range'' in the Endangered Species Act's Definitions of ``Endangered 
Species'' and ``Threatened Species'' (Final Policy) (79 FR 37578; July 
1, 2014) that provided that the Service does not undertake an analysis 
of significant portions of a species' range if the species warrants 
listing as threatened throughout all of its range. Therefore, we 
proceed to evaluating whether the species is endangered in a 
significant portion of its range--that is, whether there is any portion 
of the species' range for which both (1) the portion is significant; 
and (2) the species is in danger of extinction in that portion. 
Depending on the case, it might be more efficient for us to address the 
``significance'' question or the ``status'' question first. We can 
choose to address either question first. Regardless of which question 
we address first, if we reach a negative answer with respect to the 
first question that we address, we do not need to evaluate the other 
question for that portion of the species' range.
    Following the court's holding in Everson, we now consider whether 
there are any significant portions of the species' range where the 
species is in danger of extinction now (that is, endangered). In 
undertaking this analysis for the Western DPS, we choose to address the 
status question first--we consider information pertaining to the 
geographic distribution of both the species and the threats that the 
species faces to identify any portions of the range where the species 
is endangered.
    Habitat loss, degradation and fragmentation affect gopher tortoise 
populations in the Western DPS at a similar level rangewide. In the 
Western DPS, urbanization, climate change, and incompatible and/or 
insufficient habitat management influence the current and future 
condition of the species at a level comparable to the remainder of the 
range across the DPS. Therefore, we found that the threats are acting 
on the species relatively uniformly across the Western DPS's range. 
However, we identified one portion of the Western DPS range where the 
effects may have a more pronounced effect and, accordingly, that may 
have a different status than the remainder of the DPS. The portion we 
considered was the geographic area of the Western DPS in the State of 
Louisiana, which has seven spatially explicit local populations and 
five landscape populations. The seven local populations in the 
Louisiana portion of the Western DPS exhibit low current resiliency. 
This low resiliency and limited distribution within this geographic 
area may increase the impact of a catastrophic or stochastic event on 
the representation and redundancy of the gopher tortoise in Louisiana. 
We have identified the Louisiana portion as one that has a different 
status than the remainder of the Western DPS.
    We then proceeded to the significance question, asking whether this 
portion of the Western DPS (i.e., Louisiana) is significant. The 
Service's most recent definition of ``significant'' within agency 
policy guidance has been invalidated by court order (see Desert 
Survivors v. U.S. Department of the Interior, 321 F. Supp. 3d 1011, 
1070-74 (N.D. Cal. 2018)). In undertaking this analysis for the Western 
DPS, we considered whether the Louisiana portion of the species' range 
may be significant based on its biological importance to the overall 
viability of the Western DPS. Therefore, for the purposes of this 
analysis, when considering whether this portion is significant, we 
considered whether the portion may (1) occur in a unique habitat or 
ecoregion for the Western DPS of gopher tortoise, (2) contain high-
quality or high-value habitat relative to the remaining portions of the 
Western DPS' range, for the gopher tortoise's continued viability in 
light of the existing threats, (3) contain habitat that is essential to 
a specific life-history function for the species and that is not found 
in the other portions of the DPS, or (4) contain a large geographic 
portion of the suitable habitat relative to the remaining portions of 
the Western DPS.
    This area does not act as a refugia or an important breeding area 
for this portion. It does not contain proportionally higher quality 
habitat or higher value habitat than the remainder of the DPS. It does 
not act as an especially important resource to a particular life-
history stage for the gopher tortoise than elsewhere in the Western 
DPS.
    Overall, there is little evidence to suggest that the Louisiana 
portion of the Western DPS' range has higher quality or higher value 
habitat or any other special importance to the species' life history in 
the Western DPS. In addition, this portion constitutes a small 
proportion of the Western DPS range (approximately 17 percent of 
Western DPS. Thus, based on the best available information, we find 
that this portion of the Western DPS's range is not significant in 
terms of the habitat considerations discussed above. Therefore, no 
portion of the Western DPS's range provides a basis for determining 
that it is in danger of extinction in a significant portion of its 
range. This finding does not conflict with the courts' holdings in 
Desert Survivors v. Department of the Interior, 321 F. Supp. 3d 1011, 
1070-74 (N.D. Cal. 2018), and Center for Biological Diversity v. 
Jewell, 248 F. Supp. 3d 946, 959 (D. Ariz. 2017) because, in reaching 
this conclusion, we did not apply the

[[Page 61866]]

aspects of the Final Policy's definition of ``significant'' that those 
court decisions held to be invalid.

Determination of the Western DPS's Status

    We have determined that the western portion of the gopher tortoise 
range is a valid DPS, and the Western DPS of the gopher tortoise is 
likely to become endangered within the foreseeable future throughout 
all of its range. On the basis of this status review, we continue to 
find the western portion (Western DPS) of the gopher tortoise is a 
threatened species.

Status Throughout the Eastern DPS's Range

    We identified the eastern portion of the gopher tortoise range as a 
candidate species in the July 27, 2011, 12-month finding (76 FR 45130) 
and have included it in the Candidate Notices of Review in subsequent 
years. At the time of the 12-month finding, our assessment indicated 
the species was being impacted by the primary threat of habitat 
destruction and modification (Factor A) due to land conversion, 
urbanization, and habitat management. Other important threats to the 
species at that time included overutilization through rattlesnake 
roundups (Factor B), predation (Factor C), incompatible use of 
silvicultural herbicides (Factor E), and inadequacy of existing 
regulatory mechanisms (Factor D). We had determined disease (Factor C), 
road mortality (Factor E), and the effects of climate change (Factor E) 
to be additional stressors to the species.
    In subsequent CNORs, we reviewed the status of the eastern portion 
of the range (now Eastern DPS) and described additional information and 
conservation actions needed. In addition, we noted that the extent to 
which the many potentially viable gopher tortoise populations are 
sufficient in number, arrangement, and security to ensure the long-term 
viability of the species was unknown. In development of the SSA, we 
compiled and analyzed the best available information including 
population information and conservation measures. We also developed a 
new population viability model based on the best available information; 
this model was not considered in previous CNORs or the original 
petition finding.
    Currently, the Eastern DPS comprises the majority of gopher 
tortoise populations (approximately 84 percent) and habitat with known 
gopher tortoise occurrences (approximately 88 percent) of the gopher 
tortoise range, and, as such, the discussion of threats and the 
species' response to those threats in Status Throughout All of Its 
Range may be applied to the Eastern DPS as well. The Eastern DPS also 
includes the majority of spatially explicit local gopher tortoise 
populations across the range (84 percent or 550 populations), with 127 
populations (19 percent) exhibiting high current resiliency and 169 
populations (21 percent) exhibiting moderate resiliency (table 2). With 
many highly and moderately resilient populations widely distributed 
across the Eastern DPS's geographic area, the species' current level of 
redundancy provides the ability to withstand catastrophic events. The 
Eastern DPS includes four of the identified genetic groups for the 
species, conveying much of the species' representation and adaptive 
capacity. More than 741,330 ac (300,006 hectares) are currently known 
to be occupied by gopher tortoise in the Western DPS with more than 
14.4 million ac (5.8 million ha) of potential habitat where gopher 
tortoise occupancy is unknown. The best available information indicates 
that a greater degree of habitat management occurs in the Eastern DPS 
compared to the Western DPS. Implementation of prescribed fire has 
increased from 3 to 14 times the number of acres burned in 1994, and 44 
to 83 percent of landowners are carrying out additional beneficial 
practices for gopher tortoise (Service 2022, pp. 126-140). Therefore, 
the Eastern DPS is not currently in danger of extinction throughout its 
range.
    Accordingly, we next analyze whether the Eastern DPS is likely to 
become an endangered species within the foreseeable future throughout 
its range. In our consideration of foreseeable future, we evaluated how 
far into the future we could reliably predict the threats to these 
units, as well as the gopher tortoise's response to those threats. 
Based on the modeling and scenarios evaluated, we considered our 
ability to make reliable predictions in the future and the uncertainty 
in how and to what degree the units could respond to those risk factors 
in this timeframe. We determined a foreseeable future of 80 years for 
the Eastern DPS. The methodology and timeframe used to determine the 
foreseeable future for the Eastern DPS followed the process described 
in Status Throughout All of the Western DPS's Range, above. We analyzed 
future conditions based on input from species experts, generation time 
for the species, and the confidence in predicting patterns of climate 
warming, sea level rise, urbanization, and habitat management, enabling 
us to reliably predict threats and the species' response over time. 
Details regarding the future condition analyses are available in the 
SSA report and associated future condition model (Folt et al. 2022, SSA 
2022, appendix B).
    Rangewide threats continue to impact the Eastern DPS in the future, 
including the key drivers of habitat loss and fragmentation due to 
urbanization, climate warming, sea level rise, and habitat management. 
Conservation efforts by Federal, State, and private partners benefit 
the gopher tortoise and its habitat in the Eastern DPS and these 
actions are expected to continue into the future. Although the Eastern 
DPS (Units 2, 3, 4, and 5) is projected to decrease in the number of 
local and landscape populations in the future, 46,176 to 49,697 
individuals, 167 to 175 local populations, and 101 to 107 landscape 
populations are projected to remain across the Eastern DPS into the 
foreseeable future. These populations are distributed across the 
Eastern DPS in the foreseeable future similar to the current 
distribution.
    Based on our analysis of the five factors identified in section 
4(a)(1) of the Act, we conclude that the previously recognized threats 
to the eastern portion of the gopher tortoise range (Eastern DPS) from 
present or threatened destruction, modification, or curtailment of its 
habitat or range (Factor A) (urbanization and development, major road 
construction, incompatible and/or insufficient habitat management, and 
certain types of agriculture) are not impacting the species such that 
the species is in danger of extinction now or in the foreseeable 
future. We evaluated additional potential threats under the five 
listing factors stated above. In that evaluation, we found potential 
impacts such as URTD and other diseases (Factor C), predation (Factor 
C), overutilization (harvest and rattlesnake roundups) (Factor B), and 
nonnative invasive species (Factor E) impact individuals or 
populations, but do not have an impact at the species level at this 
time. Additionally, conservation measures and protection provided by a 
variety of conservation efforts to benefit the gopher tortoise and its 
habitat have been implemented by Federal and State agencies, 
nongovernmental organizations, private landowners, and partnerships 
across the range of the species, and we anticipate these conservation 
measures and protections will continue to benefit the gopher tortoise 
into the foreseeable future (in part due to other sensitive and 
federally listed species occurring in these areas). These conservation 
efforts and

[[Page 61867]]

regulatory mechanisms are in place across the range of the species and 
are addressing some of the identified threats by restoring, enhancing, 
or providing gopher tortoise habitat, relocating tortoises, and 
augmenting populations through captive propagation. See the SSA for a 
thorough discussion of all potential and current threats (Service 2022, 
pp. 46-102).
    Conservation efforts by the Service, State agencies, 
nongovernmental organizations, and private groups as described in 
Conservation Efforts and Regulatory Mechanisms, above, have informed 
our analysis of the species' condition by providing additional 
information regarding species abundance, density, and habitat 
conditions within the range of the species. In addition, habitat 
restoration actions and species-specific conservation measures 
including translocation of individuals and improved awareness of the 
species' needs and threats have contributed to the improved condition 
of the species. In particular, Service-approved plans or other plans 
including the gopher tortoise CCA, CCAA, rangewide conservation 
strategy with the DoD, and the Gopher Tortoise Initiative have resulted 
in the protection of gopher tortoise habitat and populations across the 
range of the species. Many of the management actions and conservation 
easements under these plans are expected to remain in place in the 
future, benefiting the species. The BMPs implemented on working forests 
benefit the gopher tortoise and its habitat; these BMPs are expected to 
continue to be implemented in the future and will continue to benefit 
the species and its habitat.
    Based on our analysis of the five factors identified in section 
4(a)(1) of the Act, we conclude that the Eastern DPS is not in danger 
of extinction or likely to become so in the foreseeable future 
throughout all of its range.

Status Throughout a Significant Portion of the Eastern DPS's Range

    Under the Act and our implementing regulations, a species may 
warrant listing if it is in danger of extinction or likely to become so 
within the foreseeable future throughout all or a significant portion 
of its range. Having determined that the Eastern DPS is not in danger 
of extinction or likely to become so in the foreseeable future 
throughout all of its range, we now consider whether it may be in 
danger of extinction or likely to become so in the foreseeable future 
in a significant portion of its range--that is, whether there is any 
portion of the species' range for which it is true that both (1) the 
portion is significant; and (2) the species is in danger of extinction 
now or likely to become so in the foreseeable future in that portion. 
Depending on the case, it might be more efficient for us to address the 
``significance'' question or the ``status'' question first. We can 
choose to address either question first. Regardless of which question 
we address first, if we reach a negative answer with respect to the 
first question that we address, we do not need to evaluate the other 
question for that portion of the species' range.
    In undertaking this analysis for the Eastern DPS, we chose to 
address the status question first. We began by identifying any portions 
of the range where the biological status of the species may be 
different from its biological status elsewhere in its range. The range 
of a species can theoretically be divided into portions in an infinite 
number of ways, so we focus our analysis on portions of the species' 
range that contribute to the conservation of the species in a 
biologically meaningful way. For this purpose, we considered 
information pertaining to the geographic distribution of (a) 
individuals of the species, (b) the threats that the species faces, and 
(c) the resiliency condition of populations. For the Eastern DPS, we 
considered whether the threats or their effects are occurring in any 
portion of the DPS' range such that the Eastern DPS is in danger of 
extinction now or likely to become so in the foreseeable future in that 
portion of the range.
    The Eastern DPS comprises the majority of gopher tortoise 
populations and habitat across the range of the species, and, 
therefore, threats that affect the species rangewide also affect the 
gopher tortoise in the Eastern DPS. We evaluated the past, ongoing, and 
anticipated threats affecting the species including habitat loss, 
degradation, and fragmentation due to land use changes from 
urbanization, climate warming, sea level rise, and insufficient and/or 
incompatible habitat management. We also considered effects from URTD 
and other diseases, predation, overutilization, and nonnative invasive 
species, and cumulative effects. Conservation efforts and regulatory 
mechanisms also influence the gopher tortoise and its habitat in the 
Eastern DPS. These factors and threats influence the gopher tortoise 
similarly rangewide; however, we identified two portions of the Eastern 
DPS range where the impact of these factors may have a more pronounced 
effect such that it may have a different status than the remainder of 
the DPS. The portions we considered were the geographic area described 
as Unit 5 (Florida) and Unit 2 (Central; west of the Apalachicola and 
Chattahoochee Rivers and east of Unit 1) in the SSA report.
    Sea level rise primarily affect populations along the coast in Unit 
5 (Florida). Although sea level rise is projected to affect coastal 
populations of gopher tortoise, the number of populations affected 
varies by location and elevation of the population, site-specific 
characteristics, and climate change scenario. Of the 21 local 
populations occurring in coastal areas rangewide, 18 of these 
populations occur in Unit 5. Of these 18 coastal populations, 5 
currently exhibit high resiliency and 13 exhibit moderate resiliency. 
Overall, Unit 5 currently has 43 populations that exhibit high 
resiliency and 50 populations that exhibit moderate resiliency. In our 
future projections, small populations in coastal areas decline in the 
same proportion as small populations throughout Unit 5 and rangewide. 
Future condition modeling projects between 58 and 62 local populations 
and 37 to 43 landscape populations will remain on the landscape in Unit 
5, including the very large populations (exceeding 1,000 individuals). 
The current and future condition analyses of gopher tortoise indicate 
sufficient resiliency, representation and redundancy in Unit 5. Given 
the species' current and future condition within this unit, we 
determined that the gopher tortoise in Unit 5 does not have a different 
status than the remainder of the Eastern DPS.
    As described in Status Throughout a Significant Portion of Its 
Range, populations in Unit 2 are generally less resilient and are 
characterized by low abundance, smaller clutch size, lower hatch rate, 
slower growth, and less extensive suitable habitat. Within the Eastern 
DPS, 26.7 percent of the populations in current low resiliency are 
found in Unit 2, which holds 5.9 percent of the abundance in the DPS. 
Although threats are similar throughout the Eastern DPS, the species' 
response is more pronounced in Unit 2 (Central) due to lower 
resiliency, and threats are having a greater impact than elsewhere in 
the DPS. For example, 14 local populations are projected to remain on 
the landscape in Unit 2 (Central) in 2100 under the medium stressors 
and less habitat management scenario. This projected decline in the 
number of populations would increase the impact of a catastrophic or 
stochastic event on the representation and redundancy in Unit 2 
(Central) Given the species' future condition within this units, we 
have identified Unit 2 (Central) within

[[Page 61868]]

the Eastern DPS as an area that has a different status than the 
remainder of the Eastern DPS.
    We then proceeded to the significance question, asking whether this 
portion of the DPS (i.e., Unit 2) is significant. The Service's most 
recent definition of ``significant'' within agency policy guidance has 
been invalidated by court order (see Desert Survivors v. U.S. 
Department of the Interior, 321 F. Supp. 3d 1011, 1070-74 (N.D. Cal. 
2018)). In undertaking this analysis for the Eastern DPS, we considered 
whether the Unit 2 (Central) portion of the Eastern DPS is significant 
based on its biological importance to the overall viability of the 
Eastern DPS. Therefore, for the purposes of this analysis, when 
considering whether this portion is significant, we considered whether 
the portion may (1) occur in a unique habitat or ecoregion for the DPS, 
(2) contain high-quality or high-value habitat relative to the 
remaining portions of the DPS, for the species' continued viability in 
light of the existing threats, (3) contain habitat that is essential to 
a specific life-history function for the species and that is not found 
in the other portions of the DPS, or (4) contain a large geographic 
portion of the suitable habitat relative to the remaining portions of 
the DPS.
    Although Unit 2 (Central) contributes to the condition of the 
species within the Eastern DPS, it does not represent a large area of 
suitable habitat relative to the remainder of the Eastern DPS. Unit 2 
(Central) holds approximately 9.2 percent of suitable habitat with 
known gopher tortoise occurrences in the Eastern DPS, and this habitat 
is of generally lower quality and is less extensive than in the 
remainder of the Eastern DPS. It does not contain proportionally higher 
quality habitat or higher value habitat than the remainder of the 
range. This area does not act as a refugia or an important breeding 
area for this portion. The area does not act as an especially important 
resource to a particular life-history stage for the gopher tortoise 
than elsewhere in the Eastern DPS.
    Overall, there is little evidence to suggest that the geographical 
area of Unit 2 (Central) of the Eastern DPS's range has higher quality 
or higher value habitat to the species' life history in the Eastern 
DPS. In addition, this unit constitutes a small portion of the gopher 
tortoise habitat in the Eastern DPS (approximately 14 percent of this 
portion of the range). Thus, based on the best available information, 
we find that this portion of the Eastern DPS's range is not 
biologically significant in terms of the habitat considerations 
discussed above. Therefore, no portion of the Eastern DPS's range 
provides a basis for determining that the species is in danger of 
extinction now or within the foreseeable future in a significant 
portion of its range. This finding does not conflict with the courts' 
holdings in Desert Survivors v. U.S. Department of the Interior, 321 F. 
Supp. 3d 1011, 1070-74 (N.D. Cal. 2018) and Center for Biological 
Diversity v. Jewell, 248 F. Supp. 3d 946, 959 (D. Ariz. 2017) because, 
in reaching this conclusion, we did not need to consider whether any 
portions are significant and, therefore, did not apply the aspects of 
the Final Policy's definition of ``significant'' that those court 
decisions held were invalid.

Determination of the Eastern DPS's Status

    Our review of the best available scientific and commercial 
information indicates that the Eastern DPS of the gopher tortoise does 
not meet the definition of an endangered species or a threatened 
species in accordance with sections 3(6) and 3(20) of the Act. 
Therefore, we find that listing the Eastern DPS of the gopher tortoise 
is no longer warranted for listing under the Act. With the publication 
of this document, the eastern portion of the gopher tortoise range (now 
the Eastern DPS) will be removed from the list of candidate species.

References Cited

    A complete list of references cited is available on the internet at 
https://www.regulations.gov and upon request from the Florida 
Ecological Services Field Office (see ADDRESSES).

Author(s)

    The primary authors of this notice are the staff members of the 
Florida Ecological Services Field Office and the Species Assessment 
Team.

Signing Authority

    Martha Williams, Director of the U.S. Fish and Wildlife Service, 
approved this action on September 20, 2022, for publication. On 
September 30, 2022, Martha Williams authorized the undersigned to sign 
the document electronically and submit it to the Office of the Federal 
Register for publication as an official document of the U.S. Fish and 
Wildlife Service.

Madonna Baucum,
Chief, Policy and Regulations Branch, U.S. Fish and Wildlife Service.
[FR Doc. 2022-21659 Filed 10-11-22; 8:45 am]
BILLING CODE 4333-15-P