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A systematic review of potential habitat suitability for the jaguar Panthera onca in central Arizona and New Mexico, USA

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In April 2019, the U.S. Fish and Wildlife Service (USFWS) released its recovery plan for the jaguar Panthera onca after several decades of discussion, litigation and controversy about the status of the species in the USA. The USFWS estimated that potential habitat, south of the Interstate-10 highway in Arizona and New Mexico, had a carrying capacity of c. six jaguars, and so focused its recovery programme on areas south of the USA–Mexico border. Here we present a systematic review of the modelling and assessment efforts over the last 25 years, with a focus on areas north of Interstate-10 in Arizona and New Mexico, outside the recovery unit considered by the USFWS. Despite differences in data inputs, methods, and analytical extent, the nine previous studies found support for potential suitable jaguar habitat in the central mountain ranges of Arizona and New Mexico. Applying slightly modified versions of the USFWS model and recalculating an Arizona-focused model over both states provided additional confirmation. Extending the area of consideration also substantially raised the carrying capacity of habitats in Arizona and New Mexico, from six to 90 or 151 adult jaguars, using the modified USFWS models. This review demonstrates the crucial ways in which choosing the extent of analysis influences the conclusions of a conservation plan. More importantly, it opens a new opportunity for jaguar conservation in North America that could help address threats from habitat losses, climate change and border infrastructure.
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A systematic review of potential habitat suitability
for the jaguar Panthera onca in central Arizona and
New Mexico, USA
ERIC W. SANDERSON,KIM FISHER,ROB PETERS,JON P. BECKMANN,BRYAN BIRD
CURTIS M. BRADLEY,JUAN CARLOS BRAVO,MELISSA M. GRIGIONE
JAMES R. HATTEN,CARLOS A. LOPEZ GONZÁLEZ,KURT MENKE
JENNIFER R. B. MILLER,PHILIP S. MILLER,CRISTINA MORMORUNNI
MICHAEL J. ROBINSON,ROBERT E. THOMAS and S HARON WILCOX
Abstract In April , the U.S. Fish and Wildlife Service
(USFWS) released its recovery plan for the jaguar Pan-
thera onca after several decades of discussion, litigation
and controversy about the status of the species in the USA.
The USFWS estimated that potential habitat, south of the
Interstate- highway in Arizona and New Mexico, had a
carrying capacity of c. six jaguars, and so focused its recov-
ery programme on areas south of the USAMexico border.
Here we present a systematic review of the modelling and
assessment efforts over the last  years, with a focus on
areas north of Interstate- in Arizona and New Mexico,
outside the recovery unit considered by the USFWS. De-
spite differences in data inputs, methods, and analytical ex-
tent, the nine previous studies found support for potential
suitable jaguar habitat in the central mountain ranges of
Arizona and New Mexico. Applying slightly modified
versions of the USFWS model and recalculating an Arizona-
focused model over both states provided additional confirm-
ation. Extending the area of consideration also substantially
raised the carrying capacity of habitats in Arizona and New
Mexico, from six to  or  adult jaguars, using the modified
USFWS models. This review demonstrates the crucial ways in
which choosing the extent of analysis influences the conclu-
sions of a conservation plan. More importantly, it opens a
new opportunity for jaguar conservation in North America
that could help address threats from habitat losses, climate
change and border infrastructure.
Keywords Ecological restoration, historical range, jaguar,
Panthera onca, rewilding, spatial model, species distribution
model, USA
Supplementary material for this article is available at
doi.org/./S
Introduction
In April , the U.S. Fish and Wildlife Service (USFWS)
released its recovery plan (USFWS, ) for the jaguar
Panthera onca after several decades of litigation and contro-
versy about the status of, and the future for, this felid in the
USA. The jaguar is listed as an endangered species in the
USA (USFWS, ) and categorized as Near Threatened
on the IUCN Red List (Quigley et al., ). Here we contrib-
ute to these debates by documenting potential habitat for
the jaguar in the central mountains of Arizona and New
Mexico, an area not considered in the latest recovery plan,
and explore the implications of this additional area for
conservation of the species as a whole.
The USFWS jaguar recovery plan described two geo-
graphical units of jaguar conservation, and recovery criteria
for each. The Pan-American Recovery Unit encompassed
all the jaguars range, from Argentina to the USA. The
Northwestern Jaguar Recovery Unit delineated jaguar
habitat in north-west Mexico and the south-west USA,
ERIC W. SANDERSON (Corresponding author, orcid.org /0000-0002-7477-0193)
and KIM FISHER Global Conservation Program, Wildlife Conservation Society,
New York, USA. E-mail esanderson@wcs.org
ROB PETERS,BRYAN BIRD and SHARON WILCOX Field Conservation, Southwest
Office, Defenders of Wildlife, Santa Fe, USA
JON P. BECKMANN and CRISTINA MORMORUNNI Rocky Mountain West Program,
Wildlife Conservation Society, Santa Fe, USA
CURTIS M. BRADLEY Geographic Information Systems, Center for Biological
Diversity, Tucson, USA
JUAN CARLOS BRAVO Mexico and Borderlands Program, Wildlands Network,
Salt Lake City, USA
MELISSA M. GRIGIONE Biology Department, Pace University, Pleasantville, USA
JAMES R. HATTEN Western Fisheries Research Center, US Geological Survey,
Cook, USA
CARLOS A. LOPEZ GONZÁLEZ Facultad de Ciencias Naturales, Universidad
Autonoma de Queretaro, Queretaro, Mexico
KURT MENKE Geographic Information Systems, Birds Eye View, Albuquerque,
USA
JENNIFER R. B. MILLER Center for Conservation Innovation, Defenders of
Wildlife, Washington, DC, USA
PHILIP S. MILLER Conservation Planning Specialist Group, IUCN, Apple Valley,
USA
MICHAEL J. ROBINSON Endangered Species Program, Center for Biological
Diversity, Silver City, USA
ROBERT E. THOMAS Bordercats Working Group, Lakewood, USA
Received  December . Revision requested March .
Accepted  May .
This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (https://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use,
distribution, and reproduction in any medium, provided the original work is properly cited.
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extending from Colima State, along the western slope of the
Sierra Madre Occidental Mountains, into the Sky Island
mountain ranges of northern Sonora, south-eastern Arizona
and south-western New Mexico. The USFWS drew the nor-
thern edge of both units at the Interstate-, suggesting this
highway represents the natural northern extent of jaguars
in the Americas (Fig. ). The northernmost recovery unit
was further subdivided into coreand secondaryareas, includ-
ing the Borderlands Secondary AreaU.S. Portion, which
comprised parts of Arizona and New Mexico south of
Interstate-. Analysis within the recovery plan suggested
that the potential carrying capacity in the USA (within the
Borderlands Secondary AreaU.S. Portion) was only six ja-
guars and, therefore, the plans recommended recovery ac-
tions focused primarily south of the USAMexico border
(USFWS, ; also see Sanderson & Fisher, ).
This conclusion is at odds with several other habitat stud-
ies covering areas north of Interstate-. Numerous photo-
graphs, physical remains, and accounts of jaguars from the
late th century to the s (Brown & Lopez González,
) demonstrate that the species formerly occurred
north of the current highway (the highway dates from
). Ironically some of these historical jaguar records
came from hunters working for the U.S. Bureau of
Biological Survey and its successor organizations, as part
of predator control efforts. Some of these government
hunters placed jaguar skulls and skins in the U.S. National
Museum (for example, Russell Culbreaths kill on the White
Mountain Apache Indian Reservation in ; USNM
). Other accounts come from ranchers, trappers, hun-
ters and tourists, including of jaguars killed near the Grand
Canyon. These observations are cited in a long series of
summary papers and books, published since the s, and
now available online (Sanderson & Fisher, ; USFWS &
Wildlife Conservation Society, ).
Because jaguar observations are infrequent and difficult
to confirm even when they do occur, and because jaguars
have been extirpated from areas where prey and vegetative
cover still remain, researchers have turned to various mod-
elling approaches to estimate the geographical distribution
of potential jaguar habitat in Arizona and New Mexico.
There have been nine such previous efforts by academic
researchers, government wildlife managers, and private
conservation organizations, published before the USFWS
() recovery plan. Some of these models have been de-
scribed in the peer-reviewed literature (e.g. Boydston &
Lopez González, ; Hatten et al., ), whereas others
have only been described in reports, despite their use in
various legal, political and scientific contexts, including the
recovery plan.
Here we conduct the first systematic review (sensu Pullin
& Stewart, ) of the existing modelling and assessment
efforts, with a focus on areas north of Interstate- in
Arizona and New Mexico (Fig. ). Not all of these studies
have been peer reviewed, so we provide sufficient in-
formation for the methods and results of each model or
assessment to be evaluated and compared (Supplementary
Material ). We also present two northward extensions of
the model used by the USFWS () analysing this previ-
ously unconsidered area, and an update of a third model
(Hatten et al., ), for a total of  treatments. We critically
compare the methods and results. Through empirical exam-
ination of convergence among the models and assessments,
we draw a boundary of a third potential recovery area,
tentatively titled the Central Arizona/New Mexico Recovery
Area.
Because legal definitions of range and status drive federal
and state action on species conservation in the USA, we
briefly summarize this history below.
History of jaguar conservation in the USA
The legal definition of jaguar range in the USA has been
disputed for nearly  years. In , jaguars were first
added to the List of Foreign Species protected under the
U.S. Endangered Species Conservation Act of  (USFWS,
), with a note that the species was found in Central and
South America. This listing ignored the numerous well-
attested, historical observations of jaguars north of the bor-
der. In , a new Endangered Species Act lead to a new
consolidated List of Endangered and Threatened Wildlife,
published in  (USFWS, ). The new list included the
jaguar with a note that its normal, known distribution
was Central and South America. Furthermore, the notice
explicitly noted that listed species, such as the jaguar, were
protected wherever found.In, USFWS gave notice
FIG. 1 Study area, with the Northwestern Recovery Unit for the
jaguar Panthera onca and subsections delineated as described in
USFWS (). The extent of the recovery unit in the USA and
Mexico is shown on the inset map.
2 E. W. Sanderson et al.
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that through an oversight, populations of jaguars (and sev-
eral other species) in the USA were not covered by the en-
dangered classifications given to the species as a whole, but
that this oversight would be rectified as quickly as possible
(USFWS, ). In , USFWS proposed listing the jaguar
as endangered in the USA, but the rule was never finalized
(USFWS, ; USFWS, ).
In , USFWS noted there was no resident or breeding
population in the USA at the time, although stragglers oc-
casionally wander into New Mexico, Arizona, and Texas,
where they are generally shot as unwanted predators
(USFWS, ). In , a rancher killed a jaguar in south-
ern Arizona (Brown & Lopez González, ). Two years
later he and another man were convicted of illegal interstate
sales and conspiracy for trying to sell the hide (and two
others) in New Mexico (Associated Press, ).
In , a recovery plan for Listed Cats of Texas and
Arizona (with Emphasis on the Ocelot) noted in its introduc-
tion that Felis (Panthera) onca have been documented as
either transient or resident in Arizona and/or Texas, but
made no specific recommendations for jaguar conserva-
tion (USFWS, ). In , a new proposal for listing
the jaguar in the USA acknowledged that A minimum of
 jaguars have been killed in Arizona since (USFWS,
). In  two different jaguars were separately photo-
graphed by hunters in clearly arid habitats in south-east
Arizona (Glenn, ;Childs,). These were the first
photographs ever taken of live jaguars in the USA. After liti-
gation initiated by the Center for Biological Diversity, in
 USFWS re-designated jaguars as endangered in the
USA (USFWS, ). But the agency also determined that des-
ignating critical habitat was not prudentbecause publication
of detailed critical habitat maps and descriptions in the
Federal Register would likely make the species more vulner-
able to prohibited activities(such as the killing in ).
Jaguars recurred north of the border, nonetheless. Cam-
era traps recorded regular use of the rugged mountains
of extreme south-eastern Arizona and adjacent south-west-
ern New Mexico (McCain & Childs, ). In , the
USFWS determined that a recovery plan would not promote
the conservation of the species, because, they argued, the
USA provided such a small fraction of habitat to the species
range-wide (USFWS, ). This argument was challenged
in court, and overturned in . Fuelling the disputes, a jag-
uar first observed in the late s, Macho B, was captured,
injured, and subsequently euthanized by Arizona state offi-
cials in  (U.S. Department of the Interior, ). Rapid
land development (Povilitis, ) and the expansion of
activity and infrastructure along the border to thwart illegal
immigration (Peters, ) added urgency to the protection
of the jaguar.
Given these events, and in response to lawsuits from the
Center for Biological Diversity and Defenders of Wildlife,
in , USFWS decided it was prudent to declare critical
habitat for the jaguar within the USA and initiated a recov-
ery planning process (USFWS, ). In , USFWS de-
signated , ha of critical jaguar habitat, in southern
Arizona and south-western New Mexico (USFWS, ),
although later a court ruling found the designation invalid
(Bies, ). A draft recovery plan was released for public
comment in  (USFWS, ). The recovery plan was fi-
nalized in  (USFWS, ) and released to the public the
following spring. After  years of controversy, the northern
edge of jaguar range had legally moved  km from the
international border to Interstate-.
Methods
We followed the procedure for systematic reviews in ecology
and conservation recommended by Pullin & Stewart (),
taking care to be transparent, objective and comprehensive
(OLeary et al., ). Our focal question was: Do scientific
models and assessments of potential habitat for the jaguar
Panthera onca in Arizona and New Mexico indicate suit-
ability in areas north of Interstate-? If so, which areas
(Fig. )? To answer this question, we developed a review
protocol based on inclusion criteria. Studies to be reviewed
must have: () examined at least some areas that fall within
the modern boundaries of the States of Arizona and/or New
Mexico, in the USA (spatial criteria), () employed objective
habitat criteria in a model or assessment based on docu-
mented aspects of jaguar ecology (scientific criteria), and
() been documented thoroughly enough that the methods
and results can be peer-reviewed (reviewable criteria). By
modelwe mean a spatial model, computed in a GIS,
using a systematic method. By assessmentwe mean any
other evaluation of jaguar status in this region that meets
the criteria above.
To locate these studies we searched using the search
terms (jaguarOR Panthera onca) AND (ArizonaOR
New MexicoOR United States) in the Web of Science
(Clarivate Analytics, Philadelphia, USA), JSTOR (Ithaka
Harbors, New York, USA), science.gov (a search engine for
U.S. Federal Government scientific publications), and exam-
ined the first  results of a search on Google Scholar
(). We also reviewed all the materials cited in the
USFWS () recovery plan and posted online by USFWS
Arizona Ecological Services (), Arizona Department of
Game and Fish (), and New Mexico Department of
Game and Fish (). Finally, we contacted individuals
who had conducted habitat studies that met our criteria,
based on our first round of review, and asked if they knew
of any additional efforts to include in this study.
For each model or assessment that met the criteria
(Fig. ), we summarized the stated purpose of each analysis,
the extent, the data inputs, a brief description of the model-
ling methods and results (Supplementary Material ), and
Habitat suitability for the jaguar 3
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FIG. 2 Comparison of jaguar habitat models for Arizona and New Mexico: (a) Sierra Institute (), (b) Sanderson et al. (b),
(c) Menke & Hayes (), (d) Boydston & Lopez González (), (e) Hatten et al. (), (f) Robinson et al. (), (g) Grigione
et al. (), (h) Theobald et al. (; note percentage thresholds defined by habitat values in Arizona and New Mexico), (i) USFWS
() (model ), (j) model  (this study), (k) model  (this study), (l) Hatten (this study).
4 E. W. Sanderson et al.
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obtained the relevant GIS layers, if possible. For the Sierra
Institute () analysis, we georeferenced and digitized
the model figure.
In a new analysis, we also extended the USFWS ()
model to the area north of Interstate- as described below.
Previously USFWS () assessed the distribution of jaguar
FIG.2 (Cont.)
Habitat suitability for the jaguar 5
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observations from all of the Northwestern Recovery Unit
(Sanderson & Fisher, ) against putative habitat variables
individually, and combined them with a simple, multivari-
ate model. Input variables were reclassed to include areas
with .and #% tree cover, intermediate, moderate and
high ruggedness (defined in Riley et al., ), and within
 km of a waterway (Hatten et al., ). Areas of high hu-
man influence (defined as a human influence index .;
Sanderson et al., a)andabove, m elevation were
excluded from the potential distribution. The technical
sub-team of the USFWS recovery team assigned weights
based on observed jaguar densities in different ecoregions
(Olson et al., ), with highest weights provided to the sub-
tropical, dry and moist forest ecoregions of Mexico. Lower
weights were assigned to pine and pineoak forest types,
and lowest weights to desert ecoregions. Ecoregions lying
entirely outside the pre-determined Northwestern Recovery
Unit boundary (notably, Arizona Mountain Forests) were ex-
cluded. The habitat suitability model was translated to jaguar
densities (jaguars/ km
) by regressing densities in study
areas against mean habitat values for those areas, with the
y-intercept forced through zero (USFWS, ). Various
iterations of this model were documented in Sanderson &
Fisher (); the final version used in the recovery plan
was labelled model .
We extended the USFWS () model over the rest of
Arizona and New Mexico by assigning the Arizona Mount-
ain Forest, Colorado Plateau Shrubland, and Colorado Rocky
Mountain Forest ecoregions (Olson et al., ) the model
weights .,. and ., respectively (model ). Jaguar ob-
servations have been reported from all of these ecoregions
historically (Brown & Lopez González, ). We also
computed a second extension, raising the elevation cut-off
from , to , m (model ). There is no known bio-
logical basis for a , m cut-off. Jaguars are known to use
areas up to , m in Jalisco, Mexico (Nunez-Perez, pers.
comm., cited in USFWS, ), and a jaguar was shot at an
elevation of ,m in the White Mountains of Arizona in
 (Brown & Lopez González, ; Davis, ). Conser-
vatively, we set our limit at , m, the approximate upper
limit of pineoak forest assemblages in Arizona(Patton et al.,
).
We also extended the Hatten et al. () model over the
entirety of Arizona and New Mexico, with some substitu-
tions of higher-quality GIS data, notably the NHDPlus
(data model version .) database (McKay et al., ), the
GAP/LANDFIRE National Terrestrial Ecosystems database
(Homer et al., ), and the -m National Elevation
Dataset (Archuleta et al., ). These improvements en-
abled the new model (Hatten, this study) to be expressed
at  m resolution.
Finally, we overlaid the models to find areas of con-
gruence by projecting to a common coordinate system
and rescaling the data to km
resolution (Fig. ) to high-
light the area we term the Central Arizona/New Mexico
Recovery Area. This research was conducted during April
December .
Results
Literature review
We reviewed  documents identified through the
literature search. Ultimately, nine papers or reports met the
inclusion criteria: Sierra Institute (), Sanderson et al.
(b), Menke & Hayes (); Boydston & Lopez González
FIG. 3 Areas of convergence of potential
jaguar habitat models in central Arizona
and New Mexico beyond the northern edge
of the Northwestern Jaguar Recovery Unit
described in USFWS ().
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TABLE 1 Comparison of data inputs to estimated jaguar Panthera onca habitat models and assessments, including the three new models in this study, covering some or all of Arizona and New
Mexico. For additional details see Supplementary Material .
Model or assessment Spatial extent
1
Categories of data inputs
Climatic
variables
Ecosystem/
land use/
land cover Elevation
Expert
opinion/
variable
choice
Human
disturbance
Jaguar
observations
Prey
base
Vegetation
cover
2
Soil
type
Topographic
variation
Water
(presence of/
distance to)
Sierra Institute (2000) Sky Islands
region
XXXXX X
Sanderson et al. (2002b) Range-wide X X
Menke & Hayes (2003) Parts of NM X X X X X X X
Boydston & Lopez
González (2005)
AZ, NM, parts
of TX & SON
XXX X XXX
Hatten et al. (2005)AZ X X X X X X X
Robinson et al. (2006)NM X X X X X X
Grigione et al. (2009) AZ, NM, TX,
SON, CHI,
COA, NLE,
TAM
XX
Theobald et al. (2017) NRU X X X X X
USFWS (2018) (model 13) NRU X X X X X X X X
Model 14 (this study) AZ, NM X X X X X X X X
Model 15 (this study) AZ, NM X X X X X X X X
Hatten (this study) AZ, NM X X X X X X X
AZ, Arizona; NM, New Mexico; TX, Texas; SON, Sonora; CHI, Chihuahua; COA, Coahuila; NLE, Nuevo Leon; TAM, Tamaulipas; NRU, Northern Recovery Unit (USFWS, ).
Remotely sensed continuous cover.
Habitat suitability for the jaguar 7
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(), Hatten et al. (), Robinson et al. (), Grigione
et al. (), Theobald et al. () and USFWS (). We
also identified some early or derivative reports (e.g. Hatten
et al., ; Sanderson & Fisher, ; Bravo & Davis, ),
but excluded them because they were largely duplicative
of other studies (Hatten et al., ; USFWS, ; and
Theobald et al., , respectively.)
Comparison of potential habitat models and assessments
for the jaguar
Comparingmodels (see details in Supplementary Material ),
we documented  categories of information used to assess
jaguar habitat potential (Table ; Supplementary Table ). All
assessments used expert opinion to some extent, although
two studies were driven almost entirely by expert consensus:
Sanderson et al. (b) and Grigione et al. (). The
range-wide assessment (Sanderson et al., b) was limited
in Arizona and New Mexico by a historical distribution
boundary from Seymour (), which stopped at the nor-
thern edge of the Sonoran and Chihuahuan deserts. The
study of Grigione et al. () supplemented Sanderson
et al. (b), using similar methods to collect observational
data and redraw areas of knowledge and potential habitat for
the jaguar farther north. These techniques have largely been
supplanted by more quantitative spatial modelling tech-
niques used in other models.
Variable selection differed across the models. All 
models/assessments considered jaguar observations of some
kind; seven considered topographic variation or terrain
ruggedness; seven considered presence of, or distance to,
water; and seven included some measure of human disturb-
ance. Six models included maps of ecosystems or land use/
land cover. Four assessments used elevation thresholds.
Only two models/assessments included explicit consider-
ation of prey base (Sierra Institute, ; Menke & Hayes,
). The three most recent models (Theobald et al., ;
USFWS, ; this study) use remotely sensed measurements
of vegetative cover. Only Boydston & Lopez González ()
used soil types and climate variables.
The treatment of variables varied in terms of grain and
extent, both spatially and temporally. The spatial extents of
models ranged from one or more parts of a state to encom-
passing multiple states in the USA and Mexico (Table ). The
spatial grain or resolution varied from  km
(Boydston
& Lopez González, )to m
(Theobald et al., ).
Three were vector-based (Sierra Institute, ; Sanderson
et al., b; Grigione et al., ); all others were raster-
based analyses. Although all models/assessments included
jaguar observations, they varied considerably as to which ob-
servations were included. As noted above, USFWS ()
only included ecoregions that overlapped the Northwestern
Recovery Unit, thereby excluding from consideration many
areas with older observations, including much of the area
analysed here. Robinson et al. ()detailed observations
from New Mexico, only one of which was used in the USFWS
() recovery plan (i.e. Glenns, , photograph from the
Peloncillo Mountains). Hatten et al. ()used sighting
records from Arizona for .Theobaldetal.()
used only observations with reliable geographical coordi-
nates for .
Modelling methods also varied. As noted above, the
Sanderson et al. (b) and Grigione et al. () assess-
ments were largely expert-driven, drawing polygons on
maps through consensual procedures. Of the spatial models,
six were variants of overlays of relevant habitat variables,
usually with some kind of weighting system, examining
areas of spatial congruence of relevant variables, including
Menke & Hayes (), Hatten et al. (), Robinson
et al. (), Theobald et al. (), USFWS () and
this study. Most of these six studies combined variables ad-
ditively to generate a habitat index; Theobald et al. ()
combined them multiplicatively. The Sierra Institute ()
used overlay techniques, but the resulting map appears to
have been hand-drawn. Boydston & Lopez González (),
in contrast, used the Genetic Algorithm for Rule Set Produc-
tion software (Sachetti-Pereira, ).
For the spatial models, nearly all were expressed in unit-
less indices of jaguar habitat suitability on different scales.
The primary exception was the model used by the USFWS
Recovery Plan (USFWS, ; i.e. model ) and its exten-
sions in this paper (models  and ), which were translated
to estimates of potential jaguar density, based on regression
analysis between the habitat index and density measure-
ments. This translation enabled estimates of potential carry-
ing capacity for jaguars in different polygonal areas, such as
the subunits of the Northwestern Recovery Unit.
Areas of congruence
Despite these differences in variable selection, spatial and
temporal extent, and model formulation, the results are re-
markably consistent about the potential jaguar habitat north
of the Interstate- highway in Arizona and New Mexico
(Figs &). The congruence is sufficient to draw a bound-
ary around what we call the jaguars Central Arizona/New
Mexico Recovery Area. For convenience, we followed the
boundaries of the Arizona/New Mexico Mountains Level
III ecoregion of the U.S. Environmental Protection Agen-
cy (Omernik & Griffith, ; see also Griffith et al., ),
which in turn is similar to the Arizona/New Mexico Forests
(Olson et al., ) and northern portions of the Madrean
Evergreen Forest ecosystem (Brown, ). We also deli-
neated a connector area from to the USFWS () North-
western Recovery Unit called the Arizona Secondary Area.
8 E. W. Sanderson et al.
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We summarized potential habitat scores (i.e. habitat suit-
ability values .) for each of the models and calculated
the area of overlap between the models and the Central
Arizona/New Mexico Recovery Area boundary in Table .
All models found some habitat in this recovery area within
their study extents, varying between  and % depending
on model method and extent. The five models that covered
.% of this area (Hatten, this study; Boydston & Lopez
González, ; Theobald et al., ; models  and ,
this study) suggested there is ,, km
of poten-
tial jaguar habitat in this region, though of variable quality.
Applying the jaguar density estimation methodology used
in the jaguar recovery plan, we estimated the carrying cap-
acity for adult jaguars to be  in the Central Arizona/
New Mexico Recovery Area, and  adults in Arizona
and New Mexico combined (Table ).
Discussion
This systematic review finds that scientific models and
assessments of potential habitat for the jaguar in Ari-
zona and New Mexico indicate suitable areas north of
Interstate- (Fig. ). Our review of the  models/assess-
ments previously conducted or reviewed here support
existence of potential jaguar habitat in a region we have
designated the Central Arizona/New Mexico Recovery
Area (Table ,Figs &). No models contradicted this find-
ing. Even the two studies that did not explicitly consider
this region (Sanderson et al., b;USFWS,;Fig. b,i,
respectively) support the notion of habitat on the margins of
their focal areas, which abut the Central Arizona/New
Mexico Recovery Area. For those exceptions, there are cor-
rections in later work. Grigione et al. (), using methods
largely similar to Sanderson et al. (b), described a
jaguar conservation unit on the Mogollon Rim (Fig. g).
Extending the USFWS () and Hatten et al. () mod-
els north, as we have done here, also suggests the potential
for additional habitat in central Arizona and New Mexico
(Fig. j,k,l).
Initial estimates of carrying capacity for models  and ,
using essentially the same model as the approved recovery
plan (i.e. model ), push the potential of a population in the
USA from six animals in Arizona and New Mexico, which the
USFWS considered non-viable, to  (Table ), which
may be viable. Other well-protected, relatively isolated jaguar
populations have persisted for decades with comparable
population sizes, such as in Corcovado, Costa Ricawith  ja-
guars; Iguazú, Argentina with  jaguars; and, Serra da
Capivara, Brazil with jaguars (see review by Zanin et al.,
). An assessment using population viability analysis, em-
ploying the USFWS () methods, is currently underway.
TABLE 2 Areas of and per cent overlap between potential jaguar habitat models and the Central Arizona/New Mexico Recovery Area. The
recovery area is , km
.
Model
Overlap between model extent & Central
Arizona/New Mexico Recovery Area (km
2
) (%)
% of area of overlap
considered potential habitat
Sierra Institute (2000) 27,256 (33) 79
Sanderson et al. (2002b) 5,025 (6) 100
Menke & Hayes (2003) 22,834 (28) 99
Boydston & Lopez González (2005) 82,422 (100) 69
Hatten et al. (2005) 57,537 (70) 82
Robinson et al. (2006) 24,885 (30) 100
Grigione et al. (2009) 39,289 (48) 51
1
Theobald et al. (2017) 76,690 (93) 42
2
USFWS (2018) (model 13) 4,942 (6) 55
Model 14 (this study) 82,422 (100) 43
Model 15 (this study) 82,422 (100) 68
Hatten (this study) 82,422 (100) 85
Another , km
(% of the recovery area) was designated for further study.
Only habitat values . in arbitrary habitat index numbers, reflecting % habitat suitability (Theobald et al., ).
TABLE 3 Estimated potential carrying capacities (number) for adult jaguars in the USA.
Geographical unit Model 13 (USFWS, 2018) Model 14 (this study) Model 15 (this study)
Borderlands Secondary AreaU.S. portion 6 6 6
Arizona Secondary Area 1 6 7
Central Arizona/New Mexico Recovery Area ,1 69 106
Elsewhere in Arizona & New Mexico 1 9 32
Total potential carrying capacity in the USA 9 90 151
Habitat suitability for the jaguar 9
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This difference in potential carrying capacity, from six to
 animals, is critical information about the potential
for jaguar conservation in the USA. If the jaguars observed
south of Interstate- are only occasional, dispersing males,
inhabiting marginal habitat (Rabinowitz, ), then jaguar
conservation in the USA should focus on Mexican popula-
tions and maintaining connectivity, as the USFWS ()
recovery plan suggests. If, however, there is sufficient poten-
tial habitat north of the Northwestern Jaguar Recovery Unit,
then comprehensive conservation efforts should also con-
sider the recovery of a self-sustaining population within
the USA, beginning with the recovery area identified here.
The recovery plan foresees the necessity to revise the bound-
aries of the recovery unit in the future, noting: While recent
survey and monitoring efforts in south-central and south-
eastern Arizona and extreme south-western New Mexico
have provided important data, as more information is
gathered on the distribution and status of jaguars within
the NRU [Northwestern Jaguar Recovery Unit] and adjacent
areas, the boundaries of the NRU may need to be expanded
or reconfigured(USFWS, ). Here we provide this
information.
Novel approaches to jaguar conservation are needed be-
cause of the politics of the international border. New physical
barriers and increased anti-immigration activity along this
boundary (per U.S. Executive Order No. ,)may
furtherconstrain natural dispersal to and from Mexico, isolat-
ing any potential jaguar population in the USA (Peters et al.,
). In such a circumstance, species recovery would require
establishment of a large population in the USA to ensure
genetic viability and demographic sustainability. Conversely,
if movements across the international boundary can be en-
hanced (Stoner et al., ), an American population might
be part of a regional metapopulation structure, contributing
to long-term viability. In either case bi-national collaboration
will be essential (USFWS, ).
This systematic review highlights some important gaps
in scientific knowledge about the jaguar in North Amer-
ica. The historical limits to the jaguars natural range in
the Americas are poorly understood, both on the contem-
porary northern and southern edges (e.g. Cuyckens et al.,
). In what is now the USA, a number of intriguing,
but difficult to interpret, historical observations exist from
areas far beyond Arizona and New Mexico, including
California, Texas, Louisiana, and other parts of the country
(Sanderson & Fisher, ). Relatedly, we also do not under-
stand how climate change will affect jaguar distributions in
the future (Povilitis, ). Culver & Hein () speculated
on the basis of mitochondrial DNA differences that perhaps
northern jaguars are better adapted to hot, dry conditions.
Since such conditions are expected to prevail in coming
decades in the south-western USA (Garfin et al., ),
future investigations of climate effects on jaguar distributions
and carrying capacity may benefit conservation efforts. At
present, only one of the habitat models reviewed (Boydston
& Lopez González, ) included climatic variables.
The existing models are generally weak on prey availabil-
ity, perhaps the most important determinant of jaguar
habitat (although see Menke & Hayes, ). Along the
international border, jaguars prey primarily on white-tailed
deer Odocoileus virginianus, javelinas Tayassu tajacu and
coati Nasua narica. Farther north, cervids may be more
important to jaguar diets, including white-tailed deer, mule
deer Odocoileus hemionus and perhaps elk Cervus elaphus.
How prey influence distribution and carrying capacity in
this area needs investigation.
Populations on the periphery of a speciesrange may
be critical to the long-term conservation of species (Lesica
& Allendorf, ), especially in a time of climate change
(Gibson et al., ; Povilitis, ). Such populations tend
to be smaller, more isolated, and more genetically and eco-
logically divergent than central ones, which confers on them
novel evolutionary potential and local ecological signifi-
cance (Leppig & White, ). The recognition of addi-
tional potential habitat in the USA will, we hope, inform
range-wide, as well as national, proposals for jaguar recovery
(Jaguar  High-Level Forum, ; USFWS, ).
The USFWS () recovery plan adopted a conservative
view with respect to the former distribution of jaguar habitat
in the USA, despite more than  years of jaguar observa-
tions and nearly decades of habitat models and assess-
ments indicating the plausibility of a wider geographical
distribution. This systematic review of these studies indi-
cates that expanding consideration to areas north of the
Interstate- highway suggests not only a stronghold of
potential habitat in Arizona and New Mexico, but a new
opportunity to restore the great cat of the Americas.
Acknowledgements We acknowledge support for this study from
the Volgenau Foundation, Carroll Petrie Foundation, and institutional
support from Defenders of Wildlife and the Wildlife Conservation
Society. We thank two anonymous reviewers and the U.S. Geological
Survey for helpful suggestions. Any use of trade, firm or product
names is for descriptive purposes only and does not implyendorsement
by the U.S. Government.
Author contributions Study design: BB, JPB, JRBM, CMB, RP, EWS;
data contributions: CMB, JCB, KF, MMG, JRH, CALG, KM, PSM, MJR,
RET, EWS; data analysis: KF, JRH, EWS; writing: all authors.
Conflicts of interest None.
Ethical standards This article abided by the Oryx guidelines on
ethical standards.
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... Historical scholarship has shown that jaguars once inhabited a much larger area (e.g., Brown & López González, 2001), including the rugged, mountain forests of Central Arizona and New Mexico. Much of this area remains potential habitat (Sanderson et al., 2021). Population viability analysis suggests that this habitat block is large enough to sustain a demographically viable population, but that connectivity to southern populations may have been lost (Miller, 2019). ...
... The proposed reintroduction area is vast, covered with suitable vegetation, and well populated with prey. As delimited by Sanderson et al. (2021), the CANRA comprises 82,406 km 2 (Figures 1 and 2), following the outlines of the "Arizona/New Mexico Mountains" ecoregion (Omernik & Griffith, 2014). This area is larger than Jaguar Conservation Units (Sanderson et al., 2002) mapped in adjacent F I G U R E 2 Land use and land cover (a) and land tenure (b) in the Central Arizona-New Mexico Recovery Area (CANRA). ...
... The dotted lines delineate existing and proposed management units, as listed in the caption of Figure 1 Mexico, including units mapped in Jalisco (29,409 km 2 ) and Sonora (13,859 km 2 ), that support viable jaguar populations on much smaller land bases. Using the same carrying capacity methods as the recovery plan (USFWS, 2019), Sanderson et al., 2021, estimated that the CANRA could potentially support 69-106 adult jaguars. Miller (2019) adapted the USFWS' population viability analysis to study a hypothetical CANRA population. ...
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Reintroduction—defined here as the return of a species to a part of its range where it has been extirpated—is a critical pathway to conservation in the 21st century. As late as the 1960s, jaguars (Panthera onca) inhabited an expansive region in the central mountain ranges of Arizona and New Mexico in the United States, a habitat unique in all of jaguar range. Here, we make the case for reintroduction, building a rhetorical bridge between conservation science and practice. First, we present a rationale rooted in the philosophy of wildlife conservation. Second, we show that the species once occupied this territory and was extirpated by human actions that should no longer pose a threat. Third, we demonstrate that the proposed recovery area provides suitable ecological conditions. Fourth, we discuss how return of the species could be a net benefit to people, explicitly recognizing a diversity of values and concerns. Fifth, we show that reintroduction is practical and feasible over a realistic time horizon. Returning the jaguar to this area will enhance the recovery of an endangered species in the United States, further its range‐wide conservation, and restore an essential part of North America's cultural and natural heritage.
... Currently, only male jaguars have been documented in the U.S. in recent decades, and conservation planners are working to design effective corridors linking populations that minimize energy expenditure. Similarly, proposed jaguar reintroduction in the U.S. (Sanderson et al., 2021) would show greater success with such considerations in remediation, especially as recovery will likely require both the reintroduction of female jaguars to Arizona (Babb et al., 2022) and migration of jaguars from Sonora (Sanderson et al., 2022). ...
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... El jaguar (Panthera onca) es el felino más grande del continente americano y se encuentra distribuido desde el norte de México hasta el norte de Argentina, con individuos que han sido eventualmente registrados en el sur de Estados Unidos (Sanderson et al., 2021). Su hábitat original, principalmente bosques por debajo de los 1500 m.s.n.m., han desaparecido con el avance de la deforestación cerca del 50% a lo largo de su distribución (Medellín et al., 2002;Sanderson et al., 2002). ...
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El jaguar (Panthera onca), felino más grande de América, se encuentra amenazado por factores como la deforestación, la cacería, reducción de sus presas y el tráfico de sus partes, y por ello se han realizado varios esfuerzos para la conservación de esta especie a nivel continental. Sin embargo, estas valiosas iniciativas tienen muchas imprecisiones en sus mapas y excluye importantes áreas para la distribución del jaguar. Para esto, basados en un mapa de cobertura de bosque y no bosque y puntos de observación de jaguares en los últimos 20 años presentamos una propuesta de mapas de distribución actual, probable e histórica, para el jaguar, con áreas que cubren entre el 60 y 47% de la superficie del Perú.
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Jaguars ( Panthera onca ) exert critical top-down control over large vertebrates across the Neotropics and have been declining due to multiple threats. Based on geospatial layers, we extracted socio-environmental variables for 447 protected areas across the Brazilian Amazon to identify protected areas that merit short-term high-priority efforts to maximize jaguar persistence. Data were analyzed using descriptive statistics, structural equations and regression modeling. Our results reveal that areas containing the largest jaguar densities and estimated population sizes are precisely those confronting most anthropogenic threats. We reveal that jaguars in the world’s largest tropical forest biome are threatened by deforestation associated with anthropogenic fires, and subsequent establishment of pastures. We provide a shortlist of protected areas that should be prioritized for short-term jaguar conservation. The future predicament of jaguar populations can only be ensured if protected areas can be proofed against downgrading and downsizing geopolitical pressures and external anthropogenic threats.
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Prepared for the 2013 National Climate Assessment and a landmark study in terms of its breadth and depth of coverage, this report blends the contributions of 120 experts in climate science, economics, ecology, engineering, geography, hydrology, planning, resources management, and other disciplines to provide the most comprehensive, and understandable, analysis to date about climate and its effects on the people and landscapes of Arizona, California, Colorado, Nevada, New Mexico, and Utah-including the U.S.-Mexico border region and the lands of Native Nations. What is the climate of the Southwest like today? What has it been like in the past, and how is it projected to change over the 21st century? How will that affect water resources, ecosystems, agricultural production, energy supply and delivery, transportation, human health, and a host of other areas? How vulnerable is the region to climate change? What else do we need to know about it, and how can we limit its adverse effects?. In addressing these and other questions, the book offers decision makers and stakeholders a substantial basis from which to make informed choices that will affect the well-being of the region's inhabitants in the decades to come.
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To preserve biodiversity we need to understand how species are distributed and which aspects of the environment determine these distributions. Human-induced changes in land-cover and loss of habitat threaten many species, particularly large carnivores, in many parts of the world. Differentiating the influence of climate and human land use on the distribution of the jaguar (Panthera onca) is important for the species’ conservation. Historically distributed from the United States to southern Argentina, the jaguar has seen its distribution range decreased at regional and local scales. Here we predict the species’ distribution range using historical records of its presence, climate variables, and MaxEnt predictive algorithms. We focus especially on its southernmost limit in Argentina to indicate the historical limits of this species, and describe its present niche in these edge populations. To estimate the effect of human activity we used a raster of land cover to restrict the jaguar’s distribution. We collected a large amount of presence records through the species’ historical range, and estimated a historical regional distribution ranging from Patagonia up to latitude-50°S. Our findings show the range of the jaguar is decreasing severely in its southern limit and also in its northern limit, and that changes in land cover/use are threats to the species. After subtracting non-suitable land-cover from the studied niche, we found the environmentally suitable area for the jaguar in the study area has decreased to 5.2% of its original size. We thus warn of the high extinction risk of the jaguar in Argentina.
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Given the proliferation of primary research articles, the importance of reliable environmental evidence reviews for informing policy and management decisions is increasing. Although conducting reviews is an efficient method of synthesising the fragmented primary evidence base, reviews that are of poor methodological reliability have the potential to misinform by not accurately reflecting the available evidence base. To assess the current value of evidence reviews for decision-making we appraised a systematic sample of articles published in early 2015 (N = 92) using the Collaboration for Environmental Evidence Synthesis Assessment Tool (CEESAT). CEESAT assesses the methodology of policy-relevant evidence reviews according to elements important for objectivity, transparency and comprehensiveness. Overall, reviews performed poorly with a median score of 2.5/39 and a modal score of zero (range 0–30, mean 5.8), and low scores were ubiquitous across subject areas. In general, reviews that applied meta-analytical techniques achieved higher scores than narrative syntheses (median 18.3 and 2.0 respectively), as a result of the latter consistently failing to adequately report methodology or how conclusions were drawn. However, some narrative syntheses achieved high scores, illustrating that the reliability of reviews should be assessed on a case-by-case basis. Given the potential importance of reviews for informing management and policy, as well as research, it is vital that overall methodological reliability is improved. Although the increasing number of systematic reviews and meta-analyses highlight that some progress is being made, our findings suggest little or no improvement in the last decade. To motivate progress, we recommend that an annual assessment of the methodological reliability of evidence reviews be conducted. To better serve the environmental policy and management communities we identify a requirement for independent critical appraisal of review methodology thus enabling decision-makers to select reviews that are most likely to accurately reflect the evidence base. ã
Technical Report
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Species richness (SR) is a tool that managers can use to include diversity in planning and decision-making and is a convenient and useful way to characterize the first level of biological diversity. A richness list derived from existing inventories enhances a manager's understanding of the complexity of the plant and animal communities they manage. Without a list of species, resource management decisions may have negative or unknown effects on all species occupying a forest type. Without abundance data, a common quantitative index for species diversity cannot be determined. However, SR data can include life his-tory information from published literature to enhance the SR value. This report provides an example of how inventory information can characterize the complexity of biological diversity in the ponderosa pine forest type in Arizona. The SR process broadly categorizes the number of plant and animal life forms to arrive at a composite species richness value. Common sense dictates that plants and animals exist in a biotic community because that community has sufficient resources to sustain life. A mixture of forest attributes maintained in time and space fundamentally supports a certain level of diversity as indicated by a richness value. As a management guideline, it is a reasonable assumption that the variety among plant communities and structures increases the potential for maintaining diverse kinds of animal habitats and resultant populations.
Technical Report
An analysis of jaguar habitat connectivity and identification of potential road mitigation locations in the Mexico-USA Northwestern jaguar Recovery Unit (NRU), with utility range wide