Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e50507, enero-diciembre 2023 (Publicado Feb. 22, 2023)

Jaguar conservation in the American continent: the role of protected

landscape and human-impacted biomes

Erik Joaquín Torres-Romero1,2,3*; https://orcid.org/0000-0002-1078-9223

Gerardo Ceballos 1; https://orcid.org/0000-0001-8374-2656

Francisco Botello4; https://orcid.org/0000-0002-1133-2831

José Ignacio González Rojas5; https://orcid.org/0000-0003-2740-7305

Anthony J. Giordano3; https://orcid.org/0000-0001-6332-2865

José Vicente López-Bao6; https://orcid.org/0000-0001-9213-998X

1. Laboratorio de Ecología y Conservación de Fauna Silvestre, Instituto de Ecología, Universidad Nacional Autónoma

de México, Circuito Exterior s/n, Ciudad Universitaria, Coyoacán, C.P. 04510, Ciudad de México, México;

ejtr23@hotmail.com (*Correspondence), gceballo@ecologia.unam.mx

2. Ingeniería en Biotecnología-Universidad Politécnica de Puebla, San Mateo Cuanalá, Juan C. Bonilla, C.P. 72640,

Puebla, México.

3. S.P.E.C.I.E.S. – Society for the Preservation of Endangered Carnivores and their International Ecological Study, P. O.

Box 7403, Ventura, CA 93006, United States of America; species1@hotmail.com

4. Departamento de Zoología, Instituto de Biología, Universidad Nacional Autónoma de México, Circuito Exterior s/n,

Ciudad Universitaria, Coyoacán, C.P. 04510, Ciudad de México, México; francisco.botello@ib.unam.mx

5. Universidad Autónoma de Nuevo León, Facultad de Ciencias Biológicas, San Nicolás de los Garza, Nuevo León, C.P.

66450. México; jose.gonzalezr@uanl.mx

6. Biodiversity Research Institute (CSIC-Oviedo University, Principality of Asturias), Oviedo University, 33600 Mieres,

Spain; jv.lopezbao@gmail.com

Received 07-IV-2022. Corrected 14-XII-2022. Accepted 08-II-2023.

ABSTRACT

Introduction: Worldwide, expanding human activities continue to be a threat to many large-bodied species,

including jaguars. As these activities continue, it is critical to understand how home range sizes will be impacted

by human-modified landscapes.

Objective: To evaluate the importance of protected and unprotected land on home-range size across their range.

Methods: We used home range data from 117 jaguars in several habitat protection categories and human biome

types. We used a Generalized Linear Mixed Model to test home range and spatial overlap with conservation

categories and human biomes.

Results: Most home-ranges were in Jaguar Conservation Units (62 %), followed by Protected Areas (21 %),

Indigenous People’s Lands (10 %) and Jaguar Movement Corridors (3 %), where 76 % of the jaguars lived inside

one the first three conservation types. However, outside of conserved land, Rangeland, Cropland, Seminatural

land and other human biomes were also important (24 % of the individuals). Jaguars in Rangeland, Cropland

and Seminatural land had the largest home ranges.

Conclusions: Although conservation land was dominant, human-impacted lands appear to play a considerable

role in satisfying the spatial requirements of jaguars.

Key words: conservation planning; human biomes; Indigenous People’s Lands; jaguar conservation units; jaguar

movement corridors; Panthera onca; protected areas.

https://doi.org/10.15517/rev.biol.trop..v71i1.50507

CONSERVATION

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e50507, enero-diciembre 2023 (Publicado Feb. 22, 2023)

INTRODUCTION

In recent decades, anthropogenic pressures

(i.e., habitat destruction, climate change, chem-

ical pollution, overharvesting, and land use

change) on the natural environment and bio-

diversity have increased worldwide (Butchart

et al., 2010). This has left a legacy of species

range contractions and extinctions (Tilman et

al., 2017; Torres-Romero et al., 2020). The

world’s mammalian large carnivores (Order:

Carnivora) are disproportionately impacted by

these pressures, and their remaining range is

increasingly overlapping with human activi-

ties (López-Bao et al., 2017; Torres-Romero

et al., 2020). Most large carnivore species

have undergone extensive range contractions

in recent human history and, are now listed as

threatened with extinction (Ripple et al., 2014;

Torres-Romero et al., 2020).

Variation in the ability of large carnivores

to share the landscape with humans is impor-

tant to both global and regional landscape

conservation planning. Previous studies have

demonstrated different evidence that illustrates

coexistence scenarios between large carnivores

and humans (Boron et al., 2016; Chapron et al.,

2014; López-Bao et al., 2017). Land abandon-

ment and/or conservation policies have facili-

tated coexistence between large carnivores and

humans in different regions, such as Europe:

i.e., the Eurasian lynx (Lynx lynx), brown bear

(Ursus arctos) and wolf (Canis lupus) (Chap-

ron et al., 2014; Cimatti et al., 2021). Sharing

the landscapes with these carnivore species

across many extensive human-dominated areas

remains, however, one of the major barriers to

large mammal’s conservation (López-Bao et

al., 2017; Lute et al., 2018).

Different conservation approaches have

been proposed to protect large carnivores (Rip-

ple et al., 2014). Some strategies have relied on

the use of physical boundaries around protected

areas, such as fences, to prevent conflicts and

reduce threats (Packer et al., 2013); or have

been focused on connecting core protected

areas for these species using habitat corri-

dors (Saura et al., 2019). Alternative strategies

increasingly seek to incorporate the persistence

of large carnivores into human-dominated

landscapes (Chapron et al., 2014; Llaneza et

al., 2018; Lute et al., 2018). This approach

RESUMEN

Conservación del jaguar en el continente americano:

papel de las áreas protegidas y biomas con intervención humana

Introducción: A nivel mundial, la expansión de actividades humanas continúa teniendo un riesgo para muchas

especies de cuerpo grande, tal como los jaguares. Conforme continúen estas actividades, es crucial entender el

impacto de paisajes modificados sobre el tamaño de su territorio.

Objetivo: Evaluar la importancia de terrenos protegidos y no protegidos sobre el tamaño de su territorio a lo

largo de su rango.

Métodos: Usamos datos de tamaño de los territorios de 117 jaguares en varias categorías de protección de

hábitats y biomas humanos. Usamos un Modelo Mixto Lineal Generalizado para probar traslapes espaciales y de

territorios con categorías de conservación y biomas humanos.

Resultados: La mayoría de los territorios estaban en Unidades de Conservación de Jaguares (62 %), seguido por

Áreas protegidas (21 %), Tierras de Pueblos Indígenas (10 %) y Corredores de Movimiento de Jaguares (3 %),

en donde el 76 % de los jaguares vivían dentro de alguna de las primeras tres modalidades de conservación. Sin

embargo, fuera de áreas protegidas, pastizales, tierras de cultivo, terrenos seminaturales y otros biomas humanos

también fueron importantes (24 % de individuos). Jaguares en pastizales, tierras de cultivo, y terrenos seminatu-

rales tuvieron territorios más grandes.

Conclusiones: Aunque las áreas de conservación fueron dominantes, áreas con impacto humano parecieron jugar

un rol considerable en satisfacer los requerimientos espaciales de los jaguares.

Palabras clave: planeación de conservación; biomas humanos; tierras de los pueblos indígenas; unidades de

conservación del jaguar; corredores de movimiento de jaguares; Panthera onca; áreas protegidas.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e50507, enero-diciembre 2023 (Publicado Feb. 22, 2023)

can sometimes lead to more frequent conflicts

between people on how to conserve large car-

nivores, and consequently, greater local pres-

sure threats on carnivores (i.e., retaliatory

killings as retribution for livestock depreda-

tions, increased vulnerability to opportunistic

but illegal hunting) (López-Bao et al., 2017).

Regardless of the conservation strategy adopt-

ed, a lack of understanding regarding how large

carnivores respond to these different conserva-

tion approaches (i.e., spatial use in relation

to landscape protection) limits our ability to

prioritize conservation planning efforts.

The jaguar (Panthera onca) is the largest

felid in the Western hemisphere. It was his-

torically distributed from the Southern United

States, southward into Northern Argentine

Patagonia (Sanderson et al., 2002). Over the

past century, habitat destruction, poaching for

illegal trade in body parts (i.e., skins, claws),

depletion of the predator’s prey base, and retal-

iatory killings following livestock depredations

(Quigley et al., 2017), have contributed to the

range contraction of the species, disappearing

across more than half of its original geographic

range (Sanderson et al., 2002). As land use

intensifies and expands across Americas, this

threat to jaguars will be compounded further

by human population growth, illegal hunting,

medicine markets, livestock production, frag-

mentation, road and other anthropogenic fac-

tors, all of which will only accelerate extinction

risk (Cullen et al. 2016; Torres-Romero et al.,

2020), and thus, effective policies are desper-

ately needed to curb these threats. A review of

suitable landscapes across Central and South

America, highlighted that protected areas are

too small and few to effectively protect jag-

uar populations (Rabinowitz & Zeller, 2010).

Because jaguars range over relatively large ter-

ritories (i.e., average home ranges of 128.6 ±

49.5 km2; González-Borrajo et al., 2017), they

are increasingly overlapping with human activi-

ties in human biomes (Figel et al., 2019; Payan

et al., 2013), which are biomes predominantly

anthropogenic, the product of human popula-

tion growth, land use and land cover change,

croplands and other human influence on eco-

systems (Ellis et al., 2010).

One conservation approach to ensure via-

ble populations of this large felid conserva-

tion is land-sharing (Johansson et al., 2016).

Although, protected areas can be critical to the

viability of low density, wide-ranging species

such as jaguars, private and communal lands

with high-quality habitat and sustainable land

use practices can be important to ensuring

long-term population viability and connectiv-

ity (Sanderson et al., 2002). In Colombia, for

example, recent studies have highlighted the

importance of unprotected areas across human-

modified landscapes to the conservation of

jaguars and their prey (Boron et al., 2016;

Payan et al., 2013). Similarly, private lands

appear to be playing a significant role in the

persistence of jaguars in the Gran Chaco near

the Paraguay-Argentina border (Giordano et

al., 2014; McBride & Thompson, 2018), south

of which jaguars are considered “functionally

extinct” (Quiroga et al., 2014). Understanding

how jaguars use core habitat areas and mixed-

use landscapes across the range of the species

could be used to improve our understanding

of which landscapes might benefit from more

sustainable practices, as well as to identify key

areas essential to jaguar persistence in human-

dominated landscapes.

In this study, we evaluated the relative

importance of different landscapes towards

the extent of jaguar home-ranges across the

Americas, including human biomes (Fig. 1).

We did this by examining different spatial

layers of land use relevant to jaguar conserva-

tion, including: (a) Jaguar Conservation Units;

(b) Jaguar Movement Corridors; (c) Protected

Areas; (d) Indigenous People’s Lands, (e) a

layer combination of potential core habitats

as Jaguar Conservation Units, Protected Areas

and Indigenous People’s Lands, all layers

pooled together, and (f) a layer related to land-

scape transformations caused by direct human

interaction classified as anthropogenic biomes,

also known as “anthromes” or “human biomes”

(henceforth, human biomes; see., Ellis et al.,

2010). Here, we analyze the home range of

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e50507, enero-diciembre 2023 (Publicado Feb. 22, 2023)

Fig. 1. Habitat protection categories and human biomes evaluated on spatial home range of jaguars (Panthera onca) in Latin

America. Abbreviations are: A. jaguar conservation units and corridors network, B. protected areas and indigenous peoples’

lands, and C. human biomes.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e50507, enero-diciembre 2023 (Publicado Feb. 22, 2023)

117 jaguars to address the following questions:

(1) What is the role of Protected Areas, Jaguar

Conservation Units, Jaguar Movement Cor-

ridors, and Indigenous People’s Lands in facili-

tating the movement of jaguars?; (2) What role

do human biomes play in facilitating movement

by jaguars?; (3) Can strategies that encompass

land-sharing across human biomes also play an

important role and if so, to what degree?

MATERIALS AND METHODS

Spatial and Landscape Data: We

acquired the most recent database on jaguar

movement from a database compiled by Mora-

to et al. (2018). This database contains 134 690

spatial locations from 117 jaguars (54 males

and 63 females) tracked using GPS technol-

ogy. These individuals were monitored in five

different countries: Brazil (N = 82), Paraguay

(N = 23), Mexico (N = 8), Argentina (N = 3)

and Costa Rica (N = 1), representing diverse

parts of the jaguar’s range. The majority of

jaguars (N = 111) were adults (> 2 years old),

four jaguars were juvenile, and two jaguars had

undetermined age (see., Morato et al., 2018).

We used information on six layers conser-

vation land protection categories considered

important for jaguar conservation, including

broad categories of human biomes. The six

major spatial layers we considered included:

(1) Jaguar Conservation Units (JCUs), defined

by Sanderson et al. (2002) as “core” habitat

areas of across the jaguar’s range, representing

~2 million km2 and believed to have stable prey

and capable of supporting at least 50 breed-

ing jaguars; (2) Jaguar Movement Corridors

(JMCs), which delineate potential routes of

dispersal, immigration, and emigration among

jaguar conservation units as described by Rabi-

nowitz and Zeller (2010) (Fig. 1); (3) Protected

Areas (PAs), which constitute ~7 million km2

of lands in Americas formally protected for

their natural, ecological or cultural values as

defined by the International Union for Conser-

vation of Nature (UNEP-WCMC, 2020), and

(4) Indigenous People’s Lands (IPLs), which

include terrestrial lands that are managed or

co-managed by Indigenous Peoples (Garnett et

al., 2018), and represented over ~4 million km2

across Latin America. Together, PAs and IPLs

constitute most of the recognized terrestrial

conservation land, habitat protection, and eco-

logically intact landscapes on Earth (Garnett et

al., 2018) (Fig. 1). We used seven distinct PAs

sub-categories, which differ in number and cat-

egory of protection depending on the enabling

laws of each country (see., International Union

for Conservation of Nature, 2020 for further

details of levels and regulatory protection),

including: Ia–Strict Nature Reserves, Ib–Wil-

derness Areas, II–National Parks, III–Natural

Monument or Features, IV–Habitat/Species

Management Areas, V–Protected Landscape/

Seascapes, and VI–Multiple Use Management

& Protected Areas, whereby the sustainable use

of natural resources is permitted.

We also examined (5) the combination

of the three largest aforementioned conserva-

tion land layers together (i.e., JCUs, PAs and

IPLs pooled; henceforth, “Lyr-COMB”), as this

overlap may indicate areas particularly impor-

tant to regional biodiversity, or larger core

habitat. Finally, we evaluated the effectiveness

of (6) human biomes (HBs) layer, which con-

stitute at least some measure of human modi-

fication, pressure, or footprint on the land via

development. We examined this last category,

which included different level and classes of

development activities such as dense settle-

ments, villages, croplands, rangeland, semi-

natural lands, and various other land uses (Fig.

1) (see., Ellis et al., 2010 for further details of

classification and description).

Spatial data analyses and home range

estimate: Because temporal autocorrelation

or serial correlation in the radiotelemetry can

underestimate the true home range size for each

animal (Swihart & Slade, 1985), as well as

bias home range size estimates due to different

collection schedules, monitoring periods, and

sampling frequencies among individuals and

years, we randomly sampled one spatial loca-

tion/day per individual (Börger et al., 2006).

This helped to mitigate against spatio-temporal

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e50507, enero-diciembre 2023 (Publicado Feb. 22, 2023)

autocorrelation, resulting in a total dataset of

19 622 independent spatial locations for sub-

sequent analyses. We then calculated annual

home ranges for each jaguar, incorporating

probabilistic estimators using kernel density

estimation (KDE, via 95 % isopleth as a rep-

resentative area; we refer hereafter to those

results obtained using this metric) using the

Home Range Tools module for ArcGIS (ESRI,

2012) a commonly used method to estimate

an animal’s home range (Laver & Kelly, 2008;

Worton, 1989).

Next, we evaluated how each jaguar’s home

range overlapped with the different habitat pro-

tection categories and land uses, previously

described. For each jaguar home-ranges (JHR)

in our dataset, we calculate the percentage of

the home range overlap where it occurred for

each individual with every selected land cat-

egory. In those cases where > 90 % of the KDE

home range was occupied by a single landscape

category (i.e., JCUs, JMCs, PAs, IPLs or Lyr-

COMB) only that spatial category was consid-

ered. Then, when KDE jaguar home-ranges did

not overlap with any formal conservation land

or habitat protection categories, we identified

the main human biomes (i.e., croplands, range-

land or seminatural lands) overlapping totally

within each home range.

We then tested for differences in jaguar

home-ranges between those located inside con-

servation lands, and the ones being outside or

called “unprotected” areas. In order to deter-

mine if a significant difference in KDE home-

range size exists between jaguars within any

land categories, we built a generalized linear

mixed model (GLMM) with gaussian distribu-

tion error and identity link to test differences

in home range size in relation to the spatial

overlap of jaguars with the different land con-

servation categories considered in this study.

We also included the interaction terms between

sex and land conservation categories. The num-

ber of locations was included as a covariate in

the model to control for potential bias associ-

ated to different sampling efforts. The country

was included as a random factor in the model

(grouping factor, several jaguars sampled by

country and mean home range size). We used

the “glmmAMDM” package to run the model

(Skaug et al., 2013).

Finally, we tested spatial variation in those

cases where KDE home ranges overlapped in

human-modified landscapes or human biomes

(i.e., croplands, rangeland and seminatural

lands); we compared differences of home-

range size between females and males in rela-

tion to each human biome using linear mixed

model (GLMM). All statistical analyses were

performed in program R 3.4.0 (R Core Team,

2014) while spatial analyses, land use mapping

and spatial comparative assessments were per-

formed using ArcGIS (ESRI, 2012).

RESULTS

Our results identified that 89 of 117 indi-

viduals (76 %; 49 females: 40 males) had a

home range size ranging from 155 ± 19 km2

and, overlapped with the combination of land

protection areas together (i.e., Lyr-COMB).

Among these 89 jaguars, the size of home range

of females and males ranged from 99 ± 14 to

224 ± 38 km2, respectively (Table 1). However,

28 of 117 jaguars (24 %; 13 females: 15 males)

utilized “unprotected” lands, i.e., outside the

boundaries of all land with some protected

status combined. Of these jaguars (N = 28),

the size of home ranges was generally larger

though still varied considerably, with females

and males ranging from 211 ± 59 to 762 ± 219

km2, respectively (Table 1). The size of home

ranges varied depending on whether the home

ranges were inside or not of the combination

conservation lands (P < 0.05) (Table 1).

The number of individuals is indicated

as n, and jaguar home-ranges are indicated as

JHR. Mean home range sizes (km2) for females,

males and both sexes, and the results as mean

± standard error (SE) are shown. In each case

(P) in bold represent the level statistically sig-

nificant. Abbreviations are: Jaguar Conserva-

tion Units (JCUs), Indigenous People’s Lands

(IPLs), Protected Areas (PAs), Jaguar Move-

ment Corridors (JMCs), and (Lyr-COMB) rep-

resent three layers categories combined (JCUs,

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e50507, enero-diciembre 2023 (Publicado Feb. 22, 2023)

IPLs and PAs). See main text for details on

how jaguar home-ranges occurred inside and

outside were depicted.

Seventy-three of 117 (62.4 %; 38 females:

35 males) JHR occurred in some part of JCUs,

where females and males JHR ranged from 101

± 18 to 232 ± 41 km2, respectively. While 44 of

117 (37.6 %; 24 females: 20 males) individual

home ranges occurred outside the designated

boundaries of JCUs, where the sizes of JHR

ranging from 157 ± 35 to 614 ± 163 km2 for

males and females, respectively, but jaguar

home-ranges do not show significant differ-

ences (P > 0.05) between inside and outside of

JCUs (Table 1).

We found fewer individuals (10 %; 7

females: 5 males) that overlapped with the

IPLs, which presented home range areas rang-

ing from 157 ± 31 to 384 ± 45 km2, respective-

ly. While more individuals (N = 105) occurred

outside of IPLs, we did not find significant

differences (P > 0.05) in range size both inside

and outside of IPLs (Table 1).

Twenty-five of 117 jaguar home-ranges

(21 %; 13 females:12 males) occurred within

PAs, where ranged from 105 ± 15 to 258 ± 57

km2, respectively. While 92 individuals (~79

%; 49 females: 43 males) were outside of PAs.

The JHR both females and males increased

their home ranges outside the boundaries of

PAs, but we did not find significant differences

(P > 0.05) both inside and outside of PAs (Table

1). On the other hand, only three individual jag-

uars (2 males: 1 female) (2.56 %) overlapped

with potential JMCs; stated differently, the

overwhelming majority of individual jaguar

movements did not overlap at all with any part

of a JMCs (97.43 %; 54 males: 60 females)

and, in both overall, we did not find significant

differences comparisons (Table 1).

Our results show that male and female jag-

uars in protected landscapes averaged smaller

home ranges than male and female jaguars

outside these conservation lands, respectively

(Table 1). Thus, some individuals did not over-

lap their home ranges at all with some catego-

ries of conservation lands considered in this

study. For example, we found that 28 jaguars

TABLE 1

Individual jaguars that occurred inside (in) and outside (out) of each land protection category

JCUs P IPLs P PAs P JMCs P Lyr-COMB P

in out in out in out in out in out

females females females females females

JHR 101 ± 18 157 ± 35 > 0.05 157 ± 31 118 ± 19 > 0.05 105±15 127 ± 22 > 0.05 83 122±18 99±14 211±59 <0.05

n38 (32.48 %) 24 (20.51 %) 7 (5.98 %) 55 (47.01 %) 13 (11.11 %) 49 (41.88 %) 1 (0.85 %) 60 (51.28 %) 49 (41.88 %) 13 (11.11 %)

males males males males males

JHR 232 ± 41 614 ± 163 > 0.05 384 ± 45 370 ± 76 > 0.05 258 ± 57 403 ± 86 > 0.05 473 ± 202 373 ± 70 >0.05 224±38 762±219 <0.05

n35 (29.91 %) 20 (17.09 %) 5 (4.27 %) 50 (42.73 %) 12 (10.25 %) 43 (36.75 %) 2 (1.71 %) 54 (46.15 %) 40 (34.18) 15 (12.82 %)

both sexes both sexes both sexes both sexes both sexes

JHR 164 ± 23 ± 83 > 0.05 251 ± 42 238 ± 39 > 0.05 179 ± 32 256 ± 44 > 0.05 343 ± 174 241 ± 36 >0.05 155±19 507±123 <0.05

n73 (62.39 %) 44 (37.60 %) 12 (10.21 %) 105 (89.74 %) 25 (21.37 %) 92 (78.63 %) 3 (2.56 %) 114 (97.43 %) 89 (76.07 %) 28 (23.93 %)

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e50507, enero-diciembre 2023 (Publicado Feb. 22, 2023)

(24 %; 15 males and 13 females) occurred in

some human biomes lands such as rangelands

(i.e., lands used mainly for livestock grazing

and pasture), followed by croplands (i.e., lands

used mainly for annual crops) and seminatural

lands (i.e., inhabited rural lands, with minor

use for permanent agriculture and settlements)

(Table 2). Based on our dataset, male jag-

uars that occurred in human biomes lands

showed the largest home ranges compared with

females with significant differences in range-

lands (females 249 km2 and males 685 km2;

P = 0.015) and croplands (females 8 km2 and

males 28 km2; P = 0.049), but non-significant

was found in seminatural lands (females 7 km2

and males 21 km2; P = 0.699).

These 28 jaguars were outside the com-

bination of the three lands with protected sta-

tus (i.e., Jaguar Conservation Units, Protected

Areas and Indigenous People’s Lands pooled)

can occur in more than one biome. Note:

Human biomes were organized into groups

according to Ellis et al. 2010 (see Ellis et al.,

2010 for further details of classification).

DISCUSSION

The results presented here correspond to

the first continental-scale evaluation of how

jaguar home ranges overlap with different land

protection status, and anthropized systems, an

important consideration given that previous

jaguar studies have underscored the importance

of protected and unprotected areas for the spe-

cies (Boron et al., 2016; McBride & Thompson

2019; Payan et al., 2013; Thompson et al.,

2021). Our results enlarge information in the

sense of landscape conservation planning for

species with large spatial requirements, which

requires the political and financial commitment

needed to implement ambitious local, regional

and continental conservation and connectiv-

ity strategies (Keeley et al., 2019; McBride &

Thompson 2019; Saura et al., 2019). Indeed,

lands with some “protected” or conservation

status have been shown to be important to jag-

uar populations and other large mammals (Prin-

gle, 2017; Torres-Romero et al., 2020). Thus,

PAs are arguably regarded as the most critical

aspects of world conservation planning efforts,

and they can be particularly critical to carnivore

populations and other vertebrates (Di Minin &

Toivonen, 2015; Wegmann et al., 2014).

On the other hand, indigenous lands are

also crucial to the sustainability and conserva-

tion of wild animal and plant communities on

Earth, occupying an even greater total area

than existing protected areas (Fa et al., 2020;

Schuster et al., 2019). Indigenous People’s

Lands are of course inclusive of land tenure

rights for semi-autonomous and autonomous

indigenous nations across ~38 million km2 in

87 countries; they further intersect with and/

or are adjacent to about 40 % of all terrestrial

protected land area, enhancing the ecological

functioning and landscapes intactness (Garnett

et al., 2018). The IPLs, despite representing

less important areas to space use and move-

ment for jaguars, still represents approximately

10.50 % of ecologically intact landscapes. Over

the past several decades, however, anthropo-

genic pressure on PAs, IPLs and, otherwise,

intact forest habitats have been accelerating

(Qin et al., 2019). In 2019, for instance, bush-

fires devastated approximately 308,048 km2

of the Amazon Basin, nearly 70 % more than

during the period of 2018; large portions

of Brazil, Venezuela, Bolivia, and Colombia

were affected (Lizundia-Loiola et al., 2020).

Other ecoregions in South America, including

the Gran Chaco and Pantanal, which include

important JCUs for jaguars, similarly suffered

devastating losses resulting from intentional

but uncontrolled fires.

Rabinowitz and Zeller (2010) originally

described 90 JCUs encompassing 1.9 mil-

lion km2, and ~98 % of JCUs overlaps with

other conservation lands. In this sense, our

results show that JCUs play a leading role in

the spatial patterns of space and movement of

jaguars in a human-dominated landscape. This

conservation lands have been proposed as criti-

cal jaguar conservation landscapes because of

their perceived importance to long-term jaguar

population viability and connectivity (Rabi-

nowitz & Zeller, 2010; Zeller, 2007; Zeller et

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e50507, enero-diciembre 2023 (Publicado Feb. 22, 2023)

TABLE 2

Percent of contribution of each human biomes vs. 28 individual home range of jaguars

Percent of contribution

Croplands Rangeland Seminatural lands

Lands used mainly for annual crops Lands used mainly for livestock grazing

and pasture

Inhabited lands with minor use for permanent

agriculture and settlements

Sex Country # ind. Residential

rainfed croplands

Populated

rainfed cropland

Remote

croplands

Residential

rangelands

Populated

rangelands

Remote

rangelands Populated woodlands Remote woodlands

Female Paraguay 1 9.73 90.27

Male Paraguay 2 0.5 99.5

Female Paraguay 3 5.15 94.85

Male Paraguay 4 100

Male Paraguay 5 12.36 87.64

Female Paraguay 6 33.61 66.39

Female Paraguay 7 100

Female Paraguay 8 32.07 67.93

Male Brazil 9 77.54 14.04 8.42

Male Paraguay 10 100

Male Brazil 11 79.29 20.71

Female Brazil 12 39.22 60.78

Female Brazil 13 21.63 19.23 59.13

Female Mexico 14 4 96

Male Paraguay 15 100

Male Brazil 16 6.8 48.54 44.66

Male Brazil 17 10.37 23.89 5.55 15 33.34 11.85

Male Mexico 18 44 56

Female Brazil 19 81.25 18.75

Female Brazil 20 8.24 88.84 2.92

Male Brazil 21 77.78 22.22

Male Paraguay 22 100

Female Paraguay 23 100

Female Paraguay 24 100

Male Brazil 25 77.25 22.75

Female Brazil 26 100

Male Brazil 27 6.12 89.69 4.19

Male Brazil 28 41.72 5 828

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e50507, enero-diciembre 2023 (Publicado Feb. 22, 2023)

al., 2011). We should reiterate here that JCUs

are not formal designations by any country and,

while many encompass national protected and

management areas, they may contain a mix of

habitat quality and land use categories.

Similarly, Rabinowitz and Zeller (2010)

identified 182 least-cost path corridors JMCs

connecting the 90 JCUs across the jaguar’s

range. We do note, however, that based on our

results, there is little evidence to suggest that

JMCs are effectively being used by jaguars.

This sort of evidence, however, requires special

attention, because JMCs are not resident range,

and it is therefore not surprising to found that

jaguars do not show a space use and move-

ment on these landscapes. As such, we recom-

mend that future field investigations turn their

attention to this critical knowledge gap, and to

evaluate their use by jaguars. Otherwise, we

face the risk of a proliferation of -paper cor-

ridors- that are never validated in the field.

On the other hand, our study highlights

the relative importance of human biomes to

jaguars. Our analyses show that throughout

their geographic distribution, jaguars include

within their home range human biomes such

as rangeland, cropland, semi-natural lands and

even villages’ lands that do not necessarily have

recognized conservation programs, but that

could be important to spatial movements of

endangered species. This shows that in addition

to the decree of protected areas, it is essential to

recognize and promote the proper management

that various societies have over their territory,

a strategy to do this is by promoting the estab-

lishment of voluntary conservation areas, even

in partially anthropized systems (for example:

areas voluntarily designated for conservation

in Mexico, Gutiérrez-Hernández et al., 2021).

In the Paraguayan Chaco, for instance,

evidence of the presence of jaguars was

encountered regularly in unprotected and/or

privately-owned mixed forest, semi-natural,

and rangeland landscapes; however, at greater

distances from protected areas, their presence

was detected less frequently than pumas (Puma

concolor), a consideration that has different

conservation planning implications for each

species (Giordano, 2015). In parts of Colom-

bia, where hunting of primary prey species and

retaliation killings were limited, jaguars were

able to use human-dominated agricultural land-

scapes (Boron et al., 2016; Payan et al., 2013).

A previous study of female cheetahs (Aci-

nonyx jubatus) showed they tended to prefer

denser vegetation types than males; they thus

seek to mitigate exposure to human pressure

through greater protective cover (Broomhall

et al., 2003). Consistent with our results, we

found that jaguars might be present in different

human-dominated land-use types more broadly

such as croplands, rangeland and seminatural-

lands (Table 2). This finding is congruent

with growing evidence that jaguars occur in

sites with higher prey biomass because there

would be sufficient food, which would increase

the human-carnivore conflict (McBride &

Thompson, 2018) due to depredations on live-

stock and pets, especially when natural prey

and habitat is lacking (Athreya et al., 2016;

Majgaonkar et al., 2019).

Therefore, effective continental, regional

and local jaguar conservation planning may

integrate the different types of human biomes

used by the species (either to favor dispersal

and connectivity between protected areas, or

to support the permanent presence of jaguars)

in the human-dominated matrix (Llaneza et

al., 2018), together with conservation lands, in

order to achieve an effective landscape-scale

conservation approach for the species.

Because of the origin of the dataset, it is

not surprising to find that the ranges of the

collared animals overlap with the categories of

protected lands, but the use of anthropized sys-

tems by the jaguar highlights the opportunity

to promote the participation of different types

of land management at the local, regional, and

continental level for the conservation of this

species. It is also important to recognize the

different conservation schemes in the territory

where landowners carry out surveillance activi-

ties, even in places with anthropized environ-

ments; cooperation between different levels of

land management and conservation is essential

to facilitate jaguar mobility through the JMCs

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e50507, enero-diciembre 2023 (Publicado Feb. 22, 2023)

maximizing the effective protective potential

of the JCUs.

Therefore, the dataset from which we

carry out the analyses presented in this work

come from research works that did not follow

the same protocols, or they did not have the

same telemetry equipment; it also evidently

comes from places where the population char-

acteristics of the jaguar allowed its capture and

subsequent monitoring, often close to areas

expressly designated for the conservation of

biodiversity (Giordano, 2015; Thompson et al.,

2021). In this sense, we recognize that the data

cannot be used to make comparisons among

sites or countries, which is not our aim, and that

it would be necessary to carry out a monitoring

effort on spatial analysis of jaguar home-ranges

in areas where there are no previous efforts to

monitor or conserve the species.

Finally, this continental network of PAs,

IPLs, and JCUs, as well as the incorporation of

HBs connected by JMCs, might theoretically

facilitate the movement of jaguars, while simul-

taneously protecting other endangered species,

and enhancing the functioning of extensive

ecological communities or entire ecosystems.

We further note that some core areas with

respect to JCUs and JMCs may have changed

with respect to land use change and human

impact since Sanderson et al. (2002) and,

Rabinowitz and Zeller (2010). Furthermore,

long-term viability of jaguars across their range

necessitates a strategy of developing functional

connectivity among key areas for the species,

on one hand, and reassesses JCUs and JMCs

on the other. Our work here therefore must be

continued by others and improved upon, so that

we might better understand how jaguars move

across fragmented and human-dominated land-

scapes at regional and continental scales.

Ethical statement: the authors declare

that they all agree with this publication and

made significant contributions; that there is

no conflict of interest of any kind; and that

we followed all pertinent ethical and legal

procedures and requirements. All financial

sources are fully and clearly stated in the

acknowledgements section. A signed document

has been filed in the journal archives.

AGRADECIMIENTOS

Alianza WWF-Fundación Telmex/Telcel.

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