Conservation in Practice
58
Conservation Biology, Pages 58–72Volume 16, No. 1, February 2002
Planning to Save a Species: the Jaguar as a Model
ERIC W. SANDERSON,*‡ KENT H. REDFORD,* CHERYL-LESLEY B. CHETKIEWICZ,* RODRIGO A. MEDELLIN,† ALAN R. RABINOWITZ,* JOHN G. ROBINSON,* AND ANDREW B. TABER*
*Wildlife Conservation Society, 2300 Southern Boulevard, Bronx, NY 10460, U.S.A. †Instituto de Ecologia, Universidad Nacional Autónoma de México, Aparto Postal 70–275, 04510 D.F., Mexico
Abstract:
International conservation planning at the end of the twentieth century is dominated by coarse-filter,supra-organismal approaches to conservation that may be insufficient to conserve certain species such as the jag-uar (
Panthera onca
). If we are to retain broadly distributed species into the next century, we need to plan explic-itly for their survival across their entire geographic range and through political boundaries while recognizing thevariety of ecological roles the species plays in different habitats. In March 1999 the Wildlife Conservation Societysponsored a priority-setting and planning exercise for the jaguar across its range, from northern Mexico to north-ern Argentina. Field scientists from 18 countries reached consensus on four types of information: (1) the spatialextent of their jaguar knowledge, (2) the known, currently occupied range of jaguars, (3) areas with substantialjaguar populations, adequate habitat, and a stable and diverse prey base, and (4) point localities where jaguarshave been observed during the last 10 years. During the exercise, these experts also conducted a range-wide as-sessment of the long-term survival prospects of the jaguar and developed an algorithm for prioritizing jaguarconservation units occurring in major habitat types. From this work, we learned that the known, occupied rangeof the jaguar has contracted to approximately 46% of estimates of its 1900 range. Jaguar status and distributionis unknown in another 12% of the jaguar’s former range, including large areas in Mexico, Colombia, and Brazil.But over 70% of the area where jaguars are thought to still occur was rated as having a high probability of sup-porting their long-term survival. Fifty-one jaguar conservation units representing 30 different jaguar geographicregions were prioritized as the basis for a comprehensive jaguar conservation program.
Planeación para Salvar una Especie: El Jaguar como Modelo
Resumen:
La planeación de la conservación internacional al final del siglo veinte esta dominada por en-foques de grano grueso, supra-organísmicas que pueden ser insuficientes para conservar ciertas especiescomo el jaguar (
Panthera onca
). Si hemos de mantener especies ampliamente distribuidas en el próximo si-glo, necesitamos planificar su supervivencia explícitamente en todo su rango geográfico a través de límitespolíticos al mismo tiempo que se reconozca la variedad de funciones ecológicas de las especies en diferenteshábitats. En marzo de 1999 la Sociedad de Conservación de Vida Silvestre promovió un ejercicio dedefinición de prioridades y de planeación para el jaguar en todo su rango de distribución, desde el norte deMéxico hasta el norte de Argentina. Científicos de 18 países llegaron a consensos en cuatro tipos de infor-mación: (1) la extensión espacial de su conocimiento del jaguar, (2) el rango conocido, actualmente ocupadopor el jaguar, (3) áreas con poblaciones importantes, hábitat adecuado y una base de presas estable y di-versa y (4) localidades en las que se han observado jaguares durante los últimos 10 años. Durante el ejerci-cio, estos expertos también hicieron una evaluación de la supervivencia a largo plazo del jaguar en todo surango y desarrollaron un algoritmo para priorizar unidades de conservación del jaguar en los tipos de hábi-tat más importantes. De este trabajo, aprendimos que el rango del jaguar conocido y ocupado se ha con-traído aproximadamente al 46% de su rango estimado
circa
de 1900. El estatus del jaguar y su distribuciónen otro 12% del rango anterior, incluyendo extensas áreas en México, Colombia y Brasil. Sin embargo, másdel 70% del área donde se piensa que todavía ocurre el jaguar fue considerada con una alta probabilidad desoportar la supervivencia a largo plazo. Se priorizaron 51 unidades de conservación representando 30 re-
giones diferentes como la base para un sólido programa de conservación del jaguar.
‡
email [email protected] submitted August 14, 2000; revised manuscript accepted April 25, 2001.
Conservation BiologyVolume 16, No. 1, February 2002
Sanderson et al. Range-wide Conservation Planning for the Jaguar
59
Introduction
Over the last 100 years the theory and practice of conserva-tion has evolved from strategies originally intended to pre-serve natural resources or awe-inspiring scenery (Callicott1990) to an intense concern for conserving biodiversity inall its facets, including genetic and species diversity and thediversity of ecosystem structure and function ( Redford &Richter 1999). This evolution has been driven by the dis-coveries of twentieth century science that have revealedthe vast diversity of biological species and the intricate andsubtle ways in which organisms interact with one anotherand with human beings—and that have thus engenderedhorror at so many species being lost through human hasteand greed (Ehrlich & Ehrlich 1981; Mann & Plummer1995). At the end of the twentieth century, the new para-digm of conservation is biodiversity writ large, includinggenetic, ecosystem, and landscape perspectives.
Simultaneous with this increasing emphasis on biodiver-sity in all its components has been an increase in the scaleof planning for conservation work, typically through mech-anisms that emphasize entities other than the populationor the species as a target for conservation effort (Noss1991; Salwasser 1991). International conservation organiza-tions, governmental and nongovernmental, have alteredtheir approach to focus increasingly on strategies that areregional to global in scope and based on conserving supra-organismal entities: hotspots of species diversity (Myers etal. 2000), globally significant ecoregions (Olson & Diner-stein 1998), ecosystems (as in ecosystem management;Boyce & Haney 1997), endemic bird areas (Slattersfield etal. 1998), and continental networks (Soulé & Terborgh1999). Such approaches seek to conserve ecosystem func-tions and the diversity of habitat types despite a lack ofknowledge of the extent of biological diversity and thecomplex array of factors that maintain it ( Hunter 1991;Franklin 1993). In short, they seek to conserve the wholewhen faced with the impossibility of knowing all the parts.
But the parts are important too. Here we provide an ex-ample of how one such important part, the jaguar (
Pan-thera onca
), can form the basis for large-scale conservationplanning. Jaguars have much to teach us about the knottyproblem of conserving broadly distributed species. Be-cause jaguars as a species range across many different na-tions and habitat types, small-scale conservation efforts se-lected ad hoc and focused over narrowly defined areashave not succeeded in stemming the tide of jaguar extirpa-tion (Weber & Rabinowitz 1996). Establishment of the firstjaguar reserve in Belize (Rabinowitz 1986) and the creationof a conservation plan for the Pantanal (Quigley & Craw-shaw 1992), although important, have not slowed the col-lapse of the jaguar’s range. Moreover, range-wide conserva-tion efforts have been inhibited by national and linguisticdifferences among conservationists, lack of knowledge onthe overall status of jaguars, and the absence of a consensuson priorities for conservation of the species.
Saving jaguars requires international, range-wide plan-ning that recognizes as a first priority ecological, not politi-cal, distinctions among jaguars. We postulate that saving aspecies means, at least, saving populations of the species inall the significantly different ecological settings in whichthey occur. As Wikramanayake et al. (1998) wrote about arelated species, “in seeking to conserve representative pop-ulations of tigers, we must consider not only the geneticdistinctiveness of tigers across the range, but also behav-ioral, demographic, and ecological distinctiveness.” Thus, itis not sufficient to pursue jaguar conservation efforts onlyin tropical forests or only in tropical forests in Brazil and Be-lize; we must begin with the range-wide context that forthe jaguar requires an international perspective.
Most species-based conservation efforts do not assumeas a starting point consideration of the entire range. Con-servation of endangered populations of the CaliforniaGnatcatcher (
Polioptila californica
) is an imperative un-der the U.S. Endangered Species Act, even though sub-stantial, unthreatened populations exist in Mexico (Zinket al 2000). Moreover, most countries do not have endan-gered species legislation of any kind, and if they do, lawsare unlikely to be consistent across the 18 nations wherethe jaguar is currently found. As a result, biological con-servation plans often respect political boundaries morethan ecological ones (Hunter & Hutchinson 1994.)
In 1999 the Wildlife Conservation Society and the Insti-tute of Ecology at the National Autonomous University ofMexico initiated a geographically based, range-wide as-sessment and priority-setting exercise for the jaguar(Medellin et al. 2001). Our goals were to comprehen-sively assess the state of knowledge about the ecology,distribution, and conservation status of the jaguar, toidentify priority areas for its conservation on a range-wide basis, and to build an international consensus forconservation of the species. This work was built on ageographic data framework that respected the kinds andqualities of information we now have while forming abaseline for future evaluations. From this information,the experts assessed the status of jaguars across the rangeand developed a prioritization mechanism to determinethe most important areas for jaguar conservation in eachregional habitat type, based on factors important for thelong-term survival of jaguars. Although the results focuson the range-wide condition of jaguars, the methodologyused and the conclusions drawn present a model for con-servation planning that could be applied to many widelyranging species.
Methods
Data Definitions
Jaguar geographic regions ( JGRs) are geographic unitsdefined by potential habitat (sensu Hall et al. 1997) and
60
Range-wide Conservation Planning for the Jaguar Sanderson et al.
Conservation BiologyVolume 16, No. 1, February 2002
bioregion across the jaguar’s historic range ( Fig. 1). Pre-sumably, the ecology of jaguars in tropical moist low-land forest is significantly different from that in xericdeserts because of differences in, for example, prey baseand habitat use. Similarly, because of regional differencesin species composition and geographic factors, the roleof jaguars in the tropical moist lowland forests of CentralAmerica is substantively different from their role in thetropical moist lowland forests of the southeast Amazon.Representing these ecological differences geographi-cally through JGRs provides a convenient, ecologicallybased unit for planning.
Each JGR is named by its geographic region, then itshabitat type (e.g., northeast Amazon/tropical moist low-land forest). The limits of the historic range were ap-proximated from Seymour (1989) as the range of jaguarsaround 1900. This historic range was subdivided into 36JGRs by lumping together North American and SouthAmerican ecoregions (Dinerstein et al. 1995) to createunits similar to the regional habitat types used in a previ-ous conservation priority-setting exercise for LatinAmerica ( Biodiversity Support Program et al. 1995).Lumping together geographic units required includingsome areas that likely were never occupied by residentjaguars, including some areas above 2000 m in theAndes and Tepuis, which may overestimate the historicrange slightly. The historic range may be slightly under-estimated on its margins, particularly in the southwest-ern United States, but we estimated that
�
5% of the areain the JGRs is subject to these problems. The total ex-tent of the historic range, represented in this way, is ap-proximately 19.1 million km
2
.Thirty-five jaguar experts from 12 nations attended the
workshop on “Jaguars in the New Millennium” (March1999) or contributed information through another ex-pert (for a list of participants, see the Appendix). Priorto the workshop, each researcher was provided a basemap for his or her self-reported area of expertise at1:2,000,000–1:4,000,000 scale, showing a preliminary setof jaguar geographic regions and basic reference infor-mation such as national boundaries, major cities, and riv-ers (Lioutty 1996). We assumed that each expert couldidentify jaguar locations on the map within 1 cm (20–40km in map units). These data were compiled in geo-graphic information system databases (Arcview–GIS),and each datum was identified with the name of thecontributing expert. At the workshop these data wereexamined systematically in regional groups to resolvecontradictions and build a consensus information base.
Four basic data types were solicited from the jaguarexperts: (1) the geographic extent of their knowledgeabout jaguar status and distribution—whether or not jag-uars are present in an area (“extent of knowledge”); (2)the area where jaguars were present as of March 1999(“known, currently occupied range”); (3) important ar-eas for jaguar conservation as defined below (“jaguar
conservation units”); and (4) point localities where jag-uars have been observed within the last 10 years (“pointobservations”) ( Fig. 1). Experts were asked to combineall observations within 20 km of the center coordinatesof the point locality. Each point observation was charac-terized by dates of first and last observation, observationmethods used, and observer.
Jaguar conservation units (or JCUs) were defined ei-ther as (1) areas with a stable prey community, currentlyknown or believed to contain a population of residentjaguars large enough (at least 50 breeding individuals) tobe potentially self-sustaining over the next 100 years, or(2) areas containing fewer jaguars but with adequatehabitat and a stable, diverse prey base, such that jaguarpopulations in the area could increase if threats were al-leviated. Jaguar conservation units were not restricted toor required to contain protected areas. After the work-shop, each JCU was given a name based on an adjoiningor encompassing protected area, river, administrativeunit, or other geographic feature.
In addition, the experts developed a geographicallycomprehensive consensus on the status of jaguars acrossthe range by assigning the following codes to entireJGRs or divisions of JGRs, as necessary. Areas that wereunknown were designated “status unknown—priorityfor survey.” Areas that were known but were no longeroccupied by jaguars were designated “no jaguars.” Forareas that were known and currently occupied by jag-uars, one of the following three classes was assigned: (1)high, (2) medium, or (3) low probability of long-termsurvival. These assignments were based on qualitativeevaluation of habitat size and connectivity, the status ofthe prey base, the status of jaguar populations, and thelevel of threat from human activity.
Prioritization of Jaguar Conservation Units
The experts were asked to weight six factors ( JCU size,connectivity, habitat quality, hunting of jaguar, huntingof prey, and population status) according to their rela-tive importance for long-term jaguar survival, keepingthe sum of all weights to 100 points. To ensure maxi-mum input, weighting schemes were developed sepa-rately by two discussion groups and then an attemptwas made to synthesize the schemes in plenary session.During the review period (described below), the au-thors determined the final weighting scheme in consul-tation with the workshop participants, as follows: JCUsize (30 points), connectivity (23), habitat quality (23),hunting of jaguars (10), hunting of prey (10), and jaguarpopulation status (4).
Each JCU was assigned to the JGR where the majority ofits area occurred. In cases where a JCU overlapped morethan one JGR, the JCU was assigned to JGRs with whichit shared over 1250 km
2
of area, the equivalent of onepoint observation. (Each point observation represents a
Conservation BiologyVolume 16, No. 1, February 2002
Sanderson et al. Range-wide Conservation Planning for the Jaguar
61
Figure 1. Spatial distribution of jaguar data across jaguar geographic regions ( JGRs; background colors of each figure): (a) extent of current knowledge about the jaguar, (b) distribution of known, currently occupied jaguar range as of March 1999, (c) distribution of jaguar conservation units ( JCUs), and (d) distribution of jaguar point observations from 1990 to 2000.
62
Range-wide Conservation Planning for the Jaguar Sanderson et al.
Conservation BiologyVolume 16, No. 1, February 2002
circular area of 20-km radius.) For those JGRs with lessthan three JCUs, the size criteria was relaxed to includeall JCUs that occurred in that JGR, no matter the amountof overlap. The total size of the JCU, not the area of JCUwithin a given JGR, was used for calculating priorities,because jaguar populations were assumed to use the en-tire JCU, not just the portion within one habitat type.
The final prioritization score for each JCU was deter-mined by multiplying the JCU’s score for each of the sixfactors by its corresponding weight and then adding thescore/weight products. The JCUs within the same JGRwere then ranked to determine the most important JCUwithin each JGR.
The final results were compiled at the workshop andwere subsequently reviewed during a post-workshop re-view period. In consultation with the appropriate ex-perts, the authors made final decisions on inconsisten-cies between data sets. All data were distributed to theparticipating experts following the workshop, and alldata were made available one year after the workshop atwww.savethejaguar.com. More extensive details on allthe methods are provided by Medellin et al. (2001).
Results
Extent of Jaguar Knowledge
The extent of knowledge about jaguars—including areaswhere jaguars are not present—covers 83% of the his-toric range of jaguars, indicating that approximately 17%of the range is unknown ( Fig. 1a). Most of the area forwhich jaguar information is lacking is in several large re-gions in Mexico (over 848,000 km
2
) and in South Amer-ica (over 2.3 million km
2
in Brazil alone). The distribu-tion of jaguar knowledge by JGR reflects the distributionof these large, unknown areas (Table 1). The Mexicanpine-oak temperate forests and the Caatinga xerics inBrazil are the least-known JGRs.
Known, Currently Occupied Jaguar Range
Jaguars are known to range over approximately 8.75 mil-lion km
2
, or 46% of their historic range, broken into 48separate areas that range in size from 114 km
2
to over 7million km
2
( Fig. 1b). Unknown areas ( Fig. 1a) were notincluded as part of the known, currently occupiedrange. The largest contiguous area of jaguar range is cen-tered in the Amazon Basin (88% of occupied range) andincludes adjoining areas in the Cerrado, Pantanal, andChaco to the south, extending to the Caribbean coast ofVenezuela and the Guianas. The Colombian Cordillerassever the connections between this contiguous rangeand a series of large ranges that stretch across CentralAmerica from the Darien to the Selva Maya in Belize,Guatemala, and Mexico. At the northern end of the range
through Mexico, known jaguar range is limited to a stripalong the western coast and three isolated observationsin the southwestern United States.
Most of the loss of occupied range has occurred innorthern Mexico, the southern United States, northernBrazil, and southern Argentina. Jaguars have been extir-pated completely from the Argentine Monte and Pampasgrasslands in southern South America and the westerngulf coastal grasslands in the United States (Table 1). Be-cause of elevation limits, jaguars are also not regularlyfound in the Pantepui or Puna montane grasslands. Jag-uars are found in 10% of the area of Mexican xerics, Par-amo, and Mexican pine-oak forests.
Jaguar Conservation Units
Based on present jaguar population size, prey base, andhabitat quality in specific areas, the experts identified 51areas (1.29 million km
2
, 6.7% of the historic range, 13%of the currently occupied range) (Fig. 1c) important tothe long-term survival of jaguars. By definition, each JCUrepresents a core population of jaguars on which con-servation might be based.
Jaguar conservation units are found wholly or partiallyin 31 of the 36 JGRs and thus represent most of the eco-logical settings where jaguars occur (Table 1). Three ofthe six JGRs not represented by a JCU are in areas wherejaguars have been extirpated or apparently were neverpresent in large numbers: Argentine monte, western gulfcoastal grasslands, and Pantepui montane grasslands.The western Andean tropical dry forests, Amazonianmangroves, and Amazonian savannas are also not repre-sented by a JCU because of lack of information aboutjaguar status in those habitat types.
Jaguar Point Observations
The experts reported 5680 observations of jaguars at535 separate localities during the last 10 years, repre-senting a total area of observation of approximately513,000 km
2
, or approximately 2.7% of the jaguar’srange that has been directly sampled ( Fig. 1d). An aver-age of 10.6 jaguar observations were made at eachpoint, indicating concentrations of research at mostpoints. Sixty percent of point observations recorded jag-uars based on at least one of the more reliable observa-tion methods: direct sighting by researcher, photograph,radiotelemetry, capture, or discovery of jaguar remains.
The density of point observations is uneven across thejaguar’s range, reflecting concentrations of research ratherthan concentrations of jaguars. The most richly studiedJGR is the Central American tropical moist forests (Table1), due largely to research in Costa Rica, Belize, and Guate-mala, although extensive research has also been con-ducted in Brazillian Cerrado and the Chaco dry tropicalforests of Bolivia and Paraguay. The JGR of the northeast
Conservation BiologyVolume 16, No. 1, February 2002
Sanderson et al. Range-wide Conservation Planning for the Jaguar
63
Table 1. Distribution of jaguar data sets by jaguar geographic region.
Extent of knowledge
(%)
c
Known, occupied
range (%)
d
Percentage of area ratedwith given probability of
jaguar long-termsurvival
h
Jaguar geographic regions ( JGRs)
a
Area ( km
2
)
b
JCU(%)
e
No.JCUs
f
No.points
g
high(%)
medium(%)
low(%)
1.1 Atlantic/Tropical Moist Lowland Forest 951,120 98 20 7 3 36 1 0 191.2 Upper Amazon/Tropical Moist
Lowland Forest 2,965,517 81 80 8 10 40 80
�
1 01.3 Northeast Amazon/Tropical Moist
Lowland Forest 1,520,518 81 81 11 5 15 80
�
1
�
11.4 Southeast Amazon/Tropical Moist
Lowland Forest 1,358,285 61 61 5 5 29 60 1 01.5 Choco-Darien/Tropical Moist
Lowland Forest 231,577 72 70 27 3 8 27 42 01.6 Central American/Tropical Moist
Lowland Forest 522,443 95 77 24 11 118 71 1 22.1 Tropical Andes/Tropical Moist
Montane Forest 756,615 91 27 5 5 38 1 22 42.2 Central American/Tropical Moist
Montane Forest 190,510 65 25 8 8 17 1 9 142.4 Venezuelan Coastal Montane
Forest/Tropical Moist Montane Forest 14,341 95 51 9 1 3 0 51 02.5 Guayana Montane Forest/Tropical
Moist Montane Forest 337,586 100 100 18 4 2 100 0 03.1 North South American/Tropical Dry Forest 163,710 95 66 3 3 6 0 41 253.2 Western Andes/Tropical Dry Forest 104,683 60 10 0 0 0 0 8 33.3 Chaco/Tropical Dry Forest 1,153,437 93 35 8 3 79 30 0 53.4 Central American/Tropical Dry Forest 55,595 86 14 5 2 6 11 2 13.5 Mexican/Tropical Dry Forest 301,289 42 11 7 5 10 9
�
1 13.6 Cerrado/Tropical Dry Forest 2,411,425 91 57 6 9 34 11 28 194.1 Mexican/Xeric 1,280,778 95 1 1 1 4 1 0 04.2 Caribbean/Xeric 123,232 98 44 1 2 2
�
1 44 04.3 Caatinga/Xeric 759,625 27 10 1 1 5 0
�
1 104.6 Argentine Monte/Xeric 409,040 100 0 0 0 0 0 0 05.1 Central American Pine
Savanna/Herbaceous Lowland Grassland 18,847 96 96 35 1 4
�
1 0 955.2 Llanos–Gran Sabana/Herbaceous
Lowland Grassland 493,095 91 90 1 5 27 1 73 165.3 Pampas/Herbaceous Lowland Grassland 1,098,214 100 0
�
1 1 0
�
1 0 05.5 Amazonian Savanna/Herbaceous
Lowland Grassland 173,461 53 53
�
1 1 4 13 0 05.6 Pantanal/Herbaceous Lowland Grassland 171,053 100 100 48 3 17 99 0 15.7 Western Gulf Coastal Grassland/Herbaceous
Lowland Grassland 42,653 100 0 0 0 0 0 0 06.1 Paramo/Herbaceous Montane Grassland 78,706 83 3
�
1 1 0
�
1 1
�
16.2 Puna/Herbaceous Montane Grassland 589,486 100 0
�
1 1 2 0
�
1
�
16.4 Pantepui/Herbaceous Montane Grassland 48,836 100 0 0 0 1 0 0 07.2 Brazillian Araucaria/Temperate Forest 220,917 100 27 6 2 9 6 0 217.3 Mexican Pine-Oak/Temperate Forest 460,465 20 8 8 6 4
�
1 5 38.1 Northern Mexico/Mangrove 7,070 75 34 20 1 2
�
1 34 08.2 Central American/Mangrove 38,237 65 35 9 9 12 6 2 258.3 Northern South America/Mangrove 20,272 53 16 4 1 0 5 8 08.4 Amazonia/Mangrove 33,235 82 82 0 0 1 74 4 08.5 Eastern South America/Mangrove 10,162 54 24 19 1 0 0 0 23
a
The JGRs are named by region/habitat type. Numeric codes cross reference Fig. 1, and GIS data sets distributed through www.savethejaguar.com.
b
Area measured in equal area azimuthal projection, central meridian -72, reference latitude 0.
c
The JGR area where status and distribution of jaguars is known.
d
The JGR area where jaguars are known to exist.
e
The JGR area with substantial jaguar populations, a stable and diverse prey base, and adequate habitat ( JCU, jaguar conservation unit).
f
Jaguar conservation units wholly or partially within each JGR.
g
Jaguar point observations in each JGR.
h
See text for details.
64
Range-wide Conservation Planning for the Jaguar Sanderson et al.
Conservation BiologyVolume 16, No. 1, February 2002
Amazon has been relatively undersampled in comparisonwith other tropical forest types. The density of sampling islowest among xeric formations and herbaceous montanegrasslands, where jaguars are relatively rare.
Range-Wide Assessment
The range-wide assessment showed variation in jaguarstatus across the range (Fig. 2). Of the jaguar’s historicrange, 18% is unknown, and jaguars are known to havebeen extirpated in an additional 37%. Within the known,currently occupied range, the probability of long-termsurvival of the jaguar in 70% of the area (over 6 millionkm
2
) is considered high. The largest of these high-proba-bility portions of the range is centered on the AmazonBasin and the adjoining Gran Chaco and Pantanal. Twodisjunct sections of the tropical moist lowland forests ofCentral America are also considered areas in which theprobability of long-term jaguar survival is high: SelvaMaya of Guatemala, Mexico, and Belize, and a narrow,continuous strip from the Choco-Darien of Panama andColombia to northern Honduras. Areas in Jalisco, the Si-erra Madre of Mexico, and in the Missiones district of Ar-
gentina were also identified as areas where the probabil-ity of long-term jaguar survival is high.
Of the currently occupied range, 18% or approxi-mately 1.6 million km
2
was classified as areas in which thelong-term survival of jaguars has medium probability.These areas are generally adjacent to high-probability ar-eas and include a large portion of the northern Cerrado,most of the Venezuelan and Colombian Llanos, and thenorthern part of Colombia on the Caribbean coast. InCentral America and Mexico, medium-probability areaswere identified in the highlands of Costa Rica and Pan-ama, southern Mexico, and the two eastern mountainranges of Mexico where jaguars occur, Sierra deTamaulipas and Sierra Madre Oriental.
The remaining parts of the range were classified as ar-eas of low probability for the long-term survival of jag-uars and thus are areas of immediate conservation con-cern. These include the Atlantic tropical moist lowlandforest and Cerrado of Brazil; parts of the Chaco in northernArgentina; the Gran Sabana of northern Brazil, Venezuela,and Guyana; parts of the coastal dry forest in Venezuela;the Central American pine savannas and mangroves alongthe Caribbean coast of Nicaragua and Honduras; parts of
Figure 2. Consensus map of jaguar status across its range.
Conservation BiologyVolume 16, No. 1, February 2002
Sanderson et al. Range-wide Conservation Planning for the Jaguar
65
the Central American tropical moist montane forest ininterior Nicaragua and Honduras and a narrow strip alongthe Pacific Coast of Mexico; and areas in the Mexicanpine-oak forests in Jalisco, Mexico.
The JGRs with the largest proportions of areas (
�
75%,area basis) with a high probability for the long-term sur-vival of the jaguar were almost all in or surrounding theAmazon Basin. Central American tropical moist lowlandforest has the largest proportion of area of high-probabil-ity in Central America and Mexico (71%). In fact, it is theonly JGR north of Colombia with
�
50% of its area cate-gorized as high-probability. Overall, only 12 out of the 36JGRs had
�
10% of their area categorized as high-proba-bility for the long-term survival of the jaguar. Twenty outof 36 had
�
1% of their area so indicated.
JCU Prioritization
Although all JCUs are important areas for jaguars, theyvary in level of threat to jaguars, size, habitat quality, andconnectivity to other JCUs. According to the weightingscheme described above, JCUs were prioritized for eachJGR, with JCUs within a given JGR compared onlyamong one another (Table 2). For example, the highest-priority JCU for the upper Amazon JGR is in and aroundAmazonia National Park, Brazil ( JCU 202). Other JCUswith higher levels of hunting or lower habitat qualityranked slightly lower.
Although all JCUs represent areas with substantial jag-uar populations, a stable prey base, and adequate habi-tat, not all JCUs occur in areas classified as high-probabil-ity for the long-term survival of the jaguar ( Table 2).Eleven JCUs had a majority of their area categorized asmedium-probability for long-term survival, and 10 werecategorized as low-probability for long-term survival. Sixof these 10 JCUs fell entirely within areas of low proba-bility of long-term survival. These JCUs are located innorthern Argentina, central Honduras, the Osa Peninsulaof Costa Rica, and the Atlantic forests of Brazil. TheseJCUs in areas categorized as low-probability for long-term survival contain the most endangered jaguar popu-lations across the range. Fortunately, many other areashave strong jaguar populations and are considered areasof high probability for the long-term survival of the jag-uar. Seventeen JCUs fell entirely within areas catego-rized as high-probability, and 23 had more than 90% oftheir area within high-probability areas.
Discussion
Although the methodology used in this exercise was pio-neered for tigers and draws generally from a host of ex-pert-driven, geographic, priority-setting exercises under-taken over the last decade, it contains a number ofinnovations that advance the methodology of geographic
priority setting, particularly for single-species-based con-servation planning. The most important innovation is alsothe simplest: planning across the complete biologicalrange of the species such that all conservation effortscould be placed in the most important context, the con-text of the species’ biology. Most current species-basedconservation plans are limited first by political bound-aries. For example, habitat conservation plans under theU.S. Endangered Species Act deal mainly with popula-tions within U.S. boundaries. Therefore, the few jaguarssighted recently in the southwestern United States are ac-corded high priority in United States conservation plan-ning, whereas from a range-wide species perspective,other populations in other parts of the range are larger(e.g., Gran Chaco of Bolivia) or more endangered (e.g., At-lantic Forests of Brazil).
Another innovation is that the data sets were nested ina geographic hierarchy that accounted for the differenttypes of knowledge we had about the species (Fig. 3).The most basic distinction separated areas in which wehad knowledge of jaguars (extent of knowledge) from ar-eas in which we lacked knowledge (unknown areas).Given the areas where jaguars are known, the next dis-tinction was to separate areas where jaguars are found(the known, currently occupied range) from those wherethey have been extirpated. Finally, in those areas wherejaguars are found, we identified where the best popula-tions exist ( JCUs) based on clearly defined criteria.
A third innovation is that we retained point observa-tions throughout the analysis, which provided an impor-tant internal check on the more subjective polygonaldata. The point observations provided the only objectiveinformation we had about jaguars: we knew that at eachpoint, at least one jaguar had been observed. Even if wechose to eliminate some types of observation, the pointdatabase enabled us to identify the observation bymethod, date, and observer. Moreover, the analysis canbe updated when significant new data become available.A mechanism is already in place to capture new informa-tion while distributing the compiled information (seewww.savethejaguar.com). Data sharing is a key compo-nent to advancing international conservation efforts.
As a result of retaining the point observations, inde-pendent observers can question the data, providing animportant check against the ever-present concern ofbias in expert-driven systems. For example, severalpoint observations fell in an area outside the extent ofknowledge in southern Brazil, an apparent inconsis-tency ( Fig. 1a & 1d). Review of these observations indi-cated that they were all
�
5 years old and not typical ofthe current situation along the rapidly changing frontierof the southern Amazon Basin.
Another innovation is that we limited conclusions aboutthe currently occupied range to only known areas. Thecustom with range maps prepared previously has been toinclude all “internal” areas if habitat exists there, even if a
66
Range-wide Conservation Planning for the Jaguar Sanderson et al.
Conservation BiologyVolume 16, No. 1, February 2002
Tabl
e 2.
Prio
ritiz
ed r
anki
ng o
f jag
uar
cons
erva
tion
units
(JC
Us)
by ja
guar
geo
grap
hic
regi
on (
JGR)
.
Are
a o
f JC
U in
JG
R(
km
2
)
d
Tota
l JC
U
are
a
(km
2
)
Per
cen
tage
of
are
a r
ate
d
wit
h g
iven
pro
ba
bil
ity
of
jagu
ar
lon
g-te
rm s
urv
iva
lE
xper
t ch
ara
cter
iza
tion
of JC
Us
in te
rms
of fa
ctors
im
port
an
t fo
r th
e lo
ng-
term
su
rviv
al of
jagu
ars
JGR
s
a
an
d J
CU
s
b
Score
c
Ran
kin
g in
JG
Rh
igh
(%)
med
ium
(%
)lo
w(%
)co
nn
ecti
vity
to
oth
er J
CU
s
e
ha
bit
at
qu
ali
tyhu
nti
ng
of
jagu
ar
hu
nti
ng
of
pre
ypopu
lati
on
sta
tus
1.1
Atl
anti
c/T
rop
ical
Mo
ist
Low
lan
d F
ore
stJC
U 2
52: A
tlan
tic
Fore
sts
(Bra
zil)
182
125
,513
30,8
430
010
0in
freq
uen
t dis
per
sal
hig
hm
uch
mu
chd
ecre
asin
gJC
U 2
50: U
pp
er R
io P
aran
á (B
razi
l)18
21
15,2
6917
,731
00
100
infr
eque
nt d
isp
ersa
lh
igh
mu
chm
uch
dec
reas
ing
JCU
257
: Mis
sio
nes
(A
rgen
tin
a, B
razi
l)11
33
26,1
3136
,716
470
53n
o d
isp
ersa
lm
ediu
mm
uch
mu
chd
ecre
asin
g1.
2 U
pp
er A
maz
on
/Tro
pic
al M
ois
t Lo
wla
nd
Fo
rest
JCU
202
: Am
azo
nia
(B
razi
l)29
21
17,9
8038
,414
100
00
freq
uen
t d
isp
ersa
lh
igh
no
ne
no
ne
stab
leJC
U 3
4: M
anu
(P
eru
)25
22
38,1
6243
,387
8812
0fr
equ
ent
dis
per
sal
hig
hso
me
som
est
able
JCU
253
: Mad
idi (
Bo
livia
, Per
u)
252
234
,836
58,8
8074
260
freq
uen
t d
isp
ersa
lh
igh
som
eso
me
stab
leJC
U 7
9: N
oel
Kem
pff
Mer
cad
o (
Bo
livia
)25
22
32,3
1568
,481
810
19fr
equ
ent
dis
per
sal
hig
hso
me
som
est
able
JCU
63:
Am
acay
acu
(C
olo
mb
ia)
252
27,
538
7,53
810
00
0fr
equ
ent
dis
per
sal
hig
hso
me
som
est
able
JCU
207
: Mac
aren
a (C
olo
mb
ia)
214
614
,041
21,0
4167
330
freq
uen
t d
isp
ersa
lm
ediu
mso
me
som
est
able
JCU
79:
No
el K
emp
ff M
erca
do
(B
oliv
ia)
252
232
,315
68,4
8181
019
freq
uen
t d
isp
ersa
lh
igh
som
eso
me
stab
leJC
U 6
3: A
mac
ayac
u (
Co
lom
bia
)25
22
7,53
87,
538
100
00
freq
uen
t d
isp
ersa
lh
igh
som
eso
me
stab
leJC
U 2
07: M
acar
ena
(Co
lom
bia
)21
46
14,0
4121
,041
6733
0fr
equ
ent
dis
per
sal
med
ium
som
eso
me
incr
easi
ng
JCU
62:
Rio
Ap
apo
ris
(Co
lom
bia
)18
27
4,14
54,
145
100
00
freq
uen
t d
isp
ersa
lh
igh
som
em
uch
stab
leJC
U 2
51: P
acay
a Sa
mir
ia (
Per
u)
179
822
,930
22,9
3010
00
0n
o d
ata
hig
hso
me
som
en
o d
ata
JCU
201
: Jav
ari (
Bra
zil)
no
dat
a9
67,5
9867
,598
100
00
no
dat
an
o d
ata
no
dat
an
o d
ata
no
dat
a1.
3 N
ort
hea
st A
maz
on
/Tro
pic
al M
ois
t Lo
wla
nd
Fo
rest
JCU
204
: Am
apá
(Bra
zil,
Fren
ch G
uian
a)25
21
71,7
4671
,754
100
00
freq
uen
t d
isp
ersa
lh
igh
som
eso
me
stab
leJC
U 2
08: V
ich
ada
(Co
lom
bia
)25
21
22,3
9422
,714
991
0fr
equ
ent
dis
per
sal
hig
hso
me
som
est
able
JCU
55:
Up
per
Ori
no
co (
Ven
ezu
ela)
252
118
,932
56,6
6510
00
0fr
equ
ent
dis
per
sal
hig
hso
me
som
est
able
JCU
200
: Rio
Jaú
(B
razi
l)n
o d
ata
438
,162
38,1
6210
00
0n
o d
ata
no
dat
an
o d
ata
no
dat
an
o d
ata
JCU
203
: Pic
o d
a N
eblin
a (B
razi
l)n
o d
ata
411
,687
11,7
5810
00
0n
o d
ata
no
dat
an
o d
ata
no
dat
an
o d
ata
1.4
Sou
thea
st A
maz
on
/Tro
pic
al M
ois
t Lo
wla
nd
Fo
rest
JCU
256
: Xin
gu (
Bra
zil)
292
132
,185
32,1
8510
00
0fr
equ
ent
dis
per
sal
hig
hn
on
en
on
est
able
JCU
202
: Am
azo
nia
(B
razi
l)29
21
20,4
3338
,414
100
00
freq
uen
t d
isp
ersa
lh
igh
no
ne
no
ne
stab
leJC
U 6
7: G
uru
pi (
Bra
zil)
133
320
,211
20,2
1110
00
0n
o d
ata
med
ium
som
eso
me
dec
reas
ing
1.5
Ch
oco
-Dar
ien
/Tro
pic
al M
ois
t Lo
wla
nd
Fo
rest
JCU
206
: Ch
oco-
Dar
ien
(C
olom
bia,
Pa
nam
a)25
21
53,3
8770
,029
936
0fr
equ
ent
dis
per
sal
hig
hso
me
som
est
able
JCU
64:
In
unda
bles
del
baj
o Sa
n J
orge
(C
olom
bia)
146
26,
916
6,91
60
100
0in
freq
uen
t dis
per
sal
med
ium
som
em
uch
dec
reas
ing
JCU
65:
San
Vic
ente
de
Chu
curi
(Col
ombi
a)75
32,
630
2,63
00
100
0in
freq
uen
t dis
per
sal
poo
rso
me
muc
hin
crea
sin
g
1.6
Cen
tral
Am
eric
an/T
rop
ical
Moi
st L
owla
nd
Fore
stJC
U 1
55: S
elva
May
a (M
exic
o,
Gu
atem
ala,
Bel
ize)
252
162
,593
63,5
5010
00
0fr
equ
ent
dis
per
sal
hig
hso
me
som
est
able
JCU
153
: So
uth
ern
Bel
ize
(Bel
ize)
252
18,
425
8,59
099
01
freq
uen
t d
isp
ersa
lh
igh
som
eso
me
stab
leJC
U 2
06: C
ho
co-D
arie
n (
Co
lom
bia
, P
anam
a)25
21
7,13
070
,029
936
0fr
equ
ent
dis
per
sal
hig
hso
me
som
est
able
con
tin
ued
Conservation BiologyVolume 16, No. 1, February 2002
Sanderson et al. Range-wide Conservation Planning for the Jaguar
67
Tabl
e 2.
(con
tinue
d)
Are
a o
f JC
U in
JG
R(
km
2
)
d
Tota
l JC
U
are
a
(km
2
)
Per
cen
tage
of
are
a r
ate
d
wit
h g
iven
pro
ba
bil
ity
of
jagu
ar
lon
g-te
rm s
urv
iva
lE
xper
t ch
ara
cter
iza
tion
of JC
Us
in ter
ms
of fa
ctors
im
port
an
t fo
r th
e lo
ng-
term
su
rviv
al of
jagu
ars
JGR
s
a
an
d J
CU
s
b
Score
c
Ran
kin
g in
JG
Rh
igh
(%)
med
ium
(%
)lo
w(%
)co
nn
ecti
vity
to
oth
er J
CU
s
e
ha
bit
at
qu
ali
tyhu
nti
ng
of
jagu
ar
hu
nti
ng
of
pre
ypopu
lati
on
sta
tus
JCU
259
: Lag
o I
zab
al (
Gu
atem
ala)
248
43,
894
5,35
395
05
freq
uen
t d
isp
ersa
lh
igh
som
eso
me
no
dat
aJC
U 1
50: T
alam
anca
(C
ost
a R
ica,
P
anam
a)21
45
6,12
514
,099
3157
12in
freq
uen
t dis
per
sal
hig
hso
me
som
ein
crea
sin
gJC
U 1
51: O
sa P
enis
ula
(C
ost
a R
ica)
182
61,
591
1,80
90
100
0fr
equ
ent
dis
per
sal
hig
hso
me
mu
chst
able
JCU
154
: Gu
anac
aste
(C
ost
a R
ica)
168
72,
017
5,32
394
60
infr
eque
nt d
isp
ersa
lm
ediu
mso
me
som
ein
crea
sin
gJC
U 1
3: I
stm
o d
e T
ehu
ante
pec
(M
exic
o)
160
89,
439
9,57
910
00
0in
freq
uen
t dis
per
sal
med
ium
som
eso
me
stab
leJC
U 7
6: L
a M
osq
uit
ia (
Ho
nd
ura
s)n
o d
ata
19,6
7930
,389
630
36n
o d
ata
no
dat
an
o d
ata
no
dat
an
o d
ata
no
dat
aJC
U 7
8: C
ord
iller
a N
om
bre
de
Dio
s (H
on
du
ras)
no
dat
a9
2,26
92,
834
800
20n
o d
ata
no
dat
an
o d
ata
no
dat
an
o d
ata
JCU
77:
Mo
nta
ña
de
Yo
ro (
Ho
nd
ura
s)n
o d
ata
975
72,
214
00
100
no
dat
an
o d
ata
no
dat
an
o d
ata
no
dat
a2.
1 T
rop
ical
An
des
/Tro
pic
al M
ois
t Lo
wla
nd
Fo
rest
JCU
253
: Mad
idi (
Bo
livia
, Per
u)
252
115
,436
58,8
8074
260
freq
uen
t d
isp
ersa
lh
igh
som
eso
me
stab
leJC
U 2
1: B
arit
u-C
alile
gua
(Arg
enti
na,
B
oliv
ia)
252
111
,654
12,5
720
010
0fr
equ
ent
dis
per
sal
hig
hso
me
som
est
able
JCU
34:
Man
u (
Per
u)
252
15,
227
43,3
8788
120
freq
uen
t d
isp
ersa
lh
igh
som
eso
me
stab
le2.
2 C
entr
al A
mer
ican
/Tro
pic
al M
ois
t M
on
tan
e Fo
rest
JCU
206
: Ch
oco
-Dar
ien
(C
olo
mb
ia,
Pan
ama)
252
12,
756
70,0
2993
60
freq
uen
t d
isp
ersa
lh
igh
som
eso
me
stab
leJC
U 2
59: L
ago
Iza
bal
(G
uat
emal
a)24
82
1,16
65,
353
950
5fr
equ
ent
dis
per
sal
hig
hso
me
som
en
o d
ata
JCU
150
: Tal
aman
ca (
Co
sta
Ric
a,
Pan
ama)
214
37,
941
14,0
9931
5712
infr
eque
nt d
isp
ersa
lh
igh
som
eso
me
incr
easi
ng
JCU
76:
La
Mo
squ
itia
(H
on
du
ras)
no
dat
a4
1,70
430
,389
630
36n
o d
ata
no
dat
an
o d
ata
no
dat
an
o d
ata
JCU
77:
Mo
nta
ña
de
Yo
ro (
Ho
nd
ura
s)n
o d
ata
41,
456
2,83
480
020
no
dat
an
o d
ata
no
dat
an
o d
ata
no
dat
aJC
U 7
8: C
ord
iller
a N
om
bre
de
Dio
s (H
on
du
ras)
no
dat
a4
566
2,21
40
010
0n
o d
ata
no
dat
an
o d
ata
no
dat
an
o d
ata
2.4
Ven
ezu
elan
Co
asta
l Mo
nta
ne
Fore
st/T
rop
ical
Mo
ist
Mo
nta
ne
Fore
stJC
U 5
4: G
uat
op
o (
Ven
ezu
ela)
132
11,
324
1,62
30
100
0in
freq
uen
t dis
per
sal
hig
hso
me
mu
chd
ecre
asin
g
2.5
Gu
ayan
a M
on
tan
e Fo
rest
/Tro
pic
al M
ois
t M
on
tan
e Fo
rest
JCU
55:
Up
per
Ori
no
co (
Ven
ezu
ela)
252
137
,733
56,6
6510
00
0fr
equ
ent
dis
per
sal
hig
hso
me
som
est
able
JCU
56:
Cau
ra (
Ven
ezu
ela)
252
118
,736
18,7
3610
00
0fr
equ
ent
dis
per
sal
hig
hso
me
som
est
able
JCU
205
: Ser
ra d
a Es
tru
tura
(B
razi
l)n
o d
ata
32,
819
2,81
910
00
0n
o d
ata
hig
hn
o d
ata
no
dat
an
o d
ata
3.1
No
rth
So
uth
Am
eric
an/T
rop
ical
Dry
Fo
rest
JCU
207
: Mac
aren
a (C
olo
mb
ia)
214
14,
074
21,0
4167
330
freq
uen
t d
isp
ersa
lm
ediu
mso
me
som
ein
crea
sin
gJC
U 5
2: C
hir
iqu
are
(Ven
ezu
ela)
182
288
497
70
1090
freq
uen
t d
isp
ersa
lh
igh
som
em
uch
stab
leJC
U 5
3: G
uar
itic
o (
Ven
ezu
ela)
533
771
1,42
80
4654
no
dat
am
ediu
mm
uch
mu
chd
ecre
asin
g3.
2 W
este
rn A
nd
es/T
rop
ical
Dry
Fo
rest
no
ne
——
——
——
——
——
——
con
tin
ued
68
Range-wide Conservation Planning for the Jaguar Sanderson et al.
Conservation BiologyVolume 16, No. 1, February 2002
Tabl
e 2.
(con
tinue
d)
Are
a o
f JC
U in
JG
R(
km
2
)
d
Tota
l JC
U
are
a
(km
2
)
Per
cen
tage
of
are
a r
ate
d
wit
h g
iven
pro
ba
bil
ity
of
jagu
ar
lon
g-te
rm s
urv
iva
lE
xper
t ch
ara
cter
iza
tion
of JC
Us
in ter
ms
of fa
ctors
im
port
an
t fo
r th
e lo
ng-
term
su
rviv
al of
jagu
ars
JGR
s
a
an
d J
CU
s
b
Score
c
Ran
kin
g in
JG
Rh
igh
(%)
med
ium
(%
)lo
w(%
)co
nn
ecti
vity
to
oth
er J
CU
s
e
ha
bit
at
qu
ali
tyhu
nti
ng
of
jagu
ar
hu
nti
ng
of
pre
ypopu
lati
on
sta
tus
3.3
Ch
aco
/Tro
pic
al D
ry F
ore
stJC
U 8
0: G
ran
Ch
aco
(B
oliv
ia, P
arag
uay
)25
21
85,7
0689
,116
100
00
freq
uen
t d
isp
ersa
lh
igh
som
eso
me
stab
leJC
U 2
4: C
hac
o (
Arg
enti
na)
192
27,
153
7,15
30
010
0in
freq
uen
t dis
per
sal
hig
hso
me
mu
chd
ecre
asin
g3.
4 C
entr
al A
mer
ican
/Tro
pic
al D
ry F
ores
tJC
U 1
54: G
uan
acas
te (
Co
sta
Ric
a)16
81
2,93
55,
323
946
0in
freq
uen
t dis
per
sal
med
ium
som
eso
me
incr
easi
ng
3.5
Mex
ican
/Tro
pic
al D
ry F
ore
stJC
U 1
01: J
alis
co (
Mex
ico
)23
81
29,4
0929
,409
615
35fr
equ
ent
dis
per
sal
hig
hso
me
mu
chd
ecre
asin
gJC
U 1
00: S
on
ora
(M
exic
o)
140
22,
020
13,8
5910
00
0in
freq
uen
t dis
per
sal
med
ium
mu
chm
uch
stab
le3.
6 C
erra
do
/Tro
pic
al D
ry F
ore
stJC
U 6
8: C
hap
ada
das
Man
gabe
iras
(B
razi
l)25
21
45,3
9745
,397
010
00
freq
uen
t d
isp
ersa
lh
igh
som
eso
me
stab
leJC
U 7
9: N
oel
Kem
pff
Mer
cad
o (
Bo
livia
)25
21
36,1
0768
,481
810
19fr
equ
ent
dis
per
sal
hig
hso
me
som
est
able
JCU
253
: Mad
idi (
Bo
livia
, Per
u)
252
18,
607
30,8
430
010
0fr
equ
ent
dis
per
sal
hig
hso
me
som
est
able
JCU
258
: Pan
tan
al (
Bra
zil,
Bo
livia
)23
84
9,10
187
,034
960
4fr
equ
ent
dis
per
sal
hig
hm
uch
som
ed
ecre
asin
gJC
U 6
9: M
aran
ho
(B
razi
l)20
25
6,25
36,
253
010
00
freq
uen
t d
isp
ersa
lm
ediu
mso
me
som
ed
ecre
asin
gJC
U 8
: Ara
guai
a (B
razi
l)19
66
29,7
9629
,846
010
00
freq
uen
t d
isp
ersa
lm
ediu
mso
me
mu
chst
able
JCU
250
: Up
per
Rio
Par
aná
(Bra
zil)
182
72,
462
17,7
310
010
0in
freq
uen
t dis
per
sal
hig
hm
uch
mu
chd
ecre
asin
gJC
U 7
: Ch
apad
a d
os
Vea
dei
ros
(Bra
zil)
156
810
,281
10,2
810
100
0in
freq
uen
t dis
per
sal
med
ium
som
eso
me
dec
reas
ing
JCU
255
: Ser
ra d
a C
apiv
ara
(Bra
zil)
146
92,
082
7,19
30
1387
infr
eque
nt d
isp
ersa
lm
ediu
mso
me
mu
chd
ecre
asin
g4.
1 M
exic
an/X
eric
sJC
U 1
00: S
on
ora
(M
exic
o)
140
110
,470
13,8
5910
00
0in
freq
uen
t dis
per
sal
med
ium
mu
chm
uch
stab
le4.
2 C
arib
bea
n/X
eric
sJC
U 2
09: L
lan
os
(Ven
ezu
ela)
196
11,
491
7,12
50
100
0fr
equ
ent
dis
per
sal
med
ium
mu
chso
me
stab
le4.
3 C
aati
nga
/Xer
ics
JCU
255
: Ser
ra d
a C
apiv
ara
(Bra
zil)
146
15,
111
7,19
30
1387
infr
eque
nt d
isp
ersa
lm
ediu
mso
me
mu
chd
ecre
asin
g4.
6 A
rgen
tin
e M
on
te/X
eric
sn
on
e—
——
——
——
——
——
—5.
1 C
entr
al A
mer
ican
Pin
e Sa
van
nas
/Her
bac
eou
s Lo
wla
nd
Gra
ssla
nd
JCU
76:
La
Mo
squ
itia
(H
on
du
ras)
no
dat
a1
6,63
330
,389
630
36n
o d
ata
no
dat
an
o d
ata
no
dat
an
o d
ata
5.2
Llan
os
- Gra
n S
aban
a/H
erb
aceo
us
Low
lan
d G
rass
lan
dJC
U 2
09: L
lan
os
(Ven
ezu
ela)
196
15,
633
7,12
50
100
0fr
equ
ent
dis
per
sal
med
ium
mu
chso
me
stab
leJC
U 5
3: G
uar
itic
o (
Ven
ezu
ela)
532
657
1,42
80
4654
no
dat
am
ediu
mm
uch
mu
chd
ecre
asin
g5.
3 Pa
mp
as/H
erba
ceou
s Lo
wla
nd
Gra
ssla
nd
JCU
257
: Mis
sio
nes
(A
rgen
tin
a, B
razi
l)11
31
1836
,716
470
53n
o d
isp
ersa
lm
ediu
mm
uch
mu
chd
ecre
asin
g5.
5 A
maz
on
ian
Sav
ann
a/H
erb
aceo
us
Low
lan
d G
rass
lan
dn
on
e—
——
——
——
——
——
—5.
6 P
anta
nal
/Her
bac
eou
s Lo
wla
nd
Gra
ssla
nd
JCU
80:
Gra
n C
hac
o (
Bo
livia
, Par
agu
ay)
252
13,
410
89,1
1610
00
0fr
equ
ent
dis
per
sal
hig
hso
me
som
est
able
JCU
258
: Pan
tan
al (
Bra
zil,
Bo
livia
)23
82
77,9
3287
,034
960
4fr
equ
ent
dis
per
sal
hig
hm
uch
som
ed
ecre
asin
g
con
tin
ued
Conservation BiologyVolume 16, No. 1, February 2002
Sanderson et al. Range-wide Conservation Planning for the Jaguar
69
Tabl
e 2.
(con
tinue
d)
Are
a o
f JC
U in
JG
R(
km
2
)
d
Tota
l JC
U
are
a
(km
2
)
Per
cen
tage
of
are
a r
ate
d
wit
h g
iven
pro
ba
bil
ity
of
jagu
ar
lon
g-te
rm s
urv
iva
lE
xper
t ch
ara
cter
iza
tion
of JC
Us
in ter
ms
of fa
ctors
im
port
an
t fo
r th
e lo
ng-
term
su
rviv
al of
jagu
ars
JGR
s
a
an
d J
CU
s
b
Score
c
Ran
kin
g in
JG
Rh
igh
(%)
med
ium
(%
)lo
w(%
)co
nn
ecti
vity
to
oth
er J
CU
s
eh
abit
at
qu
ali
tyhu
nti
ng
of
jagu
ar
hu
nti
ng
of
pre
ypopu
lati
on
sta
tus
5.7
Wes
tern
Gu
lf C
oas
tal G
rass
lan
ds/
Her
bac
eou
s Lo
wla
nd
Gra
ssla
nd
no
ne
——
——
——
——
——
——
6.1
Para
mo/
Her
bace
ous
Mon
tan
e G
rass
lan
dJC
U 2
06: C
hoc
o-D
arie
n (
Col
ombi
a,
Pan
ama)
252
15
70,0
2993
60
freq
uen
t d
isp
ersa
lh
igh
som
eso
me
stab
le6.
2 P
un
a/H
erb
aceo
us
Mo
nta
ne
Gra
ssla
nd
JCU
21:
Bar
itu-
Cal
ilegu
a (A
rgen
tin
a,
Bol
ivia
)25
21
911
12,5
720
010
0fr
equ
ent
dis
per
sal
hig
hso
me
som
est
able
6.4
Pan
tep
ui/
Her
bac
eou
s M
on
tan
e G
rass
lan
dn
on
e—
——
——
——
——
——
—7.
2 B
razi
llian
Ara
uca
ria/
Tem
per
ate
Fore
stJC
U 2
52: A
tlan
tic
Fore
st (
Bra
zil)
182
13,
060
30,8
430
010
0in
freq
uen
t dis
per
sal
hig
hm
uch
mu
chd
ecre
asin
gJC
U 2
57: M
issi
on
es (
Arg
enti
na,
Bra
zil)
113
210
,567
36,7
1647
053
no
dis
per
sal
med
ium
mu
chm
uch
dec
reas
ing
7.3
Mex
ican
Pin
e-O
ak/T
emp
erat
e Fo
rest
JCU
101
: Jal
isco
(M
exic
o)
238
110
,150
29,4
0961
535
freq
uen
t d
isp
ersa
lh
igh
som
em
uch
dec
reas
ing
JCU
100
: So
no
ra (
Mex
ico
)14
02
1,36
913
,859
100
00
infr
eque
nt d
isp
ersa
lm
ediu
mm
uch
mu
chst
able
JCU
12:
Sie
rra
Mad
re O
rien
tal (
Mex
ico
)13
33
21,6
1821
,618
010
00
infr
eque
nt d
isp
ersa
lp
oo
rso
me
som
ed
ecre
asin
gJC
U 1
02: S
ierr
a T
amau
lipas
(M
exic
o)
904
1,38
71,
387
010
00
infr
eque
nt d
isp
ersa
lm
ediu
mso
me
mu
chst
able
8.1
No
rth
ern
Mex
ico
/Man
gro
veJC
U 1
01: J
alis
co (
Mex
ico
)23
81
1,43
029
,409
615
35fr
equ
ent
dis
per
sal
hig
hso
me
mu
chd
ecre
asin
g8.
2 C
entr
al A
mer
ican
/Man
gro
veJC
U 2
06: C
ho
co-D
arie
n (
Co
lom
bia
, P
anam
a)25
21
660
70,0
2993
60
freq
uen
t d
isp
ersa
lh
igh
som
eso
me
stab
leJC
U 7
6: L
a M
osq
uit
ia (
Ho
nd
ura
s)n
o d
ata
21,
919
30,3
8963
036
no
dat
an
o d
ata
no
dat
an
o d
ata
no
dat
a8.
3 N
ort
her
n S
ou
th A
mer
ica/
Man
gro
veJC
U 2
06: C
ho
co-D
arie
n (
Co
lom
bia
, P
anam
a)25
21
772
70,0
2993
60
freq
uen
t d
isp
ersa
lh
igh
som
eso
me
stab
le8.
4 A
maz
on
ia/M
angr
ove
no
ne
——
——
——
——
——
——
8.5
East
ern
So
uth
Am
eric
a/M
angr
ove
JCU
252
: Atl
anti
c Fo
rest
(B
razi
l)18
21
1,88
730
,843
00
100
infr
eque
nt d
isp
ersa
lh
igh
mu
chm
uch
dec
reas
ing
aTh
e JG
Rs
are
na
med
by
regi
on
an
d h
abit
at
type.
Nu
mer
ic c
odes
are
in
Fig
. 1, a
nd t
he
geogr
aph
ic i
nfo
rma
tion
sys
tem
(G
SI)
da
ta s
ets
dis
trib
ute
d t
hro
ugh
ww
w.s
ave
thej
agu
ar.
com
.bTh
e JC
Us
are
na
med
usi
ng
enco
mpa
ssin
g or
adjo
inin
g ge
ogr
aph
ic f
eatu
res,
in
clu
din
g pro
tect
ed a
rea
s, r
iver
s, m
ou
nta
in r
an
ges,
an
d/o
r st
ate
s a
nd p
rovi
nce
s (
Fig.
1).
Th
e n
um
eric
codes
ref
erto
in
dic
es i
n t
he
GIS
da
ta s
ets
dis
trib
ute
d t
hro
ugh
ww
w.s
ave
thej
agu
ar.
com
.c Fi
na
l pri
ori
tiza
tion
sco
re f
or
JCU
in
giv
en J
GR
. Hig
her
va
lues
in
dic
ate
bet
ter
con
dit
ion
s fo
r lo
ng-
term
su
rviv
al of
jagu
ar.
dA
rea
mea
sure
d i
n e
qu
al a
rea
azi
mu
tha
l pro
ject
ion
, cen
tra
l m
erid
ian
-72
, ref
eren
ce la
titu
de
0.
e Con
nec
tivi
ty i
nte
rpre
ted i
n t
erm
s of
freq
uen
cy o
f dis
per
sal to
or
from
th
is J
CU
.
70 Range-wide Conservation Planning for the Jaguar Sanderson et al.
Conservation BiologyVolume 16, No. 1, February 2002
species’ status in those areas is unknown. We believe it isimportant to clearly distinguish between what is knownand what is unknown, because conservation choices de-pend on the state of knowledge. As a result, our range maphas several large “holes” because it represents only theknown, currently occupied range ( Fig. 1b).
Finally this exercise, like the preceding effort for thetiger ( Wikramanayake et al 1998), differs from otherspecies-based planning mechanisms because it is basedon an ecogeographic framework for setting priorities.The goal is not to determine the most important site forjaguar conservation overall, or the most important site ina given country, but rather to find the most importantsites for ecologically distinct populations of jaguars. Be-cause information is insufficient to define these ecologi-cal distinctions a priori, we used a geographic proxy, thejaguar geographic regions, which provided the frame-work over which the data were summarized (Table 1)and JCUs prioritized (Table 2).
All these techniques are designed to limit errors due tothe subjective nature of expert-based priority-setting sys-tems by closely tracking how well certain facts are known,where extrapolations are made, and where knowledge islacking. Moreover, these data are part of framework thatprovides to any future user the ability to reanalyze the re-sults and draw his or her own conclusions.
Whether or not the end results are “scientific” in a for-mal sense seems less important than acknowledging thelimits of what we can do when planning for speciessuch as the jaguar. The jaguar historically ranged over19.1 million km2, an area over twice that of the UnitedStates including Alaska. Within that area there are per-haps 100 professionals working to study and conservethe jaguar; of them, 35 contributed information to thiswork. Those experts speak three different languages andcome from 18 different countries. Through this exer-cise, they established a common data framework onwhich they all agreed and a broad consensus on priori-
ties for the species. The result is necessarily extensive ingeographic scope and lacking in intensive detail, but itreflects the shared state of knowledge of jaguar status,distribution, and geographic priorities.
Finally, range-wide, species-based conservation plan-ning for the jaguar, or any other broadly distributed spe-cies, complements other coarse-filter approaches to con-servation planning by testing their generality through anemphasis on single-species requirements. In this caseunfortunately, conserving supra-organismal entities suchas hotspots or ecoregions provides no guarantee of con-serving jaguars across all the ecological settings wherethey occur, because important locations for jaguars oc-cur outside hotspots and across a large number of ecore-gions (�100). Moreover, priorities determined for thejaguar alone may differ considerably from priorities de-termined through other mechanisms focused on overallbiodiversity conservation. As a result, conservation ef-forts in the Atlantic forests of Brazil might prove an inef-ficient investment for conservation of the jaguar, giventhe range of needs and opportunities for jaguar conser-vation elsewhere. No planning tool meets all goals, butdifferent planning tools can and should complement andenhance one another. With the jaguar exercise, we pro-vide a model to reintroduce species to coarse-filter, in-ternational conservation planning efforts.
Acknowledgments
We thank H. Hung, G. Raygordetsky, K. Willett, and G.Woolmer for their assistance with data entry and geo-graphic information system analysis and L. Pavajeau forSpanish translation. D. Olson, R. Sayre, and M. Grigioneprovided helpful advice for design of the priority-settingmethodology. The editors and two anonymous review-ers also provided helpful critical reviews of an early draftof this manuscript. This work was funded by grants from
Figure 3. Conceptual framework of geographic data sets for range-wide species conservation planning.
Conservation BiologyVolume 16, No. 1, February 2002
Sanderson et al. Range-wide Conservation Planning for the Jaguar 71
Jaguar Cars–US, the Prospect Hill Foundation, and soft-ware donations from the Environmental Systems ResearchInstitute (ESRI).
Literature Cited
Biodiversity Support Program, Conservation International, The NatureConservancy, Wildlife Conservation Society, World Resources Insti-tute, and World Wildlife Fund. 1995. A regional analysis of geographicpriorities for biodiversity conservation in Latin America and the Carib-bean. U.S. Agency for International Development, Washington, D.C.
Boyce, M. S., and A. Haney. 1997. Ecosystem management: applicationsfor sustainable forest and wildlife resources. Yale University Press,New Haven, Connecticut.
Callicott, J. B. 1990. Whither conservation ethics? Conservation Biology4:15–20.
Dinerstein, E., D. M. Olson, D. J. Graham, A. L. Webster, S. A. Primm,M. P. Bookbinder, and G. Ledec. 1995. A conservation assessmentof the terrestrial ecoregions of Latin America and the Caribbean.World Wildlife Fund and The World Bank, Washington, D.C.
Ehrlich, P. R., and A. H. Ehrlich. 1981. Extinction: the causes and conse-quences of the disappearance of species. Random House, New York.
Franklin, J. F. 1993. Preserving biodiversity: species, ecosystems orlandscapes? Ecological Applications 3:202–205.
Hall, L. S., P. R. Krausman, and M. L. Morrison. 1997. Importance ofstandardized terminology in habitat evaluation. Wildlife SocietyBulletin 25:761–762.
Hunter, M. L. 1991. Coping with ignorance: the coarse-filter strategy formaintaining biodiversity. Pages 266–281 in K. A. Kohm, editor. Bal-ancing on the brink of extinction. Island Press, Washington, D.C.
Hunter, M. L., and A. Hutchinson. 1994. The virtues and shortcomingsof parochialism: conserving species that are locally rare, but glo-bally common. Conservation Biology 8:1163–1165.
Lioutty, A. A. 1996. ArcAtlas: our Earth (CDROM). Environmental Sys-tems Research Institute and DATA+, Redlands, California.
Mann, C. C., and M. L. Plummer. 1995. Noah’s choice: the future of en-dangered species. Knopf, New York.
Medellin, R. A., C. Chetkiewicz, A. Rabinowitz, K. H. Redford, J. G. Robin-son, E. W. Sanderson, and A. Taber. 2001. El Jaguar en el nuevo mile-
nio: una evaluacion de su estado, deteccion de prioridades y recomen-daciones para la conservacion de los jaguares en America. UniversidadNacional Autonoma de Mexico and Wildlife Conservation Society,México, D.F.
Myers, N., R. A. Mittermeier, C. G. Mittermeier, G. A. B. da Fonseca,and J. Kent. 2000. Biodiversity hotspots for conservation priorities.Nature 403:853–858.
Noss, R. F. 1991. From endangered species to biodiversity. Pages 227–246 in K. A. Kohm, editor. Balancing on the brink of extinction. Is-land Press, Washington, D.C.
Olson, D. M., and E. Dinerstein. 1998. The global 200: a representationapproach to conserving the Earth’s most biologically valuable re-gions. Conservation Biology 12:502–515.
Quigley, H. B., and P. G. Crawshaw. 1992. A conservation plan for thejaguar Panthera onca in the Pantanal region of Brazil. BiologicalConservation 61:149–157.
Rabinowitz, A. R. 1986. Jaguar: one man’s battle to establish the world’sfirst jaguar reserve. Anchor Books, New York.
Redford, K. H., and B. D. Richter. 1999. Conservation of biodiversity ina world of use. Conservation Biology 13:1246–1256.
Salwasser, H. 1991. In search of an ecosystem approach to endangeredspecies conservation. Pages 228–265 in K. A. Kohm, editor. Balanc-ing on the brink of extinction. Island Press, Washington, D.C.
Seymour, K. L. 1989. Panthera onca. Mammalian Species 340.Slattersfield, A. J., M. J. Crosby, A. J. Long, and D. C. Wege. 1998. En-
demic bird areas of the world: priorities for biodiversity conserva-tion. BirdLife International, Cambridge, United Kingdom.
Soulé, M. E., and M. Terborgh. 1999. Continental conservation: scien-tific foundations of regional reserve networks. Island Press, Wash-ington, D.C.
Weber, W., and A. Rabinowitz. 1996. A global perspective on large car-nivore conservation. Conservation Biology 10:1046–1054.
Wikramanayake, E. D., E. Dinerstein, J. G. Robinson, U. Karanth, A.Rabinowitz, D. Olson, T. Mathew, P. Hedao, M. Conner, G. Hem-ley, and D. Bolze. 1998. An ecology-based method for defining pri-orities for large mammal conservation: the tiger as case study. Con-servation Biology 12:865–878.
Zink, R. M., G. F. Barrowclough, J. L. Atwood, and R. C. Blackwell-Rago.2000. Genetics, taxonomy, and conservation of the threatened Cali-fornia gnatcatcher. Conservation Biology 14::1394–1405.
72 Range-wide Conservation Planning for the Jaguar Sanderson et al.
Conservation BiologyVolume 16, No. 1, February 2002
AppendixExpert contributors to the workshop on “Jaguars in the New Millennium” (March 1999).
Region Expert contributor Institution
Northern Mexico Marcelo Aranda Instituto de Ecología, A.C., Veracruz, MéxicoGerardo Ceballos Universidad Nacional Autónoma de MéxicoCarlos A. López González Northern Rockies Conservation CooperativeRodrigo Medellín Universidad Nacional Autónoma de MéxicoBrian Miller Denver Zoological FoundationBill Van Pelt Arizona Game and Fish Department
Central America Eduardo Carrillo Centro Agronómico Tropical de Investigación y Enseñanza (CATIE)Sharon Matola The Belize ZooRoan McNab Wildlife Conservation SocietyCarolyn Miller Wildlife Conservation SocietyJohn Polisar Florida Museum of Natural History, University of FloridaAlan Rabinowitz Wildlife Conservation SocietyHoward Quigley The Hornocker Wildlife Research InstituteJoel Saenz Universidad Nacional, Costa Rica, HerediaChristopher Vaughan Universidad Nacional, Costa Rica, Heredia
Northern South America Ernesto Boede Centro Veterinario “Los Colorados,” VenezuelaLouise Emmons Division of Mammals, Smithsonian InstitutionAntonio González-Fernández Universidad Nacional Experimental de los Llanos
Occidentales “Ezequiel Zamora”Tadeu Gomes de Oliveira Maranho State University/Pró-Carnívoros AssociationRafael Hoogesteijn Gerente Hatos de Apur, Prohesa y Grupo de Especialistas en FelinosEdgardo Mondolfi Insituto de Zoología Tropical, Universidad Central de VenezuelaJohn Robinson Wildlife Conservation SocietyLeandro Silveira Pró-Carnívoros Association/Universidade Federal de GoiásMelvin Sunquist University of FloridaGerardo Zuloaga Villamizar Instituto de Ciencias Naturales, Universidad Nactional de Colombia
Southern South America Peter Crawshaw Jr. Centro Nacional de Pesquisa para a Conservaço de Predadores Naturais(CENAP/IBAMA)
Julio Dalponte Universidade de BrasíliaWarren Johnson Laboratory of Genomic Diversity, National Cancer InstituteAlicia Kuroiwa Tambopata Research Centre, Rainforest ExpeditionsMaria Renata Pereira Leite Duke UniversityRonaldo Morato Pró-Carnívoros Association/University of So PauloPablo Perovic Instituto de Biologia de la Altura, Universidad Nacional de JujuyKarina Schiaffino Centro de Investigaciones Ecológicas Subtropicales, Administración de
Parques Nacionales, ArgentinaDaniel Scognamillo University of FloridaAndrew Taber Wildlife Conservation Society