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A Spatial Model of Potential Jaguar Habitat in Author(s): James R. Hatten, Annalaura Averill-Murray, William E. Van Pelt Source: The Journal of Wildlife Management, Vol. 69, No. 3 (Jul., 2005), pp. 1024-1033 Published by: Allen Press Stable URL: http://www.jstor.org/stable/3803341 Accessed: 19/09/2009 18:33

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http://www.jstor.org A SPATIALMODEL OF POTENTIALJAGUAR HABITAT IN ARIZONA

JAMESR. HATTEN,1'2Arizona Game and Fish Department,Wildlife Management Division, Nongame Branch, 2221 West Green- way Road, Phoenix,AZ 85023, USA ANNALAURAAVERILL-MURRAY, Arizona Game and Fish Department,Region V, FieldOperations Division, Tucson, AZ, 85710, USA WILLIAME. VANPELT, Arizona Game and Fish Department,Wildlife Management Division, Nongame Branch, 2221 West Green- way Road, Phoenix,AZ 85023, USA

Abstract:The jaguar (Pantheraonca) is an endangered species that occasionally visits the southwestern United States from Mexico. The number ofjaguar sightings per decade has declined over the last 100 years in Arizona, USA, rais- ing conservation concerns for the species at a local and national level. In 1997, state, federal, and local govern- ments with land-management responsibilities agreed to characterize and identify potential jaguar habitat in Ari- zona and New Mexico. Specifically, the objectives of our analysis were 2-fold: (1) characterize potential jaguar habitat in Arizona from historic sighting records and (2) create a statewide habitat suitability map. We used a Geo- graphic Information System (GIS) to characterize potential jaguar habitat by overlaying historic jaguar sightings (25) on landscape and habitat features believed important (e.g., vegetation biomes and series, elevation, terrain ruggedness, proximity to perennial or intermittent water sources, human density). The amount of Arizona (%) identified as potential jaguar habitat ranged from 21% to 30% depending on the input variables. Mostjaguar sight- ings were in scrub grasslands between 1,220 and 1,829-m elevation in southeastern Arizona, in intermediately to extremely rugged terrain, and within 10 km of a water source. Conservation efforts should focus on protecting the most suitable jaguar habitat in southeastern Arizona (i.e., Santa Cruz, Pima, Cochise, Pinal, Graham counties), travel corridors within and outside Arizona, and jaguar habitat in the Sierra Madres of Sonora, Mexico. JOURNALOF WILDLIFEMANAGEMENT 69(3):1024-1033; 2005

Keywords: Arizona, Pantheraonca, conservation, Geographic Information System, GIS, habitat selection, landscape, neighborhood analysis, predictive modeling, jaguar.

Jaguars have been noted for their adaptability to In the southwestern United States, potential prey a wide variety of environmental conditions (Rabi- species include collared peccary (Tayassutajacu), nowitz and Nottingham 1986, Seymour 1989). white tailed deer (Odocoileus virginianus), mule Habitat studies in the core part ofjaguar range in- deer (Odocoileushemionus), coatis (Nasua nasua), dicate a close association with water, dense cover skunk (Mephitis spp., Spilogalegracilis), raccoon (Schaller and Crawshaw1980, Quigley and Craw- (Procyonlotor), jack rabbit (Lepus spp.), domestic shaw 1992), sufficient prey (Seymour 1989, Swank livestock, and horses. and Teer 1989), and an avoidance of highly dis- The southwestern United States and Sonora, turbed areas (Quigley and Crawshaw 1992). Mexico are at the extreme northern limits of the Jaguars have been found from sea level to 3,800 m jaguar's range that extends primarily from north- (Tewes and Schmidly 1987) but rarely over 1,000 m ern Argentina to central Mexico (Swank and Teer (Seymour 1989). They have also been found in a 1989, Sanderson et al. 2002). Over the last century, wide variety of vegetation communities from trop- the jaguar's range has been reduced to approxi- ical rainforest and flooded grassland mosaics to mately 46% of its historic (pre-1900) range due to Madrean evergreen woodland, semi-desert grass- hunting pressure and habitat loss, especially in the lands, and wooded canyons of the Sonoran southern United States, northern Mexico, north- Desert (Rabinowitz 1999, Brown and Lopez Gon- ern Brazil, and southern Argentina (Swank and zales 2001 ). Teer 1989, Sanderson et al. 2002). Jaguars have Jaguars are opportunistic feeders with more been documented occasionally in the southwest- than 85 prey species reported in their diet (Sey- ern United States since 1900, but the number of mour 1989), including mammals, reptiles, and sightings per decade has declined over the last 100 birds. Primary prey items are diurnal, terrestrial years (Brown 1983). In 1996, there were 2 jaguars mammals with a body mass >1 kg (Seymour 1989). photographed in the mountains of southeastern Arizona (Glenn 1996, Childs 1998), raising con- servation concerns for the species at local and na- tional levels and the 1 Present address: USGS, 5501 A Cook-Underwood prompting federal govern- Road, Cook, WA 98605-9717, USA. ment to list the jaguar as endangered (Federal 2 Corresponding author e-mail:[email protected] Register 1997). 1024 J. Wildl.Manage. 69(3):2005 CHARACTERIZINGPOTENTIAL JAGUAR HABITAT * Hatten et al. 1025

Representatives of local, state, and federal gov- jaguars occupied an area before capture or sight- ernments with land-management responsibilities ing, making it difficult for us to distinguish a travel in Arizona and New Mexico signed a Memoran- corridor from actual jaguar habitat. dum of Agreement (MOA) to implement a Con- To address these concerns, we screened the sight- servation Agreement forjaguar (Johnson and Van ing records for accuracy and created 3 different Pelt 1997). One of the main objectives of the Con- habitat suitability models by altering the input vari- servation Agreement was to identify potential ables (habitat or landscape features). We defined jaguar habitat in Arizona and New Mexico to help habitat suitabilitythe same as potentialjaguar habi- focus conservation efforts and identify opportu- tat (i.e., any area containing the features associated nities forjaguar habitat management. Signatories with jaguar sites as determined from our GIS of the MOA established numerous teams, includ- [ESRI, Redlands, California, USA] analysis) or ing aJaguar Scientific Advisory Group and a Habi- meeting the criteria supplied by the Jaguar Scien- tat Subcommittee. Arizona was considered the tific Advisory Group. Our first habitat suitability best starting point for mapping potential jaguar model was the most conservative because we only habitat in the United States because it contained used input variables found to be important in a the greatest number of jaguar reports north of GIS analysis. Our second model examined the Mexico during the 20th century (Rabinowitz sensitivity of the first model by relaxing some of 1999). Specifically, the objectives of our analysis the input criteria, allowing us to examine patch were twofold: (1) characterize potential jaguar connectivity and travel corridors. Our third model habitat in Arizona from historic sighting records was the most general, developed to include addi- and (2) create a statewide habitat suitability map. tional biomes that might have been missed in our GIS analysis due to spatial error or from errors of METHODS omission related to observational data. We characterized potentialjaguar habitat by over- Modeling Approach laying historic jaguar sightings and coarse-scaled In order to map potential jaguar habitat in Ari- habitat features we thought were important. We zona, we first had to identify and characterize it. took a landscape approach by focusing on coarse- The Jaguar Scientific Advisory Group recom- scaled habitat features because the jaguar sites had mended not using prey density as a criterion for highly variable positional error, making them insuf- jaguar habitat because wildlife agencies can man- ficient for a fine-scaled analysis. Coarse-scaledhabi- age wild game populations to increase in num- tat features considered important included vegeta- bers. They also recommended excluding areas tion biomes (ecosystems) and series (defined by with high human density or areas with anthro- dominant or characteristic species), elevation and pogenic impacts because jaguars are secretive an- terrain ruggedness, proximity to perennial or inter- imals and tend to avoid highly disturbed areas mittent water sources (i.e., streams, rivers, lakes, or (Quigley and Crawshaw 1992). Therefore, we springs), and human density. We used a GIS to cre- masked from our analysis areas within city bound- ate, manage and analyze the data, characterize aries, higher density rural areas visible on The- broad-scale habitat features where jaguars were ob- matic Mapper (TM) satellite imagery (30-m reso- served, and map potential jaguar habitat. lution), and agricultural areas. We obtained the outlines of cities and towns from the Arizona State Historical Database and Mapping Land Department and agricultural outlines from We examined 57 sighting records where jaguars TM imagery (Valencia et al. 1993). were supposedly sighted or killed (Appendix 1). The Jaguar Scientific Advisory Group provided The quality and reliability of the sighting records us with a list of potential habitat variables to help varied greatly, with some dating back 100 years. To steer our analysis, but most of the literature per- create a useful database for analysis, we sorted tained tojaguar habitat located far to the south of records into 3 classes based upon their reliability: Arizona in areas with dissimilar climates and bio- (1) physical evidence (n = 24), (2) firsthand ac- mes. Our task was complicated because the historic count obtained by a reliable source (interviewed sighting records contained widely varying spatial er- by a game warden or scientist [n= 7]), or (3) sec- ror (1-30 km), so we had to consider the possibil- ondhand account, which we considered unreli- ity that some of the biomes associated with jaguars able (n = 26). We also rated records according to might not be correct. Another problem was the their site description (Appendix 1 ) as excellent (n lack of information regarding the amount of time = 16), good (n = 5), fair (n = 4), or poor (n= 32) 1026 CHARACTERIZINGPOTENTIAL JAGUAR HABITAT * Hattenet al. J. Wildl. Manage. 69(3):2005

graphic maps, satellite imagery, hydrography, and shaded relief maps to visualize terrain. To re- duce positional error and create the most specific habitat suitability model, we used only the most re- liable and spatially accu- rate records (n = 25, Fig. 1). We considered sight- ing records that con- tained no physical evi- dence, were derived from secondhand accounts, or had spatial errors >8 km, insufficient for our spatial analysis. This does not mean that jaguars did not occur at some of the locations we labeled as poor or that jaguars did not visit other locations in Arizona. Hence, all spatial analyses and refer- ences to jaguar sightings refer only to the 25 sight- r ing records we analyzed.

N GIS Analysis 0 Oa80 (Im120 Ikwbndr A To calculate the dis- tance between jaguar sites and water, we cre- ated 2 that con- Fig. 1. A map displayingthe 25 jaguarsighting records we used. We onlyused sites withphys- grids ical evidence of jaguars(body, skin, or photographs)or first-handaccounts of jaguarsightings tained the distance of that were obtainedand accepted by a reliablesource (game wardenor scientist)and had an each grid cell to peren- arr.O-ntihiO nneitiJnnol Wa. i ylrnt- I1em-It AJnnJnefiUv I) a:ctpauiC II pUoiiai aCCiuIaby 8 km poor. An excellent-rated site tat features through much of the year in Arizona. contained specific information, such as the junc- To examine the relationship between jaguars and tion of 2 creeks; a good-rated site description elevation, we overlaid sighting records and Digital might say 3 km from the creekjunction, but with- Elevation Model (DEM) data (Fig. 2C). We sorted out necessarily a direction; while a fair-rated site the DEM data (30-m resolution) into 609-m description referenced a specific feature, like a (2,000-ft) elevation zones to more clearly identify creek junction or summit, but without a distance patterns in jaguar distribution. to the feature. Last, poor-rated site descriptions To examine the relationship between jaguars contained very broad locality information, like a and landscape roughness, we calculated a terrain mountain range. ruggedness index (TRI; Riley et al. 1999) for the To improve our mapping effort, we plotted whole state of Arizona (Fig. 2D). The TRI mea- jaguar sites with a GIS that displayed digital topo- sures slopes in all directions of each cell examined J. Wildl. Manage. 69(3):2005 CHARACTERIZINGPOTENTIALJAGUAR HABITAT * Hatten et al. 1027

i <610 m 611 - 1,220 m

.11 1,221 - 1.829 m _ 1.830 - 2.439 m ' 2,440 - 3,049 m I >3,050m D

X- Level - slightly rugged _ Intmeditely - moderately rugged M Highly rugged M Extremely rugged

Fig.2. Proximityto perennialor intermittentwaters (A) and springs(B), elevationzones (609-m bands [C]),and a terrainrugged- ness indexmap (D).

and thereby provides an index of overall rugged- tered and averaged) DEM data into 1-km2 cells, ness in a neighborhood, which is more descriptive and (3) we calculated the sum difference in eleva- than a simple slope surface. We calculated TRI in tion between each cell and its surrounding neigh- a 3-step process: (1) we mosaiced 1,986 30-m reso- bor cells. We divided the TRI data into 7 classes ac- lution DEMs together, (2) we aggregated (clus- cording to their relative roughness: level, nearly 1028 CHARACTERIZINGPOTENTIAL JAGUAR HABITAT * Hattenet al. J. Wildl. Manage. 69(3):2005 level, slightly rugged, in- 16 termediately rugged, moderately rugged, o c 12 highly rugged, and ex- We over- tremely rugged. 2 8I laid jaguar data with TRI data for examination. \ 4 To characterize vegeta- ao tion biomes and series occupied by jaguars, we overlaid sighting records Greatbasin Madrean RockyMountain Scrub grassland and Arizona GAP (AZ- conifer evergreen forest montaneconifer GAP) data (Halvorson woodland forest and Kunzmann 2000) 16 that had 82% overall ac- curacy at the biome level and 68% at the series level. The AZGAP is 12 based on the Brown, C B Lowe, and Pase (BLP) c: system (Brown et al. 8 1980, Brown 1994) that 0- contains 8 levels of 0) orga- Cu3 nization. The AZGAP ac- curacy assessment was 4 complicated by many fac- tors including spatial scale, boundaries, fuzzy 0 | , * , * ,.* -_ 1 and classification ranks I Pinyon- Encinal Oak-pine Pine Douglas-fir- Mixed (Kunzmann et al. 1998). -mixed grass- the AZGAP juniper However, conifer scrub cover contained the most detailed vegetation Fig.3. Frequencyof jaguarsightings in biomes (A) and vegetationseries (B). and land cover data of Arizona and provided an unprecedented opportunity to explore landscape- tains, contained 12% (n = 3) of the historic sight- scale relationships between jaguar sightings and ings. There have also been 4 jaguars sighted in vegetation biomes and series. southeastern Arizona since 1995 (no. 51, no. 52, no. 56, no. 57). Twenty-four percent (n = 6) of RESULTS jaguar sightings were dispersed along the Mogol- lon Rim from New Mexico to the Grand Canyon Jaguar Occurrence and DistributionPatterns and south to the Prescott area. The distribution ofjaguar sightings was clumped in southeastern Arizona and scattered in central GIS Overlays and northwestern Arizona (Fig. 1 ). A geographic One hundred percent (n = 25) of the sighting region surrounding Tucson (100-km radius) con- records were observed in 4 biomes (Fig. 3A). Of tained 54% (n = 16) of all jaguar sightings, in- these, 56% were observed in scrub grasslands of cluding 1 as recent as 2001, and it is comprised of southeastern Arizona, 20% in Madrean evergreen the following mountain ranges and associated forest, 12% in Rocky Mountain montane conifer grasslands: Santa Catalinas, Rincons, Santa Ritas, forest, and 12% in Great Basin conifer woodland. Baboquivaris, Tortolitas, Patagonias, Bartolos, At the vegetation series level (Fig. 3B),jaguars were Sierritas, and . A secondary observed 4.7 times more often in mixed grass-scrub concentration near Douglas, comprised of the than any other community. Related to water, 64% Chiricahua, Peloncillo, and Dos Cabezas Moun- ofjaguar records occurred within 5 km of a peren- J. Wildl. Manage. 69(3):2005 CHARACTERIZINGPOTENTIALJAGUAR HABITAT * Hattenet al. 1029 nial or intermittent creek or river, 76% within 10 to 21% when we excluded areas >10 km from km, and 84% within 20 km. Furthermore, 80% of perennial or intermittent waters and gently sloped jaguar records occurred within 2.5 km of a spring, areas (TRI < 4; Model A, Fig. 5). After considering and 96% occurred within 5 km. When we com- the experimental nature and unproven signifi- bined springs, rivers,and creeks, 100% of the sight- cance of the TRI variable that might exclude bio- ing records were within 10 km of a water source. logically important areas or mask travel corridors, Sixty percent of jaguars were observed between we reran the model without the TRI filter. This in- 1,220- and 1,829-m elevation (Fig. 4A), largely in the creased the potential habitat envelope to 28.9% of scrub grassland biome of southeastern Arizona. the state (Model B, Fig. 5). Omitting the TRI fil- The remainingjaguar sightings were between 1,036 ter reduced the patchiness of Model A and re- and 2,743 m. With respect to topography, 92% of sulted in the identification of potential travel cor- jaguar sightings occurred in intermediately rugged ridors between areas identified in Model A. to extremely rugged terrain (Fig. 4B), with the re- To compare our results to a more liberal model, mainder (8%) in nearly level terrain. we reran the model a third time (Model C, Fig. 5) to include areas that might have been visited by HabitatSuitability Mapping jaguars but missed in our earlier analysis because Four biomes contained 100% (n = 25) of jaguar of spatial error. We added the Madrean montane sites (Fig. 3A) and encompassed 37% of the state conifer forest biome because it contained vegeta- (108,222 km2). The suitability envelope decreased tion series commonly found in upper elevations of Sky Islands of southeast- 16 ern Arizona. We also added 4 riparian biomes A found in Arizona be- 12 cause jaguars might use 0 them for travel corridors. a' The addition of these 5

0- 8 biomes only marginally c) increased the amount of a potential jaguar habitat of the 4 (29.6% state). The area with the most jaguar sightings was lo- cated in southeastern I 0 610-12 1830 m 2440- Arizona (Fig. 5) in Santa 610 - 1219 m 1220- 1829 m 1830- 2439 m 2440- 3049 m Cruz, Pima, Cochise, Pinal, and Graham coun- ties. Potential habitat in that area loosely resem- 8 B bled an inverted V with the southern end sepa- rated by a swath of agri- cultural and developed 4- land and few perennial or intermittent waters. CD o, When we did not apply the TRI filter, habitat cor- 2 -- ridors formed to the south and north of the Cochise and Graham 0 - I County boundary. An- Nearly leedl Intemneiately Moderately Highly rugged Extremely other large patch of po- rugged rugged tential jaguar habitat ex- tended northwestward Fig.4. Frequencyof jaguarsightings within the 609-m elevationzones (A) and within5 of the 7 terrainruggedness indexclasses (B). from the Mogollon and 1030 CHARACTERIZINGPOTENTIALJAGUAR HABITAT * Hattenet al. J. Wildl. Manage. 69(3):2005

Gila Mountains of New Mexico, across the Mogollon Rim to the Williams-Flagstaff area. Without the TRI filter, this area just barely con- nected with another, smaller patch that ex- tended south to the Prescott area. Although these patches contained approximately 25% of the historic jaguar obser- vations, no jaguars were recorded from this area since 1964. The northern area was disconnected from the 2 habitat patches to the south by a broad band of unsuit- able land approximately 17 km wide that encom- passed the Gila River Valley near the city of Safford.

DISCUSSION Coarse-scaled Habitat Associations We found that most oc- o 49 jaguar sightings Legend ' curred between 1,220 ModelA N and 1,829 m, an eleva- M ModelB 0 60 120 Kilometers A tion band in which scrub SR -MM- ModelC I_ _ IA grasslands were com- mon. The apparent pref- Fig.5. We identifiedpotential jaguar habitat in Arizonain 3 separatemodel runs that used differ- erence of for ent inputvariables (habitat features) or criteria(roughness filter). Model A was the most spatially jaguars conservativemodel, with just 4 biomesand a terrainruggedness index (TRI) filter applied; we ex- scrub grasslands may ac- cludedlevel and slightlyrugged areas. ModelB was identicalto ModelA butwithout a TRIfilter reflect the use of applied.Model C was similarto ModelB but included5 additionalbiomes (Madreanmontane tually coniferforest and 4 All3 modelsexcluded areas thatwere >10 kmfrom water. travel corridors from the riparianbiomes). Sierra Madres of Mexico into southeastern Arizona rather than a preferred important selective factors in our analysis regard- vegetation type, or perhapsjaguars were just more less of the limitations of observational data. visible in open grasslands. Some problems with ob- There was a strong association between jaguar servational data are that animals will more often sightings and distance to water, with all sites falling be sighted in the open, especially in the daytime within 10 km of a perennial or intermittent water and in areas where more observers (e.g., ranchers, source. While we calculated these proximities as hikers) are located. All jaguar sightings in our straight line (Euclidean) distances-and in moun- database had occurred in the daytime and almost tainous terrain, they could have been considerably always in less dense vegetation. However, there further-the relatively close proximity of jaguars were no jaguar sightings in the lower elevation to water potentially explains much about their dis- deserts, which are very open and exposed; this tribution pattern in Arizona. River valleys might supports the idea that elevation and biomes were provide travel corridors for jaguars, along with J. Wildl.Manage. 69(3):2005 CHARACTERIZINGPOTENTIALJAGUAR HABITAT * Hattenet al. 1031 higher prey densities, cooler air, and denser vege- MANAGEMENTIMPLICATIONS tation than habitats. surrounding We identified a great deal of potential jaguar We found an association between occur- jaguar habitat along the Mogollon Plateau (Fig. 5), but rence and terrain that was also observed ruggedness jaguars have not been sighted north of the Gila in Mexico and and (Ortega-Huerta Medley 1999) River Valley in 40 years, supporting our hypothesis New Mexico and As the hu- (Menke Hayes 2004). that land use practices are limiting jaguar move- man in Arizona's population gently sloped valleys ment into central Arizona.Jaguar distribution pat- continues to areas grow,jaguars might avoid these terns in the last 40 years suggest that southeastern and stick to the more mountainous, rugged coun- Arizona is the most likely area for future jaguar oc- try. However, less rugged areas may serve as travel currence in the United States and conservation ef- corridors between habitat patches in neighboring forts should focus on protecting potential jaguar mountain ranges, providing that other habitat com- habitat in Santa Cruz, Pima, Cochise, Pinal, and ponents (e.g., cover, water) are available. Perhaps Graham counties (Fig. 6). The proposed conser- the most important factor explaining jaguars' ap- vation area is connected to Mexico and offers a parent preference for rugged terrain is the abun- unique opportunity for international cooperation dance of water in mountainous terrain of south- in the management and conservation of thejaguar eastern Arizona (Figs. 2A, B). at the northern extent of its range. Additional Research Needs ACKNOWLEDGMENTS The jaguar habitat suitability map (Fig. 5) pre- We thank the members of the Jaguar Conserva- sents a coarse outline of potential jaguar habitat tion Team's Habitat Subcommittee and the par- in Arizona but little in the way of fine-scale habi- ticipants in the Conservation Team's Working tat needs, and it should be viewed with caution un- Group for thoughtful comments and for their per- til more specific habitat-use data becomes avail- sistent, enthusiastic participation in this new ap- able. We constructed the 3 habitat models from proach to conserving jaguars. We also thank the observational data, and there is no accuracy cur- members of the Jaguar Scientific Advisory Group rently associated with the habitat map. We would have benefited by sorting c out the independent and Croke combined influences of landscape and habitat variables on jaguar oc- Eloy currence within a multi- variate-modeling envi- PINALCOUNTY . ronment, but our data were insufficient for this Mar approach. We need to PIMACOUNTY examine the habitat us- Tucson age of jaguars in north- ern Sonora, the closest population ofjaguars to Arizona, to refine and ground-truth our habitat asn suitability map. Further- more, identifying the travel corridors jaguars utilize from northern Sonora into Arizona and Legend '. Potential': habitat 0 50 100 Klometers A New Mexico is impera- tive to our understand- Fig.6. We identified a potential Iconservation area for jaguars in southeastern Arizona from his- ing of patch connectivity toricalsighting records and a ncillary information provided by the JaguarScientific Advisory across state and interna- Group.We developed 3 differEent habitat suitability models for jaguar, ranging from conserva- tiveto liberal.We used ModelC, ourmost liberal spatial model, in the identificationof a pro- tional borders. Iposed - - - - conservation------area.-- - -- 1032 CHARACTERIZINGPOTENTIAL JAGUAR HABITAT * Hatten et al. J. Wildl. Manage. 69(3):2005 for volunteering their time and expertise to help KUNZMANN,M. R., R J. HOBBS,AND P. S. BENNETr.1998. GAP and thematic assessment develop the habitat criteria for this model. vegetation accuracy pro- tocols using GIS and GPS technologies in Arizona. Funding for Arizona Game and Fish Department Proceedings of the 1998 Environmental Systems Re- in this was (AGFD) participation project provided search Institute Eighteenth Annual User Conference, by voluntary contributions to Arizona's Nongame San Diego, California, USA. AND Wildlife Checkoff, the AGFD's Heritage Fund (Lot- MENKE,K. A., C. L. HAYES.2004. Evaluation of the relative suitability of jaguar habitat in New tery Dollars Working for Wildlife), Arizona's Non- potential Mexico. New Mexico Department of Game and Fish Wildlife Checkoff, and Project W95 M under game Technical Report, Santa Fe, USA. the Federal Aid in Wildlife Restoration Act ORTEGA-HUERTA,M. A., ANDK. E. MEDLEY.1999. Land- (Pittman-Robertson Act). In addition, many vol- scape analysis of jaguar (Panthera onca) habitat using records in the Sierra de Mexico. unteers participated in the Jaguar Conservation sighting Tamaulipas, Environmental Conservation 26:257-269. Team and and have contributed Working Group QUIGLEY,H. B., ANDP. G. CRAWSHAW.1992. A conserva- with donated time and significantly money. tion plan for the jaguar Pantheraonca in the Pantanal of Brazil. Conservation 61:149-157. LITERATURECITED region Biological RABINOWITZ,A. R. 1999. The present status of jaguars BROWN,D. E. 1983. On the status of the jaguar in the (Panthera onca) in the southwestern United States. Southwest. Southwestern Naturalist 28:459-460. Southwestern Naturalist 44:96-100. ,editor. 1994. Biotic communities: southwestern , ANDB.G. NOTrINGHAM,JR. 1986. Ecology and be- United States and northwestern Mexico. University of haviour of thejaguar in Belize, Central America.Jour- Utah Press, Salt Lake City, USA. nal of Zoological Society of London 210:149-159. , ANDC. A. LOPEZGONZALEZ. 2001. Borderland RILEY,S.J., S. D. DEGLOR1A,AND R. ELLIOT.1999. A ter- jaguars. University of Utah Press, Salt Lake City, USA. rain ruggedness index that quantifies topographic het- , C. H. LowE, ANDC. P. PASE.1980. A digitized sys- erogeneity. Intermountain Journal of Sciences tematic classification for ecosystems with an illustrated 5:23-27. summary of the natural vegetation of North America. SANDERSON,E. W., K. H. REDFORD,C. B. CHETKIEWICZ, U.S. Forest Service, General Technical Report. RM-73. R. A. MEDELLIN,A. R. RABINOWITZ,J.G. ROBINSON, AND Rocky Mountain Forest and Range Experiment Sta- A. B. TABER.2002. Planning to save a species: thejaguar tion, Fort Collins, Colorado, USA. as a model. Conservation Biology 16:58-72. CHILDS,J.L. 1998. Tracking the felids of the borderlands. SCHALLER,G. B., ANDP. G. CRAWSHAW.1980. Movement Printing Corner Press, El Paso, Texas, USA. patterns of jaguar. Biotropica 12:161-168. FEDERALREGISTER. 1997. Final rule to extend endangered SEYMOUR,K. L. 1989. Panthera onca. Mammalian Species status for thejaguar in the United States. Federal Reg- 340:1-9. ister 62:39147-39157. SWANK,W. G., ANDJ.G. TEER.1989. Status of the jaguar. GLENN,W. 1996. Eyes of fire: encounter with a border- Oryx 23:14-21. landsjaguar. Printing Corner Press, El Paso, Texas, USA. TEWES,M. E., ANDD.J. SCHMIDLY.1987. The neotropical HALVORSON,W. L., ANDM. R. KUNZANN.2000. Assessment, felids: jaguar, ocelot, margay, and jaguarundi. Pages correction, and augmentation of the Arizona GAP veg- 697-711 in M. Novak,J. A. Baker, M. E. Obbard, and etation data: final report for the Arizona collection B. Malloch, editors. Wild furbearer management and agreement Number HAB-96-0017.U.S. Geological Sur- conservation in North America. Ontario Ministry of vey, Western Ecological Research Center, Sonoran Natural Resources, Toronto, Canada. Desert Field Station, UniversityofArizona, Tucson, USA. VALENCIA,R. A., J. A. WENNERLUND,R. A. WINSTEAD,S. JOHNSON,T. B., AND W. E. VANPELT. 1997. Conservation WOODS,L. RILEY,E. SWANSON,AND S. OLSON.1993. Ari- assessment and strategy for the jaguar in Arizona and zona riparian inventory and mapping project. 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Appendix1. The 57 sightingrecords examined for our GeographicInformation System analysis.Information on jaguarsites was obtainedfrom the publishedliterature, newspaper articles, and governmentrecords. Codea Yearb CIc LCd Location Sex 1 1901 3 2* 1902 1 2 RedingtonPass m 3 1902 3 CatalinaMountains 4 1903 3 Atascosa Mountains 5 1904 3 VerdeRiver 6 1906 3 ChiricahuaMountains f 7 1907 3 PatagoniaMountains 8 1907 3 MogollonMountains 9* 1907 1 1 South RimGrand Canyon m 10 1910 3 ChevlonCanyon f (co ntinued on next page) J. Wildl. Manage. 69(3):2005 CHARACTERIZINGPOTENTIAL JAGUAR HABITAT * Hatten et al. 1033

Appendix1. continued.

Codea Yearb CIC LCd Location Sex 11 1910 3 ChiricahuaMountains f 12* 1912 1 1 ChiricahuaMountains 13 1912 3 Sunset Mountain 14 1912 3 BozarthMesa 15 1912 3 CatalinaMountains 16 1912 3 RinconMountains 17 1913 1 4 TortolitaMountains 18 1913 3 Red Mountain m 19* 1916 1 3 CatalinaMountains m 20* 1917 1 1 Santa RitaMountains m 21 1918 3 South RimGrand Canyon 22 1918 3 Santa RitaMountains 23* 1919 1 1 Santa RitaMountains f 24 1920 2 4 RinconMountains 25 1920 2 4 Santa RitaMountains 26* 1922 1 3 RinconValley 27* 1924 1 3 MogollonPlateau 28 1926 1 4 PatagoniaMountains 29 1926 1 4 Atascosa Mountains m 30* 1926 1 2 Santa MariaMountains m 32 1929 3 Sand TankMountains 31 1930 3 ChiricahuaMountains 33* 1932 2 1 South RimGrand Canyon f 34* 1933 2 1 Atascosa Mountains m 35 1933 3 SierraEstrella Mountains 36 1934 3 Atascosa Mountains 37 1939 3 BloodyBasin 38* 1939 1 1 Atascosa Mountains m 39 1940 3 WhiteMountains 40 1947 3 Atascosa Mountains 41 1948 3 PatagoniaMountains m 42* 1949 1 2 CerroColorado Mountains f 43* 1957 1 2 WhiteRiver 44 1957 3 Red Mountain m 45* 1961 1 1 EmpireMountains m 46* 1963 1 2 WhiteMountains f 47 1964 1 4 BlackRiver m 48* 1965 1 3 PatagoniaMountains m 49* 1971 1 1 Santa Cruz River m 50* 1986 1 1 Dos Cabezas Mountains m 53* 1988 2 1 54 1988 3 SierritaMountains 55* 1993 2 1 BaboquivariMountains 51* 1996 1 1 PeloncilloMountains m 52* 1996 1 1 BaboquivariMountains 56* 1997 2 1 CerroColorado Mountains 57* 2001 1 1 PajaritoMountains m *Used in model developmentand presented in Figs. 1, 5, and 6. a Site locationson Figs. 1, 5, and 6. bThe year jaguarwas observed. c Cl = Class description:1 = physicalevidence (skin,body, photograph); 2 = reliablewitness (firsthandaccount that was obtainedand accepted by an authority[game wardenor scientist]);3 = less reliable(secondhand account). d Lc = Locationdescription: 1 = excellentlocation description (<1.7-km accuracy); 2 = good (1.7 to 5-kmaccuracy); 3 = fair (5.1 to 8-kmaccuracy), 4 = poor (>8-kmaccuracy). We did not rankclass 3 sites (Cl = 3) for locationaccuracy.