New data on the status and distribution of the bush dog (Speothos venaticus): Evaluating its quality of protection and directing research efforts

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1 BIOLOGICAL CONSERVATION 141 (2008) available at journal homepage: New data on the status and distribution of the bush dog (Speothos venaticus): Evaluating its quality of protection and directing research efforts Karen E. DeMatteo a,b,c, *, Bette A. Loiselle c a Saint Louis Zoo, Research Department, One Government Drive, St. Louis, MO 63110, USA b Saint Louis University, Department of Biology, 3507 Laclede Avenue, St. Louis, MO 63103, USA c University of Missouri St. Louis, Department of Biology, One University Boulevard, St. Louis, MO 63121, USA ARTICLE INFO ABSTRACT Article history: Received 21 December 2007 Received in revised form 2 July 2008 Accepted 11 July 2008 Available online 23 August 2008 Keywords: Carnivore Conservation GIS modeling MAXENT Survey The bush dog (Speothos venaticus), listed as CITES Appendix 1 vulnerable, is a small (5 6 kg), rarely seen canid from Central and South America. The World Conservation Union Canid Specialist Group (IUCN CSG) recommended that research with this species focus on their basic ecology so that a data-driven conservation strategy can be formulated. Information on the bush dog, however, has been lacking since standard field techniques have had little or no success with this species. The S. venaticus Status and Distribution Survey was developed in an attempt to use indirect methodologies to determine the bush dog s conservation status in the wild, its current distribution, and identify ecological needs by correlating habitat types to bush dog sightings. Survey responses and literature generated a database with 399 historic bush dog locations recorded between 1834 and These locational records were used to update the species range extent map, an important conservation planning tool. With year and precision of data accounted for, we analyzed land use coverage at known locations where bush dogs have historically been reported to evaluate the likelihood that the species persists in the area today. In addition, these locations provided training data for generation of potential distribution maps (i.e., areas of occupancy) using ecological niche modeling (i.e., Maximum Entropy) and bioclimatic data. These analyses revealed that 20% of the historic bush dog locations are associated with fragmented or altered habitat. These results allowed the status of bush dog habitat to be re-evaluated and areas that require more intensive research and protection to be identified. Ó 2008 Elsevier Ltd. All rights reserved. 1. Introduction The bush dog (Speothos venaticus), listed as CITES Appendix 1 Vulnerable (IUCN, 2004a), is a small and rarely seen canid from Central and South America (Fig. 1; IUCN, 2003) whose status, distribution, and ecological requirements are poorly understood (Eisenberg, 1989; Redford and Eisenberg, 1992; Silveira et al., 1998; Eisenberg and Redford, 1999; Michalski and Peres, 2005) and primarily based on opportunistic sightings (Defler, 1986; Strahl et al., 1992; Beccaceci, 1994; Silveira et al., 1998; Barnett et al., 2001). The World Conservation Union Canid Specialist Group (IUCN CSG) recommends that research with this species focus on gaining insight into their distribution, population density, habitat preferences, and * Corresponding author: Address: University of Missouri St. Louis, Department of Biology, One University Boulevard, St. Louis, MO 63121, USA. Tel.: ; fax: addresses: kdematteo@aol.com (K.E. DeMatteo), loiselle@umsl.edu (B.A. Loiselle) /$ - see front matter Ó 2008 Elsevier Ltd. All rights reserved. doi: /j.biocon

2 BIOLOGICAL CONSERVATION 141 (2008) Fig. 1 Map showing the overlap between the Bioclimatic Model (MAXENT V2.3.0) and historical distribution of Speothos (IUCN, 2003): Grey corresponds to habitat in the model prediction with a probability P 1 (i.e., suitable). Open hatch corresponds to species presence in the historical distribution. Grey open hatch corresponds to those areas where the model prediction and historical distribution overlap. Map contains the 399 Speothos historic locations with corresponding level of precision: open circles with black dots in the center indicate exact locations where Speothos is present, black triangles indicate estimated locations where Speothos is present, and black X s indicate estimated locations where Speothos is absent. See Appendices A and B for sources of data. behavioral ecology so that a data-driven conservation strategy can be formulated (IUCN, 2004b). However, this species has proven difficult to work with in the field. While several techniques may aid researchers in their efforts to work with this species in the field (DeMatteo and Kochanny, 2004; DeMatteo et al., 2004; Wasser et al., 2004), it will take years, if not decades, to collect the needed data. In an attempt to overcome this struggle against time, a survey aimed at collecting information on the bush dog s status, distribution, and concerns was developed and sent to researchers, conservationists, and other informed personnel. Five goals were associated with this data collection effort: (1) development of a Speothos database that could be updated and reanalyzed as new field information became available, (2) generation of updated maps of the bush dog s current and potential distribution, (3) identification of areas for focused research efforts with the bush dog, (4) evaluation of the efficacy of protected areas in providing sufficient area for self-sustaining bush dog populations, and (5) prioritization of areas that need legal protection. 2. Methods 2.1. Survey In January 2003, the S. venaticus Status and Distribution Survey was developed with one of its aims directed at understanding the current distribution of the bush dog. Variation in reporting

3 2496 BIOLOGICAL CONSERVATION 141 (2008) of bush dog historic locations was minimized by including a table requesting specific information on each observation (e.g., date, locality, coordinates, state, vegetation type) and detailed country maps ( In order to maximize the number of people that could directly respond to the Survey, it was made available in English, Spanish, and Portuguese. The Survey was sent to more than 100 people working in the area of carnivore conservation (e.g., field researchers, non-government organizations, government organizations, conservation organization, museums, universities) and posted on the IUCN CSG website ( The Survey was sent (i.e., via mail or ) to individuals in the 12 Central and South America countries that include part of the bush dog s historical distribution (Argentina [AR], Bolivia [BO], Brazil [BR], Colombia [CO], Ecuador [EC], French Guiana [FG], Guyana [GY], Panama [PA], Paraguay [PY], Peru [PE], Suriname [SR], and Venezuela [VE]), as well as to various contacts in the US and Europe. The original list of contacts was generated through the South American section of the IUCN CSG and contained a list of people that had extensive experience working with canids. A secondary list was generated via communication with this primary set of contacts and included people that they trusted and knew had first-hand knowledge of bush dogs in a specific region or local Speothos database The developed Speothos database includes all historic locations, both direct (e.g., sighting) and indirect (e.g., tracks, camera traps, hair collection, interview, and vocalizations) evidence, reported in both the Survey and literature (Appendix A). Since data were generated from respondents that had extensive experience with canids or from contacts recommended by these respondents, all reported locations were considered accurate for species identification (i.e., bush dog), independent of the type of evidence. Each location was categorized as either exact or estimated. While exact locations had specific latitude-longitude or UTM coordinates provided and were considered to have errors of less than 1 km in most cases, estimated locations were reported as general points (e.g., town) or areas (e.g., reserve). Estimated point and area locations had coordinates assigned using published resources (e.g., CALLE, previously world; NIMA, and central location within the area s boundary (e.g., state, reserve), respectively. Only exact locations were used in evaluating or modeling the bush dog s distribution with ecological (e.g., forest type, altered habitat) and climatic features (e.g., temperature) Ecological niche modeling: seasonal land coverage regions We evaluated the present-day habitat surrounding the exact bush dog locations, using two US Geological Survey (USGS) global land cover databases ( sa_int.html and that characterized seasonal land coverage into 166 habitat types. Both land cover databases were generated by remotesensed data collected at a 1-km 2 resolution between April 1992 and March Bush dog locations were rasterized to 1-km 2 area so that even if numerous bush dog locations were reported for that area, presence data were reduced to a single record per grid cell. Using ArcGIS Ò 9(ESRI, Inc., 2005) Speothos locations were then summarized as occupying intact, fragmented, or altered (i.e., cropland) habitat. Misclassification errors were minimized by matching habitat description recorded by observers, when available, with characteristics of the grid cell. Buffer zones, centered on exact historic locations, were used to evaluate the long-term stability of local bush dog populations. This stability was defined as having intact habitat in the area needed to support a self-sustaining population. Prior to establishing the buffer zones we needed to define the number of breeding individuals required to sustain a wild bush dog population and the area such a population requires; however, definitive values for both are unknown for bush dogs. Differences in the bush dog s sociality (e.g., group living versus solitary), evolutionary history, and, likely, ecology and population dynamics (e.g., dispersal pattern) prevent direct extrapolations from other Neotropical carnivores. Therefore, for this analysis we used existing Neotropical carnivore estimates (i.e., 50 breeding individuals for jaguars; Sanderson et al., 2002), estimates from other social canids (i.e., small (50 individuals) and large populations (P100 individuals) for African wild dogs; IUCN, 1997), the limited knowledge of the bush dog s socioecology, and the genetic rule used by captive managers that populations should contain a minimum of 20 breeding individuals (Lacy, 1989) as guidelines to generate two potentially viable local population size estimates for bush dogs: 25 and 50 groups (i.e., male female breeding pair). The use of group versus individual estimates was based on the fact that bush dogs are observed in a variety of group sizes (e.g., 1 10 individuals; Deutsch, 1983; Defler, 1986; Peres, 1991; Strahl et al., 1992; Silveira et al., 1998; Barnett et al., 2001) and captive studies have led to the suggestion that bush dogs live in extended family groups (Kleiman, 1972) with reproductive suppression in offspring (MacDonald, 1996). The size of the buffer zones was based on four existing bush dog home range estimates: three were developed from theoretical models using regression based on body mass ( km 2, Silveira et al., 1998;20km 2, Van Humbeck and Perez, 1998, km 2 (2 6 individuals), Beisiegel, 1999) and one based on minimum convex polygon with systematic data collection on a single radio collared family group (100 km 2,E. Lima, pers. comm.). While additional field studies are needed to address whether this breadth of home range is a reflection of variable response to local characteristics of the bush dog and its environment (e.g., group size, habitat type, degree of habitat fragmentation, prey density), preliminary field data suggests that the larger home range estimates may be more accurate. For this analysis we used an estimated home range size of 100 km 2 per group with non-overlapping home ranges between groups, as it is suspected that bush dog home ranges have little or no overlap due to both the potentially large number of prey a group of bush dogs can consume and their semi-nomadic behavior (K. DeMatteo, pers. obs.). Due to a lack of field data, it is impossible to know if bush dogs truly have non-overlapping home ranges or if there is some degree of overlap occurring; therefore, we believe the use of non-

4 BIOLOGICAL CONSERVATION 141 (2008) overlapping home ranges is a conservative approach that will avoid underestimating the area required to sustain bush dogs. Therefore, the total area needed to sustain 25 or 50 local groups of bush dogs (50 or 100 individuals, respectively) was 2500 km 2 and 5000 km 2, respectively Ecological niche modeling: bioclimatic data To estimate the potential distribution of bush dogs we used ecological niche modeling to combine data on known locations where bush dogs have been historically reported (exact locations only) and climate data. No exact locations were eliminated from modeling analyses since all points, even those prior to the 1980s, overlapped with locations reported in the last ten years. Ecological niche modeling has become an increasingly important tool for conservation biologists to provide estimates of the area occupied by species over large scales (e.g., Feria et al., 2002; Peterson et al., 2002; Draper et al., 2003; Peterson, 2003; Graham et al., 2004; Araújo and Guisan, 2006). We used climate data available from the WorldClim database ( html; Hijmans et al., 2005), which contains a minimum of 30 years ( ) of monthly temperature ( C) and precipitation (mm) measurements at a 30 arc-second resolution (approx. 1-km 2 ). We selected eight of the 19 bioclimatic variables and aggregated these data to an approximate 5-km 2 resolution for modeling: annual mean temperature, mean diurnal temperature range (mean of monthly [maximum temperature minimum temperature]), isothermality (mean diurnal temperature range/[maximum temperature of warmest month/minimum temperature of coldest month]), temperature seasonality (standard deviation 100), annual precipitation, precipitation of driest month, precipitation seasonality (coefficient of variation), and precipitation of warmest quarter. These variables were selected because of their presumed relevance to limiting distributions of bush dogs and because they were largely uncorrelated (i.e., generally r < 0.70). We used Maximum Entropy (MAXENT V 2.3.0; Phillips et al., 2006) as our ecological niche model. MAXENT is a machine learning method which uses mathematical formulation to determine the additive contribution of all variables in the model (Phillips et al., 2006). In a recent study that used independent presence/absence data distributed over multiple geographic regions, MAXENT consistently performed higher than other algorithms (e.g., GARP, DOMAIN) independent of region (Elith et al., 2006). In addition, MAX- ENT was determined to be more effective with species that have wide distributions, like the bush dog (Feria, 2007). Unlike most niche models that work with presence-only data, MAXENT can use the threshold-independent method of area under the receiver operating characteristic curve (AUC and ROC; Fielding and Bell, 1997) to evaluate the model s prediction performance. In generating the ROC, the program replaces known absences with a large number of automatically generated random points (i.e., background sample of pseudoabsences) and defines specificity by predicted area rather than commission error (Phillips et al., 2006). The program was set to run 500 iterations with a convergence threshold of , a maximum of 10,000 background points, and algorithm parameters set to auto features. We had the program randomly withhold 25% of the 256 presence locations (64 points) in order to test the model performance Speothos occurrence relative to protected/unprotected areas Two databases, the 2003 World Database on Protected Areas (WDPA03 and PróCarnívoros, a Brazilian non-governmental organization for the conservation of Neotropical carnivores (R. Cunha da Paula pers. comm.), which contain information on protected areas were used to determine the level of legal protection provided to the bush dog. The WDPA03 polygons (n = 927 total for AR, BO, BR, EC, FG, GY, PM, PY, PE, and VE or those countries where exact bush dog locations are recorded) included a wide range of protected areas recognized at both the national and international level: biosphere reserves, RAMSAR sites, world heritage sites, other protected areas or reserves recognized at the national level, including proposed areas and areas with incomplete information, and established protected areas of IUCN categories I through VI (strict nature reserve [Ia], wilderness area [Ib], national park [II], natural monument [III], habitat/species management area [IV], protected landscape/seascape [V], and managed resource protected area [VI]; IUCN, 1994). The PróCarnívoros polygons (n = 2067 total for BR) included areas classified into one of three categories: direct use where the ecosystems are nominally under total government protection (i.e., national, state, and municipal parks), indirect use where an area s resources can be exploited or managed in ways compatible with nature conservation (i.e., environmentally protected areas with flexible land use at the state and district levels, municipally regulated permanent protection areas, areas of ecological interest, state and national forests, reserves for sustainable development at the state and federal level), and areas not explicitly dedicated to nature conservation but the inhabitants tend to use them with minimal environmental impact (i.e., indigenous reserves). While the WDPA03 and PróCarnívoros databases overlap, the latter has many unique polygons that likely reflect the WDPA03 s acknowledged lack of geographic data, i.e., polygon data, for many protected areas. In addition to looking at the quality of habitat, another component of long-term stability of bush dog populations is determining how effective protected areas are at providing sufficient space to maintain self-sustaining population. As with the previous buffer zone analyses, we used buffers (ESRI, Inc., 2005) centered on exact bush dog locations with total areas of either 2500 km 2 (25 groups each requiring 100 km 2 ) or 5000 km 2 (50 groups each requiring 100 km 2 ). The difference between the percentage of buffer zones that occurred partially or completely in protected areas and those that occurred outside of protected areas was used to identify protected areas that would need to be expanded to provide complete coverage of an assumed minimum viable population of bush dog. Results were summarized for each country and across the range.

5 2498 BIOLOGICAL CONSERVATION 141 (2008) Results 3.1. Survey response The Survey added new data on the bush dog that was not previously published. Of the 138 surveys distributed, 35 (25%) were returned. Surveys contained information on all 12 countries in the bush dog s historical range. Thirty-four responses provided information on current distribution in the 12 countries (5 AR, 6 BO, 14 BR, 1 CO, 2 EC, 3 FG, 1 GY, 1 PA, 1 PY, 2 PE, 1 SR, and 1 VE); three of these surveys contained information on multiple countries (Appendix B) Speothos database Combining across all sources, the Speothos database now contains a total of 399 historic locations, 390 indicating presence and 9 marking absence across the 13 countries with in historic range of bush dogs (Fig. 1; Appendices A and B). It is unknown if these recorded absences existed throughout the history of the bush dog or if they resulted from other influences (e.g., habitat destruction, environmental changes). The historic locations were recorded between 1834 and 2004: 33 (8%) from 1834 to 1979, 23 (6%) from 1980 to 1989, 174 (44%) from 1990 to 2004, and 168 (42%) from no reported year. Within this last category, the majority of locality records were known to be from the last decade (e.g., range of years provided by respondent), but an actual date could not be obtained. Of the 399 historic locations, 143 (36%) were estimated locations and 256 (64%) were exact locations. Survey respondents provided 189 (74%) of the 256 exact locations (Appendix B). Of the 256 exact locations, 113 (44%) were direct observations. The 256 exact locations were recorded between 1842 and 2003: 2 (1%) in the 1800 s, 5 (2%) between 1960 and 1979, 19 (7%) between 1980 and 1989, 147 (58%) between 1990 and 2003, and 83 (32%) from no reported year. Within this last category, the data were categorized (direct or indirect) and included in analyses since all points, independent of year, overlapped with locations reported in the last 10 years Land use evaluation Historic locations of Speothos intersected 57 of the 166 habitat types in the seasonal land cover database. While 80% of bush dog locations were recorded in grid cells with intact habitat, almost one-quarter (20%) was considered fragmented (3%) or altered (17%). Forested areas (34.5% moist tropical evergreen rainforest, 14.5% tropical evergreen rainforest, 10% subtropical moist broadleaf forest, 6% tropical evergreen broadleaf forest, 3% humid lowland tropical rainforest, 1.0% flooded evergreen broadleaf rainforest, 0.5% marsh wetland, 0.5% moist tropical evergreen rainforest-flooded tropical evergreen rainforest [e.g., mangroves along coast], 0.5% submontane moist evergreen broadleaf forest, and 0.5% temperate evergreen broadleaf and needleleaf forest) composed the majority (71%) of intact habitat with the remainder (9%) categorized as deciduous woodland (e.g., chaco; 4%), savanna (2%), deciduous woodland-grassland mosaic (1%), shrubland (0.5%), shrubland-grassland mosaic (0.5%), grassland-forest mosaic (0.5%), and seasonally flooded savanna with palms (0.5%). At least three of these habitats were never previously associated with bush dogs: caatinga, chaco, and mangroves along the coast. Both fragmented (2% forest-grassland, 0.5% forest-shrub savanna, and 0.5% grassland-shrubland) and altered habitat (5% savanna-grassland with cropland, 4.5% forest with cropland, 3.5% shrubland-thorn woodland [e.g., caatinga]-cropland, 3% wet savanna with cropland, and 1% open woodland with cropland) were found associated with many of these intact habitat types. Fragmentation occurs throughout the bush dog s historic distribution but is most pronounced in central-eastern BO, central-eastern BR, western CO, western EC, PY, western PE, and northern VE. Not surprisingly, these higher levels of fragmentation are associated with human populated areas, which will only continue to increase in local density and area covered. This lack of habitat security was reflected as an increase in percentage of fragmented and altered habitat when individual locations were expanded to reflect the area needed to support self-sustaining groups of bush dogs. The percent of fragmented or altered habitat was similar (28% and 30%) for both breeding group estimates (25 and 50); however, the total altered area represented increased with group size (i.e., 75,149 km 2 for 25 groups versus 145,679 km 2 for 50 groups) Potential bush dog distribution from ecological niche modeling We generated a binary map (predicted suitable or unsuitable; Fig. 1) of potential bush dog distribution (i.e., areas of occupancy) by assigning all MAXENT model output values P1 as suitable environmental habitat and all values <1 as unsuitable. Using random pseudoabsence background points automatically generated by the program, the AUC of the MAXENT model was calculated to be 0.94 indicating that the prediction has a high level of accuracy in discriminating those areas where the species occurs from those where it is absent (i.e., >0.90; Swets, 1988). After conversion to an equal-area map projection, the total area predicted to be suitable for bush dogs was 14,444,725 km 2, which is approximately 58% of the total area in Central and South America. When the potential distribution based on MAXENT models is compared to the historic distribution (IUCN, 2003; Fig. 1), 77% of the potential distribution fell within the historic distribution (Fig. 1). The remaining 33% fell outside of the published historic range and included both expansions within current range countries (e.g., eastern Colombia) and expansion outside of the published historic range (Fig. 1). These new expansions extend from eastern PA through southern Mexico, portions of eastern Uruguay, and a small region in southern Chile. In addition to these expansions, there were several countries (BO, BR, CO, EC, PY, and PE) that had range contraction when comparing the model prediction to the published historic range. It should be noted that some range expansions are almost undoubtedly commission errors or false positives (e.g., Chile, Mexico), while others are concordant with location data (e.g., eastern CO, eastern PY, Santa Catarina State in SE BR).

6 BIOLOGICAL CONSERVATION 141 (2008) Speothos occurrence relative to protected/unprotected areas Overall, the percentage of buffered locations without any protection was dramatically higher than those with complete protection (Table 1). For example, for 25 and 50 groups across the 10 countries with exact locations, 31% and 25% were with no protection compared to only 2% and 1% with complete protection. The fact that the difference between these two extreme levels of protection (i.e., 0% or 100%) exists, locations with some degree of partial protection emphasizes the need to increase the connectivity between existing protected areas. 4. Discussion Ecological niche modeling can assist conservation planning, especially in cases where knowledge regarding the distribution of species is limited. The poorly understood bush dog was an ideal candidate for ecological niche modeling because of its wide distribution yet limited information on its occurrence throughout its historic range. Almost one-quarter (20%) of historical bush dog locations were associated with fragmented or altered habitat. This estimate should be considered a minimum of bush dog area that is currently altered as more than a decade has since elapsed since this land use map was developed ( ). Ecological niche model resulted in a potential distribution with high discrimination capability (AUC = 0.94); this model included some areas outside of the known range that were probably never occupied by bush dogs (e.g., southern Chile, Mexico) likely due to dispersal limitations. The understanding that the percentage of fragmented-altered bush dog habitat is only going to increase from this 20% baseline emphasizes the urgent need to determine areas that need protection and understand how this habitat variability affects the bush dog s ecological requirements. Although it is suspected that fragmentation-alteration could decrease prey levels (e.g., increased hunting pressure, exposure to disease from domestic animals) and increase the minimum required home range size for a group of bush dogs, it is unknown whether this occurs or if there is a threshold above which bush dogs cannot compensate for lower habitat quality. Until field studies can determine this ecological information, it is critical to maximize the amount of bush dog suitable habitat (i.e., 58% or just over half the total area in Central and South America) that is protected. The variation that was allowed in the current analyses of protected areas permitted a more effective evaluation of the protection afforded to the species and the extreme ranges that must be considered. It is critical to note that the actual level of legal enforcement provided to a particular area, especially those more remotely located, is extremely variable and a concern that must be addressed at all stages of conservation planning Country-specific recommendations (ecological niche modeling + occurrence relative to protected areas) CR: There is an obvious need to determine the breadth of the bush dog s distribution within CR and whether there is any connectivity between CR and populations in western PA. While CR contains reports of bush dogs, it was not one of the 12 countries included in the published historical distribution. PA: The long-term survival of bush dog populations is dependent on increasing the number of protected areas in both the eastern and western zones, especially along the borders with CR and CO, as well as the central corridor between these two zones. CO: Despite the number of protected areas in the NW region, the higher level of habitat fragmentation-alteration compared to the E indicates the need to maintain/increase the level of protection in this region. The low number of recorded locations and lack of locations in both the NW and eastern regions emphasizes the need for additional research throughout the country. Two regions that require research are: (1) the CO-VE border to determine if bush dogs are using Table 1 The buffer zone analyses which examined degree of protection summarized for the two sustainable population estimates of 10 and 25 groups and home range size 25 km 2 /group No. exact locations No. polygons 10 Groups 25 Groups 0% Protected 100% Protected 0% Protected 100% Protected All Countries Argentina Bolivia Brazil Brazil * Ecuador French Guiana Guyana Panama Paraguay Peru Venezuela The percentage of buffer zones, with bush dog exact locations as the center point, that occurred completely within and completely outside of protected areas are reported. The total number of exact bush dog locations and number of protected polygons are reported for each country. With the exception of the single analyses of the PróCarnívoros database that is marked with an asterisk (*), results were based on analyses of the WDPA03 database.

7 2500 BIOLOGICAL CONSERVATION 141 (2008) these areas of lower elevation to move E-W and bypass the physiological barrier of the Andes and (2) the western edge of PA, along the edge of the Pan American Highway, a potential connection between the N-S region of western CO, western PA, and potentially northern EC. The status of the bush dog in this latter region is in question due to the lack of reports combined with the fact that, unlike the published historic distribution, the model prediction did not include this area. VE: The numerous protected areas, high bioclimatic suitability, and widespread locations of bush dogs emphasizes the importance of VE as a potential stronghold in the bush dog s distribution; however, the connectivity between the N- S regions is a concern due to the potential physical barrier of the Guiana Highlands, the higher level of fragmented-altered habitat in the N, and few defined protected areas in the central region just NW of the Guiana Highlands along the Rio Orinoco. GY: The high level of bioclimatic suitability but low of number bush dog records across the country indicates a need for research directed at clarifying the status of bush dogs, especially along the borders of BR, SR, and VE. Conservation efforts should be focused on increasing the number of defined protected areas that have high interconnectivity, especially within the northern portion of the country where recorded bush dog locations occur among higher levels of fragmented-altered habitat. SR: Given the high bioclimatic suitability, habitat integrity, and key position in the bush dog s distribution (i.e., surrounded by FG, GY, and BR) but few defined protected areas and recorded bush dog locations, there is a clear need to couple efforts to ensure protection of suitable habitat and to determine the status of bush dogs throughout the country. FG: The widespread suitability indicated by the bioclimatic modeling suggests that research needs to be directed at the eastern and western regions to clarify the breadth of the bush dog s distribution, especially along borders with BR and SR. Conservation efforts should be focused at ensuring sufficient numbers of protected areas with high interconnectivity, especially within the northern region where fragmentation-alteration is the highest. EC: Bioclimatic modeling and habitat integrity suggests that determining the breadth of the bush dog s distribution and increasing the number and interconnectivity of protected areas in the NE region should be the focus of conservation efforts. In addition, ensuring the connectivity between the NE with areas along the borders of southern CO and northern PE needs to be addressed. The status of the bush dog in this area is unclear as the predictive model suggests that suitable areas are isolated and more widespread then the published historic distribution. Research is needed to determine the status and stability of the NW populations that are located on the edge of regions that contain few protected areas among a mosaic of fragmented-altered habitat and may be physically separated from populations in CO and PE (i.e., Andes). PE: With the exception of the SE border along BO, the status of bush dogs in N-NW region and the area along the border with BR needs to be determined. Making expansion of protected areas along the borders of BO, BR, CO, and EC a primary conservation effort will result in ensuring effective corridors between the countries. BR: In addition to securing habitat in the N-S central corridor where higher levels of fragmented-altered habitat and fewer protected areas exist, it is important that research efforts clarify the breadth and stability of the bush dog distribution across BR and with bordering countries. It needs to be determined whether the limited bush dog records that exist in regions of extremely high levels of fragmentation-alteration represent stable bush dog populations or small, isolated groups. It is important to note that three regions in BR that were previously reported in the published historic distribution but were absent in the model prediction: portions of the NE (e.g., SW Ceará, E Piauí, central Bahia, central Minas Gerais), south-central region (i.e., Mato Grosso), and western edge (i.e., Amazonas). The potential affect of this loss of area in the continuity across BR needs to be examined. In addition, it is important to note that while both the historical and current prediction for the bush dog s distribution includes all of central Amazonia, historical locations are reported only along the periphery of this region (Fig. 1). We do not believe that this pattern correlates with bush dogs being absent from the core of this but instead indicates a general lack of knowledge from this region. In fact, Sanderson et al. (2002) found a similar lack of knowledge for this region when developing a conservation plan for jaguars. We know, based on the current ecological niche model prediction, that this region contains suitable environmental conditions for bush dogs and believe that additional research efforts in the region would generate additional bush dog locations. BO: Despite the overall continuity in recorded bush dog locations and habitat integrity across this region, there is a need to increase both the number of protected areas and the connectivity among them. Research is needed to determine the full breadth of the bush dog s distribution in the northern region and the E-W-central corridor which bioclimatic modeling and high habitat integrity indicate are suitable bush dog areas and could provide important continuity with adjacent countries. Determining the status of bush dogs in southern BO is a low conservation priority since bioclimatic modeling suggests that the Andes are physiological barrier limiting the southern extreme of bush dogs in BO. PY: With the SE region of PY having both the highest number of recorded locations and bioclimatic suitability it should be the primary focus of conservation efforts, especially since some states have records reporting both presence and absence of bush dogs. Research must also clarify the status of bush dogs in the SW and central departments adjoining this region. Whether any connectivity still exists between the N- S regions is questionable due to poor habitat integrity and lack of protected areas and needs to be determined. The northern connection with BO-BR is absent in the current model prediction but present in the published bush dog distribution. AR: Levels of fragmentation-alteration suggest that bush dogs are limited to a single province in NE AR but research is needed to determine whether the connectivity suggested by bioclimatic modeling exists with PY and BR. This region at the southern extreme needs concerted conservation efforts

8 BIOLOGICAL CONSERVATION 141 (2008) between AR, BR, and PY as the extremely high levels of fragmentation-alteration in all three countries suggest long-term connectivity is in question. Predictions outside of published historic range: The predicted expansion of bush dogs into portions of Central America (i.e., CR, Nicaragua, Honduras, El Salvador, Guatemala, Belize, and southern Mexico) and South America (Uruguay and Chile) is unlikely, with the possible expansion into southern CR (see above). While the factors limiting expansion are unknown for bush dogs, it is likely that a combination, including geographical/physical barriers, disease, and biotic interaction (e.g., competition) are all important (Peterson et al., 1999; Anderson et al., 2003). If there are isolated, unknown populations of bush dogs in these regions, their long-term survival is in question due to their unknown degree of protection (i.e., occurrence in/out of protected areas) combined with high levels of habitat fragmentation-alteration throughout many of the areas. 5. Conclusions Ecological niche modeling allows point data on species location to be combined with detailed spatial information on climate and vegetation to provide insight into the bush dog s ecological needs (e.g., habitat) and physiological limits (e.g., temperature). The results of the Survey and ecological niche modeling emphasize the need for more detailed ecological studies o0n bush dogs to inform further research and conservation efforts. These models provide a preliminary estimate of suitable bush dog regions that can be modified as additional data emerge regarding their status, distribution, and basic ecological requirements. The conservation and management of the bush dog requires a large-scale approach that considers its broad ecological requirements and spatial use of areas that often cross political boundaries. Acknowledgements We thank the IUCN CSG who endorsed the project and provided web access to the survey, Luis Padilla and Tadeu Gomes de Oliveira who assisted with survey translation and Carl Soulsbury, Renata Leite, Claudio Sillero, and Matt Swarner for valuable input on the design of the survey, and PróCarnívoros Associação para Conservação dos Carnívoros Neotropicais for sharing their database on Brazilian protected areas. This project would not have been possible without all of the Survey respondents who took the time to share their knowledge for the benefit of the species: AR: Mario S. Di Bitetti (Fundación Vida Silvestre AR), Mónica V. Pia, Miguel Angel Rinas (Ministerio of Ecología RNRyT), Karina Schiaffino (CIES); BO: Joanne Devlin (Black Diamond Paving), Leonardo Maffei (WCS Proyecto Kaa Iya), Julieta Vargas (Museo Nacional de Historia Natural), Robert Wallace (WCS); BR: Brazil Natural History Museum, Otávio Borges Maia (IBAMA), Fernanda Michalski/Tadeu de Oliveira/Flavió Henrique G. Rodrigues/ Leandro Silveira (AssociaçãoPróCarnívoros), Guilherme Hemnique Braqa de Miranda (Policia Federal), Manual Ruedi (Natural History Museum of Geneva, Switzerland), Joaquim de Araujo Silva (Instituto Biotrópicos de Pesquísos em Vida Silvestre), Liliana Tiepolo (Museo Nacional Universidade Federal do Rio de Janeiro), Arnaud Desbiez (Durrell Institute for Conservation Ecology University of Kent); EC: Felipe Campos (TNC), CDC Ecuador; FG: Francois Catzeflis (Universite Sciences & Techniques), Benoit de Thoisy (Kwata NGO), Geraldine Veron (Muséum National d Histoire Naturelle-Paris); GY: Eustace Emerick Alexander (CI Guyana); PA: NB Gale (US Department of Defense); PY: José L. Cartes (Asociación Guyra Paraguay); PE: Major L. Boddicker (Rocky Mountain Wildlife Services); VE: Francisco J. Bisbal (Ministerio del Ambiente y los Recursos Naturales); AR/BR: Douglas Tent (Focus Tours, Inc.); BO/BR/SR: Museum of Natural History (London); BO/ BR/CO/PE: Teresa Pacheco (American Museum of Natural History). Appendix A List of literature used in developing the Speothos Database. This list includes literature that contains specific geographical information for bush dogs and is not meant to be viewed as a complete list of current literature on the species. Country Reference Argentina Ginsberg, J.R., Macdonald, D.W., Foxes, wolves, jackals, and dogs: an action plan for the conservation of Canids. IUCN/ SSC Canid Specialist Group, Gland, Switzerland and Cambridge, UK Jayat, J.P., Bárquez, R.M., Diaz, M.M., Martinez, P.J., Aportes al conocimiento de la distribución de los carnivoros del Noroeste de Argentina. Mastozoologia Neotropical 6, Bolivia Anderson, A., Mammals of Bolivia, taxonomy and distribution. Bulletin American Museum of Natural History 231, Emmons, L.H., Mammal fauna of Parque Nacional Noel Kempff Mercado. In: Killeen, T.J., Schulemberg, T.S. (Eds.), A Biological Assessment of Parque Nacional Noel Kempff Mercado, Bolivia. Rapid Assessment Program (RAP) Working Papers 10. Conservation International, Washington DC, USA, pp FAN-WCS, Plan de manejo, Reserva de Vida Silvestre de Rios Blanco y Negro. Fundación Amigos de la Naturaleza and the Wildlife Conservation Society, PL480. Santa Cruz, Bolivia Rumiz, D., Eulert, C., Arispe. R., Evaluacción de la diversidad de mamiferos medianos y grandes en el Parque Nacional Carraso (Cochabamba-Bolivia). Revista Boliviana de Ecologia y Conservación Ambiental 4, Wallace, R.B., Painter, R.L.E., Saldania, A.S., An observation of bush dog (Speothos venaticus) hunting behavior. Mammalia 66, (continued on next page)

9 2502 BIOLOGICAL CONSERVATION 141 (2008) Appendix A continued Country Brazil Reference Berta, A., The Pleistocene bush dog, Speothos pacivorus (Canidae) from the Lagoa Santa Caves, Brazil. Journal of Mammalogy 65, Fonseca, G.A.B, Redford, K.H., The mammals of IGBE s ecological reserve, Brasilia and an analysis of the role of gallery forests in increasing diversity. Revista Brasileira de Biologica 44, Dalponte, J.C., Estudos preliminaries sobre o cachorro do mato-vinagre, Speothos venaticus, na Estação Ecológica Serra das Araras, Estado do Mato Grosso, Brasil. Unpublished Report: Program for Studies in Tropical Conservation, University of Florida, 12pp. Dalponte, J.C., The hoary fox in Brazil. Canid News 3, Beisiegel, B.M., Ades, C., The behavior of the bush dog (Speothos venaticus Lund, 1842) in the field: a review. Revista de Etologia 4, Deutsch, L.A., An encounter between bush dog (Speothos venaticus) and paca (Agouti paca). Journal of Mammalogy 64, Emmons, L.H., Feer, F., Neotropical Rainforest Mammals: A Field Guide. University of Chicago Press, Chicago, USA Langguth, A., Gray foxes. American Life Encyclopedia 12, 267 and 279 Lund, P.W., Fortsatte bernaerkninger over Brasiliens uddöde dirskabning. Lagoa Santa d. 27 de Marts 1840 Kongelige Danske Videnskabernes Selskab Afhandlinger 9, 1 16 Malcom, J.R., Estimation of mammalian densities in continuous forest north of Manaus. In: Gentry, A.H. (Ed.), Four Neotropical Rainforests. New Haven, Yale, University Press, pp Peres, C.A., Observations of hunting by small-eared (Atelocynus macrotis) and bush dogs (Speothos venaticus) in central-western Amazonia. Mammalia 55, Silva Jr., J.S., Soares, M.C.P., An unexpected new record for the bush dog, Speothos venaticus Lund, 1842, in the Brazilian Amazonia (Carnivora, Canidae. Publicações Avulsas do Instituto Pau Brasil de História Natural 2, 7 11 Silveira, L.A., Jacomo, A.T.A., Rodrigues, F.H.G., Diniz-Filho, J.A.F., Bush dogs (Speothos venaticus) in Emas National Park, central Brazil. Mammalia 62, Voss, R.S., Emmons, L.H., Mammalian diversity in Neotropical lowland rainforests: a preliminary assessment. Bulletin American Museum of Natural History 230, Colombia Bates, M., Notes on captive Icticyon. Journal of Mammalogy 25, Defler, T.R., A bush dog (Speothos venaticus) pack in the eastern llanos of Columbia. Journal of Mammalogy 67, Hershkovitz, P., A synopsis of wild dogs in Colombia. Novedades Colombianas 3, Costa Rica Ecuador de la Rosa, C.L., Nocke, C.C., A Guide to the Carnivores of Central America. University of Texas Press, Austin, Texas Ginsberg, J.R., Macdonald, D.W., Foxes, wolves, jackals, and dogs: an action plan for the conservation of Canids. IUCN/SSC Canid Specialist Group, Gland, Switzerland and Cambridge, UK Tirira, D. (Ed.), Libro rojo de los mamiferos del Ecuador. SIMBIOE/Ecociencial Ministerio del Ambientel/IUCN, Serie Libros Rojos del Ecuador, Tomo 1. Publicación especial sobre los mamiferos del Ecuador 4, Quito, Ecuador Guyana Barnett, A., Shapley, R., Engstrom, M., Records of the bush dog, Speothos venaticus (Lund, 1842), from Guyana. Mammalia 65, Engstrom, M.D., Lim, B.K., Diversity and conservation of mammals of Guyana. In: Cabellos, G.Y., Simonetti, J. (Eds.). Diversidad y conservacion de los mamiferos neotropicales. CONABIO & UNAM, México. Parker, T.A. III, Foster, R.B., Emmons, L.B., Freed, P., Forsyth, A.B., Hoffman, B., Gill, B.D., A biological assessment of the Kanuku Mountains region of southwestern Guyana. Conservation International, Rapid Assessment Program (RAP) Working Papers, No. 5, pp Quelch, J.J., Animal life in British Guiana. Argosy Publishers, Ltd., Georgetown ter Steege, H., Boot R.G.A., Brouwer, L.C., Caesar, J.C., Ek, R.C., Hammond, D.S., Haripersaud, P.P., Van Der Hout, P., Jetten, V.G., Van Kekem, A.J., Kellman, M.A., Khan, Z., Polak, A.M., Pons, T.L., Pulles, J., Raaimakers, D., Rose, S.A., Van Der Sanden, J.J., Zagt, R.J., Ecology and logging in tropical rain forest in Guyana, with recommendations for forest management. Tropenbos Series No. 14. The Tropenbos Foundation, Wageningen, The Netherlands Paraguay Beccaceci, M.D., Bush dogs in Paraguay. Canid News 2, 17 DeMatteo, K.E., Carrillo, O., Zuercher, G.L., Ramírez, S., Smith, K., Porton, I.J., A technique for attracting bush dogs Speothos venaticus in the wild. Canid News 7.6 (Online), 1 12 ITAIPÚ Binacional, Contributión al conocimiento del Speothos venaticus, Lund, 1842 (Carnivora, canidae), 1 20 Ventre, M.M., May Detectaron Jagua Yvyguy en la Reserva Mbaracayú Un Canido en Serio Peligro de Extincion. Ecologia Suplemento Rural del Diario ABC Color Zuercher, G.L., Villalba, R.D., Records of Speothos venaticus Lund, 1842 (Carnivora, Canidae) in eastern Paraguay. Zeitschrift für Säugetierkunde 67, 1 3 Peru Alverson, W.S., Rodriguez, L.O., Moskovits, D.K. (Eds.), Peru: Biabo Cardillera Azul. Rapid Biological Inventories. The Field Museum, Chicago, USA Aquino, R., Puertas, P., Observaciones preliminares sobre la ecological de Speothos venaticus (Canidae: Carnivors) en su habitat natural. Folia Amazonica 8, Aquino, R., Puertas, P., Observations of Speothos venaticus (Candae: Carnivora) in its natural habitat in Peruvian Amazonia. Zeitschrift für Säugetierkunde 62,

10 BIOLOGICAL CONSERVATION 141 (2008) Appendix A continued Country Reference Grimwood, I.R., Notes on the distribution and status of some Peruvian mammals. Special Publication Number 21, American Committee for International Wild Life Protection and New York Society, Bronx, NY, USA INRENA (Instituto Nacional de Recursos Naturales), SINANPE (Sistema Nacional de Área Naturales Protejidas por el Estado). Ministerio de Agricultura, Peru Terborgh, J.F., Fitzpatrick, J., Lista preliminary de las aves and grandes mamíferos en los alrededores de la Estación Biológica de Cocha Cashu. Centro de Datos para la Conservación Reporte Manú, 1 12 Suriname Venezuela Husson, A.M., The mammals of Suriname. E.J. Brill, Leiden, The Netherlands Paramaribo newspaper De West 9 May 1953 (In: Husson (1978)) Sanderson, I.T., A brief review of the mammals of Suriname (Dutch Guiana), based on a collection made in In: Proceedings of the Zoological Society of London, vol. 199, pp Bisbal, F.J., Distribution and habitat association of the carnivores of Venezuela. In: Eisenberg, J.F., Redford, K.H. (Eds.), Advances in Neotropical Mammalogy. Sandhill Crane Press, Gainesville, pp Mondolfi, E., Fauna silvestre de los bosques humedos tropicales de Venezuela. In: Hamilton, S.I., Steyermark, J., Veillon, J.P., Mondolfi, E. (Eds.), Conservación de los bosques humedos de Venezuela. MARNR, Caracas, Venzuela, pp Ochoa, G.J., Molina, A., Giner, S., Inventario y studio comunitario de los mamiferos del parquet Nacional Canaima, con una lista de las especies registradas para Guyana Venezolana. Acta Cientifica Venezolana 44, Strahl, S.D., Silva, J.L., Goldstein, I.R., The bush dog (Speothos venaticus) in Venezuela. Mammalia 56, 9 13 Appendix B List of the survey respondents (n = 37) for each of the 12 countries in the bush dog s historical range. Type of evidence associated with exact locations (n = 189), those points used in ecological niche modeling and Speothos occurrence relative to protected/unprotected areas, are listed for each respondent with the number of points listed in parentheses. Country Name of respondent Institution Exact AR Dr. Mario S. Di Bitetti Fundacion Vida Silvestre Argentina Sighting (1) AR Mónica V. Pia, Bióloga Sighting (1) AR Miguel Angel Rinas Departamento Fauna y Flora, Ministerio de Ecologia, R.N.R. y T. Sighting (3) Tracks (3) AR Karina Schiaffino Centro de Investigaciones Ecológicas Sighting (2) Subtropicales (CIES) AR Douglas Trent Focus Tours Inc. BO Joanne Devlin Black Diamond Paving Sighting (1) BO Leonardo Maffei Wildlife Conservation Society Proyecto Kaa Iya Sighting (1) BO Museum of Natural History (London) Sighting (1) BO Teresa Pacheco American Museum of Natural History BO Julieta Vargas Museo Nacional de Historia Natural Various (3) BO Robert Wallace Wildlife Conservation Society Camera Trap (2) Hair Collection (1) Interview (54) Sighting (10) Tracks (8) BR Brazil Natural History Museum BR Otávio Borges Maia IBAMA BR Fernanda Michalski Associação Pró-Carnívoros Interview (1) Tracks (1) BR Guilherme Hemnique Braqa de Miranda Policia Federal BR Tadeu de Oliveira Associação Pró-Carnívoros Various (20) a Sighting (2) a BR Flavió Henrique G. Rodrigues Associação Pró-Carnívoros Sighting (1) Tracks (1) BR Manuel Ruedi Natural History Museum of Geneva, Switzerland BR Joaquim de Araujo Silva Instituto Biotrópicos de Pesquisos em Vida Silvestre Interview (1) BR Leandro Silveira Associação Pró-Carnívoros Various (5) Sighting (10) (continued on next page)

11 2504 BIOLOGICAL CONSERVATION 141 (2008) Appendix B continued Country Name of respondent Institution Exact BR Liliani Tiepolo Museo Nacional Universidade Federal do Rio de Janeiro BR Museum of Natural History (London) BR Teresa Pacheco American Museum of Natural History BR Douglas Trent Focus Tours Inc. BR Arnaud Desbiez Durrell Institute for Conservation Ecology University of Kent CO Teresa Pacheco American Museum of Natural History EC Felipe Campos The Nature Conservancy Sighting (3) EC CDC-Ecuador FG Francois Catzeflis Universite Sciences & Techniques Sighting (22) FG Benoit de Thoisy Kwata NGO Sighting (9) Tracks (2) FG Geraldine Veron Muséum National d Histoire Naturelle (Paris) GY Eustace Emerick Alexander Conservation International Guyana Sighting (1) PA N.B. Gale, DVM/MPH Retired Public Health Veterinarian US Department of Defense Sighting (3) PY José L. Cartes Asociación Guyra Paraguay Sighting (1) Vocalizations (1) Mandible (1) PE Major L. Boddicker, Ph.D. Rocky Mountain Wildlife Services Sighting (2) PE Teresa Pacheco American Museum of Natural History SR Museum of Natural History (London) VE Francisco J. Bisbal E. Ministerio del Ambiente y los Recursos Naturales Sighting (11) a Since survey response, data submitted for publication: Oliveira, T.G., in press. Distribution, habitat utilization, and conservation of the bush dog Speothos venaticus in northern Brazil. Oryx. REFERENCES Anderson, R.P., Lew, D., Peterson, A.T., Evaluating predictive models of species distributions: criteria for selecting optimal models. Ecological Modelling 162, Araújo, M.B., Guisan, A., Five (or so) challenges for species distribution modelling. Journal of Biogeography 33, Barnett, A., Shapley, R., Engstrom, M., Records of the bush dog, Speothos venaticus (Lund, 1842), from Guyana. Mammalia 65, Beccaceci, M.D., Bush dogs in Paraguay. Canid News 2, 17. Beisiegel, B.M., Contribuição ao estudo de história natural do cachorro do mato, Cerdocyon thous, e do cachorro vinagre, Speothos venaticus. Ph.D. Thesis. Instituto de Psicologia - Universidade de São Paulo, SP, Brazil. Defler, T.R., A bush dog (Speothos venaticus) pack in the eastern llanos of Columbia. Journal of Mammalogy 67, DeMatteo, K.E., Kochanny, C.O., Determining an effective and safe radio-tracking collar for bush dogs (Speothos venaticus). Canid News 7.7, 1 5 (Online). DeMatteo, K.E., Carrillo, O., Zuercher, G.L., Ramirez, S., Smith, K., Porton, I.J., A technique for attracting bush dogs (Speothos venaticus) in the wild. Canid News 7.6, 1 12 (Online). Deutsch, L., An encounter between bush dog (Speothos venaticus) and paca (Agouti paca). Journal of Mammalogy 64, Draper, D., Rossello-Graell, A., Garcia, C., Tauleigne Gomes, C., Sergio, C., Application of GIS in plant conservation programmes in Portugal. Biological Conservation 113, Eisenberg, J.F., Mammals of the Neotropics. The Northern Neotropics: Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, vol. 1. University of Chicago Press, Chicago, Illinois. Eisenberg, J.F., Redford, K.H., Mammals of the Neotropics. The Central Neotropics Ecuador, Peru, Bolivia, Brazil, vol. 1. University of Chicago Press, Chicago, Illinois. pp Elith, J., Graham, C.H., Anderson, R.P., Dudik, M., Ferrier, S., Guisan, A., Hijmans, R.J., Huettmann, H., Leathwick, J.R., Lehmann, A., Li, J., Lohmann, L.G., Loiselle, B.A., Manion, G., Moritz, C., Nakamura, M., Nakazawa, Y., Overton, J.C., Peterson, A.T., Phillips, S.J., Richardson, K., Scachetti-Pereira, R., Schapire, R.E., Soberón, J., Williams, S., Wisz, M.S., Zimmermann, N.E., Novel methods improve prediction of species distributions from occurrence data. Ecography 29, ESRI, Inc., ESRI ArcGIS Ò 9.1. Redlands, California.. Feria, A.T.P., Understanding the geographic distribution of species: An evaluation of different methods for modeling

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