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available at www.sciencedirect.com
journal homepage: www.elsevier.com/locate/biocon
New data on the status and distribution of the bush dog
(Speothos venaticus): Evaluating its quality of protection
and directing research efforts
Karen E. DeMatteoa,b,c,*, Bette A. Loisellec
a
Saint Louis Zoo, Research Department, One Government Drive, St. Louis, MO 63110, USA
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
b
A R T I C L E I N F O
A B S T R A C T
Article history:
The bush dog (Speothos venaticus), listed as CITES Appendix 1 – vulnerable, is a small (5–
Received 21 December 2007
6 kg), rarely seen canid from Central and South America. The World Conservation Union
Received in revised form
Canid Specialist Group (IUCN CSG) recommended that research with this species focus
2 July 2008
on their basic ecology so that a data-driven conservation strategy can be formulated. Infor-
Accepted 11 July 2008
mation on the bush dog, however, has been lacking since standard field techniques have
Available online 23 August 2008
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
Keywords:
conservation status in the wild, its current distribution, and identify ecological needs by
Carnivore
correlating habitat types to bush dog sightings. Survey responses and literature generated
Conservation
a database with 399 historic bush dog locations recorded between 1834 and 2004. These
GIS modeling
locational records were used to update the species’ range extent map, an important conser-
MAXENT
vation planning tool. With year and precision of data accounted for, we analyzed land use
Survey
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.
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; Sil-
veira 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.: +1 314 516 6577; fax: +1 314 516 6233.
E-mail addresses: [email protected] (K.E. DeMatteo), [email protected] (B.A. Loiselle).
0006-3207/$ - see front matter 2008 Elsevier Ltd. All rights reserved.
doi:10.1016/j.biocon.2008.07.010
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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
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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 (www.lib.utexas.edu/maps/americas). 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 (www.canids.org).
The Survey was sent (i.e., via mail or email) 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.
2.2.
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 www.calle.com/
world; NIMA, www.nima.mil/gns/html;maps) 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).
2.3.
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 (http://edcdaac.usgs.gov/glcc/
sa_int.html and http://edcdaac.usgs.gov/glcc/na_int.html)
that characterized seasonal land coverage into 166 habitat
types. Both land cover databases were generated by remotesensed data collected at a 1-km2 resolution between April
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1992 and March 1993. Bush dog locations were rasterized to
1-km2 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 ArcGIS9 (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 (4.56–
4.72 km2, Silveira et al., 1998; 20 km2, Van Humbeck and Perez,
1998, 50–100 km2 (2–6 individuals), Beisiegel, 1999) and one
based on minimum convex polygon with systematic data collection on a single radio collared family group (100 km2, 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 km2 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-
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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 km2 and 5000 km2, respectively.
2.4.
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 (http://biogeo.berkeley.edu/gis/data.
html; Hijmans et al., 2005), which contains a minimum of
30 years (1960–1990) of monthly temperature (C) and precipitation (mm) measurements at a 30 arc-second resolution (approx. 1-km2). We selected eight of the 19 bioclimatic variables
and aggregated these data to an approximate 5-km2 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, MAXENT 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 0.00001, a maximum of 10,000 back-
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ground 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.
2.5.
areas
Speothos occurrence relative to protected/unprotected
Two databases, the 2003 World Database on Protected Areas
(WDPA03 – www.biodiversity.org/wcpa) and PróCarnı́voros, 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 PróCarnı́voros 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 PróCarnı́voros 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 km2 (25 groups each requiring 100 km2)
or 5000 km2 (50 groups each requiring 100 km2). 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.
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3.
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).
3.2.
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.
3.3.
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
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(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 km2 for 25 groups versus 145,679 km2 for 50
groups).
3.4.
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 km2, 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).
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3.5.
areas
Speothos occurrence relative to protected/unprotected
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 (1992–1993). 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, expo-
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sure 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.
4.1.
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 km2/group
No. exact locations
No. polygons
10 Groups
0% Protected
All Countries
Argentina
Bolivia
Brazil
Brazil*
Ecuador
French Guiana
Guyana
Panama
Paraguay
Peru
Venezuela
256
8
81
58
58
3
33
9
3
26
8
27
927
82
50
390
2067
87
10
1
51
21
74
161
31
12
23
45
22
0
61
89
0
4
0
7
25 Groups
100% Protected
2
0
0
0
4
0
0
0
0
0
0
15
0% Protected
100% Protected
25
12
14
43
17
0
42
78
0
4
0
7
1
0
0
0
4
0
0
0
0
0
0
7
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 PróCarnı́voros database that is marked with an asterisk (*), results were based on analyses of
the WDPA03 database.
2500
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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 NS 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
1 4 1 ( 2 0 0 8 ) 2 4 9 4 –2 5 0 5
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 Piauı́, 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 NS 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
B I O L O G I C A L C O N S E RVAT I O N
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.
1 4 1 ( 2 0 0 8 ) 2 4 9 4 –2 5 0 5
2501
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
PróCarnı́voros – Associação para Conservação dos Carnı́voros
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 Ecologı́a 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 (Associação PróCarnı́voros), Guilherme Hemnique Braqa de Miranda (Policia Federal), Manual Ruedi (Natural History Museum of Geneva, Switzerland), Joaquim de
Araujo Silva (Instituto Biotrópicos de Pesquı́sos 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
Acknowledgements
We thank the IUCN CSG who endorsed the project and provided web access to the survey, Luis Padilla and Tadeu Gomes
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., 1990. 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., 1999. Aportes al conocimiento de la distribución de los carnivoros del
Noroeste de Argentina. Mastozoologia Neotropical 6, 15–30
Bolivia
Anderson, A., 1997. Mammals of Bolivia, taxonomy and distribution. Bulletin American Museum of Natural History 231, 1–
652
Emmons, L.H., 1998. 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. 129–135
FAN-WCS, 1994. 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., 1998. Evaluacción de la diversidad de mamiferos medianos y grandes en el Parque Nacional
Carraso (Cochabamba-Bolivia). Revista Boliviana de Ecologia y Conservación Ambiental 4, 77–90
Wallace, R.B., Painter, R.L.E., Saldania, A.S., 2002. An observation of bush dog (Speothos venaticus) hunting behavior.
Mammalia 66, 309–311
(continued on next page)
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1 4 1 ( 2 0 0 8 ) 2 4 9 4 –2 5 0 5
Appendix A – continued
Country
Reference
Brazil
Berta, A., 1984. The Pleistocene bush dog, Speothos pacivorus (Canidae) from the Lagoa Santa Caves, Brazil. Journal of
Mammalogy 65, 549–559
Fonseca, G.A.B, Redford, K.H., 1984. 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, 517–523
Dalponte, J.C., 1988. 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., 1995. The hoary fox in Brazil. Canid News 3, 23–24
Beisiegel, B.M., Ades, C., 2002. The behavior of the bush dog (Speothos venaticus Lund, 1842) in the field: a review.
Revista de Etologia 4, 17–24
Deutsch, L.A., 1983. An encounter between bush dog (Speothos venaticus) and paca (Agouti paca). Journal of
Mammalogy 64, 532–533
Emmons, L.H., Feer, F., 1990. Neotropical Rainforest Mammals: A Field Guide. University of Chicago Press, Chicago, USA
Langguth, A., 1972. Gray foxes. American Life Encyclopedia 12, 267 and 279
Lund, P.W., 1842. Fortsatte bernaerkninger over Brasiliens uddöde dirskabning. Lagoa Santa d. 27 de Marts
1840 Kongelige Danske Videnskabernes Selskab Afhandlinger 9, 1–16
Malcom, J.R., 1990. Estimation of mammalian densities in continuous forest north of Manaus. In: Gentry, A.H. (Ed.),
Four Neotropical Rainforests. New Haven, Yale, University Press, pp. 339–357
Peres, C.A., 1991. Observations of hunting by small-eared (Atelocynus macrotis) and bush dogs (Speothos venaticus) in
central-western Amazonia. Mammalia 55, 635–639
Silva Jr., J.S., Soares, M.C.P., 1999. 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., 1998. Bush dogs (Speothos venaticus) in Emas National Park,
central Brazil. Mammalia 62, 446–449
Voss, R.S., Emmons, L.H., 1996. Mammalian diversity in Neotropical lowland rainforests: a preliminary assessment.
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Colombia
Bates, M., 1944. Notes on captive Icticyon. Journal of Mammalogy 25, 152–154
Defler, T.R., 1986. A bush dog (Speothos venaticus) pack in the eastern llanos of Columbia. Journal of Mammalogy 67, 421–422
Hershkovitz, P., 1957. A synopsis of wild dogs in Colombia. Novedades Colombianas 3, 157–161
Costa Rica de la Rosa, C.L., Nocke, C.C., 2002. A Guide to the Carnivores of Central America. University of Texas Press, Austin, Texas
Ecuador
Ginsberg, J.R., Macdonald, D.W., 1990. 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.), 2001. 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., 2001. Records of the bush dog, Speothos venaticus (Lund, 1842), from Guyana. Mammalia 65,
232–237
Engstrom, M.D., Lim, B.K., 2002. 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., 1993. A biological assessment of the
Kanuku Mountains region of southwestern Guyana. Conservation International, Rapid Assessment Program (RAP) Working
Papers, No. 5, pp. 1–70
Quelch, J.J., 1901. Animal life in British Guiana. Argosy Publishers, Ltd., Georgetown
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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,
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Series No. 14. The Tropenbos Foundation, Wageningen, The Netherlands
Paraguay
Beccaceci, M.D., 1994. Bush dogs in Paraguay. Canid News 2, 17
DeMatteo, K.E., Carrillo, O., Zuercher, G.L., Ramı́rez, S., Smith, K., Porton, I.J., 2004. A technique for attracting bush dogs Speothos
venaticus in the wild. Canid News 7.6 (Online), 1–12
ITAIPÚ Binacional, 1988. Contributión al conocimiento del Speothos venaticus, Lund, 1842 (Carnivora, canidae), 1–20
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Zeitschrift für Säugetierkunde 67, 1–3
Peru
Alverson, W.S., Rodriguez, L.O., Moskovits, D.K. (Eds.), 2001. Peru: Biabo Cardillera Azul. Rapid Biological Inventories.
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Appendix A – continued
Country
Reference
Grimwood, I.R., 1969. 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), 1999. SINANPE (Sistema Nacional de Área Naturales Protejidas
por el Estado). Ministerio de Agricultura, Peru
Terborgh, J.F., Fitzpatrick, J., 1985. Lista preliminary de las aves and grandes mamı́feros en los alrededores de la Estación
Biológica de Cocha Cashu. Centro de Datos para la Conservación – Reporte Manú, 1–12
Suriname
Husson, A.M., 1978. The mammals of Suriname. E.J. Brill, Leiden, The Netherlands
Paramaribo newspaper ‘‘De West’’ 9 May 1953 (In: Husson (1978))
Sanderson, I.T., 1949. A brief review of the mammals of Suriname (Dutch Guiana), based on a collection made in 1938.
In: Proceedings of the Zoological Society of London, vol. 199, pp. 755–789
Venezuela
Bisbal, F.J., 1989. Distribution and habitat association of the carnivores of Venezuela. In: Eisenberg, J.F., Redford,
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Mondolfi, E., 1977. Fauna silvestre de los bosques humedos tropicales de Venezuela. In: Hamilton, S.I., Steyermark, J.,
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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 asso-
Country
Name of respondent
ciated 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.
Institution
Exact
AR
AR
AR
Dr. Mario S. Di Bitetti
Mónica V. Pia, Bióloga
Miguel Angel Rinas
Fundacion Vida Silvestre Argentina
–
Departamento Fauna y Flora, Ministerio de
Ecologia, R.N.R. y T.
Sighting (1)
Sighting (1)
Sighting (3)
AR
Karina Schiaffino
AR
Douglas Trent
Centro de Investigaciones Ecológicas
Subtropicales (CIES)
Focus Tours Inc.
BO
BO
BO
BO
BO
BO
Joanne Devlin
Leonardo Maffei
–
Teresa Pacheco
Julieta Vargas
Robert Wallace
Black Diamond Paving
Wildlife Conservation Society – Proyecto Kaa Iya
Museum of Natural History (London)
American Museum of Natural History
Museo Nacional de Historia Natural
Wildlife Conservation Society
Sighting (1)
Sighting (1)
Sighting (1)
–
Various (3)
Camera Trap (2)
Hair Collection (1)
Interview (54)
Sighting (10)
Tracks (8)
BR
BR
BR
–
Otávio Borges Maia
Fernanda Michalski
Brazil Natural History Museum
IBAMA
Associação Pró-Carnı́voros
BR
BR
Guilherme Hemnique Braqa de Miranda
Tadeu de Oliveira
Policia Federal
Associação Pró-Carnı́voros
BR
Flavió Henrique G. Rodrigues
Associação Pró-Carnı́voros
BR
BR
BR
Manuel Ruedi
Joaquim de Araujo Silva
Leandro Silveira
Natural History Museum of Geneva, Switzerland
Instituto Biotrópicos de Pesquisos em Vida Silvestre
Associação Pró-Carnı́voros
–
–
Interview (1)
Tracks (1)
–
Various (20)a
Sighting (2)a
Sighting (1)
Tracks (1)
–
Interview (1)
Various (5)
Sighting (10)
(continued on next page)
Tracks (3)
Sighting (2)
–
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1 4 1 ( 2 0 0 8 ) 2 4 9 4 –2 5 0 5
Appendix B – continued
Country
Name of respondent
Institution
Exact
BR
BR
BR
BR
BR
Liliani Tiepolo
–
Teresa Pacheco
Douglas Trent
Arnaud Desbiez
Museo Nacional Universidade Federal do Rio de Janeiro
Museum of Natural History (London)
American Museum of Natural History
Focus Tours Inc.
Durrell Institute for Conservation
Ecology – University of Kent
–
–
–
–
–
CO
Teresa Pacheco
American Museum of Natural History
–
EC
EC
Felipe Campos
–
The Nature Conservancy
CDC-Ecuador
Sighting (3)
–
FG
FG
Francois Catzeflis
Benoit de Thoisy
Universite Sciences & Techniques
Kwata NGO
FG
Geraldine Veron
Muséum National d’Histoire
Naturelle (Paris)
Sighting (22)
Sighting (9)
Tracks (2)
–
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
PE
Major L. Boddicker, Ph.D.
Teresa Pacheco
Rocky Mountain Wildlife Services
American Museum of Natural History
Sighting (2)
–
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.
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