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Pigs, Peccaries and
Hippos Status Survey and Action Plan (1993) Chapter 2.3 The White-lipped Peccary (Tayassu pecari) Ignacio J. March Status And Action Plan Summary Status categories 2 - 4 and 'indeterminate',
depending on subspecies/population. The White-lipped peccary is not considered seriously
threatened over much of its extensive range. However, the available data
suggest that most of the remaining populations of two subspecies, T. p. ringens and T. p. spiradens, of southern Mexico and Central America are
threatened to varying degrees, whilst the former population of T. p. ringens in El Salvador is
already extinct. Recent data is lacking on the current distribution and
status of T. p. equatorius, which
has much the smallest range of the five currently recognised forms, being
apparently confined to the remaining lower altitude forests west of the
Cordillera Real in south-western Colombia and north-western Ecuador. Recent
information is lacking on the distribution and status of white-lipped peccaries
in many other continental South American countries, the main stronghold for
this species. However, some reports indicate significant, recent population
declines in parts of the Amazon Basin and in the dry Chaco of south-eastern
Bolivia, western Paraguay and north-western Argentina. Peccary populations in
these areas may now be discontinuously distributed even in virgin forest. The
species is also reported to have become extinct in Uruguay about 100 years
ago, though it is probably not seriously threatened throughout much of its
range elsewhere in central and northern South America at the present time. The destruction and fragmentation of habitat is
undoubtedly the most important threat to this species, though
overexploitation by subsistence and commercial hunters has also contributed
to its decline in many areas. Priority actions for the white-lipped peccary
include: a) assessments of its current distribution and status in each
country, with a view to the development of management plans for the most threatened
populations and subspecies and the strengthening of those protected areas
which maintain important habitat and populations; b) promotion of field
studies on its habitat requirements, its ranging behaviour and its role as a
seed predator/dispersal agent; c) development of projects to encourage
self-regulation by subsistence hunters and to control or curtail large scale
commercial hunting and trade; d) the development of training courses and
workshops on research and management of peccaries; and e) a thorough review
of its generic and subspecific taxonomy at the genetic, as well as the
phenotypic, levels. Introduction The white-lipped peccary has played an important
role in the economic and cultural development of many indigenous peoples in
the neotropics and still constitutes an important resource for various tribal
and rural groups, both as a source of food and income (Olsen, 1982; Donkin,
1985). Ecologically, it is also important, since it is the only large,
terrestrial mammal in the neotropics to form large herds, often of more than
100 individuals. In addition, it has a large range, which includes or extends
into 19 Latin American countries. There have been relatively few studies conducted on
this species, and many aspects of its behaviour, ecology, habitat
requirements, population biology and status remain poorly known. However,
Sowls (1984) has provided a useful summary of most of the available
information, much of which originates from the studies of Kiltie (1980, 1981,
1982, 1983) and Kiltie & Terborgh (1983) of the species' ecology in the
rainforests of the Peruvian Amazon. Barreto & Hern ndez (1988) have
also undertaken a study of its feeding habits and behaviour in tropical
rainforest in Venezuela, and an evaluation of the species' status and habitat
in southern Mexico was conducted recently (March, 1990). The social behaviour of white-lipped peccaries is
similar to that of T. tajacu in
most respects, except for its tendency to form large herds. In common with T. tajacu, they have a rich repertoire
of vocalizations which have a role in agonistic and reproductive behaviour,
and also possibly in the cohesion of herds in dense vegetation. They use
smell to communicate and engage in reciprocal rubbing of the dorsal scent
gland, as does the collared peccary. Unlike the latter species, however,
white-lipped peccaries have not been observed marking their territories with
their scent glands, at least in captivity. Captive animals reach sexual maturity between one
and two years of age. Although distinct reproductive seasons have been
suggested, it is probable that these peccaries breed throughout the year,
with a peak in the most favourable months. In Peru, Kiltie and Terborgh
(1976) observed mating during July and August, whilst in Chiapas, Mexico,
mating has been recorded from April to November (March & Cuar¢n, 1987).
Roots (1966) recorded a gestation period of 156 to 162 days in captive
animals. As in the collared peccary (Byers & Beckoff, 1981), the modal
litter size is two, and it is probable that the dominant males are
responsible for most litters. Little work has been done on the genetics and
geographical variation of T. pecari,
and a thorough review of its systematics is required to enable a proper
re-assessment of its generic and subspecific taxonomy (see Grubb and Groves,
1993). Five subspecies are recognised at present, but the validity of some of
these are in question. The ranges of these subspecies (Fig. 2) are as
follows: Link to Fig. 2: Approximate
former distribution of the white-lipped peccary, Tayassu pecari ssp. Former and Present Distribution White-lipped peccaries are confined to the
Neotropical Region, from south-eastern Mexico in the north, through Central
America and northern and central South America, as far south as Entre Rios in
northern Argentina and Rio Grande do Sul in southern Brazil (Sowls, 1986).
Over this range it inhabits lowland forests at sea level to lower montane
forests at maximum recorded altitude of 1,900 m on the eastern slopes of the
Andes in Peru (Osgood, 1914). The species has also been reported at altitudes
of 1,500 m in Venezuela (Röhl, 1959), Panamá (Anthony, 1916), and in Atitlan,
Guatemala (Alston, 1879), although there have been no recent, verified
records at altitudes of <800 m. Its distribution is somewhat more restricted than
that of the more adaptable collared peccary, T. tajacu, with which it is almost everywhere sympatric, but it
has a much larger range than the Chacoan peccary, Catagonus wagneri, with which it is also sympatric over the
latter species' restricted distribution in the Gran Chaco. Recent data on the distribution and status of T. pecari are lacking, or inadequate
for many of the 19 countries in which the species has been recorded.
Nonetheless, it is clear that its former range has been severely reduced and
fragmented during recent decades. Available data on its present range in each
country/region, and the status of the five currently recognised subspecies may
be summarised as follows: 1 T. p.
ringens: southern Mexico (Oaxaca, Chiapas and Yucatan Peninsulas),
Guatemala, Belize, El Salvador (extinct), Honduras and northern Nicaragua. In Mexico this subspecies has probably been
extirpated from the more northerly states of Veracruz and Tabasco, though its
continued presence has been confirmed in a total of over 1.5 million ha of
protected forest in the states of Oaxaca, Chiapas, Campeche and Quintana Roo.
Within this combined area, the most important populations are in the proposed
biosphere reserve of Los Chimalapas (>400,000 ha) in Oaxaca, the Lacandona
Rainforest (which includes the Montes Azules Biosphere Reserve, >300,000
ha) in Chiapas, and in the recently established Calakmul Biosphere Reserve
(>700,000 ha) in Campeche State. A few herds may also persist in the El
Ocote Reserve (35,000 ha) and in the region of La Sepultura, Chiapas, and in
the Sian Ka'an Reserve in north Quintana Roo State, though the future of
these populations is uncertain. The neighbouring Mayan Reserve in the
Department of El Peten in Guatemala is, perhaps, now the single most
important area for this subspecies. It is also thought to occur in one or
more reserves elsewhere in Guatamala, and in some reserves in Belize,
Honduras and northern Nicaragua, but recent data from these areas are lacking
at present. White-lipped peccaries, formerly of this subspecies, are now
extinct in El Salvador. 2) T. p.
spiradens: southern Nicaragua, Costa Rica, Panama and north-western
Colombia (areas west of the Cordillera Central and Cordillera Oriental). Both T. p.
ringens (north only) and T. p.
spiradens occur in Nicaragua, where a total area of 1,451 km2 of largely
undisturbed rainforest are currently protected (FAO/PNUMA, 1989). Nicaragua
is therefore a potentially important stronghold for this species in Central
America, though their present status, and the consequences of this country's
years of strife and guerilla warfare on their populations, are not known. In
Costa Rica, the subspecies has been recorded in the Corcovado National Park
(43,735 ha ; pers. obs.) in the Osa Peninsula, and in the Oros¡ Reserve
Forest (10,147 ha), Ca¤o Negro Wildlife Refuge (9,969 ha) and the Tortuguero
National Park (18,947 ha) (A. Carr, pers. comm.). It probably also occurs in
La Amistad Biosphere Reserve, which extends across the southern border into
Panama. However, its status in the latter area, and in other reserves in
Panama is unknown, though the subspecies is now extinct on Barro Colorado
Island. Information on the present distribution and status of these animals
in Colombia is also lacking. 3) T. p.
equatorius: south-western Colombia and north-western Ecuador (west of the
Cordillera Real) only. This subspecies is reported to occur only in the
south-west corner of Colombia, in Cauca and Narino Provinces, and in the
neighbouring provinces of Esmareldas, Manabi and Pinchicha in north-western
Equador. It therefore has by far the smallest range of any of the currently
recognised forms of T. pecari, but
its present status in these areas is not known. 4/5) T. p. pecari: eastern Colombia, Venezuela,
British Guiana, Suriname, French Guina and areas north of the Amazon River in
Brazil. T. p. albirostris: eastern Peru, eastern Bolivia,
Paraguay, northern Argentina (south to Santiago del Estero and Entre Rios)
and (formerly) Uruguay, to central and south-eastern Brazil. The Amazon Basin remains by far the most important
stronghold for this species, though the increasing destruction and
fragmentation of this region are the focus of international attention and
concern; not least because only 4.28% of this region had been declared as
protected areas. Nonetheless, large populations of this species still exist
in this region, and it occurs in most of the larger and many of the smaller
parks and reserves on both sides of the Amazon River. The latter also
represents the approximate southern and northern limits of the respective
distributions of these two subspecies, neither of which can be regarded as
seriously threatened over all of their extensive ranges at present, though
the southern subspecies, T. p.
albirostris, is extinct is Uruguay where it was last recorded in 1890 (C.
Gastelmundi, pers. comm. to W. Oliver). The approximate distribution and current status of
the five subspecies are summarized in Fig. 2 and Table 3. |
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Table 3 Status of T. pecari
populations according to the categories adopted by the IUCN/SSC Pigs and
Peccaries Specialist Group. ______________________________________________________________________________________ Subspecies Country/(Region) Status Category T. p. ringens Mexico (El Ocote Reserve, north of Quintana Roo, & Sian Ka'an
Reserve) 5 T. p. ringens Mexico (Lacandona Rainforest, Chiapas; Oaxaca and Campeche) 3 - 4 T. p. ringens Guatemala (Peten)
2 - 3 T. p. ringens Belize, Honduras, Nicaragua (north)
Indeterminate T. p. ringens El Salvador
Extinct T. p. spiraden Nicaragua Indeterminate T. p. spiradens Costa Rica (north)
5 T. p. spiradens Costa Rica (south)
4 T. p. spiradens Panama
Indeterminate T. p. spiradens Colombia (north-west)
Indeterminate T. p. equatorius Colombia (south-west), Ecuador (north-west) Indeterminate T. p. pecari Colombia (east), Venezuela,
Guianas, Surinam, Brazil (north) 1 - 2 ? T. p. albirostris Brazil, Peru (east), Bolivia,
Paraguay, Argentina (north) 1 - 2 ? Uruguay (north only ?) Extinct ? = Insufficient information. |
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Habitat, Ecology and Behavior There have been erlatively
few studies conducted on this species, and many aspects of its behavior,
ecology, habitat requirements, population biology and status remain poorly
known. HOwever, Sowls (1984) has provided a useful summary of most of the
available information, much of which originates from the studies of Kiltie
(1980, 1981 a, b, c, 1982) and Kilte and Terborgh (1983) on the ecology of
the speciesin the rainforests of the Peruvian Amazon. Baretto and Hernández
(1988) have also undertake a study of its feeding habits and behavior in
tropicalrainforest in Venezuela, and an evaluation of the species' status and
habitat in southern Mexico was conducted recently (March 1990). The majority of habitat records for the species are
from humid tropical forests, though it is also occurs in various drier
habitats; the latter including dry savannas in Venezuela, the xerophitic
areas of the Chaco and in the tropical dry forest of Costa Rica (Wetzel &
Lovett, 1974; Mayer & Brandt, 1982; Vaughan, 1983; Sowls, 1984; D. Janzen
and S. Cornelius, pers. comm.). According to Sowls (1984) T. pecari is restricted to a narrower
climatic range than T. tajacu. The
optimal habitat for this species appears to be lowland rainforests,
especially in well conserved areas (>30,000 ha) where it reaches its
greatest abundancy. Conversely, its tolerance to deforested areas appears to
be minimal, as it is easily hunted out where it no longer has sufficient
cover. White-lipped peccaries usually frequent areas close
to water, and will even visit coastal beaches to forage. They are omnivorous,
but have a strong tendency towards frugivory (Husson, 1978; Kiltie, 1981b,
1981c). Their diet is comprised mostly of fruits, seeds and roots, but they
will also take invertebrates, small vertebrates, fungi and carrion. More than
40 plant species have been recorded in their diet (March, in press), and the
species may play an important role in the ecology of Neotropical forests as a
major dispersor and predator of seeds - this role being reminiscent of, and
perhaps analogous to, that of bearded pigs, Sus barbatus, in the tropical
forests of South-east Asia (Caldecott et
al., 1993). Its tendency to form large herds may also have an important
consequences on the composition of ground plant communities and the drainage
of surface water merely as a result of their intense rooting activity and the
compaction of soil caused by the passage of large herds. White-lipped peccary herds often exceed 100
individuals, though groups of as few as 5 to more than 200 individuals have
also been observed (Mayer & Brandt, 1982; March, pers. obs.). It is
possible that in certain seasons, large herds divide into smaller groups
according to the distribution and abundance of food, though the more frequent
reporting of smaller groups in some areas is probably correlated with
increased hunting pressure. The adaptive significance of forming large herds in
this species is not clearly understood, though Kiltie & Terborgh (1983)
have suggested it is primarily a predator defence strategy. In any event, the
formation of these large herds also presupposes the need to preserve
relatively extensive and continous tracts of habitat, particularly as the
movements of these herds are likely to be strongly influenced by the pattern
of food dispersion (Sowls, 1984). Kiltie & Terborgh (1983) estimated that
in the Amazon region of Peru, these herds travelled up to 10 km per day, were
able to cross wide rivers without difficulty, and had a home range of 60 to
200 sq. km. From calculations based on hunting data in this region, Bodmer et
al. (1988) obtained a local density for the species of 1.3 individuals per sq.
km, representing a biomass of 43.2 kg/sq. km. In the Matto Grosso of Brazil,
Schaller (1983) estimated a density of 1.6 animals per sq. km. The social behavior of the white-lipped peccaries is
similar to that of T. tajacu in most respects, except for its tendency
to form large herds. In common with T. tajacu, they have a rich
repertoire of vocalizations which have a role in agonistic and reproductive
behavior, and also possibly in the cohesion of herds in dense vegetation.
They use small to communicate and engage in reciprocal rubbing of the dorsal
scent gland, as does the collared peccary. Unlike the latter species,
white-lipped peccaries have not been observed marking their territories with
their scent glands, at least in captivity. Captive animals reach
sexual maturity between one and two years of age. Although distinct
reproductive seasons have been suggested, it is probable that these peccaries
breed throughout the year, with a peak in the most favourable months. In
Peru, Kiltie and Terborgh (1976) observed mating during July and August,
while in Chiapas, Mexico, mating has been recorded from April to November
(March and Cuarón 1987). Roots (1966) recorded a gestation period of 156 to
162 days in captive animals. As in the collared peccary (Byers and Bekoff
1981), the modal litter size is two, and it is probable that the dominant
males are responsible for most litters. Threats to Survival Widespread and increasing deforestation and intense
hunting pressure are the main reasons for the increasing diminution and
extinction of many populations of this species. In most Latin America
countries, the progressive destruction of habitat for agriculture and cattle
ranching, as well as timber extraction, has already accounted for much of the
species' former habitat. It has been calculated that by the end of this
century, Latin American forests will be reduced to roughly 366 million ha,
approximately half of the estimated original forest cover of 693 million ha
(Wolf, 1987). Owing to their habit of forming large herds, white-lipped
peccaries require relatively extensive and continuous areas of habitat in
order to obtain sufficient resources throughout the year. The loss and
fragmentation of their habitat also exposes them to increased hunting
pressure by facilitating their location by hunters, who can kill many
individuals in a large herd during a single encounter. White-lipped peccaries are hunted for sport, as well
as by subsistence and commercial hunters, and they are killed for their hides
as well as for meat. They have been identified as the single most important
source of meat for the Mundurucu tribe in Brazil (Murphy, 1960), the Guayakí
in Paraguay (Cadogan, 1973) and the Mayan Lacandones of Chiapas (March,
1987), as well as for many other indigenous groups who hunt them for purely
subsistence purposes (Hames, 1980). In many countries (e.g. Mexico and
Guatamala) subsistence hunters operate in most reserves and national parks,
and there is no doubt that hunting pressure is increasing over the species'
range as a whole as a result of the continuing immigration of people to
forested areas. The species has also been affected by militarization and war
in several Central and South American countries, where army personnel and
guerilla groups are often heavily dependent on wild game and can kill large
numbers of peccaries, often with automatic weapons. Despite the lack of knowledge about the species,
several countries permit sport hunting of white-lipped peccaries. In Mexico,
for example, sport hunting was permitted until at least 1990 in Campeche
State; whilst in Costa Rica it was possible to get a permit for up to 5
animals per season until 1986. In Argentina and Paraguay, sport hunting is
quite commonplace, and is controlled to only a very limited extent (A. Taber,
pers. comm.). However, sport hunting is of relatively minor significance in
comparison to the large scale commercial exploitation of the species for meat
and, especially, for hides (see Bodmer et
al., 1993). Conservation Measures Taken This species occurs in numerous protected areas
throughout its extensive range, though it is far from certain that the
existing network of protected areas is adequate to ensure the survival of
representative populations of all currently recognised subspecies or of sufficient
size to maintain viable populations of these animals in many of the smaller
nation states, especially in Central America. It is also doubtful if many of
these reserves are large enough to allow the formation of large herds of this
species. Concerns about the large number of peccary hides of
both species of Tayassu in
international trade (Broad, 1984) led to their recent (1986) inclusion on
Appendix II of CITES (Oliver, 1987; Bodmer et al., 1988b, 1993). This measure is of considerable
significance for future regulation and control of this trade, which is
potentially more damaging to T. pecari
than to T. tajacu in terms of the
species' susceptibility to commercial hunting pressures. Captive Breeding White-lipped peccaries are widely maintained in captivity
in their countries of origin, but very rarely elsewhere, mainly because of
the stringent veterinary restrictions placed on the movement of live animals
into countries with a significant domestic pork industry. The species has
been bred infrequently in captivity (Roots, 1966; McDonald and Lasley, 1978;
Frädrich, 1986), and little attempt has been made to develop cooperative
breeding programmes for this species in zoos in their countries of origin,
where replacement stock may be obtained relatively easily. The most notable exceptions to this are the captive
breeding of T. p. albirostris at
the School of Agriculture, University of Sao Paulo in Piracicaba City
(Brazil), the Santa Cruz Zoo (Bolivia), and the West Berlin Zoo (Germany),
and the pioneering programme for the locally threatened subspecies, T. p. ringens in the Regional Zoo of
Chiapas (Mexico). The Sao Paulo University project was started in 1986 and
currently (August 1991) comprises 34 individuals, all of which (including the
founder stock) are captive-bred (S. Nogueira-Filho, pers. comm. to W.
Oliver). The West Berlin stock, which comprised 18 animals in February 1991,
is derived from 5 wild-caught (in Paraguay) founders, obtained at intervals
between 1979 and 1985 (H. Frädrich, pers. comm.). By comparison, the breeding
programme for T. p. ringens in
Chiapas was started in 1985 with 5 wild-caught pairs and comprised a total
stock of 26 individuals by the end of January 1991. On at least two occasions, captive T. pecari and T. tajacu have produced F1 hybrids which were reared to maturity
(Zuckerman, 1953; Sowls, 1984), though it is not known whether these hybrids
were fertile. Additional Remarks Given that this species is a relatively large-bodied
mammal of some ecological importance (as a major seed predator/dispersal
agent), as well as being of considerable socio-economic importance to local
people, it is a potentially ideal 'flagship' species for tropical forest
conservation projects. The fact that it is also the only large terrestrial mammal
to form large herds in neotropical forests, and that it inhabits a broad
spectrum of forest types, is also of significance, since it presupposes the
need to conserve substantial tracts of forest, with all that that implies in
terms of the maintenance of representative diversity in these areas. Thus,
the development of conservation plans targetted on this species should not
only help to ensure the enhanced future management of an important local
resource, but also help to ensure the survival of diverse other, more
immediately threatened species. Conservation Measures Proposed: An Action Plan Although this species is not yet seriously
threatened throughout it extensive range, there is concern about its
continuing decline in some regions. However, the rate, severity and magnitude
of this decline, or what actions may be needed to reverse this trend, are not
always apparent owing to the paucity of accurate information on its present
status and future management needs in most countries. Any actions intended to
clarify this situation must therefore assume a high priority. By the same
token, there are many aspects of its biology which are poorly known at
present, but which are highly relevant to its future conservation, as well as
the wider issues surrounding these animals; the latter including subsistence
and commercial hunting, and national and international trade, which are
discussed in more detail elsewhere (see later text, Sections 2.5 & 2.6). Objectives: 1. To promote the effective conservation of representative
populations/subspecies of these animals and their habitats in each
country/region/biome, with particular reference to the currently most
threatened populations and subspecies. 2. To obtain more basic and applied information on
the species with a view to the development of practical management strategies
on a sound conservation basis. 3. To initiate and improve routine monitoring of
peccary population status, trends and levels of human utilisation for
subsistence and trade purposes. 4. To enable the rational and sustainable
utilization of peccary by subsistence hunters wherever appropriate (i.e.
where this is not in conflict with ultimate survival prospects of that
population). Priority Projects: 1. Assess and monitor the status and habitats of the
most threatened populations (e.g. in Mexico, Costa Rica and Panama). 2. Elaborate, supplement and disseminate existing
data on the distribution, status and habitat of the species throughout its
range. Literature and questionaire surveys need to be
conducted to assess the current distribution, status and priority management
needs of this species on a country-by-country basis, in order to the promote
field surveys in selected areas and to develop management recommendations
relating to particular populations, protected areas or other conservation
issues. 3. Strengthen existing protected areas which
maintain important populations and habitats of this species. This must include the enforcing of protective
legislation, improving the infrastructure of these areas, and promoting the
training of personnel and wildlife managers (see below). Particular problems
which need to be addressed include regulation or prevention of subsistence
hunting within designated protected areas, and coordination between neighbouring
nations for better protection of this species. For example, subsistence
hunting still occurs in most parks and reserves in Mexico and Guatamala,
including the recently gazetted Mayan Reserve (c. >800,000 ha) in the
Department of El Peten in Guatemala (perhaps the single most important
protected area for the threatened subspecies, T. p. ringens). 4. Continue existing studies, and promote further
comparative studies of the population dynamics, home range and ecology of
selected populations/herds. Priority should be given to the promotion of applied
research on this and other peccary species in Latin American universities,
research and breeding centres; thereby also facilitating the generation of
information relevant to local/national needs and issues. 5. Develop studies and projects focused on the
regulation (rather than prohibition) of subsistence hunting. White-lipped peccaries are an important
socio-economic resource for many ethnic groups and rural settlers and that
the desired regulation of this activity requires the active participation of
those people, and in some situations could be integrated with forestry
management plans. A model programme has already been implemented in Peru
(Bodmer, 1988b), which is intended to ensure its optimal utilisation through
the active monitoring and management of peccary populations and establishing
hunting quotas. 6. Implement and enforce of measures aimed at
reducing or eliminating large scale commercial hunting for trade (see
recommendations in Section 2.5). 7. Develop conservation-education programmes in
rural areas, and training courses and workshops for wildlife managers and
reserve staff. Education programmes are important in rural areas to
disseminate information on conservation of peccaries (their ecological and
socio-economic importance, existing utilisation levels, legal and
conservation status and future management needs, etc.), and other wildlife
species. These programmes should be specifically targeted at subsistence
hunters and reserve staff, as well as the wider rural communities. 8. Research on captive
breeding and reintroduction. The lack of sufficient data on the biology of T. pecari greatly hinders the
development of projects for its management, conservation and sustained use.
Research in captive animals could provide much valuable information on the
life-history, reproduction and other aspects of the biology of these animals.
This research should also address the possibility of reintroducing or
translocating animals as a means of restoring populations of this species
where necessary. 9. Conduct a thorough review of the generic and
subspecific taxonomy of this species, assess its regional variation and
enhance protection of any threatened forms. Acknowledgements This paper includes data and recommendations arising
from the project "Habitat evaluation and status of the white-lipped
peccary in Mexico", which is supported by Wildlife Conservation
International, the Program of Wildlife Management of the National University
of Costa Rica, the Program for Studies in Tropical Conservation of the
University of Florida, the U.S. Fish and Wildlife Service and the Center of
Studies for the Conservation of Natural Resources - ECOSFERA. I express my special gratitude to Philip Bubb for
help with the translation of the manuscript, to William Oliver, Andrew Taber,
Richard Bodmer, Lyle Sowls and Hans Frädrich for their valuable suggestions
and comments, to Dr. Archie Carr III, Christopher Vaughan, Joann Andrews and
Kent Redford for their support, and to Paul Vercammen and Peter Cuypers for
preparing the range map. Much of the basic information on the distribution of
the species was given by different researchers and museum curators of many
countries. I thank all of them. References Alston, E. R.
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Chacoan Peccary |