Home‎ > ‎

LC - Collared Peccary

Collared Peccary (Peccari spp.)

Status: Least Concern ver 3.1 (status for Peccari tajacu)

Pop. trend: stable

Taxonomy

Peccari tajacu Linneaus, 1758

Collared Peccary
The generic name of this species has been the subject of debate. It has been referred to as both Tayassu tajacu and Dicotyles tajacu (Jones & Manning 1992; Grubb & Groves 1993). However, recent genetic research shows that collared peccaries cluster in a separate lineage from the other modern species (Gongora & Moran 2005). This, along with taxonomic complications arising from the use of the three proposed genus names, makes Pecari tajacu the most appropriate and accepted name (Gongora et al. 2006, 2011a; Ruvinsky et al. 2011; Taber et al. 2011). DNA (Gongora et al. 2006) and morphological (Groves & Grubb 2011) data show the occurrence of two species within the collared peccary, one in North/Central and one in South America. In addition, it is probable that there is a third species in Central America. However, due to the lack of enough knowledge to differentiate the biology and ecology of each of the recently proposed species, here we refer to a single species account (Pecari tajacu) and various subspecies.


Subspecies and Distribution

Distribution

Distrbution map Collared peccary
The collared peccary is the peccary species with the largest distribution range (Sowls 1997). It occurs in Arizona, New Mexico, and Texas in the USA, a large part of Mexico, especially around the two cordilleras, most of Central America, the entire Amazon basin, the Pacific coastal forest of Colombia,Ecuador, and Peru, the llanos and lowland forest of Venezuela, the Guianas, and Suriname, all of Brazil where populations are increasingly fragmented in the south and east, and the Gran Chaco of Paraguay, Bolivia, and northern Argentina where it also occurs in the upper Parana and Paraguay river basins. In Argentina, the species is extinct in the eastern and southern portions of its original distribution. The Argentine populations of collared peccary in Misiones are isolated from the rest of the country. Some of the larger islands near the mainland in the Caribbean, such as Trinidad and Tobago, also have populations of P. tajacu. However, islands further from the mainland do not currently have peccaries. The species range has recently expanded northward in the southwestern USA (Albert et al. 2004) including into Oklahoma adjacent to Texas (Stangl & Dalquest 1990). The collared peccary is distributed from sea level to almost 3000 m a.s.l. (Sierra Las Minas, Guatemala; Gongora et al. 2011a; Taber et al. 2011).


Subspecies

The collared peccary is recognized as a single species, but there are contrasting findings about the evolutionary relationships with other extant peccary species. Phenotypic, morphological, chromosomal, and DNA data have shown some level of differentiation within the collared peccary. DNA studies have shown that sequence divergence between collared peccaries from distant geographical origins is as extreme or even higher as that between white-lipped and Chacoan peccaries (Gongora & Moran 2005). Variations in size and pelage colour, coupled with distribution data, have been the basis for proposing the existence of 14 subspecies of collared peccaries (Hall 1981; Grubb & Groves 1993), which have been divided into three subcontinental groups: ‘angulatus’ or grey Central American forms (angulatus, sonoriensis, nanus, humeralis, yucatanensis, crassus, nelsoni, and nigrescens) distributed in North and Central America; patira or blackish forms with very poorly expressed collar and dorsal stripe (patira, torvus, crusnigrum, niger, and bangsi) distributed in central Panama across South American countries near the Equatorial line including Colombia, Venezuela, Guyana, Surinam, French Guiana, Ecuador, and central Peru; and tajacu or grey to buffy subspecies with clearly marked pale collar and black dorsal stripe (tajacu) distributed from the south of some of these Equatorial countries across Bolivia, Paraguay, Brazil, and northern Argentina (Bodmer & Sowls 1993; Grubb & Groves 1993). However, comprehensive morphological, cytogenetic, genetic, and genomic studies of the genus across the Americas need to be undertaken to conclusively delineate species and subspecies and assist in defining priorities of conservation. In Colombia, the presence of four (bangsi, niger, patira, and tajacu) out of the 14 proposed subspecies above, the presence of two distinct DNA lineages and two possible different karyotypes in collared peccaries (Gongora et al. 2006, 2011b; Sabogal & Sandra 2011) certainly provide venues for interaction and hybridization between lineages in this region. Some preliminary evidence of possibly hybridization is given by the uncouplingof nuclear and mitochondrial DNA sequence variation in two specimens from Colombia, which may suggest the recent contact of isolated populations within Colombia and possible mitochondrial introgression between the North/Central and South American clades representing two possible species (Gongora et al. 2006). In addition, cytogenetic data of a single captive specimen from Colombia shows that this is heterozygous for a balanced translocation, which may represent a hybrid from the two karyotypes observed in the species (Builes et al. 2004) possibly representing the interactions between the two different DNA lineages found in collared peccaries (Gongoraet al. 2006). Further chromosome and DNA studies including from wild populations are required to better understand chromosomal polymorphisms and hybridization between collaredpeccary genetic lineages in Colombia.


Descriptive Notes

Collared Peccary
The collared peccary is the smallest of the three extant species of peccaries and ranges in total body length from 78.8 to 106 cm, height between 30 and 51 cm and a general range of 15–28 kg weight (Sowls 1997). However, some individuals have reached 42 kg in Arizona and in some areas of Peru (Taber et al. 2011). Collared peccary from Cozumel Island in the Caribbean coast of Mexico are on average the shortest of all populations with an average body length of only 82.3 cm (Merriam 1901). The collared peccary head is relatively large but ends in a smaller nasal disc than the other species. The legs are shorter than whitelipped and Chacoan peccaries. The body is covered by darkish grey hairs with a distinctive white collar that extends from the chest to the shoulders and gives its particular name to the species (Sowls 1997). However, pelage colour may differ substantially within the species even in the same area such in Colombia (Gongora et al. 2006). Variation in coat colourgoes from greyish forms in the north of the range to darker in the Central America countries and grey again in the southern extreme. In addition, their lighter collar pattern may vary from being very distinct in some individuals to barely noticeable in others (Figure 24.3). A line of dark hair starts at the back of the head and ends at the tail and is more pronounced in the juveniles (Sowls 1997; Gongora et al. 2006). Newborns and juveniles are reddish with the dark line of the back highlighted (Sowls 1997). A scent gland about 7.5 cm in diameter is located on the lower back along the mid-dorsal line, approximately 15 cm from the base of the tail. This gland emits a strong scent that can be detected several hundred metres away and that may function as a way to maintain cohesiveness among the group, or as a territorial mark (Sowls 1997; Taber et al. 2011).


Habitat

The collared peccary has the most extended distribution range of the three species and is the most adaptable; it lives in a wide array of habitat conditions from semi-arid areas in Arizona, New Mexico, and Texas to tropical perennial forest in the Amazon forest (Sowls 1997). Collared peccary tolerates temperatures as high as 45°C during the day and freezing temperatures at night in the northern extreme of its distribution. This species’ tolerance to low temperatures may explain its ability to inhabit further north than the other two species (Sowls 1997). It is welladapted to arid environments by feeding in foods rich in water contents such as cactus and also by resting in the shade during the hot hours of the day (Sowls 1997). In the northern extreme the species also inhabits montane conifer forests at higher altitudes. At the other extreme of the distribution range, the collared peccary inhabits tall, perennial tropical mature forest in the Amazon region and even pre-montane forest in the Andean foothills, with a maximum reported altitude of 2000 and 3000 m a.s.l. in Ecuador and Sierra Las Minas in Guatemala, respectively (Taber et al. 2011). The collared peccary’s ability to inhabit diverse vegetation types may help the species to survive in disturbed habitats such as highly hunted and fragmented areas. In the southern USA, collared peccaries are abundant on the outskirts of large cities. They have become accustomed to people and are often seen walking on streets, foraging through garbage bins or gardens (M. Altrichter, personal observation). In the Calakmul area of Mexico, the collared peccary is common and considered an important crop-raider in fragmented areas where people cultivate corn, squash, and other crops. Despite suffering heavy hunting pressure, it was found that collared peccaries were common in some areas of southern Mexico (Weber 2000; Naranjo 2002; Reyna-Hurtado & Tanner 2007) and in several sites of the Amazon forest (Peres 1996).


Population Densities

Despite the collared peccary being the most studied species of Tayassuidae, there are relatively few reliable estimates of population density. Density estimates of collared peccary vary enormously, from 0.05 to as high as 14 ind./km2. The factors affecting densities may be a combination of ecological and anthropogenic elements. For example, Peres (1996) showed evidence that density estimates vary according to hunting pressure. In Calakmul Biosphere Reserve, collared peccary had similar relative abundance estimates in hunted and protected sites despite being the most hunted ungulate in the area (Weber 2000; Reyna-Hurtado & Tanner 2007). However, recent estimates indicate a significant decrease in relative abundance in heavily hunted sites (Briceño-Mendes et al. 2016). It is unknown how densities of peccaries vary where they are sympatric with white-lipped and Chacoan peccaries;however, it has been observed that collared peccary densities increased at La Selva, Costa Rica, a few years after the local extinction of white-lipped peccaries (Romero et al. 2013).


Movements and Home Range

Collared Peccary
Research in movement patterns of Collared peccary goes back to early studies of Minamon (1962, cited in Sowls 1997), who marked several animals, re-trapped them and found a home range of 2.6–8.0 km2 in the Tucson Mountains of Arizona. Estimates of home range in Texas varied between 0.73 and 2.25 km2 (Ellisor & Harwell 1969), and 0.52 and 1.56 km2 (Schweinsburg 1971). In Arizona, Day (1985) found an average home range of 3.21 km² for 25 distinct herds, while in the drier Paraguayan Chaco, the home range was estimated between 1.5 and 6.8 km2 (100 per cent Minimum Convex Polygon) (Taber et al. 2011). In some tropical areas it seems that home range estimates are smaller than the arid zones of the USA, with the exception of home range estimates in the Amazon, where Fragoso (1999) estimated 10.1–11.7 km2 (100 per cent Minimum Convex Polygon) for two herds in Maraca Island in Roraima, Brazil. In northwestern Costa Rica, home ranges were between 0.64 and 1.09 km² (95 per cent Minimum Convex Polygon) for three monitored herds (McCoy et al. 1990), and Judas and Henry (1999) reported home range estimates of three herds ranging between 1.57 and 2.43 km2 (95 per cent Kernel method) from French Guiana. The average home range sizes of two radio-tracked collared peccary herds in a 21.78 km2 fragment of semi-decidous Atlantic Forest of Brazil were, respectively, 3.69 and 1.80 km2 (95 per cent Fixed Kernel method) or 4.07 and 1.07 km2 with the 95 per cent Minimum Convex Polygon method (Keuroghlian et al. 2004).In contrast to white-lipped peccaries, home-range use by collared peccaries in the Atlantic Forest of Brazil showed fewer and more diffuse centres of activity and less pronounced seasonal shifts (Keuroghlian et al. 2004).

Variation of home range area among different herds in the same region is not unusual. Some have hypothesized that larger herd sizes are correlated with larger home range sizes (Castellanos 1983; Fragoso 1999). However, McCoy et al.(1990) observed the opposite trend. Keuroghlian et al. (2004) suggested that differences in home range were due to variability of resources (habitats, fruit sources, etc.) among locations. Home range size may also vary by seasons. The collared peccary herds studied by McCoy et al. (1990) used larger areas during the dry season than during the wet season. They attributed the greater home range sizes during the dry season to a period of fruit scarcity. Keuroghlian et al. (2004) found that one herd had a larger home range in the wet season than in the dry season, while another herd showed less of a difference and an opposite trend, i.e. the dry season range was slightly larger than the wet season range. However, Ilse and Hellgren (1995) and Judas andHenry (1999) observed smaller area use when food resources were reduced and found that variation of fruit production in time and space in the French Guianas influenced collared peccary home range size. From a landscape perspective in the Atlantic Forest of Brazil, the size of the home range of collared peccary was undoubtedly affected by the territorial behaviour and minimal overlap of adjoining herds (Keuroghlian et al. 2004, 2008). Territoriality among herds has been observed by a number of authors (reviewed by Sowls 1997). Taber et al. (2011) also found that home ranges of collared peccary overlap in some degree, but herds are territorial, defending a core area.


Activity Patterns

The collared peccary seems to be a diurnal/crepuscular species that travels or moves early in the morning and late in the afternoon(Keuroghlian et al. 2004). Movement pattern varies with temperature, food and water availability, and hunting pressure (Taber et al. 2011). Castellanos (1985) found animals moving peaks between 5:00 to noon and then from 16:00 to 19:00 in theCentral Llanos of Venezuela. In the Pantanal, collared peccary activity peaked in the early evening, possibly avoiding whitelipped peccary activity peaks, which occurred in the morning (Galetti et al. 2015). In La Selva, Costa Rica, where white-lipped peccaries are locally extinct, Romero et al. (2013) found that collared peccaries were more diurnal then nocturnal. In arid environments, the collared peccary can be nocturnal to some degree to avoid intense heat (West Texas, Bissonette 1982; Chaco, Argentina, Altrichter 2005). In the semi-arid tropical forest of Calakmul, collared peccary movement peaked between 8:00 and 12:00 (Briceño-Mendes et al. 2016).


Feeding Ecology

The collared peccary is primarily frugivorous in the tropics (>60 per cent of the diet is composed of fruits), but its diet varies according to the ecosystem (Sowls 1997; Desbiez et al. 2009). Collared peccaries can consume roots, tubers, leaves, fruits, and flowers as well as invertebrates and small vertebrates (Beck 2005; Keuroghlian et al. 2008; Taber et al. 2011). In the Atlantic Forest of Brazil, fruit items represented 78 per cent in the dry and the wet seasons (Keuroghlian et al. 2008), similar to what was found in the Pantanal (Desbiez et al. 2009). In the tropical forest of Calakmul of southern Mexico, collared peccaries fed on 37 species of plants with proportions of 58 per cent fruits and 30 per cent leaves (Perez-Cortez & Reyna-Hurtado 2008). In the Atlantic Forest, diggings associated with tuber consumption were made exclusively by collared peccaries (Keuroghlian et al. 2008). In addition, there was a seasonal trend that showed higher tuber consumption in the dry compared to the wet season (dry: 63 per cent, wet: 35 per cent, n = 22 diggings associated with tuber consumption; Keuroghlian et al. 2008). Studies from other Neotropical regions reported lower percentages of fruits in peccary diets, i.e. 59–66 per cent for both white-lipped and collared peccaries in the Peruvian Amazon (Kiltie 1981; Bodmer 1989). In the Pantanal (Brazil), the diversity of fruit species consumed was similar between the two species, and diet overlap was also high (Desbiez et al. 2009). In the Chaco forest the collared peccaries fed on 128 fruit species and 79 species of seeds (Taber et al. 2011). The most common fruit in faecal samples from collared peccaries in the Atlantic Forest was a palm (Keuroghlian et al. 2008). In arid environments of the southern USA, collared peccaries depend largely on the prickly pear cactus (Opuntia spp.; Sowls 1997), which is also an important source of water, a limiting resource in this habitat. In the Calakmul Biosphere Reserve, Mexico, thr collared peccary has been observed to frequently drink, bathe and wallow in temporal water bodies (R. Reyna-Hurtado, personal observation). Collared peccary may split into temporary subgroups when foraging (Keuroghlian et al. 2008). Long-term studies have shown potential impacts of collared peccaries on seedling demographics in the Amazon. For example, Beck et al. (2010) and Reider et al. (2013) concluded that defaunation of collared peccaries, a species that has disproportional non-redundant interactions with plants via seed dispersal, predation, trampling, and herbivory, may eventuality change the forest structure and canopy tree community and affect other species.


Reproduction and Growth

Collared peccary can breed year-round with peaks from November to March in the temperate zones (Taber et al. 2011). In the Peruvian Amazon, birth peaks occur in January, July, August, and October (Taber et al. 2011). The gestation period varies between 138 and 150 days (Lochmiller et al. 1984; Sowls 1997; Mayor et al. 2006a). In the Amazon, the collared peccary female is considered to be a seasonally polyoestrous (Mayor et al. 2006b) with a pregnancy rate of 42.5 per cent (Mayor et al. 2006b). In the Peruvian Amazon, overall, gross annual productivity was 0.89 foetuses/adult femalefor collared peccaries (Gottdenker & Bodmer 1998). Reproductive production of the species in the wild was estimated at 1.12 births/ year (Mayor et al. 2006b). Both males and females have offspring with more than one partner, showing a promiscuous genetic mating system (Cooper et al. 2011). The litter size varies from one to fourt, but generally one to two newborns are seen following the mother for several months (Taber et al. 2011). Newborns are able to follow their mothers within an hour or two after being born. It has been observed that mothers actively defend their young by vocalizations and tooth chattering. Generally juveniles reach weight and height of an adult at about 40 weeks of age (Sowls 1997).


Behaviour

The collared peccary is social and lives in herds. The herd size varies largely from two to 50 animals but the common herd size is around 10 animals (Kiltie & Terborgh 1983; Robinson & Eisenberg 1985; McCoy et al. 1990; Sowls 1997; Judas & Henry 1999; Mandujano 1999; Keuroghlian et al. 2004; but see Castellanos 1983). In Arizona, herd size averages 8.5 animals (from a sample size of 127 herds; Knipe 1957 in Sowls 1997). In Calakmul Biosphere Reserve, Mexico, herd size average six from a total of 88 observations (Reyna-Hurtado, in preparation). Robinson and Eisenberg (1985) found averages of 3.1 and 6.5 animals in herds in Panama and Venezuela, respectively. The average number of individuals per herd was 8.8 and 13.5 in fragments of the Atlantic Forest highlands (Keuroghlianet al. 2004; Cullen et al. 2001, respectively). Peres (1996), studying the population status of peccaries in forests with and without hunting in the Amazon, found several small groups with averages of five to eight individuals (N = 16). In the Pantanal, group size varied between five and 10 individual (Desbiez et al. 2009).

Collared peccary herds split often into subgroups (Ilse & Hellgren 1995; Sowls 1997; Judas & Henry 1999; Keuroghlian et al. 2004). Fragoso (1999) followed two herds, one of 34 and one of 12 individuals in Maraca Island, Brazil, and observedthat these herds usually split into four subgroups. In the Atlantic Forest, Keuroghlian et al. (2004) found that radio-tracked herds would split into groups of one to three individuals during the day and foraged separately at distances between 30 and 250 m apart. Oldenburg et al. (1985) reported distances of separation of a minimum of 100 m to a maximum of 1400 m and most ofthe splits lasted less than 12 hours (Sowls 1997). Sightings of lone collared peccaries or groups of two to three individuals have frequently been reported from sites in both the Neotropics and the southwestern USA (Bissonette 1982; Castellanos 1983; Kiltie & Terborgh 1983; Green et al. 1984; Oldenburg et al. 1985; Robinson & Eisenberg 1985; Sowls 1997). Keuroghlian et al. (2004) found that collared peccaries appeared to travel more in the early morning and late afternoon, when the full herd was together. During midday, subherds would usually remain foraging in a relatively restricted area. They also spent a good part of late morning and early afternoon resting in cavities excavated at the bases of large trees, next to fallen trunks, or in other cool locations (Sowls 1997; Keuroghlian et al. 2004). This period of inactivity was also observed in the Paraguayan Chaco (Taber et al. 1994). Herd members maintained contact by vocalizing and releasing strong liquid musk from the dorsal gland.The dorsal gland of the collared peccary contains both sebaceous and sudoriferous secretory tissues that produce some 20 volatile components, some specific to males or females. They deposit scent on each other, as well as on tree trunks, rocks, and other objects (Sowls 1997; Taber et al. 2011). Eight types of vocalizations have been identified including purring, grunting, barking, woofing, growling, and squealing as well as jaw snapping (Sowls 1997; Taber et al. 2011). Byers (1980), Bissonette (1982) and Sowls (1997), among others, have described in detail the behavioural repertoire of collared peccary social communication. Some friendly interactions that arise are the touching of nasal disc as a form of greeting and the rubbing of the dorsal gland as a way to maintain group cohesiveness and member identification (Sowls 1997). Although there is no clear dominance hierarchy among members of a herd, there are frequent aggressive interactions among members that includes a behaviour named ‘squabble’ (Schweinsburg & Sowls 1972), described as when two animals face each other in a sparring position and tooth chattering accompanies the action. Tooth chattering is also used very often as intraspecific social behaviour, usually in agonistic situations (Sowls 1997).


Status in the Wild

The collared peccary is the most common of the three extant peccary species and the one with the largest distributionrange. The collared peccary tolerates disturbed areas and can survive under hunting pressure (Peres 1996; Weber 2000; Peres & Palacios 2007; Reyna-Hurtado & Tanner 2007). It is listed as Least Concern under the IUCN red list (IUCN Red List of Threatened Species. Version 2015-4. www.iucnredlist .org. Downloaded 30 December 2015) and classified in CITES Appendix II. It is the main species of a harvesting programme that exports peccary skins from the Peruvian Amazon under a certification programme that annually exports 50,000 skins approximately (Fang et al. 2008). However, its status requires monitoring, as the species may be under several threats and pressures. The two major threats to the survival of the collared peccary are over-hunting for their meat and hides and destruction of their natural habitats. These factors have already resulted in the extensive fragmentation of collared peccary populations and its extirpation over large parts of its former range (Bodmer & Sowls 1993). For example, in the areas where it was harvested under a certification programme in Peru, historical record flooding from 2010 to 2015 produced major declines on the species’ densities in vast areas within the National Park Pacaya-Samiria (Bodmer et al. 2014; R. Bodmer, personal communication). In Calakmul Biosphere Reserve, thecollared peccary displays a resilient behaviour enduring hunting pressure and has similar relative abundance in communal forest and in protected areas (Reyna-Hurtado & Tanner 2007). However, recent evidence from the same sites indicates a drastic reduction on the species’ abundance in areas of high hunting pressure (i.e. the index of relative abundance of a hunted area is 3.7 photos/1000 camera-trap days in comparison to 27.5 photos/1000 camera-trap days in non-hunting areas; Briceño-Mendes et al. 2016). This recent evidence and the continuous increase in fragmentation and hunting pressure observed in all Neotropical forests make it urgent to monitor populations of collared peccary all over its range, especially in those areas where distinct taxonomic units may exist. The collared peccary is a very important species that ecologically plays critical roles as seed predator and disperser (Beck 2005, 2006), ecosystem engineer, and soil and pond modifier (Beck et al. 2010). The collared peccary is also an important prey for large predators such as jaguar and pumas and helps maintain local livelihoods throughout its geographical distribution. Efforts must be made to ensure that the collared peccary is conserved in all areas of its original range and to reduce threats such as forest  fragmentation, hunting pressure, introduced exotic species and disease transmission. By conserving the collared peccary we ensure that this species will persist to fulfil its important ecological role and to amaze future generations as an example of adaptation and endurance in a wide range of habitats.


Text adapted from: Reyna, R., Keuroghlian, A., Altrichter, M. Beck, H. and J. Gongora (2017). Collared Peccary Peccari spp.. In: M. Meletti and E. Meijaard, editors. Ecology, Conservation and Management of Wild Pigs and Peccaries. Cambridge, UK.


Relevant Literature

Albert, S., Ramotnik, C. A., & Schmitt, C. G. (2004). Collared peccary range expansion in northwestern New Mexico. The Southwestern Naturalist 49: 524–528.

Altrichter, M. (2005). The sustainability of subsistence hunting of peccaries in the Argentine Chaco. Biological Conservation 126: 351–362.

Beck, H. (2005). Seed predation and dispersal by peccaries throughout the Neotropics and its consequences: a review and synthesis. In Forget, P-M., Lambert, J. E., Hulme, P. E. & Vander Wall, S. B. (eds.), Seed fate: predation, dispersal and seedling establishment. Wallingford: CABI Publishing, pp. 77–115.

(2006). A review of peccary–palm interactions and their ecological ramifications across the Neotropics. Journal of Mammalogy 87: 519–530.

Beck, H., Thebpanya, P. & Filiaggi, M. (2010). Do Neotropical peccary species (Tayassuidae) function as ecosystem engineers for anurans? Journal of Tropical Ecology 26: 407–414.

Bissonette, J. A. (1982). Ecology and social behavior of the collared peccary in Big Bend National Park. Scientific Monograph Series No 16. Washington, DC: US National Park Service.

Bodmer, R. E. (1989). Frugivory in Amazonian Artiodactyla: evidence for the evolution of the ruminant stomach. Journal of Zoology 219: 457–467.

Bodmer, R. E. & Sowls, L. K. (1993). The collared peccary (Tayassu tajacu). In Oliver, W. L. R. (ed.), Pigs, peccaries, and hippos: status survey and conservation action plan. Gland: IUCN.

Bodmer, R.E., Fang, T.G., Puertas, P.E., et al. (2014). Cambio climático y fauna Silvestre en la Amazonia peruana. Iquitos, Peru: Wust Editions.

Briceño-Mendes, M., Naranjo, E., Altrichter, M. & Reyna-Hurtado, R. (2016). Responses of two sympatric species of peccaries (Tayassu pecari and Pecari tajacu) to hunting in Calakmul, Mexico. Tropical Conservation Science 9: 1–11.

Builes, D. F. P., Loaiza, I. D. D., Lopez, J. B. O., Marquez, E. M. & Bock, B. C. (2004). Citogenetica basica del pecari de Collar Tayassu tajacu. Diferencias cromosomicas gruesas entre sus poblaciones. Universidad Nacional de Colombia,Medellin. Available from: www.reuna.edu.co/temporales/memorias/especies/EspeciesAnimales.htm.

Byers, J. A. (1980). Social behavior and its development in the collared peccary(Tayassu tajacu). PhD dissertation, University of Colorado, Boulder, CO.

Castellanos, H. G. (1983). Aspectos de laorganización social del baquiro de collar Tayassu tajacu en el estado Guarico, Venezuela. Acta Zoológica Venezolana 11: 127–143.

(1985). Home range size and habitat selection of the collared peccary in the state of Guarico, Venezuela. In Ockenfels, R. A., Day, G. I. & Suppleee, V. C.(eds.), Peccary workshop proceedings. Phoenix, AZ: Arizona Game and Fish Department.

Cooper, J. D., Waser, P. M., Hellgren, E. C.,Garbor, T. M. & De Wood, J. A. (2011). Is sexual monomorphism a predictor of polygynandry? Evidence from a social mammal, the collared peccary. Behavioral Ecology and Sociobiology 6: 775–785.

Cullen L. Jr. (1997). Hunting and biodiversityin Atlantic forest fragments, São Paulo, Brazil. MS thesis. Gainesville, FL: University of Florida.

Cullen, L. Jr., Bodmer R. E. & Valladares-Padua, C. (2001). Ecological consequences of hunting in Atlantic forest patches, Sao Paulo, Brazil. Oryx 35: 137–144.

Day, G. I. (1985). Javelina research and management in Arizona. Phoenix, AZ: Arizona Game and Fish Department.

de Freitas, T. T., Keuroghlian, A., Eaton, D., et al. (2010). Prevalence of Leptospira interrogans antibodies in free-ranging Tayassu pecari of the Southern Pantanal, Brazil, an ecosystem where wildlife and cattle interact. Tropical Animal Health and Production 42: 1695–1703.

Desbiez, A. L. J., Santos, S. A., Keuroghlian, A. & Bodmer, R. E. (2009). Niche partitioning between sympatric populations of native white-lipped peccary (Tayassu pecari), collared peccary (Tayassu tajacu), and introduced feral pigs (Sus scrofa). Journal of Mammalogy 90(1): 119–128.

Eisenberg, J. F. (1980). The density andbiomass of tropical mammals. In Soulé, M. & Wilcox, B. A. (eds.), Conservation biology: an evolutionary–ecological perspective. Sunderland, MA: Sinauer Associates, pp. 35–55.

Eisenberg, J. F., O’Connell, M. A. & August, P. V. (1979). Density, productivity,and distribution of mammals in two Venezuelan habitats. Vertebrate ecology in the northern Neotropics. Washington, DC: Smithsonian Instution Press, pp. 187–207.

Ellisor, J. E. & Harwell, F. (1969). Mobility and home range of collared peccary in southern Texas. Journal of Wildlife Management 33: 425–427.

Emmons, L. H. & Feer, F. (1990). Neotropical rainforest mammals. A field guide. Chicago, IL: The University of ChicagoPress.

Fang, T. G., Bodmer, R. E., Puertas, P.E., et al. (2008). Certificación de pieles de pecaríes en la Amazonia Peruana. Lima: Wust Editions.

Fragoso, J. M. V. (1999). Perception of scale and resource partitioning by peccaries: behavioral causes and ecological implications. Journal of Mammalogy 80(3): 993–1003.

Galetti, M., Camargo, H., Siqueira, T., et al. (2015). Diet overlap and foraging activity between feral pigs and native peccaries in the Pantanal. PLoS ONE 10(11): e0141459. doi:10.1371/journal. Pone.0141459

Glanz, W. E. (1982). The terrestrial mammal fauna of Barro Colorado Island: censuses and long-term changes. In Leigh, E. G., Jr, Rand, A. S. & Windsor, D. M. (eds.), The ecology of a tropical forest. Washington, DC: Smithsonian Institution Press.

Gómez, B. & Montenegro, O. (2012). Abundancia del pecarí de collar (Pecari tajacu) en dos áreas protegidas de la Guayana colombiana. Mastozoología neotropical 19: 163–178.

Gongora, J. & Moran, C. (2005). Nuclear and mitochondrial evolutionary analyses of Collared, White-lipped, and Chacoan peccaries (Tayassuidae). Molecular Phylogenetics and Evolution 34: 181–189.

Gongora, J., Morales, S., Bernal, J. E. & Moran, C. (2006). Phylogenetic divisions among collared peccaries (Pecari tajacu) detected using mitochondrial and nuclear sequences. Molecular Phylogenetics and Evolution 41: 1–11.

Gongora, J., Reyna-Hurtado, R., Beck, H., et al. (2011a). Pecari tajacu. The IUCN Red List of Threatened Species 2011: e.T41777A10562361. http://dx.doi.org/10.2305/IUCN.UK.2011-2.RLTS.T41777A10562361.en. Downloaded 5 November 2015.

Gongora, J., Biondo, C., Cooper, J., et al. (2011b). Revisiting the species status of Pecari maximus van Roosmalen et al., 2007 (Mammalia) from the Brazilian Amazon. Bonn Zoological Bulletin 60(1): 95–101.

Gottdenker, N. L. & Bodmer, R. E. (1998). Reproduction and productivity of whitelipped and collared peccaries in the Peruvian Amazon. Journal of Zoology 245: 423–430.

Green, G. E., Grant, W.E. & Davis, E. (1984).Variability of observed group sizes within collared peccary herds. Journal of Wildlife Management 48: 244–248.

Groves C. P. & Grubb, P. (2011). Ungulate taxonomy. Baltimore, MD: Johns Hopkins University Press. Grubb, P. & Groves, C. P. (1993). The Neotropical Tayassuids Tayassu and

Catagonus. In Oliver, W. L. R. (ed.), Pigs, peccaries and hippos: status survey andconservation action plan. Gland: IUCN, pp. 5–7.

Hall, E. R. (1981). The mammals of North America. New York, NY: John Wiley and Sons.

Ilse, L. M. & Hellgren, E. C. (1995). Resource partitioning in sympatric populations of collared peccaries and feral hogs in southern Texas. Journal of Mammalogy 76: 784–799.

Jones, J. K. & Manning, R. W. (1992). Illustrated key to skulls of genera of North American land mammals. Lubbock, TX: Texas Tech University Press.

Judas, J. & Henry, O. (1999). Seasonal variation of home range of collared peccary in tropical rain forests of French Guiana. Journal of Wildlife Management 63: 546–555.

Karesh, W. B., Uhart, M., Painter, L., et al. (1998). Health evaluation of white lipped peccary populations in Bolivia. Omaha,NB: American Association of Wildlife Veterinarians, pp. 445–449.

Keuroghlian, A. & Desbiez, A. L. J. (2010). Biometric and age estimation of live peccaries in the Southern Pantanal,Brazil. Suiform Soundings 9: 24–35.

Keuroghlian, A., Eaton, D. P. & Longland, W. S. (2004). Area use by white-lipped and collared peccaries (Tayassu pecari and Tayassu tajacu) in a tropical forest fragment. Biological Conservation 120: 411–425.

Keuroghlian, A., Donald, P. & Eaton, D. P. (2008). Fruit availability and peccary frugivory in an isolated Atlantic forest fragment: effects on peccary ranging behavior and habitat use. Biotropica 40: 62–70.

Kiltie, R. A. (1981). Stomach contents of rain forest peccaries (Tayassu tajacu and T. pecari). Biotropica 13: 234–236.

Kiltie, R. A. & Terborgh, J. (1983). Observations on the behavior of rain forest peccaries in Peru. Why do whitelipped peccaries form herds? Zeitschrift für Tierpsychologie 62: 214–255.

Knipe, T. (1957). Javelinas in Arizona. Wildlife Bulletin No 2. Phoenix, AZ: Arizona Game and Fish Department.

Lochmiller, R. L., Hellgren, E. C. & Grant, W. E. (1984). Selected aspects of collared peccary (Dicotyles tajacu) reproductive biology in a captive Texas herd. Zoo Biology 3: 145–149.

Low, W. A. (1970). The influence of aridity on reproduction of the collared peccary (Dicotyles tajacu; Linn.) in Texas. PhD dissertation. Vancouver: University of British Columbia.

Mandujano, S. (1999). Variation in herd size of collared peccaries in a Mexican tropical forest. The Southwestern Naturalist 44: 199–204.

Mayor, P., Guimaraes, D.A., Lopez-Gatius, F. & Lopez-Bejar, M. (2006a). First postpartum estrus and pregnancy in the female collared peccary (Tayassu tajacu) from the Amazon. Theriogenology 66: 2001–2007.

Mayor, P., Fenech, M., Bodmer, R. E. & Lopez-Bejar, M. (2006b). Ovarian features of the wild collared peccary (Tayassu tajacu) from Peruvian Northeastern Amazon. General and Comparative Endocrinology 147: 268–275.

McCoy, M. B., Vaughan, C. S. & Rodrigues, M. A. (1990). Seasonal movement, home range, activity and diet of collared peccaries (Tayassu tajacu) in Costa Rican dry forest. Vida Silvestre Neotropical 2: 6–20.

Merriam, C. H. (1901). Six new mammals from Cozumel Island, Yucatán. Proceedings of the Biological Society of Washington XIV: 99–104.

Naranjo, E. J. (2002). Population ecology and conservation of ungulates in the Lacandon forest, México. PhD dissertation, The University of Florida, Gainesville, Florida.

Nava, A. & Cullen, L. (2003). Peccaries as sentinel species: conservation, health and training in Atlantic Forest Fragments, Brazil. Suiform Soundings 35.

Oldenburg, P. W., Ettestad, P.J., Grant, W.E. & Davis, E. (1985). Size, overlap and temporal shifts of collared peccary herd territories in South Texas. Journal of Mammalogy 66: 378–380.

Peres, C. A. (1996). Population status of white-lipped Tayassu pecari and collared peccaries T. tajacu in hunted and unhunted Amazonian forest. Biological Conservation 77: 115–123.

Peres, C. A. & Palacios, E. (2007). Basinwide effects of game harvest on vertebrate population densities in Amazonian forests: implications for animal-mediated seed dispersal. Biotropica 39: 304–315.

Pérez-Cortéz, S. & Reyna-Hurtado, R. (2008). La dieta de los pecaríes (Pecari tajacu y Tayassu pecari) en la región de Calakmul, Campeche, México. Revista Mexicana de Mastozoología 12: 17–42.

Reider K. E., Carson, W. P. & Donnelly, M. A. (2013). Effects of collared peccary (Pecari tajacu) exclusion on leaf litter amphibians and reptiles in a Neotropical wet forest, Costa Rica. Biological Conservation 163: 90–98.

Reyna-Hurtado, R. & Tanner, G. W. (2007).Ungulate relative abundance in hunted and non-hunted sites in Calakmul Forest (Southern Mexico). Biodiversity and Conservation 16: 743–757.

Robinson, J. G. & Eisenberg, J. F. (1985). Group size and foraging habits of the collared peccary (Tayassu tajacu). Journal of Mammalogy 66: 153–155.

Robinson, J. G. & Redford, K. (1986). Body size, diet and population density of Neotropical forest mammals. American Naturalist 128: 665–680.

Romero, A., O’Neill, B. J., Timm, R. M., Gerow, K. G. & McClearn, D. (2013). Group dynamics, behavior, and current and historical abundance of peccaries in Costa Rica’s Caribbean lowlands. Journal of Mammalogy 94: 771–791.

Ruvinsky, A., Rothschild, M., Larson, G. & Gongora, J. (2011). Systematics and evolution of the pig. In Rothschild, M. F. & Ruvinsky, A. (eds.), The genetics of the pig. 2nd ed. Wallingford: CABI Publishing, pp. 1–13.

Sabogal, R. & Sandra, P. (2011). Filogeografía y conservación genética del pecarí de collar, Pecari tajacu en cuatro departamentos de Colombia[Phylogeography and conservation genetics of collared peccari, Pecari tajacu in four Colombiandepartments]. Maestría thesis, Universidad Nacional de Colombia.  www.bdigital.unal.edu.co/4000/#sthash.i2Bzt85s.dpuf

Schaller, G. B. (1983). Mammals and their biomass on a Brazilian ranch. Arquivos de Zoologia (Sao Paulo) 31: 1–36.

Schweinsburg, R. E. (1971). The home ranges and herd integrity of the collared peccary. Journal of Wildlife Management 35: 455–460.

Schweinsburg, R. E. & Sowls, L. K. (1972). Aggressive behavior and related phenomena in the collared peccary. Tierpsychol 30: 132–145.

Sowls, K. (1997). Javelines and other peccaries. Their biology management and use. 2nd ed. Tucson, AZ: Texas A & MUniversity Press, p. 325.

Stangl, F. B. & Dalquest, W. W. (1990). Status of the javelina Tayassu tajacu in northcentral Texas and Southern Oklahoma. Texas Journal of Science 42: 305–306.

Supple, V. C. (1983). The dynamics of collared peccary dispersion into available range. Federal Aid in Wildlife Restoration Project W-78-Arizona Game and Fish Department, Phoenix, AZ. 31 pp.

Taber, A. B., Doncaster, C. P., Neris, N. N. & Coleman, F. H. (1994). Ranging behavior and activity patterns of two sympatric peccaries, Catagonus wagneri and Tayassu tajacu in the Paraguayan Chaco. Mammalia 58: 61–71.

Taber, A., Altrichter, M., Harald, B. & Gongora, J. (2011). Family Tayassuidae (peccaries). In Wilson, D. E. & Mittermeier, R. A. (eds.), Handbook of the mammals of the world - volume 2: hoofed mammals. Barcelona: Lynx Editions, pp. 292–307.

Weber, M. (2000). Effects of hunting on tropical deer populations in Southeastern México. MSc thesis, London: Royal Veterinary College, University of London.