Peccaries

(Tayassuidae)

Class Mammalia

Order Artiodactyla

Suborder Suiformes

Family Tayassuidae

Thumbnail description
Pig-like mammals whose skin is covered in coarse bristles, with longer dorsal bristles and shorter side bristles; well-developed snout and nasal muscles; stout legs with a fused ulna and radius; relatively large, sharp, vertically directed canine teeth; a scent gland located approximately 5.9 in (15 cm) cranial to the base of the tail on the dorsal midline; a complex stomach with four sacculated compartments

Size
24.2–92.5 lb (11–42 kg)

Number of genera, species
3 genera; 3 species

Habitat
Desert, dry tropical forest, chaco, rainforest, Brazilian Atlantic forests, low Andean forests, llanos, pantanal, and cerrado

Conservation status
Endangered: 1 species

Distribution
Southwestern portion of North America; Central America; northern and central portions of South America

Evolution and systematics

The first peccaries were believed to have evolved during the Oligocene, with ancestral forms of peccary species migrating across the land bridge from North to South America around 2.5 million years ago. Platygonus is an example of an upper Pleistocene peccary genus that has been described in both North and South America, and Catagonus wagneri is proposed to be its closest extant relative. Out of the many ancestral peccary forms, there are three extant species: Tayassu pecari (white-lipped peccary), Tayassu tajacu (collared peccary), and Catagonus wagneri (Chacoan peccary).

Cladistic analysis suggest that white-lipped peccary and Chacoan peccary ancestors diverged in North America before coming to South America during the late Pliocene, and that white-lipped and Chacoan peccaries are more closely related to each other than white-lipped peccaries are to collared peccaries. Studies suggest that white-lipped and Chacoan peccaries are from a single clade that diverged about 1.7 million years ago (mya), with the white-lipped peccary clade separating from collared peccaries around 3.4–7.4.

Although common taxonomic nomenclature places white-lipped and collared peccaries in the genus Tayassu, and Chacoan peccaries in their own genus Catagonus, this classification is controversial based on evolutionary and genetic considerations. The "two-genera peccary theory" proposes that white-lipped and collared peccary are more similar to each other morphologically than to the Chacoan peccary, justifying the placement of white-lipped and collared peccaries in the genus Tayassu and the Chacoan peccary in Catagonus. The "threegenera peccary theory" breaks the Tayassuidae into three genera: Tayassu for white-lipped peccaries, Dicotyles for collared peccaries, and Catagonus for Chacoan peccaries. Genetic analysis of mitochondrial DNA supports the three-genera theory. Genetic studies place white-lipped and Chacoan peccaries in the same clade, separate from collared peccaries.

Physical characteristics

Tayassuids show few sexual dimorphisms, but skull measurements, particularly in the canine teeth and zygomatic processes, for male and female peccaries may differ. Unique

anatomical features of the Tayassuidae distinguishing them from other Suiformes include their skin, skull, and dentition, specialized digestive tract, dorsal scent gland, and hind leg structure. Peccary skin is covered in coarse bristles, with longer dorsal bristles and shorter side bristles. Color patterns differ among the three species, and there are intraspecific color variations in white-lipped and collared peccaries. Another unique feature of peccary skin is the dorsal scent gland, located approximately 5.9 in (15 cm) cranial to the base of the tail on the dorsal midline. This gland contributes to the very strong, musky odor that permeates peccary herds and meat. This dorsal gland appears to be used, in part, as a mechanism for social cohesion and herd territory definition. Additionally, peccaries do not have a thick layer of subcutaneous fat or dense fur and are poorly equipped to deal with low ambient temperatures.

Peccaries, like pigs, have well-developed snout and nasal muscles to facilitate rooting behavior. They have 38 teeth with a dental formula of I 2–2/3–3, C 1–1/1–1, P 3–3/3–3, M 3–3/3–3. Unlike swine, peccaries have interlocking canines with upper canines that point vertically downward, and a fused radius and ulna, as well as fused median metacarpals and metatarsals. Peccaries also possess a complex stomach with four sacculated compartments: two non-glandular blind sacs, a nonglandular gastric pouch, and a glandular hind stomach. This fore stomach comprises 85% of the total stomach volume of the peccary, 3–6% less than the fore stomach/stomach volume ratio of sheep or cattle. Because microbial fermentation with the formation of high volatile fatty acid levels have been observed in the peccary fore stomach, it has been proposed that their fore stomachs are a means to slow digestive passage and increase digestive efficiency for microbial fermentation of structural components of fruits, nuts, and herbaceous material. Peccaries do not have a gallbladder.

The legs are more highly evolved for running than those of the suids; the metatarsals are fused into a stout canonbone and the fifth digit is lost. The metacarpals III and IV are very stout, but unfused. The radius and ulna are co-ossified (fused together).

Distribution

The tayassuid family has a wide distribution, ranging from the southwestern portion of North America to Mexico and Central America; Pacific coasts of Colombia; tropical forests of northeastern and southeastern Peru; Venezuela and the Guianas; Amazon basin; southern Brazil; and Paraguay, northern Argentina, and lowland Bolivia. Large-bodied peccary species existed throughout the Pleistocene in North America, but vansihed with megafaunal extinction.

Habitat

Peccaries inhabit a wide range of habitat types, including desert regions of southwestern United States and northern

Mexico; moist and dry tropical forests of Central America; Central American rainforests; choco; llanos; Amazon rainforest; wetlands and forests of the Pantanal region of South America; cerrado; Brazilian Atlantic forests; and the South American dry tropical thorn forests of Bolivia, Paraguay, and Argentina known as the Chaco.

Behavior

Peccaries are social herding animals with group sizes between three to more than 500, depending on the species. Peccaries show a classical kin-selected social structure, similar to chimpanzees.

In south Texas, collared peccary herd home ranges expand from 288–300 acres (116.5–121.4 ha) in dense chaparral to 550–757 acres (222.5–306.3 ha) in open mesquite habitat. In Costa Rica, a smaller home range may be attributed to higher habitat productivity or greater food quality. However, in Brazilian Amazonian forests, collared peccaries had home ranges that were relatively large. Peripheral areas of the home range of a collared peccary herd are often shared by other herds. Home ranges of collared peccary can also vary seasonally or in relation to forage conditions.

In general, collared peccaries are diurnal, with the greatest feeding activity in southwestern United States during the morning and late afternoon until dark. Peccaries in the Paraguayan Chaco rise at dawn, then rest during the hottest part of the day. In Arizona and Texas, collared peccaries are more nocturnal in summer and primarily diurnal in winter. They are very territorial and live in mixed sex groups of adult males and females, juveniles, and sub-adults. These groups are relatively cohesive and stable. Herds often are composed of subgroups, with individuals infrequently separating from the herds. When threatened by predators such as dogs, collared peccaries disperse and flee, often running into burrows or logs.

In order to define and mark their territory, collared peccaries use aggressive behavioral interactions and mark territorial boundaries on the ground or vertical structures with secretions from their dorsal scent glands, as well as by fecal deposition. Within their territory, collared peccaries paw the ground and create small dust beds where they often rest. When territory overlaps occur between collared peccary groups, males may display aggressive behavior in areas of overlap. Aggressive behavior includes vocalizations and gestures. Aggressive vocalizations include a continuous growl, tooth clicking, squeals, and alarm-like "woofs." Vocalizations are an important component of white-lipped peccary intragroup communication. In addition to aggressive vocalizations, white-lipped peccaries also have elaborate protocols of togetherness

vocalizations. White-lipped peccaries tend to emit more aggressive vocalizations when they are spaced close together during feeding or resting as compared to when they are traveling in loosely spaced groups. When in danger, white-lipped peccaries clack their teeth, sniff loudly, and may let out a woof-like bark. Inter-individual interactions between peccaries can be quite aggressive, and consist of 14 reported agonistic behaviors, including teeth grinding and clacking, snorting, yawning or gaping, and grimacing. Fighting is relatively common in white-lipped peccary herds. In collared peccaries, the most common agonistic interaction is the squabble, which consists of two animals facing each other in a fighting position and chattering their teeth with their ears flattened back. When frightened, tayassuids bristle the hairs along their neck and back. When alarmed or when making social contact, white-lipped peccaries bristle their hairs, primarily along the dorsum and flank. Overall, intragroup aggression in collared peccary herds is less than in white-lipped peccary herds, and serious fighting is proposed to be rare within herds. Although there is relatively less aggression within collared peccary herds, there is a great deal of friendly interaction such as mutual grooming and rubbing. Similar to other species of peccaries, white-lipped peccaries exhibit allogrooming and mark each other extensively, often rubbing their head in the region of the dorsal gland of each respective partner (mutual marking).

White-lipped peccary females guard their young aggressively from other animals, including herd members, and it is very difficult to separate females from their newborns. Unlike white-lipped peccaries, collared peccary dams allow their young to nurse from other females, and young who have lost their mother may suckle other lactating females. Collared peccary newborns are also visited and touched by other members of the group. In the case of juvenile-adult interactions, a juvenile may pass underneath and between the hind legs of a dominant adult.

Feeding ecology and diet

Members of the Tayassuidae are primarily frugivorous and omnivorous. The ability of collared peccaries to subsist on a wide variety of food sources is likely an important factor in its ability to live in such a wide range of habitats. In southern Texas, prickly pear cladophils are a main staple of the collared peccary. In Texas and Arizona, Opuntia engelmannii is the most abundant food item eaten often during drought. Collared peccaries can also live in areas where there are no prickly pears. In the southwestern United States, collared peccaries need other plants for nutritional support, such as grass, forbs, acorns, pine nuts, and animal matter. They also eat plants and roots such as thistles (Cirsium arvense), gourds (Cucurbita spp.), and lechuguilla (Agave lechuguilla).

However, in Neotropical forests, collared peccaries are primarily frugivorous. Among fruits, the most common eaten by collared peccaries in the Peruvian Amazon are members of the Leguminosae, Sapotaceae, Menispermaceae, and the palms (Iriartea sp., Jessenea sp., and Astrocaryum sp.). As fruit availability decreases, collared peccaries shift to a more omnivorous diet.

When collared peccaries forage, they often root deeply into leaf litter, soil, base of trees, and mud of stream beds. Understanding the dietary composition of collared peccaries throughout their geographic range is very important, because collared peccary productivity is apparently very sensitive to their nutritional state.

White-lipped peccaries have efficient mechanisms for seed predation to exploit entire fruit resources including both the pulp and the seed. White-lipped peccaries are able to overcome the protective strategies of seeds by cracking seeds using their strong adductive jaw muscles, thick skull bones, strong resilient teeth, and interlocking canines. The strong adductive jaw muscles are attached to a large sagittal crest and

large mandible, allowing the peccary to exert massive pressure on the seeds. The thick bones of the skull can withstand the pressure required to crack palm nuts. Premolar and molar teeth of peccaries are extremely strong and difficult to dislocate from the jaw. Interlocking canines are an adaptation to prevent the jaw from dislocating during the excretion of massive pressure.

The peccaries have a sacculated fore stomach that has microbial fermentation, similar to the deer rumen. Microbial fermentation in the stomach of peccaries occurs in three blind sacs, while gastric digestion occurs in the central main sac. These blind sacs are efficient at fermenting low-fiber plant products, but not plant material with high concentrations of cell wall. The blind sacs probably help soften and digest the hard palm seeds once they are cracked into small pieces by peccaries. However, since peccaries do not ruminate, they must crack the seeds into small pieces before ingesting them into the blind sacs.

Reproductive biology

The peccary reproductive tract is bicornuate, and each uterine horn is short compared to swine, and coiled caudally. Peccaries have diffuse epitheliochorial placentation. Regarding reproductive cyclicity, peccaries are polyestrous and can breed and give birth year-round. Males are also believed to be fertile year-round. Peccary litter sizes range from one to a maximum of four precocial young, but the litter size of most peccaries is one to two. A litter the size of one is much more common than a litter size of four for all peccary species. Mating

in peccaries is not well known, but they may be promiscuous.

Conservation status

All Tayassuidae species are listed in CITES (Commission for International Trade of Endangered Species) appendices, and the Chacoan peccary is listed as Endangered by the IUCN. Hunting for bush meat is a major potential threat to peccaries. Determining sustainability of peccary harvests in hunted areas is crucial to future peccary conservation. Habitat loss, fragmentation, and degradation also pose severe threats to peccary populations. The World Conservation Union (IUCN) has established a Pig and Peccary Specialist Group. Group leaders convene and propose priority projects for study and of each species and harvest management. The IUCN Action Plan for human use of Tayassuidae advocates the development of rational and sustainable harvest management strategies by subsistence hunters, the improvement of local legislation and enforcement for harvest management, the outlaw of peccary hunting for solely commercial purposes, and to encourage the return of profits from the sale of subsistence

byproduct peccary hides. This Action Plan also recommends further studies of peccary population dynamics, harvest monitoring and trade, and the development of management programs.

Significance to humans

Tayassuids are hunted throughout their range for sport, subsistence bush meat, commercial bush meat, and for commercial sale of their pelts. They are some of the most common species used for bush meat in Central and South America. In lowland South America, peccaries are the first ranked game in terms of biomass in seven out of eight indigenous groups and three out of four mestizo communities. In 12 out of 13 studies of hunting by rural inhabitants of Amazonia, peccaries rank first in terms of mammalian biomass hunted. Their meat is consumed locally in rural sectors and sold in city markets. Peccaries are also an important source of income for many rural inhabitants of the Neotropics. Game meat comprises more than 11% of the financial returns for rural inhabitants of the Peruvian Amazon.

In addition to meat, peccary skins are also a significant source of income for many rural inhabitants of the Neotropics. From 1946–1966, more than three million collared peccary skins and more than 800,000 white-lipped peccary skins were exported from Iquitos, Peru, to international leather markets. Currently, peccary skins are in demand in Europe, particularly Germany, for the manufacture of fine leather goods such as gloves. However, in the past decade, the peccary hide trade has declined in South America. Currently, Peru is the only country exporting peccary pelts. Since the peccary pelt trade became less profitable and more controlled, the exportation of pelts fell to the current level of around 35,000 peccary skins per year.

In accord with their economic and subsistence importance, peccaries also play an important cultural role in many indigenous societies. Peccaries are often associated with spiritual guardians of game animals in many indigenous communities of Amazonia. Many indigenous peoples in the Neotropics have myths and other spiritual beliefs relating to peccaries, and raise wild-caught juvenile peccaries as pets.

Species accounts

List of Species

Collared peccary

Tayassu tajacu

taxonomy

Tayassu tajacu (Linnaeus, 1758), Mexico.

other common names

English: Javelina; French: Pecari a collier; German: Halsband-Pekari; Spanish: Sajino, chancho de monte, taitetu, jabalí, báquiro de collar, chácharo.

physical characteristics

The smallest of the three peccary species, and adult animals range in size from approximately 24–88 lb (11–40 kg). Have a characteristic "collar" of pale hairs that cross behind the neck and extend bilaterally in front of the shoulders. The rest of the body hairs are gray or black with some whitish rings on individual hairs. The dorsal snout is relatively narrow.

distribution

The most wide-ranging of the Tayassuidae occurring from the southwestern United States through Central America, the entire Amazon forest region, the Pacific coasts of Colombia, Ecuador, and Peru, and the dry tropical thorn forests (Chaco) of Paraguay, Bolivia, Brazil, and northern Argentina.

habitat

Exploit the widest range of habitats of the Tayassuids, from dry, open deserts, mesquite bosques, and oak forests in the

northern part of their range to moist and dry tropical forests in the southern part of their range. In the southwestern United States, they focus their activity in feeding areas, watering sites, and bedding grounds. In tropical forests of the Amazon, they use primarily moist terra firme habitat, and are uncommonly found in floodplain habitat.

behavior

The herd size varies depending on their habitat, from 5–30 in the deserts of the southwestern United States and 2–15 in Neotropical forests. Territorial, with relatively well-defined home ranges that vary depending on the habitat; there is little exchange of animals between herds. Although they have relatively stable territories, herds in the southwestern United States often separate into subgroups more than 328 ft (100 m) apart from one another. Occasional solitary peccaries may be observed, but is relatively rare. Diurnal, with the greatest feeding activity during the morning and late afternoon until dark; they live in mixed sex groups of adult males and females, juveniles, and sub-adults.

feeding ecology and diet

Primarily grazers and rooters, with much of their diet comprised of spiny cacti, succulent plants, a few shrubs, forbs, grasses, tubers, beans, seeds, herbivores, as well as some small lizards and mammals.

reproductive biology

Probably promiscuous. Ovarian cycles showed an average length of approximately 27.6 days. Behavioral estrus lasts for approximately of 5.7 days. Although lactational anoestrus does occur, an ovulatory postpartum oestrus has been observed. Males are fertile year-round, despite marked differences in rainfall between wet and dry seasons. Approximate age at first breeding is 16 months to two years. The average gestation period is 145 days. The average litter size is two fetuses/pregnant female.

conservation status

Legally hunted in many Central and South American countries and the United States, and are listed on Appendix II of CITES, which permits international trade in their products so long as harvests do not overexploit natural populations.

significance to humans

The most frequently exploited Tayassuidae, they are hunted from southern North America through to Argentina. Peccary meat is an important resource and popular game meat for many rural people in northeastern Peru as a source of both food and monetary income.

White-lipped peccary

Tayassu pecari

taxonomy

Tayassu pecari (Link, 1795), Cayenne, French Guiana.

other common names

English: Wari; French: Pécari à Lévres blanches; German: Weißbart-Pekari; Spanish: Huangana, pecari labiado, tropero, báquiro, puerco de monte, chancho de monte, chancho cariblanco.

physical characteristics

Adult animals weigh 55–88 lb (25–40 kg). Has a characteristic growth of white hairs under the jaw and around the mouth. Body hairs range from gray to black, and younger animals often have a reddish tan hair color. Have a relatively broad snout, and have the strongest jaws of the Tayassuidae.

distribution

Restricted to Central and South America, occurring from eastern and southern Mexico to Panama, through the Amazon regions of Colombia, Venezuela, the Guianas, Suriname, Brazil, Boliva, and Peru, the Atlantic forests and Pantanal of Brazil, and south through the tropical dry thorn forests (Chaco) of Paraguay, Brazil, and northern Argentina.

habitat

In tropical rainforests, they use lowland moist terra firme, wet terra firme, stream beds, and floodplain habitats. Often visit water sources such as ponds or streams and saltlicks, palm swamps, and wallow in mud. Excellent swimmers that tend to forage near trees, roots, and other objects, and also appear to search for seeds scatter-hoarded by rodents.

behavior

As herds move through their habitats, they give off a very strong odor, due to dorsal scent gland secretions. They make a great deal of noise by clacking their teeth, cracking palm nuts, snorting, grunting, and wheezing. They superficially graze the soil beneath the leaf litter as they forage. Herds are composed of males, females, and juveniles of various ages. They are diurnal, and forage and travel in the morning and afternoon, often resting near

water sources and saltlick wallows during the midday heat.

feeding ecology and diet

Primarily frugivorous, but they also supplement their diet with insects, particularly insect larvae and worms, carrion, amphibians, bird, and reptile eggs, and occasionally fish.

reproductive biology

May be polygynous. Ovarian cycles have an average length of 29.7 days. Behavioral oestrus has a mean of four days. Approximate age at first breeding is 1.5 years. The average gestation is 156–162 days. Breed and give birth year-around.

conservation status

Legally hunted in many Central and South American countries, and are listed on Appendix II of CITES, which permits international trade in their products so long as harvests do not overexploit natural populations. However, in many regions, populations are rare or have been locally extirpated due to over-hunting or habitat loss. Although they are locally abundant in some regions, they are apparently susceptible to the effects of hunting. Because they often travel in large herds, hunters may kill many animals at once. There are historical and anecdotal reports of indigenous peoples and colonists killing large numbers while herds were swimming across rivers.

significance to humans

Socially and economically significant to many indigenous peoples and colonists throughout the Neotropics; meat is also an important source of protein for indigenous peoples, particularly in lowland, non-flooded tropical forests. Pelts are also sold in the peccary skin trade. They also play an important role in indigenous mythology and beliefs. Indigenous peoples in Central and South America occasionally capture juvenile white-lipped peccaries and keep them as pets.

Chacoan peccary

Catagonus wagneri

taxonomy

Catagonus wagneri (Rusconi, 1930), Argentina. Believed to be a peccary species that became extinct during the Holocene, the Chacoan peccary was rediscovered by Western scientists in 1975. However, indigenous peoples in the Chaco have recognized the Chacoan peccary and other Tayassuids well before it was rediscovered. Chacoan peccaries are believed to be a relict species that survived in a thorn forest and scrub Pleistocene refugium.

other common names

English: Giant peccary; French: Pecari du Chaco; German: Chaco-Pekari; Spanish: Tágua, pecari del Chaco, chanco quimilero.

physical characteristics

Largest of the three peccary species, with adult animals weighing 66.1–94.7 lb (30–43 kg). They have relatively long, mule-like ears and large head, and well-developed sinuses, an apparent adaptation to dry, dusty environmental conditions. They have a whitish collar of hair that passes across the shoulder and extends under the chin. The rest of the body is covered in brown to gray hairs. They have relatively long bristles, giving them a shaggy appearance. Their legs are also relatively long.

distribution

The most limited geographic range among the Tayassuids, ranging in thorn forest and steppe of the Gran Chaco in Northern Argentina, southeastern Bolivia, and Paraguay.

habitat

Preferentially range in thorn forest. They show an apparent selection for forested ecosystems during wet and dry seasons, but also use open savannas to a lesser extent. They require areas of low rainfall and high temperatures, making their range the smallest of the three peccary species.

behavior

Live in stable, mixed-age and sex groups of approximately 4.5–4.6 animals; territorial. Often wallow in dust and/or mud. They are active from dawn to sunset, and move continally within their home range. Young have been observed to nurse from more than one female.

feeding ecology and diet

Cactus is a preferred food, likely due to its high water content. Although cactus is low in protein and high in oxalates, they can survive almost exclusively on this food source; they eat the giant cactus (Opuntia quimilo) known as quimil. The winter diet consists primarily of cactus (i.e., Cleistocactus baumannii, Eriocereus spp., Opuntia spp.), and flowers from Stetsonia coryune, Cereus validus, and Quiabentia chacoensi. They also eat a smaller amount of Acacia legumes, vine fruits, bromeliads, roots, and grubs, as well as soil from leaf-cutting ant mounds, likely due to its enriched mineral contents.

reproductive biology

Probably promiscuous. Reach sexual maturity at over two years of age, and have between two and three young per litter. The gestation period is five months. The farrowing season is between July and December, with most young born in August–September.

conservation status

The most endangered and least understood Tayassuid species, estimates are that the Paraguayan Chaco had a population of only 4,000 adult Chacoan peccaries in 1991. Unfortunately, little is known about Chacoan peccary population dynamics. The major threats are considered to be habitat loss to agricultural development, over-hunting, and disease. The species is listed as Endangered by the IUCN, and CITES classifies them as an Appendix I species, threatened by extinction. Appendix I species may not be traded between countries for commercial purposes. Chacoan peccaries may be exported and imported under highly regulated conditions for non-commercial purposes such as captive breeding in zoological collections. Recently, conservation groups and zoological parks have developed captive breeding programs for Chacoan peccaries in Paraguay, the United States, and Europe as a potential conservation strategy.

significance to humans

There is little information on the history of their socioeconomic and cultural significance. Despite their Endangered status, Chacoan peccaries are reported to be an occasional source of meat for local peoples and colonists in the Chaco.

Resources

Books

Bodmer, R., R. Aquino, P. Puertas, C. Reyes, T. Fang, and N. Gottdenker. Manejo y Uso Sustentable de Pecaríes in la Amazonía Peruana. Quito and Geneva: IUCN Sur and CITES, 1997.

Bodmer, R. E., L. K. Sowls, and A. B.Taber. "Economic Importance and Human Utilization of Peccaries." In Pigs, Peccaries, and Hippos, Status Survey and Conservation Action Plan, edited by W. L. R. Oliver. Gland, Switzerland: IUCN, 1993.

Emmons, L. H. Neotropical Rainforest Mammals, a Field Guide. Chicago: University of Chicago Press, 1990.

Sowls, L. K. The Peccaries. College Station: Texas A&M Press, 1997.

Periodicals

Altrichter, M., J. C. Saenz, E. Carrillo, and T. K. Fuller "Seasonal Diet of Tayassu pecari (Artiodactyla:Tayassuidae)in Corcovado National Park, Costa Rica." Revista de Biologia Tropical 48 (2002): 689–701.

Barreto, G. R., O. E. Hernandez, and J. Ojasti. "Diet of Peccaries (Tayassu tajacu and Tayassu pecari) in a Dry Forest of Venezuela." Journal of Zoology 1 (1997): 241–256.

Carrillo, E., J. C. Saenz, and T. K. Fuller. "Movements and Activities of White-lipped Peccaries in Corcovado National Park, Costa Rica." Biological Conservation 108, no. 3 (2002): 317–324.

Fragoso, J. M. V. "Home Range and Movement Patterns of White-lipped Peccary (Tayassu pecari) Herds in the Northern Brazilian Amazon." Biotropica 30 (1998): 458–469.

Kiltie, R. W. "Bite Force as a Basis for Niche Differentiation between Rain Forest Peccaries (Tayassu tajacu and Tayassu pecari)." Biotropica 14 (1982): 188–195.

Taber, A. B, C. P. Doncaster, N. N. Neris, F. H. Colman. "Ranging Behavior and Population Dynamics of the Chacoan Peccary, Catagonus wagneri." Journal of Mammalogy 74, no. 2 (1993): 443–454.

Theimer, T. C., and P. C. Keim "Phylogenetic Relationships of Peccaries Based on Mitochondrial Cytochrome b DNA Sequences." Journal of Mammalogy 79, no. 2 (1998): 566–572.

Nicole Gottdenker

Richard Bodmer, PhD