Tayassu pecari, Link, 1795

Don E. Wilson & Russell A. Mittermeier, 2011, Tayassuidae, Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona: Lynx Edicions, pp. 292-307 : 306-307

3.

White-lipped Peccary

Tayassu pecari

French: Pécari a levres blanches / German: WeilRbartpekari / Spanish: Pecari barbiblanco

Taxonomy. Sus pecari Link, 1795 ,

Cayenne, French Guiana.

Currently five subspecies are recognized, but these taxa have not been validated through DNA, and further populationlevel genetic and morphological research is needed. Cytogenetic research has revealed X-chromosome polymorphisms between specimens from the mid-western and south-western regions of Brazil, and those from Costa Rica; however, these have not been associated with subspecies distributions.

Subspecies and Distribution.

T.p.pecariLink,1795—Colombia,Venezuela,theGuianas,andNBrazil(NofAmazonRiver).

T.p.aequatoreLonnberg,1921—SWColombiaandWEcuador.

T.p.albirostreIlliger,1815—SBrazil,EPeru,Bolivia,Paraguay,andNArgentina(EJujuy,Salta,Formosa,Misiones,Chaco&SantiagodelEstero).

T.p.nngensMerriam,1901—SEMexicotoNicaragua.

T. p. spiradens Goldman, 1912 — Costa Rica to N Colombia.

Descriptive notes. Head-body 90.5-139 cm, tail 1.6-5 cm, shoulder height 40-60 cm, ear 6.4-9 cm, hindfoot 16.5-25 cm; weight 25-40 kg. No morphological dimorphism occurs between the sexes. There is one inguinal and one abdominal pair of mammary glands. The coat color varies with geographic region, ranging from dark reddishbrown to black, with white-cream coloration on the ventral region. The hair along the throat and distal rostrum is white; hence it is named the White-lipped Peccary. The pelage is bristly and there is a mane of long stiff hairs along the mid-dorsal line from the crown to the rump. When individuals encounter threats from predators or conspecifics they often erect the dorsal mane.

Habitat. Approximately 62% ofits distribution is within tropical and subtropical moist broadleaf forests (e.g. Costa Rica, Peru). Brazil alone encompasses 66% ofits total distribution. White-lipped Peccaries also utilize tropical and subtropical dry forest like the Gran Chaco of Bolivia and Argentina, savanna, dry grassland, seasonally inundated forest-savanna systems, and coastal mangroves. Their altitudinal distribution ranges from sea level to over 1900 m on the eastern slopes of the Andes. White-lipped Peccaries tend to frequent areas close to water (i.e. lakes, rivers, and wetlands) and may even visit coastal beaches to forage, for instance on turtle eggs.

Food and Feeding. White-lipped Peccaries utilize a wide variety of food resources including fruits, seeds, stems, leaves, tubers, roots, rhizomes, invertebrates, turtle and bird eggs, frogs, fishes, snakes, small mammals, and carrion. Behavioral observations, analyses of stomach contents, and feces indicate that fruit and seeds are primary food resources, followed by leaves and roots. Earthworms are also important, providing a significant protein source. Many trees drop their fruits underneath the canopy to attract frugivores, including herds of White-lipped Peccaries. They bulldoze through the leaf litter and topsoil, trampling juvenile plants, while searching for fruits and seeds. Their foraging behavior can affect understory plants and tree seedling survival, thus influencing forest structure and species composition. White-lipped Peccaries consume fruits and seeds from over 144 different plant species belonging to 38 families. In tropical moist forest the four most frequently utilized plant families are Arecaceae (24%), Fabaceae (14%), Moraceae (10%), and Sapotaceae (8%). In these habitats Arecaceae , the palm family, is their most important food, given its high abundance, highly nutritious fruit, and year-round availability, including during the dry season when food is scarce. White-lipped Peccaries have evolved a powerful mastication apparatus, with interlocking canines, thick enamel, and specialized muscles enabling a bite force of over 300 kg —the strongest amongst the peccary species. This allows them to exploit additional food resources and to destroy over 78% of the seeds they consume. This niche separation reduces interspecific competition between sympatric peccary species. White-lipped Peccaries also disperse seeds, especially of small-seeded species such as figs ( Ficus ). Given an up to three-day retention time in the gut, seeds that escaped mastication could be dispersed over distances of tens of kilometers. A few seeds, such as from moriche palm (Maunritia flexuosa), are too hard even for White-lipped Peccaries, and the animals only scrape off the fruit pulp, spitting out the seeds close to the parent tree. The peccaries may then trample the seeds into the muddy soil, protecting them from insect and fungal predation. This short-distance seed dispersal may lead to clumped species distribution, as seen in Mauritia palms. Many studies have demonstrated that trophic (seed predation, and long and short distance seed dispersal), and non-trophic (trampling and bulldozing of seedlings) peccary-plant interactions influence the recruitment, spatial distribution, and population dynamics of many plant species. By reducing competitive exclusion among plants, peccaries play an outsize role in maintaining and promoting botanical diversity in tropical forests.

Breeding. Breeding seasonality is highly variable and, among other factors, may depend on habitat and food resource availability. In Mexico and Suriname, Whitelipped Peccaries breed year-round. In Costa Rica and the Peruvian Amazon, however, peak births occur from June to September, coinciding with the fruiting season. Studies from captivity and the field indicate a gestation period of 156-162 days. The majority of births are twins; although litters of singletons and triplets are also found. The precocial neonates weigh 1.1-1. 4 kg, and have a reddish-brown coat. They stay close to their mother and are weaned around six months. Promiscuous suckling has been observed in captive White-lipped but not Collared Peccaries. Sexual maturity is reached between one and two years of age, and the life span in the wild is around 13 years.

Activity patterns. This species is primarily diurnal, with activity peaks in the morning and afternoon. However, White-lipped Peccaries extend orshift their activity towards dawn and dusk (crepuscular) and into the night when food resources are scarce (i.e. during the dry season), temperatures are high, and where human disturbance and hunting occurs.

Movements, Home range and Social organization. White-lipped Peccaries are the most gregarious terrestrial mammal species within the Neotropics. Herd sizes vary from five to over 400 individuals, with historical accounts of groups of over 1000 animals. Smaller herds tend to be found in areas with intense human hunting and/or relatively smaller fragments of suitable habitat. These peccaries often represent the highest mammalian biomass (230 kg /km?) in the ecosystems they inhabit, placing major demands on food resources and requiring continuous habitat to maintain populations. Groups contain male and female juveniles, subadults, and adults. Living in large groups helps them save energy as individuals avoid searching places recently foraged by conspecifics. It also reduces individual predation risk, and enables collective counterattacks against predators such as Jaguar (Panthera onca) and Puma (Felis concolor). Groups may break into smaller units and reunite later, depending on resource dispersion patterns. This fusion-fission behavior seems a mechanism to reduce interspecific competition for food. Genetic analyses of two populations of White-lipped Peccaries, separated by 80 linear kilometers in the southern Pantanal of Brazil, showed low levels of differentiation indicative of gene flow between these groups. However, more research is needed on population genetics, dispersal, group formation, corridor use, and migration in this species. Numerous abiotic and biotic factors, including habitat type, resource availability, and human disturbance affect their density, movement patterns, and home range size. In large protected tropical rainforests, such as Peru’s Manu National Park (over two million hectares), peccaries roam freely, and herd sizes can reach hundreds of individuals with home ranges encompassing over 200 km? Some researchers have considered White-lipped Peccaries nomadic or even migratory because herds may travel over 10 km /day. However, in fragmented landscapes in the Atlantic Forest, in southern Brazil, home ranges vary between 1879 ha and 2300 ha. Studies on smaller herds in Costa Rican tropical moist forest found home ranges between 2145 and 3780 ha; herds in the tropical semi-dry forest of the Calakmul Biosphere Reserve, Mexico, used ranges of 4360-12,100 ha. The greatest proportion of their daily activities includes feeding (34%) and moving between foraging patches (33%). Densities vary with habitat type and levels of human perturbation. In southern Brazil, densities varied between 3-9 ind/km? and 10-2 ind/km?, whereas at one site in the northern Brazilian Amazon a density of 139 ind/km? was found. In the dry forest of the Argentine Gran Chaco a density of 0-33 ind/km? was found in hunted sites versus 1-04 ind/km? in non-hunted sites. These results are similar to the 0-43 ind/km® documented in the semi-dry forest of the Calakmul Biosphere Reserve. Populations decline rapidly under intense hunting pressure, and can lead to local extinction. Like other peccaries, their vocal repertoire includes low-frequency barking, growling, moaning, high-pitched squealing, and tooth clacking. As groups forage and move through the understory, individuals emit soft vocalizations,facilitating group coherence. Alarm calls and teeth clashing are emitted when threats are detected; as multiple individuals emit alarm calls a threshold is reached at which point the whole group may take off, either together or scattering in subgroups, creating a confusing cacophony of sound and movement.

Status and Conservation. CITES Appendix II. Classified as Near Threatened on The IUCN Red List. They face multiple threats, foremost being wide-scale habitat destruction, and intensive subsistence and commercial hunting. Throughout most of their range they are a vital source of protein and income for local people. However, within the last century the distribution of White-lipped Peccaries has been reduced by over 21%. They have been driven to extinction in an area of over 1,095,666 km? and today have a high probability of long-term survival in only 47% of their range. Major population declines, including local extinction, have particularly occurred in savannas, grasslands, and tropical dry forest habitats. They are extinct in El Salvador, and their distributions have declined by over 89% in Costa Rica, 84% in Guatemala and Mexico, 63% in Argentina (recently has disappeared from Tucuman, Corrientes, and possibly Santa Fé), and 13% in Brazil. In Roraima State, northern Brazilian Amazon, there have been reports of declining densities; from 139-542 ind/km? in 1988 to 1:4-8-3 ind/km* in 1992, after a population crash probably due to an epizootic. Habitat destruction and hunting are threatening populations across 70% of their range. Conservation actions must focus on habitat protection and the regulation and management of hunting. Large continuous tracts of diverse habitats need to be conserved, and connectivity between existing fragments needs enhancementto facilitate dispersal and promote gene flow between isolated populations. Given their roles as ecosystem architects, and in structuring habitat across the Neotropics, their management and conservation needs to be founded on whole ecosystem management and precautionary principles. These principles, yet to be developed, are necessary to avoid further trophic meltdown and biodiversity loss. This species would be an ideal umbrella species for biodiversity conservation action and planning across the Neotropics.

Bibliography. Altrichter (2005), Altrichter & Almeida (2002), Altrichter & Boaglio (2004), Altrichter, Carrillo et al. (2001), Altrichter, Saenz & Carrillo (1999), Beck (2005, 2006, 2008), Beck & Terborgh (2002), Beck et al. (2010), Biondo et al. (2008), Bodmer (1991), Bodmer et al. (1997), Carrillo et al. (2002), Chebez (2008), Eisenberg & Redford (1999), Fragoso (1994, 1997, 1998a, 1988b, 1999), Giannoni et al. (1981), Hufty et al. (1973), Keuroghlian & Eaton (2008a, 2008b), Keuroghlian et al. (2004), Kiltie (1981a, 1981b, 1982), Kiltie & Terborgh (1983), March (1993), Mayer & Wezel (1987), Peres (1994, 1996), Peres & Lake (2003), Reyna-Hurtado et al. (2008), Sowls (1997), Taberet al. (2008).

Figures

1. Chacoan Peccary (Catagonus wagneri), 2. Collared Peccary (Pecari tajacu)

Subspecies and Distribution. T.p.pecariLink,1795—Colombia,Venezuela,theGuianas,andNBrazil(NofAmazonRiver). T.p.aequatoreLonnberg,1921—SWColombiaandWEcuador. T.p.albirostreIlliger,1815—SBrazil,EPeru,Bolivia,Paraguay,andNArgentina(EJujuy,Salta,Formosa,Misiones,Chaco&SantiagodelEstero). T.p.nngensMerriam,1901—SEMexicotoNicaragua. T. p. spiradens Goldman, 1912 — Costa Rica to N Colombia .