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VU - White-lipped Peccary

White-lipped Peccary (Tayassu pecari)

Status: Vulnerable A2bcde+3bcde ver 3.1
Pop. trend: decreasing


White-lipped Peccary
Taxonomy

Tayassu pecari Link, 1795

The name Tayassu pecari was first introduced by Fischer in 1814. The White-lipped peccary is currently the only species within the genus Tayassu.

The large geographic distribution of the White-lipped peccary across Central and South America through a range of habitats provides a potential opportunity to investigate the level of genetic diversity and differentiation within the species, as has been observed in the Collared peccary (Gongora et al. 2006). Thus, further research is needed to obtain a comprehensive understanding of the genetic diversity at the level of subspecies and/or species’ distribution. This would also provide critical background information to identify priorities for conservation at the genetic level. In addition, comprehensive and fine scale assessment of the confluence and potential venue for hybridization in countries such as Colombia, Brazil, and Peru, where more than one of the White-lipped peccary subspecies occurs would assist in this endeavor.


Subspecies and distribution
There has been some contrasting sub-classification (Mayer & Wentzel 1987; Grubb and Groves 1993). The latter authors propose five subspecies. These nominated subspecies are: T. p. pecari (Link, 1795) from French Guiana; T. p. aequatoris (Lönnberg 1921) found in Colombia and Ecuador; T. p. albirostis (Illiger 1815) distributed from Colombia through Argentina; T. p. ringens (Merriam 1901) found in Mexico through Nicaragua; and T. p. spiradens (Goldman 1912) which is distributed from Costa Rica through Colombia (Groves & Grubb 1993; Taber et al. 2011).


Present distribution
Distribution map White-lipped peccary

White-lipped Peccary is confined to the Neotropical region, from southeastern 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 1997). The species was introduced to Cuba in 1930 (Mayer & Wetzel 1987), but is no longer found in the wild there. It is presumed to be extirpated from El Salvador, and its range has been significantly reduced in Mexico, Central America, and South America in the last 20 years (Leopold 1959; Reyna-Hurtado et al. 2009, 2010; Altrichter et al. 2012).


Densities

White-lipped Peccary densities differ according to biome, habitat, fragmentation, and hunting. Desbiez et al. (2010) documented the importance of habitat in density estimates finding a density of 13.7 individuals/km² in forested areas and 3.0 individuals/km² in the open Cerrado. Keuroghlian et al. (2004) and Cullen (1997), both working in southern Brazil, found densities between 3.9 and 10.2 individuals/km² in tropical forest fragments. In the Brazilian Pantanal, Desbiez et al. (2004) found between 7.5 and 9.6 individuals/km² in two different locations. Altrichter (2005) estimated White-lipped Peccary densities in the Argentine Chaco between 0.33 individuals/km² in hunted sites and 1.04 individuals/km² in non-hunted sites. Reyna-Hurtado & Tanner (2007) found a similar density of 0.43 individuals/km² in the Calakmul Biosphere Reserve in Mexico.


Desriptive notes

White-lipped peccaries
White-lipped peccaries are medium-sized mammals, and superficially appear similar to some suids. However, numerous unique anatomical structures of their stomach suggest that peccaries are evolutionary more advanced than Old World suidae. The rostrum is elongated and terminates in a mobile naked disk, which includes the nostrils. Peccaries are macrosmatic and have relative small eyes with poor vision. The ears are erect, highly mobile, and oval in shape. All extant peccary species also have a unique scent gland that is located at the mid-dorsal line on the rump.

There are no obvious distinct morphological differences between genders (Mayer & Wetzel 1987; Sowls 1997; Keuroghlian 2003; Keuroghlian & Desbiez 2010). The pelage color varies with geographic region, the dorsum is typically dark red-brown to black, whereas the ventrum, medial part of legs, the throat, and distal rostrum are white-cream collared, hence: “White-lipped Peccary”. The pelage is bristly and both genders have a mane and long rigid hairs along their entire mid-dorsal section. When in distress or during aggressive behavior individuals can raise the mane (Mayer & Wetzel 1987; Sowls 1997). It is important to note that physical traits such as pelage color (adults black and subadults tan and black) and body measurements are not good indicators once the animal is over 2 years of age (Keuroghlian & Desbiez 2010). The authors further indicated that adults can been mistaken for subadults based on the presence of black and tan pelage (a trait thought to be characteristic of only young, juveniles and subadults). Similarly, animals that were all black, and assumed to be adults, were classified as sub adults upon examination of their teeth (Keuroghlian & Desbiez 2010).


Habitat

White-lipped Peccary ranges from Mexico to Argentina and over 60% of its distribution is within humid tropical forests. Though it can also be found in cloud forest, tropical moist forest, seasonally flooded palm savanna, wet and dry grasslands, mangroves, tropical dry forest, and dry thorn-forest (Mayer & Wetzel 1987; Altrichter et al. 2011; Taber et al. 2011). White-lipped Peccary occurs in habitats ranging from sea level to up to 2000 m in altitude including the eastern slopes of the Andes (Mayer & Wetzel 1987; Sowls 1997). Typically it prefers to be near waterbodies such as rivers, lakes, or water holes, especially during the dry season (Sowls 1997; Keuroghlian et al. 2008a; Reyna-Hurtado et al. 2015). Because this species lives in big cohesive groups with extensive home ranges, it requires large continuous area of undisturbed habitats (Fragoso 1998; Sowls 1997; Reyna-Hurtado et al. 2009, Altrichter et al. 2011; Taber et al. 2011, Keuroghlian et al. 2015). White-lipped peccaries are good swimmers and cross river frequently to extend their home range and activities.


Movements and home range

White-lipped Peccary travels large distances and long-term home range data indicate nonrandom seasonal movements (Keuroghlian et al. 2004; Keuroghlian et al. 2008). Key environmental variables determining movements and home range have been identified to be food availability and water sources (Altrichter et al. 2001, Keuroghlian et al. 2008a, 2008b; Reyna-Hurtado et al. 2009, 2012). Analyses of White-lipped Peccary’ s time budget (Altrichter et al. 2002) and the nutritional content of its diet (Lopez et al. 2006) suggest that low resource availability and poor nutritional quality of food affect their behavior and probably induce them to travel long distances in search of food in certain times of the year. However, home range size can also vary according to habitat, climate, and habitat fragmentation. Within the drier Cerrado ecosystem, the mean home range was 86.59 km2 (95% minimum convex polygon: MCP) in an isolated National Park (Almeida-Jácomo 2012). In the semi-arid Calakmul Biosphere Reserve, the mean home range size of 4 herds was 66.45 km2 (100% MCP) (Reyna-Hurtado et al. 2009). Keuroghlian et al. (2004) observed a herd home range of 29.51 km2 (100% MCP) for 4 subherds in an isolated Atlantic Forest fragment of 21.78 km2. In the Brazilian Amazon, mean home range of 2 herds was 57.41 km2 (100% MCP, Fragoso 1998, 2004), and Carrillo et al. (2002) reported a herd home range size of 35.33 km2 (100% MCP) for a single herd composed of several subherds in Costa Rica. In the wetlands of the Pantanal two White-lipped Peccary herds were compared, and home range area was 51% larger in an area that was deforested compared to the more pristine area (Keuroghlian et al. 2015). Reyna-Hurtado (2007) observed a high level of spatial overlap among subherds which varied with season and resources available (see also Carrillo et al. 2002; Keuroghlian et al. 2004; Almeida-Jácomo 2012).

Herds’ movement is performed at the “Landscape Scale” as described by Fragoso (1999) by covering several types of habitats. Reyna-Hurtado et al. (2012) found that groups move in a short scale (< 3 km) most of the time but occasionally do some long scale movements (>16 km) in the semi-dry Calakmul forest of Mexico. These movement patterns followed a log-normal distribution, a pattern described for species that need to visit specific habitats and feed on non-uniform spatially disperse food.

Many aspects of the movement patterns and the driving abiotic and biotic factors are still unknown. This may require multiannual study periods and probably spatial scales larger than the ones already studied. For example, indigenous communities in Bolivia report this species present and absent in approximately 10 years cycles (Vickers 1991). In other areas White-lipped Peccaries abandon established home ranges and travel further, as in Corcovado National Park, where occasionally groups leave the park and enter into the villages (Altrichter & Almeida 2002, 2009). Similarly, a radio-tracked herd in Calakmul moved out of its range for five months and later returned into the previous year home range (Reyna-Hurtado et al. 2009). Local large-scale extinctions of White-lipped peccary in pristine areas occurred in Brazil (Keuroghlian et al. 2012) and inside Manu National Park, Peru in 1970 and 2012 and it took approximately 10 years for the populations to recover. Richard-Hansen (2014) provided evidence of a population crash in French Guiana between 2000 – 2010. These extinction events occurred several times during the past decades. It is unknown what causes some of these extinction events, but a likely cause could be infectious diseases that may quickly spread over a large geographical scale (Fragoso 2004; Fang et al. 2008; Richard-Hansen et al. 2013). However, to date we have no data to test this notion.


Activity patterns

White-lipped Peccaries are primarily crepuscular, active early in the mornings and late afternoon (Mayer & Wetzel 1987; Sowls 1997). In Panama, White-lipped Peccaries also displayed nocturnal activity patterns (Mendez 1970). Whereas in Honduras, the species was primarily crepuscular, while during full moon nights they switched to more nocturnal behaviors (Idiaquez 1978). Activity patterns may be also altered with the presence of exotics, such as feral pigs (Galetti et al. 2015).


Feeding ecology

The majority of the peccaries’ s diet consists of fruits and seeds. In a review, Beck (2005) found that across its geographical range, White-lipped Peccary fed on over 144 plant species belonging to 36 families. The taxonomical rank order of consumed fruits and seeds of the five most important plant families consumed were Arecaceae (24%), Fabaceae (14%), Moraceae (10%), Sapotaceae (8%), and Chrysobalanaceae (8%). Interestingly, the palm family, Arecaceae, was the most frequently utilized taxa. The main reason for this might be that at the species level, palms have higher densities than trees and their highly nutritious fruits are available year around, including the dry season when fruits are generally scarce (Gentry 1988; Kahn 1991; Peres 1994; Henderson 1995, 2002; Terborgh et al. 1996; Pitman et al. 2001; Beck 2006). Despite that most palms have a hard-shelled endocarp that protects their seeds, peccaries coevolved a powerful mastication apparatus, with interlocking canines, thick enamel layer, and strong mastication muscles to generate a bite force of over 300 kg (Herring 1972; Kiltie 1981b, c, 1982), enabling them to pray upon most seeds (Beck 2006). Overall, White-lipped Peccary is able to destroy seeds of over 97 species from 124 plant species consumed.

However, White-lipped Peccary can also function as critical seed disperser via ingestions and defecation (endozoochory), especially for small or well-protected seeds such as Brosimun spp., Dipteryx spp., or Ficus spp. (Beck 2005, 2006). Other less well-known forms of seed dispersal include expectoration and epizoochory (Beck 2005). In the first case peccaries consume the fruit pulp but spit out the large or very hard seed which may later germinate (i.e. Mautitia flexuosa or Attalea spp.). Epizoochory occurs when seeds attach themselves via sticky saps or hooks to the fur of peccaries (i.e. Pharus virescens), and thus can be carried for long distances before falling off (Izawa & Kobayashi 1997).

Besides fruits and seeds White-lipped Peccary consumes a wide range of other resources, including leaves, tubers, roots, rhizomes, terrestrial invertebrates, turtle and bird eggs, frogs, fishes, snakes, and small mammals (citation in Beck 2005). Analyses of stomach contents and faces of peccaries indicated that fruits and seeds were the primary food resources followed by leaves and roots. Therefore, several authors consider peccaries to be primarily frugivorous or omnivorous (Kiltie 1981a; Bodmer 1989, 1991a, b; Altrichter et al. 2000, 2001; Keuroghlian & Eaton 2008a).

These myriad and unique peccary-plant interactions have critical ecological implications because they influence seed survival, predation, and dispersal, germination rates, species composition, diversity of recruiting plants, and the resulting canopy tree community (Beck 2005, 2006, 2007; Beck et al. 2013; Keuroghlian & Eaton 2009). Thus in forests where White-lipped Peccaries and other megafauna occur in lower densities or are extinct because of overhunting (“empty forest”) or habitat lost, the lack of these mammal-plant interactions can lead to dramatic changes in plant species composition and diversity, and unleash trophic cascading effects that not only affect plants but many animal species as well (Terborgh et al. 2008; Beck et al. 2013; Dirzo et al. 2014; Estes et al. 2011; Ripple et al. 2014; Galetti et al. 2015).


Reproduction and growth

Anecdotal observations (Sowls 1984) and research in the Peruvian Amazon (Gottdenker & Bodmer 1998; Mayor et al. 2009) indicate that White-lipped Peccary breeds throughout the year. Gottdenker & Bodmer (1998) suggest that this breeding pattern may be an outcome from the relatively constant rainfall and the resulting food availability year round. However, it seems that there is variation on breeding seasonality. In Central America, the species shows a distinct reproductive seasonality, with one mating peak in February and another in July (Altrichter et al. 2001). The greatest number of newborns and the peak in nursing activity were observed during July and August, when fruit availability was high (Altrichter et al. 2001). In Mexico the species has breeding seasons in April and November. Roots (1966) in Texas reported that captive White-lipped Peccary usually give birth during the rainy season.

The gestation period for White-lipped Peccary is about 180 days and the average number of offspring found from pregnant hunted animals in the amazon ranged from 1.6 to 1.7 young (Gottdenker & Bodmer 1998; Mayor et al. 2010). In the Amazon the parturition was 0.89/year (Mayor 2009). The young nurse for about six months and sexual maturity is reached between one and two years of age (Schmidt 1990; March 1993). A 1:1 fetal sex ratio was documented from pregnant individuals that were hunted in the Peruvian Amazon (Gottdenker & Bodmer 1998; Mayor et al. 2009). However, estimates based on data collected from free-ranging individuals consistently showed female-biased sex ratios. Analyzing the same herds and using paternity and maternity tests, Leite et al. (in prep.) observed that both males and females had offspring with more than one partner, showing a promiscuous genetic mating systems (Leite et al. 2013), as also observed for the Collared peccary (Pecari tajacu) (Coorper et al. 2011).


Behavior

White-lipped Peccaries display a variety of behavioral activities including foraging, ingestion, traveling, resting, wallowing, conspecific interactions, and swimming (Mayer & Wetzel 1987; Sowls 1997; Altrichter et al. 2002; Beck et al. 2010). In general, activity patters are influenced by abiotic factors such as seasonality, temperature, water availability and by biotic factors i.e. food resource availability, physiological condition of an individual, as well as by anthropogenic activities, primarily hunting, habitat destruction, and agriculture (Altrichter et al. 2002; Reyna-Hurtado et al. 2015).

Uncommon for forest dwelling ungulates, the White-lipped Peccary is highly social and lives in cohesive large groups. The strong musky scent that is emitted from the dorsal gland, is used for communication and marking territories (Mayer & Wetzel 1987; Sowls 1997). Individuals rub their lower jaw over each other’s dorsal gland and then via autogrooming distribute the scent across their bodies. These forms of reciprocal grooming lead to herd-specific scent (Schmidt 1990). White-lipped Peccary are also very vocal, including teeth clicking, low grunts, woofs, loud barks, and high frequency squeals. Vocalization also enhances group cohesion especially in dense understory vegetation (March 1993). These vocalizations can be heard over large distances and may be used for communication, group cohesiveness, and during distress. Schmidt (1990) also suggested that the white marking along the lower jaw might help in conspecific recognition, however considering some dense forests and their low light condition, its functions remains unclear.

Analysis of the species’ time budget found that White-lipped Peccary in a humid Costa Rican forest spend similar amount of daytime eating (30%) and moving (30%). The rest of the day was spent resting (28%), interacting with conspecifics (3%), and engaging in other activities. During low food availability, Altrichter et al. (2002) recorded a decrease in the time spent resting and more time foraging. Monthly variation in time spent on social interactions and the frequency of agonistic interactions seemed to be related to breeding (Altrichter et al. 2002).

Peccaries in general, but especially White-lipped Peccaries, can function as ecosystem engineers (Jones et al.1994) because they create and maintain water/mud wallows in the understory, and their rooting and foraging activities alter soil, litter and vegetation which may have cascading effects in tropical forests (Silman et al. 2003; Keuroghlian & Eaton 2009). Because of their frequent visits the soil in the wallows becomes compacted, thus their water remains longer than in natural occurring ponds (Beck et al. 2010). Wallows, especially those visited by larger White-lipped Peccary groups tend to be much bigger in size (i.e. 60 m2) than those used by the smaller family units of Collared Peccary (i.e. 10 m2). Thus wallows of White-lipped Peccary may almost represent small ponds and are critical breeding and foraging habitat for several anuran species especially during the dry season. In addition, many terrestrial bird species i.e. razor billed curacao, tinamous, and mammals such as tapir, bats, and even primate have been identified with camera traps visiting wallows either to forage or drink water (Beck pers obs).

Group size can vary largely, with anecdotal accounts of herds exceeding 2,000 animals (Mayer & Wetzel 1987), but typically containing 20-300 animals (Kiltie & Terborgh 1983; Mayer & Brandt 1987; Sowls 1997; Cullen et al. 2000; Reyna-Hurtado 2007; Desbiez et al. 2010; Keuroghlian et al. 2015; Reyna-Hurtado et al. 2015). Peccaries also display fission and fusion behavior and thus can form even larger groups. During certain seasons of the year, large herds can divide into smaller groups according to the distribution and abundance of food, although the more frequent reporting of smaller groups in some areas is probably correlated with increased hunting pressure (Ditt 2002).

Keuroghlian et al. (2004) observed that a herd of 150 White-lipped Peccaries in a fragmented area of Atlantic Forest periodically divided into 3–4 sub-herds with an average of 42 individuals each. They also observed a high frequency of switching of individuals among sub-herds and documented periodic sub-herd fusion and fission. Switching of individuals has also been observed in a current long-term (10 years) radio telemetry study in the Pantanal of Brazil (Keuroghlian et al. 2015; Keuroghlian et al. in prep), and in populations in Costa Rica (Sáenz & Carrillo 1999). In the Pantanal, Biondo et al. (2011) analyzed two different populations which were 80 km apart and found a low degree of genetic differentiation between the locations, and dispersal by both sexes (contrary to the predicted male-biased dispersal of most mammalian species (Greenwood 1980). In addition, 30% of males and females were predicted to be dispersers, which can indicate high levels of gene flow between the two different herds (Biondo et al. 2011). To test the fusion and fission between sub-herds, tests for genetic relatedness among individuals in these populations revealed a “super herd” consisting of many related individuals from different subherds (Rufo et al. in prep).


Status in the wild

Currently, the species is listed on Appendix II of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) and classified as Vulnerable on the IUCN Red List (Keuroghlian et al. 2013). The White-lipped Peccary is presently considered as endangered, or critically endangered, for some areas of Brazil and for all of Mexico by the environmental institutions of these countries (Brazil: Keuroghlian et al. 2012, Mexico: Secretaria de Medio Ambiente y Recursos Naturales-SEMARNAT, NOM-059, 2010).

The current range of White-lipped Peccary has been reduced by 20.5 percent from its historic distribution over the last 100 years (http://maps.iucnredlist.org/map.html?id=41778) including extirpations from entire countries i.e. El Salvador and Uruguay (Altrichter et al. 2012). In another 48% of its current range White-lipped Peccary remains but with reduced abundance and a low to medium probability of long-term survival. There have been major range declines in Argentina, Paraguay, southern Brazil, Colombia, Venezuela, north-east Brazil, Mexico, and Costa Rica (Altrichter et al. 2012; Keuroghlian et al. 2012). The species faces multiple threats across its range foremost wide-scale habitat destruction and degradation, commercial harvesting, unsustainable levels of subsistence hunting, and zoonotic diseases likely spread from domestic livestock (Keuroghlian et al. 2013). This species is particularly at risk in more xeric ecosystems, especially the Caatinga, Cerrado, and Pampas (Altrichter et al. 2012.). Forest fragmentation make White-lipped Peccary populations highly vulnerable to extinction because of the species’ requirement for high habitat diversity (Keuroghlian & Eaton 2008b), sources of water always available (Keuroghlian & Eaton 2008b; Beck et al. 2010; Reyna-Hurtado et al. 2012), diversity and abundance of fruiting sources (Beck 2005, 2006, Keuroghlian & Eaton 2008a; Keuroghlian et al. 2009), and relatively large home ranges (Fragoso 1998; Almeida-Jacomo 2004; Reyna-Hurtado et al. 2009). In addition, the White-lipped peccary social behavior of “protection in groups” makes it easy for hunters to spot and kill numerous individuals of the same group, especially with modern weapons (Peres 1996; Altrichter & Almeida 2002; R. Reyna-Hurtado, pers. obs.). The vulnerability of the species to human disturbance is especially problematic given the expanding industrial agriculture frontier and elongating road networks across the Neotropics that causes forest loss and fragmentation as well as allow hunters to access remote areas (Rivera 2014).


Text adapted from: Beck, H., Keuroghlian, A., Reyna, R., Altrichter, M. and J. Gongora 2017. White-lipped Peccary Tayassu pecari. In: M. Meletti and E. Meijaard, editors. Ecology, Conservation and Management of Wild Pigs and Peccaries. Cambridge, UK.


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