Tapirus terrestris, Linnaeus, 1758

Don E. Wilson & Russell A. Mittermeier, 2011, Tapiridae, Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona: Lynx Edicions, pp. 182-204 : 198-200

1.

Lowland Tapir

Tapirus terrestris

French: Tapir du Brésil / German: Flachlandtapir / Spanish: Tapir amazonico

Other common names: Brazilian Tapir, South American Tapir

Taxonomy. Hippopotamus terrestris Linnaeus, 1758 ,

“Habitat in Brasillia” i.e. Pernambuco, Brazil.

Four subspecies recognized.

Subspecies and Distribution.

T.t.terrestrisLinnaeus,1758—VenezuelathroughtheGuianastoCBrazilandNEArgentina(N&CMisiones).

T.t.aenigmaticusGray,1872—SEColombia,EEcuador,andNEPeru.

T.t.colombianusHershkovitz,1954—tropicalzonesofNColombiaintheareasofMagdalena,Bolivar,Atlantico,Cordoba,andNWAntioquia,asfarastheAtrataRiver,andasfarNastheRancheriaRiverinLaGuajira.

T. t. spegazzinii Ameghino, 1909 — SE Braz1l (Mato Grosso State), E Bolivia, Paraguay, and N Argentina to NE Santa Fe, NE Santiago del Estero, SE Jujuy, and E Salta.

Descriptive notes. Head-body 191-242 cm, tail less than 10 cm, shoulder height 83-118 cm (males), 83-113 cm (females); weight 180-300 kg. Female Lowland Tapirs are usually larger than males. Data collected from 35 Lowland Tapirs captured during a long-term telemetry study in the Atlantic Forests of Morro do Diabo State Park, Sao Paulo, Brazil, showed that the average weight of adult tapirs was 233 kg for females (200-300 kg) and 208 kg for males (180-280 kg). Measurements oftapirs in Morro do Diabo demonstrated that adult females are significantly longer andtaller than males. Full length offemales 208-242 cm, full length of males 191-223 cm; rear height of females 90-120 cm, rear height of males 89-109 cm. In captivity, newborn Lowland Tapirs usually weigh 3-2— 5-8 kg. Calves gain an average of2-27 kg per week and are completely weaned at four months of age. Growth is usually completed by 18 months of age. The color of adult Lowland Tapirs is blackish-brown dorsally with the ears edged in white; the chest, venter and limbs are dark brown; the cheeks are grizzled brown and gray. Young of all four tapir species are born dark with yellowish or white stripes and spots, a pattern that is lost after the first six months, although some vestiges of spotting may remain in young adults. The skin of Lowland Tapirs is thickest at the nape, and often covered by scars, scratches, and bruises; beneath the epidermis is a fibrous tissue layer. There is a well-developed sagittal crest that runs from the base of the muzzle to the middle of the back, which is derived from fat and soft tissues and covered by very long black hair. The short, erect mane is prominent, and is thought to help the tapirs escape predators, which seize the dorsum of the neck. This crest is not present in the other three tapir species.

Habitat. The Lowland Tapir has a broad geographic distribution and seems to be adapted to a wide range of habitat types. The species commonly inhabits tropical lowland South American moist and swamp forests, but also can be found in a wide range of other habitat types, including xeric Chaco and Cerrado forest, savanna wetlands, and lower montane forests, at elevations up to 2000 m. There are six main habitat categories where Lowland Tapirs can be found: tropical and subtropical moist broadleaf forests; tropical and subtropical moist to seasonally moist montane forests; tropical and subtropical dry forests, savannas, and shrublands; tropical and subtropical seasonally moist grasslands and savannas; montane grasslands; and mangroves. During a long-term Lowland Tapir field study in the Atlantic Forests of Morro do Diabo State Park, Sao Paulo, Brazil, tapirs strongly selected riparian environments and marshes, where they performed most of their main activities, particularly foraging. They avoided areas of agriculture and pasture land, as well as secondary growth forests. Palm forests are known to be important habitats, and they also frequent saltlicks, particularly in the Amazon. In the north-eastern region of the Brazilian Pantanal, tapirs show high preference for Acuri palm (Scheelea phalerata) forests. In the semi-deciduous Atlantic Forest, they prefer areas ofJeriva palm ( Syagrus romanzoffiana ) forests. The Jeriva is the most abundant and widespread palm species in the semi-deciduous Atlantic Forest. In the Amazon, patches of moriche palm (Mauritiaflexuosa) are used intensively by tapirs. During the study in Morro do Diabo State Park, tapirs moved fairly easily through areas of non-habitat between patches of forest. These pathways included pastures and croplands, especially sugar cane, soy bean, and corn plantations. In some instances, tapirs crossed areas of forest regeneration, degraded forest corridors, and riparian forests along small rivers. Lowland Tapirs also cross eucalyptus plantations between fragments of Atlantic Forest in Espirito Santo State, Brazil.

Food and Feeding. Due to their diet and stomach anatomy, Lowland Tapirs are closer to the browser side of the grazer-browser-frugivore continuum. There is some geographic variation in the composition of their diet. In the Peruvian Amazon, the species consumes 33% fruit and 66% leaf and fiber. The diet of Lowland Tapirs in the semideciduous Atlantic Forest consists of 37% fruit and 63% leaf and fiber. In the tropical rainforests of French Guiana their diet comprised 24-4% fruit, 22-3% leaf, and 53-2% fiber. In the Bolivian Chaco, the diet was 16-8% fruit, 62% leaf, and 21-2 % fiber. Tapirs selectively browse vegetative parts of different food plants. More than 170 species of fruits in over 60 different plant families have been identified as eaten by Lowland Tapirs throughout their range. In the Peruvian Amazon, they consumed 122 species of fruit from 33 plant families, and in the semi-deciduous Atlantic Forests of Brazil, they consumed 58 species of fruit from 23 plant families. Palm fruits are among the most important food resources, especially during the dry season when other species of fruit are less available. Mauritiaflexuosa is the single most important food item (76%) in the diet of Lowland Tapirs in the Peruvian Amazon. The distribution of these palm patches appears to strongly influence the movement patterns of tapirs in the area. In addition, tapirs are efficient dispersers of Mauritia flexuosa seeds. In Morro do Diabo State Park, Brazil, Syagrus romanzoffiana was the most frequently found fruit (18:4%) in Lowland Tapir fecal samples. In the Bolivian Chaco, Lowland Tapirs have successfully adapted to seasonal droughts, and to diets that include a large proportion of cactus fiber.

Breeding. There is very little information about the reproduction of Lowland Tapirs in the wild. Overall, the four species of tapirs have very slow reproductive cycles. Adult females usually produce a single offspring after a lengthy gestation period of 13-14 months (390-410 days). Twin births are very rare. Only one has been recorded in the AZA Studbook for Lowland Tapirs. Female tapirs are polyestrous, with an estrous cycle of 28-32 days. The estrous period lasts 1-4 days. Postpartum estrus is possible 9-27 days after the calf is born. Thus, a female Lowland Tapir may conceive within a month after giving birth. In the wild, under the best circumstances, in habitats exhibiting little seasonality in food availability, a young can be born every 14 months. In seasonally dry habitats, the interval between births may be longer. The interbirth interval for captive individuals can be as high as 18 months. Captive Lowland Tapirs are usually sexually mature by 14-24 months and conceive at an average of 3-7 years of age. Recent results coming from the AZA Studbook for Lowland Tapirs show that females reach sexual maturity at 19 months of age (1-6 years) and first reproduce at 32 months (2-7 years). In addition, the Studbook estimated that the maximum age of reproduction for the species is 15-3 years for females and 18-7 years for males. A Lowland Tapir Population and Habitat Viability Assessment (PHVA) Workshop held in 2007 modelled the dynamics of the species’ populations in the wild. Given that natural situations impose a toll on the animals, age offirst reproduction in the wild was estimated to be four years, and maximum age of reproduction 20 years for both females and males. Thus, the generation length of wild Lowland Tapirs was estimated to be eleven years. The sex ratio at birth in captivity is 3:1. The sex ratio observed in Morro do Diabo, Brazil, was 1:1-33. Monitored subsistence hunting among the Waimiri Atroari Indians in the central Amazon of Brazil recorded a sex ratio of kills of 1:1:03. Therefore, the sex ratio at birth in the wild is generally assumed to be 50% males and 50% females. Data on Lowland Tapir longevity comes from captivity and reveals that these animals live over 35 years.

Activity patterns. Tapirs are predominantly nocturnal and crepuscular. Lowland Tapir rest during the day and begin their activity after sunset, at around 18:00 h. In the semi-deciduous Atlantic Forests of Morro do Diabo State Park in Brazil, their main periods of activity are from 19:00 h to 00:00 h, with a peak between 20:00 h and 21:00 h (63% active); and from 01:00 h to 07:00 h, with a peak between 05:00 h and 06:00 h (60% active). Tapirs in Morro do Diabo are largely inactive between 11:00 h and 16:00 h. Overall, tapirs in Morro do Diabo foraged for approximately five hours in the early evening, and then seemed to rest for a few hours in the middle of the night. There followed a second foraging bout, of approximately six hours in the early hours of the day. Tapir activity patterns were analyzed separately for wet and dry season, as well for both sexes and different age classes in Morro do Diabo, and were fairly consistent. In Kaa-Iya del Gran Chaco National Park, Bolivia, tapirs are mostly active between 18:00 h and 22:00 h, and between 00:00 h and 06:00 h, with a main activity peak between 01:00 h and 06:30 h, and very little activity between 11:00 h and 15:30 h. In the Peruvian Amazon, tapir activity peaks were observed between 19:00 h and 20:00 h and between 03:00 h and 04:00 h, with the main foraging time between 21:00 h and 03:00 h. Nocturnal line-transect sampling in Morro do Diabo State Park allowed for an analysis of the effect of moonlight intensity on tapir activity. Approximately 47% of the tapir encounters occurred during the waning crescent, 21% during the new moon, 18% during the waxing crescent, and only 14% during the full moon. It was clear that tapirs in Morro do Diabo were considerably less active during the brighter phases of the moon. Previous studies suggested that a lower level of animal activity during moonlit nights is a result of higher predation risk. In El Rey National Park, Argentina, tapirs were typically diurnal, perhaps due to lack of human disturbance.

Movements, Home range and Social organization. L.owland Tapirs can move fairly easily through areas of low quality habitat (e.g. agricultural crops, pasture lands, and eucalyptus plantations) to get from one patch of forest to another. Parameters of Lowland Tapir spatial ecology reported in the literature, particularly home range size, vary widely. Tapirs are large-bodied, wide-ranging mammals that usually require considerably large home ranges. In the semi-deciduous Atlantic Forest of Morro do Diabo State Park, Brazil, tapirs use very large home ranges (4-67 km?, varying from 1-12 km? to 14-19 km?®). These home ranges have very complex internal structures, including multiple core areas, which comprise a very small proportion of the home range (50% core area = 17% of the home range; 25% core area = 6% of the home range). Little seasonal variation in size and location of home ranges and core areas was observed. These patterns are consistent for both sexes and different age classes. In the Peruvian Amazon, the home ranges of Lowland Tapirs are much smaller, ranging from 1-06 km? to 3-86 km?®, with an average of 2: 61 km ®. In Kaa-lya del Gran Chaco National Park, Bolivia, tapirs had an average home range size of 2: 48 km?*Neighboring tapirs in Morro do Diabo State Park showed very strong home range overlap between individuals (around 30%, although in some cases as high as 92%). This included strong overlap between intra and intersexual pairs, as well as between pairs of same and different age classes. Estimates of Lowland Tapir population density range from a high of 1-6 ind/km? reported for the species in Neotropical forests in undisturbed, non-hunted, or lightly hunted sites to less than 0-3 ind/km? in other regions and habitat types. In Amazonian forests, Lowland Tapirs are usually found in densities around 0-5 ind/ km?. Exceptionally high densities (3:3-3-7 ind/km?) were observed in the Amazonian dry forests of Roraima, Brazil. In the Brazilian Pantanal, tapir densities range from 0-13 ind/km? in open grassland habitat to 0-4 ind/km? in forested habitats. In the semi-deciduous Atlantic Forest, densities range from 0-21 ind/km? to 1-35 ind/km?. This wide variation in density estimates can be explained by several factors, including differences in environments, habitat types studied,levels of habitat conservation, levels of hunting, and, most importantly, differences in the methods used to estimate densities. Another factor is that the Lowland Tapir, although generally rare and elusive, can be locally common (e.g. around saltlicks, in palm forests, and near permanent water sources). Great variation in density can also reflect the ability of tapirs to adapt to different habitat types and availability of resources (food and water). Tapirs are primarily solitary. Tapir offspring normally remain with their mother for approximately twelve months. In Morro do Diabo, from a total of 81 tapir sightings obtained through nocturnal line-transect sampling, 77-78% were of one individual and 12-35% of pairs (adult female/adult male, adult female/offspring). On one occasion, three individuals were sighted. In terms of intraspecific interactions and social organization, tapirs in Morro do Diabo showed a strong home range overlap (30%) as well as overlap of core areas of use (20%) between neighboring individuals. This included strong overlap between intra and intersexual pairs, as well as between pairs of same and different age classes. The great majority of the individuals shared varying portions of their home ranges with several other individual tapirs. Thus, while tapirs in Morro do Diabo had well-defined home ranges and lived within their boundaries, no evidence was found that they defended their areas against conspecifics. In the Peruvian Amazon, Lowland Tapirs monitored through GPS telemetry regularly walked along the boundaries of their home ranges, which appeared to provide some evidence that they were defining their territory against other individuals by maintaining clear home range boundaries.

Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. Although the species has been listed as Vulnerable across its entire range, several populations in Argentina, Brazil, Colombia, and Venezuela present considerably higher levels of threat. A few examples include tapir populations in the Atlantic Forest and Cerrado biomes in Brazil, as well as Colombian and Venezuelan populations in Catatumbo, Maracaibo, and Lake Valencia watershed. The Lowland Tapir has the broadest geographic distribution of the four living species of tapirs and the species occurs in 21 different biomes in eleven countries. Historically, this species was found east of the Andes and north of the Espinal grasslands and shrublands of Argentina, throughout the Chaco, Pantanal, Cerrado, Llanos, Caatinga, Atlantic Forest, and Amazonian/Orinoco biomes. The historic distribution of the species covered approximately 13,129,874 km*. Nevertheless, populations have been severely reduced and are currently often limited to forested biomes and wetlands. The species is believed to have gone extinct in approximately 14% ofits range and the current distribution declined to 11,232,018 km*. The species has been extirpated from the dry inter-Andean valleys of the northern Andes and is becoming increasingly rare along the agricultural frontiers that are sweeping through parts of the western and southern Amazon basin. In Brazil, which constitutes a large portion of its range, the Lowland Tapir has disappeared from over one million km? (12-4% of its countrywide range). Although only about 15-20% of the Amazon has been deforested in the past 30 years, 85-90% of the Atlantic Forest has disappeared and 40% of the Pantanal has been converted to human use. Most of the Cerrado and Caatinga biomes in Brazil have been converted to agriculture and cattle ranching. As a consequence, the Lowland Tapir has been extirpated from the Caatinga; most populations in the Cerrado are small and in protected areas where illegal hunting is minimal. Some exceptions include remote areas of Cerrado (e.g. Chapada das Mangabeiras, Jalapao region in Tocantins State) where tapirs are still common. The Lowland Tapir is now either completely absent or severely fragmented across much of its historic range, with the Northern and Central Amazon as well as the remaining Pantanal ( Bolivia, Brazil, and Paraguay) becoming important strongholds as southern, eastern, and north-western populations are declining rapidly. The IUCN Red Lust published in 1996 listed Lowland Tapirs as Lower Risk/Near Threatened. Therefore, the species has deteriorated in Red List status over a period of twelve years. There is an ongoing reduction of Lowland Tapir populations estimated to be slightly greater than 30% in the past three generations (33 years). This rate of decline is predicted to continue for the next three generations. The main identified threats responsible for the decline include habitat deforestation and/or alteration; habitat fragmentation (resulting in small populations and low connectivity); hunting; cattle ranching; infectious diseases; road-kill; fire; human density; plantations of monocultures (sugar cane, soy bean); lack of patrolling of protected areas; small size of protected areas; resource extraction; and impact of tourism. Hunting is one of the most important threats. Tapirs are among the preferred game species for subsistence and commercial hunters throughout the Amazon. Estimates of tapir harvest in the State of Loreto in the Peruvian Amazon range from 15,447 to 17,886 individuals per year. Due to their individualistic lifestyle, low reproductive rate, long generation time, and low population density Lowland Tapirs rarely achieve high local abundance, which makes them highly susceptible to overhunting, and populations show rapid decline when harvested. There are a number of infectious diseases (Bluetongue, Equine Encephalitis, Infectious Bovine Rhinotracheitis, and Leptospirosis) and parasites known in Lowland Tapir populations in the Atlantic Forest and Pantanal biomes in Brazil. These diseases spread to tapirs from domestic livestock, particularly cattle and horses, and can potentially increase tapir mortality and affect reproductive rates. Another serious threat to this species is road-kill. Morro do Diabo State Park in Sao Paulo, Brazil, is crossed by a highway that, from 1996 to 2006, killed an average of six tapirs per year. Most of the tapirs killed were adult individuals capable of breeding. Road-kill is also a serious threat in the Cerrado and Pantanal biomes of Brazil. Estimates of the total population size for the species throughoutits entire range are not available. This species occurs in numerous protected areas across its range. However a large proportion of the total Lowland Tapir population is found outside the boundaries of legally protected areas, where tapirs are hunted, chased by dogs, and face many other threats. Although the species is protected legally in most countries, hunting laws are seldom enforced and have proven ineffective.

Bibliography. Affonso (1998), Aquino et al. (2001), Aquino & Calle (2003), Ayala (2002, 2003), Barongi (1986, 1993), Bodmer (1990a, 1990b, 1991a, 1995), Bodmer & Lozano (2001), Bodmer et al. (1997), Brooks et al. (1997), Chiarello (1999), CITES (2005), Cordeiro (2004), Cullen et al. (2000), Desbiez (2009), Eisenberg (1989, 1997), Emmons (1990), Emmons & Feer (1997), Flesher (2007), Fradrich & Thenius (1972), Fragoso (1997), Fragoso & Huffman (2000), Fragoso et al. (2000), Galetti et al. (2001), Gomez et al. (2005), Goncalves da Silva (2007), Haugaasen & Peres (2005), Henry et al. (2000), Herrera et al. (1999), Hershkovitz (1954), Hunsaker & Hahn (1965), Janson & Emmons (1990), Maffei et al. (2002), Mallinson (1969, 1974), Mangini (2001), Mangini & Medici (2001), Mangini et al. (2000), Medici (1999, 2001, 2002, 2010), Medici et al. (2007), Mendes-Pontes (2004), Ministerio del Medio Ambiente de Colombia (2002), Montenegro (1998, 1999, 2004), Montenegro et al. (2000), Naveda et al. (2008), Noss et al. (2003), Novaro etal. (2000), Olmos (1997), Olmos et al. (1999), Padilla & Dowler (1994), Pena et al. (1996), Peres (2000), Robinson & Redford (1986, 1991), Rodrigues et al. (1993), Salas (1996), Salas & Fuller (1996), Schaller (1983), Schipper et al. (2008), Soto (2002), Souza-Mazurek et al. (2000), Taber et al. (2008), Tobler (2008), Tobler et al. (2009), Téfoli (2006), Trolle (2003), Trolle et al. (2007), Velastin et al. (2004), Vickers (1991), Vié et al. (2009), Young (1961).

Figures

1. Lowland Tapir (Tapirus terrestris), 2. Central American Tapir (Tapirus bairdii), 3. Mountain Tapir (Tapirus pinchaque), 4. Malayan Tapir (Tapirus indicus)

Subspecies and Distribution. T.t.terrestrisLinnaeus,1758—VenezuelathroughtheGuianastoCBrazilandNEArgentina(N&CMisiones). T.t.aenigmaticusGray,1872—SEColombia,EEcuador,andNEPeru. T.t.colombianusHershkovitz,1954—tropicalzonesofNColombiaintheareasofMagdalena,Bolivar,Atlantico,Cordoba,andNWAntioquia,asfarastheAtrataRiver,andasfarNastheRancheriaRiverinLaGuajira. T. t. spegazzinii Ameghino, 1909 — SE Braz1l (Mato Grosso State), E Bolivia , Paraguay , and N Argentina to NE Santa Fe, NE Santiago del Estero, SE Jujuy, and E Salta.