Overview

Distribution

Range Description

A wide-ranging species occurring thoughout much of the bush savanna (cerrado) and dry forests of central Brazil, south from middle and upper reach of the Amazonian tributaries, the rios Juruena, Xingu and Araguaia. It extends east of the Midde and upper Rio São Francisco in the west of the states of Minas Gerais and Bahia, occupying humid forest (known as brejos, resulting from orographic rainfall), semideciduous and deciduous (floresta de caatinga) and gallery forests. It extends into eastern Paraguay and in parts of the southern and central regions of the Paraguayan chaco, west of the Río Paraguai (Stallings 1985; Stallings et al,1989). Brown and Zunino (1994) descirbed its range in north-eastern Argentina, occurring in eastern Formosa and Chaco, the extreme north-eastern section of Santa Fe, northern Corrientes and the southern part of Misiones. Alouatta guariba occurs in the Atlantic forest remnants of northern Misiones. In Bolivia, this species is found exclusively east of the Rio Beni and is absent from true Amazonian forests of northern Beni and Pando. It is patchily distributed in the rest of lowland tropical Bolivia (R.B. Wallace pers. comm. 2007). Wallace et al. (2000) recorded it from Pajaral, east of the Rio Blanco, along with A. sara. A. sara they found in low densities (0.1 groups encountered per 10 km) in floodplain forest, whereas A. caraya was restricted to semideciduous forest patches.

Aguiar et al. (2007) recorded sympatry and probable hybridization with A. guariba in riparian forest along the left margin of the Rio Paraná (between Porto Figueira and Port Camargo) in the Ilhas e Várzeas do Rio Paraná Environmental Protection Area on the state of Paraná. This region is considered to be an ecotone between Cerrado (the typical domain of A. caraya) and the Atlantic forest (A. guariba). In São Paulo, it is restricted to the right (west) bank of the Rio Paraná above the mouth of the Rio Paranaiba.

Villalba et al. (1995) reported on its probable presence in the past, and possible existence still, in the extreme north-west of Uruguay.
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Geographic Range

Alouatta caraya are found in the rainforests of central South America ranging through eastern Bolivia, southern Brazil, Paraguay, and northern Argentina.

(Walker 1999)

Biogeographic Regions: neotropical (Native )

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Physical Description

Morphology

Physical Description

Alouatta caraya are sexually dimorphic where males average 6.7 kg and females average 4.4 kg. Male body size ranges from 1.7 to 2.2 ft with tails of similar length to their body. Females' bodies average 1.6 ft with tails slightly longer than their bodies. A. caraya are also sexually dichromatic. Males usually have black hair, which gives the species the common name of Black Howler monkey. Females however have more yellow-brown or olive colored hair. Infants are born with a golden coat, which changes as the animal matures. A. caraya have long, strong prehensile tails. These tails are hairless on the underside, which allows them to be sensitive to touch and act in identifying things, much like a 5th hand. The black face is mostly hairless as well, with slightly bushy eyebrows. A. caraya have brown, medium sized eyes set in a frontal position. The muzzle is prominent and the nostrils close together. Like other howlers they have enlarged hyoid and larynx housing the vocal apparatus where the distinctive howling originates. A. caraya however, have less prominent lower jaw and bulging neck than some howlers.

(Welker et al. 1990, Walker et al. 1999, Bicca Marquez and Calegaro Marquez 1998)

Range mass: 4 to 10 kg.

Sexual Dimorphism: male larger

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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
Alouatta caraya is found in semi-deciduous forests. Today, it occupies forest patches throughout a large part of the Cerrado of central Brazil, and the Pantanal of Mato Grosso. It occupies deciduous forest (caatinga alta) in some areas in southern Piaui, western Bahia and north-western Minas Gerais. In Argentina, it specializes in semi-dry deciduous forests and gallery forests as well as the typical island forests of the Chaco (Arditi and Placci 1990; Brown and Zunino 1994). In Bolivia, its found in the Chaco and Chiquitano forests, and gallery forest and forest islands in the Beni. They can be found in secondary forest and can adapt to disturbed and degraded habitats. In Paraguay, Stallings (1995; Stallings et al. 1989) reported that A. caraya occurs throughout the eastern forests and gallery forests of the southern and western Chaco along the ríos Paraguai and Pilcomayo, and the numerous small rivers that drain the chacoan plain. It can also be found in sporadic xeric forests, continuous with gallery forests. In Rio Grande do Sul, A. caraya occurs in mixed evergreen forest (Araucaria), forest patches in savanna, and seasonal deciduous forest (Codenotti et al. 2002; Codenotti and Silva 2004).

The howler monkeys are the large leaf-eaters of the South American primate communities. The molar teeth are particularly adapted for their chewing leaves through shearing. They spend up to 70% of their day lying and sitting about quietly among the branches, fermenting leaves in their enlarged caecums. Like the spider monkeys, they are prehensile-tailed, with a naked patch of skin on the under surface at the tip. Their most characteristic feature is the deep jaw which surround the enlarged larynx and hyoid apparatus, a resonating chamber. It is with this enlarged and highly specialized voice box that they produce their howls (grunts, roars and barks). Howling sessions, usually involving the entire group, can be heard particularly in the early morning and are audible at distances of 1-2 kms (Drubbel and Gautier 1993).

Howlers are the only New World primates that regularly include mature leaves in their diet, although softer, less fibrous, young leaves are preferred when they are available. Their folivory and ability to eat mature leaves is undoubtedly one of the keys to their wide distribution and the wide variety of vegetation types they inhabit. Mature fruit is the other important food item, especially wild figs (Ficus) in many regions, but they also eat leaf petioles, buds, flowers (sometimes seasonally very important), seeds, moss, stems and twigs, and termitaria.

Behavioural ecological field studies have been carried out by Zunino (1986) and collaborators (Zunino et al. 1996, 2001) in Argentina and Bicca-Marques (1994; 2003; Bicca-Marques and Calegaro-Marques 1994) in Brazil (see also review by Neville et al. 1988).

Size:
Adult male weight mean 6.42 kg (n=58), adult female weight mean 4.33 kg (n=117) (Rumiz 1990)

Systems
  • Terrestrial
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Habitat

Alouatta caraya range varies from tropical semi-deciduous gallery forest where rains are nearly constant throughout the year, to tropical deciduous forest spotted with savanna like openings where there is a marked wet, warm season and a dry, cool season. A. caraya require forests with diverse species of plant life to supply their dietary needs. Much of their habitat is currently being diminished by destruction of these forest types. (Welker et al. 1990, Rodrigues and Marinho-Filho 1995, Kowalewski and Zunino 1997)

Terrestrial Biomes: savanna or grassland ; forest ; rainforest

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Trophic Strategy

Food Habits

Alouatta caraya are folivorous. They eat mostly leaves but do compliment their diet with fruits, buds and flowers. A. caraya rarely come down from the trees since their food source is entirely in the canopy and their water needs are met by their food. However in especially dry times they will come down to drink water in lakes or supplement their diet with marsh-living herbaceous plants

(Erwin and Mitchell 1986, Welker et al. 1990, Rodrigues and Marinho-Filho 1996)

Plant Foods: leaves; fruit; flowers

Primary Diet: herbivore (Folivore )

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Life History and Behavior

Life Expectancy

Lifespan/Longevity

Average lifespan

Status: captivity:
20.0 years.

Average lifespan

Status: captivity:
20.3 years.

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Lifespan, longevity, and ageing

Maximum longevity: 32.4 years (captivity) Observations: One wild born female was about 32.4 years old when she died in captivity (Richard Weigl 2005).
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Reproduction

Reproduction

The mating system of Alouatta caraya appears promiscuous among the members of the group.

Mating System: polygynandrous (promiscuous)

Gestation length for A. caraya is 187 days. Studies have shown that younger females have gestation length of 10 to 12 months where more mature mothers have gestation length of only 7-10 months. Females give birth to one offspring per birth and care for infants for about one full year before mating again. Infants are about 125 g at birth.

(Welker and Schafer-Witt 1990, Shoemaker 1979)

Breeding interval: Females breed once per year

Average number of offspring: 1.

Average gestation period: 187 days.

Average time to independence: 12 months.

Key Reproductive Features: iteroparous ; gonochoric/gonochoristic/dioecious (sexes separate); viviparous

Average birth mass: 187.5 g.

Average gestation period: 187 days.

Average number of offspring: 1.

Average age at sexual or reproductive maturity (male)

Sex: male:
928 days.

Average age at sexual or reproductive maturity (female)

Sex: female:
1167 days.

Females care for their young for about 12 months after they are born. Female offspring remain in their natal group and therefore stay with their mother long after they are independent.

Parental Investment: pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Female); post-independence association with parents

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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
LC
Least Concern

Red List Criteria

Version
3.1

Year Assessed
2008

Assessor/s
Fernandez-Duque, E., Wallace, R.B. & Rylands, A.B.

Reviewer/s
Mittermeier, R.A. & Rylands, A.B. (Primate Red List Authority)

Justification
This species is listed as Least Concern considering its large range, presence in several national parks, and ability to adapt to modified habitats. At present, it is not likely that the species, while declining, warrants listing in a threatened category under criterion A. Although its habitat is very fragmented, populations can live in relatively small areas and disturbed forest.

History
  • 2003
    Least Concern
    (IUCN 2003)
  • 1996
    Lower Risk/least concern
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Conservation Status

Alouatta caraya are threatened by clear-cutting and selective logging since they are heavily reliant on the biodiversity of predominantly primary forests for their diet. Some populations are more threatened than others. According to the Priority Primate Conservation Projects for the Neotropical Region from the Revised Global Action Plan for Primate Conservation, A. caraya, in the Argentine provinces of Formosa, Misiones, Salta and Corrientes are threatened and a high priority for conservation. Hunting pressure on A. caraya ranges from moderate in locations such as San Jose, Bolivia to none in northern Argentina.

(Welker et al. 1990, Mitchell and Erwin 1986, Peres 1997)

US Federal List: no special status

CITES: appendix ii

IUCN Red List of Threatened Species: least concern

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Population

Population
Although widespread, Alouatta caraya is patchily distributed, and densities vary widely. In Argentina, Brown and Zunino (1994) recorded high densities in a number of sites: Chaco forest, Formosa 111 individuals/km²; Chaco forest, Corrientes 90 individuals/km²; gallery forest, 63 individuals/km²; and inundated forest 283 individuals/km². Arditi and Placci (1990) carried out surveys in gallery forests of the Chaco in Argentina (Riacho Pilagá, Estancia Guaycolec) and found lower numbers than those reported by Brown and Zunino (1994) resulting from surveys in 1980s: 11.7 individuals/km². Dvoskin et al. (2004) repeated the surveys there in 2001 and found that numbers had increased to 26 individuals/km². The highest densities occur in flooded forests and there have been numerous surveys of Alouatta caraya in this forest type (Pope 1968; Thorington Jr et al. 1994; Rumiz 1990; Zunino et al. 1996, 2001). Codenotti et al. (2002; see also Codenotti and Silva 2004) conducted a state-wide survey of remnant A. caraya populations in Rio Grande do Sul, Brazil. Populations were found to be small, isolated, mostly single groups, and densities were consistently low. Of 13 localities, only three had densities above 2 individuals/km² (Lageado do Celso 6.5 individuals/km²; a location in the municipality of Santiago 4.0 individuals/km²; and an urban park in the municipality of São Francisco de Assis 2.3 individuals/km²).

Population Trend
Decreasing
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Threats

Threats

Major Threats
This species is threatened by habitat loss due to agricultural development for soy and cattle ranching in the Brazilian cerrado, soy in the Bolivian Chiquitano, and small-scale farms and cattle ranching in Argentina. Some subsistence hunting occurs across its range.
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Management

Conservation Actions

Conservation Actions
Alouatta caraya occurs in numerous protected areas:

Argentina
Iguazú National Park (55,000 ha) (Brown and Zunino 1994)
Pilcomayo National Park (60,000 ha) (Brown and Zunino 1994)
Chaco National Park (14,000 ha) (Brown and Zunino 1994)

Bolivia
Kaa-Iya Gran Chaco National Park (3,441,115 ha)
Otuquis Pantanal National Park (903,350 ha)
Otuquis Natural Area of Integrated Management (102,600 ha)
San Matías Natural Area of Integrated Management (2,918,500 ha)
Noel Kempff Mercado National Park (1,500,000 ha) (Wallace et al. 1998)
Ríos Blanco y Negro National Reserve (1,423,900 ha) (Wallace et al. 2000)

Brazil
Araguaia National Park (557,726 ha) (in range)
Brasília National Park (31,891 ha) (Santini 1986)
Chapada dos Veadeiros National Park 965,034 ha) (in range)
Grande Sertão Veredas National Park (241,000 ha) (in range)
Chapada Diamantina National Park (152,105 ha) (in range)
Pantanal Matogrossense National Park (136,046 ha)
Taiamá Ecological Station (914,300 ha)
Ibirapuitã State Biological Reserve (351 ha) (Marques 2003)
Ibirapuitã Environmental Protection Area (318,000 ha) (Marques 2003)
Ilha Grande National Park (108,166 ha) (Aguiar et al. 2007)
Ilhas e Várzeas do Rio Paraná Environmental Protection Area (1,003,059 ha) (Aguiar et al. 2007)

Paraguay
Cerro Cora National Park (5,500 ha) (probably extinct, Stallings 1985)
Ybicui National Park (5,000 ha) (Stallings 1985)
Tinfunque National Park (280,000 ha) (the most important protected area in Paraguay for this species; Stallings 1985)
Defensores del Chaco National Park (780,000 ha) (Stallings 1985)
Caaguazu National Park (6,000 ha) (Stallings 1985)
Kuri y National Reserve (2,000 ha) (Stallings 1985)
Yakui Protected Forest (1,000 ha) (Stallings 1985)
Nacunday Protected Forest (1,000 ha) (Stallings 1985).

It is listed on Appendix II of CITES.
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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Positive

Although there are reports of Alouatta caraya being imported to the U.S. for use as laboratory animals little else has been reported about their use. Several Black Howler monkeys can be found in zoos. A. caraya are also hunted for meat and fur.

(Muckenhirn 1976, Shoemaker 1979)

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Wikipedia

Black howler

The Black howler (Alouatta caraya) is a species of howler monkey, a large New World monkey, from northeast Argentina, east Bolivia, east and south Brazil and Paraguay. Together with the brown howler, it is the southernmost member of the Alouatta genus. Only the adult male is black; adult females and juveniles of both genders are overall whitish to yellowish-buff.[3] However, even among the adult males there are variations, and some have patches of reddish-brown or buff fur.[3]

They live in groups of three to 19 individuals (usually 7 to 9). There are usually one to three males for every seven to nine females in a group. When mating, males and females within a single group pair off.

Named for their vocalizations, they may be heard most often around sunrise. This "dawn chorus" sounds much more like roaring than howling, and it announces the howlers' position as a means to avoiding conflict with other groups. The call can be heard up to 5 km away.

These monkeys commonly sleep or rest up to 70 percent of the day,[2] making it one of the least active monkeys in the New World. Their habitat is forest, especially semi-deciduous and gallery.[2] Black Howlers are herbivorous, eating mostly leaves,[4] and occasionally fruit, such as figs. They generally prefer walking and climbing to running or leaping. The prehensile tail is very strong and acts as a fifth limb, allowing the monkeys greater versatility when climbing and allowing them greater safety in the occasional fall from a high branch. Because their limb structure makes terrestrial movement awkward, they spend most of their time in the trees and only come down for water during dry spells. Otherwise, the monkeys drink by wetting their hand on a moist leaf and then licking the water of their hand. Their lifespan is up to 20 years, but more commonly 15 years in the wild.

Female at the Apenheul Zoo

References

  1. ^ Groves, C. (2005). Wilson, D. E.; Reeder, D. M. eds. Mammal Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. pp. 148. OCLC 62265494. ISBN 0-801-88221-4. http://www.bucknell.edu/msw3/browse.asp?id=12100376. 
  2. ^ a b c Fernandez-Duque, E., Wallace, R. B. & Rylands, A. B. (2008). "Alouatta caraya". IUCN Red List of Threatened Species. Version 2011.2. International Union for Conservation of Nature. http://www.iucnredlist.org/apps/redlist/details/41545. Retrieved 19 January 2012. 
  3. ^ a b Gregorin, R. (2006). (Portuguese) Taxonomia e variação geográfica das espécies do gênero Alouatta Lacépède (Primates, Atelidae) no Brasil. Rev. Bras. Zool. 23(1).
  4. ^ National Geographic
  • Louise Emmons and Francois Feer. (1997). Neotropical Rainforest Mammals.

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