|Scientific Name:||Alouatta caraya|
|Species Authority:||(Humboldt, 1812)|
|Taxonomic Notes:||In their phylogenetic analysis using the cytochrome b gene, Nascimento et al. (2005) showed that populations of Alouatta caraya from Santa Cruz, Bolivia (Chaco) are differentiated from those in various localities in the state of Mato Grosso and (one specimen) Goiás further north. This indicates the possibility of two taxa of the Black Howler Monkey, rather than just one.|
|Red List Category & Criteria:||Least Concern ver 3.1|
|Assessor/s:||Fernandez-Duque, E., Wallace, R.B. & Rylands, A.B.|
|Reviewer/s:||Mittermeier, R.A. & Rylands, A.B. (Primate Red List Authority)|
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.
|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.
Native:Argentina (Chaco, Corrientes, Formosa, Misiones, Santa Fé); Bolivia, Plurinational States of; Brazil (Bahia, Brasília Distrito Federal, Goiás, Mato Grosso do Sul, Minas Gerais, Paraná, Rio Grande do Sul, São Paulo); Paraguay
|Range Map:||Click here to open the map viewer and explore range.|
|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).|
|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).
Adult male weight mean 6.42 kg (n=58), adult female weight mean 4.33 kg (n=117) (Rumiz 1990)
|Major Threat(s):||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.|
Alouatta caraya occurs in numerous protected areas:
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)
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)
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)
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.
Aguiar, L. M., Mellek, D. M., Abreu, K. C., Boscarato, T. G., Bernardi, I. P., Miranda, J. M. D. and Passos, F. C. 2007. Sympatry between Alouatta caraya and Alouatta clamitans and the rediscovery of free-ranging potential hybrids in Southern Brazil. Primates 48: 24-248.
Arditi, S. I. and Placci, L. G. 1990. Hábitat y densidad de Aotus azarae y Alouatta caraya en Riacho Pilagá, Formosa. Boletín Primatológico Latinoamericano 2: 29-47.
Bicca-Marques, J. C. 1994. Padrão de utilização de uma ilha de mata por Alouatta caraya (Primates: Cebidae). Revista Brasileira de Biologia 54(1): 167-171.
Bicca-Marques, J. C. 2003. How do howler monkeys cope with habitat fragmentation? In: L. K. Marsh (ed.), Primates in Fragments: Ecology and Conservation, pp. 283-303. Kluwer Academic/Plenum Publishers, New York, USA.
Bicca-Marques, J. C. and Calegaro-Marques, C. 1994. Feeding behavior of the black howler monkey (Alouatta caraya) in a seminatural forest. Acta Biologica Leopoldensia 16: 69-84.
Brown, A. D. and Zunino, G. E. 1994. Hábitat, densidad y problemas de conservación de los primates de Argentina. Vida Silvestre Neotropical 3: 30-40.
Codenotti, T. L. and da Silva, V. M. 2004. Resultados da enquete sobre ocorrência de primatas no Rio Grande do Sul, Brasil. Neotropical Primates 12(2): 83-89.
Codenotti, T. L., Silva, V. M. da, Albuquerque, V. J. de, Camargo, E. V. and Silveira, R. M. M. 2002. Distribuição e situação atual de conservação de Alouatta caraya (Humboldt, 1812) no Rio Grande do Sul, Brasil. Neotropical Primates 10(3): 132-141.
de Marques, A. A. B. 2003. Primatas. In: C. S. Fontana, G. S. Bencke and R. E. Reis (eds), Livro Vermelho da Fauna Ameaçada de Extinção no Rio Grande do Sul, pp. 499-506. EDIPUCRS, Porto Alegre, Brazil.
Drubbel, R. V. and Gautier, J.-P. 1993. On the occurrence of nocturnal and diurnal loud calls, differing in structure and duration, in red howlers (Alouatta seniculus) of French Guyana. Folia Primatologica 60: 195–209.
Dvoskin, R., Juárez, C. P. and Fernandez-Duque, E. 2003. Population density of black howlers (Alouatta caraya) in the gallery forests of the Argentinean Chaco: a preliminary assessment. Folia Primatologica 75: 93-96.
Nascimento, F. F., Bonvicino, C. R., da Silva, F. C., Schneider, M. P. and Seuánez, H. N. 2005. Cytochrome b polymorphisms and population structure of two species of Alouatta (Primates). Cytogenetic Genome Research 108: 106–111.
Neville, M. K., Glander, K. E., Braza, F. and Rylands, A. B. 1988. The howling monkeys, genus Alouatta. In: R. A. Mittermeier, A. B. Rylands, A. F. Coimbra-Filho an G. A. B. da Fonseca (ed.), The Ecology and Behavior of Neotropical Primates, Vol. 2, pp. 349–453. World Wildlife Fund, Washington, DC, USA.
Pope, B. L. 1968. Population characteristics. In: M. Malinow (ed.), Biology of the Howler Monkeys (Alouatta caraya), pp. 13-29. Karger, Basel.
Rumiz, D. I. 1990. Alouatta caraya: Population density and demography in northern Argentina. American Journal of Primatology 21: 279-294.
Santini, M. E. L. 1986. Padrões de atividade diaria de Alouatta caraya (Primates, Cebidae) reintrodizido no Parque Nacional de Brasília. In: M. T. de Mello (ed.), A Primatologia no Brasil - 2, pp. 293-304. Sociedade Brasileira de Primatologia, Brasília, Brazil.
Stallings, J. R. 1985. Distribution and status of primates in Paraguay. Primate Conservation 6: 51-57.
Stallings, J. R., West, L., Hahn, W. and Gamarra, I. 1989. Primates and their relation to habitat in the Paraguayan chaco. In: K. H. Redford and J. F. Eisenberg (eds), Advances in Neotropical Mammalogy, pp. 425-442. The Sandhill Crane Press, Inc., Gainesville, Florida, USA.
Thorington Jr., R. W., Ruiz, J. C. and Eisenberg, J. F. 1984. A study of a black howling monkey (Alouatta caraya) population in northern Argentina. American Journal of Primatology 6: 357-366.
Villalba, J. S., Prigioni, C. M. and Sappa, A. C. 1995. Sobre la posible preséncia de Alouatta caraya en Uruguay. Neotropical Primates 3(4): 173-174.
Wallace, R. B., Painter, R. L. E. and Taber, A. B. 1998. Primate diversity, habitat preferences and population density estimates in Noel Kempff Mercado National Park, Santa Cruz, Bolivia. American Journal of Primatology 46: 197–211.
Wallace, R. B., Painter, R. L. E., Rumiz, D. I. and Taber, A. B. 2000. Primate diversity, distribution and relative abundances in the Rios Blanco y Negro Wildlife Reserve, Santa Cruz Department, Bolivia. Neotropical Primates 8(1): 24–28.
Zunino, G. E. 1986. Algunos aspectos de la ecología y etología del mono aullador negro (Alouatta caraya) en habitat fragmentados. PhD Thesis, Universidad de Buenos Aires.
Zunino, G. E., Bravo, S., Ferreira, F. M. and Reisenmann, C. 1996. Characteristics of two types of habitat and the status of the howler monkey (Alouatta caraya) in northern Argentina. Neotropical primates 4: 48-50.
Zunino, G. E., González, V., Kowaleski, M. M. and Bravo, S. P. 2001. Alouatta caraya: relations among habitat density and social organization. Primate Report 61: 37-46.
|Citation:||Fernandez-Duque, E., Wallace, R.B. & Rylands, A.B. 2008. Alouatta caraya. In: IUCN 2013. IUCN Red List of Threatened Species. Version 2013.2. <www.iucnredlist.org>. Downloaded on 08 March 2014.|
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