|Scientific Name:||Alouatta pigra|
|Species Authority:||Lawrence, 1933|
|Taxonomic Notes:||The taxonomy of the howlers of Mesoamerica and the Caribbean and Pacific coasts of Colombia and Ecuador is based on Lawrence (1933), Hill (1962), Hall (1981), Froehlich and Froehlich (1986, 1987), and Cortés-Ortiz et al. (2003). Groves (2001, 2005) recognized only A. palliata (Gray, 1849) (no subspecies), A. pigra Lawrence, 1933), and A. coibensis Thomas, 1902 (no subspecies). Rylands et al. (2006) reviewed the taxonomy and distributions of Alouatta palliata, A. coibensis and A. pigra.
The taxonomic status and distribution of Alouatta pigra follows Smith (1970), Horwich (1983), Horwich and Johnson (1984). Alouatta palliata luctuosa Lawrence, 1933 listed by Hill (1962) for Belize, was not recognized by Froehlich and Froehlich (1986). Alouatta pigra luctuosa was listed by Dahl (1987) for the primates of Belize. Cortés-Ortiz et al. (2003) provided molecular genetic data indicating that Alouatta pigra should be recognized as a species distinct from A. palliata.
The revision of the Mesoamerican howler monkeys by Lawrence (1933) resulted in the black howler of the Yucatán Peninsula, Mexico, Belize, and northern Guatemala being referred to as Alouatta pigra Lawrence, 1933. Lawrence (1933) recognized the existence of a previous name, Mycetes villosus Gray, 1845, but considered that it was impossible to use because the holotype (in the Natural History Museum, London) was restricted to a damaged skull of an immature female (the skin had been lost), the type locality was imprecise and confused, and she was unable to determine to which of two forms that she named (A. palliata pigra of Mexico and Guatemala, and A. palliata luctuosa of Belize) it may have belonged. Smith (1970) argued that luctuosa was indistinguishable from pigra, but that pigra should be considered a species distinct from palliata. Napier (1976) referred to A. pigra as A. villosa and, revisiting the issue, Brandon-Jones (2006) argued that Alouatta pigra Lawrence, 1933 is a junior synonym.
|Red List Category & Criteria:||Endangered A4cd ver 3.1|
|Assessor(s):||Marsh, L.K., Cuarón, A.D., Cortés-Ortiz, L., Shedden, A., Rodríguez-Luna, E. & de Grammont, P.C,|
|Reviewer(s):||Mittermeier, R.A. & Rylands, A.B. (Primate Red List Authority)|
This species is listed as Endangered as it is estimated that this species will experience a decline approaching 60% over a period of three generations (30 years) based on past and current rates of habitat decline and continued hunting for food and capture for pets.
Alouatta pigra occurs in the Yucatán Peninsula in Mexico and Belize, extending into northern and central Guatemela. The westernmost locality given by Hall (1981) is at Frontera, in the Mexican state of Tabasco; A. palliata has been recorded just west of there along the coast, 6 miles south of Cárdenas. Further localities which define the western and southern limits to its range include 5 miles south-east of Macuspana, Tabasco, and San Mateo Ixtatán (c. 11,000 ft) in Guatemala. Smith (1970) identified the southern limits to its range in the east with three localities along the Río Motagua basin in Guatemala, including Quirigua and Zacapa (right bank of the river). Curdts (1993), on the other hand, found that the southern and south-western limits to the range of A. pigra in Guatemala were defined by the Lago de Izabal, El Golfete and the Río Dulce. He noted large numbers of A. pigra in the Río Polochic delta, entering the west end of the Lago de Izabal. These are just to the north of the Río Motagua, where Curdts (1993) identified A. palliata. Baumgarten and Williamson (2007) found A. pigra in the northern margin of Rio Dulce and Lago Izabal, and to the West of Lago Izabal. This marks the southernmost locality for A. pigra in the Sierra de las Minas.
Baumgarten and Williamson (2007) provide the most recent review of the limits of the distributions of Alouatta palliata and A. pigra in Central America and Mexico. They found that the highland massif of northern Central America (including the Sierra Madre de Chiapas and central highland of Guatemala) and its associated coniferous and subalpine vegetation forms a geographic barrier separating A. pigra from A. palliata, and defines the southern limit of A. pigra. They discussed the two contact zones between them: the broad range of overlap north of the highland massif in Mexico over the lowlands of the states of Tabasco and Campeche, and a narrow area of contiguous, non-overlapping ranges in eastern Guatemala where the highlands extend almost the Caribbean. In the first, the localities where parapatry has been observed include Macuspana, Tabasco (Smith 1970; Horwich and Johnson 1986), around Zapata, Tabasco (Horwich and Johnson 1986) and the northern point of the Laguna de Términos in Campeche (Serio-Silva et al. 2006). There is no geographical barrier separating the species, both occur in the same forests and on both sides of the Rio Usamacinta, for example (Cortés-Ortiz et al. 2003). In the second, the ranges are narrowly parapatric, separated by the Río Dulce and the Lago Izabal: A. palliata to the south and A. pigra to the north and west. South-west of Lake Izabal, A. pigra occurs in the highlands of the Sierra de las Minas, but not in the lowlands, occupied by A. palliata. Any past range overlap in this region will have been lost by the extensive oss of forest; the area is heavily farmed. Baumgarten (2006) found no evidence to support sympatry supposed previously by Horwich and Johnson (1986), Curdts (1993) and Silva-López et al. (1998).
Serio-Silva et al. (2006) reported on a detailed survey of the occurrence of A. pigra in the Yucatán Peninsula.
Native:Belize; Guatemala; Mexico (Campeche, Chiapas, Quintana Roo, Tabasco, Yucatán)
|Range Map:||Click here to open the map viewer and explore range.|
Alouatta pigra is locally common across its range, but mainly in protected areas. In Belize, this species has been recorded at densities of up to 250 individuals/km² in some areas.
Estrada et al. (2004, 2006) carried out surveys of A. pigra populations in a number of areas in Guatemala and Mexico: Calakmul Biosphere Reserve, Campeche, 15.2 individuals/km²; Yaxchilán Natural Monument, Chiapas, 12.8 individuals/km²; and Tikal National Park, Guatemala, 17.8 individuals/km²; Palenque, Chiapas 23 individuals/km². At 13 sites where Estrada et al. (2006) estimated densities of A. pigra, the mean density was 10.8±5.7 individuals/km² (range 3.5-23 individuals/km²).
|Habitat and Ecology:||
This species occurs in primary terra firma rain forest, riparian forest, seasonally flooded riparian areas, and swamps. However, it can be also found in primary and secondary forest. In areas where suitable habitat does not exist they in some cases be found in marginal habitat surrounded by pastures and urban areas (Cuarón 1991, 1997; Marsh, 1999, 2001, 2002). In Calakmul, Mexico and Tikal, Guatemala, they occur in semideciduous tropical forest, but in Yaxchilán in tall evergreen rain forest (Estrada et al. 2004). This species is mainly found in areas below 900 m, but more commonly below 400 m. However, Baumgarten and Williamson (2007) found them in locations up to 3,350 m asl. Reid (1997) reported that it is found in evergreen and semideciduous forest, remnant forest along rivers and tall second growth.
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 jaws 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 which 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. Silver et al. (1998, 2000) have studied the diet and feeding ecology of A. pigra in Belize.
Alouatta pigra lives in smaller groups than A. palliata: 2-10 individuals per group compared with 2-45 for A. palliata (mean of 12.3) (Crockett and Eisenberg 1986; Chapman and Balcomb 1998; Pavelka and Chapman 2006). Estrada et al. (2004) recorded a mean group size of 7.5±1.9 individuals (range 4-9, n=8) at Calakmul, Campeche; 6.6±2.1 individuals (range 4-10, n=8) at Yaxchilán, Chiapas; and 8.7±2.2 individuals (range 6-12, n=10) at Tikal, Guatemala. Unlike the spider monkeys, and related to the large proportion of leaves in the diet (up to 50% of the annual diet), the howler monkeys generally have quite small and broadly overlapping home ranges, of 5 ha up to 45 ha, depending on the type of habitat (Neville et al. 1988).
Infant Alouatta are probably born throughout the year in Suriname, but data are not yet sufficient to determine if there is a birth peak. In Suriname, newborn infants have been seen in March, April, and November, and January (Mittermeier 1977). Crockett and Rudran (1987a,b) examined seasonal variation in births in red howlers from northern Venezuela, and found that they were less frequent during the early wet season (weaning would occur at the time of greatest food shortage). The llanos forests are more seasonal, however, than in the Guianas, and it is possible that this is not the case elsewhere. Oestrus lasts 2-4 days, with intervals between oestrous periods of about 17 days. Interbirth intervals are generally about 16.6 months, although they may be shortened by the death of an infant to about 10.5 months (Crockett and Sekulic 1984).
Adult male weight 11.1-11.6 kg (mean 11.35 kg, n=2), adult female weight 6.3-6.6 kg (mean 6.4 kg, n=4) (Ford and Davis 1992)
Adult male weight 11.4 kg (n=2) (Peres 1994)
Adult female weight 6.43 kg (n=4) (Peres 1994).
|Major Threat(s):||The main threats to this species are deforestation, hunting (for food and for capture as pets) and disease (yellow fever epidemics) (Reid 1997; Pavelka and Chapman 2006).|
This species occurs, or may occur, in several protected areas:
Guanacaste National Park (32,512 ha) (R. Horwich pers. comm. to Rodríguez-Luna et al. 1996)
Blue Hole National Park (300 ha) (in range)
Mountain Pine Ridge (51,500 ha) (Dahl, 1987)
Cockscomb Basin Wildlife Sanctuary (40,000 ha) (Horwich et al. 1993; Horwich 1994; Koontz et al. 1994; Rodríguez-Luna et al. 1996)
Community Baboon Sanctuary (Horwich and Lyon 1990; Horwich 1994; Pavón 1994; Brockett et al. 1999, 2000; Horwich et al. 2000, 2001)
Upper Bladen (35,000 ha) (R. Horwich pers. comm. to Rodríguez-Luna et al. 1996)
Caracol Archeological Reserve (20,000 ha) (in range)
Monkey Bay Wildlife Sanctuary (Private Reserve) (443 ha) (in range)
Monkey Bay National Park (911 ha) (A. pigra reintroduced) (extinguished by yellow fever and hurricanes (1961 and 1978) (Clark and Brocket 1999)
Manatee Forest Reserve (in range)
Chiquibul National Park (107,607 ha) (Dahl 1987)
Bladen Branch Nature Reserve (39,256 ha) (Dahl 1987)
Rio Bravo Conservation and Management Area (61,513 ha) (Silva-López and Rumiz 1995)
Rio Dulce National Park (9,610 ha) (Silva-López et al. 1995; Silva-López 1998)
El Rosario National Park (1,105 ha) (in range)
Bahia Santo Tomas (1,000 ha) (in range)
Santa Rosalia (1,000 ha) (in range)
Cerro Miramundo (902 ha) (in range)
Las Victorias (82 ha) (in range)
El Reformador (60 ha) (in range)
Grutas Lanquin (in range)
Cuevas de Silvino (8 ha) (in range)
Laguna Lachua National Park (10,000 ha) (in range)
Biotopo Mario Dary Rivera Quetzal (1,153 ha) (A. pigra according to Curdts (1993) and Silva-López 1998; but A. palliata according to Silva-López et al. 1995; Matamoros and Seal 2001)
Biotopo de Chocon Machacas (7,000 ha) (Sympatric with A. palliata; Curdts 1993; Silva-López et al. 1995; Silva-López 1998; Matamoros and Seal 2001)
Biotopo San Miguel La Palotada (49,300 ha) (Sympatric with A. palliata; Curdts 1993; Silva-López et al. 1995; Matamoros and Seal 2001; possible occurrence Silva-López 1998)
Rio Escondido – Biotopo Laguna del Tigre (45,900 ha) (in range)
Dos Lagunas (45,950 ha) (in range)
Tikal National Park (57,600 ha) (Coelho et al. 1976; Schlichte 1978; Estrada et al. 2004)
Aguateca Cultural Monument (1,700 ha) (Silva-López et al. 1995; Silva-López 1998)
Ceibal Cultural Monument (1,700 ha) (in range)
Dos Pilas Cultural Monument (3,100 ha) (Silva-López et al. 1995; Silva-López 1998)
Rio Azul (28,900 ha) (in range)
Uaxactún (3,000 ha) (in range)
Ucanal (2,200 ha) (in range)
Xutilha (2,689 ha) (in range)
Ixcun Cultural Monument (400 ha)
El Peru (8,000 ha)
Biotopo San Miguel – El Zotz (42,000 ha) (in range)
Nakum (8,000 ha) (in range)
G48 Naranjo (1,200 ha)
Machaquilla Cultural Monument (2,500 ha) (in range)
Yaxha (9,000 ha) (in range)
Sierra de las Minas Biosphere Reserve (236,300 ha) (Sympatric with A. palliata; Curdts 1993; Silva-López et al. 1995; Silva-López 1998; Matamoros and Seal 2001)
Polochic (Sympatric with A. palliata; Curdts 1993; Silva-López et al. 1995; Matamoros and Seal 2001)
Sierra Lacandon National Park (200,000 ha) (in range)
Mirador / Dos Lagunas –Rio Azul National Park (147,000 ha) (in range)
El Tigre National Park (350,000 ha) (in range)
Trifinito National Park (4,000 ha)
Palenque National Park (1,771 ha) (Estrada and Coates-Estrada 1984; Mexico, SEDUE, 1989; Estrada et al. 2000; Matamoros and Seal 2001)
Tulum National Park 664 ha) (in range)
Ria Lagartos Special Biosphere Reserve (47,840 ha) (in range)
Pantanos de Centla (290,397 ha) (Matamoros and Seal 2001)
Calakmul Biosphere Reserve (Mexico, SEDUE, 1989; Estrada et al. 2004; Serio-Silva et al. 2006)
Montes Azules Biosphere Reserve (Selva Lacandona) (331,200 ha) (Mexico SEDUE, 1989; Matamoros and Seal 2001)
Sian Ka’an Biosphere Reserve (528,147 ha) (Serio-Silva et al. 2006; Matamoros and Seal 2001)
Chan-kin Protection Area for Wild Flora and Fauna (12,184 ha) (Matamoros and Seal 2001)
Yaxchilán Natural Monument (2,621 ha) (Estrada and Coates-Estrada 1984; Estrada et al. 2002, 2004; Matamoros and Seal 2001)
Bonampak Natural Monument (4,357 ha) (Estrada and Coates-Estrada 1984; Matamoros and Seal 2001)
Metzaboc (Hernández-Yãnez pers. comm., in Rodríguez-Luna et al. 1996a)
It is listed on Appendix I of CITES.
|Citation:||Marsh, L.K., Cuarón, A.D., Cortés-Ortiz, L., Shedden, A., Rodríguez-Luna, E. & de Grammont, P.C, 2008. Alouatta pigra. The IUCN Red List of Threatened Species. Version 2014.3. <www.iucnredlist.org>. Downloaded on 28 May 2015.|
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