Alouatta palliata (Gray, 1849)

Russell A. Mittermeier, Anthony B. Rylands & Don E. Wilson, 2013, Atelidae, Handbook of the Mammals of the World – Volume 3 Primates, Barcelona: Lynx Edicions, pp. 484-549 : 532-533

11 .

Mantled Howler

Alouatta palliata

French: Hurleur a manteau / German: Mantelbrtllaffe / Spanish: Mono aullador de manto dorado Other common names: Azuero Peninsula Howler (trabeata), Coiba Island Howler (coibensis), Ecuadorian Mantled/ South Pacific Blackish Howler (aequatorialis), Golden-mantled Howler (palliata), Mexican Howler (mexicana)

Taxonomy. Mycetes palliata Gray, 1849 ,

Caracas, Venezuela. Restricted by Allen in 1910 to Lago Nicaragua, Nicaragua.

Three subspecies of A. palliata are based on geography: mexicana, palliata , and aequatorialis. The forms coibensis (Coiba Island) and trabeata (Azuero Peninsula) were identified as distinct in a study of dermal ridge patterning by J. and P. Froehlich in 1986. Similar color morphs and body sizes can be found in all populations.

A study byJ. Ellsworth and G. Hoelzer in 2006, using microsatellite markers, found little genetic variability among this widely distributed species. In fact, they suggested that A. palliata may be the least genetically variable, well-sampled species known. Molecular genetic studies of mtDNA showed very low sequence diversity among subspecies and, most particularly, failed to demonstrate that coibensis was phylogenetically distinct: the difference between coibensis and palliata was less than was found among different populations of palliata . C. P. Groves in 2001 considered coibensis to be a distinct species, with trabeata as a junior synonym. The subspecies mexicana hybridizes with A. pigra in an area of sympatry in Tabasco, Mexico (c.8 km south-east of Macuspana and in other small forest fragments), and the subspecies aequatorialis is sympatric with A. seniculus in parts of north-western Colombia. Genetic analyses have indicated that crosses of the parental species result in fertile female offspring but infertile male offspring. Five subspecies recognized.

Subspecies and Distribution. A. p. palliata Gray, 1849 — NE Guatemala (Rio Motagua, and possibly along the coast a short distance to Cabo de Tres Puntas, where it meets the distribution of the Central American Black Howler, A. pigra ), E to E Costa Rica or W Panama;it is not found in El Salvador, although it may have occurred there in the past. A. p. aequatorialis Festa, 1903 — from the S distributional limits of the nominate subspecies palliata (either in E Costa Rica or W Panama) through the Serrania del Darién into W Colombia (N through basins of the rios Sint and Atrato to the Caribbean coast, and S through the Serrania del Baudo and the foothills, lowlands, and lower montane areas W of the Andes to the Pacific coast), W Ecuador, and NW Peru (extreme N of Tumbes Department). A. p. coibensis Thomas, 1902 — SW Panama (Coiba and Jicaron Is). A. p. mexicana Merriam, 1902 — S & SE Mexico (states of Veracruz, Tabasco, Campeche, Oaxaca, and N Chiapas) and Guatemala, in a swathe skirting the S of the distribution of the Central American Black Howler. A. p. trabeata Lawrence, 1933 — SW Panama (Azuero Peninsula).

Descriptive notes. Head-body 47-63 cm (males) and 46-60 cm (females), tail 60- 70 cm (males) and 55-66 cm (females); weight 4-5-9 kg (males) and 3-1-7-6 kg (females). The coat of the Mantled Howler is smooth, very short and upright, being silky black with a mantle of longer, gold or yellowish-brown fur along flanks. Backwarddirected forehead hairs meet on the crown in a slight, straight transverse crest. Adult males have a white scrotum. In the “Golden-mantled Howler” (A. p. palliata ), hair bases are walnut-colored, and the light fur along the sides is restricted to flanks. The light flank or mantle extends to the dorsum, but it is extremely variable both in color and extent, even within populations. The “Mexican Howler” (A. p. mexicana) is broadly similar in appearance to the nominate subspecies and is distinguished mainly on cranial characters. Light banded hairs are more widely distributed over the back. General color of the “South Pacific Blackish Howler” (A. p. aequatorialis) is black to dark brown, with mantle hairs shorter than in the nominate subspecies. The “Coiba Island Howler’ (A. p. coibensis) has certain consistent differences in skull structure and dermal ridges of hands and feet. It also has more strongly expressed sexual dimorphism. It is somewhat smaller than the mainland form. Fur is a duller, dark seal-brown, with the lower back paler and the veil more restricted to flanks than in the “Azuero Peninsula Howler” (A. c. trabeata). Rump and hindlimbs are walnut-brown. The Azuero Peninsula Howler is walnut-brown above, with long golden flank hairs.

Habitat. Primary and secondary rainforest from sea level to 2500 m, and riparian and dry lowland forest, coastal mangroves, and coffee plantations. The Mantled Howler prefers the upper forest canopy. Individuals occasionally travel on the ground, where they are quite capable of outrunning a human. They are also reported to swim.

Food and Feeding. Diets of Mantled Howlers vary seasonally. An average diet consists of 44-2% young leaves, 19-4% mature leaves, 18-2% flowers, 12:5% fruit, and 5-7% petioles and pulvini (a joint-like thickening at the base of the leaf).

Breeding. Breeding of Mantled Howlers is not seasonal. Gestation is 180-194 days (average 186 days), with an ovarian cycle of 11-24 days (average 16-3). Interbirth intervals are 18-25 months (average 22-5). The sex-ratio at birth is not significantly different from 1:1, but group sex ratio is skewed toward females, with an ideal ratio of 1:male:4 females. Social rank affects infant mortality; the highest ranking female tends to lose herfirst infant, and the lowest ranking females’ infants experience 80-90% mortality. Females are sexually mature at 36-40 months and give birth to their first infant 42-46 months. Females live 28-30 years, and they continue to produce infants throughout their lives. Males tend to live 25-27 years.

Activity patterns. Daily activities of Mantled Howlers are extremely variable, depending on season and food availability. A typical activity budget would be 65-74% of the time spent resting, 15-24% feeding, 10-18% traveling, and 0-4% engaged in social activity.

Movements, Home range and Social organization. Male and female Mantled Howlers leave their natal groups. Males spend 2—4 years alone and females 8-12 months. Group size is 2-44 individuals, and home ranges are 5-45 ha. Group sizes of Coiba Island Howlers are smaller than the continental populations, generally 2-9 individuals. Daily movements are 207-1267 m, depending on distribution and abundance of food sources. Groups are unimale—multifemale or multimale-multifemale. All adult males are dominant to all females, and social rank is linear, with age and social rank being inversely correlated (e.g. the youngest male and female occupy the alpha rank with older individuals occupying each declining rank).

Status and Conservation. CITES Appendix I (subspecies palliata and coibensis) and CITES Appendix II (other subspecies). Classified as Least Concern on The IUCN Red List, but including only the nominate subspecies palliata . Subspecies trabeata and mexicana are classified as Critically Endangered and coibensis and aequatorialis are classified as Vulnerable on The IUCN Red List. The Mantled Howleris threatened by deforestation, poaching, and illegal trade. Surveys in 2001-2009 in Panama located Azuero Peninsula Howlers in three provinces (Herrera, Los Santos, and part of Veraguas). Forty-five groups were counted, totaling 452 individuals with a mean of 9-6 ind/group (range 3-26); the total population was estimated at ¢.3092 individuals. It occurs in Cerro Hoya National Park in Panama. The Coiba Island Howler is restricted to the Island of Coiba, whereit is protected in Coiba Island National Park (270,175 ha). In a study of South Pacific Blackish Howlers in Colombia, a density of 20 ind/km? was recorded near EI Amargal Biological Station. They occur in Ensenada de Utria National Natural Park, Colombia, but they are rare there. They are also found in Los Katios National Natural Park, Colombia, where they are sympatric with the Colombian Red Howler (A. seniculus ). They are also found in Corcovado National Park, Costa Rica, where 0-2 groups/km® and 0-1 groups/km?*were found in Golfo Dulce. Populations of South Pacific Blackish Howlers are extremely fragmented although they can be locally common. They occur in Machalilla National Park, Los Cedros Protected Forest, and Cotacachi-Cayapas, Mache-Chindul, and Manglares Churute ecological reserves in Ecuador. The Mexican Howler occurs in a number of protected areas in Guatemala and Mexico, but its distribution is restricted and populations are generally small and fragmented.

Bibliography. Allen (1910), Baldwin & Baldwin (1976a, 1976b), Baumgarten & Williamson (2007), Carpenter (1934), Carrillo et al. (2000), Clarke (1983, 1990), Clarke & Glander (1981, 1984, 2004), Clarke & Zucker (1989), Clarke, Collins & Zucker (2002), Clarke, Crockett et al. (2002), Clarke, Zucker & Glander (1994), Cortés-Ortiz, Bermingham et al. (2003), Cortés-Ortiz, Duda et al. (2007), Ellsworth & Hoelzer (2006), Estrada (1982), Estrada & Coates-Estrada (1984, 1988), Estrada et al. (2001), Fedigan et al. (1985), Froehlich & Froehlich (1986, 1987), Glander (1978, 1980, 1992, 2006), Glander & Nisbett (1996), Groves (2001), Jones (1980), Lawrence (1933), Méndez-Carvajal (2003, 2005, 2013), Milton (1979a, 1979b, 1981a, 1982, 1996, 1998), Milton & Mittermeier (1977), Mittermeier (1977), Ramirez & Sanchez (2003), Rockwood & Glander (1979), Rowe (2000), Rylands et al. (2006), Tirira (2007), Zucker & Clarke (2003), Zucker et al. (1996).

Figures

On followıng pages Mantled Howler (Alouatta palliata) 12 Central Amerıcan Black Howler (Alouatta pigra)

Subspecies and Distribution. A. p. palliata Gray, 1849 — NE Guatemala (Rio Motagua, and possibly along the coast a short distance to Cabo de Tres Puntas, where it meets the distribution of the Central American Black Howler, A. pigra ), E to E Costa Rica or W Panama ;it is not found in El Salvador , although it may have occurred there in the past. A. p. aequatorialis Festa, 1903 — from the S distributional limits of the nominate subspecies palliata (either in E Costa Rica or W Panama ) through the Serrania del Darién into W Colombia (N through basins of the rios Sint and Atrato to the Caribbean coast, and S through the Serrania del Baudo and the foothills, lowlands, and lower montane areas W of the Andes to the Pacific coast), W Ecuador , and NW Peru (extreme N of Tumbes Department). A. p. coibensis Thomas, 1902 — SW Panama (Coiba and Jicaron Is). A. p. mexicana Merriam, 1902 — S & SE Mexico (states of Veracruz, Tabasco, Campeche, Oaxaca, and N Chiapas) and Guatemala , in a swathe skirting the S of the distribution of the Central American Black Howler. A. p. trabeata Lawrence, 1933 — SW Panama (Azuero Peninsula).