Alouatta belzebul ( Linnaeus, 1766 )

3.3.3.1 Alouatta belzebul ( Linnaeus, 1766) View in CoL

Type: Inexistent or unknown.

Type locality: Brazil. Thomas (1911) restricted the type locality to the state of Pernambuco, based on the materials collected by Marcgrave ( 1648), on which Linnaeus based his description ( Gregorin 2006).

Common name: Red-and-black howler, red-handed howler.

Alouatta belzebul has been recognized as a full species ever since Linnaeus (1766) first described it in his genus Simia . Hill (1962) recognized five subspecies: A. b. belzebul , A. b. discolor , A. b. mexianae, A. b. ululata , and A. b. nigerrima . Morphological, cytogenetic, and molecular studies have provided evidence that A. nigerrima is more closely related to A. seniculus than to A. belzebul , and here we consider it tentatively as a full species (see below). Groves (2001, 2005) considered mexianae, discolor , and ululata as synonyms of A. belzebul ; however, here we cautiously follow Gregorin (2006) who placed mexianae as a junior synonym of A. discolor and recognized belzebul , discolor , and ululata as full species based on morphological analyses of the cranium and hyoid apparatus, and pelage color pattern. Below we comment on the need to validate some of these species with genetic data.

Alouatta belzebul has a disjunct distribution with populations in the lower Amazon region in Brazil, to the south of the Rio Amazonas, as well as in isolated populations in the coastal forests of northeastern Brazil ( Langguth et al. 1987; Coimbra-Filho et al. 1995) ( Fig. 3.1 View Fig ). Genetic analyses have shown little divergence in haplotypes of populations of A. belzebul in north-east Brazil and the Amazon basin ( Bonvicino et al. 2001; Cortés-Ortiz et al. 2003), and a phylogeographic study based on the mitochondrial cytochrome b gene fails to distinguish populations from Paraíba from those of the left margin of the Rio Tocantins in the state of Pará ( Nascimento et al. 2008).