taxonID	type	description	language	source
313A88142A13F337FEF2FDEC66E5FA1D.taxon	vernacular_names	(HOWLERS, SPIDER AND WOOLLY MONKEYS AND MURIQUIS)	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A13F337FEF2FDEC66E5FA1D.taxon	diagnosis	• Medium-sized to large monkeys with long limbs or shorter limbs and more robust, howlers with enlarged voice-box and head set low on shoulders (hunched), dense and woolly or long and coarse fur, and long, strong, prehensile tail. • 80 - 160 cm. • Neotropical Region • Humid, lowland, submontane and montane, deciduous, and semi-deciduous forests.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A13F337FEF2FDEC66E5FA1D.taxon	description	• 5 genera, 25 species, 42 taxa. • 4 species Critically Endangered, 8 species Endangered, 5 species Vulnerable; none Extinct since 1600.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A11F335FF87FE706B44F344.taxon	vernacular_names	Colombian Red Howler	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A11F335FF87FE706B44F344.taxon	materials_examined	Cartagena, department of Bolivar, Colombia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A11F335FF87FE706B44F344.taxon	discussion	In 1962, W. C. O. Hill listed nine subspecies of A. seniculus: seniculus, arctoidea, stramineus, macconnell, insulanus, amazonica, Juara, puruensis, and sara. The type specimen of A. s. stramineus is a female Paraguayan Howler (Alouatta caraya) and hence not a valid name for a red howler. Populations considered to be the domain of stramineus were attributed to A. macconnelli by C. P. Groves in 2001; he concluded that macconnelli and sara were valid species but that arctoidea and juara were subspecies of A. seniculus, amazonica and puruensis were synonyms of juara, and insulanus (from Trinidad Island) was a synonym of A. macconnelli. In his study of the morphology of the Brazilian howlers (cranium, hyoid, and pelage color patterns), R. Gregorin argued that juara and puruensis should be classified as distinct species. Distributions of these two forms are poorly understood, and here (contrary to Groves regarding puruensis) they are recognized as subspecies of A. seniculus pending further studies. A genetic study by R. Stanyon and coworkers in 1995 concluded that arctoidea was a full species, and it is considered so here. Three subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A11F335FF87FE706B44F344.taxon	distribution	Subspecies and Distribution. A. s. seniculus Linnaeus, 1766 — Colombia wherever there is forest (except along the Pacific coast and in the desert of the Guajira Peninsula in the far N), NW Venezuela (around Lake Maracaibo), Amazonian Brazil (N of the Solimoes and S of the Rio Negro), E Ecuador and E Peru (E of the Andes, E of the Rio Huallaga, to the upper rios Maranon, Napo, and Putumayo). A. s. juara Elliot, 1910 = W Amazonian Brazil S of the Solimoes, extending W into Amazonian Peru, in the Jurua Basin and, considering the form amazonica to be a synonym, extending across the Rio Solimoes to the interfluvium of the rios Japura and Negro; the W limit defining its distribution is not known. A. s. puruensis Lonnberg, 1941 — Brazilian Amazon, from the Rio Jurua Basin E to the lower Rio Madeira and the middle Rio Aripuana, it extends E across the upper Aripuana to the Rio Teles Pires, to the S is restricted to the N of the Rio Abuna, a left bank tributary of the Rio Madeira that extends along the N border of Bolivia; its occurrence is uncertain in the N of Rondonia State, S of the Rio Ji-parana, although it occurs in S Rondonia, to the Rio Mamoré-Guaporé, in the Serra da Pacaas Novos, and E through the Serra dos Parecis to the Rio Teles Pires.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A11F335FF87FE706B44F344.taxon	description	Descriptive notes. Head-body 51 - 63 cm (males) and 48 - 57 cm (females), tail 57 - 80 cm (males) and 52 - 69 cm (females); weight 5 - 4 - 9 kg (males) and 4 - 1 - 7 kg (females). The nominate subspecies of the Colombian Red Howler (seniculus) is strongly sexually dimorphic. Overall, it is golden-toned to coppery-red on the body, contrasting with a maroon head, shoulders, limbs, and proximal part of the tail. Crown hairs run completely forward, meeting backward-directed forehead hairs in a forwardly concave “ V. ” In general, males are much bigger than females and often have a blackish beard, limbs, and tail. Coat color ranges from old gold to deep golden-red, with the head (including the beard), shoulders, limbs, and proximal part of the tail deep maroon — the latter becoming paler along its length and with the tip nearly as light as the body. The face is black. Some individuals from Venezuela are almost entirely purplish-red. The “ Jurua Red Howler ” (A. s. juara) is not sexually dichromatic. It is a dark reddish-brown, with a small area in the middle of the back a little lighter in color (orangey-rufous). Limbs and base of the tail are dark rufous to black. Thetail is paler, more golden, from the middle to the tip. The “ Purus Red Howler ” (A. s. puruensis) is sexually dichromatic. Males are dark rufous (red brown) with a golden color on the upper dorsum (around the shoulder blades), whereas females are golden-orange, with distal portions of their limbs, base of theirtails, and their beards dark rufous.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A11F335FF87FE706B44F344.taxon	biology_ecology	Habitat. All types of forested environments including mangroves along the Caribbean coast, humid, semi-deciduous, and deciduous forest, and cloud forest (including oak forest in Colombia). Colombian Red Howlers occur in gallery and riparian forests in the Eastern Llanos of Colombia. They are found from sea level to cloud forest at elevations up to 3200 m in the Central Andes, Colombia. Although occurring in all forest types in the Amazon, Colombian Red Howlers are sometimes very rare or absent in terra firma forests. They prefer productive alluvial plains and flooded forest, particularly those flooded annually by silt-rich white water (varzea). Their preferences for flooded forests and mineral licks were given as the reason for their comparative rarity at numeroussites surveyed in Rondonia. Colombian Red Howlers were absent or very scarce in terra firma forests where species of Lagothrix and Ateles were still abundant in the interface between Amazon forest and savanna vegetation in southern Rondonia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A11F335FF87FE706B44F344.taxon	food_feeding	Food and Feeding. Colombian Red Howlers in Venezuela and Colombia eat fruit (70 % of ingested plant material by dry weight), with young leaves, flowers, and mature leaves making up the rest. A study by H. L. Queiroz in the varzea of Lago Mamiraua, Mamiraua Sustainable Development Reserve in Brazil, estimated a diet of 47 - 3 % fruits (ingested with insects), 25 - 6 % mature leaves, 19 - 9 % young leaves, and 7 - 2 % miscellaneous items, including flowers. Fig trees, especially Ficus trigona (Moraceae), often dominate the diet, supplying young and mature leaves and fruits. Weevil larvae in fig fruits (average 2 - 3 larvae / fruit) also provide protein. Besides Ficus, the four most important plants in annual diets of Colombian Red Howlers at Lago Mamiraua were Coussapoa (Urticaceae) providing young and mature leaves and fig-like fruit, Piranhea trifoliata (Euphorbiacae) providing leaves, Neoxythece elegans (Sapotacae) providing fruit, and Pithecellobium corymbosum (Fabaceae) providing young leaves. At Tinigua National Natural Park, Colombia, most of the fruit in the diets of Colombian Red Howlers came from species of Brosimum, Ficus, Pseudolmedia, and Castilla (all Moraceae). In cloud forest in Huila in the Colombian Andes, the annual diet of a group of Colombian Red Howlers was 44 - 5 % young leaves, 42 - 4 % fruits, 7 - 5 % mature leaves, 5 - 4 % flowers, and 0 - 2 % leaf petioles. Ficus (nine species), Cecropia (Urticaceae), Morus (Moraceae), and Quercus (Fagaceae) were the principal genera in the diet.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A11F335FF87FE706B44F344.taxon	breeding	Breeding. Female Colombian Red Howlers reach sexual maturity at 43 - 54 months, and they give birth to their first infant at ¢. 60 months after a gestation of 186 - 194 days. Births are usually single and are not seasonal. At Pacaya-Samiria National Reserve in Peru, births occur throughout the year, with the majority in the wet season (October-April) and only a few in the dry season (May-September). Receptive periods for breeding last 2 — 4 days, with intervals of c. 17 days. Interbirth intervals are 16 - 28 months, although they may be shortened by the death of an infant to ¢. 10 - 5 months. Only females carry infants; they cling to the mothers’ ventrum until they are about one month old and are then carried on the back. By four months, they venture off their mothers to explore, and by six months old, they are carried only at moments of alarm.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A11F335FF87FE706B44F344.taxon	activity	Activity patterns. Colombian Red Howlers spend 67 - 78 % of their day resting, 10 - 12: 7 % feeding, and 4 - 4 - 5 - 6 % traveling. At Lago Mamiraua, they spend 76 - 3 % oftheir day resting, 16 % traveling, 6 % feeding, and 1 - 7 % engaging in social or other activities. A similar pattern was noted for a group in cloud forest (elevation 2300 m): 78 - 5 % resting, 12 - 7 % feeding, 5 -: 6 % traveling, and 3 - 2 % engaging in social and other activities. Colombian Red Howlers generally occupy the middle and upper forest canopy, but occasionally they go to the ground to feed on soil or cross between forest patches. They swim across rivers and sleep in large emergenttrees.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A11F335FF87FE706B44F344.taxon	biology_ecology	Movements, Home range and Social organization. Groups of Colombian Red Howlers are multimale — multifemale or unimale — multifemale, with 6 - 13 individuals. A population studied over many years by P. Soini in Pacaya-Samiria National Reserve had mixed-sex groups, all-male groups, and solitary individuals. Mixed-sex groups had 2 - 11 individuals (exceptionally 13) and averaged 5 - 5 individuals, with 1 - 2 adult males and 1 - 3 adult females. Their home ranges were 6 - 9 ha. Sizes of nine groups in semi-deciduous mesic forest patches and gallery forest in the Eastern Llanos of Colombia were 3 - 9 individuals, with 1 - 3 adult males and home ranges of 10 - 26 ha. At Caparu on the Rio Apaporis in Colombia, a group of seven individuals, with one adult male and two adult females, used a very large home range of 182 ha, with daily movements of 370 - 2200 m (average 1150 m / day). Both sexes leave their natal groups. Violent fights can accompany the entry of a new dominant male into a group. A harpy eagle (Harpia harpyja) took an adult 7 - 3 - kg male in Manu National Park and Biosphere Reserve in Peru. Queiroz and J. M. Ayres estimated densities of Jurua Red Howlers in varzea around Lago Mamiraua at the mouth of the Rio Japura at 38 - 39 - 5 ind / km?. A density of 36 ind / km? was recorded in riparian and flooded forest in Pacaya-Samiria National Reserve. C. Peres compared primate communities in terra firma forest and varzea along the Rio Jurua, Brazil. At five varzea sites, densities of Jurua Red Howlers were 24 - 7, 35 - 7, 89 - 7, 15 - 6, and 35 - 7 ind / km?. In terra firma forest, howlers were sometimes not seen during censuses even though they occurred there. Otherwise, densities at nine locations were 1 - 9 - 2 ind / km?. Densities of 1 - 1 ind / km? and 4 ind / km? were estimated at two non-hunted sites of nutrient-poor terra firma forest on the lower rios Caqueta and Apaporis in Colombia; howlers did not occur at a third site.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A11F335FF87FE706B44F344.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The IUCN Red List, including all three subspecies. The Colombian Red Howler is widespread and locally common, but itis intensively hunted for meat in some areas. It occurs in numerous and many large protected areas, including Mamiraua and Amana state sustainable development reserves and Jau, Pico da Neblina, and Serra do Divisor national parks in Brazil; Amacayacu, Tinigua, Chiribiquete, El Tuparro, Paramillo, and Sierra de la Macarena national natural parks and Nukak and Puinawai national natural reserves in Colombia; and Pacaya-Samiria National Reserve and Manu National Park and Biosphere Reserve in Peru.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F334FF5AFE9667F0FC78.taxon	vernacular_names	Ursine Red Howler	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F334FF5AFE9667F0FC78.taxon	materials_examined	Venezuela. Restricted by Cabrera in 1957 to the valley of Aragua, state of Aragua, near Caracas.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F334FF5AFE9667F0FC78.taxon	discussion	The diminutive form occurring on the island of Trinidad, and formerly on Tobago (insulanus) is considered a junior synonym. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F334FF5AFE9667F0FC78.taxon	distribution	Distribution. Endemic to Trinidad I and Venezuela E of Lake Maracaibo, along the coast from Falcon State to Miranda State, the northern Mts (Cordillera Oriental, Cordillera Central, and Coro System), and extending S through the Llanos to the Rio Orinoco; it may extend into Arauca State, Colombia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F334FF5AFE9667F0FC78.taxon	description	Descriptive notes. Head-body 45 - 65 cm, tail 55 - 68 cm; weight 6 - 8 kg (males) and 4 - 5 - 7 kg (females). The Ursine Red Howleris generally golden-toned to coppery-red on the body, contrasting with a maroon head, shoulders, limbs, and proximal part of the tail. There is no contrasting “ saddle. ” Crown hairs run completely forward, meeting backward-directed forehead hairs in a forwardly concave “ V. ” Sexes are similar, although males are generally much larger than females and often have a blackish beard, limbs, and tail. Red howlers on Trinidad are smaller: head — - body 43 - 5 - 48 - 5 cm (males) and 42 - 8 - 45 - 3 cm (females), tail 55 - 61 cm (males) and 53 - 60 cm (females); weight 4.7 - 6 - 1 kg (males) and 4 - 1 - 5 - 7 kg (females) from data provided by K. Glander. They are not sexually dimorphic, and their pelage is almost uniformly reddish brown.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F334FF5AFE9667F0FC78.taxon	biology_ecology	Habitat. Deciduous forest patches, open woodland, and gallery forest of the Llanos at elevations of 10 - 1160 m, and may exist in cloud forests along coastal Venezuela up to elevations of 2000 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F334FF5AFE9667F0FC78.taxon	food_feeding	Food and Feeding. Diets of Ursine Red Howlers may consist of up to 70 % leaves, with fruits, flowers, and other plant parts making up the remainder.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F334FF5AFE9667F0FC78.taxon	breeding	Breeding. Ovarian cycles of Ursine Red Howlers are c. 17 days. When a femaleis receptive, a dominant male stays close to her in consortship for several hours or up to several days. Consortships are briefer with unfamiliar males that have only recently entered a group. Females maintain their distances from unfamiliar males, only briefly approaching them and soliciting mating. Males and females show sexual solicitation, involving tongue flicking. In multimale groups, the dominant male has priority access to females, although sometimes females copulate with a subordinate male or even males from neighboring groups. Births are normally single, after a gestation of 186 - 194 days. The annual birth rate in a population at Hato Masaguaral in the Venezuelan Llanos was 0 - 68 births / female / year. Births are not seasonal. Interbirth intervals are 10 - 5 - 26 months (average 16 - 6). The age at sexual maturity is 58 - 66 months for males and 43 - 54 months for females. Females give birth for the first time at about five years old, and males rarely sire offspring before they are seven years old.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F334FF5AFE9667F0FC78.taxon	activity	Activity patterns. Ursine Red Howlers rest during 67 - 78 % of the day, feed during 10 - 12 -: 7 %, and travel during 4 - 4 - 5 - 6 %, the remainder in other activities.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F334FF5AFE9667F0FC78.taxon	biology_ecology	Movements, Home range and Social organization. Groups of Ursine Red Howlers are unimale — multifemale or multimale-multifemale. In semi-deciduous forest patches of the Llanos, group sizes are 3 - 13 individuals, with means of 6 - 3 - 7 - 6 individuals; in open woodland, they are 4 - 16 individuals, with means of 8 - 5 - 10 - 5 individuals; and in semideciduous gallery forest, they are 4 - 14 individuals, with means of 5 - 9 - 8 - 1 individuals. Groups contain 1 - 2 adult males and generally 2 - 3 adult females, with adult male to adult female sex ratios of 1: 1 - 5 - 2 - 3. Home ranges in gallery forest are relatively large, c. 25 ha, with daily movements of 290 - 655 m (average 542 m). In open woodland, home ranges are smaller, 4 - 7 ha, with daily movements of 20 - 840 m (average 340 - 445 m in different studies). Home range overlap can be as high as 64 %. Two groups in Trinidad had eight and nine individuals, with four and three adult males, respectively. Their home ranges were 6 - 7 ha. Studies of roaring contests between groups and between groups and solitary males indicate that besides defending resources (establishing dominance where they meet rather than defending territorial boundaries), they are the way males defend and maintain ownership of groups of females. Resident females also deter entry of females from other groups with roaring contests. Male and female Ursine Red Howlers disperse. Males disperse singly, with half-brothers, or even as father-son pairs. The majority (62 %) of males disperse before reaching sexual maturity, but ages of dispersal vary (2 - 3 - 19 years); they all leave behind living mothers, sisters, or daughters in the group. The company of a father or brother is evidently a factor in dispersal of immature males; only adult males over five years old disperse singly. When entering a group, which generally involves vicious fights, in a coalition of relatives, males have longer tenure than those that do not. Male invasions are sometimes followed by infanticide, with the new male killing infants less than six months old. Tenures of dominant breeding adult males average 5 - 1 - 6 - 7 years. Groups of Ursine Red Howlers generally contain a maximum of four females. Female immigration is infrequent, and they are rarely able to enter groups after dispersing because resident females prevent them. In cases when they do, the adult female composition has been reduced by the death of a resident female. Females typically breed in their natal group, or they form new groups by pairing up with a solitary male. Because the age when females first breed is about five years (equivalent to the average time of tenure of a breeding male), they rarely have the opportunity to mate with their father, so incest avoidance likely is not the cause of female emigration. Rather, it is probable that competition for resources among females is the cause — the number of breeding positions in a group being limited. Densities in semi-deciduous forest patches and semi-deciduous gallery forest are similar at 25 - 54 ind / km? *, but in open woodland, densities can be higher at 83 - 118 ind / km ”.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F334FF5AFE9667F0FC78.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. The Ursine Red Howler is widespread and locally common but intensively hunted for meat in some areas. Population size on Trinidad is not known, but it occurs in three wildlife sanctuaries: Trinity Hills, Bush Bush, and Central Range. Surveys in Trinidad in 1994 indicated 24 groups in all, with average group sizes of 3 - 5 - 4 - 6 individuals and densities of 11 - 83 ind / km? Further surveys in 1997 indicated that the population remained stable. Archeological evidence from Tobago has shown that red howlers used to occur there but have been extirpated.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F333FA57FBA860C9FDA1.taxon	vernacular_names	Guianan Red Howler	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F333FA57FBA860C9FDA1.taxon	materials_examined	coast of Demerara (Guyana).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F333FA57FBA860C9FDA1.taxon	discussion	There is much regional variation, with albinism and melanism also reported. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F333FA57FBA860C9FDA1.taxon	distribution	Distribution. The Guianas, N Brazil (E of the Rio Negro and N of the Rio Amazonas, also on Gurupa I in the Amazon estuary), and S Venezuela (Rio Cassiquiare to the Rio Orinoco).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F333FA57FBA860C9FDA1.taxon	description	Descriptive notes. Head-body 51 - 63 cm (males) and 48 - 57 cm (females), tail 57 - 80 cm (males) and 52 - 69 cm (females); weight 5 - 4 - 9 kg (males) and 4 - 1 - 7 kg (females). The Guianan Red Howler is uniformly dark rufous-brown, with a yellowish-brown back (this tone occasionally restricted to a golden “ saddle ”). Arms to elbows and legs to thighs are orange red. There is a dark dorsalstripe, and the tail becomes paler distally.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F333FA57FBA860C9FDA1.taxon	biology_ecology	Habitat. High rainforest, riverbank high forest, marsh forest, swamp forest, Pterocarpus — Tabebuia swamp, savanna forest, mountain savanna forest, and ridge forest. The Guianan Red Howler is also sometimes seen in liana forest, swamp scrub (Annona Chrysobalanus formation), and mangrove forest, although it appears to be quite rare in the latter two. Guianan Red Howlers occasionally enter plantations. They spend most of their time in the middle and upper forest canopy, although they will use all levels and may even go to the ground to cover short distances between trees. They occur from sea level to elevations of 1200 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F333FA57FBA860C9FDA1.taxon	food_feeding	Food and Feeding. The diet of the Guianan Red Howler was been studied in detail by C. Julliot at Nouragues Field Station, French Guiana. It feeds many on different kinds of fruits, flowers, and young leaves. It also eats mature leaves, immature fruits, moss, bark, and soil of termitaria. Studies by C. Julliot and D. Sabatier recorded the use of 195 plant species from 47 families. The majority of the diet was young leaves (54 %); mature fruits (21 - 5 %), typically with juicy pulp, yellow, red and orange drupes and berries; and flowers (12: 6 %). Fruit dominates the diet in February-May (wet season) and October (Goupia glabra, Goupiaceae). Flowers are eaten in July and December, including those of Micropholis cayennensis (Sapotaceae), Eperua falcata (Fabaceae), Odontadenia (Apocynaceae), and Maripa scandens (Convolvulaceae). Sapotaceae and Moraceae are important families in the diet of Guianan Red Howlers. They disperse seeds of more than 95 % of the fruits that they eat.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F333FA57FBA860C9FDA1.taxon	breeding	Breeding. Gestation is 186 - 194 days. Interbirth interval averages 16 - 6 months, although they may be shortened to ¢. 10 - 5 months by the death of an infant. Sexual maturity is reached at 43 - 54 months for females, who give birth to theirfirst infant at c. 60 months. Births are usually single and are not seasonal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F333FA57FBA860C9FDA1.taxon	activity	Activity patterns. The activity budget of a group of Guianan Red Howlersin terra firma in a 1000 - ha forestjust north of Manaus, Amazonas State, Brazil, was as follows: resting, 70 - 9 %, feeding 15 - 7 %, traveling 11 %, and social and other behaviors 2: 4 %.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F333FA57FBA860C9FDA1.taxon	biology_ecology	Movements, Home range and Social organization. Guianan Red Howlerslive in small groups that usually include one adult male, one or more adult females, and one or more juveniles and infants. Group size is 2 - 8 individuals. Home ranges in French Guiana can be as large as 45 ha, with daily movements of up to 1645 m. During the formation of Lake Guri after the construction of the hydroelectric dam on the Rio Caroni, Venezuela, researchers witnessed the demise of all but one (a juvenile) of a group of six Guianan Red Howlers. They were killed over seven months by a Jaguar (Panthera onca). Harpy eagles (Harpia harpyja), and no doubt other large raptors, prey on Guianan Red Howlers.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A10F333FA57FBA860C9FDA1.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. The Guianan Red Howler is widespread and locally common, but it is intensively hunted in some areas and deforestation due to cattle ranching occurs in the southern part of its distribution. It occurs in a number of very large protected areas, particularly in Suriname and Brazil.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF82F6256919FD65.taxon	vernacular_names	Amazon Black Howler	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF82F6256919FD65.taxon	materials_examined	Brazil. Restricted by Cabrera in 1957 to Patinga, Amazonas.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF82F6256919FD65.taxon	discussion	Formerly believed to be a subspecies of A. belzebul. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF82F6256919FD65.taxon	distribution	Distribution. NC Brazil (states of Amazonas and Para), S of the Rio Amazonas between the Rios Madeira and Tapajos; there are outlying populations at Oriximina and Obidos on the N bank the Rio Amazonas and at Lago Janauaca, just S of the Rio Solimoes between the rios Coari and Purus.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF82F6256919FD65.taxon	description	Descriptive notes. Head-body 58 - 63 cm (males) and 48 - 5 - 64 - 8 cm (females), tail 64 - 69 cm (males) and 56 - 5 - 65 cm (females); weight 4 - 9 - 8 kg. The Amazon Black Howler is a large, entirely black species, with minimal sexual dichromatism. Unlike the Red-handed Howler (A. belzebul), which has a backward-directed toupee that meets a forward-directed stream from the nape whorl, the Amazon Black Howler has a short transverse crest that runs across the middle to the back of the crown. Morphology of the hyoid apparatusis distinct from that of the Red-handed Howler.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF82F6256919FD65.taxon	biology_ecology	Habitat. Tall evergreen terra firmaforest, seasonally inundated forest and forest patches in savanna areas, and secondary forest in abandoned rubber plantations.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF82F6256919FD65.taxon	food_feeding	Food and Feeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF82F6256919FD65.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF82F6256919FD65.taxon	activity	Activity patterns. The Amazon Black Howler is no doubt diurnal and arboreal, butit has not been studied in the wild.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF82F6256919FD65.taxon	biology_ecology	Movements, Home range and Social organization. The Amazon Black Howlerlives in unimale — - multifemale groups.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF82F6256919FD65.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The IUCN Red Lust. Little information is available for the Amazon Black Howler. It occurs in Amazonia National Park to the east of the Rio Tapajos.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF87FDC1633CF6C2.taxon	vernacular_names	Bolivian Red Howler	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF87FDC1633CF6C2.taxon	materials_examined	Province of Sara, department of Santa Cruz, Bolivia, 450 m above sea level.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF87FDC1633CF6C2.taxon	discussion	Molecular genetic and cytogenetic analyses demonstrated that A. sara is a distinct taxon in the red howler group. It is a sister species to A. seniculus, believed to have diverged from a common ancestor c. 2 - 4 million years ago. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF87FDC1633CF6C2.taxon	distribution	Distribution. Bolivia, from the Pando Department S along the Andean Cordillera and E into C Bolivia including the entire Rio Beni Basin and E as far the Mamoré-Guaporé interfluvium.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF87FDC1633CF6C2.taxon	description	Descriptive notes. Head-body 54 - 71 - 2 cm, tail 51 - 5 - 60 cm; weight 6 - 9 kg. The Bolivian Red Howleris similar in size to the Colombian Red Howler (A. seniculus), but with a smaller hyoid. Fur is brick-red above, with limbs, head, and proximal part of the tail slightly darker (i. e. more rufous).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF87FDC1633CF6C2.taxon	biology_ecology	Habitat. Tropical forest, including riverine forest and seasonally flooded forests on the Chiquitano Shield, up to elevations of 1000 m. To the east of the Rio Beni, it is sympatric with the Paraguayan Howler (A. caraya) in some areas, but the Bolivian Red Howler seems to prefer humid forests and seasonally flooded forests along major rivers while the Paraguayan Howler prefers drier, semi-deciduous forest and gallery forest in areas of savanna and chaco. There is a single report of both species occupying adjacent trees in igapo (black-water seasonally inundated forest) at Lago Caiman along the Rio Iténez (= Guapore).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF87FDC1633CF6C2.taxon	food_feeding	Food and Feeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF87FDC1633CF6C2.taxon	breeding	Breeding. There is no specific information available for this species, but breeding and reproduction are likely to be similar to the Colombian Red Howler.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF87FDC1633CF6C2.taxon	activity	Activity patterns. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF87FDC1633CF6C2.taxon	biology_ecology	Movements, Home range and Social organization. Based on surveys, the Bolivian Red Howler occurs in unimale-multifemale or multimale-multifemale groups, with 1 - 7 individuals. No data are available on home ranges or daily movements.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F333FF87FDC1633CF6C2.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. Bolivian Red Howlers are hunted for meat in some regions. They occur in Amboro, Carrasco, Madidi, and Noel Kempff Mercado national parks, Beni Biological Station and Biosphere Reserve, and Pilon Lajas Biosphere Reserve and Communal Lands. They are rare in the Rios Blanco y Negro National Reserve; encounter rates during surveys at various sites ranged from zero to 0 - 4 groups / 10 km. It is possible that they are the red howlers occurring in Isiboro Sécure National Park.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F332FA9EFC9C6004F584.taxon	vernacular_names	Red-handed Howler	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F332FA9EFC9C6004F584.taxon	materials_examined	Brazil. Restricted by O. Thomas in 1911 to Pernambuco, Brazil, and subsequently restricted by A. Cabrera in 1957 to the Rio Capim, eastern Para.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F332FA9EFC9C6004F584.taxon	discussion	In 1962, W. C. O. Hill listed five subspecies of A. belzebul (belzebul, discolor, ululata, mexianae, and nigerrima), but C. P. Groves considered all but mnigerrima as junior synonyms. Following R. Gregorin in 2006, belzebul, discolor, nigerrima, and ululata are recognized here as distinct species. The identity of mexianae is uncertain; Gregorin believed it to be a synonym of A. discolor, whereas M. Fernandes in his 1994 review of howlers of the Marajo Archipelago concluded it was a synonym of A. belzebul. Although considered an essentially Amazonian species, there are relict populations in the Atlantic Forest of north-eastern Brazil, and molecular and cytogenetic genetic analyses have shown that A. belzebul formsa sister group with A. guariba; the two are believed to have split from a common ancestor c. 4 million years ago. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F332FA9EFC9C6004F584.taxon	distribution	Distribution. Lower Amazonian Brazil S of the Rio Amazonas in the states of Para (also on Mexiana, Caviana, and Marajo Is in the Amazon estuary), Maranhao, Tocantins, and Mato Grosso, crossing the rios Tocantins and Araguaia W as far as the Rio Xingu-Iriri, and isolated populations in NE Brazil in the coastal regions of the states of Rio Grande do Norte, Paraiba, Pernambuco, and Alagoas (it is probable that its distribution was once continuous through the states of Ceara and Piaui to the Amazonian population). Its southern distributional limits in Para coincide with the change from tall forest to open formations such as the cerrado (bush savanna), caatinga (xeric scrub), and the babassu palm (Orbygnia) forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F332FA9EFC9C6004F584.taxon	description	Descriptive notes. Head-body 58 - 65 cm (males) and 37 - 50 cm (females), tail 56 — 70 cm (males) and 45 - 57 cm (females); weight 6 - 5 - 8 kg (males) and 4 - 8 - 6 - 2 kg (females). The Red-handed Howler has a short-haired toupee on the brows that meets a forward-directed stream (from a whorl between the nape and withers) in a low transverse crest on front of the crown. This forward stream diverges to the sides to run laterally. It is primarily black, but hands, feet, tip of the tail, and sometimes forehead and back are reddish-brown to yellow. In some areas, all individuals of one or both sexes may be uniformly black or red. The scrotum is rusty-red.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F332FA9EFC9C6004F584.taxon	biology_ecology	Habitat. Primary, seasonal semi-deciduous terra firma, wet evergreen and flooded forest, and some disturbed forest. The Red-handed Howler prefers the upper levels of the forest canopy.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F332FA9EFC9C6004F584.taxon	food_feeding	Food and Feeding. A north-eastern population of Red-handed Howlers, studied in the Fazenda Pacatuba, Paraiba, by C. Bonvicino, was relatively frugivorous, eating fruits of 34 species, seeds of five species, flowers of five species, leaves ofsix species, and gum exuded from seed pods of Parkia pendula (Fabaceae). As is typical for howlers, fruits of the Moraceae family, especially Ficus and Brosimum, figured prominently in the diet, as did Myrtaceae (Eugenia, Campomanesia) and Fabacae (Inga). In the wet season when fruits are abundant, Red-handed Howlers spent 915 % if their of time eating fruit, 2 % young leaves, 2 % mature leaves, and 4 - 5 % other items, and in the dry season, 43 - 9 % fruit, 40 - 8 % flowers, 11 % young leaves, and 4 - 3 % mature leaves. Diets of Red-handed Howlers also were also studied in tall (canopy 40 m), terra firma forest in the Amazonian part its range, in the Caxiuana National Forest, just east of the lower Rio Xingu in Para. The diet of one group studied by M. Jardim from September (middle of the dry season) to March (middle of the wet season) indicated a diet c. 70 % fruit, 18 % young leaves, 11 % mature leaves, and 1 % flowers. A subsequent study of the annual diets of two groups showed a predominance of leaves (58: 5 % of the diet overall), undoubtedly because it covered the late wet season and early dry season when fruit is relatively scarce. Diets were 40 - 5 % new leaves, 36 - 5 % fruit, 13 - 6 % leaves of unknown age, 4 - 9 % flowers, and 4 - 5 % mature leaves. Leaf consumption increased when fruits were scarcer (May-August). For one of the groups, c ¢. 50 % of the fruit feeding records came from just three species: Ficus guianensis (Moraceae) 23 - 2 %, Ocotea cf. cernua (Lauraceae) 11: 6 %, and Virola sebifera (Myristicaceae) 9 - 8 %. Red-handed Howlers also eat seeds of Vouacapoua americana (Fabaceae) and soil from termitaria during the dry season.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F332FA9EFC9C6004F584.taxon	breeding	Breeding. Two births of Red-handed Howlers were recorded in the study in Paraiba: one in early November (early dry season) and the other in late August (late wet season). Two births were also recorded in an Amazonian population. Births occurred in the early afternoon. One of the females was seen to eat the placenta. One infant was carried ventrally for the first month and then on the mother’s back. The other infant was carried dorsally from the moment it was born. These two infants died in their second and fourth months, one possibly because of a botfly infestation.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F332FA9EFC9C6004F584.taxon	activity	Activity patterns. During the dry season in Paraiba, Red-handed Howlers rested for 57 - 7 % of the day, traveled for 19 - 9 %, fed for 119 %, called for 3 - 4 %, and groomed and played for 5 - 6 % (other activities for 1 - 5 %). During the wet season, they rested more (60 - 65 %), traveled (18 - 4 %) and fed less (8 - 2 %), vocalized more (9: 3 %), and engaged in social activities less (3: 6 %). In the dry season, the amount of time feeding was quite consistent through the day, but in the wet season, there were morning (09: 00 - 11: 00 h) and afternoon (13: 00 - 15: 00 h) peaks. In the dry season, Red-handed Howlers were active from ¢. 06: 00 h to ¢. 18: 00 h, but in the wet season, they tended to begin their day earlier (c. 05: 00 h) and retire earlier (c. 15: 00 h). Bouts of howling were bimodal, with a strong peak in the early morning (04: 00 - 07: 00 h) and a lesser peak in the late afternoon (16: 00 - 17: 00 h). Red-handed Howlers travel and feed in the middle and upper forest canopy and sleep on the broad branches of tall emergenttrees.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F332FA9EFC9C6004F584.taxon	biology_ecology	Movements, Home range and Social organization. Red-handed Howlers live in unimale — multifemale or multimale-multifemale groups. In north-eastern Brazil, four groups had 5 - 14 individuals; all groups had a single male, exceptfor the largest group of 10 - 14 individuals that had two males. All groups had 2 - 3 adult females. Home ranges of two groups, both with 6 - 8 individuals, were 4 - 7 ha and 9 - 5 ha. In Caxiuana National Forest, three groups of Red-handed Howlers had single males and 5 - 9 individuals; home ranges were 13 - 5 - 18 ha. They used a larger portion of their home ranges in the wet season than in the dry season, associated with a more diverse diet in terms of numbers of species and trees they visited for food. The number oftrees visited in the wet season was nearly twice that in the dry season. Daily movements are similar in the two seasons, with a mean of 1328 m (range 997 - 2296 m) in the dry season and 1621 m (range 861 - 2409 m) in the wet season. Tayras (Eira barbara) were observed accompanying a group of Red-handed Howlers over a few days, eventually attacking an infant. The adult male gave alarm calls continuously but made no attempt to defend the infant. The infant was wounded, but because the Tayras moved off when they saw the human observer, it was saved. Tayras were also seen attacking a subadult female while on the ground. A number of deaths recorded at the time of these attacks were likely caused by Tayras. On an island in the man-made lake of Tucurui in Para, sightings and deaths of Red-handed Howlers suggested that a Tayra family swam across a channel to reach the island.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A17F332FA9EFC9C6004F584.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. The Red-handed Howler is widespread and can be locally common in the eastern Amazonian states of Para and Maranhao. It is very rare in north-eastern Brazil, where the total population is estimated at ¢. 300 individuals in eleven isolated locations: seven in Paraiba, two in Rio Grande de Norte, one in Pernambuco, and one in Alagoas. The largest population of ¢. 80 individuals occurs in a small area of Atlantic Forest in Pacatuba, Paraiba, surrounded by sugar cane plantations. Red-handed Howlers were reintroduced in Guaribas Biological Reserve (4322 ha), Paraiba, with translocated individuals from Sapé and Santa Rita in Paraiba and the release of local pets; the population numbered twelve individuals in 2007. The eastern Amazon in southern Para and the north of Mato Grosso has suffered devastating deforestation over the last 20 years; the timber industry, mining, infrastructure development, and agroindustry have taken a heavy toll on the forests and their associated wildlife. The Red-handed Howler is the largest primate in the region and heavily hunted. It is protected in Caxinana National Forest, what remains of the damaged Gurupi Biological Reserve, and Tapirapé Biological Reserve to the south — all in Para.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F332FF5EF5E469C2F675.taxon	vernacular_names	Spix’s Howler	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F332FF5EF5E469C2F675.taxon	materials_examined	Forte Curupa (= Gurupa), Para, Brazil.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F332FF5EF5E469C2F675.taxon	discussion	The type locality is on the southern bank of the Rio Amazonas, not on the island of Gurupa, which is occupied by A. macconnelli. Formerly believed to be a subspecies of A. belzebul. Although C. P. Groves in 2001 considered A. discolor to be a synonym of A. belzebul, R. Gregorin in 2006 concluded it was a distinct species based on the morphology of the cranium and hyoid apparatus and pelage color and patterns. On the left bank of the Rio Santa Helena, it is sympatric with A. seniculus puruensis. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F332FF5EF5E469C2F675.taxon	distribution	Distribution. Lower Amazonian Brazil S of the Rio Amazonas, from the right bank of the rios Tapajos and Juruena to the rios Xingu and Iriri; its distribution extends E of the Rio Xingu near its mouth to Gurupa I, and S at least as far as the left bank of the Rio Santa Helena (left bank tributary of the Rio Teles Pires in Mato Grosso State), and the Serra do Cachimbo in Para State.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F332FF5EF5E469C2F675.taxon	description	Descriptive notes. Head — body 57 - 91 - 5 cm (males) and 46 - 5 - 57 cm (females), tail 65 — 67 - 5 cm (males) and 60 - 5 - 65 cm (females); weight 6 - 5 - 8 kg (males) and 4 - 8 - 6 - 2 kg (females). General color of Spix’s Howler varies from dark brown to black, but a dorsal band, hands, feet, and tip of the tail are rufous-chestnut or mahogany red. The dorsal band is absent or week in young individuals. Spix’s Howlers have distinctly long lateral hairs.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F332FF5EF5E469C2F675.taxon	biology_ecology	Habitat. Tall terra firma forest, igapo (seasonally flooded forest along clear or black rivers), palm swamps, and secondary forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F332FF5EF5E469C2F675.taxon	food_feeding	Food and Feeding. The diet of Spix’s Howler was studied by L. Pinto at Paranaita, Rio Santa Helena, Mato Grosso, for one year (1999 - 2000). The group of 7 - 9 individuals used 197 individual plants and ate soil from seven arboreal termitaria. Plant food sources comprised 67 species in 24 families; 17 species of Fabaceae accounted for 37 % of the feeding records. An additional 17 species were Moraceae, including Ficus (four species), Brosimum, Castilla, Helicostylis, Maquira, and Pseudolmedia, and accounted for 29 % of the plant feeding records and 38 % of the fruit feeding records. Spix’s Howlers ate flowers, particularly from three species of Bignoniaceae (62 % of the records). Ripe fruits were predominant in the diet in all months but one, and their contribution to the diet was similar between wet and dry seasons. Most fruits were available for one or two months. Fruits of Dialium guianense (Fabaceae) were eaten for six months, and they provided 26 % of the fruit part of the diet and 15 - 5 % of the overall diet. Fruits of this species were abundant in the dry season and made up most of the diet at that time. Young and mature leaves of Acacia ct. polyphylla were eaten in every month, contributing 0 - 4 - 21 - 2 % to the monthly diet. The annual diet was 55 % ripe fruit and some unripe fruit (0 - 6 %), 19 - 8 % young leaves, 5 - 7 % flowers, 5 % mature leaves, 4 - 4 % bark, 3 - 2 % decaying wood, 2: 3 % live wood, 0 - 5 % soil from termitaria, and small amounts of twigs, petioles, and orchid pseudobulbs. Compared with the diets of other howlers, Spix’s Howler has one of the highest rates of frugivory.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F332FF5EF5E469C2F675.taxon	breeding	Breeding. L. Pinto observed copulations of Spix’s Howlers with an average duration of 40 seconds, preceded by the male offering to groom the female. She reported three births in April, August, and September 2000. One of births occurred in a tall tree with its canopy covered with vines.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F332FF5EF5E469C2F675.taxon	activity	Activity patterns. In the study at Paranaita, the daily activity budget over the year was 587 % resting, 20 % feeding, 14 - 2 % traveling, 4 % moving around in trees, and 3 - 1 % engaged in social interactions and other activities. Spix’s Howlers tend to retire to their sleeping trees earlier in the wet season. Most sleeping trees had a canopy covered with tangles of vines, where the individuals could be well hidden.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F332FF5EF5E469C2F675.taxon	biology_ecology	Movements, Home range and Social organization. Groups of Spix’s Howlers are unimale-multifemale or multimale-multifemale, with 2 - 9 individuals. The group of 7 - 9 individuals studied in Paranaita initially had an adult male, two adult females, one subadult male, one subadult female, and two female infants. In the course of 13 months, in addition to changes in ages of some members, three infants were born and the subadult male emigrated. The home range of this group was 69 ha, mostly in terra firma forest but also including areas of Euterpe palm swamp and igapo. Seasonal migration was observed during peak fruiting in igapo during the flood season (January — April). Daily movements of this group were 176 - 1425 m, with an average of 761 m / day throughout the year and with no seasonal variation.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F332FF5EF5E469C2F675.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. There is only one study of the Spix’s Howler in the wild. Hunting and deforestation are occurring throughoutits distribution, and it is particularly threatened by the advancing agricultural frontier along the Santarém-Cuiaba Highway, where the land is logged, deforested, and turned into cattle ranches and industrial plantations (e. g. soy bean). Tapajos, Itaituba I, Itaituba II, and Altamira national forests are within its distribution, but they are notstrictly protected.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F331FA50F5AC69FBFD65.taxon	vernacular_names	Maranhao Red-handed Howler	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F331FA50F5AC69FBFD65.taxon	materials_examined	Brazil, Miritiba (= Humberto de Campos), northern Maranhao.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F331FA50F5AC69FBFD65.taxon	discussion	Formerly believed to be a subspecies of A. belzebul. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F331FA50F5AC69FBFD65.taxon	distribution	Distribution. NE Brazil in remnant forest patches in the N of the states of Maranhao, Piaui, and Ceara, from the municipality of Humberto de Campos E to the Serra da Ibiapaba in the municipality of Corean; it is also found in the municipalities of Ilha Grande do Piaui, Caxing 6, Campo Maior, and Jatoba do Piaui in the S of its range.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F331FA50F5AC69FBFD65.taxon	description	Descriptive notes. Head-body of adult male type specimen 56 - 5 cm, tail 56 cm. No specific data are available for body weight. The Maranhao Red-handed Howler is sexually dichromatic. The male is black, with rufous to reddish brown hands and feet, tail tip and flanks. The female is yellowish-brown with sparse grayish hairs, giving an olivaceous appearance.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F331FA50F5AC69FBFD65.taxon	biology_ecology	Habitat. Patches of tall forest along the coast, semi-deciduous forests in the caatinga (dry forest scrub), transitional Babassu palm (Orbygnia) forests in Maranhao, and mangroves on the coast of Piaut and Maranhao.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F331FA50F5AC69FBFD65.taxon	food_feeding	Food and Feeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F331FA50F5AC69FBFD65.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F331FA50F5AC69FBFD65.taxon	activity	Activity patterns. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F331FA50F5AC69FBFD65.taxon	biology_ecology	Movements, Home range and Social organization. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A16F331FA50F5AC69FBFD65.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List. The Maranhao Red-handed Howler is classified as critically endangered on the Brazilian list of threatened species. It is the least studied of all howlers and occupies a region with a long history of human occupation and high levels of forest loss and hunting. Today, hunting is seen as the most serious threat. Surveys in 2004 - 2006 by staff of the Brazilian government's National Center for Research and Conservation of Brazilian Primates mapped 19 localities where Maranhao Red-handed Howlersstill survive in very small numbers. It is possible that it occurs on the Ubajara National Park that protects a narrow strip of forest on the eastern slopes of the Serra da Ipiapaba in Ceara. Populations of Maranhao Red-handed Howlers have been found in the environmental protection areas of Serra da Ipiapaba in Ceara and Piaui and Delta do Parnaiba that extends across Piaui and Ceara into Maranhao.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33FFF96FE98672FF67E.taxon	vernacular_names	Brown Howler	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33FFF96FE98672FF67E.taxon	materials_examined	Brazil. Restricted by Cabrera in 1957 to the Rio Paraguacu, Bahia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33FFF96FE98672FF67E.taxon	discussion	The taxonomy of A. guariba is poorly understood. It is phenotypically variable and descriptions of the pelage color and patterns are insufficient to identify them in the wild. The validity of separating northern and southern populations as distinct subspecies is hampered by its extreme rarity in the northern part ofits distribution, supposedly the domain of the nominate subspecies. Following his study of cranial morphology and pelage patterns of the Brazilian howlers, R. Gregorin in 2006 argued that the two subspecies of the Brown Howler should be considered as distinct species. He used the name fusca of E. Geoffroy Saint-Hilaire in preference to Humboldt's guariba, indicating that the name guariba is a junior synonym of A. belzebul. Molecular genetic studies by E. Harris and colleagues in 2005 identified distinct haplotypic groups in the southern part of the distribution: one in the states of Rio Grande do Sul and Santa Catarina and the other in the states of Espirito Santo and Rio de Janeiro. It is possible that these represent two subspecies or even distinct species. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33FFF96FE98672FF67E.taxon	distribution	Subspecies and Distribution. A. g. guariba Humboldt, 1812 — Brazilian Atlantic Forest from the Rio Sao Francisco (although the Rio Paraguacu is the northernmost known locality) to the Rio Doce; however, based on his studies of morphology and pelage color patterns Gregorin in 2006 indicated that it extends from the Rio Paraguacu in Bahia State, S along the coast across the lower Rio Jequitinhonha extending inland somewhat into Minas Gerais State, S through the Rio Doce valley as far the Rio Paraiba do Sul Basin in Rio de Janeiro. A. g. clamitans Cabrera, 1940 — Brazilian Atlantic Forest S of Rio Doce (but Gregorin has recently identified howlers of the upper Rio Doce, Rio Matip 6 and Caratinga, and the middle and upper Rio Jequitinhonha as marking the N limits of this subspecies) through the S of Espirito Santo to Rio de Janeiro, Sao Paulo, Parana, Santa Catarina, and Rio Grande do Sulstates (S limit is Rio Camaqua Basin in Rio Grande do Sul, 31 ° 10 ” S, where the Atlantic Forest gives way to the Pampas grasslands), and inland in the forested E of Minas Gerais (“ Zona da Mata ”) and the Brazilian Highlands; also in NE Argentina (E of the Mts in Misiones Province to ¢. 28 ° S).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33FFF96FE98672FF67E.taxon	description	Descriptive notes. Head-body 50 - 60 cm (males) and 44 - 54 cm (females), tail 52 - 67 cm (males) and 48 - 57 cm (females); weight 5 - 3 - 7 - 2 kg (males) and 4 - 1 - 5 kg (females). The Brown Howleris black or dark brown to reddish above, with the fur having a rather golden sheen to it. There is a triangular or transverse crest toward the front of the crown, where forward-pointing crown hairs meet short, backward-pointing forehead hairs. A heavy facial beard is present in both sexes. Females are generally paler overall. The “ Northern Brown Howler ” (A. g. guariba) is not sexually dichromatic; both sexes are normally red-fawn, with females usually somewhat duller in color. Males from Minas Gerais are brown with an orange-brown rump, and females are yellow-brown. The “ Southern Brown Howler ” (A. g. clamitans) is generally dark reddish-brown. Males tend to be lighter colored than females. Males from Sao Paulo are orange-red to redbrown with a red belly, while males from Santa Catarina and Rio Grande do Sul are bright red-orange, with dark brown feet. Females are dark brown or blackish, or with the back frosted with orange or yellow-brown.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33FFF96FE98672FF67E.taxon	biology_ecology	Habitat. Lowland, submontane, evergreen, and semi-deciduous, seasonal, and secondary Atlantic coastal forest, including Araucaria (Araucariaceae) pine forest in the southern part of the distribution in Parana, Santa Catarina, and Rio Grande do Sul, from sea level to elevations of 700 m; not found in mangrove forests. Brown Howlers prefer to live in the vicinity of streams, and although they mainly use the upper forest canopy, they frequently descend to the ground to drink.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33FFF96FE98672FF67E.taxon	food_feeding	Food and Feeding. A year-long study by S. Mendes in 850 - ha forest in the Caratinga Biological Station in Minas Gerais, Brazil, showed that the diet of the Brown Howler was largely folivorous, but seasonal. In the dry season, leaves accounted for 77 - 7 % of the feeding records, flowers 10 - 9 %, buds 10 %, and fruits 1 - 4 %. In the wet season, leaves accounted for 63 - 5 %, fruits 29 - 8 %, flowers 5 - 8 %, and buds 0: 9 %. In the dry season, leaves in the diet were mostly mature (67 - 7 %), with young leaves accounting for only 6: 5 % (25 - 7 % undetermined). In the wet season, mature leaves accounted for 30 - 4 % and young leaves 48 - 7 %. Insect foraging was not observed, although insects were certainly eaten along with fruits and leaves. Another study by A. Chiarello in a 250 - ha seasonal, semi-deciduous secondary forest in the Mata de Santa Genebra Reserve, near Campinas in Sao Paulo also recorded a high percentage of leaves (73 %) in the diet, with flowers accounting for 13 % and fruits 6 % (8 % unidentified). Sixty-eight plant species were included in the diet. Fruits were a consistently small part of the diet throughoutthe year, but flowers assumed a greater importance (24 - 5 % ofthe diet) in the dry season. At least 59 % of the leaves in the diet were young. Brown Howlers were very selective and ate leaves from relatively few species. Predominant species were Celtis iguanaea (Ulmaceae), Aspidosperma polyneuron (Apocynaceae), and three members of the Fabaceae, Cassia ferruginea, Inga uruguayensis, and Piptadenia gonoacantha. A study in less diverse Araucaria (Araucariaceae) pine forest in the municipality of Balsa Nova, Parana, by J. Miranda recorded 34 plant species in the diet. Leaves accounted for 57 - 3 %, fruits 41 %, and flowers 1 - 7 % of the feeding records during one year. Ocotea (Laureaceae) was important for fruits and leaves and, as had been found in a study in Rio Grande do Sul by R. Silveira, Brown Howlers also ate fruits of exotic cultivated trees, such as Diospyros kaki (caqui, Ebeneceae) and Eriobotrya japonica (loquat, Rosaceae). A similar diet was noted in seasonal, semi-deciduous forest at Ibipora, Parana in a study by L. Aguiar and colleagues: 50 - 5 % leaves, 47 - 9 % fruits, 1 - 4 % flowers, and 0 - 2 % stems.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33FFF96FE98672FF67E.taxon	breeding	Breeding. Births of Brown Howlers occur throughout the year. Gestation is reported to be 190 days, with an interbirth interval of 21 months. Based on observations at Balsa Nova, an infant is first carried ventrally. After one month, it begins to move onto its mother’s back. Even in the first month, it leaves its mother momentarily, but serious independent exploration is seen only, and rather suddenly, from the fourth to fifth months. When an infant is four months old, it spends ¢. 20 % ofits time exploring, but by the fifth month, this increases to almost 90 %, with relatively little interaction with its mother. When an infant Brown Howler was two months old, it was sometimes carried by other group members, including a juvenile male, a subadult male, and an adult female. Tidbits of solid food were eaten in the second month, when it also hung from a branch with its prehensile tail for the first time. At two months of age, it also drank from a puddle alongside its mother. Play with a juvenile was seen when the infant was four months old. Males are sexually mature at c. 61 months and females at c. 43 months.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33FFF96FE98672FF67E.taxon	activity	Activity patterns. Daily activity patterns of Brown Howlers vary, with 10 - 21 % of the time spent feeding, 56 - 77 % resting, 11 - 24 % traveling, and 1 - 4 % in social activity.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33FFF96FE98672FF67E.taxon	biology_ecology	Movements, Home range and Social organization. Unimale — unifemale and multimale — - multifemale groups of Brown Howlers have been reported. Groups generally have 5 - 11 individuals, with 1 - 3 adult males and 1 - 4 adult females. Nineteen groups in Caratinga Biological Station had 3 - 10 individuals (average 6 - 8). Twenty-five groups at Serra da Cantareira in Sao Paulo averaged 5 - 8 individuals. Home ranges were 10 - 33 ha, and daily movements were 15 - 1564 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33FFF96FE98672FF67E.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List, including the subspecies clamitans, but the nominate subspecies guariba is classified as Critically Endangered. The conservation status is confused because exact distributional limits of Southern and Northern brown howlers are not clearly understood. Although Brown Howlers occur in the Atlantic Forests of Brazil and northeastern Argentina, a region with a high rate of historical forest loss, it remains widely distributed, is present in many protected areas, and is not declining at a rate sufficient to qualify for a threat category. It is probable that the Brown Howler extended north as far as the Rio Sao Francisco in the past, but it has been entirely extirpated from the northern part of its former distribution. Today, the northern limit is forests remaining in southern Bahia where it is found in small numbers in just a handful of localities. Primary threats are widespread forest loss and fragmentation, hunting for its meat in some areas, and disease epidemics (yellow fever brought from Africa). Nevertheless, the Brown Howler is able to survive in even quite small forest fragments if there is no hunting. Selective logging for prized Araucaria pine is a threat in the southern part of its distribution. In Rio Grande do Sul, itis an important “ flagship ” species for conservation of the state’s remaining forests and even protection of forest patches in the urban district of the capital, Porto Alegre.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33EFA91F5AA6629F5F3.taxon	vernacular_names	Paraguayan Howler	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33EFA91F5AA6629F5F3.taxon	materials_examined	Paraguay.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33EFA91F5AA6629F5F3.taxon	discussion	Molecular genetic studies have shown that this species is sister to the Amazonian red howlers (A. seniculus, A. sara, and A. macconnelli); they diverged from a common ancestor c. 4 million years ago. A. caraya hybridizes with A. guariba in certain narrow zones of contact in northern Argentina (Misiones Province) and southern Brazil (states of Parana and Rio Grande do Sul). Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33EFA91F5AA6629F5F3.taxon	distribution	Distribution. C Brazil (S of the states of Para, Tocantins, Maranhao — upper Rio Paranaiba — and Piaui, also in W Bahia, Rondonia, Mato Grosso — N to Rio Arrais, W to the Pantanal in the Rio Paraguay Basin, Goias, Minas Gerais, and Mato Grosso do Sul states; in the S, it occupies the bush savanna (cerrado) regions of the Rio Parana Basin in the N and NW of Sao Paulo State, and the W ofthe states of Parana, Santa Catarina, and Rio Grande do Sul), N Argentina (Misiones, Corrientes, E of Formosa and Chaco, and N of Santa Fe provinces, in the basins of the rios Parana, Uruguay, and Paraguay), Paraguay (E forests and in some areas of the Chaco where it occupies gallery forests along the rios Paraguay and Pilcomayo), and E & SE Bolivia (as far S as the Yungas of Chuquisaca and Tarija departments and the headwaters of the rios Pilcomayo and Pilaya); its distribution may also extend into the NW of Uruguay.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33EFA91F5AA6629F5F3.taxon	description	Descriptive notes. Head — - body 60 - 65 cm (males) and ¢. 50 cm (females), tail 60 - 65 cm (males) and 54 - 5 - 60 cm (females); weight 5 - 3 - 9 - 6 kg (males) and 3 - 6 - 6 - 5 kg (females). Adult Paraguayan Howlers are sexually dichromatic, but both sexes are born blonde with hair that gradually changes to adult colors. As adults, mature males are generally uniformly black, and females and young of either sex are pale grayish-yellow or golden-brown. Male Paraguayan Howlers from Bahia and Goias are black, while those from Mato Grosso and Parana have a brown back and hind parts, in addition to a black head and forequarters. Males from Sao Paulo and Minas Gerais, meanwhile, are brown-black with yellowish hands, feet, belly, and tail tip. Underparts of adult females are sometimes tinged with orange, and there is a darker brown to grayish line that runs from the crown along the back to the base of the tail. In all individuals, the face is invariably dark, fur is stiff and lengthy, and the beard is prominent. Hairs on the crown grow entirely forward from a whorl on the nape to overhang the brows, with the exception of a small upright fringe behind the brows that meets the forward hairs in a small crest. The scrotum is rust-red.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33EFA91F5AA6629F5F3.taxon	biology_ecology	Habitat. Much of the range of the Paraguayan Howler is in the cerrado of central Brazil and semi-arid caatinga forest scrub in north-eastern Brazil, where they use gallery and riparian forest and patches of seasonal semi-deciduous and deciduous forests (mesic Atlantic Forest, “ cerradao, ” and arboreal caatinga). In southern Rondonia, they are associated with highland savanna habitats and gallery forest in the Guaporé grasslands. In Paraguay, they occur in eastern forests and, in certain areas of the Chaco, gallery forests along the rios Paraguay and Pilcomayo and numerous small rivers draining the Chacoan plain. In Bolivia, they are found in riparian forest in the Chaco along the Rio Parapeti, the inundated and terra firma lowland and submontane dry Chiquitania forest. In the southern part ofits distribution, high numbers of Paraguayan Howlers occur in flooded (seasonally or irregularly) and highly productive riparian forest in the basins of the rios Parana and Paraguay in northern Argentina and the Pantanal in Bolivia and Mato Grosso, Brazil. Periodic flooding results in a continual deposition of alluvial sediments. The diversity of trees is not high; the flooding regime selects for those that can tolerate having their roots underwater for prolonged periods (occasional very prolonged floods can result in widespread die off of trees). A botanical survey of a 1 - ha plot on the island of Brasilera in the Rio Parana, Argentina, contained just 23 tree species. The flora of these islandsis evidently not rich, but it is optimal for Paraguayan Howlers. Some populations in Argentina include pine plantations in their home ranges.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33EFA91F5AA6629F5F3.taxon	food_feeding	Food and Feeding. In a study by S. Bravo and A. Sallenave on the fluvial island of Brasilera, the annual diet of Paraguayan Howlers included parts of 22 plant species from 15 families, mainly fruits, leaves (young and mature), and flowers. Annually, mature leaves made up 51 % ofthe diet, ripe fruits 19 % (but as high as 71 % in some months), young leaves 13 %, flowers 13 %, and buds 4 %. Other items eaten in small quantities included petioles, buds, and aquatic plants. In the cerrado, Paraguayan Howlers eat roots of the marsh-living spikesedge Eleocharis (Cyperaceae), which are also a good source of water. When fruits, young leaves, or flowers were unavailable, they ate mature leaves, forming the large majority of the diet in May-July. It is possible that palatable mature leaves were a scarce commodity at this time, as evidenced by the groups splitting into two subgroups each day, coming together only in the late afternoon to sleep. Animal protein came from arthropods consumed with leaves (e. g. aphids or galls) or fruits such as Ocotea (Lauraceae) and Ficus (Moraceae). Figs contained galls inside their fruits. Each month, the principal food item of Paraguayan Howlers was heavily infested, and in the case of Ocotea, they evidently chose to feed from trees suffering from these infestations. Fruits of Ocotea diospyrifolia, for example, tended to be infested by curculionid beetles (weevils). Howlers ingesting these fruits kill parasitic beetles but do not harm seeds, which are effectively dispersed. Paraguayan Howlers feed more in the more infested trees, even going into a neighboring group’s home range to find them. Insect-infested food items accounted for ¢. 25 % of the diet during the year. These Paraguayan Howlers were living in very productive forests. In terra firma forests nearby, leaves can make up 70 % of the annual diet. A year-long study of several populations of Paraguayan Howlers in north-eastern Argentina found that leaves made up 64 - 82 % of the diet, fruits 12 - 19 %, pine cones 6 - 7 %, flowers 0 - 3 %, bark 0 - 7 %, and other items 0 - 1 %. This is the only howler in which some populations feed on pine cones. They can also be seed predators. In Brazil, they open immature, oblong, woody pyxidia of Cariniana estrellensis (Lecythidaceae), picking them off the tree, and holding them so that they can bash them sideways, while raising the arm above the head to do so. After each strike, they look at the fruit to see if the operculum is open. When it is open, they pull it out and raise the pyxidium above their head and pour the seeds into their mouth. Howlers fed on these fruits at a time when other fruits and leaves were scarce, and males aggressively defended the tree against other howlers when feeding.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33EFA91F5AA6629F5F3.taxon	breeding	Breeding. Male Paraguayan Howlers reach sexual maturity at 24 - 37 months and females at 35 - 42 months. In captivity, females first reproduce at 35 - 42 months old. Gestation is ¢. 180 days. The ovarian cycle is ¢. 20 days. Annual birth rate in the wild in northern Argentina has been recorded at 0 - 89 infants / female / year. Births occur throughout the year in most populations, but they are seasonally facultative with extreme changes in food availability. In terra firma gallery forest in northern Argentina, conceptions tend to occur in the wet season when food is abundant, with births occurring in the dry season when food is scarce so that weaning is timed to coincide with food abundance (peaking in April-August). In flooded forest, however, food availability is much less seasonal, and births tend to occur throughout the year. Alloparental care is common. Males carry infants and helpjuveniles, and females suckle infants and even juveniles that are not their own. A case of adoption (the mother died) has also been recorded. Weaning occurs at 9 - 12 months. Interbirth intervals in captivity are 7 - 15 - 5 months, and in the wild, they average 15 - 9 months.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33EFA91F5AA6629F5F3.taxon	activity	Activity patterns. Activity patterns of Paraguayan Howlers are extremely variable, depending on group size and food availability. They generally rest 46 - 77 % of the day, travel 2 - 30 %, feed 13 - 18 %, engage in social activities 3 - 9 %, and other behaviors 1 - 5 %. In the study of groups on Brasilera Island, two groups rested 57 % ofthe day, fed 19 %, traveled 16 %), engaged in social behaviors (mainly allogrooming and play) 6 %, and confronted other groups 2 %.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33EFA91F5AA6629F5F3.taxon	biology_ecology	Movements, Home range and Social organization. Groups of Paraguayan Howlers are unimale-multifemale or multimale-multifemale, with 3 - 22 individuals. In general, groups are quite small, and high densities are often characterized by numerous small groups in small home ranges (2 - 4 ha). Groups are large in seasonal deciduous and semi-deciduous forests of islands in the Rio Parana in northern Argentina. They are believed to result from difficulties in dispersing in riparian forest islands and forest patches. Two groups studied on the island of Brasilera (Rio Parana) had 16 and 22 individuals: one with two adult males and five adult females and the other with two adult males and six adult females. In Brazil, between the rios Paranapanema and Ivinhema, 13 groups had 6 - 18 individuals (average 11 - 5), with 1 - 3 adult males. Seventeen groups on the island of Tragadero Sur also in the Rio Parana had 4 - 14 individuals, with 1 - 9 adult males and 1 - 5 adult females. In terra firma areas and especially dry forests, groups of Paraguayan Howlers tend to be smaller. At a site in the eastern Argentinean Chaco, seven groups had 2 - 10 individuals (average 5 - 7), with densities of 12 - 26 ind / km? Home range size varies from less than 2 ha to 117 ha, with daily movements of 75 - 1666 m. In riparian forests of the Chaco in Kaa-lya del Gran Chaco National Park, Bolivia, densities were 15 - 187 ind / km? ®. The density on the island of Brasilera in Argentina was very high at 425 ind / km? *. When home ranges are very small, daily movements are as low as 85 m but up to 1500 m, and group encounters are frequent, up to six a day. Both sexes disperse. Juvenile and subadult females spend some time on the periphery of their natal group’s home range before they are evicted by adult females that vocalize and threaten and chase them. Infanticide has been reported after a male enters a group and takes over as the dominant male. Females do not enter established groups, but form new groups with dispersing males in new territories. Females form a dominance hierarchy, with the youngest being the most dominant.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1BF33EFA91F5AA6629F5F3.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. The Paraguayan Howler is a wide-ranging species and in some areas common. Nevertheless, over large expanses ofits distribution, populations have been reduced or lost as suitable habitat becomes highly fragmented due to conversion for farmland, including commercial plantations (tees and crops) and pasture. Enormous numbers of Paraguayan Howlers are lost when rivers are dammed. The Paraguayan Howler is wide-ranging and adaptable; in some regions, it can be found in small, semi-natural and degraded forests, able to thrive on cultivated species such as bamboos, eucalyptus, tangerines, peaches, guavas and timber trees such as Melia azedarach (Meliaceae) and Schizolobium parahyba (Fabaceae). They are generally not hunted for food (their flesh is considered bitter), but the plush of the male is much sought after by local people to make caps, bags, and blankets.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1AF33DFA46F5176969FB4D.taxon	vernacular_names	Mantled Howler	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1AF33DFA46F5176969FB4D.taxon	materials_examined	Caracas, Venezuela. Restricted by Allen in 1910 to Lago Nicaragua, Nicaragua.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1AF33DFA46F5176969FB4D.taxon	discussion	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.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1AF33DFA46F5176969FB4D.taxon	distribution	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).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1AF33DFA46F5176969FB4D.taxon	description	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.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1AF33DFA46F5176969FB4D.taxon	biology_ecology	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.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1AF33DFA46F5176969FB4D.taxon	food_feeding	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).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1AF33DFA46F5176969FB4D.taxon	breeding	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.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1AF33DFA46F5176969FB4D.taxon	activity	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.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1AF33DFA46F5176969FB4D.taxon	biology_ecology	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).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1AF33DFA46F5176969FB4D.taxon	conservation	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.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A19F33CFA90FABC66C7FB00.taxon	vernacular_names	Central American Black Howler	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A19F33CFA90FABC66C7FB00.taxon	materials_examined	Uaxactun, Petén, Guatemala.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A19F33CFA90FABC66C7FB00.taxon	discussion	The Central American Black Howler is genetically distinct and is believed to have diverged from A. palliata c. 3 million years ago. It is sympatric with A. palliata in Tabasco State, Mexico, and Chocon Machacas Biotope and Mario Dary Rivera Biosphere Reserve in Guatemala. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A19F33CFA90FABC66C7FB00.taxon	distribution	Distribution. SE Mexico (Tabasco, Campeche, Yucatan, Quintana Roo, and S Chiapas, states), Belize, and N & C Guatemala; the S limits of its distribution are defined by Lago de Izabel, El Golfete, and Rio Dulce in Guatemala.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A19F33CFA90FABC66C7FB00.taxon	description	Descriptive notes. Head-body 65 - 71 cm (males) and 52 - 64 cm (females), tail 60 - 69 cm (males) and 50 - 54 cm (females); weight 7 - 5 - 9 - 1 kg (males) and 4 - 6 - 7 kg (females); all measurements from Belize. In 1935, A. Murie reported that the largest of three male Central American Black Howlers that he shot in Guatemala weighed 11 - 1 kg (the only male weight he provided). No individuals have been found that weigh that much since. Fur of the Central American Black Howler is notably long, soft, and dense. Adult males and females are black with traces of brown on shoulders, cheeks, and back. There is a slight crest on the crown; fur comes forward from a whorl on the nape and then suddenly becomes very short and upright, with a slight parting going back to the whorl. Males older than four months have a conspicuous white scrotum.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A19F33CFA90FABC66C7FB00.taxon	biology_ecology	Habitat. Primary and secondary forest, high-elevation and lowland rainforest, and semi-deciduous evergreen and riparian lowland forest; riparian and seasonally flooded forests are particularly favored. Like the Mantled Howler (A. palliata), the Central American Black Howleris often seen in the vicinity of Mayan archaeological sites.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A19F33CFA90FABC66C7FB00.taxon	food_feeding	Food and Feeding. Diets of Central American Black Howlers vary seasonally. The feeding ecology of two groups was studied by S. Silver and colleagues for a year in the Community Baboon Sanctuary on the banks of the Belize River. They fed on 74 plant species, 53 of which were trees and the remainder epiphytes or lianas. Fruits and young leaves, and, to a lesser extent, mature leaves of nine species of Ficus (Moraceae) were eaten. The average annual diet of the two groups was 40 - 8 % fruits (monthly range 10 - 5 - 64 - 5 %), 37 - 2 % young leaves (18: 1 - 64 - 6 %), 10 - 6 % flowers, 7 - 9 % mature leaves (0 - 2 - 31 - 4 %), and 3 - 5 % other items such as pulvini (a joint-like thickening at the base of the leaf) and leaf petioles (leaf stalks). Thirty-two percent of their feeding time was dedicated to Ficus (monthly range 6 - 65 % of feeding records). They ate mature leaves from four species: Ficus maxima and FE insipida (both Moraceae), Sapindus saponaria (Sapindaceae), and Cecropia (Urticaceae). Flowers were an important source of food when fruits were most scarce in the early dry season.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A19F33CFA90FABC66C7FB00.taxon	breeding	Breeding. Births of Central American Black Howlers occur throughout the year. When a female is receptive, a male follows her around, resting beside her, in contact, and sniffing her urine. Females show anogenital rubbing on a branch, which males sniff with interest. Males put their heads up and rub their chins and chests on branches at this ime. The female gazes at the male and flicks her tongue slowly prior to copulation. Infants are initially carried on the ventrum. Males and females, besides the mother carry young (allomothering). The transfer of young is described as passive: a female approaches and remains near the mother and infant, looking, sniffing, and occasionally touching the infant, and then extends her neck or chest over the infant and coddles it onto her body. Interest from allomothers peaks twice, once when infants are 2 - 3 weeks old and again when they are 6 - 10 weeks old. Experienced mothers tend to carry infants more during the first peak, and males and inexperienced females during the second peak. Infants begin to spend more time off their mothers and allomothers at 9 - 10 weeks old.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A19F33CFA90FABC66C7FB00.taxon	activity	Activity patterns. The Central American Black Howler is diurnal and arboreal. A yearlong study in Belize demonstrated that activity patterns are flexible. A typical profile is 61 - 9 % resting (range 50 - 67 - 7 %), 24 - 4 % feeding (19 - 1 - 31 - 2 %), 9 - 8 % resting (6: 6 - 13 - 5 %), 2: 3 % engaged in social affiliative behaviors (playing and grooming), and 1 - 6 % vocalizing.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A19F33CFA90FABC66C7FB00.taxon	biology_ecology	Movements, Home range and Social organization. Groups of Central American Black Howlers are unimale — multifemale or multimale — multifemale, with 3 - 9 individuals. Twenty groups of Central American Black Howlers at Palenque, Chiapas, Mexico (humid, evergreen forest) averaged seven individuals (range 2 - 12), with 1 - 4 adult males and 1 - 4 adult females. Home range sizes were 25 - 50 ha and varied with group size. Males disperse, and group takeovers by males are violent. Infanticide has been observed following the immigration of a new male and the expulsion of the previous dominant male. In Belize, densities are 8 - 22 ind / km?. In Tikal, Guatemala, densities are low at 5 - 9 ind / km?, but they are higher at 16 - 5 ind / km? * in the Muchukux forest in Quintana Roo and 23 ind / km? * at Palenque, Chiapas, Mexico.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A19F33CFA90FABC66C7FB00.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The [UCN Red List. The Central American Black Howler continues to decline in numbers because of severe loss of habitat over much of its range. It is still occasionally hunted, and suffers from yellow fever epidemics and hurricanes. It can reach very high numbers when not hunted, even near human settlements. A remarkable initiative for their conservation is that of the Community Baboon Sanctuary in Belize, where a pioneer program of community land management with subsistence farmers has been set up to specifically accommodate the needs of the Central American Black Howler. Management agreements guarantee permanence of mosaics and corridors of forests, including riparian forest, and leave intact key food resources of howlers, while allowing for the traditional slash-and-burn agriculture by local villagers. Effective ladder bridges have been put in place where roads separate the forest canopy. In 1990, the Sanctuary included more than 100 people in seven villages over an area of more than 46 km? and the program has grown since then. From 1985 (when the program began) to 1999, the population of Central American Black Howlers grew by 8 - 9 ind / year, from 840 to more than 2000 individuals. The density estimate in 1999 was 178 ind / km? ®. Villagers have also benefitted from tourists interested in seeing howlers.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1DF338FF80FE73622DFAB5.taxon	vernacular_names	Central American Spider Monkey	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1DF338FF80FE73622DFAB5.taxon	materials_examined	type locality unknown. Restricted by R. Kellogg and E. Goldman in 1944 to San Juan del Norte (= Greytown), Nicaragua.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1DF338FF80FE73622DFAB5.taxon	discussion	Ateles geoffroy: 1 s highly variable, both individually and geographically, and its taxonomy may still require revision. Seven subspecies are typically recognized: geoffro, azuerensis, Jfrontatus, grisescens, ornatus, vellerosus, and yucatanensis. The form grisescens, however, is of doubtful validity, having never been observed in the wild and being the subject of a single rather poor account regarding a captive specimen. P. H. Napier in her 1976 catalogue of the primate collection in the British Museum (Natural History) argued that the form panamensis is ajunior synonym of ornatus. There is a hybrid zone between the subspecies ornatus and A. fusciceps in Panama. Seven subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1DF338FF80FE73622DFAB5.taxon	distribution	Subspecies and Distribution. A. g. geoffroyi Kuhl, 1820 — S & SE Nicaragua (coastal region around San Juan del Norte or Martina Bay, probably ranging across the lowlands to the vicinity of Lake Managua and Lake Nicaragua on the Pacific coast); possibly in N Costa Rica. A. g. azuerensis Bole, 1937 — SC Panama, known only from the forested mountains of the W side of the Azuero Peninsula (Veraguas Province) in the vicinity of Ponuga, where it appears to be isolated; it may also occur to the W along the Pacific coastto the Burica Peninsula, near the Panamanian and Costa Rican border. A. g. frontatus Gray, 1842 — N & W Nicaragua and NW Costa Rica. A. g. grisescens Gray, 1866 — S Panama along the Pacific coast in the valley of the Rio Tuyra and SE through the Serrania del Sapo of extreme SE Panama into the Cordillera de Baudo of NW Colombia. A. g. ornatus Gray, 1871 — C & E Costa Rica, and Panama (from Chiriqui Province to the Serrania de San Blas E of the Canal Zone). A. g. vellerosus Gray, 1866 — E & SE Mexico (E San Luis Potosi, Veracruz, Tabasco, E Oaxaca, and Chiapas states), Guatemala (including the highlands), El Salvador, and Honduras (along the N coastto the lowlands of La Mosquitia in Gracias a Dios Department). A. g. yucatanensis Kellogg & Goldman, 1944 — SE Mexico (forests of the Yucatan Peninsula), NE Guatemala, and adjoining parts of Belize; intergrading in S Mexico (Campeche State) and Guatemala with vellerosus.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1DF338FF80FE73622DFAB5.taxon	description	Descriptive notes. Head — - body 39 - 63 cm (males) and 31 - 45 cm (females), tail 70 - 86 cm (males) and 64 - 75 cm (females); weight 7 - 4 - 9 kg (males) and 6 - 9 - 4 kg (females). The Central American Spider Monkey is extraordinarily variable in its coloration, from yellowish through reddish to blackish-brown on the back and flanks, although there is usually black on the head and limbs and white on the muzzle or around the eyes. Forward-directed hairs from the nape form a sort of cowl that ends in a triangular crest above the brows, and cheek hairs stand out prominently. Roughly from north to south through their distributions, the subspecies appear as follows. Dorsal surfaces of the “ Mexican Spider Monkey ” (A. g. vellerosus) ranges from black to dark brown, exceptfor a lighter band across the lumbar region, and contrasts strongly with its lighter abdomen and inner limbs. Exposed, flesh-colored skin is often present around the eyes. The “ Yucatan Spider Monkey ” (A. g. yucatanensis) is a relatively small, slender subspecies. Its fur is brownish-black on the head, neck, and shoulders, becoming lighter brown on the lower back and hips and contrasting with its silvery-white underside, inner limbs, and sideburns. “ Geoffroy’s Spider Monkey ” (A. g. geoffroyi) is silvery to brownish-gray on the back, upper arms, and thighs. A variable amount of black exists on elbows, knees, upper arms, and lower legs, while hands and feet are always black. Its chest coloration is similar to the back, but the lower abdomen may be somewhat golden. The top of its head is dark, sometimes black, with mixed light and dark hairs directed forward, often forming a distinct light band over the forehead. Its face is black, and flesh-colored “ spectacles ” around the eyes are common. Lighter side whiskers are often present. The tail is bicolored. The “ Black-browed Spider Monkey ” (A. g. frontatus) is similar in color pattern to the nominate subspecies, although somewhat darker. The “ Ornate Spider Monkey ” (A. g. ornatus) is a uniformly darker brown than either Geoffroy’s Spider Monkey or the Black-browed Spider Monkey. Its back is golden brown and the face, top of the head, forearms all around, outersides of legs, hands, and feet are black. Its underside does not contrast strongly with the dorsal coat color. The back of the “ Azuero Spider Monkey ” (A. g. azuerensis) is grayish-brown and somewhat darker than the underside. As in the Ornate Spider Monkey, outer surfaces of its limbs are black, but the top of its head and neck are either black or brownishblack.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1DF338FF80FE73622DFAB5.taxon	biology_ecology	Habitat. Primary and secondary lowland rainforest; evergreen, semi-deciduous and cloud forest; and mangrove swamp. The Central American Spider Monkey will also use deciduousforest (Santa Rosa National Park, Costa Rica).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1DF338FF80FE73622DFAB5.taxon	food_feeding	Food and Feeding. In a nine-month study of the diet of the Yucatan Spider Monkey by J. Cant in Tikal National Park in Petén, Guatemala, 57 % of their time was spent feeding on fruit, 14 - 1 % leaves, 10 - 5 % seeds (almost entirely unripe seeds of the breadnut or Maya nut tree, Brosimum alicastrum, Moraceae), 4 - 9 % unidentified reproductive parts of Brosimum (female flowers, very young fruit, and perhaps male flowers), 2 % caterpillars (infested young leaves of Brosimum), and 1 - 4 % buds (11 - 1 % other items). Depending on the month, fruit comprised up to 84 % of the diet and Brosimum seeds 57 %. Ninety-seven percent of the leaves eaten were young, and 61 % of young leaves and 68 % of mature leaves were from Brosimum. Caterpillars were believed to be the only source of animal protein, except for insects ingested incidentally with leaves and fruits such as figs. Brosimum was very important; 56 % of the diet was made up ofits fruits, seeds, leaves, and petioles. In upland forests of Tikal, species of Brosimum are very abundant: 11 % ofall trees and 21 % of the basal area. Other species ofsignificance providing fruits were Spondias mombin (Anacardiacaeae), Dendropanax arboreus (Araliaceae), Manilkara achras (Sapotaceae), a species of Ficus (Moraceae), and Cupania prisca (Sapindaceae). A study of the Black-browed Spider Monkey by C. Chapman in the deciduous forest of Santa Rosa National Park indicated a diet of 77 - 8 % fruit, 9 - 8 % flowers, 7 -: 3 % young leaves, 2: 6 % buds, 1 - 3 % insects, and 1 - 2 % mature leaves. Figs made up 29 - 2 % of the diet, and the sweetjuicy fruits of Muntingia calabura (Muntingiaceae) 16 - 1 %. Leaves, rather than fruits, made up 70 - 80 % of the diet in the early dry season (January-February). Overall, fruit comprised 14 - 100 % ofthe diet, depending on the month. Fruit was also the major componentof the diet of the Ornate Spider Monkey studied by C. Campbell on Barro Colorado Island, Panama: fruit 81 - 7 % (monthly range 69 - 91 %), leaves 16 - 7 % (6 - 32 %), flowers 1 % (0 - 9 %), and insects 0 - 6 % (0 - 2 %).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1DF338FF80FE73622DFAB5.taxon	breeding	Breeding. The ovarian cycle of the Central American Spider Monkey is 22 - 7 days, and menstrual bleeding, not always visible, lasts for 2 - 4 days. Males evidently show investigative behaviors by sniffing places where a female has been sitting, sniffing her urine, and holding and sniffing her elongated clitoris. These behaviors probably allow a male to assess the reproductive condition of a female. Pairs are more likely to mate when a female is cycling, but they also mate when she is not. Males are aggressive toward females, and mating may often occur by coercion. There is no specific behavior on the part of the female that indicates receptivity. Gestation is 226 - 232 days. Sex ratio at birth of Black-browed Spider Monkeys at Santa Rosa National Park is 1: 1. High numbers of infants have been recorded in the dry season (December — - May) in Santa Rosa. On Barro Colorado Island, most births occur in May-December, with conceptions when fruit is abundant or increasing in availability in December — May. Infants are entirely black, and their coat color lightens appreciably during their first five months. Infants are weaned at 24 - 26 months old, or more. Juveniles continue to associate with their mothers even when a new infant is born and only begin to move around independently when they are c. 3 years old. Females are sexually mature at c. 6 - 5 years old, and they first give birth at c. 7 years old. Interbirth intervals were 34 - 7 months at Santa Rosa and 32 months on Barro Colorado. Individuals maylive up to 27 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1DF338FF80FE73622DFAB5.taxon	activity	Activity patterns. A general activity budget of Black-browed Spider Monkeys at Santa Rosa National Park over three years was feeding 33 - 5 %, traveling 32 - 6 %, resting 24 - 6 %, and engaging in social and other behaviors 9 - 8 %.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1DF338FF80FE73622DFAB5.taxon	biology_ecology	Movements, Home range and Social organization. Group sizes of Central American Spider Monkeys are 16 - 24 individuals. Mean daily movement on Barro Colorado Island was 1203 m (range 460 - 2400 m); daily movements were greater when fruit was abundant in the early and middle wet season. In another study on Barro Colorado, the home range of a group of 20 - 24 Ornate Spider Monkeys was 963 ha. Their daily movements averaged 2055 m (maximum 4500 m). At Santa Rosa, the home range of a group of 42 Black-browed Spider Monkeys was 42 ha, and their daily movement averaged 1297 m. Female Central American Spider Monkeys disperse from their natal group, but males are generally philopatric. Males are dominant to females. Adult males are more aggressive, show more territorial behavior, and are more socially cohesive than females. Males patrol home range borders, generally in slightly larger subgroups than is typical when they are in other parts of their home range. Adult females are more vocal, more submissive, less social, and more dispersed than adult males. Females tend to travel alone with their infants and juveniles in groups of 3 — 4 individuals. Adult males travel together but form consortships with females when they are receptive. Densities of 24 - 5 - 45 ind / km? were estimated for the Yucatan Spider Monkey in seasonal dry forests of Tikal and they can be as high as 89 - 5 ind / km? in undisturbed moist forests, such as those at Punta Laguna, Quintana Roo State, Mexico.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1DF338FF80FE73622DFAB5.taxon	conservation	Status and Conservation. CITES Appendix I (subspecies frontatus and panamensis, the latter considered here a synonym of ornatus) and CITES Appendix II (other subspecies). Classified as Endangered on The IUCN Red List. Subspecies geoffroyi, azuerensis, and vellerosus are classified as Critically Endangered, ornatus and yucatanensis as Endangered, Jfrontatus as Vulnerable, and grisescens as Data Deficient on The IUCN Red List. The Central American Spider Monkey is threatened mainly by habitat loss and fragmentation, with several subspecies having been severely affected. Nevertheless, several large areas of relatively continuous habitat occur in Selva Maya (Mexico, Guatemala, and Belize), the Atlantic zone of Nicaragua and Honduras, and along the Atlantic coast and the Darien in Panama. It is also affected by pet trafficking and hunting for meat in some areas. [tis relatively common in a few parks, but locally extinct in some areas. Geoffroy’s Spider Monkey is known to occurin just one protected area, Arenal Volcano National Park in Costa Rica. Population surveys of the Azuero Spider Monkey confirmed the existence of 145 individuals in two of the three provinces where it once occurred (Veraguas and Los Santos); it is extinct in Herrera Province. It is still present in Cerro Hoya National Park and Montuoso and Tronosa forest reserves in Panama. The Black-browed Spider Monkey occurs in Barra Honda, Guanacaste, Palo Verde, Rincon de la Vieja, and Santa Rosa national parks in Costa Rica. The Ornate Spider Monkey occurs in at least 22 protected areas including Braulio Carrillo, Cahuita, Chirripo, Corcovado, and Manuel Antonio national parks and Carara Biological Reserve in Costa Rica. The Mexican Spider Monkey occurs in Palenque National Park and El Triunfo, Montes Azules, and Los Tuxtlas biological reserves in Mexico. The Yucatan Spider Monkey is found in Cockscomb Basin Wildlife Sanctuary in Belize; Tikal and Rio Dulce national parks and Sierra de las Minas Biosphere Reserve in Guatemala; and Tulum and Canon del Sumidero national parks and Calakmul, Ria Celestin, and Ria Lagartos biosphere reserves in Mexico. The “ Hooded Spider Monkey ” (A. g. grisescens) may occur in Canglon Forest Reserve in Panama, according to its supposed distribution.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF338FF47FAD164A5F9A1.taxon	vernacular_names	Brown-headed Spider Monkey	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF338FF47FAD164A5F9A1.taxon	materials_examined	South America. Restricted by R. Kellogg and E. Goldman in 1944 to Ecuador, Hacienda Chinipamba, near Penaherrera, Imbabura Province at 1500 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF338FF47FAD164A5F9A1.taxon	discussion	The form rufiventris is sometimes considered a distinct species. A hybrid zone exists in Panama between rufiventris and A. geoffroyi ornatus, even though cytogenetic studies by M. A. A. Medeiros and coworkers, published in 1997, indicated that rufiventris was reproductively isolated from A. geoffroyi. In a review in 1975, P. G. Heltne and L. M. Kunkel argued that the form robustus is a junior synonym of rufiventris. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF338FF47FAD164A5F9A1.taxon	distribution	Subspecies and Distribution. A. [. fusciceps Gray, 1866 — NW Ecuador, W of the Andes, historically in the provinces of Esmeraldas and Carchi, from the Colombian border S to the Cordillera de Colonche in the Guayas Province (most S record is in “ Puente sobre el rio Chimbo, ” Chimborazo Province), but today it is restricted to two remnant populations (Awa Ethnological Reserve N of the Rio Mira, and Cotacachi-Cayapas Ecological Reserve and adjacent Los Cedros Protected Forest and surrounding forests S of the Rio Mira). A. f. rufiventris Sclater, 1872 — E Panama (Atlantic slope) and W Colombia from the Uraba region in NW Antioquia, N through Cordoba, Sucre, and N Bolivar departments (N distributional limit on the S bank of the Canal del Dique, Cartagena), W of the Rio Cauca to the coast, E to the lower Rio Cauca along the W bank to SC Antioquia (the Cerro Pirre or the Rio Tucuti marks the border with A. geoffroyi grisescens), and S to the Cordillera Occidental of the Andes in SW Colombia (the most southerly record is Barabacoas, Narino Department).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF338FF47FAD164A5F9A1.taxon	description	Descriptive notes. Head-body 37 - 64 cm (males) and 30 - 54 cm (females), tail 71 - 86 cm (males) and 66 - 77 cm (females); weight ¢. 8 - 9 kg (males) and c. 8 kg (females). Fur of the Brown-headed Spider Monkeyis coarse and dark, with varying amounts of brown on cheeks and crown and sparse white hairs on lips and chin. Cheek hairs meet those of the crown in a small eye-to-ear crest. The nominate subspecies is brownishblack above with a yellow-brown anterior crown, grading from brown to black on the nape. It often has a white moustache and beard. The “ Colombian Black Spider Monkey ” (A. f. rufiventris) is glossy-black, except for a few whitish or golden hairs on cheeks and muzzle, and a slight brownish tinge on the forehead. Skin of the face is black. The genital area is reddish.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF338FF47FAD164A5F9A1.taxon	biology_ecology	Habitat. [Lowland to lower montane primary rainforest. In Ecuador, the Brown-headed Spider Monkey inhabits tropical lowland and subtropical humid forests at elevations of 100 - 1700 m. It survives today in the pre-montane wet forest of Cotacachi-Cayapas Ecological Reserve at elevations of 300 - 1200 m, butitis very scarce at higher elevations of 2500 m. The Colombian Black Spider Monkey occurs in dry, humid, and cloud forest, occupying the greatest range of forest habitats of any Colombian spider monkey. It occurs as high as 2000 - 2500 m above sea level in the Cordilllera Occidental.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF338FF47FAD164A5F9A1.taxon	food_feeding	Food and Feeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF338FF47FAD164A5F9A1.taxon	breeding	Breeding. Interbirth intervals are at 22 - 36 months (average 33 months). Infants have a pink face and red ears. They are dependent on nursing for ¢. 10 months and suckle until they are 18 - 20 months old. Individuals may live up to 24 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF338FF47FAD164A5F9A1.taxon	activity	Activity patterns. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF338FF47FAD164A5F9A1.taxon	biology_ecology	Movements, Home range and Social organization. Densities of Brown-headed Spider Monkeys, based on calls, were 7 - 5 ind / km? at 332 m above sea level to 0 - 9 ind / km? at 1570 m. In 2007, X. Cueva and W. Pozo surveyed five sites in the buffer zone of the Cotacachi-Cayapas Ecological Reserve and recorded densities of 0 - 6 - 16 - 6 ind / km?, with an average subgroup size of 3 - 8 individuals.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF338FF47FAD164A5F9A1.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Critically Endangered on The IUCN Red List, including both subspecies. Forest loss and hunting have devastated populations of both subspecies of the Brown-headed Spider Monkey. The nominate subspecies is protected by national law in Ecuador. It is restricted to three small forested areas: Awa Ethnological Reserve, Cotacachi-Cayapas Ecological Reserve, and adjacent Los Cedros Protected Forest in north-western Ecuador. Logging, road construction, and hunting are damaging forests and associated wildlife in Awa. At Los Cedros, density is 1 - 2 ind / km? ®. A recent attempt to recount the population there failed because numbers were so low. Total population of the nominate subspecies is believed to be in the low hundreds. Based on calculations from satellite photos, more than 30 % of the habitat of the Colombian Black Spider Monkeys along the Pacific coast in Colombia has been lost in the past ten years. Ground-truthing of these data found that only 2: 5 % of viable secondary forest is left in this region. In Panama, there is a lower human population and likely higher habitat availability, but more research is needed to determine the status of the Brown-headed Spider Monkey and the threats to its populations. Its habitats are entirely fragmented in the southern part of it distribution, and there are only two known locations where it occurs in the Choco region in Colombia (Las Orquideas and Los Katios national natural parks). It may also occur in Chagres and Darién national parks in Panama. Parks such as Los Katios and Las Orquideas are believed to have very few spider monkeys because of indigenous hunting, and the density may be decreasing in Los Katios.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF327FA5DF9C56612FA15.taxon	vernacular_names	Black Spider Monkey	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF327FA5DF9C56612FA15.taxon	materials_examined	Peru. Restricted by R. Kellogg and E. Goldman in 1944 to the Rio Comberciato, affluent of the Rio Urubamba, Cuzco, Peru.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF327FA5DF9C56612FA15.taxon	discussion	Hybridization reportedly occurs with A. belzebuth in a restricted zone south of the Rio Maranon in Peru, and there is also a zone of apparent interbreeding with A. marginatus on the east bank of the upper Rio Tapajos, in Brazil. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF327FA5DF9C56612FA15.taxon	distribution	Distribution. Amazonian Brazil, S of the Rio Amazonas-Solimoes, W of the rios Tapajos-Teles Pires to the Rio Ucayali in NE Peru (where it is replaced by A. belzebuth on the left bank of the lower Ucayali); it crosses the middle Ucayali S of the Rio Cushabatay (a left bank tributary of the Ucayali), extending to the interfluvium of the rios Ucayali and Huallaga, then S along the Cordillera Oriental, and into N & C Bolivia S to ¢. 17 ° S, and from there, NE through Noel Kempff Mercado National Park.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF327FA5DF9C56612FA15.taxon	description	Descriptive notes. Head-body 45 - 60 cm (males) and 40 - 52 cm (females), tail 80 - 88 cm (males) and 70 - 80 cm (females); weight ¢. 7 kg (males) and c. 5 kg (females). The Black Spider Monkey is similar to the Red-faced Black Spider Monkey (A. paniscus) in overall appearance but slightly smaller and shorter-haired, with largely black facial skin. Older individuals have some depigmenation around the eyes. Fur of adults is entirely black, except for a silvery genital patch and, sometimes, a few white hairs on muzzle, cheeks, and forehead. Infant Black Spider Monkeys have whitish skin around their eyes, pink feet, and sparse fur that appears grayish.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF327FA5DF9C56612FA15.taxon	biology_ecology	Habitat. Primary lowland terra firma rainforest, lowland subtropical semi-deciduous forest, and riparian and flooded forest. In Noel Kempff Mercado National Park, Black Spider Monkeys are observed most often in tall forest but also in “ saternejal ” forest along the forest-savanna border, in the vicinity of small forest streams, that suffers periodic flash floods. They are found in transitional forest (forest to savanna) in southern Rondonia, Brazil. At Bolivia's Beni Biological Station and Biosphere Reserve, they are restricted to high forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF327FA5DF9C56612FA15.taxon	food_feeding	Food and Feeding. Black Spider Monkeys have been studied in Manu National Park and Biosphere Reserve, Lago Caiman in Noel Kempff Mercado National Park, and La Chonta and Guayaros forest reserves in Bolivia. At Lago Caiman, R. Wallace recorded a diet of 85 - 8 % fruit, 10 - 7 % flower buds and flowers, and the remainder items such as invertebrates, leaf galls, bark, and fungi. Seventy-eight plant species from c. 31 families made up the diet. Fruit made up more than 70 % of the diet in all but one ofthe eleven months of the study (63 % in July at the end of the dry season). Important species providing fruits included Ampelocera ruizii (Ulmaceae), Spondias mombin (Anacardiaceae), Ficus americana (Moraceae), two species of Brosimum (Moraceae), and the palm Euterpe precatoria. Black Spider Monkeys ate seeds of a few species in unripe fruits, particularly non-fleshy fruits, in the dry season — believed to be an important source of protein. In a year-long study by A. Felton and coworkers at La Chonta, fruits made up 82 % of the diet, leaves 13 % (buds and young leaves), and flowers 4: 7 %; the remainder was palm hearts, rotting wood, aerial roots, stalks, and bark. The plant part of the diet consisted of 63 species from 37 families; Moraceae was the most important in terms of number of species and time spent feeding (61: 2 % of the feeding records). They also ate caterpillars on leaves of Terminalia oblonga (Combretaceae). Fruits of Ficus boliviana and FE trigona, flowers and flower buds of Pseudolmedia laevis (all Moraceae), and fruit of a species of Myrciaria (Myrtaceae) comprised almost 50 % of the feeding records throughout the year. Lipid-rich fruits of Virola sebifera (Myristicaeae), although scarce, were highly favored. In 98 % ofall fruit feeding events, seeds were ingested whole and defecated; seeds of palm fruits were spat out. Fruit and figs comprised more than 70 % of the diet in all months except June (53 %), and they ate both ripe and unripe fruits. Ripe and unripe figs (unripe particularly from F boliviana rather than F trigona) were a staple item throughout the year. Black Spider Monkeys preferred ripe figs to other ripe fruit, and unripe figs were eaten even when ripe figs and fruit of other species were available. Unripe figs were apparently selected because they were nutritionally rewarding for their proteins and lipids. At Manu in the diverse forests of the western Amazon, the diet for three months in the dry season was ¢. 80 % fruit, c. 17 % leaves, c. 2 % flowers, and c. 1 % leaf petioles.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF327FA5DF9C56612FA15.taxon	breeding	Breeding. Births of Black Spider Monkeys at Manu occur throughout the year, but there is a peak in the early wet season. Infant survivorship up to one year old has been estimated at 67 %. Interbirth intervals are c ¢. 34 - 5 months. There is evidence that females are able to adjust the sex ratio before birth. Females have a dominance hierarchy that may inhibit the ability of other females to raise male offspring. High-ranking females tend to give birth to male infants more than do the lower ranking females, who produce more female offspring. Infanticide involving a subadult male killing a 4 - week old infant was recorded in the population in Manu. Infanticide may be an adaptive behavior of males to reduce the interbirth interval from 2 - 3 - 5 years to less than a year.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF327FA5DF9C56612FA15.taxon	activity	Activity patterns. Activity budgets of Black Spider Monkeys at Lago Caiman varied throughout the year, determined by changes in dispersion of fruiting trees and fruit abundance. In the dry season (April-August), resting occupied 45 - 50 % of the day, traveling 22 - 28 %, and feeding 17 - 24 %. In the wet season (September — March), Black Spider Monkeys rested less (37 - 42 %), traveled more (30 - 40 %), and spent about the same amount of time feeding, varying between months from ¢. 12 % to 23 %. Social and other activities were relatively constant at ¢. 5 %. In the dry season, they have smaller daily homes ranges, feeding more on leaves and fewer, less widely dispersed fruit sources. At Lago Caiman, Black Spider Monkeys typically move and feed in the upper forest canopy (91 % of all sightings), but they were often seen in the middle canopy and emergent trees. A similar activity budget was recorded for two groups of Black Spider Monkeys in Manu: resting 45 %, traveling 26 %, and feeding 29 %. At the Rio Jiparana in the state of Rondonia, Brazil, they spent ¢. 75 % of their time in the lower and middle canopy 10 - 20 m above the ground. About 14 % of the observations were in the top of the canopy at 2 - 25 m above the ground.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF327FA5DF9C56612FA15.taxon	biology_ecology	Movements, Home range and Social organization. Black Spider Monkeys have a fission-fusion social organization, with large groups (sometimes called communities) usually traveling and foraging in subgroups of females and their young, occasionally with a male or multiple males. All male groups are also observed. Group sizes are 37 - 55 individuals. A group of 55 individuals at Lago Caiman had 15 adult males and 15 adult females — an unusual sex ratio because most groups of Black Spider Monkey are skewed toward females. At Manu, two groups each had five males and 15 and 16 females. An average subgroup size of 3 - 3 individuals was recorded from a number of localities in Rondonia, and subgroup sizes at Manu average 3 - 1 individuals (range 2 - 9), along with solitary individuals. Lago Caiman has a variable subgroup size of c. 6 individuals. Females disperse, males are philopatric. Males associate more with other males in the group than with females, but males show more alloparental care than females. Males are territorial, and intergroup interactions occur at home range boundaries, marked by aggressive calling and “ ook-barking ” when two groups confront each other with 50 - 150 m between them. Sometimes males, with fur raised (piloerection), charge and chase each other. After one group retreats, ook-barking can continue for an hour or so. Home range sizes are 153 - 231 ha in Manu, 295 ha at Lago Caiman, and 340 ha at La Chonta Forest Reserve. Mean daily movements are 1977 m (465 - 4070 m) at Manu and 2338 m (460 - 5690 m) at Lago Caiman. Males travel farther during the day than females. Over eleven months, males in a group at Lago Caiman traveled an average of 2546 m / day and females 1972 m / day. They also traveled faster: males averaged 957 m / hour and females 778 m / hour. This difference was believed to result from home range border patrolling by males. Males tended to travel in larger subgroups (average 8 - 5 individuals) when near home range borders than when in more central parts of their home range (7 - 4 individuals). Adult females occupied, and preferentially used, a core area of the group’s home range. In Bolivia, densities of Black Spider Monkeys are usually 5 - 25 ind / km?. Densities in the Noel Kempff Mercado National Park (Lago Caiman) are as high as 32 ind / km? ®. Density of 31 ind / km? was recorded in terra firma lowland forest in Manu National Park and Biosphere Reserve, Peru. High densities are correlated positively with habitat heterogeneity. Densities in Brazil vary according to forest type: 3 - 1 - 9 - 6 ind / km? in terra firma forests and 2 - 6 - 3 - 6 ind / km? in varzea (white-water flooded forest). Surveys along the Rio Jurua found densities in non-hunted areas as high as 25 ind / km?, but at four hunted locations, Black Spider Monkeys were too rare to be counted at two and densities at the other two were only 0 - 6 - 1 - 3 ind / km?. Average density was 4 - 2 ind / km? * at eight non-hunted sites in the Amazon Basin and 0 - 7 ind / km? at ten moderately to heavily hunted sites.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A1CF327FA5DF9C56612FA15.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The [UCN Red List. The Black Spider Monkey is widespread and common where not hunted for its meat, but it has been extirpated or is threatened by hunting and habitat loss in many areas. It exhibits little or no adaptability to human pressures. Forests in the southern part of its distribution in the states of Rondonia, Mato Grosso, and Acre, in particular, are being devastated along the agricultural frontier moving northward through the Brazilian Amazon. Cattle ranching and forest loss are also widespread in northern Bolivia and south-eastern Peru. Additional threats include hunting and deforestation associated with highway development projects, agricultural expansion for soy production, mining activities in the Peruvian Amazon, and habitat degradation from selective logging, which may affect key fruiting species and forest structure and open up new areas for hunting. The Black Spider Monkey is known to occur in ten protected areas in Bolivia (Ambor 6 National Park, Beni Biological Station and Biosphere Reserve, Carrasco National Park, Isiboro Sécure National Park, Madidi National Park, Manuripi-Heath National Reserve, Noel Kempff Mercado National Park, Pilon Lajas Biosphere Reserve and Communal Lands, and Rios Blanco y Negro National Reserve); 14 in Brazil (Abufari Biological Reserve, Amazonia National Park, Cunia Ecological Station, Guaporé Biological Reserve, Iqué Ecological Station, Jara Biological Reserve, Jutai-Solimoes Ecological Reserve, Mamiraua Sustainable Development Reserve, Mapinguari National Park, Pacaas Novos National Park, Rio Acre Ecological Station, Samuel Ecological Station, Serra da Cutia National Park, and Serra do Divisor National Park); and three in Peru (Bahuaja-Sonene National Park, Manu National Park and Biosphere Reserve, Pacaya-Samiria National Reserve). It may also be found in the Tambopata National Reserve in Peru.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A03F326FA8AFA70631FF819.taxon	vernacular_names	Red-faced Black Spider Monkey	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A03F326FA8AFA70631FF819.taxon	materials_examined	South America, Brazil. Restricted by E. Geoffroy Saint-Hilaire in 1803 to French Guiana.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A03F326FA8AFA70631FF819.taxon	discussion	The border between this species and A. belzebuth is the Rio Branco in Roraima State. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A03F326FA8AFA70631FF819.taxon	distribution	Distribution. The Guianas (E of the Essequibo River, but excluding the lowland coastal plains) and NE Brazil (N of the Rio Amazonas and E of the rios Negro and Branco); its occurrence W of the Essequibo and in Venezuela is doubtful.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A03F326FA8AFA70631FF819.taxon	description	Descriptive notes. Head-body 51 - 5 - 58 cm (males) and 42 - 66 cm (females), tail 72 — 85 - 2 cm (males) and 64 - 93 cm (females); weight average 9 - 1 kg (males, n = 20) and 8 - 4 kg (females, n = 42). Male Red-faced Black Spider Monkeys are typically larger and more robust than females. Fur is long, silky, and glossy-black, although much sparser on the underside. The tail is thickly furred for two-thirds of its length and then tapering sharply toward its tip — a characteristic that distinguishes the Red-faced Black Spider Monkey from all other spider monkeys. Adults have a naked, pink or reddish face, with sometimes a few short, silvery or white hairs on the muzzle. Pointed tufts of hair protrude sideways from in front of each ear.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A03F326FA8AFA70631FF819.taxon	biology_ecology	Habitat. Primary high forest; rarely in edge or degraded forest and Euterpe palm swamp. In Suriname, the Red-faced Black Spider Monkey is almost entirely restricted to the high forests of the interior and enters the coastal plain only in the western part of the country, although historical accounts suggest that populations may have ranged closer to the coast. In Suriname, it rarely enters river edge forest, which may be a response to increased hunting pressure along waterways. In Guyana, it also occurs mainly in high forest. To what degree Red-faced Black Spider Monkeys extend to the coastal region of French Guiana today is unknown, although sightings in coastal forests were made in the early 1900 s. In Brazil, it occurs in lowland, submontane, and montane (Serra da Pacaraima) forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A03F326FA8AFA70631FF819.taxon	food_feeding	Food and Feeding. Like all spider monkeys, the Red-faced Black Spider Monkey is highly frugivorous and feeds largely on the mature, soft parts of a very wide variety of fruits, which comprise 80 % or more ofits diet and are found mainly in the emergent trees and upper part of the forest canopy. It also eats young leaves and flowers (especially at times offruit shortage in the early dry season), young and mature seeds, floral buds, pseudobulbs, aerial roots, bark, decaying wood, fungi, honey, and on occasion small insects such as termites and caterpillars, Many fruits, even quite large ones, are swallowed whole, but in other cases, the outer layer is bitten off, and the aril or mesocarp is swallowed along with the seed or seeds. Dueto its method of feeding and subsequent defecation (endozoochory), the Red-faced Black Spider Monkey has been found to be a significant seed disperser for nearly 140 tree and liana species. Seeds of an additional ten species are dispersed by the monkeys carrying them off some distance before dropping them. Seeds of only about two dozen food species are ruined when eaten (seed predation).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A03F326FA8AFA70631FF819.taxon	breeding	Breeding. Although young individuals can be observed throughout the year, the Redfaced Black Spider Monkey has a clearly defined birth season in Suriname. Newborn infants are seen only in November — January, indicating that they are born at the end of the long dry season and during the short wet season. Nevertheless, the few data available from other areas indicate lack of a distinct breeding season. Young resemble adults in coloration. Lifespan in the wild may exceed 30 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A03F326FA8AFA70631FF819.taxon	activity	Activity patterns. The Red-faced Black Spider Monkey is diurnal and arboreal, primarily an inhabitant of the upper forest canopy and emergent trees. It also enters the middle and lower canopy, but it is very rarely seen in the understory. The day typically includes two resting periods: 08: 00 - 10: 00 h and 12: 00 - 14: 30 h. More time is spent resting during the dry season when fruit is scarce. Feeding activity is highest for the first two hours of the morning and the last two hours before entering sleeping trees. Sleeping trees are alwaystall emergents, well free of a contiguous canopy, with a broad, open crown, either small-leaved, leafless, or in leaf flush, horizontally branched, and located strategically near feeding sites for the following days or, better still, in the next day’s feeding site.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A03F326FA8AFA70631FF819.taxon	biology_ecology	Movements, Home range and Social organization. Red-faced Black Spider Monkeyslive in groups of up to 20 - 30 individuals, but they are typically observed traveling, feeding, or resting in smaller subgroups and rarely seen all together. Within the home range, subgroups may travel 500 - 5000 m / day, depending on its size and composition, weather, season, and distribution of available food sources. During the dry season, individuals tend to travel less, especially when leaves become a more significant component of the diet. They appear to maintain discrete, non-overlapping home ranges of a few hundred hectares that are defended by males. In the Raleigh Falls-Voltzberg Reserve, Suriname, density of Red-faced Black Spider Monkeys was 8 - 2 ind / km?. Densities were 7 - 14 ind / km? at Nouragues, French Guiana and 2 - 4 - 6 - 2 ind / km? at three sites in Guyana.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A03F326FA8AFA70631FF819.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. The Red-faced Black Spider Monkey is widespread and common where not hunted for its meat, but locally extirpated or threatened by hunting and habitat loss in many areas. Habitat loss, although not as severe in the more northerly parts of its distribution, has been particularly notable in Brazil. Hunting remains a threat everywhere, especially French Guiana and even in protected areas. The Red-faced Black Spider Monkey is reported from or believed to occur in numerous and many large protected areas including Cabo Orange and Tumucumaque national parks in Brazil, Central Suriname Nature Reserve in Suriname, Kaw and Nouragues nature reserves and Guiana Amazonian Park in French Guiana, and Iwokrama Forest Reserve and Kaieteur National Park in Guyana.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F326FF57F8726B6EFABA.taxon	vernacular_names	White-whiskered Spider Monkey	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F326FF57F8726B6EFABA.taxon	materials_examined	Brazil, Rio de Janeiro. Restricted by R. Kellogg and E. Goldman in 1944 to Cameta, Rio Tocantins.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F326FF57F8726B6EFABA.taxon	discussion	The restriction to Cameta was evidently erroneous or perhaps based on a pet animal. There is no evidence for its occurrence east of the Rio Xingu. There is a zone of apparent interbreeding between this species and A. chamek on the east bank of the upper Rio Tapajos; such individuals are typical A. marginatus in their pelage characters. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F326FF57F8726B6EFABA.taxon	distribution	Distribution. Endemic to the Brazilian Amazon (states of Mato Grosso and Para), S of the lower Rio Amazonas between the rios Tapajos and Teles Pires (right banks) in the W and the Rio Xingu (left bank) in the E; the upper Rio Teles Pires and the Serra do Cachimbo mark the S limits to its distribution.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F326FF57F8726B6EFABA.taxon	description	Descriptive notes. Head-body 50 - 70 cm (males) and 35 - 58 cm (females), tail 75 - 90 cm (males) and 62 - 77 cm (females); weight c. 10 - 4 kg (males) and c. 5 - 8 kg (females). The White-whiskered Spider Monkeyis relatively small. Its fur is entirely black except for a triangular white patch on the forehead and whitish cheek whiskers. Juveniles have a very pale, flesh-colored face that becomes red in adults.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F326FF57F8726B6EFABA.taxon	biology_ecology	Habitat. Primary lowland rainforest, and riparian, marsh, semi-deciduous, and dry savanna forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F326FF57F8726B6EFABA.taxon	food_feeding	Food and Feeding. In Cristalino State Park, northern Mato Grosso, the White-whiskered Spider Monkey feeds mainly on ripe fruits and young leaves especially from species ofViolaceae, Fabaceae, and Rutaceae. They eat seeds of Brosimum guianense (Moraceae) and soil from termite nest tunnels.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F326FF57F8726B6EFABA.taxon	breeding	Breeding. No information is available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F326FF57F8726B6EFABA.taxon	activity	Activity patterns. Preliminary observations of White-whiskered Spider Monkeys in Cristalino State Park indicate an activity budget of ¢. 44 % traveling, 28 - 6 % resting, 12 - 2 % feeding, and 15 - 2 % engaged in other activities.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F326FF57F8726B6EFABA.taxon	biology_ecology	Movements, Home range and Social organization. The White-whiskered Spider Monkey is the least known of the Amazonian spider monkeys. A group of at least 24 individuals was seen in surveys in Tapajos National Forest. No estimates of density have been published.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F326FF57F8726B6EFABA.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The [UCN Red List. The small distribution of the White-whiskered Spider Monkey, the smallest of any of the Amazonian spider monkeys, is bisected by the Transamazon Highway along which there has been considerable settlement and by the Cuiaba-Santarém Highway. It is particularly susceptible to hunting. Accelerated expansion of the agricultural frontier, creating enormous areas of soy bean plantations that accompany the paving of the Cuiaba-Santarém Highway, is a major threat. Surveys are currently ongoing throughout it distribution. It is known to occur in five Brazilian national forests (Altamira, [taituba I, Itaituba II, Tapajos, and Xingu), offering some degree of protection. It also occurs in Upper Xingu National Indigenous Park and, reportedly, on a few private ranches. Surveys along the lower Rio Tapajos, including Tapajos National Forest, show that its forests are fragmented and degraded in many areas. It is not found in forest fragments of less than 100 ha, and it is extinct or at least extremely rare in the northern part of its distribution along the Tapajos, including the northern one-third of the national forest. In two sites (c. 3 ° 20 ’ S), it was found at relative abundances of 0 - 14 and 0 - 7 sightings / 10 km. It was also found in very low numbers along the southern edge of the national forest adjacent to the Transamazon Highway. Its absence from many areas is ascribed to selective logging (perhaps removing key tree species for the monkeys) and hunting. It occurs in the Cristalino State Park (184,900 ha) in northern Mato Grosso in the southern part of its range.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F325FA6EFAED6B56FDA9.taxon	vernacular_names	White-bellied Spider Monkey	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F325FA6EFAED6B56FDA9.taxon	materials_examined	type locality unknown. This species hybridizes with A. chamek in a restricted zone south of the Rio Maranon, between this river and the Rio Ucayali.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F325FA6EFAED6B56FDA9.taxon	discussion	Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F325FA6EFAED6B56FDA9.taxon	distribution	Distribution. Not well known, but includes S Venezuela (upper Orinoco Basin, limited by the Llanos / forest interface to the W and by the left bank of the Rio Caura in the E), NW Brazil (N of the Rio Solimoes as far as the mouth of the Rio Japura, E to the Rio Negro, throughout the region N of the Rio Negro and W of the Rio Branco in Amazonas and Roraimastates), E Colombia (in the piedmont of the Cordillera Oriental as far N as the Rio Upia drainage in Boyaca Department, including the region of Sierra de la Macarena, E Meta Department, and S to the Coehmani rapids of the Rio Caqueta in the SE of Caqueta Department; it is rare W of the Rio Yari and has been seen on the upper Rio Mesay, opposite the mouth of the Rio Pira-Parana on the right bank of the Rio Apaporis, and the lower right bank of the Apaporis), E Ecuador and NE Peru (restricted to the S of the Rio Napo, extending into N Peru between the rios Putumayo and Amazonas, along the left bank of the Rio Ucayali, and including the basins of the rios Napo, Tigre, Maranén, Pastaza, Pacaya, and Samiria, S as far as the Rio Cushabatay, a tributary of the Rio Ucayali).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F325FA6EFAED6B56FDA9.taxon	description	Descriptive notes. Head-body 46 - 50 cm, tail 74 - 81 cm; weight 8 - 3 kg (males, n = 10) and 7 - 9 kg (females, n = 16). The White-bellied Spider Monkey is blackish with a contrasting yellowish-white underside, inner surfaces of limbs, backs of thighs, and underside of the tail. The face is hairless and usually dark, and there is a triangular white or golden patch on the forehead in ¢. 33 % of the population. Infant White-bellied Spider Monkeys have a reddish face and dark fur.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F325FA6EFAED6B56FDA9.taxon	biology_ecology	Habitat. Primary montane tropical, subtropical forest, lowland riparian, marsh, and semi-deciduous forest. The White-bellied Spider Monkey is dependent on high primary forest and is generally associated with terra firma clay soils.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F325FA6EFAED6B56FDA9.taxon	food_feeding	Food and Feeding. Feeding ecology of White-bellied Spider Monkeys has been studied at a number of sites in northern Brazil (Ilha de Maraca, Rio Uraricoera), Ecuador (Yasuni National Park), Colombia (Tinigua and Sierra de la Macarena national natural parks), and southern Venezuela (Rio Tawadu, a tributary of the Rio Nichare in the El Caura Forest Reserve). Like all spider monkeys, the White-bellied Spider Monkey is highly frugivorous, feeding largely on the mature soft parts of a wide variety offruits. In the extremely botanically diverse forest in the western Amazon, at Yasuni, they eat fruits of more than 238 species. At Maraca Ecological Station, they fed in more than 468 trees in one year, with just a few (28) contributing more than 1 % the feeding records. In these studies, average percentages of fruits (largely ripe fruit) in the diet were 73 - 92 %. Diets are supplemented with mostly young leaves from trees and lianas (particularly in the dry season), flowers, seeds, aerial roots, animal prey (either caterpillars or insects ingested incidentally with fruits), and rotting wood. They occasionally eat soil (geophagy) from termitaria and the ground at mineral licks where they also drink the slightly salty water. Fruits are seasonally abundant; many trees and lianas fruit in the early to middle wet season, and fruits are most scarce in the late wet and early dry season. At Maraca, White-bellied Spider Monkeys travel less in the dry season (October-April), feeding on a relatively small number oftrees for prolonged periods. In the wet season (May-September), with numerous trees in fruit, they range over a larger area, traveling farther and feeding in shorter bouts in a large number oftrees.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F325FA6EFAED6B56FDA9.taxon	breeding	Breeding. Births of White-bellied Spider Monkeys appear to be spread throughout the year, which seems to be typical for this genus. Single births are the rule, but twins occasionally occur. A female at Sierra de la Macarena that gave birth to twins was monitored to see how she and her infants fared. She evidently struggled, resting more and feeding and traveling less than females with single infants or no infants. She relied more on flowers and leaves when others were relying on fruits. The twins grew more slowly than a singleton would have, were slower in their social development, and were small for their age. Individuals may live 20 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F325FA6EFAED6B56FDA9.taxon	activity	Activity patterns. Activity budgets of White-bellied Spider Monkeys are determined in large part by dispersion, seasonal abundance, and quality of foods in each forest. Activity budgets at Maraca Ecological Station, Yasuni, and Sierra de la Macarena are resting 45 - 61 %, feeding 17 - 22 %, traveling 10 - 25 %, and social and other behaviors 1 - 3 %. At Maraca, White-bellied Spider Monkeys rested less in the wet season (64 % in the dry month of November and 32 % in July). They traveled more and spent more time feeding (average 22 - 5 % of the day) in the wet season than in the dry season (12 %). This might have been related to the fact that they eat less fruit (slow to harvest) and more leaves (quicker to harvest) in the dry season. At Sierra de la Macarena, they spend c. 70 % oftheir time feeding and traveling in the lower and middle canopy at 12 - 24 m and c. 10 % resting and interacting socially in the upper forest canopy at 24 - 30 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F325FA6EFAED6B56FDA9.taxon	biology_ecology	Movements, Home range and Social organization. Social organization of White-bellied Spider Monkeys has been studied at several sites in Colombia, Brazil, and Ecuador. Group sizes are 15 - 30 individuals, typically with 3 - 6 adult males and 5 - 11 adult females. They generally travel and feed in separate subgroups centered on matrilines of each adult female in the subgroup. Malesjoin these subgroups or travel separately, alone or in small groups. Subgroups are fluid in their composition and generally have 2 - 5 individuals. Sizes of subgroups tend to be larger when fruits are most abundant. Group members come together infrequently. When visiting salt lick sites in Sierra de la Macarena National Natural Park, 1 - 2 individuals at a time go to the ground to lick soil and drink, while others remain in the trees. It is thought that those in the trees are being vigilant for predators because salt licks are frequently visited by peccaries (Pecari and Tayassu), tapirs (Tapirus), brocket deer (Mazama), and agoutis (Dasyprocta) and are haunts for Jaguars (Panthera onca) and Ocelots (Leopardus pardalis) as a result. At other times, the entire group (or at least larger numbers) is seen together when noisily and excitedly confronting neighboring groups. Such intergroup encounters occur at salt licks when they are near the borders of home ranges. The home range size of a group of 19 - 23 individuals at Maraca was 316 ha. Home range sizes at Sierra de la Macarena vary among different groups and can be as large 388 ha. At Maraca, daily movements during a year averaged 1750 m; they were shorter in the dry season (average 984 m) and longer in the wet season (2160 m). At Yasuni, daily movements varied from 723 m to a record 6039 m (annual average 3311 m). At Sierra de la Macarena and Maraca, male White-bellied Spider Monkeys move more widely within a group’s home range than do females. At Maraca, mean home range size was 140 ha for any particular male in the group and 87 - 5 ha for females. A group of 30, with five adult males and ten adult females, at Sierra de la Macarena occupied a home range of 163 ha; males had home ranges of 83 - 93 ha and females 55 - 66 ha. Home ranges of males overlapped widely (71 - 87 %), whereas home ranges of females overlapped less (39 - 64 %). Males travel more along boundaries of their home ranges, and travel faster and farther when doing so, indicating that they might be monitoring boundaries and also females in their separate home ranges. Males travel in slightly larger subgroups when they are on the home range border. The difference in the home range sizes of the two sexes are related to differences in diet. Males eat more (widely dispersed) fruit, and females eat more leaves (in smaller home ranges and perhaps of necessity requiring a higher protein diet). At Yasuni, a group of 22 - 17 individuals occupied a home range of 314 ha, and curiously, sizes of individual home ranges of males and females did not differ; both were large at 152 ha for males and 149 ha for females. Females did not appear to have semi-exclusive homes ranges, as has been found in the otherstudies; overlap for both sexes was 80 %. Males did not use the boundary area of the home range more than females. Densities of 15 - 18 ind / km? * were estimated in the Sierra de la Macarena. Researchers at Sierra de la Macarena witnessed predation of an adult male White-bellied Spider Monkey by a Jaguar and a failed attack on an adult female by a Puma (Puma concolor).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A02F325FA6EFAED6B56FDA9.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The [UCN Red List. The White-bellied Spider Monkey is threatened by deforestation and hunting for its meat, to the point of local extirpation. In south-eastern Colombia, habitat is lost due to forest clearance for coca plantations, accompanied by fumigation that results in defoliation of contiguous forests. In Peru, the White-bellied Spider Monkey has been extirpated over large areas, and remnant populations are subjected to strong hunting pressure in some regions. In Ecuador, it is rare near human settlements, being hunted for food and in demand for pets, but it can be common in undisturbed forests. In the Colombian Amazon, the White-bellied Spider Monkey is considered the most endangered primate, its distribution coinciding with the most heavily colonized parts of the region, yet it remains fairly common within protected areas. It occurs in Pico da Neblina and Serra da Mocidade national parks and a number of ecologicalstations in Brazil; Amacayacu, Cahuinari, Cordillera de los Picachos, Cueva de los Guacharos, La Paya, Sierra de la Macarena, Tinigua, and Serrania de Chiribiquete national natural parks in Colombia; Podocarpus, Sumaco-Napo Galeras, and Yasuni national parks and Cayambe-Coca and Cofan-Bermejo ecological reserves in Ecuador; Pacaya-Samiria National Reserve in Peru; and Duida-Marahuaca, Jaua-Sarisarinama, Parima Tapirapeco, Serrania de la Neblina, and Yapacana national parks in Venezuela.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A01F324FA8CFDDC69FBFCCA.taxon	vernacular_names	Variegated Spider Monkey	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A01F324FA8CFDDC69FBFCCA.taxon	materials_examined	valley of the Rio Magdalena, Colombia. Restricted by R. Kellogg and E. Goldman in 1944 to La Gloria, Rio Magdalena, elevation 45 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A01F324FA8CFDDC69FBFCCA.taxon	discussion	Two subspecies are recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A01F324FA8CFDDC69FBFCCA.taxon	distribution	Subspecies and Distribution. A. h. hybridus 1. Geoffroy Saint-Hilaire, 1829 — N Colombia (from the right bank of the Rio Magdalena, N to the S slopes of the Sierra Nevada de Santa Marta and the N of the Serrania de Perija, and in the middle Rio Magdalena Valley at least to the departments of Caldas and Cundinamarca; two populations on the slopes of the Cordillera Oriental on the Venezuelan border (one in the Rio Catatumbo watershed in the Norte de Santander Department and the other in the NE piedmont forest in Arauca Department) and N & NW Venezuela (along the SE part of the Cordillera de la Costa in the states of Miranda and probably Vargas, and on both sides of the Venezuelan Andesin the states of Zulia, Tachira, Mérida, Trujillo, Portuguesa, Apure, and Barinas, in the E in the piedmont and lowland forests of San Camilo and Ticoporo, and in the Win the piedmont forest around S Lake Maracaibo to the Sierra de Perija). A. h. brunneus Gray, 1871 — N Colombia in the departments of Antioquia, Bolivar, and Caldas, between the lower rios Cauca and Magdalena.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A01F324FA8CFDDC69FBFCCA.taxon	description	Descriptive notes. Head-body 47 - 50 cm (males) and 45 - 48 cm (females), tail 76 — 81 cm (males) and 74 - 76 cm (females); weight 7 - 9 - 8 - 6 kg, (males, n = 2) and 7 - 5 - 10 - 5 kg (females, n = 7). The Variegated Spider Monkey is pale wood-brown above, with a darker brown head, and whitish or buffy below, including inner sides of limbs and ventral surface of the tail. It has a distinct white triangular patch on the forehead. Some individuals have grayish-blue eyes. In the nominate subspecies, hindlimbs are usually lighter and similar in color to the back, but knees tend to be darker. The belly varies from white with brownish shades to a light yellow. Ventral hair ofthe tail ranges from yellowish to buffy light brown. In the “ Brown Spider Monkey ” (A. h. brunneus), exterior color of limbs, back, and feet is dark brown. The belly varies from whitish to yellowish, contrasting sharply with the brownish sides, tail, and head.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A01F324FA8CFDDC69FBFCCA.taxon	biology_ecology	Habitat. Evergreen, semi-deciduous, riparian, and montane tropical forests at elevations of 20 - 700 m (nominate subspecies in Venezuela).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A01F324FA8CFDDC69FBFCCA.taxon	food_feeding	Food and Feeding. The diet of the Variegated Spider Monkey in riparian forest fragment at the Hacienda San Juan de Carare on the east bank of the Rio Magdalena, Colombia, was unusual in its relative lack offruit. Averaging observations of males and females, time spent feeding on different items was 48 % ripe fruits, 41 - 42 % leaves, 45 % flowers, 2 % decayed wood, and 4 - 5 % termite soil, seeds and other items. This diet is probably nottypical but a result of a 5 - year isolation in a forest fragment. Geophagy during frequent visits to a mineral lick has been reported for a group of Variegated Spider Monkeys in El Paujil Nature Reserve in the Serrania de las Quinchas. Camera trapping recorded visits of 1 - 29 individuals at the lick on 81 of 379 days. A list of potential feeding trees for the Variegated Spider Monkey in the region ofVenezuela's El Avila National Park was compiled, but no formalstudy ofits feeding ecology was carried out there.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A01F324FA8CFDDC69FBFCCA.taxon	breeding	Breeding. Newborn Variegated Spider Monkeys were observed in August in the Magdalena Valley, but no formal study of reproductive behavior has been done.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A01F324FA8CFDDC69FBFCCA.taxon	activity	Activity patterns. Average activity budgets of 9 - 11 Variegated Spider Monkeys (males and females combined) at the Hacienda San Juan de Carare were c. 41 % resting, ¢. 29 % feeding, c. 25 % of the day traveling, and ¢. 5 % engaged in social and other activities.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A01F324FA8CFDDC69FBFCCA.taxon	biology_ecology	Movements, Home range and Social organization. Subgroupsizes ofVariegated Spider Monkeys averaged 4 - 5 individuals in Serrania de San Lucas in southern Bolivar. Median subgroup size at Hacienda San Juan de Carare during one year was three adults, and subgroups of less than four individuals accounted for 73 % of the observations. Males were more aggressive to females than to males. Males approached females more often but were more aggressive to them. This aggression was offset to some extent by females grooming the males. Aggression between females was uncommon, perhaps due to the relative lack of fruit and the amount ofleaves in the diet reducing competition. Interspecific infanticide and aggression toward infants were reported atthis site, with male Variegated Spider Monkeys attacking infant Colombian Red Howlers (Alouatta seniculus) and infant Humboldt’s White-fronted Capuchins (Cebus albifrons). Two infanticides and all but one of seven attacks on infants were perpetrated by a subadult male. The single attack on an infant Humboldt’s White-fronted Capuchin was carried out by two adult male Variegated Spider Monkeys. These primates were restricted to a 70 - ha forest fragment, and although the reasons for this behavior were unclear, it might have been related to food competition, a pathology, or redirected aggression. Densities of Variegated Spider Monkeys were 9 - 14 ind / km? in Serrania de San Lucas and 8 - 2 - 9 - 6 ind / km? at Cerro Bran, near Porto Rico, Bolivar.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A01F324FA8CFDDC69FBFCCA.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Critically Endangered on The IUCN Red List, including both subspecies. Principal threats to the nominate subspecies of the Variegated Spider Monkey in Colombia and Venezuela are habitat destruction and hunting for food. Cattle ranching has significantly reduced the extent of lowland forest in Zulia and in piedmont forests of the Serra da Perija where heavy hunting also occurs. In central Venezuela, large expanses once covered by primary forest are now covered by secondary vegetation, resulting in the apparent extirpation of the Variegated Spider Monkey in this region by 2001. Unfortunately, buffer zones surrounding protected areas with known populations of the nominate subspecies (e. g. Guatopo National Park) continue to be cleared for slash-and-burn agriculture and human settlements. Logging also threatens lowland forests of the eastern Andes, San Camilo, and Ticoporo. The Brown Spider Monkey has a small geographic range in a region where forest loss, degradation, and fragmentation are widespread. Remaining populations in areas of Antioquia (e. g. Segovia, Remedios, Maceo, Yondo, Puerto Berrio, and Zaragoza) are surrounded by human settlements, with less than 10 % of the original habitat remaining. As a result, the viability of these populations of Brown Spider Monkeysis questionable, and local extirpations are highly probable in the absence of protection. Potential sites for protected areas remain in the Serrania de San Lucas and parts of Nechi in the south of Bolivar and the north of Antioquia departments. The nominate subspecies occurs in Sierra Nevada de Santa Marta, Sierra Nevada del Cocuy, Catatumbo-Bari, and Tama national natural parks in Colombia and Guatopo, El Avila, and Sierra de Perija national parks and Caparo Forest Reserve in Venezuela. It may also occur in Colombia’s Chingaza National Natural Park. Brown Spider Monkeys are found in the proposed Serrania de San Lucas National Natural Park, and in Ciénagas de Barbacoas which is a priority for the creation of a protected area.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A05F320FF6EFE27632AFDA1.taxon	vernacular_names	Humboldt’s Woolly Monkey	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A05F320FF6EFE27632AFDA1.taxon	materials_examined	Colombia, Rio Guaviare, above the mouth of the Rio Amanaveni, Guainia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A05F320FF6EFE27632AFDA1.taxon	discussion	Although the genus was revised by C. P. Groves in 2001 to include four species (L. lagothricha, L. lugens, L. poeppiggii, and L. cana) rather than the one species with four subspecies as earlier defined by] J. Fooden in 1963, Colombian studies of phenotype, karyotypes, and the mitochondrial COII gene strongly suggest that at least lagothricha and lugens belong to the same species and should be considered subspecies of L. lagothricha. The subspecies lagothricha appears to intergrade with lugens west of the Rio Caguan in the Caqueta Department, Colombia. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A05F320FF6EFE27632AFDA1.taxon	distribution	Subspecies and Distribution. L. lL. lagothricha Humboldt, 1812 — SE Colombia, NE Ecuador, NW Brazil, and N Peru, from the Cordillera Oriental, N of the rios Napo and Amazonas, to the Rio Uva, a N bank affluent of the Rio Guaviare (Vichada Department, Colombia), E to the Rio Orinoco and upper Rio Negro to the Venezuelan border and NW Brazil; it is not known if it occurs in Venezuela. L. [. lugens Elliot, 1907 — Colombia, E of the Rio Atrato, throughout the Cordillera Oriental, W of the Rio Ariari (Meta Department), and parts of the Cordillera Central to the northernmost parts of the Serrania de San Lucas (SE Bolivar & N Antioquia departments), in some places up to elevations of 3000 m, and northward in the piedmont forest (now mostly destroyed) to the Rio Arauca (Arauca Department); there is no evidence thatits occurs in neighboring SW Venezuela.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A05F320FF6EFE27632AFDA1.taxon	description	Descriptive notes. Head-body 46 - 65 cm (males) and 46 - 58 cm (females), tail 53 — 77 cm (males) and 62 - 72 cm (females); weight of males ¢. 7 kg (but when dominant they become more robust and can weigh as much as 10 kg) and of females 5 - 7 kg. Humboldt’s Woolly Monkey is quite variable in color, from very light blond (locally known as “ churucos blancos ” or white woollies) in the Amazon Basin to a very uniform dark brown (almost black) in the Cordillera Central. Generally, individuals are some shade of brown above, sometimes with a slightly paler head, dark gray hands and feet, and often ajet-black underside of long chest hair in adults. The area between shoulder blades can be paler than shoulders, and there is occasionally some ticking on lower legs, arms, and tail. There may be a coronalstripe of lighter tone or even blond on the head in both subspecies. In the Amazon, some individuals have blond tufts around the base of the tail. Males are larger than females, with longer canines, and well-developed masseter muscles on either side of the head. The clitoris is as long as or longer than the penis, and adult testicles are very prominent. The original description of the two subspecies is generally inadequate to distinguish them because of variability of tones and overlap of characteristics purported to belong to each. Pelage morphs of the “ Brown Woolly Monkey ” (L. l. lagothricha) in the lowlands are actually a mosaic (not a cline) of different morphs that vary from very light blond to very dark brown with many individuals in between the two extremes. Generally, the “ Colombian Woolly Monkey ” (L. I. lugens) tends to have darker tones of brown than are typical of the Brown Woolly Monkey, culminating in the holotypic charcoal animals of the Cordillera Central of the Andes. Many individuals of both subspecies have ticked or agouti forearms and forelegs, long black chest hair (in adults), and coronalstripes.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A05F320FF6EFE27632AFDA1.taxon	biology_ecology	Habitat. Primary, secondary, flooded, and non-flooded lowland rainforest and primary and secondary montane forest up to elevations of 3000 m. Humboldt’s Woolly Monkeys use the middle and upper forest canopy. They can be found in wide (several kilometers) Colombian piedmont gallery forests, but most of these populations have been extirpated in what are now destroyed and fragmented forests.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A05F320FF6EFE27632AFDA1.taxon	food_feeding	Food and Feeding. Humboldt’ ' s Woolly Monkeys feed mainly on ripe, fleshy, endozoochorous fruits, supplemented with young leaves, flowers, gums, seeds, palm nuts, and invertebrate prey. They exploit well over 200 species of fruits during the year; 185 species were tabulated in a study in eastern Colombia, while more than 220 species were identified in diets in forests west of the Rio Duda, western Colombian Amazon. In the study in eastern Colombia where soils are very poor, the diet consists of 78 - 9 % fleshy fruits, 11: 4 % leaves, 4 - 9 % invertebrates, 4 - 3 % seeds, 0 - 3 % bark, 0 - 1 % flowers, and 0 - 1 % tendrils. The most important species were Chrysophyllum amazonicum (Sapotaceae) at 10 - 3 % of the fruit-feeding records, Manilkara amazonica (Sapotaceae) at 5 %, Iriartea ventricosa (Arecaceae) at 2: 2 %, Cheiloclinium sp. (Celastraceae) at 22 %, and Protium sp. (Burseraceae) at 2 %. The most important families were Sapotaceae, Moraceae, and Fabaceae, followed by 38 other families. At the Rio Duda site, the most interesting differences were that Moraceae was the most important family rather than Sapotaceae (ninth in importance) and the consumption of invertebrates (12 %) was much higher at Rio Duda. These differences might be explained by highersoilfertility, because species of Moraceae seems to increase in importance on better soils and the availability of invertebrates might increase where better soils support increased primary production. Another explanation for higher invertebrate consumption could be differences in data collection techniques. Humboldt’s Woolly Monkeys tend to spend the most time feeding on sweet and succulent fruits from abundant plants that produce large fruit crops. During fruit scarcity, they prefer large trees that fruit during times of scarcity and that have fruits with a high pulp-to-seed ratio, low lipid content, and clumped distributions. Results from one study suggest that Humboldt’s Woolly Monkeys are generalized, opportunistic frugivores that avoid some fruits with secondary compounds.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A05F320FF6EFE27632AFDA1.taxon	breeding	Breeding. Female Humboldt’s Woolly Monkeys are sexually mature at c. 6 years of age. They generally leave their natal group for another group at this time, although some evidence suggests that dominant females remain in their natal group. Mating begins soon after a female enters a new group. Males sometimes approach and follow receptive females, but females frequently follow males and solicit mating. Mating occurs throughout the year, resulting in births year-round, but with a peak when food is plentiful during the early rainy season (March — May) in the eastern Colombian Amazon and late rainy season to early dry season (July-December) in the western Colombian Amazon. All sexually mature males have access to the group’s females and copulate with them, but the number of copulations is positively correlated with the linear hierarchy among males. During copulation, both sexes give a special vocalization (click or “ tooth chatter ”), a rhythmical movement of the teeth with maximal exposure. Copulation is dorsal ventral, with the male behind the female and the female reclining on the branch. Copulation lasts 6 - 7 minutes. All adult males may mate with a receptive female, but the dominant male copulates most. Dependent young climb over the copulating individuals and look at the pair’s genitalia. Females emigrate from their group to a neighboring group at c. 6 years of age and usually begin copulating soon after. There are very low levels of aggression toward new females, perhaps because they are quickly accepted by adult males. A single offspring is born after a gestation of 223 - 225 days. In a twelve-year study, females were found to have a 23 - 36 - day reproductive cycle, with a birth interval averaging 36 - 7 months. Following birth, the female has a period of sexual inactivity averaging 23 - 4 months, but sexual activity, when reassumed, continues for an average of 7 - 2 months before she conceives again. Copulation can occur at any time, but there are times when females copulate more frequently; this is in December — May in the western Colombian Amazon. During copulation periods, cycles last 2: 3 - 3 - 1 days (when copulation is frequent) interspersed by 11 - 3 - 14 - 7 - day intervals with no mating. After conception, copulations become non-cyclic. Births are usually singleton, occasionally twins. Neonates are nursed for up to a year. They are carried ventrally at first but move to the mothers’ back after c. 2 weeks. Adult males sometimes participate, carrying an infant on their back and actively guarding it. Both sexes are fully mature at c. 8 years, but the first parturition is usually at c. 9 years. Humboldt’s Woolly Monkeys may live for more than 30 years; the population under study at the Rio Duda site in Colombia includes a 31 - year-old individual.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A05F320FF6EFE27632AFDA1.taxon	activity	Activity patterns. Humboldt’s Woolly Monkeys are diurnal and arboreal, spending most of their time in the middle and upper forest canopy. Activity budgets calculated in two studies were resting 29 - 9 % and 23 - 2 %, moving 38 - 8 % and 34 - 5 %, foraging 25 - 8 % and 36 - 2 %), and other activities 5 - 5 and 6 - 1 % (the first values from a population in a very infertile, poor forest and the second from a population in a rich, fertile forest closer to the Andes). The woolly monkeys evidently were able to travel and forage less and rest more in rich forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A05F320FF6EFE27632AFDA1.taxon	biology_ecology	Movements, Home range and Social organization. Humboldt’s Woolly Monkeys live in patrilineal social groups; only females move between groups, while males are philopatric. Depending on the availability of ripe fruit, a group moves 1000 - 3500 m / day and occupies home ranges of 169 - 800 ha. Average daily movement for a group of 24 individuals in eastern Colombia was 2880 m in a home range of 760 ha. A group of 17 individuals in the Rio Dudasite, western Colombian Amazon, traveled 1633 m / day and occupied a smaller home range of 169 ha. Differences in home range size and daily movement of the two groups were probably due to differences in the quality of the two forests where they were found.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A05F320FF6EFE27632AFDA1.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List, only as the Humboldt’s Woolly Monkey. The Colombian Woolly Monkey is classified as a distinct species (L. lugens) and Critically Endangered on The IUCN Red List. Humboldt’s Woolly Monkeys are intensively hunted, especially by indigenous people, because their meatis considered delicious and their fat is used in cooking and for supposed medicinal purposes. They are also frequently kept as pets, with the mother typically being shot to obtain the infant. Humboldt’s Woolly Monkeys are known to occur in Pico da Neblina National Park and Juami-Japura Ecological Reserve in Brazil; Amacayacu, Cahuinari, La Paya, and Serrania de Chiribiquete national natural parks and Nukak National Natural Reserve in Colombia; and Sumaco-Napo Galeras National Park, Cayambe-Coca and Cofan-Bermejo ecological reserves, and Cuyabeno Wildlife Reserve in Ecuador. Nevertheless, at least in Brazil and Colombia, indigenous peoples have the right to hunt them even in protected areas. In Colombia, presence of any indigenous settlement eventually results in extirpation of Humboldt’s Woolly Monkeys in a radius of several kilometers.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F320FF42FDC46608FD8B.taxon	vernacular_names	Gray Woolly Monkey	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F320FF42FDC46608FD8B.taxon	materials_examined	Brazil. Restricted by |. Fooden in 1963 to the south bank of Rio Solimoes near the mouth of Rio Tefe.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F320FF42FDC46608FD8B.taxon	discussion	A. Humboldt has been credited at times with the name ofthis species, but E. Geoffroy Saint-Hilaire is the rightful authority. Humboldt in 1812 (dated 1811 but actually published a year later) credited Geoffroy Saint-Hilaire for his 1812 publication in Tome 19 of Annales du Muséum d Histoire Naturelle, where this species’ name first appeared. Lagothrix cana was earlier considered a subspecies of L. lagothricha, along with the subspecies lugens and poeppigii. Because of clear phenotypic differences between the four, C. P. Groves in 2001 regarded them as separate species. The forms lagothricha and lugens are in fact not clearly different and not easy to distinguish (very variable), but poeppigii and cana do have a distinctive phenotype. So in this treatment, L. cana is considered a [phylogenetic] species, although new molecular evidence shows similar genetic markers for all taxa mentioned. A newly discovered population in Madidi National Park, northern Bolivia, may be a distinct taxon, or an isolated population of tschudii. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F320FF42FDC46608FD8B.taxon	distribution	Subspecies and Distribution. L. c. cana E. Geoffroy Saint-Hilaire, 1812 — Brazilian Amazon, S of the Rio Amazonas-Solimoes, between the rios Jurua and Tapajos-Juruena, but restricted to the left bank of the Rio Madeira above the mouth of the Rio Jiparana (a right bank tributary), and to the right bank of the Rio Jiparana, S to ¢. 12 ° S, and W along the N (left bank) of the Rio Abuna on the upper Rio Madeira. L. c. tschudii Pucheran, 1857 — SE Peru, its range is poorly known butis believed to extend between the rios Pachitea and Ucayali, S from the Rio Inuya, W as far as the Rio Inambari, and S to both sides of the Rio Madre de Dios into the Rio Tambopata Basin, to the frontier with Bolivia, with an isolated population in Madidi National Park.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F320FF42FDC46608FD8B.taxon	description	Descriptive notes. Head-body 50 - 65 cm (males) and 45 - 58 cm (females), tail 62 — 80 cm (males) and 53 - 72 cm (females); weight of males c. 7 kg (but when dominant they become more robust and can weigh as much as 10 kg) and of females 5 - 7 kg. The Gray Woolly Monkey is generally grayer than Humboldt’s Woolly Monkey (L. lagothricha), with the pelage ticked all over on the upper side. Males are larger than females, with relatively shortertails, longer canines, and well-developed masseter muscles on either side of the head. Clitoris is as long as or longer than penis, and testicles of adult males are clearly evident. “ Geoffroy’s Woolly Monkey ” (L. ¢. cana) is generally grayish-brown, with a contrastingly dark gray head. Hands, feet, and tail are a darker brown, and the underside is blackish-gray, often with a dark reddish tinge. Individuals from lowland areas tend to be pale fawn-gray with darker extremities. The “ Peruvian Woolly Monkey ” (L. c. tschudii) is very dark (deep blackish-gray) with a tinge of red. Head, limbs, and tail are black.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F320FF42FDC46608FD8B.taxon	biology_ecology	Habitat. Primary and some secondary lowland and highland rainforest in upland and flooded forests. The Gray Woolly Monkey prefers terra firma forest and uses the middle and upper forest canopy.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F320FF42FDC46608FD8B.taxon	food_feeding	Food and Feeding. Gray Woolly Monkeys are principally frugivores. In the plant part of their diet, they eat ripe and unripe fruits, seeds and gums (80 - 7 %), young leaves (14 - 4 %), flowers (3 - 1 %), petioles (17 %), and mature leaves (0 - 1 %). They also occasionally eat insects. At one Amazonian site, Gray Woolly Monkeys included more than 225 species of plants in their diets. Species of Moraceae, Sapotaceae, and Fabaceae accounted for 43 % of their food species and 63 % of their time spent feeding. Depending on food availability, groups sometimes split into feeding parties; they average 3 - 5 animals, but range from one to 22. Groups are more cohesive when fruit is more abundant.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F320FF42FDC46608FD8B.taxon	breeding	Breeding. No specific information is available for this species, but aspects of breeding are no doubt very similar to those of Humboldt’s Woolly Monkey.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F320FF42FDC46608FD8B.taxon	activity	Activity patterns. No specific information is available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F320FF42FDC46608FD8B.taxon	biology_ecology	Movements, Home range and Social organization. One study recorded a home range of more than 935 ha for a group of 39 - 41 individuals (excluding dependent infants). The study lasted only eight months and the home range was undoubtedly larger, possibly 1200 ha. Larger groups have been seen, although it is possible that they are two groups traveling in tandem, something which has been documented for Humboldt’s Woolly Monkey.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F320FF42FDC46608FD8B.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List, including Geoffroy’s Woolly Monkey. The Peruvian Woolly Monkey is classified as Data Deficient on The IUCN Red List. Gray Woolly Monkeys are intensively hunted, their meat is considered delicious, and their fat is used in cooking and for medicinal purposes. They are also frequently kept as pets, with the mothertypically being shot to obtain the infant. Geoffroy’s Woolly Monkey occurs in numerous protected areas in Brazil, including Amazonia and Pacaas Novos national parks. The Peruvian Woolly Monkey occurs in Manu National Park and Biosphere Reserve and possibly in Tambopata National Reserve, in Peru, and Madidi National Park in Bolivia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F32FFA59FDFD60A8F8C6.taxon	vernacular_names	Poeppig’s Woolly Monkey	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F32FFA59FDFD60A8F8C6.taxon	materials_examined	Brazil, banks of the Rio Maranon. Restricted by A. Cabrera in 1957 to Peru, lower Rio Huallaga, north of Yurimaguas, Loreto.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F32FFA59FDFD60A8F8C6.taxon	discussion	The revision of the genus Lagothrix by] J. Fooden in 1963 placed L. poeppigii as a subspecies of L. lagothricha (along with lugens and cana). Because of its clear phenotypic difference, poeppigii was regarded as a distinct species by C. P. Groves in 2001. An orange-colored woolly monkey was depicted in E. Cruz Lima’s 1945 book Mammals of Amazonia, (plate XVI). It was based on a captive juvenile of unknown provenance. The wild population was discovered by C. Peres in 2003, west of the town of Eirunepé on the upper Rio Jutaizinho, the west bank of the Rio Jurua in the south-western Brazilian Amazon. An extremely light-colored variant is pale ocherous-blond, with the color of the head being distinctly lighter than that of the back. The arms, legs, hands, feet, tail tip, and ventral surface are ocherous-white. The facial skin is pitch black, and the muzzle and chin are bordered with white whiskers. Both forms remain to be officially described and may in fact represent distinct subspecies or species. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F32FFA59FDFD60A8F8C6.taxon	distribution	Distribution. From the highlands of the Cordillera Oriental in E Ecuador and N Peru, extending E, S of rios Napo and Amazonas, to ¢. 70 ° W, 5 ° S to the Rio Jurud in W Brazil, and S, W of the Rio Ucayali, to the Rio Pachitea in EC Peru.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F32FFA59FDFD60A8F8C6.taxon	description	Descriptive notes. Head-body 46 - 65 cm (males) and 46 - 58 cm (females), tail 62 — 80 cm (males) and 53 - 72 cm (females); weight of males c. 7 kg (but when dominant they become more robust and can weigh as much as 10 kg) and of females 5 - 7 kg. Poeppig’s Woolly Monkeys range in color from light yellow-gray through every shade of brown to almost black, although generally with a silvery sheen. Head, hands, feet, chest, and groin are invariably black, with the remainder of the belly being deep reddish. Males are larger than females with larger canines and well-developed masseter muscles on either side of the head. The clitoris is as long as or longer than the penis, but adult males have easily seen testicles. A rare color morph is light yellow-gray with a dark head and extremities. The most silvery individuals come from the highest elevations along the western edge of its range. Those from the eastern (lowland) part of the distribution most likely represent a distinct subspecies.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F32FFA59FDFD60A8F8C6.taxon	biology_ecology	Habitat. Primary and some secondary lowland and high-elevation rainforest. Poeppig’s Woolly Monkeys use the middle and upper forest canopy.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F32FFA59FDFD60A8F8C6.taxon	food_feeding	Food and Feeding. The diet of Poeppig’s Woolly Monkey is similar to that of Humboldt’s Woolly Monkey (L. lagothricha). Two habituated groups in eastern Ecuador had an annual diet of 76 - 4 % fruits, 10 % insects, 7 - 2 % leaves, 3 - 3 % flowers, 1 - 3 % undetermined plant parts, 0 - 9 % aroid epiphyte pith, 0 - 5 % immature seeds, 0 - 3 % vertebrates including frogs, and 0 - 1 % fungi. Offruits eaten, 95 % were ripe or nearly so, and 80 % of the leaves were new or meristematic tissue of tendrils and growing points of epiphytic vines. Plant foods in the diet corresponded to a minimum of 231 morpho-species, belonging to at least 77 genera and 45 families. Offruits eaten, the diet was strongly biased toward a small number of important species. The nine most frequently selected genera made up more than 50 % ofall fruit feeding records: 12: 9 % Inga (Fabaceae), 8: 5 % Ficus (Moraceae), 7 - 1 % Spondias (Anacardiaceae), 4 -: 6 % Trichilia (Meliaceae), 4 - 2 % Otoba (Myristicaceae), 3: 8 % Coccoloba (Polygonaceae), 3 -: 3 % Quararibea (Bombacaceae), 3: 2 % Pourouma (Urticaceae), 2: 4 % Byrsonima (Malpighiaceae), 2 - 1 % Paullinia (Sapindaceae), 2 - 1 % Guatteria (Annonaceae), and 2 - 1 % Matisia (Bombacaceae). More than 75 % of all fruit feeding records came from just 31 genera from 22 families, and more than 50 % of the fruit were from just seven families: 15 - 3 % Fabaceae, 10 - 8 % Moraceae, 8 -: 2 % Anacardiaceae, 5 - 3 % Bombacaceae, 5 - 2 % Mpyristicaceae, 4 - 6 % Meliaceae, and 3 - 9 % Sapindaceae. Proportions of fruit in the diet varied from highs of 87: 5 % in March and 88: 7 % in July to a low of 63: 6 % in December. During months of low fruit consumption, Poeppig’s Woolly Monkeys ate more flowers and leaves. Males ate significantly more insects and other prey than either females or juveniles; adult males ate insects 10 - 7 % of the time and subadult males 13 %. Adult females ate insects only 7: 5 % of the time and subadult females 7 - 3 %. Feeding party size was 1 - 11 individuals (average 2 - 9). In this study, Poeppig’s Woolly Monkey generally eat from large trees; mean trunk diameter at breast height of all feeding trees was 43 - 2 cm. Another study in the same part of Ecuador recorded 104 species provided fruit, constituting about 74 % of the diet, with the remaining 10 % being leaves, 5 % flowers, and 11 % insect eggs, larvae, spiders, ants, caterpillars, termites, beetles, katydids, bees, cicadas, hemipteran bugs, millipedes, tree frogs, and lizards.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F32FFA59FDFD60A8F8C6.taxon	breeding	Breeding. Copulation of Poeppig’s Woolly Monkey is often preceded by solicitations from the female who uses a click (or “ tooth chatter ”) vocalization when 10 m or less from the male. The male sometimes answers the female in the same way, which sometimes continues through mating. Males may ignore solicitations from females. Nineteen of 44 mounts recorded were performed by adult males, 24 by subadult males, and one by a male of undetermined maturity. The average mount lasts c. 4 minutes, based on 25 mounts. Multiple males often mate with a single female during her period of receptivity on the same day or on separate days. Male Poeppig’s Woolly Monkeys are very tolerant of each another, but other adult females often harassed a copulating pair, sometimes interrupting them. Stereotyped scent marking using chest and anogenital areas is performed by males and females and seems to be associated with mating, although its role is not well understood. A single young is born after a gestation of ¢. 225 days.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F32FFA59FDFD60A8F8C6.taxon	activity	Activity patterns. Poeppig’s Woolly Monkeys are diurnal and arboreal. A time budget calculated for two groups was 35 - 5 % moving, 19 - 5 % eating, 17 - 3 % foraging, 17 - 1 % resting, 8: 7 % engaged in social activity, 1 - 4 % engaged in other activities, and 0 - 5 % engaged in agonistic behavior. There were monthly variations in activities, particularly for foraging and resting, correlated with fruit abundance. They foraged and rested more when fruit was abundant and rested less when fruit was scarce.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F32FFA59FDFD60A8F8C6.taxon	biology_ecology	Movements, Home range and Social organization. Groups of Poeppig’s Woolly Monkeys are multimale-multifemale, generally with 23 - 25 individuals. Genetic evidence indicates that most females disperse from their natal group, although some males also probably disperse. Two groups studied during one year moved an average of 2642 m / day and 2447 m / day in home ranges of ¢. 120 ha and 108 ha. The groups had core areas in their home ranges, which were used intensively. High home range overlap was typical, e. g. 45 %, 47 % and 70 % for three groups. Densities of Poeppig’s Woolly Monkeys in eastern Ecuador are the highest (so far) for any Lagothrix population at c. 31 ind / km?.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A04F32FFA59FDFD60A8F8C6.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The [UCN Red List. Poeppig’s Woolly Monkeys are hunted, their meat being considered delicious and the fat is used in cooking and for medicinal purposes. They are frequently kept as pets by Indians and Amazon settlers. Hunters target the mother to obtain the infant. They are sold openly as bushmeat in markets in Iquitos, despiteits illegality. There is reason to believe Poeppig’s Woolly Monkeys have declined by at least 30 % over the past 45 years (three generations) due primarily to hunting and habitat loss. They are known to occur in nine protected areas: Serra do Divisor National Park, Jutai-Solimoes Ecological Reserve, and Javari-Buriti Area of Outstanding Ecological Interest in Brazil; Podocarpus, Sangay Sumaco Napo-Galeras, and Yasuni national parks in Ecuador; and the Pacaya-Samiria National Reserve and ARC Comunal Tamshiyacu Tahuayo in Peru.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32FFF6DF7FF69AEF752.taxon	vernacular_names	Peruvian Yellow-tailed Woolly Monkey	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32FFF6DF7FF69AEF752.taxon	materials_examined	Peru, banks of the Amazon. Restricted byJ. Fooden in 1963 to Puca Tambo, 1550 m, 80 km east of Chachapoyas, San Martin.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32FFF6DF7FF69AEF752.taxon	discussion	This species is monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32FFF6DF7FF69AEF752.taxon	distribution	Distribution. N Peru in a few scattered locations on the E slopes of the Andes, along the Maranon-Huallaga divide, mainly confined to Amazonas and San Martin regions, but also in small areas of Huanuco and La Libertad regions.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32FFF6DF7FF69AEF752.taxon	description	Descriptive notes. Head-body 40 - 54 cm, tail 56 - 63 cm; weight 8 - 3 - 10 kg (males) and 5 - 7 kg (females). Pelage of the Peruvian Yellow-tailed Woolly Monkey is notably long and thick (particularly on legs), giving the body a robust, muscular look. It is normally a deep mahogany or coppery color, although darker on the lower back, nape, and extremities. A yellow band runs beneath the last one-half of the tail around the callous. There is a buffy triangular patch on the nose, with the apex between the eyes. The face is otherwise brown. Limbs are relatively short. Males have significantly longer canines than females; they also have a tuft of yellowish or brownish fur on the scrotum. Females have a smaller tuft around the vulva, with a clitoris that is as long as or longer than the penis. As in spider monkeys (Ateles), juvenile females can be differentiated by observing the clitoris.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32FFF6DF7FF69AEF752.taxon	biology_ecology	Habitat. Steep primary pre-montane and montane cloud forest in rugged terrain at elevations of 1400 - 2700 m, where there are deep gorges and ravines. Peruvian Yellowtailed Woolly Monkeys can survive in regenerating secondary forest if human disturbance is minimal. Regional temperatures are 4 - 25 ° C, with high humidity. They spend most of their time in the canopy and subcanopy.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32FFF6DF7FF69AEF752.taxon	food_feeding	Food and Feeding. Peruvian Yellow-tailed Woolly Monkeys feed primarily on ripe fruits, along with young leaves, flowers, lichens, leaf bases of bromeliads, epiphyte roots and bulbs, and possibly some arthropods. They infrequently raid maize crops.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32FFF6DF7FF69AEF752.taxon	breeding	Breeding. A single offspring is born every 3 - 5 years after a gestation of ¢. 7 - 5 months. Young lack the yellow tail band and the pubic tuft of the adults. Both sexes reach full maturity at ¢. 8 years of age, although successful reproduction may take place long before that. Infants are seen in groups throughout the year, but births peak in the wet season.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32FFF6DF7FF69AEF752.taxon	activity	Activity patterns. In a study at La Esperanza, a 700 - ha disturbed forest west of the Rio Imaza, three groups of Peruvian Yellow-tailed Woolly Monkeys studied for 15 months had the following activity budget: 29 % of the day traveling, 29 - 8 % feeding, 26 - 2 % resting, 2: 3 % engaged in social activities, and 12 - 7 % engaged in other activities. Adult females and females with infants spent more time feeding than males or juveniles. Females with infants spent the least time resting, and juveniles spent the most time engaged in social activities such as play.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32FFF6DF7FF69AEF752.taxon	biology_ecology	Movements, Home range and Social organization. Peruvian Yellow-tailed Woolly Monkeys live in multimale-multifemale groups of three to more than 20 individuals. Densities at La Esperanza were 8 - 3 - 9 - 3 ind / km? ® in 2008 - 2009, and group sizes averaged 8 - 9 individuals, with 1 - 3 adult males, 1 - 6 adult females, and 0 - 6 juveniles and infants. Lone males have been observed. They usually forage in smaller groups. A group of eleven individuals had a home range of 69 ha.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32FFF6DF7FF69AEF752.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Critically Endangered on The IUCN Red List. The Peruvian Yellow-tailed Woolly Monkey is one of the world’s rarest mammals. Until the 1950 s, the remoteness of its small remaining habitat afforded some protection, but since then much of it has been destroyed due to policies that promoted colonization through road building and the establishment of human settlements during the 1960 s and 1970 s. The area where it occurs has the highest rate of human immigration and deforestation of the entire country, and its habitat is greatly fragmented and disturbed due to clearance for agriculture and cattle ranching. It appears to be highly sensitive to alterations in its habitat, and group sizes today are probably unnaturally small because of hunting. Only a tiny population survives today. In 2009, only ¢. 2000 km * of suitable habitat for the Peruvian Yellow-tailed Woolly Monkey remained, and it has probably been extirpated from much of the area because of hunting. In 2012, the population was estimated at no more than 1000 mature individuals. Despiteits legal protection, itis still hunted by local communities for sport, and its meat and fur. Mothers are also shot so that infants can be taken and sold as pets, although they usually do not survive. Peruvian Yellow-tailed Woolly Monkeys are also persecuted for their alleged damage to crops. It occurs in five national protected areas: Rio Abiseo National Park, Alto Mayo Protected Forest, Cordillera de Colan National Sanctuary, Chayu Nain Communal Reserve, and Rio Nieva Reserved Zone. It also occurs in complementary protected areas: the conservation concessions of Alto Huayabamba and Abra Patricia, Private Area of Conservation Abra Patricia-Alto Nieva, Private Area of Conservation Hierba Buena-Allpayacu, and Huamanpata Conservation Area. Although there are many protected areas where Peruvian Yellow-tailed Woolly Monkeys occur, populations are small, and long-term management plans are needed to ensure their ongoing viability.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32EFA64F58C6992FD4E.taxon	vernacular_names	Southern Muriqui	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32EFA64F58C6992FD4E.taxon	materials_examined	Brazil. Restricted by C. Vieira in 1944 to Rio de Janeiro.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32EFA64F58C6992FD4E.taxon	discussion	Two subspecies were formerly recognized as arachnoides and hypoxanthus. C. P. Groves in 2001 classified them as separate species. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32EFA64F58C6992FD4E.taxon	distribution	Distribution. SE Brazil, through the coastal Serra do Mar in the states of Rio de Janeiro, Sao Paulo, and NE of Parana, the N limits are the Serra da Mantiqueira and the rios Paraiba and Paraiba do Sul.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32EFA64F58C6992FD4E.taxon	description	Descriptive notes. Head-body 49 - 6 cm (one male) and 47 - 9 cm (one female), tail 72: 6 cm (one male) and 75 - 3 cm (one female); weight 10 - 2 kg (one male) and 8 - 5 kg (one female). These measurements are from the Fazenda Barreiro Rico, Anhembi, Sao Paulo. Two adult males recorded by C. Vieira, one from Juquia and the other from Ubatuba, both in Sao Paulo, measured: head-body 63 cm and tail 65 cm. Sexual dimorphism of the Southern Muriquiis absent, except for small differences in lengths of canine teeth; males have longer canine teeth. Coat coloris very distinctive, being predominantly beige, with light, or dark brown or light gray-brown coloration. Geographic and regional variations in coat pattern exist. Southern Muriquis retain black pigmentation of the face, palms, and soles of the feet from infancy into adulthood. Adult males and females develop only minor depigmentation, characterized by small pink or white spots in the pubic region and sometimes on the face. Juveniles and adults have a pronounced round belly, due to their extensive small and large intestines for leaf digestion. They have long arms, a metacarpal hook with no external thumb, and a long prehensile tail, with the final third being glabrous with a sensorial tactile surface. Males have a large conspicuous scrotum, and females have an elongated cylindrical clitoris.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32EFA64F58C6992FD4E.taxon	biology_ecology	Habitat. Evergreen, dense, montane and submontane, semi-deciduous and deciduous forest at 40 - 1500 m above sea level in the Brazilian Atlantic Forest. Southern Muriquis originally occupied seasonal semi-deciduous forests of the Sao Paulo Plateau, but today only one population remains there, in forest patches in the Fazenda Barreiro Rico.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32EFA64F58C6992FD4E.taxon	food_feeding	Food and Feeding. Diets of the Southern Muriqui vary seasonally. They are strictly herbivorous. They have not been seen to eat insects or small animal prey, but insects are undoubtedly ingested incidentally with fruits and leaves. They are predominantly frugivorous, but leaves are also an important part of their diet. Studies at foursites in four different forests in Sao Paulo indicate a diet ofripe and unripe fruits (45 - 73 - 2 %), young leaves (3: 5 - 12 %), mature leaves (1 - 7 - 37 %), and flowers (2 - 6 %). They occasionally eat tree bark, vines, and branches, and they also eat soil. At Carlos Botelho State Park in Sao Paulo, they feed on 138 different plant species. They are more folivorous (leaves make up to 67 % of the diet) in degraded semi-deciduous forest fragments at Fazenda Barreiro Rico on the lowland plateau, where fruits are scarce. Southern Muriquis sometimes “ camp out ” together in varying numbers (2 - 42 individuals) in the late afternoon to sleep near a large patch offruiting trees and remain there for two or more days.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32EFA64F58C6992FD4E.taxon	breeding	Breeding. Southern Muriquis have long developmental periods. If similar to the Northern Muriqui (B. hypoxanthus), puberty occurs at c. 7 years of age. Reproduction is seasonal, with mating occurring mainly during the rainy season (October-November). Births occur throughout the year, but there is a peak in the dry season (May-June). The ovarian cycle is c. 21 days, and gestation is ¢. 216 days.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32EFA64F58C6992FD4E.taxon	activity	Activity patterns. Southern Muriquis typically feed most in the early morning and late afternoon. They rest for some time after the first morning meal, which is followed by traveling and foraging until late morning. In the dry cold months of the austral winter (June-July), they rest more in the morning and frequently may spend the entire morning in the same place. Resting increases in the hot austral summer, especially during the hottest hours of the day in the early afternoon. When fruit is more abundant in winter months, they are more active and travel and feed throughout the afternoon.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32EFA64F58C6992FD4E.taxon	biology_ecology	Movements, Home range and Social organization. Southern Muriquis are primarily arboreal; they spend a negligible amount of time on the ground, and then just to drink water or, rarely, eat soil. They live in multimale-multifemale groups but, at certain times of the year, split into subgroups to forage and travel. Subgroups are fluid in their composition. Daily movement patterns are determined strongly by the distribution of trees with edible fruits. Home range in the pristine continuous forest of Carlos Botelho State Park is ¢. 340 ha. Sometimes they travel across their home range in a single day; daily movements vary from ¢. 600 m in dry cold months up to 5100 m in hot wet months (average 2267 m). Male and female Southern Muriquis have different daily home ranges, which may be associated with energetic demands of reproduction. Females travel more during the day to acquire alternative foods, which may be explained by their nutritional requirements for reproduction. Females disperse when close to puberty (6 - 7 years old). Infanticide has never been recorded. Most of the time, Southern Muriquis are not aggressive and have high levels of tolerance for each other. Nevertheless, non-affiliative behaviors such as inter-individual chasing, attempts at biting, body-touching and body-grabbing occur, and a coalitionary lethal killing was once observed in Carlos Botelho. Six individuals mounted a fatal attack on an older male. Reasons for the killing were thought to be related to resource competition for food or reproductive females. The main predators include large cats such as the Puma (Puma concolor) and the Jaguar (Panthera onca), medium-sized cats such as the Ocelot (Leopardus pardalis) and Geoffroy’s Cat (Leopardus geoffroyi), and small cats such as the Oncilla (Leopardus tigrinus) and the Jaguarundi (Puma yagouaroundi). The black hawkeagle (Spizaetus tyrannus) can also take infants. Southern Muriquis give alarm calls, which they also use for snakes such as Bothrops and Micrurus.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0BF32EFA64F58C6992FD4E.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. Southern Muriquis are classified as endangered on the Brazilian List of Threatened Species. Their numbers have declined from an estimated 400,000 individuals before 1500 to less than 2500 in 2008 because of widespread forest loss and hunting. They are hunted for cultural and recreational purposes and, notably today, by illegal palm heart harvesters while camping in the forest. The upper tail muscles of the Southern Muriqui are particularly prized for food, and this culinary preference persists among hunters and palm harvesters. The current population status of the Southern Muriqui is not well known, and there are ongoing studies to this end by the non-governmental organization Associacao Pro-Muriqui as part of a national action plan developed by the Brazilian government. It is now extinct in several protected and unprotected areas where it was once abundant, and the main reasons were illegal hunting and lack of law enforcement. Southern Muriquis are thought to occur in 18 public and private protected areas, but all but four are too small, or the habitat is too degraded, to fully protect their populations. Recent estimates indicate that 1300 Southern Muriquis remain in disjunct populations in the states of Sao Paulo and Rio de Janeiro, with three smaller, fragmented populations in Parana (Castro) and the interior of Sao Paulo (Fazenda Sao Sebastiao in the Serra da Mantiqueira and Fazenda Barreiro Rico on the Sao Paulo Plateau). The largest populations occur in the largest tract of continuous forest of the Atlantic Forest Biome, the World Heritage Site of the Southern Reserves of the Atlantic Forest, which includes the contiguous Carlos Botelho (37,432 ha), Intervales (45,000 ha), and Alto Ribeira (55,000 ha) state parks and Xitué State Ecological Station (3095 ha). Carlos Botelho State Park has a population of ¢. 500 individuals.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0AF32DFA58FCB968B9F6C6.taxon	vernacular_names	Northern Muriqui	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0AF32DFA58FCB968B9F6C6.taxon	materials_examined	Brazil, Bahia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0AF32DFA58FCB968B9F6C6.taxon	discussion	Two subspecies were formerly recognized, arachnoides and hypoxanthus. C. P. Groves in 2001 classified them as separate species. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0AF32DFA58FCB968B9F6C6.taxon	distribution	Distribution. Historically ranged through the Atlantic Forest in the states of Bahia, Espirito Santo, Minas Gerais, and Rio de Janeiro, excluding only lowland forests in the extreme S of Bahia and N Espirito Santo, the N limit of the distribution was probably the Rio Jequirica or the right bank of the Rio Paraguacu, and the S limit is still poorly defined but is probably the Serra da Mantiqueira in S Minas Gerais, near the state boundaries with Rio de Janeiro and Sao Paulo where it meets the distribution of the Southern Muriqui (B. arachnoides).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0AF32DFA58FCB968B9F6C6.taxon	description	Descriptive notes. Head-body 47 - 8 - 49 - 7 cm (males) and 46 - 1 - 51 - 4 cm (females), tail 73 - 4 - 77 - 3 cm (males) and 73 - 8 - 81 cm (females); weight 9 - 2 - 9 - 6 kg (males) and 6 - 9 - 8 - 8 kg (females); sample of three individuals in all cases. These measurements were taken from living individuals captured in Fazenda Esmeralda, Rio Casca, Minas Gerais. Male and female Northern Muriquis are similar in size, and there is no sexual dimorphism in the size of their canines. Two male and four female specimens in the British Museum from Engenheiro Reeve, Espirito Santo, listed by P. H. Napier in 1976, were head-body 58 - 61 cm (males) and 54 - 5 - 60 cm (females), tail 67 - 69 cm (males) and 74 - 80 cm (females). Although muriquis are generally hailed as the largest New World monkeys, these measurements do not exceed those of large spider monkeys such as the White-bellied Spider Monkey (Ateles belzebuth) and the Red-faced Black Spider Monkey (A. paniscus), or even a large male Central American Black Howler (Alouatta pigra). There is likely natural geographical variation in the body size of Northern Muriquis, and nutritional status and time of the year may result in disparate measurements. Pelage of the Northern Muriqui is very distinctive and predominantly beige, with light or dark brown or light gray-brown coloration. Northern Muriquis are born with a black face, but at sexual maturity, the face and genitalia lose their pigmentation and become spotty pink. Juveniles and adults have a pronounced swollen belly, accommodating extensive small and large intestines for the digestion of leaves. Adaptations for suspensory locomotion and tail-assisted brachiation include long arms, a metacarpal hook, and a long prehensile tail. Males have a large conspicuous scrotum, and females have an elongated cylindrical clitoris. Northern Muriquis have a vestigial thumb, which is diagnostic in differentiating them from Southern Muriquis that lack the thumb.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0AF32DFA58FCB968B9F6C6.taxon	biology_ecology	Habitat. Evergreen, dense, semi-deciduous and deciduous forest at 250 - 1350 m above sea level. The majority of the information available on the ecology, behavior, reproduction, and demography of the Northern Muriqui comes from the work of K. B. Strier and her colleagues since 1982 in seasonal, semi-deciduous forest in Caratinga Biological Station (Reserva Particular do Patrimonio Natural Feliciano Miguel Abdala), Minas Gerais. Habitats in Caratinga include relatively intact and mature forest (canopy 30 - 35 m), disturbed (degraded, logged) mature forest, swathes of secondary forest in advanced stages of succession (canopy 25 m), and young secondary growth. G. Fonseca in 1986 found that Northern Muriquis preferred late successional secondary forest. They used the intact forest as much as would be expected based on availability and avoided degraded mature forest. Contrary to expectations considering their size, they are not averse to entering young secondary growth — no doubt allied to their predilection for young leaves.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0AF32DFA58FCB968B9F6C6.taxon	food_feeding	Food and Feeding. The diet of a group of Northern Muriquis monitored over 14 months at Caratinga was 51 % leaves, 32 % fruits (mostly ripe), 11 % flowers, and bark, bamboo, buds, and ferns (6 %). Diets varied with the availability and abundance of food resources throughout the year. Sixty three plant species from 57 genera were identified as food sources. They ate flowers of Apuleia praecox (Fabaceae) and Adenocalymma marginatum (Bignoniacae), and nectar and pollen of Mabea fistulifera (Euphorbiaceae). Diets were similar between the sexes, except that the females tended to eat more flowers.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0AF32DFA58FCB968B9F6C6.taxon	breeding	Breeding. At Caratinga, breeding of Northern Muriquis is seasonal. Females usually disperse while still adolescent at c. 6 years old (average 73 - 4 months, range 64 - 78 months). A female that stayed in her natal group had her first infant at ¢. 7 - 5 years old, but age at first reproduction of dispersing females is prolonged. Entering another group takes a while, and following immigration, sexual activity is delayed a minimum of eleven months. Most females undergo puberty only after they have transferred to another group. On average, they give birth to their first infant at 33 - 34 months after joining the group. Ovarian cycles last c. 21 days, but there are no visible signs of sexual receptivity. Pheromonal cues, especially from urine, are probably important, and males sniff the females’ genitalia. They pull on the females’ pendulous clitoris, which, rubbing on branches as it does, is probably important in distributing information about her sexual condition. Sexual interactions are concentrated in the early and middle wet season, and births peak in the dry season (April-September). Nearly two-thirds (66 %) of 37 births recorded from 1982 to 1994 at Caratinga occurred in June-August and 86 % in the dry season. Given c ¢. 7 months of gestation, conceptions are concentrated in the wet season (October — March). The interval between parturition and the first post-partum mating averages 23 - 5 months, and mating continues for an average of 5 - 6 months. Females rarely continue their sexual activity into the dry season, so those that do not conceive in the wet season can experience a year’s delay in reproduction (mating again only in the next wet season). Non-lactating females do not cycle throughout the year. Stimuli involved in the shut-down of ovarian cycling in the dry season may be shorter day lengths, declining female condition, or inclusion or exclusion of items in the diet that may affect fertility. Females that conceive early in the wet season (to give birth early in the dry season) resume sexual activity sooner (average 22 - 2 months) than females that conceive late in the wet season (23 - 9 months) or in the early dry season (24 - 3 months). Birth intervals average 36 - 4 months. Giving birth in the dry season is believed to be advantageous because it is not a period when fruit is abundant, which implies much traveling; relying on leaves, daily movements are shorter. Carrying an infant becomes more tiring as they grow, and mothers are able to benefit from increased food abundance in the early wet season when infants are getting larger but are still being carried most of the time. Male Northern Muriquis interact very little with infants. Weaning begins when infants are c. 18 months old. Infants born in the dry season of one year are weaned in the wet season of the next year when fruit, young leaves, and flowers are abundant.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0AF32DFA58FCB968B9F6C6.taxon	activity	Activity patterns. Activity budgets of Northern Muriquis at Caratinga are relatively constant, averaging 49 % resting, 29 % traveling, 19 % feeding, and 3 % engaged in social and other activities. The pattern during the day is variable, but they there is a distinct habit of feeding more in the late afternoon. Resting and feeding are inversely related. They tend to be more active during the warmest hours of the day when it is cooler in the dry winter months (April-September), and they are more active in the coolest hours during the hotter summer months of the wet season (October-March). In the dry season, mature leaves make up a greater portion of their diet, and they travel less. In the wet season, when fruits, flowers, and young leaves are more abundant, they tend to travel more. Male and female activity budgets do not differ, but adults tend to rest more and immature animals are more active. Lactating females spend more time feeding than those without nursing infants. A 7 - month study by R. Lemos de Sa of a group of 15 Northern Muriquis at the Fazenda Esmeralda recorded an activity budget of 62 % resting (monthly range 56 - 68 %), 18 % feeding (11 - 24 %), 16 % traveling (11 - 24 %), and 4 % engaged in social and other activities (1 - 5 %). Time spent feeding was similar to that found at Caratinga, but times spent resting (more) and traveling (less) were undoubtedly related to their smaller home range in a forest of only 44 ha. They spent more time feeding in the dry season (19 - 23 % of the day) when food was scarcer than in the wet season when food was more abundant (9 - 15 %).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0AF32DFA58FCB968B9F6C6.taxon	biology_ecology	Movements, Home range and Social organization. Northern Muriquis live in multimale — multifemale groups as a large as ¢. 60 individuals. In a 14 - month study at Caratinga (1983 - 1984), a group of 23 - 27 individuals, with six adult males and eight adult females, had a home range of more than 168 ha in an 800 - ha forest. Their, daily movements averaged 1283 m (141 - 3403 m). About 46 % of the home range was also used by a neighboring group. Home range use by this group was studied again in 1998 - 1999. It had grown to 57 - 63 individuals, including 13 - 14 adult males and 19 adult females. Home range size had increased to 309 ha. Daily movements remained quite constant despite the increase in home range size, averaging 1313 m (200 - 2835 m). At thissize, the group tended to split into subgroups that traveled and foraged independently. Subgroups were larger when food was more abundant in the wet season (average 41 - 8 individuals) than in the dry season (36 - 6 individuals) when fruits were scarce. Males generally stay in their natal group, but females disperse to other groups before they become sexually active. Intergroup encounters provide stimulus for emigrations and immigrations, and the process is evidently stressful. Females trying to enter a new group are not welcomed by resident females. They spend some time peripheral to the group and are chased and harassed. Sometimes they return for a time to their natal group. Even when tolerated, they tend to remain peripheral to a new group for some months, interacting most with the new group’s juveniles and adolescent males. Philopatric males have a high degree of relatedness among themselves, and certain individuals tend to consistently travel together. Females travel with their offspring of various ages. Northern Muriquis are promiscuous. Males and females are co-dominant, and they show no aggression in either sexual or non-sexual contexts. When aggression does occur, it is generally mild. Female choice probably makes it of little advantage for males to compete physically. Sexual competition is believed to be enacted through sperm competition rather than through overt contests. Males have large testicles that produce voluminous quantities of sperm.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
313A88142A0AF32DFA58FCB968B9F6C6.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Critically Endangered on The IUCN Red List. Northern Muriquis are also classified as critically endangered on the Brazilian List of Threatened Species. Once widespread in the Atlantic Forest, their numbers are estimated at ¢. 1200 individuals. Habitat fragmentation and hunting in the most heavily populated states of Brazil are the reasons for their decline. Threats to remaining populations include low genetic diversity and the isolation of the remaining populations. Today, they are found in fourteen separate populations, most of them formally protected. A national action plan was developed by the federal government in 2011, which sets out priorities and needs for research and conservation of Northern Muriquis. These include increasing habitat availability and providing connectivity among the forest patches where they survive. The Rio deJaneiro Primate Center has an effective breeding program, but with few individuals that were pets or rescues. There is no immediate reason to have individuals taken from the wild for captive breeding. A number of non-governmental organizations and the Brazilian government are working to save and nurture extant wild populations. Northern Muriquis occur in Caparao, Alto Cariri and Itatiaia national parks; Augusto Ruschi and Mata Escura biological reserves; Serra do Brigadeiro, Rio Doce, Ibitipoca, and Alto Cariri state parks; Mata dos Muriquis Wildlife Refuge; Reserva Particular do Patrimonio Natural Feliciano Miguel Abdala (Caratinga Biological Station), Mata do Sossego, and Fazenda Duas Barras private reserves. They are also found in a number of small, privately owned forest patches in Santa Maria de Jetiba, Espirito Santo. The extraordinary recovery of the population in Feliciano Miguel Abdala reserve, from c. 40 individuals in the early 1980 s to well over 200 individuals today, testifies to the resilience of the Northern Muriqui, given adequate protection and space.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Atelidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 484-549, DOI: http://doi.org/10.5281/zenodo.5727205
