Dusky titi
Callicebus moloch

TAXONOMY

Suborder: Haplorrhini
Infraorder: Simiiformes
Family: Pitheciidae
Subfamily: Callicebinae
Genus: Callicebus
Species: C. aureipalatii, C. baptista, C. bernhardi, C. brunneus, C. caligatus, C. cinerascens, C. cupreus, C. discolor, C. donacophilus, C. dubius, C. hoffmannsi, C. modestus, C. moloch, C. oenanthe, C. olallae, C. ornatus, C. pallescens, C. stephennashi

Other names: titi monkey, luca luca, lucachi; callicèbe (French); springaffe (German); zogue-zogue (Spanish); C. aureipalatii: golden palace monkey, Madidi titi monkey; C. baptista: Baptista lake titi; C. bernhardi: Prince Bernhard's titi monkey; C. brunneus: brown titi monkey; brun springapa (Swedish); C. caligatus: booted titi, chestnut-bellied titi; brunbröstad springapa (Swedish); C. cinerascens: ashy black titi, ashy titi, ashy-grey titi; askgrå springapa (Swedish); C. cupreus: coppery titi monkey, red titi monkey; callicébe roux (French); roter springaffe, kopferfürbiger springaffe (German); kopparspringapa, röd springapa (Swedish); C. discolor: red titi; socayo, songo songo (Spanish); C. donacophilus: Bolivian grey titi, Bolivian titi monkey, white-eared titi; vitörad springapa (Swedish); C. dubius: doubtful titi monkey, dubious titi, Hershkovitz's titi; vitpannad springapa (Swedish); C. hoffmannsi: Hoffmann's titi monkey; gulbröstad springapa, Hoffmanns springapa (Swedish); C. modestus: Bolivian titi, modest titi, Rio Beni titi, mono tití (Spanish); Boliviansk springapa, gråbrun springapa (Swedish); C. moloch: dusky titi, red-bellied titi; grijze springaap (Dutch); titi molock (French); grå springapa, mörk springapa, orabassn (Swedish); C. oenanthe: Andean titi monkey, Rio Mayo titi; sugkamat (Aguaruna); bergsspringapa (Swedish); C. olallae: Beni titi monkey, Olalla's titi, Ollala brothers' titi; mono tití (Spanish); benispringapa, Olallas springapa (Swedish); C. ornatus: ornate titi monkey; C. pallescens: white-coated titi; C. stephennashi: Stephen Nash's titi monkey.

The taxonomic arrangement of this genus is debated (see van Roosmalen et al. 2002). Groves (2001) places titis into four groups, subsuming the Personatus group of van Roosmalen et al. (2002) under the Moloch group. Kobayashi (1995) and van Roosmalen et al. (2002) place the titis into five taxonomic groups; Torquatus, Personatus, Moloch, Cupreus and Donacophilus groups. However, the species in the Personatus group according to Kobayashi (1995) and van Roosmalen et al. (2002) are separated from the other nearest Callicebus species by around 1,000 km (621.4 mi) (van Roosmalen et al. 2002). In addition, the members of their Torquatus group are differentiated from the other groups by habitat and ecology (van Roosmalen et al. 2002). For these reasons, the members of the Torquatus and Personatus groups according to Kobayashi (1995) and van Roosmalen et al. (2002) will be addressed elsewhere and only the members of their Moloch, Cupreus and Donacophilus groups as defined by Kobayashi (1995) and van Roosmalen et al. (2002) will be considered here. The Cupreus group contains C. caligatus, C. cupreus, C. discolor, C. dubius, C. ornatus, and C. stephennashi. The Moloch group consists of C. baptista, C. bernhardi, C. brunneus, C. cinerascens, C. hoffmannsi, and C. moloch. Finally, the Donacophilus group is made up of C. donacophilus, C.modestus, C. oenanthe, C. olallae, and C. pallescens (van Roosmalen et al. 2002). The recently described C. aureipalatii shows physical similarities to members of the Cupreus group but its distribution borders that of members of the Moloch group (Wallace et al. 2006). In this factsheet when titi monkeys are referred to, data about species from the Torquatus and Personatus groups are omitted. In addition, the discussed species are grouped together, primarily due to a lack of information about the majority of the species and presumable similarities in ecology and behavior.

MORPHOLOGY

Titi monkeys are small to medium-sized primates, typically about the size of a rabbit (Hershkovitz 1990; Kobayashi 1995). They do not have prehensile tails and are not sexually dimorphic (Hershkovitz 1990). Pelage and body coloration differs markedly between species of titi monkeys, but all species typically have thick fur or are somewhat shaggy in appearance (Hershkovitz 1990). All species can be differentiated based on differences in pelage, body color and pattern (Hershkovitz 1988). Body and extremity pelage can range from blackish, brown, grey to orange, red, or yellow and combinations of the colors. In addition, some species have dorsal surfaces contrasting with the rest of their body, often a lighter shade, while others are fairly uniform in color across their body. The tail sometimes contrasts with the rest of the body and some species have contrasting ear tufts (Rowe 1996).

C. donacophilus is characterized by body and limbs that range from grey agouti to orange agouti with entirely orange chest and belly and white ear tufts. C. cupreus sideburns, ventral surfaces and inner extremities are reddish. This coloration contrasts sharply with the animal's dorsal surfaces including its tail and head, which are brown. C. moloch is colored in much the same pattern as C. cupreus but its sideburns, ventral surfaces and inner extremities are light-orange to orange. In addition, its dorsal surfaces are greyish to pale-brown agouti (van Roosmalen et al. 2002). The coloration of the forehead and sideburns is either the same as the head or is contrasting in color. The skin of the face, ears and genitals is blackish (Hershkovitz 1990). See van Roosmalen et al. 2002 for more information about variation between titi species.

There is no sexual dimorphism in titi monkeys. The head and body of C. donacophilus males averages 311mm (12.2 in) while females average 340mm (13.4 in). In C. moloch, these measurements average 333mm (13.1in) for males and 331mm (13.0 in) for females. The tail is longer than the head and body combined (Hershkovitz 1990). Body weight is variable, ranging from around 800 g (1.8 lb) to around 1200 g (2.6 lb). C. moloch males range in weight from 850-1200 g (1.9-2.6 lb), averaging 1017 g (2.2 lb) while females range from 700-1020 g (1.5-2.2 lb), but average 877 g (1.9 lb). C. cupreus males average 1106 g, ranging from 1000-1175 g (Hershkovitz 1990).

In captivity, C. donacophilus, C. cupreus, and C. moloch have all lived to older than 25 years and both sexes of differing species have reached that age (Weigl 2005).

Titi monkeys use short leaps to move about the lower levels of the forest, including the understory and brush layer although titis will enter the main canopy (Youlatos 1999; Lawler et al. 2006). Much of the lower levels of the forest are discontinuous, that is, the animals are not able to move without leaping small distances. The majority of these leaps are only for small distances, often less than several body lengths (Lawler et al. 2006). They can be best described as quadrupedal, moving through their environment primarily through walking, clambering, and leaping, supplemented by bounding and climbing (Youlatos 1999). Titi monkeys are only rarely seen on the ground, but when traveling on the ground, titis can move relatively quickly, utilizing a bounding movement which may see them leap as high as a meter off of the ground (Fragaszy 1979; Kinzey 1981). Titis prefer horizontal supports which are less than 5cm in diameter and their tails never touch what they are walking on (Welker et al. 1998a; Youlatos 1999).

RANGE

CURRENT RANGE MAPS (IUCN REDLIST):
Callicebus aureipalatii | Callicebus baptista | Callicebus bernhardi | Callicebus brunneus | Callicebus caligatus | Callicebus cinerascens | Callicebus cupreus | Callicebus discolor | Callicebus donacophilus | Callicebus dubius | Callicebus hoffmannsi | Callicebus modestus | Callicebus moloch | Callicebus oenanthe | Callicebus olallae | Callicebus ornatus | Callicebus pallescens | Callicebus stephennashi

Titi monkeys are widely distributed across South America, found in Bolivia, Brazil, Colombia, Ecuador, Paraguay, and Peru. The species under consideration here are for the most part found south of the Amazon River and west of the Brazilian cerrado, or grassland (Hershkovitz 1988; van Roosmalen 2002). In the west, the range is limited by altitude and the rise of the Andes and the distribution extends as far south as Paraguay, between the Pilcomayo and Paraguay Rivers (Hershkovitz 1988).

C. baptista, C. bernhardi, C. cinerascens, C. dubius, C. hoffmannsi, C. moloch, and C. stephennashi are only found in Brazil (Hershkovitz 1990; van Roosmalen 2002). Species native to only Bolivia include C. donacophilus and C. aureipalatii, which is restricted to the northwest of the country (Hershkovitz 1990; van Roosmalen 2002; Wallace et al. 2006). Both C. olallae and C. modestus are also only found in Bolivia, but share a restricted distribution, confined only to the upper Rio Beni drainage basin. C. brunneus is also found in Bolivia, but its range extends to Brazil and Peru as well. C. oenanthe is only found in northern Peru. C. caligatus is found in Brazil and Peru and C. cupreus is found in Brazil, Peru, and Ecuador. C. discolor is found in Colombia, Ecuador, Peru and possibly Brazil, although its presence has not been confirmed there. C. ornatus is only found in eastern Colombia and is separated by about 350 km (217.5 mi) from any other member of the Cupreus group by C. torquatus. C. pallescens is the only species of titi monkey found in Paraguay, although it is also found in Brazil (Hershkovitz 1990; van Roosmalen 2002).

HABITAT

The habitat of the titi monkeys described here are somewhat varied, ranging from evergreen neotropical terra firma rain forest, to floodplain forest, to inundated forest, to forest edge, swamp edge and "broken up" forest areas (Janson pers. comm. cited in Kinzey 1981; Kinzey 1981; Warner 2002). They prefer dense vegetation, often choosing the thickest available including dense tangles, thickets, thorny underbrush and areas that are wet or even waterlogged on the ground as well as areas near streams and lakes (Moynihan 1976; van Roosmalen et al. 2002; Felton et al. 2006). Titis like gallery forest, high forest, liana forest, and treefall edges as well as secondary growth and patches, often in the middle of savanna and near streams (Moynihan 1976; Youlatos 1999). They also are found in flooded forest and dense, low forest with broadleaved trees and bushes (Moynihan 1976).

Titis can also be found in highly disturbed habitats (Ferrari et al. 2000). Among the titis, several species have distinctive habitat preferences not shared by other species. C. donacophilus is found in gallery forests areas nearby or associated with open areas such as grasslands and swampy grasslands while C. cinerascens habitat is associated with cerrado woodlands, an intermediate area between forest and savannah (Ferrari et al. 2000). Titis will also enter and utilize stands of imported bamboo. If present, bamboo can be quite important to the species, probably because titis prefer dense vegetation (Moynihan 1976; Warner 2002). In Colombia, titis can occur in the piedmont as high as 500m (1640.4 ft) altitude while in Peru, C. oenanthe can be found in cloud forests above 800m (2624.7 ft) (Hernández-Camacho & Cooper 1976; Aquino & Encarnación 1994).

Near the eastern end of the contiguous titi distribution in the Brazilian Amazon in the Amazon (Tapajos) National Park, rainfall averages 175.4 cm (69.1 in) annually with a dry season between July and November and the highest rainfall between February and March (Branch 1983). Further west, in Peru, the dry season is somewhat earlier, occurring between May and September or early October (Terborgh 1983; Warner 2002). However, the timing of the dry and wet seasons can vary considerably from year to year (Terborgh 1983). Average temperatures at the Cocha Cashu study site in Peru can range from a high of around 30°C (86°F), usually between August, September and October to typical lows between 14-16°C (57.2-60.8°F) (Terborgh 1983). Rainfall at Cocha Cashu averages around 200 cm (78.7 in) annually, mostly between October and April (Wright 1996).

ECOLOGY

Titi monkeys are predominantly frugivores who will also eat some invertebrates and other parts of plants (Wright 1989). Titis will eat over a hundred different species of fruit and plants (Wright 1985). Other researchers describe them as strongly omnivorous (Izawa & Yoneda 1981). Due to their predominantly vegetarian diet, a significant portion of their day is spent at rest, from less than 25% in C. discolor in Ecuador (including periods when they are resting out of view) to 60% in C. moloch in Peru (Terborgh 1985; Carrillo-Bilbao et al. 2005; Lawrence 2007). Based on time spent feeding during the dry season in Peru, titis eat 70% fruit, 26% leaves, and less than 1% insects (Kinzey 1978). They prefer fruits from small trees because they can economically exploit foods concentrated as such, and they will often eat fruit that is not ripe (Wright 1989). Titis will eat moths, butterfies, cocoons, spiders, and ants, generally less than 2cm (.8 in) in size and are able to capture flying insects from the air (Meritt 1980; Crandlemire-Sacco 1988; Wright 1985; 1989). When it comes to prey however, titis will only rarely seek out prey and prefer to scan their environment for their quarry in a stationary manner (Crandlemire-Sacco 1988; Wright 1989). In addition a large proportion of the diet is leaves, especially young leaves and leaf buds, which supply protein (Terborgh 1985; Crandlemire-Sacco 1988). A large proportion of the titi's feeding time is spent in smaller trees often with small crowns less than 10 m in diameter (Wright 1984a; 1985). They will not eat exudates (Crandlemire-Sacco 1988). In addition, there is an increase in feeding time on leaves, including vine and bamboo leaves, during the dry season (Wright 1989; Lawrence 2007). During lactation, the proportion of insects in the diet of females increases, probably due to an increased protein need (Herrera & Heymann 2003).

The titi is diurnal (Mason 1968). Titis will rise early in the morning after sunrise and will remain awake and active until around sunset or slightly before (Kinzey 1978; Wright 1989). However, the length of daily activity varies with the seasons, with titis rising before dawn in the warmer months when more fruit is available and sometimes staying in the sleeping site for up to four hours after sunrise in colder months when fruits are harder to find (Wright 1989). The day is typically divided up into two main feeding sessions, one in the morning and one in the afternoon, separated by a period of rest at midday (Kinzey 1978; Mason pers. comm. cited in Kinzey 1978). The last several hours of the day are often spent intensely feeding on leaves (Janson pers. comm. cited in Kinzey 1981). Total daily activity lasts an average of 11.5 hours with feeding averaging around 2.7 hours per day (Kinzey 1978).

Sleeping sites are typically in vine tangles located on small branches which are at least about 15 meters (49.2 ft) above the ground. Sites are sometimes reused from night to night and are always more than 100 meters (328.1 ft) from the edge of the home range (Kinzey 1981; Wright 1995). During sleeping, group members are close to one another, within one meter (Mason 1968). In addition, during rest and sleeping, titis will huddle together and entwine their tails in a characteristic fashion (tail twinning) (Mason 1968; Robinson 1979a; Kinzey 1997).

Typical day range can vary between 425 m (1394.4 ft) and 1152 m (3779.5 ft), but usually averages towards the lower end of that range (Mason 1968; Kinzey 1981; Wright 1985; 1989; Polanco-Ochoa & Cadena 1993). However, due to weather and exceptional circumstances, the day range can be significantly shorter or longer, having been observed as low as 150 m (492.1 ft) and as high as 1450 m (4757.2 ft) (Wright 1985). Home ranges are also variable but generally small, varying between less than .005 km² (.002 mi²) up to .14 km² (.05 mi²) (Mason 1968; Kinzey 1978; Terborgh 1983; Polanco-Ochoa & Cadena 1993; Carrillo-Bilbao et al. 2005; Lawrence 2007). However, some estimates place the home range somewhat higher, between .1 km² (.04 mi²) and .2 km² (.08 mi²) (Izawa & Yoneda 1981). In the dry season, when less fruit is available to titis, the daily path is reduced by up to two-thirds to conserve energy (Patricia Wright cited in Terborgh 1985).

Titi monkeys can live in the same habitats as a number of other primates including marmosets (Callithrix sp.), tamarins (Saguinus sp.), squirrel monkeys (Saimiri sp.), saki monkeys (Pithecia sp.), capuchins (Cebus sp.), owl monkeys (Aotus sp.), howler monkeys (Alouatta sp.), woolly monkeys (Lagothrix sp.), and spider monkeys (Ateles sp.) (Wright 1988; Youlatos 1999). Larger sympatric species will chase the smaller titis from food sources, including larger fruit trees (Wright 1996). Titis coexist with other sympatric species as is the case with C. moloch ranging with S. fuscicollis due to differential habitat utilization and diet (Crandlemire-Sacco 1988). Nevertheless, if possible, titi monkeys will avoid all other primates (Wright 1985).

The most important predators of titi monkeys are raptors. Several different sizes and species of raptors are capable of and have been observed attacking a titi, including the Guianan crested eagle (Morphnus guianensis) and the Ornate hawk-eagle (Spizaetus ornatus) (Terborgh 1983). Felids may also be a predator of the titi monkey, but arboreal mammals make up only a small and unimportant part of their diet and are only taken in exceptional circumstances (Emmons 1987). Arboreal snakes are also present in some titi habitats and may also be a predator (Wright 1985; Cisneros-Heredia 2007). Tufted capuchins (Cebus apella) have rarely been observed to predate titis. Sampaio & Ferrari (2005) observed a tufted capuchin killing and subsequently consuming an infant of C. moloch. In addition, Lawrence (2007) reported that a large female C. brunneus juvenile died after it had been attacked by tufted capuchins while feeding in the same tree. Titis avoid predators primarily through crypsis and hiding (Terborgh 1983).

Content last modified: December 19, 2007

Written by Kurt Gron. Reviewed by Gabriela de Luna.

Cite this page as:
Gron KJ. 2007 December 19. Primate Factsheets: Dusky titi (Callicebus moloch) Taxonomy, Morphology, & Ecology . <http://pin.primate.wisc.edu/factsheets/entry/dusky_titi>. Accessed 2015 March 2.