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Alouatta pigraMexican black howler monkey

By Kimberly Lau

Geographic Range

Alouatta pigra is found in Belize, northern Guatemala, and southeastern Mexico ("Belize Zoo", 2007; Gonzalez-Kirchner, 1998). They are endemic to the largest tropical rain forest region of Mesoamerica, Selva Maya, which encompasses over 4 million hectares of land in these three countries (Estrada et al., 2004). They are the only howler monkey species on the Yucatan peninsula, and are found in a reduced area of the Yucatan (Estrada et al., 2004; Gonzalez-Kirchner, 1998). They inhabit the Mexican states of Campeche, Quintana Roo, and Chiapas (Estrada et al., 2004). Populations have been found and heavily studied at the Mayan sites of Calakmul and Yaxchilán, Mexico, and Tikal National Park, Guatemala (Estrada et al., 2004). They are widespread throughout Belize. ("Belize Zoo", 2007; Estrada, et al., 2004; Gonzalez-Kirchner, 1998)

Habitat

The habitat of Alouatta pigra is varied, but is typically tropical forests. They are known from lowland, deciduous, semi-deciduous forests, and evergreen forests (Belize Zoo, 2007; Gonzalez-Kirchner, 1998). They are known from pristine tropical rainforest to riverine forests, and can be normally found at low elevations in riparian forests of less than 400 m, although black howlers have been found at elevations higher than 500 m (Estrada et al., 2004). In riverine habitats they usually avoid riverbank areas and inhabit inland areas (Estrada et al., 2004). They prefer larger trees and usually inhabit the middle to upper canopy (Gonzalez-Kirchner, 1998). Although they have a wide habitat range, A. pigra has lower rates of success in areas disturbed by humans (Gonzalez-Kirchner, 1998). ("Belize Zoo", 2007; Estrada, et al., 2004; Gonzalez-Kirchner, 1998)

  • Average elevation
    400 m
    1312.34 ft

Physical Description

Alouatta pigra is one of the largest monkeys in the Americas. It averages 70 cm long, excluding the tail, which is slightly longer than its body length ("Belize Zoo", 2007). It is the largest and most sexually dimorphic of the howler monkeys, with an average weight of 11.4 kg for males and 6.4 kg for females (Fleagle, 1999). They are characteristically covered by black hair, which is a bit longer near the throat, and they have white genitalia (Treves, Drescher, & Snowdon, 2003). Males have a pink scrotum, which distinguishes Alouatta pigra from Alouatta palliata (Horwich, 1983a). Young retain pale fur until 9 to 10 weeks of age (Treves, Drescher, & Snowdon, 2003). Like other members of Atelidae, A. pigra has a prehensile tail with gripping pads (Fleagle, 1999). ("Belize Zoo", 2007; Fleagle, 1999; Horwich, 1983a; Treves, et al., 2003)

  • Sexual Dimorphism
  • male larger
  • Average mass
    6.4 - 11.4 kg
    lb
  • Average length
    70 cm
    27.56 in

Reproduction

Generally, only the dominant males or other males with high social status in the troop will copulate with females. The alpha male copulates more frequently and with more females at the peak of their cycle, which lasts two to four days (Van Bell, 2006). As a result of the social standing necessary to have the privilege of mating, males often mate later in life than females (Van Bell, 2006). Females seem to have reproductive success at a younger age, and, regardless of social position, copulate multiple times per cycle (Van Bell, 2006).

Van Bell’s (2006) description of the mating behavior of Alouatta pigra begins with the male sniffing the urine of the female and licking the female genitalia to detect her stage in the estrus cycle. To demonstrate interest, both the male and the female flick their tongue in and out of their mouth. Observational studies also show that females may incite mating by grabbing the hairs on a male' face. During the mating period, the pair mutually stay together for a few days and perform multiple copulations. The male usually mounts the female, holding on to her shoulders with his hands, and sometimes will use his feet to grasp as well. Generally, copulation lasts 30 seconds to one minute (Horwich, 1983b). (Horwich, 1983b; Van Bell, 2006)

Sexual maturity is reached at the age of four years old ("Belize Zoo", 2007). However, males from 3 to 4 months of age already have a pale pink pigmented scrotum with descended testes (Horwich, 1983a). Researchers have been unable to detect any visual indications of the female estrus cycle in Alouatta, and there has not been much research on reproductive behavior and biology in general. Most research on reproduction is based on hormones collected from fecal matter in conjunction with information about the ages of individuals from which the fecal samples were collected (Van Bell, 2006). The female estrus cycle lasts from 11 to 24 days, peak time for conception lasts 2 to 4 days (Van Bell, 2006). ("Belize Zoo", 2007; Horwich, 1983a; Van Bell, 2006)

Mexican black howler monkeys do not exhibit seasonality in reproduction, possibly because their diet of leaves and unripe fruits is not seasonal in availability (Crockett & Rudran, 1987). There are slightly fewer births when new leaves and fruits emerge (Van Bell, 2006). Pregnancy lasts about 180 days. Offspring of both sexes often do not stay with their original troop (Kitchen, Horwich, & James, 2004). (Crockett and Rudran, 1987; Kitchen, et al., 2004; Van Bell, 2006)

  • Breeding interval
    Mexican black howler monkeys reproduce yearly.
  • Breeding season
    Mexican black howler monkeys may breed throughout the year.
  • Average number of offspring
    1
  • Average gestation period
    180 days
  • Range age at sexual or reproductive maturity (female)
    3 to 4 years
  • Range age at sexual or reproductive maturity (male)
    3 to 4 years

Mexican black howler monkeys have single births ("Belize Zoo", 2007). Males in this species commit infanticide, possibly with the aim of limiting the sizes of their troops (Knopff & Pavelka, 2006). Females care for their young for 12 months after birth, providing direct care and protection from predators and adult males. Parenting is an individual role, as other members of the troop normally do not help raise offspring. Mothers are rarely far away from their newborns and are watchful for dangers to their young (Treves, Drescher, & Snowdon, 2003). However, as their young grow past the neonatal and infant stage, they are accorded a much greater range of freedom. ("Belize Zoo", 2007; Knopff and Pavelka, 2006; Treves, et al., 2003)

  • Parental Investment
  • altricial
  • pre-fertilization
    • provisioning
    • protecting
      • female
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female
  • pre-independence
    • protecting
      • female
  • post-independence association with parents

Lifespan/Longevity

The average lifespan of Mexican black howler monkeys is 20 years ("Primate Info Net", 2004). ("Primate Info Net", 2004)

  • Average lifespan
    Status: wild
    20 years

Behavior

Mexican black howler monkeys live in groups of 4 to 11 individuals (Estrada, 2004; Gonzalez-Kirchner, 1998). Adults make up about two thirds of a group, and the sex ratio is almost equal, about 4:3, although these numbers can vary quite widely (Estrada et al., 2004). Groups are more terrestrial than those of other howler monkeys. Aside from sleeping, Mexican black howler monkeys spend most of their time resting (about 70%), followed by feeding (18%), moving (5%), and socializing (1.2%) (Behie & Pavelka, 2005). Another study showed that they spend about 77% of their time resting, followed by about 15% feeding, 5% moving, and less than 1% socializing (Behie & Pavelka, 2005). Their long periods of inactivity are to promote the digestion of leaves, as they lack many adaptations common to other folivorous mammals (Kitchen, Horwich, & James, 2004). (Behie and Pavelka, 2005; Estrada, et al., 2004; Kitchen, et al., 2004)

A typical group consists of one dominant alpha male, a few females, their offspring, and occasionally extra males (Treves, Drescher, & Snowdon, 2003). Subordinate males are probably involved in group defense (Kitchen, Horwich, & James, 2004). Occasional lone males have been recorded (Estrada et al., 2004). These solitary males generally are trying to start their own groups (although they have also been found to usurp alpha males from already established groups) and can be used as indications of nascent groups. Groups that are recently formed are generally much smaller than longer-established groups (Gonzalez-Kirchner, 1998). (Estrada, et al., 2004; Gonzalez-Kirchner, 1998; Kitchen, et al., 2004; Treves, et al., 2003)

Generally, Mexican black howler monkeys have densities that range from 6.3 to 89.5 individuals per square km, lower than most other howler monkey species (Estrada et al., 2004; Gonzalez-Kirchner, 1998). Group sizes are also smaller than most other howler monkeys (Knopff & Pavelka, 2006). Current research focuses on which factors, including social considerations, their inactive lifestyle, or intragroup food competition, limit group size. One hypothesis is that infanticide limits group sizes (Knopff & Pavelka, 2006). It is not clear, however, if it is small group size that causes high rates of infanticide, or vice versa. Also, reproductive success decreases when group size increases, naturally limiting group size (Knopff & Pavelka, 2006). (Estrada, et al., 2004; Gonzalez-Kirchner, 1998; Knopff and Pavelka, 2006)

Mexican black howler monkeys uses arboreal quadrupedalism as their main mode of locomotion (Fleagle, 1999). (Fleagle, 1999)

Home Range

The size of the territories is dependent on the number of monkeys in the group. Territory sizes are from 3 to 25 acres and are announced by howling vocalizations. ("Belize Zoo", 2007)

Communication and Perception

Mexican black howler monkeys announce and defend group territories through howling vocalizations ("Belize Zoo", 2007). They are highly social animals Adult howlers are less social, while infant and adolescent howlers engage in most of the social interactions (Behie & Pavelka, 2005). Adults in a group will often howl when approached by one or a few solitary males who may be trying to take over the alpha spot in the troop (Kitchen, Horwich, & James, 2004). Other threats include these solitary males trying to commit infanticide and thus lowering the reproductive success of the current alpha male (Kitchen, Horwich, & James, 2004). Howling can last for over an hour and is low frequency and loud: about 88 decibels at a 5 m distance (Kitchen, Horwich, & James, 2004). The energy used in howling is still less, and the risk smaller, than in physical confrontation. When more than one male is present howling occurs in staggered rounds and, as a result, howling can be used to indicate the number of males present (Kitchen, Horwich, & James, 2004). This information is useful to a group in deciding whether or not to approach another group and initiate a physical confrontation (Kitchen, Horwich, & James, 2004). The howls of females are aurally different from those of males, but they also participate in group howling. ("Belize Zoo", 2007; Behie and Pavelka, 2005; Kitchen, et al., 2004)

Food Habits

Mexican black howler monkey diet is varied. They are strict herbivores, consuming many plant parts, including flowers, fruits, and leaves ("Belize Zoo", 2007). All Alouatta species are highly folivorous. Alouatta pigra is one of the more frugivorous species in this genus. Their preferred food is fruit and they usually eat leaves only in conditions where frugivory is unfavorable (Behie & Pavelka, 2005). This allows them to have a flexible, adaptable diet that can adjust in cases of habitat change. Generally, a little more than 36% of their diet consists of fruits, followed by about 30% mature leaves, 25% new leaves and buds, and 5% flowers (Behie & Pavelka, 2005). Although they consume a significant amount of leaves, Alouatta pigra has a primitive digestive tract more suitable for frugivory; breaking down leaves therefore requires a longer digestion time, which explains the long inactive periods in their activity budget (Kitchen, Horwich, & James, 2004). ("Belize Zoo", 2007; Behie and Pavelka, 2005; Kitchen, et al., 2004)

  • Plant Foods
  • leaves
  • fruit
  • flowers

Predation

Mexican black howler monkeys can howl to frighten away predators or alert others of a predator's presence. Mothers aggressively protect their young for a year against predators and other dangers (Kitchen, Horwich, & James, 2004). The greatest predatory threat to young black howlers is infanticide by adult males. Otherwise, there are few natural predators for Mexican black howler monkeys. Occasionally they are preyed upon by jaguars, pumas, and harpy eagles (Knopff & Pavelka, 2006). Other cats, large birds of prey, and large arboreal snakes are also potential predators (Treves, Drescher, & Snowdon, 2003). Humans have been known to illegally catch individuals for pets, and they are occasionally eaten by humans (Kitchen, Horwich, & James, 2004). (Kitchen, et al., 2004; Knopff and Pavelka, 2006; Treves, et al., 2003)

Ecosystem Roles

Mexican black howler monkeys are often found living in close conjunction with Ateles geoffroyi and other spider monkeys (Estrada et al., 2004). They are also important for the role they play in seed dispersal (Van Bell, 2006). Their frugivorous diet has helped disperse the seeds of some rainforest plants. Although they do not shy away from human contact and their presence is quite obvious due to their howling, Mexican black howler monkeys rarely interfere with human crops. (Estrada, et al., 2004; Estrada, 2006; Van Bell, 2006)

  • Ecosystem Impact
  • disperses seeds

Economic Importance for Humans: Positive

Mexican black howler monkeys are important members of the ecosystems in which they live. Their howling is a unique element of their Mesoamerican forests and their presence can attract ecotourism.

Economic Importance for Humans: Negative

Mexican black howler monkeys occasionally raid crops, although this behavior is rare (Estrada, 2006). (Estrada, 2006)

  • Negative Impacts
  • crop pest

Conservation Status

Mexican black howler monkeys are endangered by hunting and habitat destruction. The forests in which they live and feed are often converted into pastures or agricultural plots. Human influence as a whole is causing significant fragmentation of A. pigra habitats (Estrada, 2006). Other factors resulting in declining populations are: disease, abnormal rainfall, human capture for pets, and natural disasters such as hurricanes (Gonzalez-Kirchner, 1998). In areas affected negatively by humans, Mexican black howler populations become smaller both in numbers and in area (Gonzalez-Kirchner, 1998). They have been listed by the IUCN as an endangered species since 2003. They were previously (as recently as 2000) considered lower risk/least concern (Cuarón et al., 2003). The U.S. Fish and Wildlife Service lists Alouatta pigra as a threatened species. In the next 30 years, the total population is predicted to decrease 74% (Cuarón et al., 2003). Primates from this region have been hunted and captured as pets, causing their populations to decline further (Estrada et al., 2004). (Cuaron, et al., 2003; Estrada, et al., 2004; Estrada, 2006; Gonzalez-Kirchner, 1998)

Other Comments

Alouatta pigra is commonly known as Mexican black howler monkeys, Guatemalan howler monkeys, and saraguatos.

Contributors

Tanya Dewey (editor), Animal Diversity Web.

Kimberly Lau (author), Yale University, Eric Sargis (editor, instructor), Yale University.

Glossary

Neotropical

living in the southern part of the New World. In other words, Central and South America.

World Map

acoustic

uses sound to communicate

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

arboreal

Referring to an animal that lives in trees; tree-climbing.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

chemical

uses smells or other chemicals to communicate

choruses

to jointly display, usually with sounds, at the same time as two or more other individuals of the same or different species

diurnal
  1. active during the day, 2. lasting for one day.
dominance hierarchies

ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates

ecotourism

humans benefit economically by promoting tourism that focuses on the appreciation of natural areas or animals. Ecotourism implies that there are existing programs that profit from the appreciation of natural areas or animals.

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

folivore

an animal that mainly eats leaves.

frugivore

an animal that mainly eats fruit

herbivore

An animal that eats mainly plants or parts of plants.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

polygynous

having more than one female as a mate at one time

rainforest

rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.

riparian

Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

tactile

uses touch to communicate

terrestrial

Living on the ground.

territorial

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

threatened

The term is used in the 1994 IUCN Red List of Threatened Animals to refer collectively to species categorized as Endangered (E), Vulnerable (V), Rare (R), Indeterminate (I), or Insufficiently Known (K) and in the 1996 IUCN Red List of Threatened Animals to refer collectively to species categorized as Critically Endangered (CR), Endangered (EN), or Vulnerable (VU).

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

year-round breeding

breeding takes place throughout the year

References

2007. "Belize Zoo" (On-line). Black Howler Monkey: Alouatta pigra. Accessed May 03, 2007 at http://www.belizezoo.org/zoo/zoo/mammals/how/how1.html.

2004. "Primate Info Net" (On-line). The Life Spans of Nonhuman Primates. Accessed May 08, 2007 at http://pin.primate.wisc.edu/aboutp/phys/lifespan.html.

Behie, A., M. Pavelka. 2005. The Short-Term Effects of a Hurricane on the Diet and Activity of Black Howlers (Alouatta pigra) in Monkey River, Belize.. Folia Primatol., 76: 1-9.

Crockett, C., R. Rudran. 1987. Red howler monkey birth data I: seasonal variation.. American Journal of Primatology, 13: 347-368.

Cuaron, A., P. de Grammont, L. Cortes-Ortiz, G. Wong, J. Silva. 2003. "2006 IUCN Red List of Threatened Species" (On-line). Alouatta pigra. Accessed May 08, 2007 at www.iucnredlist.org.

Estrada, A. 2006. Human and Non-human Primate Co-existence in the Neotropics; a Preliminary View of Some Agricultural Practices as a Complement for Primate Conservation.. Ecological and Environmental Anthropology, 2: 17-29.

Estrada, A., L. Luecke, S. Van Bell, et al.. 2004. Survey of black howler (Alouatta pigra) and spider (Ateles geoffroyi) monkeys in the Mayan sites of Calakmul and Yaxchilan. Primates, 45: 33-39.

Fleagle, J. 1999. Primate Adaptation and Evolution. London: Elsevier Academic Press.

Gonzalez-Kirchner, J. 1998. Group Size and Population Density of the Black Howler Monkey (Alouatta pigra) in Muchukux Forest, Quintana Roo, Mexico.. Folia Primatol., 69: 260-265.

Horwich, R. 1983. Breeding behaviors in black howler monkey (Alouatta pigra) of Belize.. Primates, 24: 222-230.

Horwich, R. 1983. Species Status of the Black Howler Monkey, Alouatta pigra, of Belize. Primates, 24: 288-289.

Kitchen, D., R. Horwich, R. James. 2004. Subordinate Male Black Howler Monkey (Alouatta pigra) Responses to Loud Calls: Experimental Evidence for the Effects of Intra-group Male Relationships and Age.. Behavior, 141: 703-723.

Knopff, K., M. Pavelka. 2006. Feeding Competition and Group Size in Alouatta pigra. International Journal of Primatology, 27: 1059-1078.

Treves, A., A. Drescher, C. Snowdon. 2003. Maternal Watchfulness in Black Howler Monkeys (Alouatta pigra). Ethology, 109: 135-146.

Van Bell, S. 2006. Studies of hormonal cycles, sexual behavior and the phenology of reproduction in howler and spider monkeys.. Accessed May 03, 2007 at http://www.primatesmx.com/fecalhormones.htm.

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