Propithecus coquereli
Coquerel's sifaka
(Also: Coquerel's mouse lemur)

Coquerel’s sifakas (Propithecus coquereli) are endemic to Madagascar. This species resides in dry deciduous forests found to the north and east of the Betsiboka River. It has been reported as far north as Bealanana, as far south as Ambato-Boeni, and to the east in the vicinity of Antetemasy. It can be found in Ankarafantsika National Park as well as the Bora Special Reserve. Documented sightings have occurred in the coastal mangroves of Baie de Mahajamba. ("Coquerel's Sifaka", 2011; "Propithecus coquereli", 2008; Pastorini, et al., 2001; Rakotoarisoa, et al., 2006)

Coquerel’s sifaka inhabit old growth and secondary growth forests of mixed deciduous and evergreen trees, from sea level to 300 m above sea level. They also travel through scrub habitat when traveling between fragmented forest patches. ("Coquerel's Sifaka", 2011; "Propithecus coquereli", 2008)

Coquerel’s sifakas have dorsal pelage and limbs that are predominately white. They have large chocolate-brown markings on the front of the arms, thighs and chest, which may differ slightly in size and placement. The fur is quite dense. The back may be a pale silver-gray or brown, while the tail ranges in color from silver-gray to white. Short white hairs cover the muzzle and the face is black. They have small, black ears that protrude through the surrounding fur. Males can be differentiated from females by their gular (throat) gland, which stains the surrounding skin and hair, as well as the dark red-brown color of the perianal skin. Coquerel’s sifaka range in mass from 3.7 to 4.3 kg. ("Coquerel's Sifaka", 2011; Jolly, 1966; Kappeler, 1991; Pastorini, et al., 2001; Rakotoarisoa, et al., 2006; Richard, 2003; Tattersall, 1982)

Coquerel’s sifakas have synchronized estrous which occurs during January and February. Exact timing can be predicted from the two day flushing of the vulva found in females in the precopulatory period. Females mate with intragroup males or males from visiting groups. They appear to be polyandrous, which may serve to confuse paternity and impede male infanticide. Males have been witnessed fighting over access to estrous females, however, the victor isn't always chosen to mate. Coquerel’s sifakas appear to continue reproducing regardless of senescence. The oldest reproducing individual on record was 24 years old, and animals have been known to reproduce the year they die. (Bastian and Brockman, 2007; "Coquerel's Sifaka", 2011; Grieser, 1992; "Propithecus coquereli", 2008; Jolly, 1966; Kappeler, 1991; Richard, 1978; Richard, 2003; Taylor, 2008)

Gestation in Coquerel’s sifakas lasts for approximately 162 days. Typically, a single infant is born during the dry season, which occurs during June and July. Newborns weigh between 85 and 115 g at birth, with an average weight of 100 g. Infants cling to the mother's venter during travel until they are about 1 month old, at which point they move to the dorsum. Infants are weaned during the wet season at approximately 5 to 6 months of age and are completely independent after 6 months. Most individuals reach adult size in 1 to 5 years, depending on habitat conditions and forage availability. Estimated age of sexual maturity for both males and females is reportedly 2 to 3.5 years of age. Females have been known to give birth for the first time at 3 years of age, while others have others have been reported to have their first offspring at the age of 6. Hybridizations can occur between closely related species such as Propithecus verreauxi, which was once considered a subspecies of Coquerel’s sifakas. (Bastian and Brockman, 2007; "Coquerel's Sifaka", 2011; Grieser, 1992; "Propithecus coquereli", 2008; Jolly, 1966; Kappeler, 1991; Richard, 1978; Richard, 2003; Taylor, 2008)

Coquerel's sifakas have a maximum of 2 offspring per group per year, presumably due to the high costs of reproduction. Females give birth every other year and must increase their basal metabolic rate before and during parturition. As with most mammals, the most energetically expensive aspect of reproduction is lactation, which occurs during the dry season and lasts for 5 to 6 months. Although rare, males and juveniles have been observed carrying infants. Information on paternal care is limited, however, the highest ranking male in the group offers limited support to females and their young. (Bastian and Brockman, 2007; "Coquerel's Sifaka", 2011; Grieser, 1992; "Propithecus coquereli", 2008; Jolly, 1966; Kappeler, 1991; Richard, 1978; Richard, 2003; Taylor, 2008)

In the wild, Coquerel's sifakas live between 27 and 30 years. The oldest known individual in the wild was 30 years old. In captivity, they live between 25 and 30 years, and the oldest known captive individual, held at the Duke Lemur Center, lived to be 31 years old. ("Coquerel's Sifaka", 2011; Richard, 2003; Taylor, 2008)

Coquerel’s sifakas are diurnal and live in groups ranging from 3 to 10 individuals. They are mainly arboreal and maintin a vertical posture while using the back legs to leap more than 6 m in a single bound. Their primary mode of locomotion is commonly referred to as “vertical clinging and leaping”. At rest, the body is held vertically pressed to a tree trunk or branch. Occasionally, they descend from the canopy to move between forest fragments or to obtain fallen food. When on the ground, they hop bipedally with their long arms held out for balance. Allo-grooming plays an important role in developing and maintaining social relationships and intragroup hierarchies. Play is often seen during the wet season. ("Coquerel's Sifaka", 2011; "Propithecus coquereli", 2008; Kubzdela, et al., 1992; Richard, 1978; Richard, 2003; Taylor, 2008)

Coquerel’s sifakas are matriarchal. Females commonly remain with their natal group while males are forced to disperse upon reaching maturity. Males often switch groups, and it is uncommon for them to stay with a single group for more than 8 years. Occasionally, females leave their natal group, however, this only occurs if there are large number of unmated adult females. Females are generally dominant to males and may use aggression to secure preferred forage or to regulate sexual admittance. If males do not obey, females may lunge, smack or bite the back of the neck or limbs. Male submission is exhibited via posturing, which includes cowering, rolling his tail between his legs, baring teeth in a grimace, or leaping out of the females way. Males may also exhibit submission through vocalizations, which primarily includes soft calls. Group membership and female dominance does not appear to diminish with age. ("Coquerel's Sifaka", 2011; "Propithecus coquereli", 2008; Kubzdela, et al., 1992; Richard, 1978; Richard, 2003; Taylor, 2008)

Coquerel's sifakas have home ranges between 40,000 and 90,000 m^2, with a core territory of 20,000 to 30,000 m^2. Core territories are used about 60% of the time. ("Propithecus coquereli", 2008; Richard, 1978; Richard, 2003)

Coquerel’s sifakas engage in auditory, visual and olfactory communication. Alarm calls used for aerial predators are often described as roaring barks and growls. General alarm calls, which sound like "sifaka" with an explosive clicking sound at the end, are used to alert group members of terrestrial predators. When separated from their group, Coquerel's sifakas emit a loud, extended wail. Facial expressions and body postures include a play face where the mouth is held open in a silent laugh, and head jerks where the head is thrown quickly back while calling when facing a predator. Polymorphic trichromacy, which allows them to see a full range of colors, was recently discovered in Coquerel’s sifaka. Both males and females use of the anogenital region, the area between the anus and the genitala, for scent marking. Males also use the gular gland for scent marking branches and tree trunks. Both males and females scant mark with urine as well. Males touch the end of the penis to a tree trunk while clinging to it and move up the trunk about 50 cm leaving a line of urine. Vertical trunk marking is less common in females; however, they may press their body to the tree trunk while they climb a short ways up to leave a similar mark as males. The markings are thought to display sex of the marker as well as reproductive status of females since markings greatly increase in frequency during mating season. (Fichtel and P. van Schaik, 2006; Hayes, et al., 2005; Jolly, 1966; Richard, 2003; Veilleux and Bolnick, 2009)

Coquerel’s sifaka eats immature and mature leaves, seeds, flowers, fruit and bark. The majority of their diet consists of leaves, and their teeth are well-adapted for slicing and grinding plant material. Their total diet includes 75 to 100 different plant species; however, 60 to 80% of the time they feed on only 10% of these species. It has been suggested that Coquerel’s sifakas are opportunistic feeders as dominant forage plants change with season. Their enlarged cecum and colon helps facilitate digestion of their highly fibrous diet. Undigested beans have been found in feces, and it has been proposed that nourishment is obtained from the casing rather than the bean itself. Captive individuals at the Duke Lemur Center are primarily fed shining leaf sumac and mimosa. Between 30 to 40% of the day is spent foraging, with peak foraging activity occurring during morning, midday, and late-afternoon. Foraging bouts are separated by rest, and when foraging, they remain within their territories and spend the majority of the time within a core area. The majority of aggression found between sexes is related to feeding. Females commonly exhibit dominance during foraging bouts. Female dominance during feeding likely plays an important role during gestation and lactation. Females usually exercise dominance by controlling access to preferred food or feeding areas by being the first to feed or feeding until satisfied and then allowing males to access the food (Campbell, et al., 2001; "Coquerel's Sifaka", 2011; Hayes, et al., 2005; Jolly, 1966; Kubzdela, et al., 1992; Richard, 1978; Richard, 2003)

Coquerel’s sifakas are preyed upon by hawks and other raptors, constrictor snakes as well as the puma-like fossa, the largest mammalian carnivore found on Madagascar. Aerial predators would be of most danger to the infants. Introduced predators include feral dogs, African wildcats, European wildcats, large Indian civets and Egyptian mongooses. The most imminent threat to Coquerel’s sifakas is humans, which hunt them for food and sport. In the past, the Malagasy people did not hunt sifakas because it was considered “fady” or taboo; however, there are reports that hunger is overpowering this custom. Populations of Coquerel's sifakas that have been hunted in the past flee from humans; if not, they may give a general alarm call. Numerous reports describe being approached by a group of Coquerel’s sifakas on the ground. All members of the group give alarm calls and head jerks while approaching humans. One report documented a group coming within 3 to 5 m. Between alarm calls, they were said to stare and weave their heads back and forth. Roaring barks are made for aerial predators and "sifaka" calls are made for terrestrial predators. Neighboring groups often return alarm calls after searching the local area for potential predators. (Butler, 2008; Fichtel and P. van Schaik, 2006; Jolly, 1966; Richard, 1978)

Coquerel’s sifaka are prey for a number of native and introduced vertebrate predators. As seed predators, Coquerel’s sifaka may help disperse seeds as well. There is no information available regarding parasites of this species. (Jolly, 1966; Richard, 1978)

Coquerel’s sikafas have been the subject of many studies that may provide insight on the evolutionary history of primates, specifically that of humans. They have also been the subject of various research efforts, including those investigating the evolution of color vision, female dominated society, evolution of paternal care, and causes of speciation. Coquerel's sifakas are commonly hunted by the people of Madagascar. In addition, because lemurs are endemic to Madagascar, the emerging eco-tourism industry benefits significantly from their presence. (Grieser, 1992; Kappeler, 1991; Kubzdela, et al., 1992; Mayor, et al., 2004; Pastorini, et al., 2001; Veilleux and Bolnick, 2009)

There are no known adverse effects of Coquerel’s sifaka on humans'.

Coquerel’s sifakas are classified as an endangered species on the IUCN's Red List of Threatened Species. The most immediate threat is habitat loss due to deforestation and slash and burn farming. Trees are either cleared for farming, raising livestock or for charcoal production. In addition to reducing the amount of potential habitat for Coquerel's sifakas, deforestation also reduces forage availability. Hunting pressure is also a major concern. Coquerel's sifakas are currently found in two protected areas: the Ankarafantsika National Park and the Bora Special Reserve. However, illegal hunting is thought to be common in these areas as well. Increased predation by introduced species has negatively impacted this species as well. PAW (Projects for Animal Welfare) of Madagascar was founded in 2011 to combat the threat of introduced cats and dogs. The group is a non-profit that seeks to spay and neuter the population of cats and dogs on the island so that they will not threaten the native wildlife. Coquerel's sifakas are listed under Appendix 1 by CITES (the Convention on International Trade in Endangered Species of Wild Fauna and Flora). (Butler, 2008; "Coquerel's Sifaka", 2011; "Propithecus coquereli", 2008; Richard, 2003)

Coquerel’s sifakas were considered a subspecies of Verreaux’s sifaka. The popular children’s show Zoboomafoo is based on a real Coquerel’s sifaka named Jovian who lives at the Duke Lemur Center. ("Coquerel's Sifaka", 2011; "Propithecus coquereli", 2008; Mayor, et al., 2004)