Coquerel's Sifaka

The **Coquerel's sifaka** (*Propithecus coquereli*) is a diurnal, medium-sized lemur of the sifaka genus *Propithecus*. It is native to Madagascar. The Coquerel's sifaka was once considered to be a subspecies of the Verreaux's sifaka, but was eventually granted full species level.

## Description

Coquerel's sifaka is a vertical clinger and leaper with long, powerful hind legs and an upright posture. It has a head-body length of 42–50 cm and a tail length of 50–60 cm. The total mature length (including tail) is approximately 93 to 110 cm. Adult body mass is typically around 4 kg. The dorsal pelage and tail are white, with maroon patches on the chest and portions of the limbs. The coat is generally dense. Its face is bare and black except for a distinctive patch of white fur along the bridge of the nose. Its naked ears are also black, and its eyes are yellow or orange. The bottom of the lemurs hands and feet are black, while the thighs, arms, and chest are a chocolate brown. Like all lemurs, Coquerel’s Sifaka’s have a toothcomb. They use it for grooming and sometimes scraping fruit off a pit.

## Distribution

This species occurs at altitudes of less than 300 ft in the dry deciduous forests of northwestern Madagascar, including coastal forests. It occurs from the Betsiboka River, up to the Maevarano River, and are common in large area between these rivers. A recent study reported its southeast-most presence in Ambalanjanakomby along the Betsiboka River. A extensive survey of the species distribution conducted in 2009, 2010 and 2011 led to the confirmation of its presence in most forest fragments between the two above-mentioned rivers. Nevertheless, its eastern distribution limits are unclear, between the Sofia and Bemarivo rivers, the species has twice been reported to be absent (Table 1). Similarly, the southern part of the inter-river system between the Bemarivo and Betsiboka rivers, where little is known about the presence of the species, requires surveys.

Groups of this species have a home range area amounting to 4-9 hectares. A 2014 line transect distance sampling work in Ankarafantsika National Park suggests that population densities ranges from 5 to 100 ind/km²) and significant (negative) effects of road, and forest edge, and/or a (positive) effect of river proximity on densities. The population size may be ~47,000 individuals in the Ankarafantsika National Park. However, the species is frequently seen around villages and in areas dominated by introduced tree species.

## Diet

Coquerel's sifaka has an herbivorous diet that varies by season. In the wet season, it eats immature leaves, flowers, fruit, bark, and dead wood. In the dry season, it eats mature leaves and buds. It may browse nearly 100 plant species, but the majority of its feeding time will be concentrated on about 10% of these. Since it has a very fibrous diet, Coquerel's sifaka has an enlarged cecum and extremely long colon that helps facilitate digestion. These lemurs spend between thirty and forty percent of their day foraging, especially in the morning, midday, and evening. Females often take leadership roles during foraging and exert their dominance by eating the preferred food or denying the males food until they are satisfied. These lemurs are beneficial to the environment because they aid seed dispersion and serve to populate the plant life. Captive Coquerel’s sifkas eat shining leaf sumac and mimosa.

## Behavior

Coquerel's sifaka lives in matriarchal groups of about three to ten individuals. It is diurnal and primarily arboreal. Much is known about its behavior from observations in the wild and in captivity.

### **Social Structure**

Coquerel’s sifaka spend the majority of its time in areas of just two or three hectares. However, they can live in areas with four to eight. Even though their home range may overlap with other groups of sifaka, they avoid each other to avoid aggression. When friendly Coquerel's sifakas meet, they greet by rubbing their noses together. Matriarchy is rare in the animal kingdom as a whole but common among lemurs. A matriarchal system is particularly pronounced in Coquerel's sifaka. All adult and even most subadult females are dominant over males.

Females have preferential access to food and other resources. When a female is browsing an area or tree, a male waits for her to finish before he moves there to feed. If he gets in the way of the female, she may lunge, smack, or bite him. The male then exhibits submissive behavior by rolling his tail between his legs, chattering softly, and baring his teeth in a grimace before quickly leaping out of her way. When mating, Coquerel's sifaka commonly practices polyandry. A female may choose to mate with only one male, but most often she will mate with several, from other visiting groups as well as from her own. Males compete for access to sexually receptive females. However, the winner of a fight will not necessarily be the one she selects to breed with. The criteria by which she chooses a mate are evidently more complex.

In some other animals, polyandrous mating is thought to raise the chances of successful fertilization, but this does not appear to be the case in Coquerel's sifaka. Instead, polyandry is thought to be advantageous because when paternity is confused, the likelihood of male infanticide decreases.

### **Reproduction**

Female Coquerel's sifakas choose who they mate with whether it be intragroup males or males from outside groups. They have synchronized estrous in January and February. Infants are born in June and July after a gestation period of about 162 days. Normally, one infant is born during Madagascar’s dry season (June–July). Newborn lemurs have an average weight of 100 grams, though it can vary between 85 and 115 grams. An infant will cling to its mother's chest until about a month or so after birth, then transfer to her back. Infants are weaned and become fully independent by about six months of age. Adult size is reached anywhere from one to five years.

Males and females become sexually mature around two- to three-and-a-half years old, though some do not have their first offspring until they are six. Hybrid species have been known to occur with some species. One is the *Propithecus verreauxi*.

### **Locomotion**

In the trees, Coquerel's sifaka moves by vertical clinging and leaping. It maintains an upright posture when at rest or when propelling itself between branches or trunks. This style of arboreal locomotion is characteristic of most, if not all, lemurs. This particular lemur can leap from tree to tree anywhere up to 35 feet. They have the extraordinary ability to leap to spiny trees and precisely place their hands and feet so that they will not hurt themselves.

Occasionally Coquerel's sifaka will descend to the ground to cross open spaces. Its terrestrial locomotion is unique to its species. Like Verreaux's sifaka, it moves in a series of bipedal hops with its arms thrown out to the sides for balance. However, whereas Verreaux's sifaka bounds sideways and crosses its legs one in front of the other, the Coquerel's sifaka bounds forward, like a kangaroo. It leans in the direction of its jump to achieve forward momentum.

A study at Duke University’s Primate Center examined feeding behaviors of captive sifakas to determine their handedness. Given chopped fruits and vegetables, adult male and female sifakas showed a predominant preference for left-handedness, while younger sifakas alternated hands to grab food. The study reveals that Coquerel’s sifakas gain dexterity and hand preference with age, diverging only slightly by gender.

### **Communication**

Coquerel's sifaka uses a variety of auditory, visual, and olfactory signals to communicate. ‘Sifaka' is a Malagasy name that comes from the lemurs’ characteristic “shif-auk” sound. The first syllable is a low growl that "bubbles" in the throat, and the second is a clicking sound like an amplified hiccup. The "shih-*fak*" call is used to warn fellow group members of a potential ground predator or to threaten enemies and intruders. Coquerel's sifaka is highly territorial.

Contact calls used when groups are traveling include soft grunts and growls. If a sifaka is separated from its group members, it may emit a long, loud wail to find them. One visual signal which Coquerel's sifaka uses to communicate is a rapid backward jerking of the head. This is a threatening action which may accompany the "shih-*fak*" call.

Sifakas also rely heavily on scent for communication. Males typically scent-mark using a gland in their throats, which they will rub back and forth along branches. Females are more likely to scent-mark with anogenital glands. It is not entirely clear what information is conveyed in these scents, beyond the demarcation of territory.