Overview

Brief Summary

Biology

Verreaux's sifaka is a diurnal, social species existing in groups that are typically 5 members strong (4). These groups are mixed, and females appear to be dominant over males, who are the dispersive sex. The mating system is poorly understood, but in small groups only one female breeds each year. The season occurs in late January and early February and births are 6 months later; during the mating season males may fight between themselves for access to females (6). The single young develops rapidly, initially riding on its mothers belly and moving to her back after a month; young are fully independent at around 6 months of age (6). Groups inhabit home ranges that vary in size depending on the resources available; a core territory within this range is usually defended against neighbouring groups (6).  Sifakas spend the majority of their time in the treetops, travelling via vertical clinging and leaping from one tree to the next (2). This method of locomotion can take them as far as 10 m in one leap (2). To cross open spaces they descend to the ground and adopt what looks like a skipping 'dance' on their hind legs with forearms outstretched for balance (2). The diet varies with season but encompasses a wide range of leaves, fruit, flowers and bark (3). Group members are vigilant for predator attacks, and aerial predators such as the Madagascar harrier-hawk (Polyboroides radiatus) excite different calls than those for predators on the ground (6).
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Description

Verreaux's sifaka is a large lemur; the species is split into 4 subspecies each of which has a distinctive appearance. All members of this species have lustrous creamy white fur but they differ significantly in other markings (6). Verreaux's sifaka (Propithecus verreauxi verreauxi) has a dark brown crown and white tufted ears, in some males the upper chest may also have a reddish-brown tinge, and the face and muzzle of both sexes are very dark (6). Coquerel's sifaka (P. v. coquereli) sports deep maroon thighs and arms, and the colour also extends across the chest. The face is black although a patch of white fur extends down the bridge of the nose; the ears are also black (6). Decken's sifaka (P. v. deckeni) is all white although the fur may sometimes have a golden tinge; the face is black and has a more rounded shape (6). The crowned sifaka (P. v. coronatus) gains its name from its dark chocolate-brown head and throat. The upper body has a golden sheen and the black face is squarer than those of the other subspecies (6). All 4 sifakas have a typical body plan with long, powerful hindlimbs and large hands and feet, allowing them to leap from tree to tree in their arboreal habitat (4). The tail of this species is particularly long and may be used for balance. Sifakas earn their common name from their 'shifak' sounding alarm call (5).
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Comprehensive Description

Verreaux's Sifaka is a large long-tailed lemur with creamy white fur, a dark brown crown, and white tufted ears. Powerful hind limbs and large hands and feet allow sifakas to leap from tree trunk to tree trunk in a type of locomotion called vertical clinging and leaping.

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Distribution

Range Description

This species is endemic to the island of Madagascar. The Tsiribihina River is believed to be the northern limit of the range, ranging as far as Tolagnaro and the Andohahela National Park in the south-east. Sea level to 1,300 m (Mittermeier et al. 2008).
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Geographic Range

Verreaux's sifakas are found in the western and southwestern regions of Madagascar.

Biogeographic Regions: ethiopian (Native )

Other Geographic Terms: island endemic

  • Walker, E. 1968. Mammals of the World. Baltimore: Johns Hopkins Press.
  • Nowak, R. 1999. Walker's Primates of the World. Baltimore: Johns Hopkins University Press.
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Range

Verreaux's sifaka is endemic to Madagascar and the 4 subspecies occur in apparently isolated and distinct ranges in the south and west of the island (2). Coquerel's sifaka has the northernmost distribution along the northwest coast; next is the crowned sifaka, the most endangered of the 4 subspecies and inhabiting the smallest range (8). Decken's sifaka occurs in a band in the west of the country and it is possible that in its northern limit this species is sympatric with the crowned sifaka (6). P. v. verreauxi has the widest distribution of any of the sifakas of Madagascar and is found in the remaining forests in the south and southwest of the island, and west to the Tsiribihina River (6).
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Physical Description

Morphology

Physical Description

Verreaux's sifakas have distinct coloration with white fur and a hint of yellow contrasting their hairless, black face. Their hands and feet are also black. They have a long tail that ranges from 43 to 56 cm in length, roughly the same length as their body (45 to 55 cm). Verreaux's sifakas are not sexually dimorphic, as both males and females weigh between 3 and 7 kg.

Range mass: 3 to 7 kg.

Range length: 45 to 55 cm.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: sexes alike

Average basal metabolic rate: 3.738 W.

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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
This is a diurnal lemur that inhabits tropical dry lowland and montane forest, including spiny bush, brush-and scrub thickets, and riparian forests, and is also recorded in lowland rainforest in the south-east. It tends to live in small to medium multi-male groups that range from 2-14 (average five to six individuals), with home ranges sometimes exceeding 10 ha. Breeding is seasonal, with mating taking place in January and February. Infants are almost completely independent at six months (Mittermeier et al. 2008, and references therein). The age at which sexual maturity is reached varies with habitat. For example, in the spiny forests of Beza Mahafaly fewer than half the females have reproduced by six years of age (Richard et al. 2002), but three-year-old females are routinely seen with newborns at Berenty (Jolly 1966).

Systems
  • Terrestrial
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Verreaux's sifakas are primarily arboreal and are found in deciduous and evergreen forests. However, they are widespread and can also be found in wet and dry habitats throughout southwest Madagascar.

Habitat Regions: tropical ; terrestrial

Terrestrial Biomes: forest ; rainforest ; scrub forest

  • Dewar, R., A. Richard. 2007. Evolution in the Hypervariable Environment of Madagascar. Proceedings of the National Academy of Sciences of the United States of America, 104/34: 13723-13727.
  • Wade, D. 1996. Deforestation and Its Effects in Highland Madagascar. Mountain Research and Development, 16/2: 101-116.
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This species is characterized by four distinct subspecies that occur in apparently isolated and distinct ranges in the southern and western parts of Madagascar. Verreaux's Sifakas prefer habitats with large trees in various forest types from dry, spiny deciduous woodlands to evergreen forests.

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Inhabits spiny and dry deciduous forests (2).
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Trophic Strategy

Food Habits

Verreaux's sifakas are herbivorous and feed primarily on leaves, bark, and flowers. When abundant, fruit may also be incorporated into their diet. Nutrition levels directly coincide with season and rainfall amounts, with food of higher nutritional value being most abundant during the wet season.

Plant Foods: leaves; wood, bark, or stems; fruit; flowers

Primary Diet: herbivore (Folivore , Lignivore)

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Associations

Ecosystem Roles

Verreaux’s sifakas are an important prey item for fossas (Cyptoprocta ferox), stray dogs (Canis lupus familaris), and harrier hawks (Polyboroides radiatus). Also, they are strict herbivores and considered to be important seed dispersers.

Ecosystem Impact: disperses seeds

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Predation

Verreaux's sifakas commonly use alarm calls to warn group members of the presence of a predator. Different alarm calls are given for aerial (harrier hawk, Polyboroides radiatus) and terrestrial predators, which includes fossas (Cyptoprocta ferox) and stray dogs (Canis lupus familaris). Alarm calls are performed by males and females of all ranks in the social hierarchy and are thought to be most effective in group settings. As group size increases the number of individuals able to spot potential predators increases. As a result, group cohesion is an important aspect of deterring predators. It is also thought that large groups are able to intimidate potential predators more easily than small groups.

Known Predators:

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Life History and Behavior

Behavior

Communication and Perception

Verreaux's sifakas communicate over long distances via clear, deep barks. Barks sound similar to the word “sifaka” and are produced only when intruders are nearby. Barks are generally made by the group leader, and if group cohesion is ever threatened by an outsider, growling or barking is produced to ward off intruders.

Scent marking is an additional form of communication used by Verreaux's sifakas. It serves multiple purposes including marking territory, making one's presence known (specifically females in estrus), claiming food or territory, attracting mates, and may be used during non-physical competition. Scent marking is a versatile and important way of communicating in Verreaux's sifakas.

Communication Channels: acoustic ; chemical

Other Communication Modes: scent marks

Perception Channels: visual ; acoustic

  • Palagi, E., D. Antonacci, I. Norscia. 2008. Peacemaking on Treetops: First Evidence of Reconciliation from a Wild Prosimian (Propithecus verreauxi). Animal Behaviour, 76/3: 737-747.
  • Trillmich, J., C. Fichtel, P. Kappeler. 2004. Coordination in Group Movements in Wild Verreaux's Sifakas (Propithecus verreauxi). Behaviour, 141/9: 1103-1120.
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This species is primarily a frugivore or fruit eater, but its diet changes with the seasons. In the rainy season, Verreaux’s Sifakas eat more fruits and flowers, whereas in the dry season they consume mainly dead wood, bark, and leaves – a habit which may help them absorb more water.

Verreaux's sifaka is a social species living in groups of 3-13 members. These groups are mixed and females appear to be dominant over males. Breeding females give birth to a single black, hairless offspring which clings to its mother's belly for 3 to 4 weeks, before moving to her back.

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Life Expectancy

Lifespan/Longevity

Little is known about the lifespan of Verreaux's sifakas in the wild, as the residents of Madagascar rarely come into contact with them. Studies have found that they have a surprisingly low parasitic load, which may result in increased lifespan relative to other closely related primates. The average lifespan of captive Verreaux's sifakas is 18 years with a recorded maximum of 23.5 years.

Range lifespan

Status: captivity:
23.5 (high) years.

Average lifespan

Status: captivity:
18 years.

Average lifespan

Status: captivity:
18 years.

Average lifespan

Sex: male

Status: captivity:
20.6 years.

Average lifespan

Status: captivity:
18.2 years.

Average lifespan

Status: captivity:
18.0 years.

Average lifespan

Sex: female

Status: captivity:
18.2 years.

  • Muehlenbein, M., M. Schwartz, A. Richard. 2003. Parasitologic Analyses of the Sifaka (Propithecus verreauxi verreauxi) at Beza Mahafaly, Madagascar. Journal of Zoo and Wildlife Medicine, 34/3: 274-277.
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Lifespan, longevity, and ageing

Maximum longevity: 30.5 years (captivity) Observations: One male of the *coquereli* subspecies lived 30.5 years at the Duke University Primate Center (Richard Weigl 2005). Infant survival of old females drops, but a decline in fertility has been reported in some of the oldest females (Pochron et al. 2004).
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Reproduction

Verreaux's sifakas form social hierarchies, which are subject to change during the mating season as females only breed with dominant males. Subordinate males challenge dominant males, often resulting in intense competition and fighting.

Verreaux's sifakas scent mark as a form of communication. Females scent mark to get the attention of males. Subordinate males scent mark to get the attention of females, and dominant males scent-mark to claim territory. Scent-marking often results in conflict among males.

Mating System: polygynous

Verreaux's sifakas breed from late January through March. Following implantation, gestation lasts 130 to 141 days. Typically, mothers give birth to only one offspring per year, which occurs between June and September. Average birth weight is 40 g. Mothers carry young close to their abdomen and chest for the first 2 to 3 months of the offspring's life. At about 3 months old, young move to their mothers back until they reach 5 to 6 months of age and are weaned at about six months. Young sifakas reach adult size at 21 months and are sexually mature by two and a half years old.

Breeding interval: Once a year

Breeding season: January to March

Average number of offspring: 1.

Range gestation period: 4 to 5 months.

Average birth mass: 40 g.

Average weaning age: 180 days.

Average time to independence: 2.5 years.

Average age at sexual or reproductive maturity (female): 2.5 years.

Average age at sexual or reproductive maturity (male): 2.5 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); viviparous

Average birth mass: 72.25 g.

Average number of offspring: 1.

Average age at sexual or reproductive maturity (male)

Sex: male:
912 days.

Average age at sexual or reproductive maturity (female)

Sex: female:
912 days.

Verreaux's sifakas live in social groups and young are cared for by adults. A mother's position in the social hierarchy affects the social status of her young and parental care continues until young reach full size at about 21 months. Females stay with the group, while males either stay with the group or leave to form their own. Mothers carry newborn sifakas near their chest and abdomen for the first 2 to 3 months after birth and then on her back until offspring reach about 6 months of age.

Parental Investment: male parental care ; female parental care ; post-independence association with parents; extended period of juvenile learning; maternal position in the dominance hierarchy affects status of young

  • Nowak, R. 1999. Walker's Primates of the World. Baltimore: Johns Hopkins University Press.
  • Lewis, R. 2005. Sex Differences in Scent-Marking in Sifaka: Mating Conflict or Male Services?. American Journal of Physical Anthropology, 128/2: 389-398.
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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
VU
Vulnerable

Red List Criteria
A2cd

Version
3.1

Year Assessed
2008

Assessor/s
Andrainarivo, C., Andriaholinirina, V.N., Feistner, A., Felix, T., Ganzhorn, J., Garbutt, N., Golden, C., Konstant, B., Louis Jr., E., Meyers, D., Mittermeier, R.A., Perieras, A., Princee, F., Rabarivola, J.C., Rakotosamimanana, B., Rasamimanana, H., Ratsimbazafy, J., Raveloarinoro, G., Razafimanantsoa, A., Rumpler, Y., Schwitzer, C., Thalmann, U., Wilmé, L. & Wright, P.

Reviewer/s
Mittermeier, R.A. & Rylands, A.B. (Primate Red List Authority)

Contributor/s

Justification
Listed as Vulnerable as the species is thought to have undergone a reduction of more than 30% over the past 30 years (assuming a generation length of 10 years) due primarily to a decline in area and quality of habitat within the known range of the species and due to levels of exploitation.

History
  • 2000
    Vulnerable
  • 1996
    Vulnerable
  • 1990
    Vulnerable
    (IUCN 1990)
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Verreaux's sifakas are rapidly losing their natural habitat. Due to slash-and-burn agriculture, the deciduous forests of Madagascar are severely threatened. Additionally, Malagasy forests have been subjected to commercial logging, overgrazing by livestock, and charcoal manufacturing. Fortunately, captive breeding has been successful in Verreaux's sifakas, which, according to the IUCN Red List, are considered "vulnerable".

US Federal List: no special status

CITES: appendix i

State of Michigan List: no special status

IUCN Red List of Threatened Species: vulnerable

  • Loudon, J., M. Sauther, K. Fish, M. Hunter-Ishikawa, Y. Ibrahim. 2006. One Reserve, Three Primates: Applying a Holistic Approach to Understand the Interconnections Among Ring-Tailed Lemurs (Lemur catta), Verreaux's Sifaka (Propithecus verreauxi), and Humans (Homo sapiens) at Beza Mahafaly Special Reserve, Madagascar. Ecological and Environmental Anthropology, 2/2: 54-74.
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IUCN Red List Status: VULNERABLE

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Status

Classified as Vulnerable (VU - A2cd) on the IUCN Red List 2004 (1), and listed on Appendix I of CITES (7). Subspecies: Verreaux's sifaka (Propithecus verreauxi verreauxi) classified as Vulnerable (VU-A2cd); Coquerel's sifaka (P. v. coquereli) classified as Endangered (EN - A2c, B1+2bc); Decken's sifaka (P. v. deckeni) classified as Vulnerable (VU - A1c); crowned sifaka (P. v. coronatus) classified as Critically Endangered (CR - B1+2bc, C2a) (1).
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Population

Population
Significant variation in population densities has been noted in different forest types, but even very small forest patches can support sizeable populations of this sifaka. Population densities have been estimated at 47 individuals/km² in the degraded forests of Bealoka, at 150-200 individuals/km² at Berenty (Jolly et al. 1982; O’Connor 1987) and at 860 individuals/km² at Antserananomby; however, the population density at the last mentioned site had declined to about 49/km² as of 2004 (Kelley et al. 2007).

Population Trend
Decreasing
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Threats

Major Threats
Despite this species' wide distribution, the two principal habitats upon which it depends for survival -- spiny desert and riparian forest -- are under threat due to the need for timber, charcoal and fuelwood (Sussman and Richard 1986). Although hunting of P. verreauxi is fady (taboo) to several of the tribes living in its range (e.g., Antandroy, Mahafaly), it is hunted by other tribes (e.g., Sakalava) and immigrants to the region (Goodman and Raselimanana 2003). In the Isalo region, this lemur is known as sifaka-bilany or “sifaka of the cooking pot,” but it is unclear whether this is because of its popularity as a food item or because of the sooty black appearance of individuals from this part of the species’ range (Mittermeier et al. 2008).
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These engaging lemurs are threatened by habitat destruction throughout their range. The dry deciduous forests of the west are being cleared for timber extraction, firewood and charcoal production and resulting fragments of habitat are also at risk from deliberate fires (6). Hunting of the species is a taboo in some areas but does still occur in others and is made easier by the relatively open habitat (6). Of the 4 subspecies, the crowned sifaka is the most endangered and population estimates vary between just 100 and 1,000 individuals (1).
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Management

Conservation Actions

Conservation Actions
This species is listed on Appendix I of CITES. This species occurs in five national parks (Andohahela, Isalo, Kirindy-Mitea, Tsimanampetsotsa, and Zombitse-Vohibasia), two special reserves (Andranomena and Beza-Mahafaly), two private reserves (Analabe and Berenty), the Kiridindy CFPF, and a number of unprotected forests (Mittermeier et al. 2008). As of 2007, there are no animals held in captivity.
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Conservation

Verreaux's sifaka is protected from international trade by its listing on Appendix I of the Convention on International Trade in Endangered Species (CITES) (1). Individuals have been successfully bred in captivity (4), and there are estimated to be around 24 of these sifakas currently held in captivity in the United States and Europe, although which subspecies these represent is uncertain (8). Within Madagascar, P. v. verreauxi is known to occur in at least 7 protected areas, Coquerel's and Decken's sifakas in at least 2, but at present the crowned sifaka has not been confirmed from any of the protected areas within its range (6). More data is urgently needed on the distribution and natural ecology of the lesser-studied subspecies and protected reserves are vital to preserve the remaining, if fragmented, habitat of the species as a whole.
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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Negative

There are no known adverse effects of Verreaux's sifakas on humans.

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Economic Importance for Humans: Positive

Verreaux’s sifakas are the most extensively studied of the Malagasy prosimians and have been the subject of a wide range of research topics. These topics include, but are not limited to bipedalism in mammals, the evolution of social hierarchies, the causes and consequences of female dominance in mammals, the evolution and ecological consequences of male dimorphism, the causes and consequences of decreased parasitic loads, and the evolution of characteristics that are necessary for living in the variable environments of Madagascar. Verreaux's sifakas are also an important component of the unique Madagascar ecosystem, which attracts tourists from around the globe.

Positive Impacts: ecotourism ; research and education

  • Lewis, R., P. Kappeler. 2005. Seasonality, Body Condition, and Timing of Reproduction in Propithecus verreauxi verreauxi in the Kirindy Forest. American Journal of Primatology, 67/3: 347-364.
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Wikipedia

Verreaux's sifaka

Verreaux's sifaka (Propithecus verreauxi), or the white sifaka, is a medium sized primate in one of the lemur families, Indriidae. It lives in Madagascar and can be found in a variety of habitats from rainforest to western Madagascar dry deciduous forests and dry and spiny forests. The fur is thick and silky and generally white with brown on the sides, top of the head, and on the arms. Like all sifakas, it has a long tail that it uses as a balance when leaping from tree to tree. However, its body is so highly adapted to an arboreal existence that on the ground its only means of locomotion is hopping. The species lives in small troops which forage for food.

There are four sub-species of this kind of lemur. There are many things unknown about Verreaux's sifaka, so their life span in the wild has not been approximated, but in captivity they generally live to up to 18 years old.[citation needed]

Anatomy[edit]

In adulthood the full head and body length is between 42 and 45 cm (17 and 18 in). The tail of a fully grown Verreaux's sifaka grows to be between 56 and 60 cm (22 and 24 in) long. In terms of weight, adult females reach 3.4 kg (7.5 lb) on average, and adult males 3.6 kg (7.9 lb).[citation needed]

Verreaux's sifaka has a relatively low, flat braincase. The face is broader than that of most other Indriids, but its snout is reduced. This species of sifaka is also distinguished by its unique dentition. Its dental formula is 2.1.2.32.0.2.3. The upper incisors are very small and are slightly angled inward towards the gap between I1 and I2. In the mandible, Verreaux's sifaka displays the stresirhine characteristic: the toothcomb. Formed by the procumbent lower incisor and canine, the toothcomb projects past the front margin of the mouth. P. verreauxi also presents the high, shearing molar crests of a folivore, helping to shread the leaves, fruit, and flowers that it eats. Postcranially, Verreaux's Sifaka has a low intermembral index that ranges from 63-66. It has a broader ribcage than most other prosimians and has many lumbar vertebrae lending it a lot of flexibility. The pelvic is high and narrow and the acetabulum is relatively shallow, also allowing for greater flexibility. Like other Indriids, P. verreauxi has a short calcaneus, pointed nails, and slightly webbed hands and feet.[citation needed]

Diet[edit]

Verreaux's sifaka is mostly folivorous

Verreaux's sifaka forage for food in the troop it lives in, primarily in the morning and late afternoon, so they can rest during the hottest part of the day. They are herbivores; leaves, fruit, bark and flowers are typical components of the diet. However, they are mostly folivorous (leaves represent the majority of the diet over the year, especially in the dry season) and they seem to choose food items based on quality (lower tannin content) rather than on availability.[3]

Behaviour[edit]

Verreaux's sifaka are diurnal and arboreal, and engage in sunbathing with outstretched arms and legs. Verreaux's sifaka move through the trees by clinging and leaping between vertical supports. They are capable of making remarkable leaps through the trees - distances of 9-10m are not uncommon. On the ground, they hop bipedally.[citation needed] They live in family groups, or troops, of 2-12, which may consist of one male and female, or many males and females together. Group and population sex ratio can be more or less skewed toward males.[4][5] Many groups seem to be effectively harem groups with a single dominant male unrelated with resident female(s).[6] They have a home range of 2.8.5ha, and although they are territorial, it is the food source they will defend rather than the territory's boundaries, as often boundaries overlap. Females are dominant over males, forming a matriarchal society.

Females use anogenital secretion mainly for territory demarcation whereas males seem to use specialized secretions (via anogenital and throat glands) more for sexual "advertisement" than for territorial purposes.[7][8] Males show bimorphism, by showing either a clean or stained chest, derived from throat gland secretions and smeared on surfaces by rubbing the upper part of the chest.[9] Stain-chested males engage in the most active marking, and chest staining seems to be related to testosterone levels.[10]

Males and females were found to engage in a biological market, exchanging grooming for grooming during the non-mating period, and grooming ("offered" by males) for reproductive opportunities (sexual access "offered" by females) during the mating period.[11] A study found that females copulate more with stained-chested than with clean-chested males. On the other hand, clean-chested males, with a lower scent-releasing potential, usually offer more grooming to females. This “grooming for sex” tactic allows males with a clean chest to get to copulate with females, even if at low rate.[12]
It has also been discovered that sifaka dyads often engage in post-conflict reunions after aggressive episodes: reconciliation occurs more frequently when food is not involved and for low intensity aggressions.[13] In this species play behavior persists into adulthood where it is used, especially by stranger males during the mating period, as an ice-breaking mechanism to reduce xenophobia [14]

Reproduction[edit]

Females give birth to one infant after a gestation period of 130 days, between June and August. For the first 6–8 weeks, the infant clings to the mother's stomach, but for the following 19 weeks, it clings to her back.[citation needed]

Ecology and Conservation status[edit]

Currently this species is considered to be Vulnerable by the IUCN. In the small spiny forest fragments of South Madagascar, sifaka abundance appears to be influenced by the proportion of large trees (diameter at breast height >=5 cm) and by the abundance of the plant species Allouadia procera,[15] a key species of the spiny forest habitat.[16]

References[edit]

  1. ^ Groves, C. P. (2005). Wilson, D. E.; Reeder, D. M, eds. Mammal Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. p. 121. OCLC 62265494. ISBN 0-801-88221-4. 
  2. ^ Andriaholinirina, N., Baden, A., Blanco, M., Chikhi, L., Cooke, A., Davies, N., Dolch, R., Donati, G., Ganzhorn, J., Golden, C., Groeneveld, L.F., Irwin, M., Johnson, S., Kappeler, P., King, T., Lewis, R., Louis, E.E., Markolf, M., Mass, V., Mittermeier, R.A., Nichols, R., Patel, E., Rabarivola, C.J., Raharivololona, B., Rajaobelina, S., Rakotoarisoa, G., Rakotomanga, B., Rakotonanahary, J., Rakotondrainibe, H., Rakotondratsimba, G., Rakotondratsimba, M., Rakotonirina, L., Ralainasolo, F.B., Ralison, J., Ramahaleo, T., Ranaivoarisoa, J.F., Randrianahaleo, S.I., Randrianambinina, B., Randrianarimanana, L., Randrianasolo, H., Randriatahina, G., Rasamimananana, H., Rasolofoharivelo, T., Rasoloharijaona, S., Ratelolahy, F., Ratsimbazafy, J., Ratsimbazafy, N., Razafindraibe, H., Razafindramanana, J., Rowe, N., Salmona, J., Seiler, M., Volampeno, S., Wright, P., Youssouf, J., Zaonarivelo, J. & Zaramody, A. (2014). "Propithecus verreauxi". IUCN Red List of Threatened Species. Version 2014.1. International Union for Conservation of Nature. Retrieved 2014-06-16. 
  3. ^ Norscia, I., Carrai, V., Borgognini-Tarli, S.M. (2006). "Influence of Dry Season and Food Quality and Quantity on Behavior and Feeding Strategy of Propithecus verreauxi in Kirindy, Madagascar". International Journal of Primatology 27: 1001–1022. doi:10.1007/s10764-006-9056-x. 
  4. ^ Richard, A.F. (1985). "Social boundaries in a Malagasy prosimian, the Sifaka (Propithecus verreauxi)". International Journal of Primatology 6: 553–568. doi:10.1007/BF02692288. 
  5. ^ Norscia, I., Palagi P. (2008). "Berenty 2006: Census of Propithecus verreauxi and possible evidence of population stress.". International Journal of Primatology 29: 1099–1115. doi:10.1007/s10764-008-9259-4. 
  6. ^ Kappeler, P.M., Schäffler, L. (2007). "The lemur syndrome unresolved: extreme male reproductive skew in sifakas (Propithecus verreauxi), a sexually monomorphic primate with female dominance". Behavioral Ecology and Sociobiology 62: 1007–1015. doi:10.1007/s00265-007-0528-6. 
  7. ^ Lewis, R.J. (2005). "Sex differences in scent-marking in sifaka: Mating conflict or male services?". American Journal of Physical Anthropology 128 (2): 389–398. doi:10.1002/ajpa.20206. PMID 15795894. 
  8. ^ Lewis R.J. (2006). "Scent marking in Sifaka : No one function explains it all". American Journal of Primatology 68 (6): 622–636. doi:10.1002/ajp.20256. PMID 16715510. 
  9. ^ Lewis R.J. and van Schaik C.P. (2007). "Bimorphism in Male Verreaux’s Sifaka in the Kirindy Forest of Madagascar". International Journal of Primatology 28: 159–182. doi:10.1007/s10764-006-9107-3. 
  10. ^ Lewis R.J. (2009). "Chest Staining Variation as a Signal of Testosterone Levels in Male Verreaux's Sifaka". Physiology & Behavior 96: 586–592. doi:10.1016/j.physbeh.2008.12.020. 
  11. ^ Norscia, I., Antonacci, D., Palagi, E. (2009). "Mating First, Mating More: Biological Market Fluctuation in a Wild Prosimian". In Brosnan, Sarah Frances. PLoS ONE 4 (3): e4679. doi:10.1371/journal.pone.0004679. PMC 2650411. PMID 19262737. 
  12. ^ Dall'Olio S., Norscia I., Antonacci D., Palagi E. (2012). "Sexual Signalling in Propithecus verreauxi: Male "Chest Badge" and Female Mate Choice". In Proulx. PLoS ONE 7 (5): e37332. doi:10.1371/journal.pone.0037332. 
  13. ^ Palagi, E., Antonacci, D., Norscia, I. (2008). "Peacemaking on treetops: first evidence of reconciliation from a wild prosimian (Propithecus verreauxi)". Animal Behavior 76: 737–747. doi:10.1016/j.anbehav.2008.04.016. 
  14. ^ Antonacci, D., Norscia, I., Palagi, E. (2010). "Stranger to Familiar: Wild Strepsirhines Manage Xenophobia by Playing". In Iwaniuk, Andrew. PLoS ONE 5 (10): e13218. doi:10.1371/journal.pone.0013218. PMC 2951354. PMID 20949052. 
  15. ^ Norscia, I., Palagi, E. (2011). "Fragment quality and distribution of the arboreal primate Propithecus verreauxi in the spiny forest of south Madagascar". Journal of Tropical Ecology 27: 103–106. doi:10.1017/S0266467410000519. 
  16. ^ Elmqvist, T., Pyykonen, M., Tengo, N., Rakotondrasoa, F., Rabakonandrianina, E., Radimilahy, C. (2007). "Patterns of Loss and Regeneration of Tropical Dry Forest in Madagascar: The Social Institutional Context". In Somers, Michael. PLoS ONE 2(5): e403. doi:10.1371/journal.pone.0000402. 
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