Coquerel's sifaka

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Coquerel's sifaka
Propithecus coquereli 02.jpg
CITES Appendix I (CITES) [2]
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Strepsirrhini
Family: Indriidae
Genus: Propithecus
Species:
P. coquereli
Binomial name
Propithecus coquereli
Propithecus coquereli range map.svg
Distribution of P. coquereli [1]
Synonyms
  • damonisGray, 1870

Coquerel's sifaka (Propithecus coquereli) is a diurnal, medium-sized lemur of the sifaka genus Propithecus . It is native to northwest Madagascar. Coquerel's sifaka was once considered a subspecies of Verreaux's sifaka but was eventually granted full species status. It is listed as Critically Endangered on the IUCN Red List due to habitat loss and hunting.

Contents

In popular culture, the title character in the children's TV show Zoboomafoo is a Coquerel's sifaka. The species was named after French entomologist Charles Coquerel.

Description

The dorsal pelage and tail of Coquerel's sifaka are white, and it has 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. [4] [5] The bottom of the lemur's hands and feet are black, while the thighs, arms, and chest are chocolate brown. Like all lemurs, Coquerel's sifaka has a toothcomb, used for grooming and sometimes scraping fruit off a pit. [6]

Distribution

This species lives at altitudes less than 300 ft (91 m) in the dry deciduous forests of northwestern Madagascar, including coastal forests. [7] It is found from the Betsiboka River up to the Maevarano River, and in large area between these rivers. Extensive surveys of the species' distribution conducted in 2009, 2010, and 2011 led to the confirmation of its presence in most forest fragments between these rivers. [7] Nevertheless, its eastern distribution limits are unclear. Between the Sofia and Bemarivo Rivers, the species has twice been reported to be absent (Table 1 [7] ). 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. [7]

Map of P. coquereli distribution Map of Propithecus coquereli distribution.tif
Map of P. coquereli distribution

Groups of this species have a home range area amounting to 4–9 ha (9.9–22.2 acres). [8] A 2014 work [9] in Ankarafantsika National Park suggests that population densities range from 5 to 100/km2 (13 to 259/sq mi) 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. [9] However, the species is frequently seen around villages and in areas dominated by introduced tree species. [7]

Diet

This species' herbivorous diet 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. [10] It may browse nearly 100 plant species, but the majority of its feeding time is concentrated on about 10% of these. [11] Since it has a very fibrous diet, Coquerel's sifaka has an enlarged cecum and extremely long colon that helps facilitate digestion. [12] These lemurs spend 30-40% of their day foraging, especially in the morning, midday, and evening. Females often lead 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. Captive Coquerel's sifakas eat shining leaf sumac and mimosa. [13]

Behaviour

Coquerel's sifaka lives in matriarchal groups of about three to ten individuals. [7]

Clinging to tree trunk Sifaka in Madagascar (cropped).jpg
Clinging to tree trunk

It is diurnal and primarily arboreal. Much is known about its behavior from observations in the wild and in captivity. [12]

Social structure

Coquerel's sifakas spend the majority of their time in areas of just 2–3 ha (4.9–7.4 acres). However, they can live in areas with 4–8 ha (9.9–19.8 acres). Though their home range may overlap with other groups of sifakas, they avoid each other to avoid aggression. [14] 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. [12]

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. [12]

When mating, Coquerel's sifaka commonly practices polyandry. A female may choose to mate with only one male, but most often she mates 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 for breeding. The criteria by which she chooses a mate are evidently more complex. [6]

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. [11]

Reproduction

Female Coquerel's sifakas choose their mate(s). [15] They have synchronized estrus in January and February. [12] Infants are born in June and July after a gestation period around 162 days. Normally, one infant is born during Madagascar's dry season (June–July). Newborn lemurs have an average weight of 100 g (3.5 oz), though it can vary between 85 and 115 g (3.0 and 4.1 oz). An infant clings to its mother's chest until about a month or so after birth, then transfers to her back.

Infants are weaned and become fully independent around six months of age. Adult size is reached at one to five years. [10]

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. Hybrids have been known to occur with some species. One is P. verreauxi . [16]

Locomotion

Coquerel's sifaka in the wild at Anjajavy Forest Coquerel's sifaka propithecus coquereli.jpg
Coquerel's sifaka in the wild at Anjajavy Forest
The terrestrial locomotion of Coquerel's sifaka Coquerel's sifaka lemur (Propithecus coquereli) terrestrial locomotion.jpg
The terrestrial locomotion of Coquerel's sifaka

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. [12] This particular lemur can leap from tree to tree up to 35 feet (11 m). [17] It has the extraordinary ability to leap to spiny trees and precisely place its hands and feet so that it will not hurt itself.

Occasionally, Coquerel's sifaka descends 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. [10]

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. Coquerel's sifakas gain dexterity and hand preference with age, diverging only slightly by gender. [18]

Communication

Coquerel's sifaka uses a variety of auditory, visual, and olfactory signals to communicate. [19] "Sifaka" is a Malagasy name that comes from the lemurs' characteristic "shif-auk" sound. [20] 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. [21]

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. [11]

One visual signal which Coquerel's sifaka uses to communicate is a rapid backward jerking of the head. This is a threatening action that may accompany the "shih-fak" call. [10]

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. [11]

A study of sifaka vocalizations found that roaring barks are associated with anti-raptor responses, in which the Coquerel's sifakas looked up and climbed down. [22] The "tchi-fak" vocalizations was associated with anti-terrestrial responses, in which the sifakas looked down and climbed up. [22] The meaning of growls seemed to vary by population, where a population subject to significant raptor predation associated the growls with anti-raptor responses, but another population associated growls with mild disturbance. [22]

Scientific importance

Coquerel's sifakas, like many lemurs, have been studied to help scientists learn about the evolutionary history of primates, including humans. They have been the subject of those researching evolution of color vision, paternal care, matriarchal primate societies, and causes of speciation. [12]

Conservation status and threats

Coquerel's sifaka is found in only two protected areas in Madagascar, the Ankarafantsika National Park and the Bora Special Reserve. It is a critically endangered species, according to the IUCN's Red List of Threatened Species, and it is listed in CITES Appendix I. [2] The principal threats to its existence are deforestation, habitat fragmentation, and hunting pressure. The local people often clear trees to produce new farming land, especially in the marshes where rice can be grown. In northwestern Madagascar, deforestation results from annual burning to create new pastureland for livestock. Trees are also cut for the production of charcoal. [14]

Many local Malagasy traditions prohibit hunting of Coquerel's sifaka. One such taboo derives from a legend of a sifaka saving the life of a boy who has fallen out of a tree. The story goes like this:

A little boy heads into the forest to find some honey. He spots a hive in a high tree and he ascends it. As he is about to reach in to collect the honey, he is immediately attacked by bees. The surprise causes the boy to lose his grip on the tree branch, and he falls to what is almost certainly his death. As the boy plummets toward the earth, a large lemur suddenly appears, swoops in, and catches the boy, saving his life. Ever since that day, lemurs became sacred to the Malagasy and it is said that anyone who kills one shall have extreme misfortune. [23]

These protective taboos are breaking down with cultural erosion and immigration. [14]

This lemur is now hunted for bushmeat, but humans are not the only threat. The introduction of foreign species, especially cats and dogs, has hurt the Coquerel's sifaka. Projects for Animal Welfare encourages the neutering and spaying of the cats and dogs on the island to protect the native wildlife. [1] Even the protected areas where the Coquerel's sifaka occurs offer it little protection. It is hunted even within Ankarafantsika, and the Bora Special Reserve has become seriously degraded. [10]

Predation

Many animals prey on Coquerel's sifakas. Hawks and other raptors attack them from above, while constrictor snakes and the fossa threaten them from the ground. Introduced predators such as feral dogs, African cats, European cats, mongooses and civets also prey on them. However, of all these creatures, humans are the biggest hazard. Though killing the lemurs was taboo, [12] Coquerel's sifaka see humans now as a threat and may give out an alarm call to warn the others. [12]

Some lemurs are unaware of the danger humans pose and will approach humans on the ground. To intimidate predators they do recognize, the lemurs announce the threat with a warning call, and stare at the threat, shaking their heads back and forth. [24]

Cultural references

The titular lemur on the PBS Kids television program Zoboomafoo is portrayed by a Coquerel's sifaka [12] named Jovian. Jovian lived at the Duke Lemur Center, where the show was originally filmed until he died from kidney failure at the age of 20 on November 10, 2014. [25] His son Charlemagne, known as "Charlie", lives at the center, with his family group of other Coquerel's sifakas. [26]

Related Research Articles

<span class="mw-page-title-main">Indriidae</span> Family of lemurs

The Indriidae are a family of strepsirrhine primates. They are medium- to large-sized lemurs, with only four teeth in the toothcomb instead of the usual six. Indriids, like all lemurs, live exclusively on the island of Madagascar.

<span class="mw-page-title-main">Sifaka</span> Genus of lemur

A sifaka is a lemur of the genus Propithecus from the family Indriidae within the order Primates. The name of their family is an onomatopoeia of their characteristic "shi-fak" alarm call. Like all lemurs, they are found only on the island of Madagascar. All species of sifakas are threatened, ranging from endangered to critically endangered.

<span class="mw-page-title-main">Indri</span> Genus of lemurs

The indri, also called the babakoto, is one of the largest living lemurs, with a head-body length of about 64–72 cm and a weight of between 6 and 9.5 kg. It has a black and white coat and maintains an upright posture when climbing or clinging. It is monogamous and lives in small family groups, moving through the canopy, and is herbivorous, feeding mainly on leaves but also seeds, fruits, and flowers. The groups are quite vocal, communicating with other groups by singing, roaring and other vocalisations. Besides humans, it is the only mammal found that can use rhythm.

<span class="mw-page-title-main">Lemur</span> Clade of primates endemic to the island of Madagascar

Lemurs are wet-nosed primates of the superfamily Lemuroidea, divided into 8 families and consisting of 15 genera and around 100 existing species. They are endemic to the island of Madagascar. Most existing lemurs are small, have a pointed snout, large eyes, and a long tail. They chiefly live in trees and are active at night.

<span class="mw-page-title-main">Verreaux's sifaka</span> Species of lemur

Verreaux's sifaka, or the white sifaka, is a medium-sized primate in one of the lemur families, the Indriidae. Critically Endangered, it lives in Madagascar and can be found in a variety of habitats from rainforest to dry deciduous forests of western Madagascar and the spiny thickets of the south. Its 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, on the ground its only means of locomotion is hopping. The species lives in small troops which forage for food.

<span class="mw-page-title-main">Giant mouse lemur</span> Genus of small primates from Madagascar

The giant mouse lemurs are members of the strepsirrhine primate genus Mirza. Two species have been formally described; the northern giant mouse lemur and Coquerel's giant mouse lemur. Like all other lemurs, they are native to Madagascar, where they are found in the western dry deciduous forests and further to the north in the Sambirano Valley and Sahamalaza Peninsula. First described in 1867 as a single species, they were grouped with mouse lemurs and dwarf lemurs. In 1870, British zoologist John Edward Gray assigned them to their own genus, Mirza. The classification was not widely accepted until the 1990s, which followed the revival of the genus by American paleoanthropologist Ian Tattersall in 1982. In 2005, the northern population was declared a new species, and in 2010, the World Wide Fund for Nature announced that a southwestern population might also be a new species.

<span class="mw-page-title-main">Fossa (animal)</span> Cat-like, carnivorous mammal endemic to Madagascar

The fossa is a slender, long-tailed, cat-like mammal that is endemic to Madagascar. It is a member of the carnivoran family Eupleridae.

<span class="mw-page-title-main">Diademed sifaka</span> Species of lemur

The diademed sifaka, or diademed simpona, is an endangered species of sifaka, one of the lemurs endemic to certain rainforests in eastern Madagascar. Along with the indri, this species is one of the two largest living lemurs, with an average weight of 6.5 kg and a total adult length of approximately 105 centimetres (41 inches), half of which is its tail. Russell Mittermeier, one of the contemporary authorities on lemurs, describes the diademed sifaka as "one of the most colorful and attractive of all the lemurs", having a long and silky coat. P. diadema is also known by the Malagasy names simpona, simpony and ankomba joby. The term "diademed sifaka" is also used as a group species designation formerly encompassing four distinct subspecies.

<span class="mw-page-title-main">Golden-crowned sifaka</span> A medium-sized lemur with mostly white fur, prominent furry ears, and a golden-orange crown

The golden-crowned sifaka or Tattersall's sifaka is a medium-sized lemur characterized by mostly white fur, prominent furry ears, and a golden-orange crown. It is one of the smallest sifakas, weighing around 3.5 kg (7.7 lb) and measuring approximately 90 cm (35 in) from head to tail. Like all sifakas, it is a vertical clinger and leaper, and its diet includes mostly seeds and leaves. The golden-crowned sifaka is named after its discoverer, Ian Tattersall, who first spotted the species in 1974. However, it was not formally described until 1988, after a research team led by Elwyn L. Simons observed and captured some specimens for captive breeding. The golden-crowned sifaka most closely resembles the western forest sifakas of the P. verreauxi group, yet its karyotype suggests a closer relationship with the P. diadema group of eastern forest sifakas. Despite the similarities with both groups, more recent studies of its karyotype support its classification as a distinct species.

<span class="mw-page-title-main">Menabe Antimena</span> Protected area in Menabe, Madagascar

Menabe Antimena is a protected area in Menabe region of western Madagascar. It belongs to the municipalities of Tsimafana, Beroboka, Bemanonga, Belon'i Tsiribihina, and Tsaraotana.

<span class="mw-page-title-main">Silky sifaka</span> A large lemur from Madagascar

The silky sifaka is a large lemur characterized by long, silky, white fur. It has a very restricted range in northeastern Madagascar, where it is known locally as the simpona. It is one of the rarest mammals on Earth. The silky sifaka is one of nine sifaka species, and one of four former subspecies of diademed sifaka (P. diadema). Studies in 2004 and 2007 compared external proportions, genetics, and craniodental anatomy supporting full species status, which has generally been accepted.

<span class="mw-page-title-main">Kirindy Mitea National Park</span> National park in Madagascar

The Kirindy Mitea National Park is a national park on the coast of the Mozambique Channel, in south-west Madagascar. The 72,200 hectares park contains many endemic animals and plants and claims to have the greatest density of primates in the world.

Maningoza Special Reserve is a 9,826 hectares wildlife reserve near Besalampy in Madagascar. It was created in 1956 to protect the many endemic plants and animals, and it also contains some of the last remaining areas of dry deciduous forest on the island.

<span class="mw-page-title-main">Perrier's sifaka</span> Species of lemur

Perrier's sifaka is a lemur endemic to Madagascar. It was formerly considered to be a subspecies of diademed sifaka It has a very small range in northeastern Madagascar where its habitat is dry deciduous or semihumid forest. Part of its range is in protected areas. It is an almost entirely black sifaka and measures about 90 cm (35 in), half of which is a bushy tail. Females are slightly larger than males.

<span class="mw-page-title-main">Crowned sifaka</span> Species of mammal

The crowned sifaka is a sifaka endemic to western Madagascar, a part of the world where nature and its biological diversity faces enormous and devastating consequences resulting from anthropogenic activities. It is a species of lemur belonging to the Indriidae family, it is of comparable size to the Golden-crowned sifaka and up to a meter in length, of which 47-57 centimeters are tail. The species is an arboreal vertical climber and leaper whose diet consists of leaves, fruits and flowers. It is threatened by habitat destruction caused by human activities and is currently classified as critically endangered by the IUCN. Conservation planning needs to take local people needs and views into account in order to be successful over the long term.

<span class="mw-page-title-main">Milne-Edwards's sifaka</span> Species of lemur

Milne-Edwards's sifaka, or Milne-Edwards's simpona, is a large arboreal, diurnal lemur endemic to the eastern coastal rainforest of Madagascar. Milne-Edwards's sifaka is characterized by a black body with a light-colored "saddle" on the lower part of its back. It is closely related to the diademed sifaka, and was until recently considered a subspecies of it. Like all sifakas, it is a primate in the family Indriidae.

<span class="mw-page-title-main">Madagascar succulent woodlands</span>

The Madagascar succulent woodlands are a xeric shrublands ecoregion in southwestern and central western Madagascar. Native plants survive in the arid climate and long dry season with adaptations like succulent leaves, water storing trunks, photosynthetic stems, and dropping leaves during the dry season. The ecoregion is threatened by various human activities.

<span class="mw-page-title-main">Makay Massif</span> Mountain range in Madagascar

The Makay Massif is a mountain range in western Madagascar.

References

  1. 1 2 3 Louis, E.E.; Bailey, C.A.; Sefczek, T.M.; King, T.; Radespiel, U.; Frasier, C.L. (2020). "Propithecus coquereli". IUCN Red List of Threatened Species . 2020: e.T18355A115572275. doi: 10.2305/IUCN.UK.2020-2.RLTS.T18355A115572275.en . Retrieved 19 November 2021.
  2. 1 2 "Checklist of CITES Species". CITES. UNEP-WCMC. Retrieved 18 March 2015.
  3. ↑ Groves, C. P. (2005). Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore: Johns Hopkins University Press. p. 120. ISBN   0-801-88221-4. OCLC   62265494.
  4. ↑ Kappeler, P. M. (1991). "Patterns of sexual dimorphism in body weight among prosimian primates". Folia Primatologica. 57 (3): 132–146. doi:10.1159/000156575. PMID   1794769.
  5. ↑ Tattersall, I. (1982). The Primates of Madagascar. Columbia University.
  6. 1 2 Pochron, S. T.; Wright, P. C. (2005). "Dance of the sexes: a lemur needs some unusual traits to survive in Madagascar's unpredictable environment". CBS. Archived from the original on 25 September 2015. Retrieved 12 March 2012.
  7. 1 2 3 4 5 6 Salmona, Jordi (2014). "Extensive survey of the Endangered Coquerel's sifaka Propithecus coquereli". Endangered Species Research. 25 (2): 175–183. doi: 10.3354/esr00622 . hdl: 10400.7/348 .
  8. ↑ Petter, J.-J. (1962). "Recherches sur l'écologie et l'éthologie des lémuriens malgaches". Mémoires du Muséum National d'Histoire Naturelle, Série A, Zoologie. 27: 1–146.
  9. 1 2 Kun-Rodrigues C, Salmona J, Besolo A, Rasolondraibe E, Rabarivola C, Marques TA, Chikhi L (2014). "New density estimates of a threatened sifaka species (Propithecus coquereli) in Ankarafantsika National Park". American Journal of Primatology. 76 (6): 515–528. doi:10.1002/ajp.22243. hdl: 10400.7/558 . PMID   24443250. S2CID   11185235 . Retrieved 18 Feb 2015.
  10. 1 2 3 4 5 Mittermeier, R.A.; Konstant, W.R.; Hawkins, F.; Louis, E.E.; et al. (2006). Lemurs of Madagascar . Illustrated by S.D. Nash (2nd ed.). Conservation International. ISBN   1-881173-88-7. OCLC   883321520.
  11. 1 2 3 4 Richard, Alison (2003). "Propithecus, Sifakas". In Steven M. Goodman; Jonathan P. Benstead (eds.). The Natural History of Madagascar. Chicago: University of Chicago. pp. 1345–1348. ISBN   0-226-30306-3.
  12. 1 2 3 4 5 6 7 8 9 10 Oldenkamp, R. (2011). "Propithecus coquereli". Animal Diversity Web. Retrieved 18 March 2012.
  13. ↑ Campbell, J.; Glenn, K.; Grossi, B.; Eisemann, J. (2001). "Use of Local North Carolina Browse Species to Supplement the Diet of a Captive Colony of Folivorous Primates (Propithecus sp.)". Zoo Biology. 20 (6): 447–461. doi:10.1002/zoo.10007.
  14. 1 2 3 Garbutt, N. (2007) Mammals of Madagascar: A Complete Guide. A&C Black Publishers Ltd.
  15. ↑ Kubzdela, K.; Richard, A.; Pereira, M. (1992). "Social Relations in Semi-Free Ranging Sifakas (Propithecus verreauxi coquereli) and the Question of Female Dominance". American Journal of Primatology. 28 (2): 139–145. doi:10.1002/ajp.1350280206. PMID   31941220. S2CID   84398797.
  16. ↑ Taylor, L. (2008). "Old Lemurs: Preliminary Data on Behavior and Reproduction from the Duke University Primate Center", pp. 319-333 in J Fleagle, C Gilbert, eds. Elwyn Simons: A Search for Origins. New York: Springer Science Business+Media, LLC.
  17. ↑ Iverson, Erik (26 July 2012). "Coquerel's Sifaka." Archived 2012-07-12 at the Wayback Machine Mongabay.
  18. ↑ Mason, Angela; Wolfe, Linda; Johnson, Jeffrey (1995). "Hand Preference in the Sifaka (Propithecus verreauxi coquereli) During Feeding in Captivity". Primates. 36 (2): 275–280. doi:10.1007/BF02381353. S2CID   21949388.
  19. ↑ Greive, Bradley Trevor. Priceless: The Vanishing Beauty of a Fragile Planet. Kansas City, MO: Andrews McMeel Pub., 2003. Print.
  20. ↑ Richard, A. (1978). Behavioral Variation: Case Study of a Malagasy Lemur. Cranbury, New Jersey: Associated University Presses, Inc.
  21. ↑ Jolly, Alison (2004). Lords and Lemurs: Mad Scientists, Kings with Spears, and the Survival of Diversity in Madagascar. Boston: Houghton Mifflin. Archived from the original on 2005-03-04.
  22. 1 2 3 Fichtel, Claudia; Kappeler, Peter M. (2011). "Variation in the Meaning of Alarm Calls in Verreaux's and Coquerel's Sifakas (Propithecus verreauxi, P. coquereli)". International Journal of Primatology. 32 (2): 346–361. doi:10.1007/s10764-010-9472-9. PMC   3047677 . PMID   21475394.
  23. ↑ Smith, Dave (29 December 2011). "Lemurs Killed, Eaten More in Madagascar as Taboos Fade." Archived 2012-02-03 at the Wayback Machine International Business Times.
  24. ↑ Fichtel, C.; van Schaik, C. P. (2006). "Semantic Differences in Sifaka (Propithecus verreauxi) Alarm Calls: A Reflection of Genetic or Cultural Variants?". Ethology. 112 (9): 839–849. Bibcode:2006Ethol.112..839F. doi:10.1111/j.1439-0310.2006.01239.x.
  25. ↑ Jovian Archived 2014-11-14 at the Wayback Machine . Duke Lemur Center
  26. ↑ Duke Lemur Center. (2011). "Coquerel's Sifaka".
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