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Fossa (animal)

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This article is about the animal with the common name Fossa. For the taxonomic genus, see Malagasy Civet. For other uses, see Fossa.

Fossa[1]
Scientific classification
Kingdom:
Animalia
Phylum:
Chordata
Class:
Mammalia
Order:
Carnivora
Family:
Eupleridae
Genus:
Cryptoprocta

Bennett, 1833
Species:
C. ferox
Binomial name
Cryptoprocta ferox
Bennett, 1833
Range map showing the Fossa's distribution in Madagascar. Areas in red mark its distribution and mostly run along the outer edge of the island.
Distribution of the Fossa
(Cryptoprocta ferox)[3]
Synonyms
  • typicus A. Smith, 1834

The Fossa (Cryptoprocta ferox) (Template:Pron-en or /ˈfɒsə/[4]) is a cat-like, carnivorous mammal that is endemic to Madagascar. It is a member of the Eupleridae, a family of carnivorans closely related to the mongoose family (Herpestidae). Its classification has been controversial because its physical traits resemble those of cats, yet other traits suggest a close relationship with viverrids (most civets and their relatives). Its classification, along with that of the other Malagasy carnivores, influenced hypotheses about how many times mammalian carnivores have colonized the island. With genetic studies demonstrating that the Fossa and all other Malagasy carnivores are most closely related to each other (forming a clade, recognized as the family Eupleridae), it is now thought that euplerids colonized the island once approximately 18 to 20 million years ago.

The Fossa is the largest mammalian carnivore on the island of Madagascar, though it has been compared to a small cougar. Adults have a head-body length of 70–80 cm (28–31 in) and weigh between 5.5–8.6 kg (12–19 lb), with the males being larger than the females. It has semi-retractable claws and flexible ankles that allow it to climb up and down trees head-first, and also support jumping from tree-to-tree. The Fossa is unique within its family for the shape of its genitalia, which share traits with those of cats and hyenas.

The geographic range of the species is widespread, though population densities are usually low. It is found solely in forested habitat, and actively hunts both day and night. Over 50% of its diet consists of lemurs, the endemic primates found on the island, though tenrecs, rodents, lizards, birds, and other animals are also documented as prey. Mating usually occurs in trees on horizontal limbs and can last for several hours. Litters range from one to six pups, which are born blind and toothless (altricial). Infants wean after 4.5 months and are independent after a year. Sexual maturity occurs around three to four years of age, and life expectancy in captivity is 20 years. The Fossa is listed as Vulnerable by the International Union for Conservation of Nature (IUCN). It is generally feared by the Malagasy people and is often protected by their taboo, known as fady. The greatest threat to the species is habitat destruction.

Taxonomy

The Fossa was formally described in 1833 by Edward Turner Bennett. The genus name Cryptoprocta refers to how the animal's anus is hidden by its anal pouch (crypto- is Greek for "hidden", and procta is Greek for "anus").[5] The species name ferox is Latin for "fierce" or "wild."[6] Its common name is Malagasy and can be spelled fossa or fosa.[5] Due to shared physical traits with civets, mongooses, and cats (Felidae), its classification has been controversial. Bennett originally placed the Fossa as a type of civet in the family Viverridae, a classification that long remained popular among taxonomists. Its compact braincase, large eye sockets, retractable claws, and specialized carnivorous dentition, have also led some taxonomists to associate it with the felids.[7] In 1939, William King Gregory and Milo Hellman placed the Fossa in its own subfamily within Felidae, the Cryptoproctinae. George Gaylord Simpson placed it back in Viverridae in 1945, still within its own subfamily, yet admitted it had many cat-like characteristics.[5][8]

The Fossa has a cat-like appearance, resembling a small Cougar.[5]

In 1993, Géraldine Veron and François Catzeflis published a DNA hybridization study suggesting that the Fossa was more closely related to mongooses (family Herpestidae) than to cats or civets.[7][8] However, in 1995, Veron's morphological study once again grouped it with Felidae.[8] In 2003, molecular phylogenetic studies using nuclear and mitochondrial genes by Anne Yoder and colleagues showed that all native Malagasy carnivorans share a common ancestry that excludes other carnivores (meaning they form a clade, making them monophyletic) and are most closely related to Asian and African Herpestidae.[9][10][11] To reflect these relationships, all Malagasy carnivorans are now placed in a single family, Eupleridae.[1] Within Eupleridae, the Fossa is placed in the subfamily Euplerinae with the Falanouc (Eupleres goudoti) and Malagasy Civet (Fossa fossana), but its exact relationships are poorly resolved.[1][11][9]

An extinct relative of the Fossa was described in 1902 from subfossil remains and recognized as a separate species, Cryptoprocta spelea, in 1935. This species was much larger than the living Fossa, but otherwise similar.[5][12] Across Madagascar, people distinguish two kinds of Fossa—a large fosa mainty ("black fossa") and the smaller fosa mena ("reddish fossa")—and a white form has been reported in the southwest. It is unclear whether this is purely folklore or related to variation with sex or age, or whether there is indeed more than one species of living Fossa.[13][12]

Phylogeny of Eupleridae within Feliformia[11]
Feliformia 

(other feliforms)

Viverridae (civets, genets, and their relatives)

Hyaenidae (hyenas)

Herpestidae (mongooses)

Eupleridae (Malagasy carnivores)

Phylogeny of Malagasy carnivores (Eupleridae)[9]
Eupleridae 
Cryptoprocta

C. ferox (Fossa)

†C. spelea (Giant Fossa)

Fossa (Malagasy Civet)

Eupleres (Falanouc)

Description

The Fossa appears as a diminutive form of a large felid, such as a Cougar,[13] but with a slender body and muscular limbs,[7] and a tail nearly as long as the rest of the body.[13] It has a mongoose-like head,[7] relatively longer than that of a cat,[13] although with a muzzle that is broad[7] and short,[13] and with large but rounded ears.[5][13] It has medium brown eyes set relatively wide apart with pupils that contract to slits. Like many carnivorans that hunt at night, its eyes reflect light; the reflected light is orange in hue.[7] Its head-body length is 70–80 cm (28–31 in) and its tail is 65–70 cm (26–28 in) long. There is some sexual dimorphism, with adult males (weighing 6.2–8.6 kg (14–19 lb)) being larger than females (5.5–6.8 kg (12–15 lb)).[13] Smaller individuals are typically found north and east on Madagascar, while larger ones to the south and west,[5] likely due to Bergmann's rule. Unusually large individuals weighing up to 20 kg (44 lb) have been reported, but there is some doubt as to the reliability of the measurements.[13]

Cranium (dorsal, ventral, and lateral views) and mandible (lateral and dorsal views)

Both males and females have a short, straight pelage that is relatively dense[13] and is without spots or patterns.[5] Females are a reddish-brown dorsally and colored a dirty cream ventrally. Males are similarly colored, although some have an orange coloration to their abdomen from gland secretions when in rut,[13] although some researchers have failed to observe this. Darker, blackish individuals are known,[13][5] as are whitish ones,[13] but there is no consensus as to whether these are melanistic and leucistic individuals,[5] folklore, or simply general pattern distinctions. Locals call the reddish morph fosa mena, and the larger blackish morph fosa mainty.[13] The tail tends to be lighter in coloration than the sides. Juveniles are either gray or nearly white.[5]

Several of the animal's physical features are well adapted to climbing through trees.[7] It uses its tail to aid in balance and has semi-retractable claws that it uses to climb trees in its search for prey.[13] It has semiplantigrade feet,[5] switching between a plantigrade-like gait (when arboreal) and a digitigrade-like one (when terrestrial).[14] The soles of its paws are nearly bare and covered with strong pads.[5] The Fossa has very flexible ankles that allow it to readily grasp tree trunks so as to climb up or down trees head first or to leap to another tree.[7] Captive juveniles have been known to swing upside down by their hindfeet from knotted ropes.[7]

The Fossa has several scent glands, although the glands are less developed in females. Like herpestids it has a perianal skin gland inside an anal sac which surrounds the anus like a pocket. The pocket opens to the exterior with a horizontal slit below the tail. A gland on its chest produces a reddish substance that stains the abdominal fur of males in rut. Other glands are located near the penis or vagina, with the penile glands emitting a strong odor. Like the herpestids, it has no prescrotal glands.[5]

One of the more interesting physical features of this species is its external genitalia. Males have an unusually long penis and baculum, reaching to between its forelegs when erect. The glans extends down the shaft about halfway and is spiny except at the tip. In comparison, the glans of felids is short and spiny, while that of viverrids is smooth and long.[5] The female Fossa exhibits transient masculization, starting at about 1-2 years of age, developing an enlarged, spiny clitoris that resembles a male's penis. The enlarged clitoris is supported by an os clitoridis,[13] which decreases in size as the animal grows.[14] Females do not have a pseudo-scrotum,[13] but they do secrete an orange substance that colors their underparts, much like the secretions of males.[15] Hormone levels (testosterone, androstenedione, dihydrotestosterone) do not seem to play a part in this transient masculization, as those levels are the same in masculinized juveniles and nonmasculinized adults. It is speculated that the transient masculization either reduces sexual harassment of juveniles females by adult males, or reduces aggression from territorial females.[13] While other mammal species (such as the Spotted Hyena) have a pseudo-penis,[16] none of them are known to have theirs diminish in size as the animal grows.[15]

Overall, the Fossa has features in common with three different carnivoran families, leading researchers to place it and other members of the Eupleridae alternatively in Herpestidae, Viverridae, and Felidae. Felid features are primarily those associated with eating and digestion, including teeth shape and facial portions of the skull, the tongue, and the digestive tract,[5] typical of its exclusively carnivorous diet.[7] The remainder of the skull most closely resembles those of genus Viverra, while the general body structure is most similar to that of various members of Herpestidae. The permanent dentition is 3.1.3-4.13.1.3-4.1, with the deciduous formula being similar but lacking the fourth premolar and the molar. The Fossa has a large, prominent rhinarium similar to that of viverrids, but has comparatively larger, round ears, almost as large as those of a similarly sized felid. Its facial vibrissae (whiskers) are long, with the longest being longer that its head. Like some mongoose genera, particularly Galidia (which is now in the Fossa's own Eupleridae family) and Herpestes (of Herpestidae), it has carpal vibrassae as well. Its claws are retractile, but unlike those of Felidae species, they are not hidden in skin sheaths. It has three pairs of nipples (one inguinal, one ventral, and one pectoral)[13] to support a litter of two to four young.[5]

Habitat and distribution

The Fossa has the most widespread geographical range of the Malagasy carnivores, and is generally found in low numbers throughout the island in remaining tracts of forest, preferring pristine undisturbed forest habitat. It is also encountered in some degraded forests, but in lower numbers, reflecting the declining populations in those areas. Although the Fossa is found in all known forest habitats throughout Madagascar, including the western, dry deciduous forests, the eastern rainforests, and the southern spiny forests,[17] it is seen more frequently in the trees of humid forests rather than dry forests. This may be because the reduced canopy in dry forests provides less shade, and also because the Fossa seems to travel more easily in humid forests.[7] It is absent from those areas with heaviest habitat disturbance and, like most of Madagascar's fauna, is absent from the central high plateau of the country.[17]

The Fossa has been found across several different elevational gradients in undisturbed portions of protected areas throughout Madagascar. In the Réserve Naturelle Intégrale d'Andringitra, evidence of the Fossa has been reported at four different sites ranging from 810 to 1,625 m (2,657 to 5,331 ft).[18] Its highest known elevational range was reported at 2,000 m (6,600 ft);[19] its presence high on the Andringitra Massif was subsequently confirmed in 1996.[18] Similarly, evidence has also been reported of the Fossa at the elevational extremes of 440 m (1,440 ft) and 1,875 m (6,152 ft) in the Andohahela National Park.[20] The presence of the Fossa at these locations indicates its ability to adapt to various elevations, consistent with its reported distribution in all Madagascar forest types.[17]

Behavior

Fossa are active both day and night (cathemeral).

The Fossa is active during both the day and the night and is considered cathemeral; activity peaks may occur early in the morning, late in the afternoon, and late in the night.[13] The animal generally does not reuse sleeping sites, but females with young do return to the same den.[13] The home ranges of male Fossas in Kirindy Forest are up to 26 km2 (10 sq mi) large, compared to 13 km2 (5.0 sq mi) for females. These ranges overlap—by about 30% according to data from the eastern forests—but females usually have separated ranges. Home ranges grow during the dry season, perhaps because less food and water is available. A Fossa has been observed moving a straight-line distance of 7 km (4.3 mi) in 16 hours.[13] The animal's population density appears to be low: in Kirindy, where it is thought to be common, its density has been estimated at one animal per 4 square kilometres (1.5 sq mi).[7]

Except for mothers with young and occasional observations of pairs of males, the species is solitary.[5][7] Fossas communicate using sounds, scents, and visual signals. Vocalizations include purring, a threatening call,[5] and a call of fear, consisting of "repeated loud, coarse inhalations and gasps of breath".[7] A long, high yelp may function to attract other Fossas. Females mew during mating and males produce a sigh when they have found a female.[5][7] Throughout the year, animals produce long-lasting scent marks on rocks, trees, and the ground using glands in the anal region and on the chest.[7][5][13] They also communicate using face and body expression, but the significance of these signals is uncertain.[5] The animal is aggressive only during the mating, and males in particular fight boldly. After a short fight, the loser flees and is followed by the winner for a short distance.[5]

Diet

The Fossa is a carnivore that hunts small to medium-sized animals, from fish to birds. One of eight carnivorous species endemic to Madagascar, the Fossa is the only predator capable of preying upon adults of all extant lemur species[21] and is the predominant predator of lemurs.[22] In addition, the Diademed Sifaka (Propithecus diadema) has been reported as a prey species, even though it is about 90% of the weight of the average Fossa.[7] Reports of its dietary habits demonstrate a wide variety of prey selectivity and specialization depending on habitat and season; diet does not vary by sex. While the Fossa is thought to be a lemur specialist in Ranomafana National Park,[23] its diet is more variable in other rain forest habitats. The diet of the Fossa in the wild has been studied by analyzing their distinctive scats, which resemble gray cylinders with twisted ends and measure 10–14 cm (3.9–5.5 in) long by 1.5–2.5 cm (0.6–1.0 in) thick.[24] Scat collected and analyzed from both Andohahela and Andringitra contained lemur matter, and rodents. Eastern populations in Andringitra incorporate the widest recorded variety of prey, including both vertebrates and invertebrates. Vertebrates consumed ranged from reptiles to a wide variety of birds, including both understory and ground birds, and mammals, including insectivores, rodents, and primates. Invertebrates eaten by the Fossa in the high mountain zone of Andringitra include insects and crabs.[18][20] One study found that vertebrates comprised 94% of the diet of Fossas, with lemurs comprising over 50%, as well as tenrecs (9%), lizards (9%), and birds (2%). Seeds, which comprised 5% of the diet, may have been in the stomachs of the lemurs eaten, or may have been consumed with fruit taken for water, as seeds were more common in the stomach in the dry season. Even other large prey items, with the average prey size being 40 grams (1.4 oz), in contrast to the average prey size of 480 grams (17 oz) in humid forests and over 1,000 grams (35 oz) in dry deciduous forests.[13] In a study of Fossa diet in the dry deciduous forest of western Madagascar, more than 90% of prey items were vertebrates, and more than 50% were lemurs. The primary diet consisted approximately six lemur species and two or three spiny tenrec species, along with snakes and small mammals.[24] Generally, the Fossa preys upon larger lemurs and rodents in preference to smaller ones.[25]

Prey are obtained by hunting either on the ground or in the trees. During the non-breeding season the Fossa hunts individually, but during the breeding season hunting parties may be seen, and these may be pairs or later on mothers and young. One member of the group scales a tree and chases the lemurs from tree to tree, forcing them down to the ground where the other is easily able to capture them.[13] Long-term observations of the Fossa's predation patterns on rainforest sifakas suggest that the Fossa hunts in a subsection of their range until prey density is increased, then moves on.[26] The Fossa has been reported to predate on domestic animals, such as goat and small calves, and especially chickens. In captivity, the Fossa consumes between 800 and 1,000 grams (28 and 35 oz) of meat a day. {{citation}}: Empty citation (help) Food taken in captivity include amphibians, birds, insects, reptiles, and small- to medium-sized mammals.[5]

This wide variety of prey items taken in various rainforest habitats is similar to the varied dietary composition noted[18][20] occurring in the dry forests of Western Madagascar, as well. As the largest endemic predator on Madagascar, this dietary flexibility combined with a flexible activity pattern[21] has allowed[18][20] it to exploit a wide variety of niches available throughout the island, making it a potential keystone species for Madagascar ecosystems.[17]

Breeding

Fossa illustration circa 1927

Most of the details of reproduction in wild populations are from the western dry deciduous forests; whether certain of these details are applicable to eastern populations will require further field research.[13] Mating typically occurs during September and October,[5] although there are reports of it occurring as late as December,[13] and can be highly conspicuous.[7] Intromission usually occurs in trees on horizontal limbs about 20 m (66 ft) off the ground. Frequently the same tree is used year after year, with remarkable precision as to the date the season commences. Trees are often near a water source, and have limbs strong enough and wide enough to support the mating pair, about 20 cm (7.9 in) wide. Some mating has been reported on the ground as well.[13]

As many as eight males will be at a mating site, staying in close vicinity to the receptive female. The female seems to choose the male she mates with, and the males compete for the attention of the female with a significant amount of vocalization and antagonistic interactions. The female may chose to mate with several of the males, and her choice of mate does not seem to have any correlation to the physical appearance of the males.[13] To stimulate the male to mount her, she gives a series of mewling vocalizations. The male mounts from behind, resting his body on her slightly off-center,[13] a position requiring delicate balance; if the female were to stand, the male would have significant difficulty continuing. He places his paws on her shoulders[7] or grasps her around the waist and often licks her neck.[13] Mating may last for nearly three hours.[15] This lengthy mating is unusual due to the physical nature of the male's erect penis, which has backwards-pointing spines along most of its length.[15] Fossa mating includes a copulatory tie,[13] which may be enforced by the male's spiny penis.[15] The tie is difficult to break if the mating session is interrupted.[13] Copulation with a single male may be repeated several times, with a total mating time of up to fourteen hours, while the male may remain with the female for up to an hour after the mating. A single female may occupy the tree for up to a week, mating with multiple males over that time. Also, other females may take her place, mating with some of the same males as well as others.[13]

The birthing of the litter of one to six[14] (typically two to four)[5] takes place in a concealed location, such as an underground den, a termite mound, a rock crevice, or in the hollow of a large tree[13] (particularly those of the Commiphora genus).[7] Contrary to older research, litters are of mixed sexes.[5][13] Young are born in December or January, making the gestation period 90 days,[5] with the late mating reports indicating a gestational period of about six to seven weeks.[13] The newborns have nearly white fur, are blind and toothless, and weigh no more than 100 g (3.5 oz).[5][13] After about two to three weeks the cubs' eyes open,[5] they become more active, and their fur darkens to a pearl gray.[13] The cubs do not take solid food until three months old,[15] and do not leave the den until they are 4.5 months old; they are weaned shortly after that.[5][13] After the first year, the juveniles are independent of their mother.[13] Permanent teeth appear at 18 to 20 months.[5][13] Physical maturity is reached by about two years old,[15] but sexual maturity does not happen for another year or two,[5][13] and the young may stay with their mother until they are fully mature. Lifespan in captivity is up to or past 20 years of age, possibly due to the slow juvenile development.[14]

Human interactions

The Fossa has been assessed as "Vulnerable" by the IUCN Red List since 2008, as its population size has probably declined by at least 30% over the last 21 years; previous assessments have included "Endangered" (2000) and "Insufficiently Known" (1988, 1990, 1994).[2] The species is dependent on forest and thus threatened by the widespread destruction of Madagascar's native forest, but is also able to persist in disturbed areas.[13][7] Several diseases have been isolated from the Fossa, some of which are thought to have been transmitted by feral dogs and cats.[13]

Taboo, known in Madagascar as fady,[27] offers protection for the Fossa and other carnivores.[28] In the Marolambo District (part of the Atsinanana region in Toamasina Province), the Fossa has traditionally been hated and feared as a dangerous animal. It has been described as "greedy and aggressive", known for taking fowl and piglets, and believed to "take little children who walk alone into the forest". Some do not eat it for fear that it will transfer its undesirable qualities to anyone who consumes it,[27] but the animal is also taken for bushmeat.[13] Near Ranomafana National Park, the Fossa, along with several of its smaller cousins and the introduced Small Indian Civet (Viverricula indica), are known to "scavenge on the bodies of ancestors", which are buried in shallow graves in the forest. For this reason, eating these animals is strictly prohibited by fady. However, if they wander into villages in search of domestic fowl, they may be killed or trapped. Small carnivore traps have been observed near chicken runs in the village of Vohiparara.[28]

References

  1. ^ a b c Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 559–561. ISBN 978-0-8018-8221-0. OCLC 62265494.
  2. ^ a b Template:IUCN2008
  3. ^ Garbutt 2007, pp. 211–214.
  4. ^ Croke, V. "The Deadliest Carnivore". Discover. Retrieved 2010-05-12.
  5. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag Köhncke, M.; Leonhardt, K. (1986). "Cryptoprocta ferox" (PDF). Mammalian Species (254): 1–5. Retrieved 19 May 2010.
  6. ^ Borror 1960, p. 39.
  7. ^ a b c d e f g h i j k l m n o p q r s t u Hawkins 2003, pp. 1360–1363.
  8. ^ a b c Yoder & Flynn 2003, pp. 1253–1256.
  9. ^ a b c Yoder, A.D.; Burns, M.M.; Zehr, S.; Delefosse, T.; Veron, G.; Goodman, S.M.; Flynn, J.J. (2003). "Single origin of Malagasy Carnivora from an African ancestor" (PDF). Nature. 421: 734–737. Retrieved 19 May 2010.
  10. ^ Veron, G.; Colyn, M.; Dunham, A.E.; Taylor, P.; Gaubert, P. (2004). "Molecular systematics and origin of sociality in mongooses (Herpestidae, Carnivora)" (PDF). Molecular Phylogenetics and Evolution. 30: 582–598. doi:10.1016/S1055-7903(03)00229-X. Retrieved 19 May 2010.
  11. ^ a b c Barycka, E. (2007). "Evolution and systematics of the feliform Carnivora". Mammalian Biology. 72 (5): 257–282. doi:10.1016/j.mambio.2006.10.011.
  12. ^ a b Goodman, S.M.; Rasoloarison, R.M.; Ganzhorn, J.U. (2004). "On the specific identification of subfossil Cryptoprocta (Mammalia, Carnivora) from Madagascar" (PDF). Zoosystema. 26 (1): 129–143.
  13. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am an ao ap aq ar as Goodman 2009, Family Eupleridae (Madagascar Carnivores)
  14. ^ a b c d Mueller, J.; Sironen, A.; Lukas, K.E. (2007). "Infant development and behavior in the Fossa Cryptoprocta ferox" (PDF). Small Carnivore Conservation. 37: 11–17.
  15. ^ a b c d e f g Macdonald 2009, pp. 668–669.
  16. ^ Drea, C.M.; Place, N.J.; Weldele, M.L.; Coscia, E.M.; Licht, P.; Glickman, S.E. (2002). "Exposure to naturally circulating androgens during foetal life incurs direct reproductive costs in female spotted hyenas, but is prerequisite for male mating" (PDF). Proceedings of the Royal Society B. 269: 1981–1987. doi:10.1098/rspb.2002.2109.
  17. ^ a b c d Dollar 2007, pp. 63–76.
  18. ^ a b c d e Goodman, S.M. (1996). "The carnivores of the Reserve Naturelle Integrale d'Andringitra, Madagascar". Fieldiana Zoology (85): 289–292. ISSN 0015-0754.
  19. ^ Albignac 1973, pp. 1–206.
  20. ^ a b c d Goodman, S.M.; Pidgeon, M. (1999). "Carnivora of the Reserve Naturelle Integrale d'Andohahela, Madagascar". Fieldiana Zoology (94): 259–268. ISSN 0015-0754.
  21. ^ a b Dollar, L. (1999). "Preliminary report on the status, activity cycle, and ranging of Cryptoprocta ferox in the Malagasy rainforest, implications for conservation" (PDF). Small Carnivore Conservation. 20: 7–10.
  22. ^ Wright, P.C. (1995). "Demography and life history of free ranging Propithecus diadema Edwardsi in Ranomafana National Park, Madagascar" (PDF). International Journal of Primatology. 16 (5): 835–854. doi:10.1007/BF02735722.
  23. ^ Wright, P.C.; Heckscher, S.K.; Dunham, A.E. (1997). "Predation on Milne Edward's sifaka (Propithecus diadema edwardsi) by the fossa (Cryptoprocta ferox) in the rainforest of southeastern Madagascar". Folia Primatologica. 68 (1): 34–43. doi:10.1159/000157230.
  24. ^ a b Hawkins; Racey, P.A. (2008). "Food habits of an endangered carnivore, Cryptoprocta ferox, in the dry deciduous forests of western Madagascar". Journal of Mammalogy. 89 (1): 64–74. doi:10.1644/06-MAMM-A-366.1.
  25. ^ Rasoloarison, R.M.; Rasolonandrasana, B.P.N.; Ganzhorn, J.U.; Goodman, S.M. (1995). "Predation on vertebrates in the Kirindy Forest, western Madagascar" (PDF). Ecotropica. 1: 59–65. Retrieved 2010-05-21.
  26. ^ Irwin, M.T.; Raharison, J.L.; Wright, P.C. (2009). "Spatial and temporal variability in predation on rainforest primates: do forest fragmentation and predation act synergistically?". Animal Conservation. 12 (3): 220–230. doi:10.1111/j.1469-1795.2009.00243.x.
  27. ^ a b Ruud 1970, p. 101.
  28. ^ a b Jones, J.P.G.; Andriamarovolona, M.A.; Hockley, N.J. (2007). Taboos, social norms and conservation in the eastern rainforests of Madagascar (PDF). 9th International BIOECON Conference on "Economics and Institutions for Biodiversity Conservation". Retrieved 19 May 2010. {{cite conference}}: External link in |conferenceurl= (help); Unknown parameter |conferenceurl= ignored (|conference-url= suggested) (help)
Books cited
  • Albignac, R. (1973). Faune de Madagascar (in French). Vol. 36 Mammifères. Carnivores. pp. 1–206. ASIN B000LPMXS6.
  • Goodman, S.M.; Benstead, J.P., eds. (2003). The Natural History of Madagascar. University of Chicago Press. ISBN 0-226-30306-3.
Chapter 13 – Mammals
  • Yoder, A.D.; Flynn, J.J. (2003). "Origin of Malagasy Carnivora". pp. 1253–1256. {{cite book}}: Invalid |ref=harv (help); Missing or empty |title= (help)
  • Hawkins, C.E. (2003). "Cryptoprocta ferox, Fossa, Fosa". pp. 1360–1363. {{cite book}}: Invalid |ref=harv (help); Missing or empty |title= (help)
  • Primate Anti-Predator Strategies (Developments in Primatology: Progress and Prospects). Springer. 2007. ISBN 978-0-387-34807-0. {{cite book}}: |editor1-first= missing |editor1-last= (help); |editor2-first= missing |editor2-last= (help); Unknown parameter |editor1_last= ignored (help); Unknown parameter |editor2_last= ignored (help)
  • Dollar, L.; Ganzhorn, J.U.; Goodman, S.M. (2007). "Primates and other prey in the seasonally variable diet of Cryptoprocta ferox in the Dry deciduous forest of Western Madagascar". pp. 63–76. doi:10.1007/978-0-387-34810-0. {{cite book}}: Invalid |ref=harv (help); Missing or empty |title= (help)
  • Ruud, J. (1970). Taboo: A Study of Malagasy Customs and Beliefs (2nd ed.). Oslo University Press. ASIN B0006FE92Y. {{cite book}}: Invalid |ref=harv (help)
Wikimedia Commons has media related to Cryptoprocta ferox.
Wikispecies has information related to Cryptoprocta ferox.
  • ARKive - images and movies of the Fossa (Cryptoprocta ferox)

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