The extant native carnivores of Madagascar form a monophyletic group and together comprise the family Eupleridae (there are also three carnivore species that have been introduced to Madagascar by humans: the domestic dog [Canis lupus familiaris]; the Wildcat [Felis silvestris], which was introduced in the 19th century and is not uncommon in natural forest habitats; and the Small Indian Civet [Viverricula indica], which is generally found in heavily degraded and open edges or at the forest edge). Goodman (2009) recognized 8 euplerid species in seven genera and noted that at least one more form was likely to be recognized as a distinct species after further investigation.
The carnivores of Madagascar have been surrounded by considerable taxonomic confusion over the years as a result of what is now believed to be striking convergent evolution between this highly isolated carnivore lineage in Madagascar (currently separated from the mainland by around 400 km of open water) and the evolution of carnivores in other parts of the world. This convergence in features resulted in various members of the Madagascar carnivore lineage resembling cats (family Felidae), civets (family Viverridae), or mongooses (family Herpestidae) from "off-island" as they adapted to fill similar ecological niches (similar striking examples of convergent evolution can be seen in the similarities between various species in the Australian marsupial lineage and "ecological equivalents" evolving elsewhere). The recent molecular phylogenetic data indicating that all the Madagascar carnivores have a single carnivore ancestor rather than multiple ones implies a scenario requiring just a single ancient colonization from the distant continent of Africa rather than several. Unfortunately, although researchers have worked to identify shared derived morphological characters (i.e., "synapomorphies") uniting this group, which would support the monophyly indicated by the molecular data, no such characters have been identified. The sister group to the Eupleridae is the Herpestidae; sister to (Eupleridae + Herpestidae) is Hyaenidae (Agnarsson et al. 2010; Eizirik et al. 2010).
Goodman (2009) reviewed the history of the taxonomic treatment of Madagascar's native carnivores.The largest and best known living euplerid, the Fosa (Cryptoprocta ferox), is nearly the size of a small Puma and was at one time believed to be a felid (at other times it was placed in its own family or considered to be a viverrid or herpestid). The Spotted Fanaloka (Fossa fossana) and Falanouc (Eupleres goudotii) were believed to be viverrids and the Ring-tailed Vontsira (Galidia elegans), Broad-striped Vontsira (Galidictis fasciata), Grandidier's Vontsira (Galidictis grandidieri), Brown-tailed Vontsira (Salanoia concolor), and Narrow-striped Boky (Mungotictis decemlineata) were all believed to be herpestids. Today, species in the first three genera above are included in the subfamily Euplerinae whereas the mongoose-like species in the latter four genera are placed in the subfamily Galidiinae. By the middle of the 19th century, six of the eight euplerids were known to scientists and formally described. The Narrow-striped Boky was described in 1867 and Grandidier's Vontsira in 1986 (this latter species has a conservation status of Endangered, with a range of less than 500 km2 that is essentially a single location in extreme southwestern Madagascar). The Giant Fosa (Cryptoprocta spelea) is known only from subfossil remains and apparently went extinct within the past few thousand years.
The basic biology of some euplerids, such as the vontsiras, remains almost unknown. Conservation assessments are difficult for most euplerids due to a dearth of information, but the enormous loss of forest habitat in Madagascar during the latter half of the 20th century surely had a negative impact impact on all euplerid species, although some more than others. In addition to threats from habitat loss, most (possibly all) euplerids are consumed as bushmeat. They are all generally viewed by local people as vermin and blamed for preying on domestic animals, especially fowl, although much of this predation can actually be attributed to the introduced Small Indian Civet. The small geographic range of Grandidier's Vontsira may be due more to natural ecological conditions rather than to human impacts on the environment.
(Goodman 2009 and references therein)
Euplerids tend to have slender bodies with relatively small heads and pointed rostra, but fossas look more cat-like and have with blunt snouts. The thick, soft pelage is gray or brown and is spotted or striped in all but Eupleres and Cryptoprocta. The foot posture is plantigrade or digitigrade. The male is larger than the female.
Euplerids live in various habitats, from humid forests, marshes, bogs, and swamps to deserts and savannahs.
Like other carnivores, euplerids can perceive visual, acoustic, chemical and tactile signals. They communicate via scent in the form of glandular secretions, such as in scent marks, and through various cries, groans and other vocalizations.
Euplerids are primarily carnivorous, consuming small mammals, birds, reptiles, frogs, as well as insects, crustaceans and other invertebrates. Eupleres goudotii specializes in eating earthworms and other intertebrates, while Fossa fossana and Galidia elegans may eat fruit as well as animal matter.
Humans, domestic dogs and possibly other predators, such as birds of prey and large snakes, eat euplerids. The fossa is Madagascar's largest native predator and, except for humans, is at the top of the food chain. Cryptic coloration (spots and stripes on neutral backgrounds) probably conceals most species well. The eastern falanouc runs or freezes when disturbed, while the fossa releases a foul-smelling substance from its anal glands when it is alarmed.
Mating systems in euplerids vary with social structure. Malagasy civets form monogamous pairs. Fossas are strictly solitary, suggesting a polygynous or polygynandrous mating system. Species in Galidia, Mungotictis and Salanoia live alone or in pairs, suggesting that they are monogamous within, but not across, breeding seasons. Species in Eupleres and Galidictis live alone, in pairs or in small family groups, which may indicate monogamy. Mungotictis individuals live in small groups with several adults of each sex, but it is not known if all adult group members breed. Euplerids have definite breeding seasons, lasting 2-8 months, depending on species. Usually 1-2 young are born after 3 months, but the fossa can have up to 4 young. Weaning takes place between 2 and 4.5 months. Females of some genera, such as the fossa, select dens where they bear and nurse their young. Narrow-striped mongooses live in family groups of several adults, juveniles, and young, so offspring have an association with their parents beyond weaning. Lifespans of euplerids in the wild are unknown. Fossas may live 20 years in captivity.
The fossa may attack poultry. All euplerid species are considered to be threatened, due to habitat destruction, predation by dogs, hunting by humans for meat and competition from non-native species. The IUCN considers Fossa fossana, Galidia elegans, Galidictis fasciata, and Salanoia concolor to be vulnerable and Cryptoprocta ferox, Eupleres goudotii, Galidictis grandidieri, and Mungotictis decemlineata to be endangered, with C. ferox, E. goudotii and F. fossana on on Appendix II of CITES.
Family Eupleridae Subfamily Euplerinae: Fossa (Cryptoprocta ferox), Giant fossa (C. spelea) (extinct) Eastern falanouc (Eupleres goudotii), Western falanouc (E. major), Malagasy civet (Fossa fossana) Subfamily Galidiinae: Ring-tailed mongoose (Galidia elegans), Broad-striped Malagasy mongoose (Galidictis fasciata), Grandidier's mongoose (G. grandidieri), Narrow-striped mongoose (Mungotictis decemlineata), Brown-tailed mongoose, (Salanoia concolor), Durrell's mongoose (S. durrelli)
The family Eupleridae, the Malagasy carnivores, consists of eight species and seven genera in two subfamilies. The diversity of form and function in this family is such that some have suggested it be split into several families.
- Gaubert, P., W. Wozencraft, P. Cordeiro-Estrela, G. Veron. 2005. Mosaics of convergences and noise in morphological phylogenies: What's in a viverrid-like carnivoran?. Systematic Biology, 54(6): 865-894.
- Wilson, D., D. Reeder. 2005. Mammal Species of the World. Baltimore and London: Johns Hopkins University Press.
Species in the family Eupleridae are restricted to the island of Madagascar.
Biogeographic Regions: ethiopian (Native )
Other Geographic Terms: island endemic
- Nowak, R. 1999. Walker's Mammals of the World. Baltimore and London: The Johns Hopkins University Press.
Aside from molecular synapomorphies, euplerids have few traits in common. They tend to have slender bodies with relatively small heads and pointed rostra, although fossas (Cryptoprocta ferox) are more cat-like in appearance, with blunt snouts. Head and body length ranges from 250 mm in Mungotictis and Salanoia to 800 mm in adult male Cryptoprocta. The thick, soft pelage is gray or brown, and spotted or striped in all but Eupleres and Cryptoprocta. The foot posture is plantigrade or digitigrade.
Other Physical Features: endothermic ; homoiothermic; bilateral symmetry
Sexual Dimorphism: male larger
Euplerids live in a variety of habitats, from humid forests, marshes, bogs, and swamps, to deserts and savannahs.
Habitat Regions: temperate ; tropical ; terrestrial
Terrestrial Biomes: desert or dune ; savanna or grassland ; forest ; rainforest ; scrub forest
Wetlands: marsh ; swamp ; bog
- Schreiber, A., R. Wirth, M. Riffel, H. Van Rompaey. 1989. Weasels, Civets, Mongooses, and their Relatives. An Action Plan for the Conservation of Mustelids and Viverrids. Gland, Switzerland: IUCN.
Euplerids are primarily carnivorous, consuming small mammals, birds, reptiles, frogs, insects, crustaceans, and other invertebrates. Eupleres goudotii is specialized for eating intertebrates such as earthworms. Fossa fossana and Galidia elegans may include some fruit in their diets in addition to animal matter.
Primary Diet: carnivore (Eats terrestrial vertebrates, Eats eggs, Insectivore , Eats non-insect arthropods); herbivore (Frugivore ); omnivore
Euplerids that eat fruit are primary consumers, and all euplerids are secondary and higher-level consumers, because they eat a variety of animal matter. They, in turn, are eaten by humans and domestic dogs, and possibly other predators as well. Cryptoprocta ferox is the largest native predator on Madagascar and, except for humans, is at the top of the food chain.
No information is available on the specific predators of euplerids, besides humans and domestic dogs, both of which are not native to Madagascar. Cryptic coloration in the form of spots and stripes on neutral backgrounds probably conceals most species well. Eupleres goudotii is known to either run or freeze when disturbed, and Cryptoprocta ferox releases a foul-smelling substance from its anal glands when it is alarmed. Large birds of prey or large snakes are potential predators.
- humans (Homo sapiens)
- domestic dogs (Canis lupus familiaris)
Anti-predator Adaptations: cryptic
Life History and Behavior
Like other carnivores, euplerids can perceive visual, acoustic, chemical, and tactile signals. Communication is through scent in the form of glandular secretions and through a variety of cries, groans, and other vocalizations.
Communication Channels: acoustic ; chemical
Other Communication Modes: scent marks
Perception Channels: visual ; tactile ; acoustic ; chemical
Euplerid lifespans in the wild are unknown. Cryptoprocta holds the longevity record in captivity, at 20 years.
Mating systems in Eupleridae vary, as social structure varies from species to species. Fossas form monogamous pairs, while species in the genera Galidia, Mungotictis, and Salanoia are either found alone or in pairs, suggesting that they are monogamous within, but not across, breeding seasons. Species in the genera Eupleres and Galidictis live alone, in pairs, or in small family groups, which also might indicate monogamy. Mungotictis individuals live in small groups with several adults of each sex, but it is not known whether all of the adults within a group breed. Cryptoprocta individuals are strictly solitary, suggesting a polygynous or polygynandrous mating system.
Mating System: monogamous ; polygynous ; polygynandrous (promiscuous)
Euplerids have definite breeding seasons, which vary by species and last anywhere from two to eight months. Gestation lasts around three months. Usually there are just one or two young per litter, though Cryptoprocta can have up to four. Weaning takes place between two and four and a half months.
Key Reproductive Features: semelparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization ; viviparous
Like all eutherian mammals, euplerid females nuture their young through a placenta until the young are born. They then provide their offspring with milk for two to four and a half months. Females of some genera, such as Cryptoprocta, select dens in which to bear and nurse their young. Mungotictis individuals live in family groups of several adults, juveniles, and young; thus, offspring have an association with their parents beyond weaning.
Parental Investment: pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Male, Female); pre-independence (Provisioning: Female, Protecting: Male, Female); post-independence association with parents
- Nowak, R. 1999. Walker's Mammals of the World. Baltimore and London: The Johns Hopkins University Press.
Molecular Biology and Genetics
Statistics of barcoding coverage
Specimens with Sequences:5
Specimens with Barcodes:0
Species With Barcodes:3
All species in this family are threatened. Fossa fossana, Galidia elegans, Galidictis fasciata, and Salanoia concolor are considered vulnerable by the IUCN, and Cryptoprocta ferox, Eupleres goudotii, Galidictis grandidieri, and Mungotictis decemlineata are considered endangered. Cryptoprocta ferox, Eupleres goudotii, and Fossa fossana are on Appendix II of CITES. Habitat destruction in the form of deforestation is the main cause of their decline, though hunting by humans and predation by dogs also impact populations.
- IUCN, 2006. "2006 IUCN Red List of Threatened Species" (On-line). Accessed September 19, 2006 at www.redlist.org.
Relevance to Humans and Ecosystems
Cryptoprocta ferox individuals sometimes attack poultry. There are no reports of other euplerids negatively affecting humans.
Euplerids are commonly hunted for their meat.
Positive Impacts: food
Eupleridae is a family of carnivorans endemic to Madagascar and comprising 10 known living species in seven genera. The best known species is the fossa (Cryptoprocta ferox), in the subfamily Euplerinae. All species of Euplerinae were formerly classified as viverrids, while all species in the subfamily Galidiinae were classified as herpestids.
Recent molecular studies indicate the 10 living species of Madagascar carnivorans evolved from one ancestor that is thought to have rafted over from mainland Africa 18-24 million years ago. This makes Malagasy carnivorans a clade. They are closely allied with the true herpestid mongooses, their closest living relatives. The fossa and the Malagasy civet (Fossa fossana) are believed to be the most ancient surviving species within this group.
Taxonomy and phylogeny
Historically, the relationships of the Madagascar carnivorans have been contentious, but molecular evidence suggests they form a single clade, now recognized as the family Eupleridae. The hyena family Hyaenidae is a sister taxon of the euplerid and herpestid clade, and when grouped together with the viverrids and felids, as well as some smaller groups, forms the feliform (cat-like carnivores) clade.
The evolutionary divergence between the herpestids and the euplerids dates back to the Oligocene;. At that time, feliforms shared many similarities, particularly between the cats and the viverrids. Palaeoprionodon (within superfamily Aeluroidea), found in Europe and Asia from the late Eocene or early Oligocene, looked similar to the modern fossa, while Proailurus, an extinct form of cat, exhibited many viverrid-like characteristics. Despite these similarities in the fossil record, the modern Malagasy carnivores are distinctly different, with the Euplerinae and Galidiinae subfamilies bearing similarities with civets and mongooses, respectively. Species in Euplerinae (including the fossa, falanouc, and Malagasy civet) have auditory regions similar to those of viverrids, while those in Galidiinae have auditory regions similar to those of herpestids. Based on this trait, Robert M. Hunt Jr. proposed in 1996 that Madagascar was colonized twice, once by viverrids and once by herpestids. However, the genetic studies by Yoder and colleagues in 2003 suggested a single colonization event occurred by a primitive herpestid ancestor, which was quickly followed by adaptive radiation. The common ancestor arrived from Africa, probably by rafting, during the late Oligocene or early Miocene (24–18 Mya), though Philippe Gaubert and Veron estimated a divergence date of 19.4 Mya (16.5–22.7 Mya).
- Family Eupleridae
- Subfamily Euplerinae
- Subfamily Galidiinae
|Phylogeny of Malagasy carnivorans (Eupleridae)|
|Phylogeny of Eupleridae within Feliformia|
- Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. pp. 532–628. ISBN 978-0-8018-8221-0. OCLC 62265494.
- Flynn, J; Finarelli, J. A.; Zehr, S; Hsu, J; Nedbal, M. A. (April 2005). "Molecular phylogeny of the carnivora (mammalia): assessing the impact of increased sampling on resolving enigmatic relationships". Syst. Biol. 54 (2): 317–37. doi:10.1080/10635150590923326. PMID 16012099.
- Yoder, AD; Burns, A.D.; Zehr, M.M.; Delefosse, S.; Veron, T.; Goodman, G.; Flynn, S.M. (2003). "Single origin of Malagasy Carnivora from an African ancestor" (PDF). Nature 421 (6924): 734–737. doi:10.1038/nature01303. PMID 12610623. Retrieved 19 May 2010.
|last8=in Authors list (help)
- Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. pp. 559–561. ISBN 978-0-8018-8221-0.
- Yoder, A.D.; Flynn, J.J. (2003). "Origin of Malagasy Carnivora". In Goodman, S.M.; Benstead, J.P. The Natural History of Madagascar. University of Chicago Press. pp. 1253–1256. ISBN 0-226-30306-3.
- Barycka, E. (2007). "Evolution and systematics of the feliform Carnivora". Mammalian Biology 72 (5): 257–282. doi:10.1016/j.mambio.2006.10.011.
- Köhncke, M.; Leonhardt, K. (1986). "Cryptoprocta ferox" (PDF). Mammalian Species (254): 1–5. Retrieved 19 May 2010.
- Gaubert, P.; Veron, G. (2003). "Exhaustive sample set among Viverridae reveals the sister-group of felids: the linsangs as a case of extreme morphological convergence within Feliformia" (PDF). Proceedings of the Royal Society B 270 (1532): 2523–2530. doi:10.1098/rspb.2003.2521. PMC 1691530. PMID 14667345.
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