Mongooses and fossa

(Herpestidae)

Class Mammalia Order Carnivora Suborder Feloidea Family Herpestidae

Thumbnail description

Small to medium-sized carnivores with long bodies, short legs, and highly tapered snouts; fur is often grizzled and brown or gray in color

Size

Number of genera, species

20 genera; 41 species

Habitat

Savanna, forest, open woodland, and semi-desert

Conservation status

Endangered: 6 species; Vulnerable: 5 species; Data Deficient: 1 species

World Map Where Pangolins Live

Distribution

Africa, Asia, Madagascar, and southern and eastern Mediterranean region

Distribution

Africa, Asia, Madagascar, and southern and eastern Mediterranean region

Evolution and systematics

Though once considered in the same family as civets and genets (Viverridae), mongooses are now recognized as a separate family (Herpestidae). This family is comprised of 21 genera, including about 41 species, depending on the authority. Several authors have suggested the division of Her-pestidae into three subfamilies and recent molecular evidence supports this division. The subfamilies include: (1) the Her-pestinae, primarily solitary or pair-living mongooses (genera: Atilax, Bdeogale, Cynictis, Herpestes, Ichneumia, and Rhynchogale); (2) the Mungotinae, small, highly social mongooses (genera: Crossarchus, Dologale, Helogale, Liberiictis, Mungos, Paracynictis, and Suricata); and (3) the Galidiinae, consisting of all endemic Malagasy carnivores (genera: Cryptoprocta, Eupleres, Galadictis, Galidia, Mungotictis, and Salanoia).

Despite the notable variation in physical appearance, the endemic carnivores of Madagascar arose from a single African ancestor 24-18 million years ago (mya). Their common ancestry and close phylogenetic relationship to the mongooses requires placement of the large cat-like fossa (Cryptoprocta ferox) within the Herpestidae, even though, superficially, it bears little resemblance to its mongoose kin. In Africa and Asia, herpestid evolution may have been constrained by competition with other carnivore families that dominated several specialized niches. As a result, mongoose radiation in Africa and Asia involved numerous, but rather subtle adaptations and specializations in diet and habitat, most evident in morphology of ears, muzzle, whiskers, jaw, and teeth. However, in Madagascar, the lack of canids, felids, or mustelids allowed a wider radiation of body forms and niches, from small, squirrel-like insectivores (Mungotictis) to large, arboreal predators (Cryptoprocta ferox).

Physical characteristics

Mongooses are small to medium-sized carnivores. Body lengths vary from 7 to 31 in (18 to 80 cm) and weights range

A meerkat (Suricata suricatta) community. (Photo by Harald Schutz. Reproduced by permission.)
Fossa (Cryptoprocta ferox) are the largest native predators of Madagascar. (Photo by Harald Schütz. Reproduced by permission.)
An eastern Malagasy ring-tailed mongoose (Galidia elegans elegans) feeding on rodent. (Photo by Harald Schütz. Reproduced by permission.)

panded and is as large or larger than the entotympanic element. A median lacerate foramen is present. The teeth, numbering 34-40, are distinctive for each genus. Carnassials are well developed and most species have an internal cusp on the third upper premolar that is variable in size and is vestigial in some species.

Distribution

The family is a widespread successful group but confined to the Old World. Most mongooses are African, occupying the entire continent except for the Sahara. Only one genus (Herpestes) exists in Asia, ranging from the Philippines and Borneo to Southeast Asia, south China, Sri Lanka, India and Arabia. It is also present in southern Spain, Portugal, and the from 7 oz to 26.5 lb (200 g-12 kg). Most have long, slender bodies and relatively short legs. The ears are small and rounded and the snout is highly tapered. Eyes of most species contain ovular-shaped horizontal pupils. The fur is generally coarse and grizzled and the color often correlates with the local soil, indicating the importance of camouflage. The feet, legs, tail or tail tip are commonly a different hue. A few species have coats bearing stripes (Mungotictis, Galadictis, Suricata, Mungos), or ringed tails (Galidia), but the majority lack strongly marked coats. Mongooses have four to five toes that do not bear retractile claws, although they are semi-retractile in Cryptoprocta. Undersides of the feet tend to be hairless. Unlike the civets and genets, mongooses lack perineal civetone glands; however, a well-developed anal pouch containing at least two glandular openings is present in all species. Scent deposits from the anal pouch function as communication for both solitary and highly social species.

The structure of the auditory bulla is important in characterizing the family. There is a clear demarcation between the entotympanic and ectotympanic parts of the auditory bulla, which is perpendicular to the long axis of the skull (it is oblique in Viverridae). The ectotympanic element is ex-

Yellow mongoose (Cynictis penicillata) mother and young in Etosha National Park, Namibia. (Photo by Jen and Des Bartlett. Bruce Coleman, Inc. Reproduced by permission.)
A banded mongoose clan (Mungos mungo) foraging in the early morning in Botswana. (Photo by Clem Haagner. Bruce Coleman, Inc. Reproduced by permission.)

Near East. The Galidiinae are restricted to the large island of Madagascar. The small Indian mongoose, Herpestes javini-cus, was introduced by man in the 1800s to Fiji, the Hawaiian Islands, and several islands in the West Indies for biological control of rats in sugarcane plantations. However, they are now considered pests in these areas as their unspe-cialized diets have rendered them a threat to many native birds and reptiles, as well as to domestic fowl.

Habitat

Mongooses are primarily terrestrial but the marsh mongoose (Atilax paludinosus), Bengali water mongoose (Herpestes palustris), and ring-tailed mongoose (Galidia elegans) are semi-aquatic, spending much time in streams and rivers hunting for aquatic invertebrates. Madagascar's fossa is arboreal and equipped with semi-retractile claws, and a long tail for movement and balance in the trees. The slender mongoose (Her-pestes sanguinus), though chiefly terrestrial, is also adept at climbing trees in search of food.

Mongooses inhabit a variety of ecotypes including forest, open woodland, savanna, semi-desert, and desert. The Mungotinae are restricted to open habitats with the excep tion of the cusimanses (Chrossarchus sp.) and the Liberian mongoose (Liberiictis kuhni), which inhabit the rainforest interior. The Herpestinae occupy a wider range of habitats from rainforest to semi-desert. Madagascar's Galidiinae can be found in rainforest, dry forest, and spiny desert.

Sleeping dens and birth dens often include tree hollows and tree falls in forested regions, and rock crevices, earth holes, and termitaries in open terrain. For many species, presence of denning sites are very important and may be more limiting for a population than food resources.

Behavior

Most mongooses are solitary but some species live in pairs and several form large, stable social groups that forage and den together. The gregarious species of mongooses have social systems that rival only the primates in diversity and complexity. Group living seems to have evolved for reasons unlike the larger social carnivores that primarily benefit from communal hunting. Predation pressure seems more likely to have favored group living in mongooses. Social mongooses tend to be small, diurnal species inhabiting open areas, characteristics that make them extremely vulnerable to predation. Liv-

A dwarf mongoose (Helogale parvula) emerging from its burrow. (Photo by Animals Animals ©Michael Dick. Reproduced by permission.)

ing in a group provides some protection and early warning of a predator attack. It has also been proposed that diet is relevant to the development of sociality in these animals. The high abundance and renewability of invertebrate prey reduces the cost of sharing a territory. Mongooses preying upon vertebrate prey tend to be solitary, as the presence of another individual may interfere with a hunt. Diurnal habits and open habitat also facilitate keeping contact, an important part of social living. There are no nocturnal social species; noisy social interactions at night may pose a risk as nocturnal predators often rely on keen sense of hearing. The social cusimanses do not live in open habitat but are able to maintain contact in the dense rainforest understory by giving constant whistling calls while traveling.

Galidictis

A western Malagasy broad striped mongoose (Galidictis grandidieri) grooming. (Photo by Harald Schütz. Reproduced by permission.)

Another advantage to group living in mongooses is increased efficiency in the care of young. In most species, young are born rather helpless and leaving them alone in a den while foraging may be risky. In several species, such as the dwarf mongoose (Helogale parvula), the helpless young are guarded by babysitters while the rest of the group forages. Group members also help feed the young by bringing them insects or worms to eat. For species that occupy no permanent den site, such as the cusimanse, young are not able to keep up with the group for several weeks and must be carried to different foraging spots. Individuals in the group take turns carrying the young from place to place and also help feed them.

For many of the highly social species, such as the yellow mongoose (Cynictis penicittata) and dwarf mongoose, a strict hierarchy develops with a single dominant breeding pair. Sexual dimorphism is minimal in most genera and dominance is based more on age, size, and assertiveness than on gender. Female dominance is not uncommon.

The size of groups is variable among species. Three to eight individuals have been recorded in Liberian mongoose groups, and up to 40 have been recorded for the banded mongoose (Mungos mungo). Pair living is also common. In the ring-tailed mongoose of Madagascar (Galidia elegans), strong pair bonds are formed between mates who forage and den together. Even the "solitary" species are occasionally seen foraging in pairs, and some species such as the Cape gray mongoose (Herpestes pulverulentus) and the white-tailed mongoose (Ichneumia albicauda) may forage individually but den with family members. Communal denning may offer better protection and added warmth and reduces competition for denning sites.

All species communicate through scent marking with the anal glands. Some species also have cheek and chest glands.

An Indian mongoose (Herpestes nyula) killing a cobra. (Photo by E. R. Degginger. Bruce Coleman, Inc. Reproduced by permission.)

A western Malagasy broad striped mongoose (Galidictis grandidieri) grooming. (Photo by Harald Schütz. Reproduced by permission.)

An Indian mongoose (Herpestes nyula) killing a cobra. (Photo by E. R. Degginger. Bruce Coleman, Inc. Reproduced by permission.)

When a jackal approaches, banded mongooses move towards it in a tightly knit group, creating the appearance of a larger animal from which the jackal will retreat. (Illustration by Marguette Dongvillo)

Chemicals in the anal pouch may constitute individual signatures and indicate reproductive condition, sex, and/or dominance rank. Mongooses commonly mark territory borders, den sites, food resources, and even other group members. Social species have a complex vocal repertoire with several distinct calls.

Feeding ecology and diet

Most herpestids are opportunistic predators, feeding on small vertebrates including rodents, birds, reptiles, and frogs, and various invertebrates such as insects, snails, crabs, and worms. All species are predacious, but a few species also eat plant matter such as fruits and tubers.

Many mongoose species can open bird eggs by hurling the egg towards a hard fixed object with the forepaws, usually between the hind legs. The banded mongoose has been observed hurling stones in the same manner at an ostrich egg that was too large to throw.

Mongooses are most famous for preying on venomous snakes such as cobras, and some species have actually developed resistance to snake venom. The Egyptian mongoose (Herpestes ichneumon) and the Indian gray mongoose (Herpestes

A Malagasy narrow striped mongoose (Mungotictis decemlineata) feeding on excavated lizard eggs. (Photo by Harald Schütz. Reproduced by permission.)

edwardsii) have both been found resistant to hemorrhagic and neurotoxic snake venoms and can withstand up to 13 times the normal lethal dose for mammals.

Although most mongooses are opportunistic feeders on vertebrate and invertebrate prey, a few species have quite specialized diets. The Liberian mongoose has specialized dentition and reduced jaw musculature as an adaptation for eating almost exclusively large, burrowing earthworms. The fossa is specialized with its large canines and cat-like morphology to prey upon large arboreal lemurs.

Reproductive biology

Mating system is variable by species (monogamous or polygamous). Sexual maturity is generally achieved at two years of age. However, the small Indian mongoose may reproduce as early as 10 weeks. Most species are polyestrus, having two or more litters per year. A few are highly seasonal, breeding only when food is most abundant. Copulation is usually preceded by increased frequency of scent marking and some form of chasing or ritualized fighting. During mating, the male clasps the female's back just forward of the pelvic region with his forepaws and grips the back or side of her neck with his mouth without biting. Litter sizes range from one (common in the Galidiinae) to six, with four being most common. All species use a birthing den than may consist of a burrow, termitary, hollow log, or tree. Young are born blind and with little hair in most species, with eyes not opening until week two. In contrast, offspring of the narrow-striped mongoose (Mungotictis decemlineata) are born fully furred with eyes open at birth, and walk by day three.

Conservation status

Though generally a successful group, several species are at risk due to loss of habitat. Six species are listed as Endan gered and five species are listed as Vulnerable by the World Conservation Union (IUCN). Several more may also be at risk and many are lacking sufficient data to be evaluated. All eight Galidiinae species are threatened (four Vulnerable and four Endangered) because of the high levels of habitat destruction and fragmentation occurring in Madagascar. The Liberian mongoose and Jackson's mongoose of Africa, and the Bengali water mongoose of India have been classified as threatened because of habitat loss. Until recently, mongooses had been largely ignored by the scientific community and as a result, few data exist on population sizes or distributions. Of the mongooses that have not yet been evaluated, those restricted to undisturbed forests or other specialized habitats subject to human disturbance are most likely to be at risk.

Significance to humans

The mongoose has had a long relationship with humans, perhaps starting with the ancient Egyptians. It is found in frescoes and reliefs dating as far back as 2800 B.C. The animal was often embalmed in Egypt and believed to embody a number of gods. It may also have been used for hunting birds. The mongoose's ability to fight snakes gave rise to numerous fables; the oldest and most familiar stories found in the fifth book of the Panchatantra, a group of Sanskrit tales dating from approximately 100 B.C. The mongoose is portrayed as a hero, protecting man from cobra, a story later recounted in Rudyard Kipling's famous tale Rikki Tikki Tavi.

Mongooses are easily tamed and frequently kept as pets in both Asia and Africa, as companions and to keep away venomous snakes. They are hunted as food in some parts of Africa.

Marsh Mongoose
A marsh mongoose (Atilax paludinosus) on guard in Marsai Mara, Kenya. (Photo by Peter Davey. Bruce Coleman, Inc. Reproduced by per-

1. Ring-tailed mongoose (Galidia elegans); 2. Fossa (Cryptoprocta ferox); 3. Liberian mongoose (Liberiictis kuhni); 4. Dwarf mongoose (Helogale parvula); 5. Small Indian mongoose (Herpestes javanicus). (Illustration by Marguette Dongvillo)

Ring-tailed mongoose

Galidia elegans

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