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
7–31 in (18–80 cm); 7 oz–26.5 lb (200 g–12 kg)

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

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 Herpestidae into three subfamilies and recent molecular evidence supports this division. The subfamilies include: (1) the Herpestinae, 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

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 expanded

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

Near East. The Galidiinae are restricted to the large island of Madagascar. The small Indian mongoose, Herpestes javinicus, 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 unspecialized 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 (Herpestes 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 exception 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. Living

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.

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 penicillata) 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.

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

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 Endangered 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.

Species accounts

List of Species

Ring-tailed mongoose

Galidia elegans

subfamily

Galidiinae

taxonomy

Galidia elegans Geoffroy, 1837, Madagascar.

other common names

French: Galidie élégante; German: Ringelschwanzmungo; Spanish: Mangosta de cola anillada; Malagasy: Vontsira mena.

physical characteristics

Body length 11.8 in (30 cm); tail 8.3 in (21 cm); weight 2 lb (900 g). Fur is soft russet-brown. Tail has five or six black rings.

distribution

Northern, eastern, and west-central Madagascar.

habitat

Lives in humid forests.

behavior

Diurnal. Living in pairs or small family groups lead by female.

feeding ecology and diet

Diet consists of small mammals, birds, frogs, reptiles, aquatic invertebrates, fish, insects, and millipedes.

reproductive biology

Monogamous. One young is born, fully furred, with eyes closed. Mature at one year.

conservation status

Listed as Vulnerable due to habitat loss.

significance to humans

None known.

Fossa

Cryptoprocta ferox

subfamily

Galidiinae

taxonomy

Cryptoprocta ferox Bennett, 1833, Madagascar.

other common names

Malagasy: Fosa; French: Fousa; German: Frettkatze; Spanish: Fossa gatuno.

physical characteristics

Largest herpestid, 5–6.6 ft (1.5–2 m) with tail. Weight 15.4–26.5 lb (7–12 kg). Coat is reddish brown. Slender body and short legs, square muzzle and round ears. Canines and carnassials are well developed.

distribution

All of Madagascar except High Plateaux.

habitat

Lives in forest and woodland.

behavior

Crepuscular and nocturnal, solitary. Lives in the trees and on the ground.

feeding ecology and diet

Diet includes lemurs, small mammals, birds, reptiles, amphibians, and insects.

reproductive biology

Polygamous. Females have one litter per year of two to four young born November to January. Weaning at 4.5 months. Adulthood reached at three years.

conservation status

Listed as Endangered due to habitat loss and human persecution.

significance to humans

None known.

Small Indian mongoose

Herpestes javanicus

subfamily

Herpestinae

taxonomy

Herpestes javanicus (Geoffroy, 1818), Java.

other common names

English: Javan gold-spotted mongoose; French: Petite mangouste indienne; German: Indien Goldstaub-Manguste; Spanish: Mangosta javanés.

physical characteristics

Body length 15.4 in (39 cm); tail 10.2 in (26 cm); weight 1.8 lb (800 g). Coat is brown to reddish brown, speckled with black and gray hair tips.

distribution

Malayan and Indo-Chinese Peninsulas, Sumatra, Java. Introduced to West Indies, Hawaiian Islands, Fiji, Comores, Costa Rica, Japan, and Mauritius.

habitat

Habitat generalist. Found in arid plains to wet tropical forests.

behavior

Diurnal and solitary. Home ranges are 0.62 mile (1 km) wide.

feeding ecology and diet

Feeds on invertebrates, birds, rodents, and reptiles, including venomous snakes.

reproductive biology

Females reproductive as early as 10 weeks. Breed two or three times per year. Litter of two to four young weaned in one month. Mating system is not known.

conservation status

Not threatened.

significance to humans

Where introduced, responsible for extinction of several endemic island species, killing of poultry, and the spread of rabies. Estimated to cause $50 million in damage every year in Puerto Rico and the Hawaiian Islands alone.

Dwarf mongoose

Helogale parvula

subfamily

Mugotinae

taxonomy

Helogale parvula (Sundevall, 1846), South Africa.

other common names

Swahili: Kitafe; French: Mangouste nain; German: Zwergichneumon; Spanish: Mangosta enana.

physical characteristics

Body length 7–11 in (18–28 cm); tail 5.5–7.5 in (14–19 cm); weight 7–10 oz (200–300 g). Grizzled fur is reddish brown. Adults have juvenile features, including a short muzzle and large head.

distribution

Ethiopia to Angola and eastern South Africa.

habitat

Lives in dry savanna and woodland.

behavior

Diurnal and highly social. Group is led by matriarch.

feeding ecology and diet

Eats mostly insects, and other arthropods, but some small vertebrates are consumed.

reproductive biology

Monogamous, only alpha pair breed. Cooperative care of young by group members. Up to three litters per year, each up to six young.

conservation status

Not threatened.

significance to humans

None known.

Liberian mongoose

Liberiictis kuhni

subfamily

Mungotinae

taxonomy

Liberiictis kuhni Hayman, 1958, Liberia.

other common names

French: Mangouste de Liberia; German: Liberia-kusimanse; Spanish: Mangosta de Liberia.

physical characteristics

Body length 15.7–17.7 in (40–45 cm); tail 7–8.7 in (18–22 cm); weight 4.4–5 lb (2–2.3 kg). Coarse brown coat with grizzled, stiff guard hairs. Black stripe on neck. Tapering tail. Snout is long and mobile, extending beyond lower lip.

distribution

Liberia, western Ivory Coast, and southern Guinea.

habitat

Found in riverine and swamp forest.

behavior

Diurnal. Living in small family groups of four to eight individuals.

feeding ecology and diet

Specialist on large burrowing earthworms and other soft invertebrates.

reproductive biology

Breeding thought to occur during the rainy season (May– September) during peak earthworm availability. Mating system is not known.

conservation status

Listed as Endangered due to habitat degradation and hunting pressures.

significance to humans

Subject to hunting for bushmeat.

Resources

Books

Albignac, R. "The Carnivores." In Madagascar. (Key Environments), edited by Alison Jolly, Philippe Oberle, and Roland Albignac. Oxford: Pergamon Press, 1984.

Ewer, R. F. The Carnivores. Ithaca, NY: Cornell University Press, 1973.

Gittleman, J. Carnivore Behavior, Ecology and Evolution. Vols. 1 & 2. Ithaca, NY: Cornell University Press, 1989.

Kingdon, J. The Kingdon Field Guide to African Mammals. San Diego: Academic Press, 1997.

Periodicals

Bdolah, A., E. Kochva, M. Ovadia, S. Kinamon, and Z. Wollberg. "Resistance of the Egyptian Mongoose to Sarafotoxins." Toxicon 35 (1997): 1251–1261.

Yoder, A., et al. "Single Origin of Malagasy Carnivora from an African Ancestor." Nature 421 (2003): 734–737.

Amy E. Dunham