Civets, Genets, and Linsangs (Viverridae)

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Civets, genets, and linsangs

(Viverridae)

Class Mammalia

Order Carnivora

Family Viverridae


Thumbnail description
Small to medium carnivores resembling the Mustelidae but limited to the Old World; body and tail are long, legs are short, and face is elongated with a pointy nose and conspicuous ears; claws are short and sharp; many species with long, banded tails

Size
13–35 in (33–88 cm), 1.3–30 lb (0.6–13 kg)

Number of genera, species
20 genera; 34 species

Habitat
Forests, steppes, and brushland

Conservation status
Critically Endangered: 1 species; Endangered: 4 species; Vulnerable: 6 species; Data Deficient: 3 species

Distribution
Western Europe, Africa, Arabia, and Southeast Asia

Evolution and systematics

The Viverridae is an old and primitive carnivore family for which the fossil record is scarce. First appearance of Viverridae in Europe and Asia occurs in the early Oligocene and in Africa in the early Miocene. Because of incomplete records, the place of origin of viverrids in the Old World is unknown. Extant species resemble fossil forms, suggesting that skeletal morphology and tooth structure has remained unchanged for 40–50 million years. The diversity within the family is explained by the variety of the niches occupied, from truly terrestrial, partly to mostly arboreal, to aquatic. The mongooses were once considered in the Viverridae under a different subfamily, the Herpestinae. However, most authorities now recognize mongooses under the family Herpestidae.

The current Viverridae is divided into six subfamilies. The Euplerinae consists of two species in two genera, the falanouc (Eupleres goudotii) and the Malagasy civet (Fossa fossana). The Hemigalinae includes the otter civet (Cynogale bennettii) and three species of palm civets. The Nandiniinae has only one species, the African palm civet (Nandinia binotata), whereas the Paradoxurinae consists of five genera and seven species of palm civets as well as the binturong (Arctictis binturong). Finally, the largest subfamily is the Viverrinae, which consists of seven genera and 20 species of genets (Genetta and Osbornictis), civets (Civettictis, Viverra, and Viverricula), and linsangs (Poiana and Prionodon). Authorities are still debating the position of one species, the fossa (Cryptoprocta ferox), but it is herein considered with the mongooses in the Herpestidae.

Physical characteristics

Civets, genets, and linsangs are characterized by long, sinewy bodies, short legs with strong, sharp claws, long tails, and elongated heads with pointed snouts. Ears typically are fairly large and erect. Most species have short fur with, a slender

appearance, and a long, non-prehensile tail that may exceed the length of the body. One species, the binturong, differs from most viverrids by having exceptionally long fur, stocky appearance, and a prehensile tail. Viverrids have five fingers and five toes. Perineal glands are present, and well developed in numerous species, especially Civettictis civetta, Viverra zibetha, and Viverricula indica. Pelage color is uniform in the binturong, but most species have dark spots, bands, or stripes, and many have banded tails.

Viverrids display adaptations based on the niche they occupy. Terrestrial species such as Civettictis civetta, Viverra zibetha, and Eupleres goudotii have digitigrade feet, non or semi-retractile claws, short tails, and fur with spots or stripes. Arboreal or semi-arboreal species can be separated into two groups, agile species that leap or jump (e.g., Genetta servalina) or less-agile species that grasp branches and move more slowly (Arctictis binturong, Paguma larvata, and Paradoxurus sp.). Agile arboreal species have digitigrade feet, long tails, and spotted fur, whereas less agile species have plantigrade feet, long tails, but uniform fur. Long tails occur in all arboreal species

and help animal maintain their balance. Aquatic species have bare soles, plantigrade feet, uniform pelage, and long tails.

Distribution

The Viverridae are confined to the Old World, and occupy parts of western Europe, most of Africa including Madagascar, Southeast Asia, and Malaysia.

Habitat

The Viverridae occupy tropical forests and densely vegetated areas. Possibly the most flexible species is the common genet (Genetta genetta), which can be found in deciduous forests, plantations, steppes, and parks.

Behavior

The Viverridae are shy animals that are primarily nocturnal. They are solitary, or live in pairs or small groups. Most species are good climbers, some are almost exlusively arboreal (Paradoxurus hermaphroditus and Poiana richardsonii, for example), and two species, the otter civet and the aquatic genet (Osbornictis piscivora), are aquatic. Many species such as Genetta genetta use secretions from their anal glands to scent mark their territories. Scent marks are deposited either passively when moving through vegetation, or actively by squatting and rubbing the anal region on the ground or on prominent objects. Scent marks likely allow assessment of social status, individual recognition, kin recognition, and sexual receptiveness. Vocalizations are well developed in some species.

The behavior and ecology of most species of Viverridae are poorly known, and the family is probably the least known of all carnivores. Most species are nocturnal and shy, and occupy dense vegetation. Because few species are of commercial value to humans, and because most are secretive, few studies have been devoted to gaining a better understanding of this group. Knowledge of spacing patterns and population estimates are unavailable for most species, and much current knowledge arises from opportunistic observations or specimen collections for museums, or from captive animals kept as pets or in zoological gardens.

Feeding ecology and diet

All species are opportunistic feeders, although some specialization occurs toward frugivory in palm civets, to a mostly carnivorous diet in genets. The Viverridae are excellent predators and mostly use their sense of sight and hearing to find prey.

Reproductive biology

Breeding occurs seasonally or throughout the year. Some groups may bear two litters per year, and litter size is one to six. Young are born blind but furred. Little else is known of their reproduction or mating system.

Conservation status

Of the fourteen species of Viverridae listed on the IUCN Red List, one species, the Malabar civet (Viverra civettina) is listed as Critically Endangered, and two species (otter civet and falanouc) as Endangered, mostly because of habitat destruction, predation, and illegal hunting. Many species such as Owston's palm civet (Chrotogale owstoni), Sulawesi palm civet (Macrogalidia musschenbroekii), and Jerdon's palm civet (Paradoxurus jerdoni) are listed as Vulnerable. Three additional species, the Johnston's genet (Genetta johnstoni), Abyssinian genet (Genetta abyssinica), and the aquatic genet (Osbornictis piscivora) are listed as Data Deficient because of lack of information on their abundance and population trends.

Significance to humans

The main importance of this carnivore family to humans is in the production of civet oil, the product from the anal glands of species in at least three genera (Civettictis, Viverra, and Viverricula). The musk, often referred to as "civet," has a sweet smell and is used mostly for the perfume industry and for medicinal purposes. In the seventeenth and eighteenth century, civet was highly sought after for the perfume industry. Today, synthetic replacements have caused the market to decline, but captive civets are still held for civet production. Most of the production originates in Ethiopia. The civet is removed from captive animals by squeezing the protruding anal pockets. Males produce a stronger and better quality civet, and production from individual animals averages 0.14–0.67 oz (4–19 g) per week.

Some species of Viverridae are considered pests in some areas for depredation of poultry. Many species are kept as pets to control rodents and insects around households. Skins of some animals may be used locally by indigenous people, but no species are harvested commercially for their pelts.

Species accounts

List of Species

African civet
Common genet
Aquatic genet
African linsang
African palm civet
Binturong
Otter civet
Falanouc

African civet

Civettictis civetta

subfamily

Viverrinae

taxonomy

Viverra civetta (Schreber, 1776), Guinea.

other common names

English: Civet cat; French: Civette africaine; German: Afrika Zibetkatze; Spanish: Cibeta.

physical characteristics

Body length 27–33 in (67–84 cm), tail 13–19 in (34–47 cm), weight 22–38 lb (10–17 kg). It has medium-length legs and a rather long, erectile dorsal mane. The ash-gray to yellowish basic coloration has numerous black-brown spots on the sides of the body.

distribution

Sub-Saharan Africa.

habitat

Forests and open grassy areas.

behavior

Solitary, territorial, and predominantly nocturnal, African civets rest in dense vegetation during the day. Scent marking is common with secretions from the perineal glands, but vocal communication is limited. Longevity may reach 14 years.

feeding ecology and diet

Opportunistic omnivore. African civets consume mainly fruits, rodents, insects, reptiles, birds, amphibians, and carrion.

reproductive biology

Breeding occurs year round, gestation is 60–81 days, litter size one to four young. Young are born fully furred, and eyes open at birth or shortly thereafter. Weaning occurs at 14–16 weeks, and sexual maturity is reached after one year. Mating system is not known.

conservation status

Not threatened.

significance to humans

African civets are easily tamed, and probably are the most popular animal for the production of civet.


Common genet

Genetta genetta

subfamily

Viverrinae

taxonomy

Viverra genetta (Linnaeus, 1758), Spain.

other common names

English: European genet, small-spotted genet; French: Genette commune, genette européenne; German: Ginsterkatze; Spanish: Genetta.

physical characteristics

Body length 17–22 in (43–55 cm), tail 13–16 in (33–51 cm), weight 3–6 lb (1.5–2.5 kg). The body is elongated, with a long tail. The head is small and has a long snout and medium-sized ears. Legs are short. The fur has a light basic coloration and dark spots, and the tail is banded.

distribution

Spain, Portugal, France, Arabia, north Africa, and in sub-Saharan savannas.

habitat

Forests, savannas, shrublands.

behavior

European genets are probably the best known of all Viverridae. Nocturnal and solitary, common genets shelter in dense vegetation and hollow trees during the day. Genets are excellent climbers, and often climb down head first. Home ranges vary in size from 1–3 mi2 (2–8 km2), and overlap. Genets scent mark with the secretion from their perineal glands, and marks allow assessment of social status and individual recognition. Several different vocalizations are used.

feeding ecology and diet

Opportunistic, genets consume mainly rodents, birds, amphibians, fruits, reptiles and insects.

reproductive biology

Mating occurs from January to September, but peaks in February or March. Gestation lasts 10–11 weeks, litter size is one to four. Young are born with hair, but belly is naked. Eyes open after eight days. Sexual maturity is reached after two years, and longevity may exceed 16 years. Mating system is not known.

conservation status

Only one subspecies, G. g. isabelae, is rare and listed by the IUCN.

significance to humans

None known. Genets are easily tamed, and readily breed in captivity.


Aquatic genet

Osbornictis piscivora

subfamily

Viverrinae

taxonomy

Osbornictis piscivora J. A. Allen, 1919, Zaire.

other common names

English: Congo water civet, fishing genet; French: Genette aquatique; German: Wasser-Schleichkatze.

physical characteristics

Body length 18–20 in (44–50 cm), tail 13–16 in (34–41 cm), weight 3 lb (1.5 kg).

distribution

Democratic Republic of the Congo (Zaire).

habitat

Heavily forested areas in proximity to water.

behavior

Solitary.

feeding ecology and diet

Mainly fish, possibly amphibians and crustaceans.

reproductive biology

Nothing is known.

conservation status

Listed by the IUCN as Data Deficient.

significance to humans

Meat may be consumed by local people.


African linsang

Poiana richardsonii

subfamily

Viverrinae

taxonomy

Genetta richardsonii (Thomson, 1842), Fernando Po (Equatorial Guinea).

other common names

English: Oyan; French: Linsang africain, poiana; German: Pojana.

physical characteristics

Body length 13–15 in (33–38 cm), tail 14–15 in (36–38 cm), weight 1.4 lb (650 g). The tail is longer than the body and is covered by a thick fur; it has 12 dark rings and a dark tip. The yellow basic coloration is covered by numerous small, black spots.

distribution

Sierra Leone, Ivory Coast, Gabon, Cameroon, Congo, and Fernando Po.

habitat

Forests.

behavior

Nocturnal, rests on tree limbs or in deserted squirrel nests during the day.

feeding ecology and diet

Nuts, insects, and birds.

reproductive biology

Litter size two to three. Mating system is not known.

conservation status

Not threatened.

significance to humans

Skins used by indigenous people in some areas.


African palm civet

Nandinia binotata

subfamily

Nandiniinae

taxonomy

Viverra binotata (Gray, 1830), Africa, Ashantee (Ghana).

other common names

French: Nandinie d'Afrique; German: Pardel-Roller.

physical characteristics

Body length 18–23 in (44–58 cm), tail 19–25 in (46–62 cm), weight 4–11 lb (2–5 kg). The hair is long and very thick, varying in color from yellowish gray-brown to brown. The upper side of the body is darker. The nape of the neck has three dark longitudinal stripes, and there are numerous dark spots on the upper body side. The shoulder has a white blotch. The upper side of the tail is banded. The feet are very short and have naked soles.

distribution

Tropical Africa from Senegal and Angola to southern Sudan and eastern Africa, and south to Zimbabwe.

habitat

Tropical forests.

behavior

Arboreal and nocturnal. Density of 13 per mi2 (5 per km2). Territories marked with scent. African palm civets use their forefeet much more skillfully than other viverrids; they approach the fine movements and dexterity of raccoons (Procyonidae).

feeding ecology and diet

Mostly fruits, but also rodents, birds eggs, insects. Animal prey is recognized chiefly by its movement.

reproductive biology

Probably polygynous. Births peak in May and October, gestation is 64 days, litter size usually two, but may reach four.

conservation status

Not threatened.

significance to humans

Occasionally tamed and kept as pets.


Binturong

Arctictis binturong

subfamily

Paradoxurinae

taxonomy

Viverra binturong (Raffles, 1821), "Malacca."

other common names

English: Bear cat; French: Binturong; German: Binturong.

physical characteristics

Body length 24–38 in (61–97 cm), tail 22–35 in (56–89 cm), weight 20–30 lb (9–14 kg). The body is compact and the feet are short with naked soles. The hair is rough and loose, with a blackish color. The binturong is the only viverrid with a prehensile tail.

distribution

India, Nepal, Thailand, Malaysia, and Indochina.

habitat

Dense forests.

behavior

Nocturnal and mostly arboreal, the binturong runs like a bear when it is on the ground, putting the entire sole of the foot down. The slinking motion that is normally characteristic of viverrids is not found in the binturong.

feeding ecology and diet

Fruits, leaves, birds, carrion, and fish.

reproductive biology

Most births occur from January to March, gestation 84–99 days, litter size one to six. Both parents care for the young. Mating system is not known.

conservation status

Not threatened.

significance to humans

May be domesticated and kept as pets. Common as a zoo animal.


Otter civet

Cynogale bennettii

subfamily

Hemigalinae

taxonomy

Cynogale bennettii Gray, 1837, Sumatra.

other common names

English: Water civet; French: Civette-loutre de Sumatra; German: Mampalon; Spanish: Cibeta nutria.

physical characteristics

Body length 23–27 in (57–68 cm), tail 5–8 in (13–21 cm), weight 6.5–11 lb (3–5 kg). The body is heavy and compact. The black legs are short and have naked soles and greatly curved claws. All feet are webbed, but the hind legs have less webbing than the forefeet. The broad, flat nose is well supplied with vibrissae. The nasal openings are on top of the nose, and the ears are rounded. The hair is yellowish gray-brown with a black-brown throat and lower lip. The chin and a spot over the eyes are yellowish white.

distribution

North Vietnam, Malaysia, Sumatra, Java, and Borneo.

habitat

Streams and swampy areas.

behavior

Solitary, good climber, but not strong swimmer. Longevity may reach five years.

feeding ecology and diet

Crustacens, mollusks, fish, birds, small mammals, and fruits.

reproductive biology

Litter size is two to three. Mating system is not known.

conservation status

Listed as Endangered by the IUCN.

significance to humans

None known.


Falanouc

Eupleres goudotii

subfamily

Euplerinae

taxonomy

Eupleres goudotii Doyère, 1835, Madagascar.

other common names

English: Malagasy mongoose, slender falanouc, small-toothed mongoose; French: Euplère de Goudot; German: Kleinfalanuk; Spanish: Fanaloca, mangosta dentipequeño.

physical characteristics

Body length 18–26 in (45–65 cm), tail 9–20 in (22–50 cm), weight 4–9 lb (2–4 kg). The fore limbs are short and weak, with small paws. The hind legs are longer. The claws are very long, curved, and laterally compressed. There are no anal or perineal glands.

distribution

Madagascar.

habitat

Humid, lowland forests.

behavior

Nocturnal and terrestrial, it sleeps in burrows or crevices during the day. The people of southeastern Madagascar use falanouc tails for ornamental clothing.

feeding ecology and diet

Falanoucs dig for food and feed on insects, earthworms, and other invertebrates.

reproductive biology

Mating occurs in July or August, litter size is one or two. Mating system is not known.

conservation status

Listed as Endangered by the IUCN.

significance to humans

None known.

Common name / Scientific name / Other common namesPhysical characteristicsHabitat and behaviorDistributionDietConservation status
Owston's palm civet Chrotogale owstoni Spanish: Hemigalo de OwstonBody and base of tail have alternating and sharply contrasting dark and light transverse bands, longitudinal stripes present on neck. Underparts are pale buffy. Head and body length 20–25 in (50.8–63.5 cm), tail length 15–19 in (38.1–48.2 cm).Species is largely terrestrial, but has been seen in trees. Prefers densely vegetated habitats near water sources in both primary and secondary forests. Mating usually occurs in January and March. Each female has one to two litters, each containing one to three offspring.Southern Yunan and southwest Guangxi provinces in China; northern Vietnam; and northern Laos.Heavily consists of earth-worms; however, small vertebrates, invertebrates, and some fruit may also be included in their diet.Vulnerable
Hose's palm civet Diplogale hosei Spanish: Hemigalo de HoseColoration is dark brown or black, underparts are grayish, yellowish, white, or slightly rufescent. Buffy gray patch from eye to cheek. Tail is not banded, but dark throughout. Head and body length 26 in (66 cm), tail length 11.8 in (30 cm).Montane forest and is largely terrestrial. Partly arboreal, climbs well. Two young per litter.Borneo.Mainly orthopterans, worms, and other invertebrates.Vulnerable
Banded palm civet Hemigalus derbyanus Spanish: Hemigalo franjeadoNarrow, median dark streak on head, two broad stripes from neck to elbow, two imperfect stripes at base of tail. Coloration is whitish to orange buff, usually lighter and more buffy underneath. Head and body length 16.1–20.1 in (41–51 cm), tail length 10–15.1 in (25.5–38.3 cm).Tall forest. Terrestrial, partly arboreal, climbs well. Two young per litter.Tenasserim, Malay Peninsula, Sumatra and certain small islands to west, and Borneo.Mostly orthopterans, worms, and other invertebrates. Also eat fruit.Not threatened
Small-toothed palm civet Arctogalidia trivirgata Spanish: Civeta de los palmaresColoration of upperparts is tawny, from dusky grayish tawny to bright orange tawny. Head is dark gray and paws are brown. White stripe on muzzle, three brown or black stripes on back. Very long tail. Head and body length 17–20.9 in (43.2–53.2 cm), tail length 20.1–24 in (51–61 cm).Dense forests, sometimes coconut plantations, but avoids human settlements. Nocturnal and arboreal. Breeding continues throughout year. Two or three young per litter.Assam to Indochina and the Malay Peninsula, and on Sumatra, Bangka, Java, Borneo, and numerous small nearby islands of the East Indies.Omnivorous diet, such as squirrels, birds, frogs, insects, and fruit.Not threatened
Sulawesi palm civet Macrogalidia musschenbroekii Spanish: Civeta celebianaUpperparts are light brown chestnut to dark brown. Underparts range from fulvous to white, with a red breast. Gray patches on face (cheeks and above eyes). Brown spots and bands present on sides and lower back. Short, close fur, whorl in neck. Head and body length 39.4 in (100 cm), tail length 23.6 in (60 cm).Montane and lowland forests, and in scrubby grassland. Good climber.Sulawesi.Rodents and fruit.Vulnerable
Masked palm civet Paguma larvata Spanish: PagumaWhite and black mask covers face. General color is gray tinged with buff, orange, or yellowish red. No stripes or spots on body. Head and body length 20–30 in (50.8–76.2 cm), tail length 20–25 in (50.8–63.6 cm).Forests and brush country. Raises young in tree holes. Arboreal and nocturnal. Solitary. May be two breeding periods per year.Kashmir to Indochina and the Malay Peninsula, in much of eastern and southern China, on the Andaman Islands, Taiwan, Hainan, Sumatra, and Borneo.Small vertebrates, insects, and fruit.Not threatened
Palm civet Paradoxurus hermaphroditus Spanish: MusangColoration is gray to brown, entirely masked by black tips of guard hairs. Patter of dorsal stripes and lateral spots. Patter consists of white patches and white band across forehead. Backward direction of hairs on neck. Head and body length 17–28 in (43.2–71 cm), tail length 16–26 in (40.6–66 cm).Tropical forests. Reproduction occurs throughout the year. Litter size is two to four young. Nocturnal and arboreal.Kashmir in the west to the Philippines in the east; from southern China and the Himalayas in the north to the Greater Sundas and many lesser Sunda Islands in the south.Primarily frugivorous, feeding on berries and pulpy fruits. Also eats reptiles, eggs, and insects.Not threatened
Blotched genet Genetta tigrina Spanish: Jineta de motas grandesColoration is white, gray, or buff. Spots and dark markings cover pelage from shoulders to base of tail. Spots vary from black to rust. Short legs, long body with white-ringed, black-tipped tail. Head and body length 19.3–23.6 in (49–60 cm), tail length 16.5–21.3 in (42–54 cm).Tropical rainforest, tropical deciduous forest, tropical scrub forest, tropical savanna, and grasslands. Very little known about reproductive patterns. Litter size ranges from one to five offspring. Nocturnal and solitary. Arboreal and nimble. Vocalization important.South Africa and Lesotho.Small rodents, birds, reptiles, fruit, and invertebrates.Not threatened
Common name / Scientific name / Other common namesPhysical characteristicsHabitat and behaviorDistributionDietConservation status
Banded linsang Prionodon linsang Spanish: Linsang rayadoColoration varies from whitish gray to brownish gray and becomes creamy on underside. Dark patter of four or five broad, transverse black or dark brown bands across back. One large stripe on each side of neck. Dark spots on sides of body and legs. Head and body length (35–45 cm), tail length (30.4–42 cm).Forests. Nocturnal and arboreal. Nests made of sticks and leaves. No clear breeding season. Two young per litter.Western and southern Thailand, Malay Peninsula, Sumatra, Bangka, Java, and Borneo.Small mammals, birds, eggs, and insects.Not threatened
Indian civet Viverra zibetha English: Large Indian civet; Spanish: Civeta hindú grandeColoration is gray or brown. Black spots cover body, white stripes on neck. Usually two white stripes and three black stripes. Tail has black rings. Limbs are black. Average body weight 11–24.3 lb (5–11 kg), tail length 13 in (33 cm).Scrub and densely forested areas. Females are polyestrous, breeding throughout the year. Two litters per year, each producing as many as four offspring. Females raise young alone. Solitary, nocturnal animals. Terrestrial, communicate through glandular secretions.Nepal and eastern India to southeastern China and Malay Peninsula.Carnivorous, feeding on birds, frogs, snakes, small mammals, and chickens. They also eat fruit, roots, eggs, and have been recorded eating fish and crabs.Not threatened
Oriental civet Viverra megaspila English: Large-spotted civet; Spanish: Civeta de motas grandesLong, loose fur elongated in median line of body, forming low crest or mane. Black spots on grayish or tawny ground color. Black and white stripes on sides of neck and throat. Head and body length 23–37.4 in (58.5–95 cm), tail length 11.8–19 in (30–8.2 cm).Wide variety of habitats in forest, brush, and grassland. Stay in dense cover during day. Nocturnal, mainly terrestrial, good climbers. Generally solitary, one to four offspring per litter.Peninsular India, Myanmar to Indochina, and Malay Peninsula.Small mammals, birds, snakes, frogs, insects, eggs, fruit, and some roots.Not threatened

Resources

Books

Ewer, R. F. The Carnivores. Ithaca, NY: Comstock Publishing, 1998.

Gittleman, J. L. Carnivore Behavior, Ecology, and Evolution. Vol. 2. Ithaca, NY: Cornell University Press, 1996.

Macdonald, D. W. The New Encyclopedia of Mammals. Oxford: Oxford University Press, 2001.

Nowak, R. M. Walker's Mammals of the World. 6th ed. Baltimore: John Hopkins University Press, 1999.

Wilson, D. E., and D. M. Reeder. Mammal Species of the World: A Taxonomic and Geographic Reference. 2nd ed. Washington DC: Smithsonian Institute Press, 1993.

Periodicals

Larivière, S., and J. Calzada. "Genetta genetta." Mammalian species 680 (2001): 1–6.

Ray, J. C. "Civettictis civetta." Mammalian Species 488 (1995): 1–7.

van Rompaey, H. "Osbornictis piscivora." Mammalian Species 309 (1998): 1–4.

Veron, G. "Pads Morphology in the Viverridae (Carnivora)." Acta Theriologica 44 (1999): 363–376.

Veron, G., and S. Heard. "Molecular Systematics of the Asiatic Viverridae (Carnivora) Inferred from Mitochondrial Cytochrome b Sequence Analysis." Journal of Zoology and Systematics Evolution Research 38 (2000): 209–217.

Serge Larivière, PhD