Mustelidaebadgers, otters, weasels, and relatives

Di­ver­sity

Mustel­idae is the largest fam­ily within Car­nivora and is com­prised of 56 species in 22 gen­era. Mem­bers of this fam­ily in­clude weasels, stoats, pole­cats, mink, marten, fish­ers, wolver­ines, ot­ters, bad­gers and oth­ers. While many au­thors have tra­di­tion­ally con­sid­ered skunks a sub­fam­ily within Mustel­idae, re­cent mol­e­c­u­lar ev­i­dence in­di­cates that skunks do not lie within the mustelid group and in­stead are rec­og­nized as a sin­gle fam­ily, Mephi­ti­dae, a sys­tem­atic un­der­stand­ing which is ac­cepted here (Dra­goo and Hon­ey­cutt, 1997; Flynn et al., 2005; Marmi et. al., 2004; Sato et. al., 2003; Sato et. al., 2004). (Dra­goo and Hon­ey­cutt, 1997; Flynn, et al., 2005; Marmi, et al., 2004; Nowak, 1991; Sato, et al., 2003; Sato, et al., 2004; Vaughan, et al., 2000; Whitaker and Hamil­ton, 1998)

Mustelids in­habit all con­ti­nents ex­cept Aus­tralia and Antarc­tica, and do not occur on Mada­gas­car or oceanic is­lands. Mem­bers of this group can be found in di­verse habi­tats, which in­clude both ter­res­trial, aquatic and ma­rine en­vi­ron­ments. Mustelids are mainly car­niv­o­rous, with var­i­ous mem­bers of the fam­ily ex­ploit­ing a great di­ver­sity of both ver­te­brate and in­ver­te­brate prey. Mustelids are gen­er­ally pro­fi­cient hunters; some weasels can take prey larger than them­selves. Mem­bers of this fam­ily often hunt in bur­rows and crevices, and some species have evolved to be­come adept at climb­ing trees (e.g., marten) or swim­ming (e.g., sea ot­ters, mink) in search of prey. (Nowak, 1991; Sato, et al., 2003; Vaughan, et al., 2000; Whitaker and Hamil­ton, 1998)

Gen­er­ally, mustelids have elon­gate bod­ies with short legs and a short ros­trum, as typ­i­fied by weasels, fer­rets, mink, and ot­ters. Wolver­ines and bad­gers have broader bod­ies. An order of mag­ni­tude dif­fer­ence in size ex­ists be­tween the small­est and largest mustelid species. The small­est species is the least weasel (Mustela ni­valis), weigh­ing be­tween 35 and 250 grams. Wolver­ines (Gulo gulo) and sea ot­ters (En­hy­dra lutris) reach 32 kg and 45 kg, re­spec­tively. All mustelids have well de­vel­oped anal scent glands, which serve var­i­ous func­tions, in­clud­ing ter­ri­to­r­ial mark­ing and de­fense. (Hutch­ings and White, 2000; Nowak, 1991; Vaughan, et al., 2000)

Ge­o­graphic Range

Mem­bers of the fam­ily Mustel­idae in­habit all con­ti­nents ex­cept Antarc­tica and Aus­tralia. They do not occur on Mada­gas­car or oceanic is­lands, but have been in­tro­duced to New Zealand. (Dra­goo and Hon­ey­cutt, 1997; Sato, et al., 2003; Vaughan, et al., 2000)

Habi­tat

Mustel­idae are dis­trib­uted from the arc­tic to the trop­ics and oc­cupy nearly all ter­res­trial habi­tats. Sev­eral species are semi- or nearly fully aquatic and in­habit fresh­wa­ter rivers and streams, as well as coastal ma­rine wa­ters. (Nowak, 1991; Vaughan, et al., 2000)

Phys­i­cal De­scrip­tion

Adult mustelids range in size from 114 mm and 25 g (least weasel) to over 1 m and 45 kg (sea ot­ters). These an­i­mals are gen­er­ally long-bod­ied with short legs. Most species have slen­der bod­ies, but some, like bad­gers (Mustelinae, Mustelinae) and wolver­ines have much broader bod­ies. The skull is elon­gate with a rel­a­tively short ros­trum. Adult males are gen­er­ally about 25 per­cent larger than fe­males of the same species. The ears are short, as are the legs, each of which bears five dig­its. The claws do not re­tract and, in dig­ging species, are es­pe­cially ro­bust. Mustelids are dig­it­i­grade or planti­grade. The den­tal for­mula varies among species: 3/3, 1/1, 2-4/2-4, 1/1-2 = 28-38. The ca­nines are long, and the car­nas­sials are well-de­vel­oped. The upper mo­lars are often nar­row in the mid­dle, giv­ing them an hour­glass shape. Mustelids have a pow­er­ful bite; in many species, the large post­g­le­noid process locks the lower jaw into the upper, caus­ing the lower jaw to only move in the ver­ti­cal plane, with­out any ro­tary mo­tion. (Nowak, 1991; Vaughan, et al., 2000; Whitaker and Hamil­ton, 1998)

  • Sexual Dimorphism
  • male larger

Re­pro­duc­tion

Mat­ing sys­tems vary both within and among species. Many species are polyg­y­nous and/or promis­cu­ous. Some species are so­cial, while oth­ers are soli­tary. So­cial or­ga­ni­za­tion can vary within species as well. Mustelids re­quire pro­longed pe­ri­ods of cop­u­la­tion to in­duce ovu­la­tion of an un­fer­tilzed egg. As a re­sult, cop­u­la­tion may last for sev­eral hours be­fore fer­til­iza­tion can be suc­cess­ful. (Am­stislavsky and Ter­novskaya, 2000; John­son, et al., 2000; Vaughan, et al., 2000)

Most mustelids breed sea­son­ally, but the length of the re­pro­duc­tive pe­riod varies among species. Day length often dic­tates the onset of the breed­ing sea­son, which usu­ally lasts 3 to 4 months. Many mustelids un­dergo de­layed im­plan­ta­tion, with the fer­til­ized em­bryo tak­ing up to 10 months (e.g. Meles meles) to im­plant in the uterus in some species. En­vi­ron­men­tal con­di­tions such as tem­per­a­ture and day length de­ter­mine when im­plan­ta­tion oc­curs. Mustelids that live in more sea­sonal cli­mates are more likely to ex­hibit de­layed im­plan­ta­tion. Fol­low­ing im­plan­ta­tion, ges­ta­tion typ­i­cally lasts 30 to 65 days. Fe­males give birth to a sin­gle lit­ter each sea­son, the size of which varies within and among species. For ex­am­ple, sables have an av­er­age lit­ter size of 2.2, but can give birth to any­where from 1 to 7 pups. The moun­tain weasel av­er­ages 8.7 pups per lit­ter, but can have be­tween 3 and 14 young in a sin­gle bout of re­pro­duc­tion. Gen­er­ally, mustelids are al­tri­cial, being born small and blind. They reach sex­ual ma­tu­rity be­tween 8 months and two years fol­low­ing birth. (Am­stislavsky and Ter­novskaya, 2000; Nowak, 1991; Thom, et al., 2004; Vaughan, et al., 2000)

Young are gen­er­ally born in an alri­cial state, re­quir­ing ex­teni­sive care and pro­tec­tion from their mother. Young mustelids typ­i­cally are able to care for them­selves when they are about two months old. Fe­males de­fend ter­ri­to­ries in order to ac­quire enough re­sources to care for their young and most often nurse and pro­tect them in a bur­row or den. (John­son, et al., 2000; Nowak, 1991)

  • Parental Investment
  • altricial
  • precocial
  • pre-fertilization
    • provisioning
    • protecting
      • female
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female
  • pre-independence
    • provisioning
      • female
    • protecting
      • female

Lifes­pan/Longevity

Mustelids typ­i­cally live be­tween 5 and 20 years in the wild. (Nowak, 1991)

Be­hav­ior

Mem­bers of the fam­ily Mustel­idae are ei­ther di­ur­nal or noc­tur­nal. Many of the long, nar­row-bod­ied species are quick and agile, and move in a bound­ing, scam­per­ing fash­ion. The broader-bod­ied forms have a more lum­ber­ing gait. Some species are adept climbers, while oth­ers are ex­cel­lent swim­mers. Many species spend a great deal of time on the ground, search­ing for food in crevices, bur­rows, or under cover. Many species shel­ter in bur­rows. (Nowak, 1991; Vaughan, et al., 2000)

So­cial be­hav­ior varies both within and among species, and may vary in re­la­tion to local en­vi­ron­men­tal con­di­tions such as food avail­abil­ity. For ex­am­ple, Eu­ro­pean bad­gers are known to form groups with sev­eral males and fe­males that are all re­pro­duc­tively ac­tive within the group. Yet in other parts of their range, Eu­ro­pean bad­gers may live soli­tar­ily or in pairs.​Many species are ter­ri­to­r­ial for at least part of the year, with in­di­vid­u­als com­pet­ing over hunt­ing areas or ac­cess to mates (e.g., Mustela er­minea). (John­son, et al., 2000)

Com­mu­ni­ca­tion and Per­cep­tion

Vi­sion and hear­ing are im­por­tant in Mustel­idae, but ol­fac­tion is par­tic­u­larly well de­vel­oped. In ad­di­tion to using scent cues to find food, scent-mark­ing is the main form of com­mu­ni­ca­tion in this fam­ily. Se­cre­tions from well-de­vel­oped scent glands func­tion in ter­ri­to­r­ial in­ter­ac­tions, in­di­cate re­pro­duc­tive state, and are used in other so­cial con­texts. The de­gree and func­tion of scent mark­ing varies among species, and ac­cord­ing to so­cial and en­vi­ron­men­tal con­di­tions within species. (Hutch­ings and White, 2000; Nowak, 1991; Vaughan, et al., 2000)

Food Habits

Mustelids are pri­mar­ily car­niv­o­rous, but some species may at times eat plant ma­te­r­ial. A wide range of an­i­mal taxa are preyed upon by var­i­ous mem­bers of this fam­ily; many mustelids are op­por­tunis­tic feed­ers rather than spe­cial­ists. How­ever, many mustelids are es­pe­cially adept at cap­tur­ing small, mam­malian prey. Weasels, for ex­am­ple, are ca­pa­ble of chas­ing and cap­tur­ing ro­dents in their bur­rows. Ot­ters are well-adapted to chas­ing and cap­tur­ing aquatic prey, in­clud­ing fish, crus­taceans, and other aquatic in­ver­te­brates. Mustelids hunt in a va­ri­ety of ter­res­trial, aquatic, and ar­bo­real habi­tats. Some species reg­u­larly prey on an­i­mals larger than them­selves. Some species have been known to store food (e.g., Mustela, Gulo). (Nowak, 1991; Vaughan, et al., 2000)

Pre­da­tion

Mustelids are gen­er­ally small car­ni­vores, and are there­fore sub­ject to pre­da­tion by larger car­ni­vores such as canids with which they co-oc­cur. They may also fall prey to large snakes (Ser­pentes), rap­tors (Fal­coni­formes), and owls (St­rigi­formes). Some mustelids se­crete nox­ious chem­i­cals to dis­cour­age preda­tors. In some of these species, apose­matic color pat­terns can help ward off preda­tors. (Bright, 2000; Nowak, 1991)

Ecosys­tem Roles

Mustelids mainly im­pact their com­mu­ni­ties through their ef­fects on prey pop­u­la­tions. Many species limit ro­dent and bird pop­u­la­tions. In some cases, mustelids limit ro­dents that are con­sid­ered pests, in other cases, mustelids threaten rare bird species. Some species, such as sea ot­ters (En­hy­dra lutra) are key­stone preda­tors, en­hanc­ing the di­ver­sity of their com­mu­nity by keep­ing highly com­pet­i­tive prey species in check. Honey bad­gers (or ra­tels, Mel­livora capen­sis) have de­vel­oped com­men­sal re­la­tion­ships with both hu­mans and honey guides (In­di­ca­tor in­di­ca­tor), using both to aid in the lo­ca­tion of bee colonies. (Bright, 2000; Mac­don­ald and King, 2000; Nowak, 1991)

Com­men­sal/Par­a­sitic Species

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Many mustelids help con­trol ro­dent pop­u­la­tions that are con­sid­ered to be pests. In ad­di­tion, many are hunted and/or raised for their pelts, which are often con­sid­ered highly valu­able (e.g., the pelts of mink and sable). Some species have been do­mes­ti­cated and are traded as pets (e.g., fer­rets). (Bright, 2000; Nowak, 1991)

  • Positive Impacts
  • pet trade
  • body parts are source of valuable material
  • controls pest population

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Some mustelids are con­sid­ered pests, ei­ther for harm­ing poul­try live­stock, for threat­en­ing other species in the wild, or for trans­mit­ting dis­eases. Eu­ro­pean bad­gers have been im­pli­cated in the trans­mis­sion of bovine tu­ber­cu­lo­sis. Cat­tle may be­come in­fected from graz­ing on land where bad­gers have defe­cated. Up to 20% of bad­gers carry the dis­ease in areas where bovine tu­ber­cu­lo­sis is a prob­lem. Since 1975, bad­gers have been culled in the United King­dom, but there is no conl­cu­sive ev­i­dence that it has helped con­trol bovine TB. As mam­malian car­ni­vores, mustelids can also be in­fected by, and trans­mit, ra­bies. (Bright, 2000; Gough and Rush­ton, 2000; Nowak, 1991)

Con­ser­va­tion Sta­tus

Some mustelid species are con­sid­ered highly threat­ened by the IUCN, while other species are so abun­dant that they are con­sid­ered pests. Ap­prox­i­mately 38% of all species of Mustel­idae are con­sid­ered threat­ened,which is a much higher pro­por­tion than mam­mals in gen­eral (15%). Habi­tat de­struc­tion is a se­ri­ous risk to species with re­stricted habi­tat re­quire­ments such as ot­ters and martens. Smaller car­ni­vores that are re­stricted to small habi­tat frag­ments may also be at risk to pre­da­tion by larger car­ni­vores that can more eas­ily move among frag­ments. Hunt­ing has been a prob­lem for some species, while oth­ers, par­tic­u­larly trop­i­cal mustelids, do not seem to be de­clin­ing as a re­sult. En­dan­gered mustelids in­clude: Colom­bian weasels (Mustela fe­lipei), Eu­ro­pean mink (Mustela lutre­ola), In­done­sian moun­tain weasels (Mustela lutre­olina), ma­rine ot­ters (Lon­tra fe­lina), south­ern river ot­ters (Lon­tra provo­cax), sea ot­ters (En­hy­dra lutris), and giant Brazil­ian ot­ters (Pteronura brasilien­sis). Sea mink (Neo­vi­son macrodon) be­came ex­tinct in re­cent times. (Bright, 2000; Gough and Rush­ton, 2000)

Nu­mer­ous re-in­tro­duc­tion pro­grams for var­i­ous mustelid species have met with mixed suc­cess. Gen­er­ally, "soft" re-in­tro­duc­tions, those that allow the an­i­mals to ac­cli­mate to their new sur­round­ings while in a tem­po­rary en­clo­sure, are more suc­cess­ful than "hard" re-in­tro­duc­tions, in which cap­tive-bred an­i­mals are re­leased di­rectly into the wild. Black-footed fer­rets (Mustela ni­gripes) are con­sid­ered ex­tinct in the wild, al­though sev­eral re-in­tro­duc­tion pro­grams are un­der­way. (Bright, 2000)

  • IUCN Red List [Link]
    Not Evaluated

Other Com­ments

The ear­li­est mustelid fos­sils are from the Old World and have been dated to the Oligocene (33.5 – 23.8 mya) or mid-Miocene (23.8 – 5.3 mya). There is de­bate re­gard­ing which fos­sils from these epochs rep­re­sent pos­si­ble an­ces­tral forms that led to Mustel­idae and which fos­sils rep­re­sent the first mod­ern mustelids.

Con­trib­u­tors

Matt Wund (au­thor), Tanya Dewey (ed­i­tor), An­i­mal Di­ver­sity Web.

Glossary

Australian

Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.

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Ethiopian

living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.

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Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

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Neotropical

living in the southern part of the New World. In other words, Central and South America.

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Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

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acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

aposematic

having coloration that serves a protective function for the animal, usually used to refer to animals with colors that warn predators of their toxicity. For example: animals with bright red or yellow coloration are often toxic or distasteful.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

bog

a wetland area rich in accumulated plant material and with acidic soils surrounding a body of open water. Bogs have a flora dominated by sedges, heaths, and sphagnum.

brackish water

areas with salty water, usually in coastal marshes and estuaries.

carnivore

an animal that mainly eats meat

causes or carries domestic animal disease

either directly causes, or indirectly transmits, a disease to a domestic animal

chaparral

Found in coastal areas between 30 and 40 degrees latitude, in areas with a Mediterranean climate. Vegetation is dominated by stands of dense, spiny shrubs with tough (hard or waxy) evergreen leaves. May be maintained by periodic fire. In South America it includes the scrub ecotone between forest and paramo.

chemical

uses smells or other chemicals to communicate

coastal

the nearshore aquatic habitats near a coast, or shoreline.

cosmopolitan

having a worldwide distribution. Found on all continents (except maybe Antarctica) and in all biogeographic provinces; or in all the major oceans (Atlantic, Indian, and Pacific.

delayed implantation

in mammals, a condition in which a fertilized egg reaches the uterus but delays its implantation in the uterine lining, sometimes for several months.

desert or dunes

in deserts low (less than 30 cm per year) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity. Vegetation is typically sparse, though spectacular blooms may occur following rain. Deserts can be cold or warm and daily temperates typically fluctuate. In dune areas vegetation is also sparse and conditions are dry. This is because sand does not hold water well so little is available to plants. In dunes near seas and oceans this is compounded by the influence of salt in the air and soil. Salt limits the ability of plants to take up water through their roots.

diurnal
  1. active during the day, 2. lasting for one day.
endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

estuarine

an area where a freshwater river meets the ocean and tidal influences result in fluctuations in salinity.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

fossorial

Referring to a burrowing life-style or behavior, specialized for digging or burrowing.

freshwater

mainly lives in water that is not salty.

insectivore

An animal that eats mainly insects or spiders.

intertidal or littoral

the area of shoreline influenced mainly by the tides, between the highest and lowest reaches of the tide. An aquatic habitat.

introduced

referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

keystone species

a species whose presence or absence strongly affects populations of other species in that area such that the extirpation of the keystone species in an area will result in the ultimate extirpation of many more species in that area (Example: sea otter).

marsh

marshes are wetland areas often dominated by grasses and reeds.

molluscivore

eats mollusks, members of Phylum Mollusca

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

natatorial

specialized for swimming

nocturnal

active during the night

oriental

found in the oriental region of the world. In other words, India and southeast Asia.

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pet trade

the business of buying and selling animals for people to keep in their homes as pets.

piscivore

an animal that mainly eats fish

polar

the regions of the earth that surround the north and south poles, from the north pole to 60 degrees north and from the south pole to 60 degrees south.

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

polygynous

having more than one female as a mate at one time

rainforest

rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.

riparian

Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).

saltwater or marine

mainly lives in oceans, seas, or other bodies of salt water.

scent marks

communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them

scrub forest

scrub forests develop in areas that experience dry seasons.

seasonal breeding

breeding is confined to a particular season

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

stores or caches food

places a food item in a special place to be eaten later. Also called "hoarding"

suburban

living in residential areas on the outskirts of large cities or towns.

swamp

a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.

tactile

uses touch to communicate

taiga

Coniferous or boreal forest, located in a band across northern North America, Europe, and Asia. This terrestrial biome also occurs at high elevations. Long, cold winters and short, wet summers. Few species of trees are present; these are primarily conifers that grow in dense stands with little undergrowth. Some deciduous trees also may be present.

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

terrestrial

Living on the ground.

territorial

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

tundra

A terrestrial biome with low, shrubby or mat-like vegetation found at extremely high latitudes or elevations, near the limit of plant growth. Soils usually subject to permafrost. Plant diversity is typically low and the growing season is short.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

young precocial

young are relatively well-developed when born

Ref­er­ences

Am­stislavsky, S., Y. Ter­novskaya. 2000. Re­pro­duc­tion in Mustelids. An­i­mal Re­pro­duc­tion Sci­ence, 60-61: 571-581.

Bright, P. 2000. Lessons from lean beasts: con­ser­va­tion bi­ol­ogy of the mustelids. Mam­mal Re­view, 30: 217-226.

Dra­goo, J., R. Hon­ey­cutt. 1997. Sys­tem­at­ics of mustelid-like car­ni­vores. Jour­nal of Mam­mal­ogy, 78: 426-443.

Flynn, J., J. Finarelli, S. Zehr, J. Hsu, M. Ned­bal. 2005. Mol­e­c­u­lar phy­logeny of the Car­nivora (Mam­malia): As­sess­ing the im­pact of in­creased sam­pling on re­solv­ing enig­matic re­la­tion­ships. Sys­tem­atic Bi­ol­ogy, 54: 317-337.

Gough, M., S. Rush­ton. 2000. The ap­pli­ca­tion of GIS-mod­el­ling to mustelid land­scape ecol­ogy. Mam­mal Re­view, 30: 197-216.

Hutch­ings, M., P. White. 2000. Mustelid scent-mark­ing in man­aged ecosys­tems: im­pli­ca­tions for pop­u­la­tion man­age­ment. Mam­mal Re­view, 30: 157-169.

John­son, D., D. Mac­Don­ald, A. Dick­man. 2000. An analy­sis and re­view of mod­els of the so­cio­bi­ol­ogy of the Mustel­idae. Mam­mal Re­view, 30: 171-196.

Koepfli, K., R. Wayne. 2003. Type I Sts mark­ers are more in­for­ma­tive than cy­tochrome b in phy­lo­ge­netic re­con­struc­tion of the Mustel­idae (Mam­malia: Car­nivora). Sys­tem­atic Bi­ol­ogy, 52: 571-593.

Mac­don­ald, R., C. King. 2000. Bi­ol­ogy of mustelids: re­views and fu­ture di­rec­tions. Mam­mal Re­view, 30/3-4: 145.

Marmi, J., J. Lopez-Gi­raldez, X. Domingo-Roura. 2004. Phy­logeny, evo­lu­tion­ary his­tory and tax­on­omy of the Mustel­idae based on se­quences of the cy­tochrome b gene and a com­plex repet­i­tive flank­ing re­gion. Zo­o­log­ica Scripta, 33: 481-499.

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