A Review of the Literature on the Worldwide Distribution, Spread of, and Efforts to Eradicate the Coypu (Myocastor coypus) Author(s): Jacoby Carter and Billy P. Leonard Source: Wildlife Society Bulletin, Vol. 30, No. 1 (Spring, 2002), pp. 162-175 Published by: Wiley on behalf of the Wildlife Society Stable URL: http://www.jstor.org/stable/3784650 . Accessed: 01/08/2014 14:48

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp

. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected].

.

Wiley and Wildlife Society are collaborating with JSTOR to digitize, preserve and extend access to Wildlife Society Bulletin.

http://www.jstor.org

This content downloaded from 158.135.136.72 on Fri, 1 Aug 2014 14:48:22 PM All use subject to JSTOR Terms and Conditions 162 LITERATURE REVIEW OF COYPUS WORLDWIDE A review of the literature on the worldwide distribution,spread of, and effortsto eradicate the coypu (Myocastorcoyp us)

JacobyCarter and BillyP. Leonard AbstractWe conducteda literature review of coypu (Myocastor coypus) introduction and eradi- cationefforts worldwide. The coypu (also called nutria) has been introduced from its ori- ginsin SouthAmerica to everycontinent except Australia and Antarctica. While per- ceivedin some regions as a valuableresource, inmost regions the are considered a pestspecies. Coypus have caused damage to water control structures, crops, and marsh systemsand are considered a disease host. Eradicationefforts have met with varying degreesof success. For those efforts tobe successful,the coypu populations must be - latedto prevent reintroduction, harsh winters are necessary to reducetheir populations, andintensive trapping efforts must be sustaineduntil the last coypu has been removed.

Keywords coypu, eradication, exotic species, forbearer, herbivory, Myocastor coypus, nonindige- nousspecies, nutria, trapping,

A tahnkaform poem transliteratedfrom Japanese: The last review of coypus worldwide was by "Shirasagimo Sugamomo Koi mo kechirashite, Aliev (1967). Since then coypus have been intro- Sasagase gawa wo Nutoriayuku." duced into severalother countries and theirstatus has changed. Our purpose was to review the his- -Ito Shinsuke,Okayama, Japan tory of their introductionoutside their native Asahi Shimbun,14 April1996 range,and effortsto controlor eradicatethem in The Englishtranslation: regions where they have become feral. There is documentationfor the introductionof coypus into "Pushingaway white herons,ducks, and carp, every continent and major temperate region the nutriagoes his way in the RiverSasagase." except Australia,Antarctica, and New Zealand (A. The coypu (Myocastor coypus), also called Buchanan,TasmanianHerbarium, Australia, personal nutria,is an aquatic rodentnative to SouthAmerica communication;T Robinson,Ministry of Research, south of 230 latitude,including Argentina, Bolivia, Science, and Technology,New Zealand, personal southern Brazil, Chile, Paraguay,and Uruguay communication). (Ehrlich1967, Banfield 1974, Kinler et al. 1987,Kin- The firstattempt at coypu farmingoccurred in ler 1992a). Coypus have long been valued as France in the early 1880s, but the firstextensive furbearers,which has led to theirintroduction into establishmentof coypu farmsoccurred in South manycountries outside of theirnative range. Americain the 1920s (Evans 1970). The success of

Addressfor Jacoby Carter: United States Geological Survey, National Wetlands Research Center, 700 CajundomeBlvd., Lafayette, LA 70506, USA;e-mail: jacoby-carter~usgs.gov. Address for Billy P. Leonard:Department of Biology,University of , Lafayette,LA 70504, USA;present address: United States Forest Service, Rocky Mountain Research Station, 333 BroadwaySE, Suite1 15, Albuquerque, NM 87102,USA.

WildlifeSociety Bulletin 2002, 30(1):162-175 Peeredited

This content downloaded from 158.135.136.72 on Fri, 1 Aug 2014 14:48:22 PM All use subject to JSTOR Terms and Conditions Literaturereview of coypus worldwide * Carterand Leonard 163 these operations led to Table1. Coypuintroduction and statusby country in the eastern hemisphere. expansion of coypu farm operations in and Region-Country Date ofintroduction Method of introduction Current status NorthAmerica (Evans 1970). Africa These farms became the Kenya 1950 escape,released extinct source for wild populations Zimbabwe,Zambia, and Botswana before1958 unknown neverestablished of coypus aroundthe world. EastAsia Coypus from these farms China 1960s escape,released present often escaped, or in many Japan 1910 escape present cases theywere deliberately SouthKorea unknowndate unknown present releasedinto the wild to pro- Thailand 1993 escape neverestablished vide a or to CentralAsia and theMiddle East remove aquatic vegetation Armenia 1940 released present1967 (Dozier 1952, Evans 1970, Azerbaijan 1930 to 1932 released present1967 Kinler1992b, Bounds 2000). Georgia 1930 to 1932 released present1967 In many systems coypus Israel 1948-1966 escape,released present1967 have become a nuisance Russia 1926 released present Tajikistan 1949 released present1967 species because their feed- Turkey before1984 released present1967 activities marsh ing destroy Turkmenistan 1930 and 1932 released present1967 vegetation, their burrows Europe undermine water control Austria 1935 escape present structures,and theyfeed on Belgium 1930s escape present agricultural crops (Lin- Bulgaria unknown unknown present scombe et al. 1981, Grace TheCzech Republicand Slovakia 1992, Linscombe and Kinler before1 950s escape present 1997). Coypus also have Denmark 1930s and40s escape extinct England 1929 escape eradicated1989 been associated with para- Finland before1967 escape 1990s extinct sites that affecthumans and France 1882 escape present livestock(Moutou 1997) and Germany 1926 escape,released 1 930s present can adverselyaffect wildlife Greece before1948 escape present (Gebhardt 1996). However, Hungary unknown escape present in manycountries the coypu Ireland before1967 unknown extinct is considered a welcome Italy 1928 escapeby 1960 present resourceas a furbearer(Aliev Netherlands 1930 escapeby 1940 present 1967). Theirclassification as Norway before1946 unknown extinct a pest oftendepends on the Poland unknown escape 1948 present price of theirpelts (Lowery Romania unknown unknown present 1974). High furprices and Spain before1967 escape extinct Sweden before1 967 unknown extinct subsequent trappingreduce Switzerland after1967 unknown present the population, mitigating TheFormer Federal Republic of Yugoslavia some of the environmental before1967 escape present damage that coypus can cause, and bringmoney into the local economy(Kinler et systemswhere they are considered a nuisance al. 1987). Low furprices usuallyresult in a reduced species. fur harvest,which allows coypu populations to exceed the carrying capacity of their habitat (Gosling and Baker 1987, Kinleret al. 1987, Carter Methods et al. 1999). We reviewedpublished literature on BetweenMarch and August1998 we searchedfor coypus and queried expertsto summarizecoypus' articleson Myocastorcoypus using electronic data- worldwidedistribution and expansionpatterns. We bases, includingCambridge Scientific Abstracts, The also documented effortsto eliminatethem from Web of Science,Wildlife Worldwide, and Wilsondisk

This content downloaded from 158.135.136.72 on Fri, 1 Aug 2014 14:48:22 PM All use subject to JSTOR Terms and Conditions 164 WildlifeSociety Bulletin 2002, 30(1):162-175

CD-ROM. We also performedInternet searches on Results the WorldwideWeb using the Metacrawlersearch engine. Search termsincluded "Myocastot;""coy- The distributionof coypus outside theirhome pus,""coypu," and "nutria." After compiling a bibliog- range in South America resulted mainly from raphyof articles located through the database search attemptsto breed themfor their fur (Harris 1956, and fromprevious research, we acquiredadditional Aliev 1967, Kinler 1992a). In favorablehabitats informationusing reference tracing, focusing on arti- coypus oftenescaped and became feral. In some cles that discussedMyocastor coypus distribution, countriesthey have been released deliberatelyand introduction,control, or eradication. Additional managedin a semicaptivestate (i.e., not caged but informationrequests were posted on severalelec- placed in areas where the habitatis managedinten- tronicbulletin boards. Severalexperts (L. Foote Uni- sivelyfor their benefit) (Aliev 1967). In othercoun- versityof Alberta,Canada; M. Gosling,Ministry of tries,escaped coypushave not survivedin the wild. Agriculture,Fisheries and Food,England; G. Piero, l'Is- In general these are places with harsh winters. titutoNazionale per la Fauna Selvatica,Italy) were Coypus are generallyconsidered vulnerable to cold contacteddirectly. These expertsalso providedboth weather(Ehrlich 1967, Gosling et al. 1983,Kinler et informationand additionalbibliographic resources. al. 1987, Reggianiet al. 1995). However,they also All of the articles were reviewed for date of exhibita behavioralflexibility that allows themto coypu introduction,distribution, method of control surviveand move into areas thatwere previously or eradication,economic cost or benefit,and dam- consideredtoo cold forthem to exist in the wild age. We summarizedthe informationon a map (Ehrlich 1962, 1967; Doncaster and Micol 1990). (Figure1) to assistvisualization. Some countries,after assessing the economic and

Escape or reliea Extinct Rangeexpansion Unknown Neveestablied Countryoforigin - Erdirdlote Year Indicates date of knownIntroduction. Figure1. A color-codedmap of coypu distributions around the world by Carey Hamburg and Jacoby Carter.

This content downloaded from 158.135.136.72 on Fri, 1 Aug 2014 14:48:22 PM All use subject to JSTOR Terms and Conditions Literaturereview of coypus worldwide * Carterand Leonard 165 environmentalburden that uncontrolled wild coy- Hemisphere.It is implied,but is by no means clear, pus have placed on theirsociety, have labeled the thatthe map representedwild populations.For pur- animala pest species and attemptedto eradicateit poses of this studywe assumed thatit did, unless (Gosling and Baker 1989). In otherinstances coy- evidence to the contrarywas presented. Because pus are not considered enough of a problem to amount of informationon coypu introductionsin warrantan eradicationcampaign, or eradicationis NorthAmerica is moreextensive than for elsewhere viewed as an unattainablegoal (Wilner 1982). In in the world,this review is divided into 2 parts, fact,during the 1950s at leastone UnitedStates Fish the EasternHemisphere and NorthAmerica. Table 1 andWildlife Service biologist (Dozier 1952) regard- summarizesinformation by countryon when and ed coypus as less of a nuisanceand moreof an eco- how coypus were released and theircurrent status nomic benefit than native (Ondatra for the Eastern Hemisphere;Table 2 summarizes zibethicus). North American information.What follows are Primarymethods of eradicationfall into 3 cate- regional and national summariesof coypu intro- gories:trapping, poisoning, and shooting(Gosling duction,distribution, control, and eradication. 1981, Gosling et al. 1988, Stevens 1992, Moutou 1997). These methods are often most effective Eastern hemisphere when used in combinationwith baitingand artifi- Africa cial islandsor rafts(Wilner 1982, Bakerand Clarke We did not finda comprehensivereview of coy- 1988, LeBlanc 1994). Aliev's (1967) review had a pus in Africa.They have been reportedin Kenya breedingdistribution map of coypus in the Eastern and in an area of south-centralAfrica where the countries of Botswana, Zambia, and Zimbabwe meet (Table 1). Kinleret al. (1987) placed themin Africa,and Woods et al. (1992) assertedthat they were presentin EastAfrica. Kenya. Coypuswere importedto the Kinangop Plateaufor fur farming in 1950 (Harperet al. 1990). By 1965 some individualshad escaped and arrived at Lake Naivasha,and by 1970 a large population was established (Harper et al. 1990). Pythons (Python rebae) were introduced for biological control,but they were ineffective(Harper et al. 1990). However,for reasons unknown,the coypu populationwas decliningthere in 1980 and none have been seen since 1984 (Harperet al. 1990). Zimbabwe, Zambia, and Botswana. Aliev (1967) reportedcoypus in Zimbabweand possibly Botswanaand Zambia,although the date of intro- duction is unknown.According to J.duToit (Mam- mal ResearchInstitute, University of Pretoria,South Africa,personal communication),coypus are not feralanywhere in southernAfrica, and he suggested thatthis was due to the highpredator density. Eastia We foundno comprehensivereviews of coypus in EastAsia. Most of the informationwas scattered in a varietyof sources coveringspecific countries. Coypus are known to have been introducedinto China,Japan, Korea,Thailand, and Taiwan (Table 1). Most of the informationis anecdotal. China. Coypus were introducedinto China in the early1960s forfur farming (W. S. Xie, Institute A coypuin a Louisianamarsh. Photo by Thomas G. Hargis. of Zoology,Chinese Academy of Science, China,

This content downloaded from 158.135.136.72 on Fri, 1 Aug 2014 14:48:22 PM All use subject to JSTOR Terms and Conditions 166 WildlifeSociety Bulletin 2002, 30(1):162-175

Table2. Coypuintroduction and statusby state and provincein the North America. China. Coypus are now consid- ered an agriculturalpest (Xie and Date of Methodof Li 1999). Region-Country introduction introduction Currentstatus Japan. Coypus were intro- UnitedStates duced into Japan in 1910 (Miura Alabama 1949 released present 1976). The Japanese military Arkansas 1940s unsuccessfulrelease encouraged breeding them on after1960 rangeexpansion present small fur farms because of the 1899 escape eradicated Colorado before1978 escape extinct high qualityof theirpelts (Miura Delaware unknown rangeexpansion present 1976). When pelt pricesdropped, Florida 1950s rangeexpansion, the coypus were eitherkilled or escape,& release present released (Miura 1976). Since Georgia before1952 release present 1963,coypus have been huntedas Idaho beforel978 escape extinct an eradication and control Illinois unknown escape extinct method (Miura 1976). Japan's before1978 escape eradicated Indiana largest concentrationsoccur in Kansas before1978 unknown extinct the Okayama prefecture(Miura Kentucky before1970 released neverestablished Louisiana 1930S released,escape present 1976). 1930s escape present South Korea. Coypus have Michigan 1930s escape neverestablished been documentedin South Korea Minnesota before1978 escape neverestablished (Han et al. 1999), although the Mississippi before1 971 rangeexpansion date of introductionand present & released present statusare unknown. Missouri before1978 unknown extinct Thailand. The governmentof Montana before1981 unknown extinct Thailandviews coypus as a prob- Nebraska before1978 escape neverestablished lem They NewMexico before1970 escape extinct (Kanwanich 1998). NorthCarolina 1941 introduced present were imported from China in Ohio 1937 escaped neverestablished 1993 for breeding,and by 1998 Oklahoma unknown unknown present were seen in thewild (Kanwanich Oregon 1941 escape present 1998). The local residentshave Texas 1941 escape,released eaten some,but no organizedcon- & rangeexpansion present trolprogram has been implement- Tennessee before1996 rangeexpansion present ed (Kanwanich 1998). The gov- 1939 unknown neverestablished ernmentdebated the prohibition Virginia before1978 rangeexpansion present 1930s escape present of importingadditional animals Canada into the country (Kanwanich BritishColumbia before1943 escape present 1998) and decided not to promote Ontario unknown unknown present coypu farmingoperations (Rid- NovaScotia before1 978 unknown neverestablished montri1998,1Tunyasiri 1998). Quebec after1927 unknown present CentralAsia and theMiddle East Mexico Coypus have been introduced NorthernMexico on TexasBorder throughoutcentral Asia and Israel before1980 rangeexpansion present (Table 1). The last reviewof coy- pus in Asia and the Middle East was by Aliev(1967). personalcommunication). Originally introduced to Armenia. Aliev(1967) reportedcoypus in Arme- northern China, they also were introduced to nia. A trial of raisingcoypus semicaptivein the southernChina (Xie and Li 1999). Laterattempts wild was begun in 1940 (Aliev 1967). Semicaptive were made to export them fromChina to other raisingof coypus,as practicedin the Caucasus and Asian countriesfor fur farming (Kanwanich 1998). in EasternEurope, consists of managingthe habitat Aftera drop in furprices, coypus were released or so that the animals do not experience the detri- escaped and now wild populationsexist in South mentaleffects of harshwinters. Ponds are drained

This content downloaded from 158.135.136.72 on Fri, 1 Aug 2014 14:48:22 PM All use subject to JSTOR Terms and Conditions Literaturereview of coypus worldwide * Carterand Leonard 167

and vegetativecover is provided. In some places most recentreview of theirdistribution in Europe coypus are even fed over winter (Ehrlich 1962, is by Mitchell-Joneset al. (1999). Aliev 1967,Suvegova et al. 1994). Coypu acclimati- Austria. Coypus have been bred in Austria(Lau- zation was a success on the upper sectionsof irri- rie 1946). The captureof free-livingcoypus began gationcanals near springs(Aliev 1967). in 1935 (Aliev 1967). Wild populations are still Azerbaijan. Vinogradovand Argiropulo(1941) present(Mitchell-Jones et al. 1999). reportedcoypus acclimatizedin Azerbaijan. Ani- Belgium. Coypus have been bred in captivity mals were released from 1930 to 1932 on reser- since the 1930s and are now feral(Laurie 1946, voirs (Aliev 1967). Breeders raised a pedigreed Aliev 1967, Litjens 1980). The population was groupcalled Azerbaijanwhite coypus (Aliev 1967). healthyenough to have dispersersmove intoneigh- Georgia. Coypus were introducedin Georgia boringHolland (Litjens 1980). between 1930 and 1932 (Aliev 1967, Vinogradov Bulgaria. Aliev(1967) gave no indicationof coy- and Argiropulo1941). Duringperiods of highpop- pus in Bulgaria.Mitchell-Jones et al. (1999) report- ulationdensity, coypus have leveledthe herbaceous ed the animalsalong the borderswith Greece and cover in marshyareas (Laurie 1946). Romania. Israel. Coypus are presentin Israel. From 1948 The Czech Republicand Slovakia. Accordingto to 1966 theywere observedliving in varioushabi- Aliev (1967) and Kinleret al. (1987), coypus were tats and under various environmentalconditions raisedin captivityin Czechoslovakia.They are cur- (Ehrlich 1967). Originally,they were raised on rentlyin both republics,with more populationsin farms;now escaped, they cause damage to fish- the Czech Republic(Mitchell-Jones et al. 1999). ponds (Aliev 1967). Denmark. Stubbe (1989) reportedthat coypus Kazakhstan. Coypus were released on reser- were observedin the wild in Denmarkin the 1930s voirsin Kazakhstanbetween 1930 and 1932 (Aliev and 40s but succumbedto the harshwinters there. 1967,Vinogradov and Argiropulo1941); theircur- Currentlyno coypus are reported in Denmark rentstatus is unknown. (Mitchell-Joneset al. 1999). Russia. Coypuswere successfullyintroduced on England. The firstcoypus were importedinto a large scale in 1926 (Laurie 1946). Farmswere GreatBritain in 1929 forfur farms (Laurie 1946). established in the KrasnodarTerritory of Russia The firsttrapping campaigns were in the early from 1933 to 1934 (Aliev 1967). Coypus were 1940s (Laurie 1946), and effortsto reduce their released between 380 and 450 N latitude and numbersoccurred in the 1960s (Norris 1967a, b). became acclimatizedin lakes and floodplainsof In 1981 a 10-yeareradication campaign employing largerivers in thatregion (Vinogradov and Argirop- 24 trapperscommenced (Gosling and Baker1987). ulo 1941). On 10 January1989 no coypushad been trappedin Tajikistan. A successfulexperiment was set up 21 months and the trapping campaign was by governmentagencies to acclimatizecoypus to declared a success and terminated(Gosling and reservoirsin the valleyof the RiverVakhsh in 1949 Baker 1989). (Aliev 1967). Finland. Aliev's(1967) rangemap indicatedwild Turkey. Coypus were establishedin large reed coypu populationshere. In the early1990s coypus swamps and fensin the northeasterncorner of the escaped fromfur farms and a wild populationexist- country(Kinler et al. 1987); theircurrent status is ed in the southof Finlandnear Turku (K.Jutila, per- unknown. sonal communication).Coypus are currentlylisted Turkmenistan.Coypus were released on reser- as a game animal (Hunters' Centralorganisation voirs between 1930 and 1932 (Aliev 1967, Vino- 2000). However,they are now classifiedas extinct gradovand Argiropulo1941); theircurrent status is in thewild (Mitchell-Joneset al. 1999). It is hypoth- unknown. esized thatthey could not toleratethe harshwin- Europe ters(K. Jutila,personal communication). Coypus have been introducedthroughout most France. Coypus were introducedinto Franceas of Europe (Table 1). In generalthey are considered early as 1882, and fur farmingbegan in earnest a pest in western European countries and a between 1925 and 1928 (Bourdelle 1939). Some resourcein easternEuropean countrieswhere fur coypus escaped captivityand became feral(Bour- is morevalued. MostEuropean studies of coypusin delle 1939). By 1979 theywere officiallyclassified the wild come fromItaly, France, and England.The as pests (Reggianiet al. 1993). From1974 to 1985,

This content downloaded from 158.135.136.72 on Fri, 1 Aug 2014 14:48:22 PM All use subject to JSTOR Terms and Conditions 168 WildlffeSoctety Bulletin 2002,30(1): 162-175 theyincreased in numberand are now the subject Italy,including Sicily and Sardinia,and are presently of many eradication campaigns (Lagaude 1975, regarded as a pest because of the damage they Abbas 1991, Moutou 1997), in part because they cause to rice farms(Velatta and Ragni1991a and b; host liver flukes (Fasciola hepatica) and are Gariboldi1993; DeCiechi and Prigioni1997; Cocchi thoughtto spread the disease to cows (Moutou and Riga 1999). Italy'semphasis is on controlling 1997, Menard et al. 2000). Coypus are found the populationsto reduce damage (Spacone et al. throughoutFrance (Moutou 1997), but the popula- 1991) tion tends to be concentratedin centraland west- Netherlands. Coypus were introduced here central France (Lagaude 1975, Dagault and around 1930 for fur farmingand by 1940 were Saboureau 1990). observedin the wild (Litjens1980). Because they Germany. Coypuswere firstintroduced to East- damage levees and the sugarbeet crop,they were ern Germanyin 1926 (Stubbe 1992). Some time considereda candidatefor eradication by European afterwardthey escaped captivity,and by 1935 small agencies (Litjens 1980, Barends 1998). Controlis wild colonies began to appear in the Elbe-Trave by trapping(Litjens 1984, Barends 1998). Despite Canal; however, these colonies were generally populationlosses fromtrapping and harshwinters, short-lived(van den Brink1968, Stubbe 1992,Geb- they persist in the Netherlandsbecause thermal hardt1996). West Germanyhad a huntingseason pollutionin riversallow some to surviveharsh win- forcoypus by the 1950s and was attributingeco- tersand because theyimmigrate from Belgium and nomicdamage (e.g.,undermining of dams) to them Germany(Litjens 1980). (Stubbe 1989). Even so, releases continuedand in Norway. Captivecoypu breedinghas been prac- 1991 over 400 coypuswere illegallyreleased in the ticed in Norway(Laurie 1946). Aliev's(1967) dis- Brandenburg region of East Germany (Stubbe tributionmap did not show breedingpopulations 1992). The range of coypus continuesto expand, present.It is unknownwhether they ever escaped, and recentlythey appeared in the Dresden area but currentlyno wild populationsexist (Mitchell- (Feileret al. 1999). Todaypopulations are generally Joneset al. 1999). low because theyare keptin check by cold winters Poland. Coypusare bredin captivitybut also are and are caughtin the heavyhunting for muskrats, managed in a semiwild state on ponds (Ehrlich which are morepopulous and are consideredmore 1962,Kinler et al. 1987,Labecka 1990). The animals of a pest thancoypus (Stubbe 1989,1992; Gebhardt were firstobserved in the wild in 1948 (Ehrlich 1996). They are consideredmore of a pest in the 1967). In a semicaptivesystem, ponds are drained formerWest Germany than in the formerEast Ger- in winterto protectthe coypus fromfreezing on many,where trapping is greater (Stubbe 1989, the ice (Ehrlich1962). Today coypu farmsare an 1992; Gebhardt1996). importantindustry in Poland (Niedzwiadek et al. Greece. Coypus were raised in captivityin 1988), and wild populations are also present Greece (Aliev 1967). Between 1948 and 1966 they (Mitchell-Joneset al. 1999). were observed in the wild in a varietyof habitats Romania. Aliev's(1967) reviewdid not indicate such as ponds,lakes, ditches, rivers, swamps, marsh- the animalswere in Romania. In his reviewof coy- es, meadows, and wooded areas (Ehrlich 1967). pus in Germany,Stubbe (1989) reportedthat they There are currentlywild coypu populationsin the had been observedin the wild in Romania. Popu- northernpart of Greece (Mitchell-Joneset al. 1999). lations are on the southernborder with Bulgaria Hungary. Coypus have been farmedin Hungary and along the Black Sea (Mitchell-Joneset al. 1999). (Sztojkovet al. 1982, Kinleret al. 1987, Salyiet al. Spain. Coypus were bred in Spain,and Aliev's 1988) and today are presentin southernHungary (1967) rangemap showed wild stockpresent. Cur- on the border(Mitchell-Jones et al. 1999). rentlythey are not in the wild (Mitchell-Joneset al. Ireland. Aliev's(1967) rangemap showed their 1999). presence here,but he providedno furtherinforma- Sweden. Aliev's(1967) rangemap indicatedwild tion. Mitchell-Joneset al. (1999) reportedthem as populationsof coypus here. They continueto be not being present. raisedon furfarms, but Mitchell-Joneset al. (1999) Italy. Coypus were firstimported into Italyin indicatedthat wild populationsno longerexisted. 1928 forcommercial use (Cocchi and Riga 1999), Switzerland. Mitchell-Joneset al. (1999) report- and were firstreported in the wild in 1960 (Reg- ed coypus within Switzerland,although Aliev giani et al. 1993). They have spread throughout (1967) did not.

This content downloaded from 158.135.136.72 on Fri, 1 Aug 2014 14:48:22 PM All use subject to JSTOR Terms and Conditions Literaturereview of coypus worldwide * Carterand Leonard 169

The Former Federal Republic of Yugoslavia. Pursley1978), and it is increasing(Bounds 2000). Aliev (1967) reportedthat coypus were raised in Arkansas. Coypuswere importedhere in the late captivityin Yugoslavia, although he did not indicate 1940s as weed controlagents (Evans 1983). They where. Currentlythey are foundin Macedonianear mustnot have formed a viablepopulation at thattime the borderswith Greece and Albania,Croatia, and because theywere not noted again untilthe early the currentborders of the Federal Republic of 1960s,when theyreentered the state by range expan- Yugoslavia(Mitchell-Jones et al. 1999). sionfrom Louisiana (Bailey and Heidt1978). Theyare now feralthroughout much of Arkansas(Bailey and NorthAmerica Heidt 1978) and are increasing(Bounds 2000). The coypu is widelyestablished in NorthAmeri- California. The firstcoypus importedinto the ca (Table 2). The firstrecorded effort to establisha UnitedStates were to ElizabethLake, California, for populationin NorthAmerica was at ElizabethLake, furfarming in 1899 (Evans 1970). Althoughthis California,in 1899; the effortfailed because the attemptwas not successful,subsequent importa- animalsdid not reproduce(Evans 1970). The first tions musthave been made because by 1940 Cali- successful reproductionoccurred in Quebec in forniahad a smallferal population (Schitoskey et al. 1927 (Evans 1970). We could findno attemptsto 1972). Conditionsthere are generallynot favorable introducecoypus into Mexico. Most introductions forcoypus in thewild. A smalleradication program in the UnitedStates and Canada were the resultsof was successful,and Deems and Pursley (1978) escapes fromfur farms and occurredin the 1930s reportedthe animalswere eradicatedby 1978. and 1940s. Coypus were introducedin the United Colorado. Coypuswere importedinto Colorado Statesto removenuisance aquatic vegetation. for fur farming,and Deems and Pursley (1978) UnitedStates reported a small feral population. However, During the 1930s coypu ranches were estab- Bounds'(2000) surveyfound no coypus there. lished in Louisiana,Michigan, New Mexico, Ohio, Delaware. Bounds (2000) reportedthat coypu Oregon,Utah, and Washington. These ranchesfailed populationsin Delaware were expanding.We have duringWorld War II because of poor pelt prices no informationon when they mighthave been (Kinleret al. 1987),and the animalswere oftensim- introduced;however, it seems likelythey came in plyreleased. Startingin the 1930s and 1940s,entre- fromMaryland. There have been feralcoypu popu- preneurs promoted coypus as weed cuttersand lationsin Marylandsince the 1930s (Paradiso1969, transplantedthem throughout the Southeast(Dozi- Wilneret al. 1979), and there are no barriersto er 1952, Evans 1970). State and federalagencies coypu movementbetween the two states. purposelyreleased coypus in Alabama,Arkansas, Florida. Coypus moved into Florida by range Georgia, Kentucky,Maryland, Mississippi, Okla- expansion and via escapes or releases fromfur homa,and inlandTexas and Louisiana to promote farms in the 1950s (Brown 1975). They have the furtrade and controlaquatic vegetation(Dozi- escaped captivityor have been released and are er 1952,Evans 1970). now feral(Brown 1975, Deems and Pursley1978, Coypus have been introducedor have migrated Bounds 2000). into30 states(Table 2,Evans 1970,Deems and Purs- Georgia. Coypuswere introducedhere for weed ley 1978,Bounds 2000). The mostrecent survey of controlby stateand federalagencies (Dozier 1952; coypus in the United States was completed by Evans 1970,1983). They are now feral(Deems and Bounds (2000), who contacted federal wildlife Pursley 1978) and their population is stable refugemanagers and representativesof statenatu- (Bounds 2000). ral resource agencies to determinepresence and Idaho. A smallferal population existed (Deems status of coypus in theirrespective jurisdictions. and Pursley 1978) but is now reported extinct The respondentsreported coypus in 15 states. (Bounds 2000). Most studies of coypus in the wild in the United Illinois. Coypus were reported in Illinois Stateshave takenplace in Louisianaand Maryland. (Kennedyand Kennedy1998). Bounds'(2000) sur- Alabama. Coypus were broughtinto Alabama vey did not findany. by the state'sDepartment of Conservationand by Indiana. Coypuswere importedinto Indiana for privateindividuals in 1949 as weed controlagents fur farming;some escaped but were eradicated (Lueth 1949, Dozier 1952, Evans 1970). A viable (Deems and Pursley1978). feralpopulation now existsin Alabama(Deems and Kansas. Coypus were importedinto Kansas for

This content downloaded from 158.135.136.72 on Fri, 1 Aug 2014 14:48:22 PM All use subject to JSTOR Terms and Conditions 170 WildlifeSociety Bulletin 2002, 30(1):162-175 fur farming(Deems and Pursley 1978). Bounds' kept coypu densitiesbelow a level at which signif- (2000) surveyfound none. icantdamage could occur. Since thattime, the drop Kentucky.Coypus were introducedby stateand in furprices has reduced huntingpressure and a federalagencies forweed control,but theydid not succession of mild wintershas occurred. Coypu survive (Evans 1970, 1983; Deems and Pursley populationsare now consideredtoo high,and stud- 1978). ies are underwayto determinethe best strategyfor Louisiana. Coypus were firstintroduced near eradicating them from Maryland's wetlands New Orleans in the early 1930s, but they were (Haramisand Colonas 1999). quicklytrapped out (Evans 1970, Baileyand Heidt Michigan. Coypuswere firstfarmed in Michigan 1978). The species was broughtback intoLouisiana in the 1930s (Evans 1970, 1983); althoughsome in 1938 forfur farming, and some escaped in 1940 were accidentallyreleased, they did not survivein by burrowingout of pens and climbingover fences the wild (Deems and Pursley1978, Bounds 2000). damagedby a hurricane(Evans 1970,Lowery 1974, Minnesota. Coypus have been presentin Min- Bailey and Heidt 1978). These animals quickly nesotaon furfarms (Hazard 1982). Some were acci- expanded theirrange, and in less than2 yearscoy- dentallyreleased into the wild but did not survive pus were reportedin Lacassine NationalWildlife (Deems and Pursley1978). Refugenear Lake Arthur,approximately 78 km by Mississippi. Coypus enteredthis statevia range water from their purported release site (Dozier expansion fromLouisiana and importationby pri- 1952). Coypus are feralin the stateand are chiefly vate individuals(Wolfe 1971, Deems and Pursley controlledby trappingand (Lowery 1974, 1978, Kennedyand Kennedy1998). Theirpopula- Deems and Pursley 1978, Wolfe and Bradshaw tion is stable(Bounds 2000). 1987). Duringthe 1960s and 1970s,coypu popula- Missouri. Coypus were present here (Deems tions were controlledby furtrappers; however, as and Pursley1978, Kennedy and Kennedy1998), but furprices dropped in the 1980s,so did the number Bounds (2000) reportedthat they no longerare. of coypus caught (Kinler et al. 1987; Louisiana Montana. Hall (1981) mentioned coypus in Departmentof Wildlifeand Fisheries,unpublished Montana,but Bounds' (2000) surveyreported they report). The drop in trappingresulted in a large are no longerpresent. increasein the coypu populationand loss of marsh Nebraska. Coypus were reportedin Nebraska, due to overgrazing(Kinler et al. 1987; Louisiana but none have survivedin the wild (Deems and Departmentof Wildlifeand Fisheries,unpublished Pursley1978, Hall 1981). report).To increasetrapping pressure, the stateof New Mexico. Coypus were introducedhere for Louisianais now promotingcoypu meat as a low- furfarming (Evans 1970, 1983; Deems and Pursley fat,nutritious product for retail and specialtymar- 1978) and there were small feral populations kets (E. Mouton,Louisiana Departmentof Wildlife (Deems and Pursley 1978). However, Bounds and Fisheries,personal communication). (2000) reportedthey are no longerpresent. Maryland. In the late 1930s and early 1940s, North Carolina. In 1941 a huntingclub intro- coypu care and breedingexperiments were con- duced coypus to HatterasIsland and theyquickly ducted by the United States Armyat Blackwater spread(Milne and Quay 1967). Deems and Pursley NationalWildlife Refuge in DorchesterCounty (Par- (1978) reported a small viable feral population. adiso 1969). Priorto 1949 the stateof Maryland did Currentlyit is expanding(Bounds 2000). not keep accurate records of annual furcatches. Ohio. Coypus were importedfor fur farming in Paradiso (1969) reported that in 1949 trappers 1937 (Bednarik 1961). Some escaped or were reported4 coypustaken, but no otheranimals were released into the wild. No wild populationscur- reporteduntil 1956. Wilneret al. (1979) reported rentlyexist (Bednarik 1961, Deems and Pursley thatferal coypus were firstdetected in 1952. By 1978,Bounds 2000). 1969 Paradiso (1969) stated that coypus existed Oklahoma. Coypushave been recordedin Okla- only in DorchesterCounty and were precariously homa and are presentin low numbers(Deems and established. Ten years later,Wilner et al. (1979) Pursley1978). Theirpopulation is currentlystable reportedthat eradication efforts were unsuccessful (Bounds 2000). and thatthe populationhad dispersedthroughout Oregon. Coypus were importedfor fur farming the state into all suitablehabitats. In 1982 Wilner in 1937; some escaped and feralones were official- (1982) noted thattrapping and long,cold winters ly recorded as early as 1941 (Larrison 1943).

This content downloaded from 158.135.136.72 on Fri, 1 Aug 2014 14:48:22 PM All use subject to JSTOR Terms and Conditions Literaturereview of coypus worldwide * Carterand Leonard 171

Currentlya viable wild population exists (Deems Quebec. The earliestrecord of successfulrepro- and Pursley1978) and is expanding(Bounds 2000). ductionof fur-farmed coypus in NorthAmerica was Texas. A hurricaneis thoughtto have scattered reportedfrom Quebec, withanimals imported from coypus into east Texas from Louisiana in 1941 Germanyin 1927 (Evans 1970). They were feralin (Evans 1983). Startingaround 1947-1948, private the Ottawa River drainage of Quebec (Banfield individualsintroduced the animalsthroughout east 1974). Texas because of theirvalue as a furbearerand rep- Mexico utationas a "weed cutter"(Dozier 1952). They are A range map presented by Schwandt-Arbogast now reportedin everyregion of Texas except the (2000) showed coypusin northernMexico, in states highplains (Deems and Pursley1978, Hollander et along the borderwith Texas and along the Gulfof al. 1992). Recentlythey have moved up the Rio Mexico. Eyewitnesseshave reportedseeing them in GrandeValley as fareast as Big Bend NationalPark Rio Grande Valley on both sides of the United and are threateningwetlands along the river. States-Mexicoborder in the Big Bend region (J. Bounds (2000) reportedthe populationas stable. Carter,personal observation). Because oftheir distri- Tennessee.The firstcoypu was observedhere in bution pattern,we hypothesizethat the animals 1996; there is a small wild population (Kennedy enteredMexico through range expansion fromTexas. and Kennedy 1998), reported to be expanding (Bounds (2000). Utah. Coypus were importedfor fur farming in Discussion 1939 (Evans 1970, 1983), and althoughsome may The coypuhas been introducedto everycontinent have escaped, they did not survive in the wild outsideof its native South America except Antarctica (Deems and Pursley1978, Bounds 2000). and Australia.It has become establishedin new areas Virginia. It is hypothesizedthat coypus expand- outside its native range primarilyby 3 processes: ed intoVirginia from North Carolina (Deems and deliberateintroductions into new areas,escape or Pursley1978, Pagels 1989). Currentlythe popula- releasefrom fur farms, and rangeexpansion. Though tion is stable(Bounds 2000). not all initialcoypu invasions are successful,as in the Washington.Coypus were importedin the late case of Arkansas,subsequent invasions may succeed 1930s and early1940s forfur farms (Larrison 1943), (Baileyand Heidt1978, Evans 1983). and feralindividuals were reportedas earlyas 1941 The primaryreason coypus are broughtto a new (Larrison1943). Currentlya viablewild population area is fortheir pelt. Anotheris theirability to con- exists(Deems and Pursley1978) and is expanding trolaquatic weeds (Dozier 1952;Evans 1970,1983), (Bounds 2000). Canada BritishColumbia. Coy- pus were introducedfor furfarming, and the earli- est recordof feralcoypus is from 1943 (Holdom 1944). A feralpopulation is stillpresent (Deems and Pursley1978). Ontario. Coypus are feralin the Ottawa River drainageof Ontario (Ban- field1974). Nova Scotia. Coypus wereimported for fur farm- ingand somewere acciden- tallyreleased into the wild, butcurrently no wildpopu- lationsare knownto exist Coypuexclosures on BlackwaterNational Wildlife Refuge in Maryland illustrate the effects of (Deems and Pursley1978). coypuon marshloss. Phototaken November 2000 byJacoby Carter.

This content downloaded from 158.135.136.72 on Fri, 1 Aug 2014 14:48:22 PM All use subject to JSTOR Terms and Conditions 172 WildltfeSociety Bulletin 2002, 30(1):162-175 but this abilityhas limitedeffectiveness because cient habitat,coypu eradication efforts the animals do not necessarilyeat targetedweed have seldom been successful. species and often find nontargetedplants more The first successful coypu eradication effort palatable (Evans 1970, 1983). Earlystudies in the occurredin California(Evans 1970). However,the southern United States indicated that coypus best-documentedexample of a successfuleradica- caused onlyminor or localized marshdamage and tion campaign is from England (Gosling 1989). theywere consideredless destructivethan native Eradicationis difficultbecause veryfew coypus are muskrats(Dozier 1952, Harrisand Webert 1962). needed to establisha populationif suitablehabitat However,those early studies were done in years is present,and once established,their populations with extensivetrapping. As pelt prices fell,so did can grow quickly. Once large populationsare in trappingactivity and coypu populationsincreased place,it is difficultto assembleenough resources to (Kinleret al. 1987;Louisiana Department of Wildlife completelyeliminate them. Thus, coypu popula- and Fisheries,unpublished report). Later studies tions that are isolated from reinvasionand are indicate that coypus can cause extensive marsh reduced to a manageablelevel by adverseweather loss (Wilneret al. 1979, Foote and Johnson1993, are most susceptibleto eradication.The 3 condi- Linscombeand Kinler1997). Problemsassociated tionsneeded fora successfuleradication effort are: with high coypu populationsfall into severalcate- the populationshould be isolatedfrom sources of gories:destruction of marshhabitat, destruction of new immigration,inclement weather or harshcon- water controlstructures such as dykesand levees, ditionsare needed to reduce largepopulations, and destructionof agriculturalcrops, and the factthat trappingmust be continueduntil no coypusare left. the animalscan serveas repositoriesof a varietyof These factorspresent some interestingchoices diseases (Evans 1983,Howerth et al. 1994, Moutou for management. In south Louisiana,the coypu 1997,Linscombe and Kinler1997). Economicdam- populationis large and well dispersed. Even if an age caused by coypus has promptedgovernment appropriate weather event occurred to greatly agencies to implementcontrol or eradicationpro- reduce the population,the probabilityis high that grams(Gosling 1989). However,coypus are gener- individualswould escape an intenseremoval cam- alistfeeders, have a highbirthrate, and theirhabitat paignor reinvadefrom neighboring states. An erad- requirementsare not very rigid (Evans 1983). ication campaign is much more likelyto be suc- Because of these factors,control and eradication cessfulin Big Bend NationalPark, because the park are oftenhard to implement. contains limitedcoypu habitat and the invasion Not all of the countriesthat have importedcoy- routeinto it is througha narrow,controllable corri- pus have had problems with them. In eastern dor,the Rio Grande. Europe and the Caucasus, the coypu is highly Coypu distributionthroughout North America regardedfor its pelt (Aliev 1967). These countries presentssome interestingissues. The animalshave have breeding programsto produce high-quality been introducedto 30 statesbut currentlyare pres- pelts and have establishedpedigrees (Aliev 1967). ent in only 15, mostlysouthern states. The states So while Louisianaand Argentinahave the greatest where coypus do not currentlyexist probably rep- harvest of feral animals,the highest quality and resent areas where wintersare harsh or wetland quantityof coypu pelts in the world come from habitatis limited. Harsh winters cannot be the semicaptivepopulations in easternEurope and the entirestory, however, because coypus are reported Caucasus (Kinleret al. 1987). in the Ottawa River drainage in the Canadian Cold winters seem to be the most effective provincesof Quebec and Ontario. It is also inter- mechanismto limitcoypu populationexpansion in estingto note thatcoypus are not reportedas being thewild (Ehrlich1967, Stubbe 1992, Bounds 2000). in SouthCarolina, even thoughthey are presentor For example, though coypus were either intro- expandingin neighboringGeorgia and NorthCar- duced or escaped into the wild in all the Scandina- olina (Bounds 2000). vian countries,currently no wild populationsexist Coypushave been transportedall overthe world in the region (Mitchell-Joneset al. 1999). In con- due to perceptionsabout theireconomic value and trast,muskrats, an ecologically similarbut more our limitedunderstanding of the effectsthat non- cold-hardyspecies fromNorth America,are suc- indigenousspecies can have on habitats.Although cessful throughoutScandinavia (Mitchell-Jones et some areas have benefited,others have suffered. al. 1999). In regionswith mild wintersand suffi- Once coypus are introduced,their eradicationis

This content downloaded from 158.135.136.72 on Fri, 1 Aug 2014 14:48:22 PM All use subject to JSTOR Terms and Conditions Literaturereview of coypus worldwide * Carterand Leonard 173

not always possible. In fact,without the 3 condi- DAGAULT, N., AND M. SABOUREAU. 1990. Caracteristiquesde la tions listedabove, eradicationis not possible with reproductiondu Myocastor (Myocastor coypus M.) male currenttechnology, and managingthe population dans la regiondu MaraisPoitevin. Canadian Journal of Zool- ogy 68:1584-1589. (In Frenchwith English abstract). to reduce damage to habitatand crops is the best DECIECHI, R. AND C. PRIGIONI. 1997. Distributionand ecology of thatcan be achieved. the coypu (Myocastorcoypus) in the valley of the Ticino River (NorthwesternItaly). Atti Della Societa Italiana Di Acknowledgments. We thank L. Foote, M. Scienze NaturaliE Del Museo Civico Di StoriaNaturale Di Gosling,and G. Piero for theiradvice and sugges- Milano 138:13-23. (Fromabstract). DEEMS E. E, AND D. PURSLEY, editors. 1978. NorthAmerican tions. We thankP. Klerks, A. Martucci,K. Jutila,and furbearers:their management, research and harveststatus in N.Takimotofor providing translations of articles. We 1976. InternationalAssociation of Fishand WildlifeAgencies thankC. Hamburg,S. Hartley,and N. Gormanousfor in cooperation with the MarylandDepartment of Natural assistancein preparingour graphics. We thankJ. Resources-WildlifeAdministration. University of Maryland, Buys and L. Broussardfor their assistance with College Park,USA. DONCASTER, C. P, AND T. MICOL. 1990. Responseby coypusto cat- researchand retrieval of journalarticles. We thank astrophicevents of cold and flooding.Holarctic Ecology 13: K. Yokoto-Carterfor providing us with a copy and 98-104. translationof the Tahnka. J. Carter thanks his branch DOZIER, H. L. 1952. The presentstatus and futureof nutriain the chiefs(past and present),E. Proffitt,R. Best,and V. southeaststates. Proceedingsof the AnnualConference of Burkett,for their support. B. Leonardthanks Dr. P the SoutheasternAssociation of Game and FishCommission- ers 6: 368-373 Klerksfor his advice and support,and his wife,E EHRLICH, S. 1962. Experimenton the adaptationof nutria to win- Leonard,for her patience and understanding.The terconditions. Journal of Mammalogy43:418 UnitedStates Geological Survey's National Wetlands EHRLICH, S. 1967. Field studiesin the adaptationof nutriato sea- ResearchCenter provided financial support for this sonal variations.Mammalia 31:347-360. project. EVANS,J. 1970. About nutriaand theircontrol. UnitedStates Bureau of Sport Fisheries and Wildlife,Denver Wildlife ResearchCenter, Denver, Colorado, USA. EVANS,J. 1983. Nutria. Pages B61-B70 in Preventionand con- Literaturecited trolof wildlifedamage, R. M.Timm,editor. Great Plains Agri- ABBAS,A. 1991. Feedingstrategy of coypu. Journalof Zoology culturalCouncil,Wildlife Resource Committee and Nebraska 224:385-401. CooperativeExtension Service Instituteof Agricultureand ALIEv,E F 1967. Numericalchanges and the populationstruc- NaturalResources, University of Nebraska,Lincoln, USA. ture of the coypu,Myocastor coypus (Molina,1782), in dif- FEILER, A., H.-J.KAPISCHKE, K. MISSBACH, M. WILHELM, AND U. ferentcountries. SeaugetierkunklicheMitteilungen 15:238- ZOEPHEL. 1999. The mammalsof Dresden and its surround- 242. ings(Mammalia). Third contribution to theMammal Fauna of BAILEY,J. W, AND G. A. HEIDT. 1978. Range and statusof the Saxony. FaunistischAbhandlungen (Dresden) 21:341-375 nutriaMyocastor coypus, in Arkansas. Proceedingsof the (fromabstract). ArkansasAcademy of Science 32:25-27. FOOTE,A. L., AND L. A. JOHNSON. 1993. Plantstand development BAKER,S. J.,AND C. N. CLARKE.1988. Cage trappingcoypus (Myo- in Louisiana coastal wetlands:nutria grazing effects. Pro- castor coypus) on baited rafts.Journal of Applied Ecology ceedings of theThirteenth Annual Meeting of the American 25: 41-48. Society of Wetland Scientists,May 31-June 6 1992, New BANFIELD,A. W F 1974. The Mammalsof Canada. Universityof Orleans,Louisiana, USA. Toronto,Toronto,Ontario, Canada. GARIBOLDI, A. 1993. La Nutria(Myocastor coypus) in Lombardia. BARENDS,E 1998. Muskratand coypu: beautifulanimals, but a SupplementAlle RicercheDi Biologia Della Selvaggina21: nuisance. Levende Natuur99:12-17 (fromabstract). 259-262. (Italianwith English abstract). BEDNARIK,K. E. 1961. Nutriain Ohio. Ohio Divisionof Wildlife, GEBHARDT, H. 1996. Ecological and economic consequences of Columbus,USA. introductionsof exotic wildlife(birds and )in Ger- BOUNDS,D. L. 2000. Nutria:an invasivespecies of nationalcon- many.Wildlife Biology 2: 205-211. cern. WetlandJournal 12:9-16. GOSLING,L. M. 1981. The dynamicsand controlof a feralcoypu BOuRDELLE,E. 1939. Americanmammals introduced into France population. Pages 1806-1825 in WorldwideFurbearer Con- in the contemporaryperiod, especially Myocastor and Onda- ferenceProceedings. 3-11 August1980, Frostburg, Maryland, tra. Journalof Mammalogy20:287-291. USA. BRoWN,L. N. 1975. Ecologicalrelationship and breedingbiolo- GOSLING, M. 1989. Extinctionto order.New Scientist121:44-49. gy of the nutria(Myocastor coypus) in the Tampa,Florida, GOSLING, L. M., S. J.BAKER, AND J.R. SKINNER. 1983. A simulation area. Journalof Mammalogy56:928-930. approach to investigatingthe response of a coypu popula- CARTER,J., A. L. FOOTE,AND L. A. JOHNSON. 1999. Modelingthe tionto climaticvariation. EPPO Bulletin13:183-192. effectof nutria(Myocastor coypus) on wetland loss. Wet- GOSLING, L. M., AND S. J.BAKER. 1987. Planningand monitoring lands 29:209-219. an attemptto eradicatecoypus fromBritain. Pages 100-1 13 COCCHI,R., AND E RIGA. 1999. NutriaMyocastor coypus (Moli- in the Proceedingsof a Symposiumheld at the Zoological na, 1782). Iconografia Dei MammiferiD'Italia Isituto Societyof London.28-29 November 1986, London,United Nazionale Per La Fauna Selvatica38 (In Italian). Kingdom.

This content downloaded from 158.135.136.72 on Fri, 1 Aug 2014 14:48:22 PM All use subject to JSTOR Terms and Conditions 174 WildlIfeSociety Buetin 2002,30(1):162-175

GOSLING,L. M., ANDS. J.BAKER. 1989. The eradicationof muskrats Ministryof NaturalResourcesToronto, Canada. and coypus fromBritain. Biological Journal of the Linnean LABECKA,S. 1990. Studieson the relationshipsbetween mating Society38: 39-51. time and whelping and reproductionindices in nutria. GOSLING,L. M., S. J.BAKER, AND C. N. CLARK. 1988. An attemptto Roczniki Naukowe Zootechniki 17:169-178. (Polish with removecoypus (Myocastorcoypus) froma wetlandhabitat Englishabstract). in East Anglia.Journal of Applied Ecology25:49-62. LAGAUDE,V 1975. Le ragondinen Camargue. Phytoma-Defense GRAc(E,J.B. 1992. The impactof nutria(Myocastor coypus) on Des Cultures27:18-22. GulfCoast wetlands;Symposium Introduction. Pages 70-74 LARRISON,E. J. 1943. Feralcoypus in the PacificNorthwest. The in Proceedings of the thirteenthannual meeting of the Murrelet24:3-9. AmericanSociety of Wetland Scientists. 31 May-6June 1992, LAURIE,E. M. 0. 1946. The coypu (Myocastorcoypus) in Great New Orleans,Louisiana, USA. Britain.Journal of AnimalEcology 15:22-34. HALL,E. R. 1981. The mammalsof NorthAmerica. Second edi- LEBLANC,D. J. 1994. Nutria.Pages B71-B80 in S. E. Hygnstrom, tion. JohnWiley & Sons,New York,New York,USA. R. M.Timm, and G. E. Larson,editors. Prevention and control HAN,S. H., D. S. KONG,ANDJ. Y CHA. 1999. Exoticanimal species of wildlifedamage. Universityof Nebraska Cooperative in South Korea. The Ecological Societyof Americaeighty- Extension,Lincoln, USA. fourthannual meeting. 8-12 August1999, Spokane,Washing- LINSCOMBE,G., ANDN. KINLER. 1997. A survey of vegetative dam- ton,USA. (Abstract). age caused by nutriaherbivory in the Baratariaand Terre- HARAMIS,M., AND R. COiLONAS.1999. The effectof nutria (Myocas- bonne Basins.Barataria-Terrebone National Estuary Program. tor coypus) on marsh loss in the lower eastern shore of No. 31. NichollsState UniversityThibodaux, Louisiana, USA. Maryland:An enclosurestudy. United States Geological Sur- LINSCOMBE,G., N. KINLER,AND V WRIGHT.1981. Nutria population vey Internet article http://www.pwrc.usgs.gov/resshow/ densityand vegetativechanges in brackishmarsh in coastal nutria.htm(Date accessed: 11 February,2001). Louisiana. Pages 129-141 in Proceedingsof theWorldwide HARPER,D. M., K. M. MAVUTI,AND S. M. MUCHIRI. 1990. Ecology FurbearerConference. 3-11 August 1980, Frostburg,Mary- and managementof Lake Naivasha,Kenya, in relationto cli- land,USA. matic change, alien species' introduction,and agricultural LITJENs,B. E. J. 1980. De beverrat,Myocastor coypus (Molina), development.Environmental Conservation 17:328-336. in Nederland.Lutra 23:45-53. (Dutch withEnglish abstract) HARRIS,V T 1956. The nutriaas a wild furmammal in Louisiana. LITJENS,B. E. J. 1984. De beverratMyocastor coypus in Neder- NorthAmerican Wildlife Conference 21:474-485. lands limburgen aangrenzendegebieden. Lutra 27: 208 HARRIS,V T. ANDF WEBERT. 1962. Nutriafeeding activity and its (Dutch). effecton marshvegetation in southwesternLouisiana. Spe- LOWERY,G. H. 1974. The mammalsof Louisianaand its adjacent cial ScientificReport,Wildlife No. 64. UnitedStates Fish and waters. LouisianaState University, Baton Rouge, USA. WildlifeService, Washington, D.C., USA. LUETH,E X. 1949. The firstyear of nutriainvestigation on the HAzARDt,E. B. 1982. The mammalsof Minnesota. Universityof MobileDelta. Proceedingsof the AnnualConferences South- Minnesota,Minneapolis, USA. eastern Association of Game and Fish Commissioners3: HOLDOM,M. W 1944. Additionalnotes on feralcoypus. The 98-104. Murrelet25:30. MENARD,A., M. LHOSTIS, G. LERAY,S. MARCHANDEAU,M. PASCAL,N. HOLLANDER, R. D., R. N. ROBERTSON, AND R.J. KINUCAN. 1992. First ROUDOT,V MICHEL,AND A. CHAUVIN.2000. Inventory of wild record of the nutria,Myocastor coypus, in the Trans-Pecos rodentsand lagomorphsas naturalhosts of Fasciola hepati- regionof Texas. TexasJournal of Science 44:119. ca on a farm located in a humid area in Loire Atlantique HOWERTH,E. W, A. J. REEVES,M. R. McELVEEN,AND F W AUSTIN. (France). Parasite7:77-82. (Fromabstract). 1994. Surveyfor selected diseases in nutria(Myocastor coy- MILNE,R. C., ANDT. L. QUAY. 1967. The foods and feeding habits pus) in Louisiana.Journal of Wildlife Diseases 30:450-453. of the nutriaon HatterasIsland, North Carolina. Proceedings HUNTERS' CENTRALORGANISATION. 2000. http://www.mkj-jco.fi/ of the AnnualConference of the SoutheasternAssociation of m_sanasto.htm.(Date accessed: 22 October,2001). Game and Fish Commissioners20:112-123. KANWANICH,S. 1998. The argumentsfor and againstbreeding MITCHELL-JONES,A. J., AMORI,G., BOGDANOWICZ,W, KRYSTUFEK,B., nutrias. Bangkok Post Perspective, 8 February 1998 REIJNDERS,P J. H., SPITZENBERGER,E, STUBBE,M., THISSEN,J. B. Bangkok,Thailand. Internet http://www.bangkokpost.net M., VOHRALIK,V, ANDZIMA, J. 1999. The atlas of European (Date accessed: 22 October 2001). mammals.Academic, London, United Kingdom. KENNEDY,M. L., ANDP. K. KENNEDY. 1998. Firstrecord of nutria, MIURA,S. 1976. Dispersalof nutria in OkayamaPrefecture. Jour- Myocastorcoypus (Mammalia: Rodentia) in Tennessee. Brim- nal ofthe Mammalogical Society of Japan 6:231-237 (Japanese). leyana25:156-157. MOUTOU,E 1997. Aquatic and semi-aquaticmammals intro- KINLER,N. W 1992a. Biologyand ecology of nutria.Page 75 in duced into France.Risks and Consequences. BulletinFran- Proceedingsof the thirteenthannual meetingof the Ameri- cais De La Peche Et De La Pisciculture344/345:133-139. can Societyof Wetland Scientists 31 May- 6 June1992, New (Frenchwith English abstract). Orleans,Louisiana, USA. NIEDZWIADEK,S., J. A. N. ZAJAC,AND P BIELANSKI.1988. The state KINLER,N. W 1992b. Historicaland biologicaloverview. Pages and prospects for coypu breeding. BiuletynInformacyjny 8-14 in Nutria and management symposium. InstytutZootechniki 36:21-26 (fromabstract). LouisianaCooperative Extension Service, 8-9 October 1992, NORRIS,J. D. 1967a. The controlof coypus (Myocastorcoypus Baton Rouge,USA. Molina) by cage trapping. Journalof Applied Ecology 4: KINLER, N. W, G. LINSCOMBE, AND P R. RAMSEY. 1987. Nutria.Pages 167-189. 326-343 in M.Novak,J. A. Baker, M.E. Obbard, and B. Malloch, NoRlos, J.D. 1967b. A campaignagainst feral coypus (Myocas- editors. Wild furbearermanagement and conservationin tor coypus Molina) in Britain. Journal of Applied Ecology 4: NorthAmerica. The OntarioTrappers Association, Ontario 191-199.

This content downloaded from 158.135.136.72 on Fri, 1 Aug 2014 14:48:22 PM All use subject to JSTOR Terms and Conditions Literaturereview of coypus worldwide * Carterand Leonard 175

PAGELS,J. 1989. Of mice and men and othermammals. Virginia 115-116 in J.Salkind, translator. Fauna of the U.S.S.R. The Wildlife50: 16-21. SmithsonianInstitute and The NationalScience Foundation, PARADIso,J.L. 1969. Mammalsof Maryland.Pages 117-119 in Washington,D.C., USA. NorthAmerica Fauna, No. 66. Bureau of SportFisheries and WILNER,G. R. 1982. Nutria.Pages 1059-1076 in J.A. Chapman Wildlife,Washington,D.C., USA. and G. A. Feldhamereditors, Wild mammalsof NorthAmeri- REGGLANI,G., L. BoiTANI,S. D'ANTONI,AND R. DE STEFANO. 1993. ca. JohnHopkins University, Baltimore, Maryland, USA. Biologyand controlof the coypu in the Mediterraneanarea. WILNER,G. R., J.A. CHAPMAN,AND D. PURSLEY. 1979. Reproduc- SupplementAlle Ricerche Di Biologia Della Selvaggina21: tion,physiological responses, food habits,and abundance of 67-100. nutriaon Marylandmarshes. Wildlife Monographs 65. REGGIANI, G., L. BoiTANI,AND R. DE STEFANO. 1995. Population WOLFE,J. L. 1971. Mississippiland mammals.Mississippi Game dynamicsand regulationin the coypu Myocastorcoypus in and Fish Commission,Jackson,USA. CentralItaly. Ecography 18:138-146. WOLFE,J. A., ANDD. K. BRADsHAw.1987. Alligatorfeeding habits: RIDMONTRI, C. 1998. AgricultureMinistry won't promotenutria new data and a review. NortheastGulf Science 9:1-8. farming.Bangkok Post on-line. 10 February1998. http:// WOODS, C. A., L. CONTEREAS,G. WILNER-CHAPMAN,AND H. P WHID- www.bangkokpost.net(Date accessed: 22 October,2001). DEN. 1992. Myocastorcoypus. MammalianSpecies No. 138. SALYI,G., V SZTOJKOV,AND M. ILILBERTNE. 1988. vetch XIE, Y ANDZ. Li. 1999. Invasivespecies in China-an overview. (Coronilla varia L.) poisoningof the coypu. Magyarkaraar- Global DiversityForum-South and SoutheastAsia 1, 24-26 tudomaanyiLapja 43: 313-316 (Hungarian with English October 1999,Columbo, Sri Lanka. abstract). SCHITOSKEY,E, J.EVANS, AND G. K. LA VOE. 1972. Statusand con- trolof nutriain California.Pages 15-17 in Proceedingsof the fifthVertebrate Pest Conference. 7-9 March1972, Fresno, California,USA. SCHWANDT-ARBOGAsT,A. 2000. Coypu (Nutria)Myocastor coypus, ~~~~~~~~~~~~~~~~~~~~~~Fv-^^...... Burke Museum of NaturalHistory and Cultureweb page, University of Washington, Seattle, Washington, USA. http://www.washington.edu/burkemuseum/mammalogy/ myco.html,(Date accessed: 22 October 2001). ::vc::A... *- - s> ...... a;: - .:: SPACONE, R., M. MORELLINI,AND E. BRUNO. 1991. Controlmethods fora naturalizedpopulation of Nutria(Myocastor coypus). SupplementAlle Ricerche Di Biologia Della Selvaggina19: 725 (Italian). STEVENS, W E 1992. USDA animal damage controlnutria and muskratmanagement symposium. Pages 52-53.8-9 October Louisiana State UniversityAgricultural Center. Louisiana JacobyCarter (right) has been an ecologistat the UnitedStates CooperativeExtension Service, Baton Rouge,USA. GeologicalSurvey's National Wetlands Research Center in STUBBE, M. 1989. Die NutriaMyocastor coypus (Molina). Buch Lafayette,Louisiana, since 1995. Dr. Carterreceived a BS in Der Hege 1(Haarwild):630-639 (German). zoologyand a BS inbotany from the University ofWashington, Seattle in 1982 In in STUBBE, M. 1992. Die NutriaMyocastor coypus in Den Ostlichen 1992 he receivedan MS ecology from Cornell in and in 1995 a Ph.D in Deutschen Bundeslandern. SemiaquatischeSaugetiere 9.6: University Ithaca, NewYork, wildlifemanagement from the Universityof Massachusetts, 80-97 (Germanwith Englishabstract). Amherst. His primaryresearch interest is in using simulation SUVEGOVA,K., D. MERTIN,E. ORAVCOVA,AND J. RAFAY. 1994. Probi- models as a tool forecological research. Currentresearch proj- otic use in the managementof coypus. AivocisnaVyroba39: ectsinclude determining per-capita impact of nutriaon marsh 239-245 (Polish with Englishabstract). loss, modelingforest lost in the Xalapa regionof Mexico, mon- SzTojKov,V, E OSZTOTICS,AND L. MOLNAR.1982. A disease caused itoringand modelingsubmerged aquatic vegetation in brackish by Treponemabyodysenteriae in a large-scalecoypu farm. watersof the Gulfof Mexico, modelingharvest impacts on map MaygarAllatorvosok Lapja 37:94-97 (Hungarianwith Eng- turtles(Graptemys sp.) in northLouisiana, and evaluatingfire lish abstract). behavior models in coastal marshes. He is also interestedin new toaid in research.Two TUNYASIRI,Y 1998. Chuan says no to nutria farmingplea. developing technologies ecological recentprojects are evaluatingthe use of light-inducedfluores- 12 February 1998. Bangkok Post on-line. Http://www cence to assess plant healthand a new design fornutria exclo- .bangkokpost.net(Date accessed: 28 February2002). sures.A memberof The Wildlife Society since 1990, Dr. Carter VAN DEN BRINK, E H. 1968. A fieldguide to the mammalsof is also secretaryof the Asian Ecology Section and chair-electof Britainand Europe. Houghton Mifflin,Boston, Massachu- the InternationalAffairs Section of the EcologicalSociety of setts,USA. America. Billy P. Leonard (left)recently received his M.S. in VELATTA, E, AND B. RAGNI. 1991a. The coypus, Myocastorcoy- biologyfrom the Universityof Louisiana,Lafayette, where he pus, population of the Lake Trasimeno(Umbria). Supple- was also awardeda B.S. in resourcebiology and biodiversity. mentAlle RicercheDi Biologia Della Selvaggina 19(Decem- Billyis primarilyinterested in avian ecology as itrelates to habi- tat time will be at a new ber):311-326 (Italian). preferences.By press Billy working job doingbird surveys for the US ForestService. VELATTA,F, AND B. RAGNI. 1991b. Distributionof Myocastor coy- pus in Umbria(Central Italy). SupplementAlle RicercheDi BiologiaDella Selvaggina16:633-636 (Italian). VINOGRADOV,B. S., AND A. I. ARGIROPULO. 1941. Nutria. Pages

This content downloaded from 158.135.136.72 on Fri, 1 Aug 2014 14:48:22 PM All use subject to JSTOR Terms and Conditions