Differences in habitat use between the native Eurasian beaver and the invasive North American beaver in Finland

1 23 Biological In asions ISSN 1387-3547 Biol In asions DOI 10.1007/s10530-019-01919-9 Di e ences in habi a use be ween he na i e Eu asian bea e and he in asi e No h Ame ican bea e in Finland Riikka Alakoski, Kaa ina Kauhala & Vesa Selonen 1 23 You a icle is published unde he C ea i e Commons A ibu ion license which allows use s o ead, copy, dis ibu e and make de i a i e wo ks, as long as he au ho o he o iginal wo k is ci ed. You may sel - a chi e his a icle on you own websi e, an ins i u ional eposi o y o unde ’s eposi o y and make i publicly a ailable immedia ely. ORIGINAL PAPER Di e ences in habi a use be ween he na i e Eu asian bea e and he in asi e No h Ame ican bea e in Finland Riikka Alakoski .Kaa ina Kauhala .Vesa Selonen Recei ed: 9 July 2018 / Accep ed: 14 Janua y 2019 ÓThe Au ho (s) 2019 Abs ac Habi a equi emen s la gely de e mine he dis ibu ion and abundance o a species. An in asi e species can he e o e h ea en he su i al o a na i e species, i he wo species a e simila in niche use. In Finland, he dis ibu ion o he in asi e No h Ame ican bea e (Cas o canadensis)is app oaching he ange o he na i e Eu asian bea e (Cas o ibe ) possibly c ea ing a h ea o he la e . We compa ed he habi a use o he na i e and in asi e bea e species in Finland in he main dis ibu ion o he species and wi hin a smalle a ea whe e he species li e in sympa y. We compa ed he used habi a s ( olume o bi ch and o he deciduous ees and dis ance o ag icul u al and u ban a eas) a bea e lodges and a andom loca ions in he a ailable ipa ian habi a wi h (condi ional) logis ic eg ession models. Resul s indica ed ha he na i e bea e lodges we e loca ed close o ag icul u e han hose o he in asi e bea e . The olume o bi ch was also sligh ly g ea e nea he lodges o he na i e bea e han hose o he in asi e bea e . Howe e , habi a use o bo h o he species seemed qui e lexible, because he habi a nea lodges did no di e much om he a ailable habi a . We conclude ha he p obabili y ha he No h Ame ican bea e will in ade he dis ibu ion a ea o he Eu asian bea e in Finland depends, a leas pa ly, on he abili y o he o me o li e in p oximi y o ag icul u al a eas. Howe e , me hods o he han hose ela ed o managing habi a quali y may be he bes app oach o con olling he in asi e species. Keywo ds Alien species Cas o ibe Cas o canadensis Habi a use Moni o ing coun s Ci izen science In oduc ion Habi a equi emen s la gely de e mine he dis ibu- ion and abundance o species. Two e y simila species, which sha e he same ecological niche, seldom coexis in he same a ea o a long ime (K ebs 1972; Eh le ´n and Mo is 2015). A se ious p oblem can de elop when an alien species is in oduced o an a ea whe e a e y simila na i e species al eady exis s (Ebenha d 1988). In he wo s case, he in asi e species may ou -compe e he na i e one. In asi e species may also ansmi diseases o pa asi es o na i e ones (e.g. Kauhala 1996; Sainsbu y e al. 2000), o hey may al e he habi a s o he na i e species (Rowe and Gill 1985). They can hyb idize wi h na i e species (e.g. Ebenha d 1988; Geno esi e al. 2009), R. Alakoski (&)V. Selonen Depa men o Biology, Uni e si y o Tu ku, 20014 Tu ku, Finland e-mail: [email p o ec ed] K. Kauhala Na u al Resou ces Ins i u e Finland Luke, I a ¨inen Pi ka ¨ka u 4 A, 20520 Tu ku, Finland 123 Biol In asions h ps://doi.o g/10.1007/s10530-019-01919-9(0123456789().,- olV)(0123456789().,- olV) and in asi e p eda o s may ha e a de imen al e ec on na i e p ey animals (e.g. Ebenha d 1988; Kauhala 1996; Woods e al. 2003; Banks e al. 2008). An in asi e species po en ially h ea ening a na i e species is he No h Ame ican bea e (he ea e NA bea e , Cas o canadensis), which is pa ly sympa ic wi h he na i e Eu asian bea e (Cas o ibe )in Finland (Kauhala and Tu kia 2013; Kauhala and Ka inen 2018). The Eu asian bea e was hun ed o ex inc ion in Finland in 1868 (G ani 1900; Lah i 1972; Lah i and Helminen 1974) bu was ein oduced in he 1930s when 17–19 indi iduals we e b ough om No way (e.g. Lah i and Helminen 1969; Ha ¨ ko ¨nen 1999). Se en NA bea e s we e also in o- duced (Lah i and Helminen 1969,1980; E mala e al. 1989), because a he ime hei s a us as a dis inc species was no known. Bo h species we e eleased in o se e al a eas in Finland, bu he Eu asian bea e popula ion su i ed (only one pai ) and s a ed o inc ease slowly only in Sa akun a in sou hwes e n Finland, a egion whe e no NA bea e s we e in o- duced (Fig. 1). NA bea e s lou ished especially well in eas e n Finland (Fig. 1; E mala 1996). The ange and popula ion size o he NA bea e inc eased as e han ha o he Eu asian bea e , eaching an es ima ed popula ion size o [10 000 in 2017 (Kauhala and Ka inen 2018). The p esen ange o he NA bea e co e s mos o eas e n and cen al Finland, and spo adically Lapland, whe eas ha o he Eu asian bea e is es ic ed o a smalle a ea in wes e n Finland (mainly Sa akun a), and i s popula ion size is es i- ma ed o be 3300–4500 (Fig. 1; Kauhala and Ka inen 2018). The ange o he NA bea e has sp ead wes wa ds in ecen decades and pa ly o e laps he dis ibu ion o he Eu asian bea e in h ee a eas, i.e., in he egions o Pi kanmaa, Pohjanmaa and Lapland (Fig. 1; Kauhala and Tu kia 2013; Kauhala and Ka inen 2018; Iso ou u e al., unpubl. da a). I is possible ha he wo species ha e li ed close o each o he o some decades in Pi kanmaa, and a p esen , hey e en li e in he same i e sys ems and ha e, a leas on wo occasions, been ound nea he same lodge (Kauhala and Ka inen 2018). Because he Eu asian bea e did no su i e a e he o iginal in oduc ions in a eas whe e bo h bea e species we e p esen , i is assumed ha he sp ead o he NA bea e is a h ea o he Eu asian bea e (Liukko e al. 2016). The Eu asian bea e is classi ied as ‘nea h ea ened’ in Finland (Liukko e al. 2016). The NA bea e has la ge li e s, which may be he eason o a mo e apid g ow h a e o he popula ion and may gi e i he ad an age when he species mee (Pa ke e al. 2012). Bo h bea e species a e monogamous and e i o- ial (Wilsson 1971; Nole and Rosell 1994), and he ecological niches o hem a e ai ly simila . Bo h species eed mainly on deciduous ees (Collen and Gibson 2001; Danilo e al. 2011; Pa ke e al. 2017); and in summe , e es ial and aqua ic he bs a e also pa o hei die (Wilsson 1971; Lah i and Helminen 1974; Nole and Rosell 1994; Danilo e al. 2011). A ma u e bea e couple builds a lodge, a bank bu ow, o a combina ion o he wo, wi h an unde wa e en ance (Wilsson 1971; Lah i and Helminen 1974;Mu ¨lle - Schwa ze 2011). The c i ical wa e dep h needed o a lodge may be abou 40 cm (Rosell and Pa ke 1996; Baskin 2011); ha is, he lodges o he bea e s a e always by he wa e a ea. A bea e amily can occupy se e al lodges du ing he summe , bu only one lodge is used du ing he win e (Lah i and Helminen 1974). Bea e s a e cen al-place o age s (e.g. Haa be g and Rosell 2006), and hey commonly o age wi hin a 50-me e adius om he lodge, which can be de ined as hei co e a ea. Bea e s can, howe e , mo e up o 250 m om wa e in sea ch o good o aging ees (Smi h e al. 1994;Mu ¨lle -Schwa ze 2011). One amily g oup needs app oxima ely 1–2 km o sui - able habi a along a wa e cou se o sho eline (de ined as hei e i o y, Ha man 1994). Bea e s usually inhabi o es ed a eas, bu may also use ag icul u al a eas, especially a eas ha a e mosaics o ields and o es pa ches along a i e sys em. The ange o he Eu asian bea e in Finland co e s especially la ge ag icul u al a eas, and hey can make dams and bank bu ows in small di ches be ween wo ields and e en o age in ields (Kauhala and Ka inen 2018). We ha e also ecei ed obse a ions o bea e s in small owns o illages om he public, which indica es ha hey do no always a oid human p esence. We compa ed he habi a use o he wo bea e species wi hin hei co e a eas a ound lodges and on a la ge e i o y scale, and compa ed he habi a use wi h o he habi a s a ailable in he landscape (Johnson 1980). We used ci izen-science da a on bea e lodge loca ions in he main dis ibu ion a ea o he species in Finland. We p edic ed, based on ea lie s udies (e.g. Lah i and Helminen 1974; Ha man 1994; Nelne and Hood 2011), ha (1) he habi a use o he wo bea e 123 R. Alakoski e al. species would be ai ly simila and hey would a o habi a pa ches wi h deciduous o es s. Howe e , he e s ill may be a di e ence be ween he species in he amoun o bi ches and o he deciduous ees nea hei lodges, o example, due o di e ences in he amily s uc u e be ween he species (Pa ke e al. 2012). We u he p edic ed (2) ha he wo bea e species would be ound o di e in hei habi a use in ela ion o ag icul u al and u ban a eas, because he na i e bea e li es in an ag icul u e-domina ed landscape, whe eas he NA bea e li es in a o es -domina ed a ea wi h a spa se human popula ion. We discuss he possible ole o habi a equi emen s in he conse a ion o he Fig. 1 Map o he cu en dis ibu ion o bea e obse a ions in Finland and he success ul in oduc ion si es o he Eu asian and he No h Ame ican bea e , om whe e he species began o sp ead (unsuccess ul in oduc ion si es a e no included). The s udy a ea consis ed o he dis ibu ion a eas o bea e s, excep Lapland (g ay a ea in he map). Eu asian bea e s in Lapland ha e appa en ly dispe sed o Finland om Sweden. The zoomed pic u e is om he a ea in Pi kanmaa, whe e he species a e pa ly sympa ic 123 Di e ences in habi a use na i e Eu asian bea e and in con olling he in asi e NA bea e in Finland. Me hods S udy a ea The s udy a ea consis ed o he dis ibu ion a eas o bea e s in Finland, wi h he excep ion o Lapland (Fig. 1). The landscape in Finland consis s la gely o coni e ous and mixed o es s wi h app oxima ely 168 000 lakes wi h an a ea o a leas 500 m 2 (S a is ics Finland 2018a) and app oxima ely 20,000 km o i e s (Biodi e si y 2018). Fo es s co e app oxi- ma ely 75% o he land a ea and he Sco s pine (50%; Pinus syl es is), he No way sp uce (30%; Picea abies) and bi ches (17%; Be ula sp.) a e he mos common ee species (Luke 2018a). Ten pe cen o he land co e is composed o inland wa e s, 9% o ag icul u al a eas and 3.5% o u ban a eas, i.e. popula ion cen e s including esiden ial, indus ial and comme cial a eas (Biodi e si y 2018). The human popula ion densi y is highe whe e he main dis ibu- ion o he Eu asian bea e is, in Sa akun a in sou hwes e n Finland, wi h 28.18 ind./km 2 , han he human popula ion densi y o he a ea whe e he main dis ibu ion o he No h Ame ican bea e is, in eas e n Finland, wi h 3.66, 9.18, 10.32 and 14.71 ind./ km 2 in Kainuu, No h Ka elia, E ela ¨-Sa o and Pohjois-Sa o, espec i ely (S a is ics Finland 2018a). Ag icul u e is mo e dominan in Sa akun a wi h 16.7% o he a ea being ag icul u al land in 2013, compa ed o eas e n Finland wi h 1.4, 3.9, 3.8 and 7.2% being ag icul u al land in Kainuu, No h Ka elia, E ela ¨-Sa o and Pohjois-Sa o, espec i ely (Luke 2018b). C op cul i a ion is mo e common in Sa a- kun a compa ed o eas e n Finland, whe e ca le a ming is mo e common. Lakes a e mo e abundan in eas e n Finland (also called he Lake Dis ic ) wi h inland wa e s co e ing a minimum o 12.1% o he a ea o Kainuu o a maximum o 25.5% o E ela ¨-Sa o, when in Sa akun a he p opo ion is 5.3% (Ja ¨ iwiki 2018). Da a o bea e s Lodge si es we e ob ained om moni o ing coun s ca ied ou by he Finnish Game and Fishe ies Resea ch Ins i u e (Na u al Resou ces Ins i u e Fin- land, Luke since 1.1.2015). Bea e lodge coo dina es we e collec ed by hun e s du ing he all o 2013 and 2014 and he sp ing o 2015 (B omme e al. 2017; cases o a si e ha ing da a o mul iple yea s we e omi ed om he da a). The en i onmen al a iables used in he s udy we e he same o hese yea s. Hun e s usually know hei hun ing a eas well, and his me hod should supply a comp ehensi e p opo ion o lodges, bu ob iously no all lodges could be de ec ed wi h he ci izen science app oach. Only inhabi ed win e ing lodges we e epo ed o es ima e he numbe o bea e amily g oups in each game managemen dis ic . An occupied win e lodge can be ecognized by a ood cache nea he lodge (Mu ¨lle -Schwa ze 2011). The bea e species could no be iden i ied in he ield by a ci izen scien is collec ing he lodge- loca ion da a. Ins ead, he iden i ica ion o bea e species was based on he his o ic dis ibu ion o bea e s in Finland (Fig. 1), DNA analyses om wood chips collec ed nea bea e lodges and skull mo - phome y om hun ed bea e s (Kauhala and Timonen 2016). The coo dina es o 758 and 628 Eu asian bea e and No h Ame ican bea e lodges, espec i ely, we e analyzed in GIS (ESRI A cMap 10.2.2; ESRI 2011) wi h land co e a iables. The geog aphic in o ma ion o s eams 20 m (as polylines) and lakes and la ge i e s (as polygons) we e added om he da a o he Na ional Land Su ey o Finland ( opog aphic map 1:100,000) (Maanmi auslai os 2/2015), and lakes and la ge i e s we e ans o med o polylines ( om now on, all a e called wa e cou ses). We lands we e pa ly included in hese da a, as he e a e no la ge sepa a e we land a eas in he Finnish landscape (excep pea bogs). Only lodges ha we e wi hin 50 m om a wa e cou se based on GIS we e included o emo e possible e o s in coo dina es and o emo e cases whe e lodges we e in small s eams no digi ized in GIS. This was necessa y because he e i o y bu e s we e gene a ed along wa e cou ses (see below). Based on his, 328 lodges o he Eu asian bea e and 159 lodges o he No h Ame ican bea e we e excluded om he analysis. The Eu asian bea e li es in a mo e ag icul u e-domina ed landscape wi h small di ches, which is p obably why mo e o his species’ lodges we e in undigi ized wa e cou ses. We did no ha e a eason o suspec his emo al o he lodges wi h unce ain loca ions would a ec ou 123 R. Alakoski e al. analysis because he numbe o lodges o bo h species was s ill qui e high, and he Eu asian bea e was no o e ep esen ed in ela ion o he NA bea e . On he con a y, some quali y checks o he lodge loca ions p o ided by he ci izen scien is s should be done. Only win e ing lodges should ha e been epo ed, bu lodges we e some imes e y close o each o he . The e o e, lodges ha we e close han one km o ano he lodge along a wa e cou se we e coun ed as he same amily g oup’s lodges (Ha man 1994), because amily g oups’ home anges do no usually o e lap (Ko belo a ´e al. 2016). Only he cen e mos lodge in a amily g oup’s a ea was included in he analysis. In he end, we used coo dina es o 428 lodges o he Eu asian bea e and 466 lodges o he NA bea e ha we e sui able o gene a ing he bu e s. Habi a a iables in co e a eas, e i o ies and a ailable en i onmen The habi a a iables used in he analyses we e: he olume o bi ches, olume o o he deciduous ees, dis ance om u ban a eas and dis ance om ag icul- u al a eas. The olumes o bi ches and o he decid- uous ees (m 3 /ha) we e compu ed using he ee olume da a om he o es in en o y da a o he Finnish Fo es Resea ch Ins i u e in 2013 (Luke 2015). The da a gi es he olume o bi ches (Be ula pendula, B. pubescens and B. nana) and o he deciduous ees as one g oup, including he Eu asian aspen (Populus emula), alde (Alnus incana and A. glu inosa), Eu opean moun ain ash o owan (So bus aucupa ia) and he goa willow (Salix cap ea) (Yli alo 2013) wi hin 16 m 916 m squa es. The dis ances om he u ban a eas and ag icul u al a eas we e he nea es neighbo a i hme ic dis ance om he lodge/ andom poin . The ag icul u al a ea and u ban a ea we e added om he 2013 Co ine land co e da a o Finland wi h 20 m 920 m squa es (SYKE 2/2015). The ag icul- u al a ea included classes 2111–2441 (a able land, pas u es e c.). The u ban a ea included classes 1111–1424 (u ban ab ic, indus ial uni s, dump si es e c.) and 4122 (pea bogs in comme cial use). Roads we e no included as possible sou ces o dis u bance, because oads loca ed closes o lodges we e mainly he smalles oads wi h e y low a ic. Habi a a iables we e measu ed in wo scales: wi hin a co e a ea a ound lodges and wi hin a e i o y. Co e a eas we e calcula ed by bu e ing he lodges wi h a adius o 50 m (e.g. Mu ¨lle -Schwa ze 2011). We calcula ed he olumes o bi ches and o he deciduous ees pe hec a e wi hin he co e a ea and he dis ances om he lodge o he nea es ag icul u al and u ban a eas. In he second scale, he e i o y ex ended a maximum o 500 m om he co e a ea along all wa e cou ses connec ed o he co e a ea and 250 m om he sho eline o he lake o om he middle o he s eam, i.e., he maximum o aging dis ance on land acco ding o Mu ¨lle -Schwa ze (2011) (Fig. 2). Wi h a maximum dis ance o 500 m, each e i o y included sho eline (on bo h sides o he wa e cou se) wi h a mean leng h o 2050 m o he Eu asian bea e ( ange 484–5654 m) and 2438 m o he NA bea e ( ange 327–9746 m). We also calcu- la ed he olumes o bi ches and o he deciduous ees pe hec a e wi hin he e i o y. The nea es neighbo dis ances o he e i o y om he ag icul u al and u ban a eas we e calcula ed using he a e age a i h- me ic dis ance o 100 andom poin s placed wi hin he e i o y (Fig. 2). To compa e habi a a iables wi hin co e a eas and e i o ies wi h hose a ailable in he landscape in habi a s close o ipa ian zone, we laid andom poin s in he p oximi y o all wa e cou ses in he landscape. We used andom poin s ha we e a maximum o 250 m om a wa e cou se ( om now on called a wa e cou se a ea) and did no o e lap wi h a bea e e i o y. Random poin s we e laid wi hin each municipali y in Finland wi h bea e lodge coo dina es om he 2013 municipali y di ision om he da a o he Na ional Land Su ey (Maanmi auslai os 2/ 2015). Municipali ies we e selec ed as he a ailable landscape a eas because he sizes o he municipali ies a e o sui able size o he bea e s o po en ially dispe se ( he median size o a municipali y in Finland is 750 km 2 ). The numbe o andom poin s o each municipali y was compu ed by mul iplying he wa e - cou se a ea as hec a es (excluding bea e e i o ies) in a municipali y wi h he highes numbe o lodges ( wo species sepa a ely) pe wa e cou se a ea ound in all municipali ies. The numbe o andom poin s was hen mul iplied so ha hei o al numbe s we e app oxima ely en old compa ed o he numbe o lodges: 4452 o he Eu asian and 4874 o he NA bea e . We compu ed he olume o deciduous ees (bi ches and o he deciduous ees sepa a ely) pe hec a e a each andom poin (in e sec ing wi h a 16 m 916 m squa e wi h ee olume), and he 123 Di e ences in habi a use sho es a i hme ic dis ances be ween andom poin s and he nea es u ban and ag icul u al a eas. Da a om he sympa ic Pi kanmaa a ea A sepa a e compa ison was done o he bea e obse a ions wi hin he egion whe e bo h species a e sympa ic ( he egion o Pi kanmaa; Fig. 1). A o al o 50 coo dina es, 25 o bo h species, we e used o his analysis. Coo dina es included bea e lodges (15 Eu asian (E); 7 NA), eeding si es (4 NA), si es whe e d oppings we e ound (1 NA) and si es whe e bea e s we e sho (13 E; 10 NA; ecei ed om hun e s who sen us bea e skulls o species de e mina ion, Kauhala and Timonen 2016) ( om now on called ac i i y si es. Volumes o bi ch and o he deciduous ees wi hin 250 m om each bea e ac i i y si e and dis ances o ag icul u al and u ban a eas om 100 andom poin s in he = 250 m bu e we e calcu- la ed. Only he 250-m bu e ( e i o y scale) was selec ed o his analysis, because no all ac i i y si es desc ibed he accu a e loca ion o he lodge. Fig. 2 A GIS image o he lodge si e, he co e a ea and he e i o y compu ed along he wa e cou ses. Examples a e om wo Eu asian bea e lodges 123 R. Alakoski e al. S a is ical analysis Analysis o he habi a use in co e a eas and e i o ies o each bea e species We compa ed he co e a eas and he e i o y bu e s using condi ional logis ic eg ession analyses sepa- a ely o he wo bea e species (Duchesne e al. 2010). Wi h condi ional analyses, we could ma ch he co e a eas and e i o ies o each lodge o he analysis, ha is, he esponse a iable was he ma ched pai o used (co e a ea) and a ailable ( e i o y). Explana o y a iables we e habi a cha ac e is ics, i.e., olumes o bi ches and o he deciduous ees (m 3 /ha) and dis- ances o ag icul u al and u ban a eas (mo e abou condi ional analysis, see e.g. Duchesne e al. 2010). Fo a compa ison o co e a ea/ e i o y e sus andom poin s in he landscape, we could no o m ma ched pai s (condi ional model). Ins ead, we buil wo logis ic eg ession models, whe e he binomial esponse a iable was he used habi a (1) o he co e a ea o e i o y e sus a ailable habi a (0) in he landscape, ep esen ed by andom poin s along he sho e o all wa e cou ses in he municipali y. Explana- o y a iables we e habi a cha ac e is ics, i.e., ol- umes o bi ches and o he deciduous ees (m 3 /ha) and dis ances o ag icul u al and u ban a eas. In addi ion, he municipali y was included as a andom a iable in he model. Analysis o he di e ences in habi a use be ween he Eu asian bea e and he No h Ame ican bea e To analyze whe he he habi a use di e ed be ween he species, we combined he da a o bo h bea e s and pe o med models simila o hose desc ibed abo e (co e s. e i o y, co e s. landscape, e i o y s. landscape). Condi ional logis ic eg ession was used when compa ing he co e a ea o e i o y, and binomial logis ic eg ession when compa ing he co e a ea and e i o y o en i onmen . In addi ion, we included he in e ac ion e ms be ween he class a iable ‘species’ (1 Eu asian Bea e , 2 NA bea e ) and he habi a a iables in he models. The in e ac ion e ms we e included sepa a ely o each habi a a iable. Thus, we could es i habi a use di e ed be ween he Eu asian and he NA bea e . Because his analysis o he wise epea s he abo emen ioned analysis, we epo only he obse ed signi ican esul s o in e ac ion e ms. Analysis o he habi a use in he egion whe e species a e sympa ic Fo he compa ison o habi a use o he in asi e and na i e bea e s in an a ea whe e hey a e sympa ic (Pi kanmaa), we buil a model we e species (1 Eu asian bea e , 2 NA bea e ) was he esponse a iable. Explana o y a iables we e habi a cha ac- e is ics, i.e. olumes o bi ches and o he deciduous ees (m 3 /ha) and dis ances o ag icul u al and u ban a eas. In addi ion, ype o obse a ion was included as a class a iable in he model (ac i i y si es: 1 lodge, 2 o he ype o obse a ions). In his analysis, we assumed ha he a ailabili y o habi a s was simila o NA bea e s and Eu asian bea e s, because hey we e li ing in he same a ea. Resul s Habi a use in co e a eas and e i o ies Co e a eas o bo h species included s a is ically signi ican ly mo e bi ch han did e i o ies in he model (Table 1). The olume o o he deciduous ees was lowe in he co e a eas han in he e i o ies o he NA bea e (Table 1). When compa ing co e a eas and e i o ies o andom poin s a ailable in he ipa ian landscape, he only s a is ically signi ican di e ence was ha he Eu asian bea e ’s co e a eas and e i o ies we e close o ag icul u al a eas han andom poin s we e (Table 1). The co e a ea and he e i o y o he NA bea e did no di e signi ican ly om he landscape in any o he measu ed a iables (Table 1). Di e ences in habi a use be ween he Eu asian bea e and he No h Ame ican bea e The dis ance om ag icul u al a eas, compa ed o he a ailable en i onmen ( andom poin s) di e ed be ween he na i e and in asi e bea e (in e ac ion e m be ween bea e species and dis ance o ag icul- u e; co e: F 1,10109 = 4.88, p= 0.03; e i o y: F 1,10110 = 4.04, p= 0.04; Table 2); ha is, he mean dis ances o ag icul u al a eas we e g ea e in he NA 123 Di e ences in habi a use