Ecology in a human-dominated World - - - - 187 sk, Dept of sk, Dept ń L. across a spatial gra spatial a across L. Castor fiber Castor widerska and S. Saath, Univ. of Gda Univ. widerska and S. Saath, Ś widerska, Szymon Saath widerska, Szymon Saath Ś , spatial habitat structure, space dependent habitat selection, , spatial habitat structure, Castor fiber Castor 187–200 –––––––––––––––––––––––––––––––––––––––– Open Online article Authors. This is an © 2018 The Commons This is an open access articleCreative under the terms of the in any and reproduction License, which permits use, distribution Attribution cited. is properly the original work medium, provided habitat selection was affected by different scale-related phenomena related 1) to central phenomena scale-related by different habitat selection was affected clumped distribution of woody plants in the which resulted place foraging behaviour, Finally, local population density and woody plant cover. and 2) within the territory, the largest spatial scale studied (e.g. 3) habitat selection occurs independently across to the limited natal dispersal range of due watersheds), which was probably between the animals. Keywords: delta River Vistula data with 183 randomly selected sites not occupied by the species. The beavers pre the species. The beavers not occupied by data with 183 randomly selected sites plants, including shrubs, of woody habitats with high availability and avoided ferred Within a single family terri habitats, such as arable lands. modified anthropogenically a colony distance from with increasing observed woody plant cover we decreasing tory, of assessment the on depend preferences habitat beaver that suggests which centre, tested We habitat elements. of preferred both the abundance and spatial distribution the importance habitat selection with principal coordinates beaver of spatial scale in scale explained that the geographical showed of neighbour matrices analysis, which density explained in habitat composition, while the local beaver 46.7% of the variation related were that gradients spatial main two found We variability. this of 10.3% only related to the larg spatial scale: first, the most importantto the broad gradient was and the woody plant cover independent of and was sites beaver distances between est locally more The second most important gradient appeared site density. local beaver beaver indicate that European results Our and was associated with these variables. We evaluated habitat selection by European beaver beaver European by habitat selection evaluated We on the family territory) local (within scale. Based dient from ecoregional to a broad, territories along assessed the habitat composition of 150 beaver we aerial photography, these and compared delta (northern River Poland) Vistula the main water bodies of the 42 sk, Poland. – A. Weydmann, Univ. of Gdańsk, Inst. of Oceanography, Gdynia, Poland. AZ and AW also at: AZB AZ and AW Gdynia, Poland. of Oceanography, of Gdańsk, Inst. Univ. Weydmann, – A. sk, Poland. ń

Jens-Christian Svenning Svenning Jens-Christian 2018 8 August Accepted doi: 10.1111/ecog.03621 A. Kelt Douglas Editor: Subject Editor-in-Chief: Ecography 42: 187–200, 2019 analysis and software, Gdynia, Poland. – R. Pudełko, Inst. of Soil Science and Plant Cultivation State Research Inst., Dept of Bioeconomy and Systems and Systems of Bioeconomy Dept Inst., Research State Cultivation and Plant of Soil Science Inst. – R. Pudełko, Gdynia, Poland. analysis and software, Poland. Puławy, Analysis, A. Zwolicki (http://orcid.org/0000-0003-2710-681X) ([email protected]), K. Moskal, J. K. Moskal, ([email protected]), (http://orcid.org/0000-0003-2710-681X) A. Zwolicki Ecology Gda and Zoology, Vertebrate Adrian Zwolicki, Rafał Pudełko, Katarzyna Moskal, Joanna Rafał Pudełko, Adrian Zwolicki, and Agata Weydmann by European beaver European by The importance of spatial scale in habitat selection importance of spatial The Research www.ecography.org ECOGRAPHY doi: 10.1111/ecog.03621 Ecology in a human-dominated World after whichitwasdriven totheverge ofextinction by rodent inEurope and wascommonuntiltheMiddle Ages, (Fryxell 2001,Campbell et al. 2005). andreproductionconsequences forthesurvival ofacolony and has long-lasting which asettlementdecisioniscrucial, lived and territorial herbivores, such asbeavers role,Habitat especiallyinlong- selection playsanimportant time lost toreproduction duringthisprocess (Morris 1987). linked toemigrationand,hence,canbemodelledas the 2013). At themacrohabitat orlandscapescale,thecostsare for exploiting the habitat or potentially better ones (Cassini optimal microhabitat patches,whenthistimecouldbeused to aforagingstrategyandtimespentmoving through sub within thescaleofhabitatpatches,animalcostsare linked 1980,Cassini2013).Duringtial scale(Johnson selection withspa levels, andthecostsofhabitatselectionalsovary 1 km (Schaefer et al.2008). responded totheirenvironment atthescaleofapproximately while red deer 2009b), (Mayor et al. most evidentatdistancesupto15 km bou mobility andthesensesemployed, suchassight.Hence, cari tion occurs varies among species and is influenced by their 2015, Dupke et al. 2017).Thespatialscale over whichselec 2009b, Mancinelli et al. vegetation phenology(Mayor et al. herbivorous animalsismainlyrelated tofoodavailability and tribution (Morris 1992).The valuation oflocal resources in individuals, which are often smaller than a population dis capabilitiesandbyown thedispersaldistancesof sensory environmental features orqualityislimitedbothby their Moreover, thearea over whichanyindividualisabletoassess (Fretwell andLucas 1969,Rosenzweig 1981,Morris 2003). avoidance, aswell as intra-andinter-specificcompetition tion isalsomodified by bioticinteractionssuchaspredator Davidson 2000,Cassini2013).Theprocess ofhabitatselec distribution of resources often changes spatially (Morris and environmental conditionsare scale-dependentbecausethe (Morris 2003).Additionally, preferences towards certain availability ofresting andhidingplaces,orbreeding areas cies, including food quality and availability, but also the of resources mostlyrelated totheforagingecologyofaspe tion mustrely onaqualitative assessmentofdifferent types 2009a, Cassini2013). al. et theme ofhabitatselection(Schneider2001,Mayor scale-dependent, includinganimalbehaviourandtherelated increasingly clearthatnumerous ecologicalphenomenaare metrics oftenchangeswithdifferent geographicalscales,itis 1992). Rooted intheconceptthatvariation inecological 1989,Steelesince the1980sand‘90s(Wiens 1991,Levin ingly appreciated, andhasbeenquantitatively expressed ofspatialscaleinecologyhasbeenincreasThe importance Introduction 188 The European beaver Habitat selection at broad scales constrains that at lower All animalsconsumeresources. Therefore, habitatselec Rangifer tarandusRangifer Cervus elaphus Cervus habitatselectioninNewfoundland was Castor fiber inthePyrenees Mountains L.isthelargestnative Castor , for ------attractive territories(Nolet andRosell 1995, 1994,Hartman density graduallyincreases, forcing beavers tooccupyless- be selected.Aspreferred habitats are occupied,population tions are relatively sparseandoptimalhabitatsare thefirst to Pinto et al. 2009,John et al. 2010,Swinnen et al. 2017). ent stagesofapopulationdevelopment 1995, (Hartman for characterizingtheprocess ofhabitatselectionindiffer al.2012),makingthemausefulmodelspecies et (Rosell beavers have rapidly colonized their historical distribution In of absence, recent decades, after nearly a half-century capacity(Bartak et al.population approaches carrying 2013). lation growth patternwithacharacteristicdampeningasthe 2009). In thedispersalphase,beavers follow alogisticpopu or freshly cuttrees (Nolet andRosell 1998, John andKostkan leaving visiblemarks oftheiroccupancy, suchaslodges,dams, the speciesdispersesandoccupieshabitatalongwatercourses, zation by beavers inEurope isrelatively well documented,as 1986, Dzi been successfullyreintroduced ( ulations were fullyextirpatedby 1945,buttheyhave since over-hunting forfurandcastoreum. In Poland, beaver pop (IDD; Fretwell andLucas (1969). beavers meetthepredictions ofanidealdespoticmodel must considerexistingterritories. Consequently, European tat selectivityare interrelated because asettlementdecision 2001).Moreover,tion (Fustec et al. territoriality andhabi plants onriverbanks isneededtomaintainabeaver popula cover, whichimplies thatextensive growth ofpreferred tat quality, and family home ranges depend on vegetation sity ofabeaver populationisinfluencedlargely by habi parental colony (Collen andGibson 2001).The localden beavers fail toestablish territories, they may return to the 2017),oriftheseyoung to theageof7yr(Mayer et al. areas,grate tonew althoughtheymaydelaydispersalup 2006).Older young (2–3yr)generallyemi (Rosell et al. the previous year, andanykitsborninthecurrent year which consistsofamonogamousadultpair, yearlings born tect resources. They normally live as a family unit (colony), cover withinfamilyterritories. quality couldberelated ofwoodyplant totheproportion and Busher 1979, Tyurnin 1983). Therefore, beaver habitat consequently, (Jenkins theyare forwintersurvival crucial growth, woodyplantsare oftentheonlyfood source, and 2009).Outside theperiodofvegetative 2009, Pinto et al. al.2001,John andKostkan et al.1994,Fustec et (Nolet this is mostly represented by willow-scrub communities uous woody plants, and in European riparian habitats, foodsources, which,forbeavers,particular comprisedecid even years after a few of colonization (Campbell et al. 2005). as earlyarrivals occupylargerandhigher-qualityterritories, role indeterminingthesizeimportant ofthefamilyterritory, of parental pairsintounoccupiedareas islikelytoplayan 2010).Thesequenceofarrival 2001,John et al. Fustec et al. During theinitialphasesofrecolonization, beaver popula Beavers actively mark anddefendtheirterritoriestopro Habitat selectionistheresult ofthepreference for ę ciołowski andGozdziewski 1999).The recoloni Ż urowski andKasperczyk ------Ecology in a human-dominated World

2 ------189 uławy Fens uławy Fens Ż and has an aver 2 ledziowy, Linawa, and Linawa, and ledziowy, Ś no Reserve (a), and an example of no Reserve ż uławy Fens system, we also studied the we system, uławy Fens Ż ski channels. ń no Reserve, located on the east side of the Vistula Vistula of the no Reserve, located on the east side ż Within the Within related to fluvial processes and climate similarities, results in results in and climate similarities, to fluvial processes related 2013). 2000, Cassini (Kondracki homogeneous vegetation Vistula in the all main watercourses covers study area Our a mosaic of habitat qualities, from delta and includes River to highly modi River, Vistula such as the semi-natural areas Piaskowy, the including areas, fied Panie Dru and mires extensive by surrounded and 1a) (Fig. delta River While the landforms and the canal-system of swamp-forest. Material and methods Material area Study across in a northerly direction flows River Vistula The the largest unregulated is 1000 km. It for over Poland the river Sea and at its outlet to the Baltic in Europe, river or Delta Vistula called the forms a wide delta km of ca 2320 an area 1), which covers (Fig. density of 145 inhabitants per km age human population is rela Delta Vistula the (Pruszak et al. 2005). Geologically, rela by is covered the flat area (ca 6000 yr old); young tively is located between homogenous alluvial soils, and it tively sandbars to the and to the east and west hills of glacial origin (30%) is located below north. a large part Because area of the channels, ditches, and pump systems of rivers, sea level, the past over been constructeding stations have in the area and channels in the delta area rivers centuries. The several forming a network directions, in different and flow spread and generally shallow which are of multiple watercourses very The delta has homogenous geomorphol slow-flowing. genesis the with along that, structure soil and geology, ogy, ------Kuhl. John and Kostkan and Kostkan John Kuhl. C. canadensis The goal of our study was to assess the importanceThe goal of our study was to assess of Habitat selection is a hierarchical process based on behav based process is a hierarchical selection Habitat Figure 1. Beaver territories and randomly distributed sites in the Vistula River delta, including the Dru River Vistula territories and randomly distributed sites in the Beaver Figure 1. (b). Nogat River point located in the centre buffer designation as applied to orthophotomaps a beaver around enon, we selected the Vistula River delta, which is an anthro delta, which River Vistula selected the enon, we recently that was quite pogenically modified environment beavers. European by recolonized lation density and differences in habitat quality between in habitat quality between lation density and differences limitations (2nd distant water bodies or animal dispersion place to study this phenom selection). As a suitable order might be determined by the central place foraging strategy place foraging the central determined by might be geographi scale), to a broad selection of Johnson’s order (3rd scale, which could depend on the local popu cal ecoregion spatial scale in habitat selection by European beaver across across beaver European spatial scale in habitat selection by mostly to the a local scale related a gradient of scales, from which territory, distribution of woody plants within a family habitats in 2nd-order selection, and did not test beaver pref selection, and did not test beaver habitats in 2nd-order distribution of certain to the spatial resources relative erences within a territory. (2009), in the Morava River, described 2nd and 3rd-order 2nd and 3rd-order described River, (2009), in the Morava non-random. were habitat preferences selection, in which between not describe the spatial variability they did However, and 3rd order of Johnson’s scale (Beier and Barrett 1987, 1987, and Barrett scale (Beier of Johnson’s order and 3rd Hartman 1996, Fustec et Campbell al. 2001, et 2005, al. Pinto et al. 2009, John et al. for both the European 2010) and Canadian beaver for particular (4th items within these habitat patches food The importance and abiotic factors of both biotic order). 2nd selection mostly across for habitat has been recognized distribution of a species (1st order) to the home range or ter the home range or to of a species (1st order) distribution ritory patches within of habitat the utilization order), (2nd and the preferences order), (3rd these home ranges/territories ioural decisions governed by the assessment or usage of assessment or usage the by governed ioural decisions four described (1980) Johnson scales. different at resources the geographical from which range of this hierarchy, orders Ecology in a human-dominated World den, and4)geographicalcentres offeedingremains (e.g. 1)winterfoodsupply,importance): 2)beaver lodge,3)main on thekeysignsoftheiractivity, whichare (inthisorder of The presence ofbeavers inagiven area canbedetectedbased Beaver colony status (Simunkova and Vorel 2015). sites) and treated as the central points of familyhome ranges as evidenceofrecently occupiedterritories(colonies,beaver andscentmounds, weretion basedonobservations regarded caches. Thelattertwosigns,alongwithadditionalinforma stripping, dams, burrows, building of lodges, and winter food mounds, canals,feedingstations,cuttrees andbranches,bark receivers (accuracy up to60mfrom thewaterwere registered by Garmin GPS Donkor andFryxell 1999). over thisdistancefrom 1980,Fryxell water(Jenkins 1992, a sharpdeclineintheactivitiesofthesesemiaquaticanimals dicular totheshoreline; previous studieshave documented the waterbodiesandtheirsurroundings upto60mperpen banks andalongwatercourses from acanoe. We explored checked forevidenceofbeaver activity, mainlyfrom the Supplementary materialinAppendix 1fordetails),were Reserve (totalshoreline lengthof794.51km,seethe delta (rivers, channels,andponds),aswell astheDru which covered allmainwaterbodiesofthe Vistula River piles, and scent mounds, are clearly visible. The studyarea, traces oftheirpresence, suchastree cuts,girdles, food ver activityinlateautumn,winter, andearlyspring, when dates were chosenaccording totheseasonsofhighbea Nov 2011–Apr 2012andOct 2012–Apr 2013.The Fieldwork wascarriedoutduringtwoconsecutive seasons: Fieldwork 70 km southofthestudyarea (Aszyk1994). Wda River basin,which flows intothe Vistula approximately troduced. Thenearest introduction occurred in1979the rivers ofthe Vistula River basin,where beavers hadbeenrein source ofthispopulationwasnaturaldispersionfrom other was found in 2002 (Ciechanowski et al. 2013). One potential the Dru Delta (AszykandKistowski 2002).Thefirstsignsofbeaver in ver familieswere reported ofthe inthewestern part Vistula began in the mid-90s (Gromadzka 1996). In 2001, four bea natural habitat. a suitablelocationtostudybeaver behaviourinanoptimal, (PLH280028 andPLB280013,respectively), whichmakes it under both theHabitats Directive and theBirds Directive Union network of protected areas Natura 2000 and listed tected nature thatislocatedwithintheEuropean reserve it similar to the watercourses of the Vistula Delta, it is a pro Dru 190 The geographicpositions ofallsignsbeaver activity The recolonization ofbeavers inthe Vistula Delta likely ż no Reserve(thelakeandthesurrounding canals)make ż no Reserve were notedin2001,andthefirstlodge ~ 5m); thisincludedtracks,faeces,scent ż no no ------supply (key sign no. 1), a family lodge (key sign no. 2) was such that,forexample, in the absence of any winter food ence of key signs, according to above order of importance, of digital maps. The colony centre was defined bythe pres marking mounds,andalltypesoffeedingsignswiththeuse a visualanalysisofthedistributionkeysigns,scent during tracking.Thiswasfollowed by verification basedon In thefield,potentialboundaries were defined preliminarily to determineacentre andtheborders ofafamilyterritory. other fresh signs(mostlyslidesandindicationsoffeeding). with fresh signsofactivity)andusuallycontainednumerous (most frequently awinterfoodsupplyand/orfamilylodge those inwhichatleastoneofthefresh keysignswaspresent ondatafrom selectedbeaverperformed sites,characterized as functional status.Allanalysespresented inthisstudywere beaver activitybutwithoutanykeysigns;and3)ofacertain signsof from new othersignsandfamilyterritorieswithafew the presence clearlyseparated ofoldsignsonly;2)uncertain, ity, colonieswere assignedtothree types:1)abandoned,with Based onthepresence ofoldandfresh signsofbeaver activ heaps oftree brancheslocatedontheshoreline). orshrub Dru covered theselectedwatercourses andtheshores ofLake and mosaickingofthe photos totheorthophotomap, which during image triggering enabled semi-automatic calibration 0.2 and0.7m.Maintenance opticalaxis ofthevertical the ground resolution oftheresulting photoswasbetween era inthevisibleband.Due totheavailable flightaltitude, photography wastakenwithaFujifilm IS-1digitalcam 172 aircraft (flightstakenon10and11 Sept 2012). Aerial gyro-stabilized photographic platformattachedtoaCessna study area, we tookaseriesofaerialphotographsusing habitat preferences, andevaluate foodresources over the To documentthecurrent stateoflanduse,assessbeaver Aerial photography GIS analyses defined asacentre point,etc. tographs. An analysis of the accuracy of the orthophotomap tographs. Ananalysisofthe accuracy oftheorthophotomap locations inthefieldthat were readily identifiedinaerialpho ground by measuring,withaGPSreceiver, theposition of Image calibrationreference pointswere determined onthe the photographsdidnotrequire adigitalelevation model. phy andthealmostcompletelyflatterrain,calibration of to thealtitude(1500–2000ma.g.l.)ofaerialphotogra pixel resolution wassetat0.5m.Due oftheorthophotomap Mapping usedthePUWG 1992coordinate system,andthe transformation methodwithnearest-neighbour resampling. the Raster/Georeferencer module. We appliedapolynomial 1.8.0 release). Selected aerialphotos were calibratedusing beaver habitat were inQuantum GIS(Lisboa performed Digital imageinterpretation and spatial analysisofthe Data processing An important stepindistinguishingthebeaver siteswas An important ż no, aswell asabuffer zone of60m. - - - - - Ecology in a human-dominated World - - - 191 species decreased rapidly up to 500 m from m from rapidly up to 500 species decreased Salix A detrended correspondence analysis (DCA) was used to correspondence A detrended To test 3rd-order selection, which describes the local scale test 3rd-order To In addition, to investigate the spatial land use structure land use structure the spatial investigate addition, to In dom sites, as indicated by the SIMPER analysis performed the SIMPER sites, as indicated by dom A2). Table 1 Appendix material (Supplementary beforehand gradients in the data environmental the theoretical explore A3) and: 1) to Table 1 material Appendix (Supplementary and random sites, the beaver between illustrate variability in composition use land the in variability the calculate to 2) and random sites (Anderson et al. 2008). SIMPER (similarity 2008). SIMPER and random sites (Anderson et al. run analysis was Bray- percentages) to decompose the average at beaver each land use type dissimilarities between Curtis 1993). The analysis operates at the and random sites (Clarke in a in our case, resulting, variable, dependent of each level of the groups between of mean dissimilarity value percentage us to quantify the This allows each land use type separately. a certain contribution of percent land use type to differences and random sites, and thus, to identify the beaver between Land use differences. the likely major contributors to these per transformed and standardized square-root class data were 2006). and Gorley sample (Clarke selection of land use composition pattern within the 2nd-order applied sites and randomly selected sites, we beaver between for all (rmANOVA) analysis of variance measures a repeated and 201–500 multiple ring buffers (0–100, 101–200 m). in the land use class data, the number of zeros reduce To of sizes area due to the different which could bias the results cover land we combined the following buffers, the repeated as trees as well classes into higher categories: shrubs, forest, combined with the were and shrubs (categories 3 to 5 above) grassland, fallow land, arable while plants category, woody (categories 1, 2, 6 and 7, respectively) land and urban area The group category. combined with the anthropogenic were major as understood preferences, beaver on based was ing ran and beaver the between differences to contributors study in a similar riparian habitat, the buffer included the the buffer similar riparian habitat, study in a the partmost exploited where home range, the beaver of usage of 1994). the den (Nolet et al. ring distinguished multiple we 0–500 m buffers, within the that zones concentric of successive buffers (nested polygons both in m 201–500 and 101–200 0–100, of overlap) not do and only measured we all cases In and random areas. beaver 60 m linear buffer along land use data within the compared are by this method The buffer ranges extracted the shoreline. 1b. in Fig. presented Statistical analyses selec (2nd-order preferences habitat beaver general test To of all land use types the composition compared tion), we and randomly selected beavers in territories occupied by permutational multivariate a using unoccupied territories with 999 permutations) (PERMANOVA analysis of variance 0–100, 101–200, design, where measures with a repeated sites beaver tested jointly between and 201–500 m rings were ------500 m from the nearest beaver beaver the nearest 500 m from 1%. We distinguished eight land distinguished We 1%. > ≤ To compare areas occupied by beavers with those not with those beavers occupied by areas compare To Because beavers exploit habitats only in the immedi exploit habitats beavers Because 2001, Fustec et al. 2003). As such, in many cases this buffer to a according However, territory. the entire did not cover based on previous findings that the average beaver territory beaver findings that the average based on previous 2 km (hence, of shore 1 km of river included approximately line) (Hartman 1994, Rosell et al. and Hovde 1998, Rosell nated a buffer of 0–500 m around the centres of both the the centres nated a buffer of 0–500 m around to points; selected randomly 183 the and territories beaver within the 60 m buffer these include only the area reiterate, of this buffer was 1b). The size (Fig. adjacent to the shoreline that were used for comparison with occupied territories. used for comparison with that were classes within the designated land use by assess coverage To desig we areas, selected randomly in and territories beaver inhabited, a series of random points were selected along the inhabited, a series of random points were and studied watercourses 1a) (Fig. resulted in the selection of 183 areas This territory. (Cassini 2013). Therefore, in our study, habitat composition in our study, (Cassini 2013). Therefore, land use patches in the proportionis defined as different of the 60 m linear buffer along a shoreline. small canals, and tributaries of other watercourses. The land small canals, and tributaries of other watercourses. habi as described be could distinguished were that types use their surroundings tat components or patches differing from oped tracts, and unused parts and urban par of agricultural by any type of covered – all artificial areas area’ cels); 7) ‘urban land roads, buildings, paved infrastructure: anthropogenic riverbeds, – old water bodies’ ‘small 8) fills, and fenced areas; covered mostly by natural vegetation (this class also covers (this class also covers natural vegetation mostly by covered difficult to define as other land use patterns, that are areas roads, paths, undevel including flood embankments, ground was registered as a point layer; thus, it was possible to record thus, it was possible to record as a point layer; was registered in all in detail all objects important food resources to beaver – non-shrub area land’ ‘fallow defined classes (polygons)); 6) shrub clusters; 4) ‘forest’ – detached complexes where trees trees where complexes – detached shrub clusters; 4) ‘forest’ small clus and – single trees and shrubs’ dominated; 5) ‘trees the larger complexes located far from ters of shrubs that were (this latter class classified as class 3 or 4 in polygon format often ploughed, which further facilitated the recognition of often ploughed, which further the recognition facilitated – all cadastral agricultural this land use class); 2) ‘grassland’ – large 3) ‘shrubs’ and pastures; used as meadows parcels their actual surfaces were their actual surfaces were (cadas fields arable cultivated all – land’ ‘arable 1) classes: use most were fields during the survey the tral parcels; (Sept), process was conducted as precisely as possible with a special was conducted as precisely process The quality land use classes. vegetation focus on separating the that performed showed fragments selected control 60 on and polygons surfaces the digitized the between of differences Fig. 1b), similar to that employed during fieldwork. Based on Based fieldwork. during that employed to similar 1b), Fig. a comprehensive within this buffer, the aerial photography digitalisation. This manual by was prepared map land cover ate vicinity of water, all further analyses were restricted to a restricted further all were analyses of water, ate vicinity studied watercourses m buffer along the linear 60 (Jenkins 2009; and Kostkan 1999, John and Fryxell 1980, Donkor was carried out on an independent sample of ground control control sample of ground out on an independent was carried less of (RMSE) error square mean root a showed and points than 5 m. Ecology in a human-dominated World model, thiswaytheefficiency valuesofeach variable were to compare bothCCAand VP withthesameunconstrained eigenvalues. Also,we usedfouraxes inallpresented models axes intheordination modelswasdefinedbasedontheir of theDCA(terBraak andŠmilauer 2012).Thenumberof environmental factorby thatexplainedby thefirstfouraxes ing thepercentage ofthevariability explainedby agiven explained non-random variability in the data (%) by divid variables from theconstrainedordination models (CCA, VP) 2002). sites (Økland 1994, Borcard and Eilertsen and Legendre calculated previously by PCNM, and the density of beaver of datavariation betweenpartitioning the spatial variable, canonical correspondence analyses(CCA),resulting inthe strained ordination, which inthisstudyfollowed unimodal independent variables. VP couldbecalculatedbasedoncon titioned) fractionofvariation indata,explainedby different This way, itispossibletoassess theuniqueandshared (par ables inrelation tothesetofmultivariate response variables. percent ofexplainedvariation) ofdifferent independent vari powerattempts toresolve theexplanatory (describedby the material Appendix 1 Table A5). Variation partitioning tribution andonthedensityofbeaver sites(Supplementary composition amongbeaver sitesdependsontheirspatialdis totesthow analysis(VP) variation inhabitat partitioning independent variables inconstrainedordination analyses. eigenfunctions),whichmaybeused as variables (orthogonal are setofindependentspatial thendecomposedintoanew culated from spatialdata(e.g.latitudeandlongitudevalues), which are presented astheEuclidean distancematrixand cal sites, especiallytheneighbouringones.Thesedistances, variables are usedtodeterminedistancesbetween studied 2009).In thePCNMmethod,spatial ation (Legendre et al. or fine)onthemodelspeciescommunityunderconsider significance ofdifferent spatialscales(broad, intermediate, eigenfunctions), italsoprovides toassessthe anopportunity 2002). Based ontheorder oftheprincipalcoordinates (PCO, ables orfrom communityprocesses (Borcard andLegendre nates eitherfrom thephysicalforcing ofenvironmental vari to quantifythespatialstructure ofecologicaldatathatorigi applied toasetofsiteswithgoodsamplingarea coverage as Moran’s Eigenvector Maps (MEM).Thismethodcanbe pal coordinates ofneighbourmatrices(PCNM),alsoknown 2010) and marine systems (Weydmann et al. 2014) is princi al. 2009, Peres-Neto and Legendre et terrestrial (Legendre developed methodthathasbeenappliedsuccessfullytoboth the influenceofscaleonspeciesorcommunities.A recently research, anumberofstudieshave assessedandquantified Šmilauer 2012). land useclasseswithinthe0–500mbuffer(ter Braak and Appendix 1 Table A4).BothDCAmodelswere basedonall efficiency ofconstrainedmodels material (Supplementary the beaver sitesonly, whichwasthenusedtocalculatethe 192 We calculated the efficiency with which environmental Therefore, we used thePCNMmethodwith variation Due ofspatialscaleinecological totheimportance ------STATISTICA-Features in STATISTICA ver. 13.1( UK), whileSpearman’s rankcorrelation andrmANOVA PRIMER ver. 6.0(Plymouth Marine Laboratory, Plymouth, to findthebestfitofmodel. models (GAMs)withtheAkaikeinformationcriterion(AIC) present theserelationships, we employed generalized additive a colonycentre were usedasthegeographicallocation. To neighbouring colonies,whilethelatitudeandlongitudeof oftheoverlappingportion area ofthe500 mbuffersbetween rank correlation. Thelocaldensitywascalculatedasthepro plants andanthropogenic Spearman’s landuse,we performed areas of neighbouring beaver colonies) and cover of woody beaver sites(forthe0–500mbufferbetween the overlapping the familywisetypeIerrors. comparisons, we usedHolm’s (1979)correction tocontrol with theMonte Carlotest(999permutations). For multiple data. The verification ofstatisticalhypotheseswasperformed log-transformation [ coherent between themodels.In allordination techniques,a composition oflandcover (habitattypes)withincreasing beaver sitesandrandomdiffered significantly inthe 101–200 m,and201–500m rings;p the beaver vsrandomsitesinallranges (pairs inthe0–100m, post hoccomparisonsshowed significantdifferences between nested in beavers vs random factor, p PERMANOVA repeated measures analysis(IDofsites inhabited by beavers (n beaver sites (n Significant differences inland usecomposition between Habitat selection found inthechannels(0.05–0.07colonieskm (0.17) and Wisła Królewiecka (0.16).Thelowest densitywas the canals),followed by therivers: Szkarpawa(0.18), Nogat colonies km River andthepondswithinitsfloodplains(0.410.87 highest beaver densitieswere alsorecorded forthe Vistula and Dru number ofterritorieswere notedinthe Vistula River (n (Supplementary materialAppendix 1 Table A1).Thegreatest 150 recently occupiedterritoriesinthe Vistula River Delta Based on10540signs ofbeaver activity, we distinguished Density anddistributionofbeaver sites Results Canoco ver. 5.10(terBraak andŠmilauer 2012). VP withPCNM,DCA,CCAandGAMswere in performed lowest numberwasfoundinchannels(Linawa(3),Panie PERMANOVA and SIMPER analyses were in run To determinetherelationship between localdensityof ż Ś no Reserve(30),andtheNogat River (25),whilethe ledziowy (2each),and Czarna Łacha(1);Fig. 1a).The –1 , respectively), Dru =

150) andrandomlydistributedsitesnot x’

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183) were revealed based on the + < 1)] wasusedtonormalize the www.statsoft.com/Products/ ż no Reserve (0.39,0.86on = =

0.001; Table 1a). The 0.001). Additionally, 0.001). –1 ). = 49), ń ski - Ecology in a human-dominated World

------193 – – p p 0.001 0.001 0.001 0.001 0.001 0.004 0.001

< < < < – – F 3.5 6.4 13.3 62.7 46.6 10.4 17.7 Pseudo-F no and the Vistula and Vistula the and no ż – MS MS 601.5 4891 3570.8 2125.8 3819.8 78926 56560.9 67027.0 3.02) the beaver territories were rela territories were 3.02) the beaver

= SS SS distance in Table 1b). Table distance in 4251.5 9782 7141.6 × 67027.0 78926 56560.9 395790.0 1264400.0 1745300.0 3.12, Axis2 = 1 2 1 2 1 2 df df 331 658 994 The above results suggest that there were other factors, such as were results suggest that there The above distant water between and differences spatial autocorrelation Dru Lake between (especially bodies systems), that influenced both habitat selection river Nogat variation geographical 3b). This and habitat composition (Fig. density (described was tested in comparison with local beaver variation by of the 0–500 m buffer) overlap as the percentage partitioning(VP) based was which analysis, PCNM the with revealed a significantly higher average cover of woody plants, cover a significantly higher average revealed lands, in all rings within of anthropogenic cover and lower with 2), Fig. 1b, (Table sites random to relative sites beaver in highest values the the proportion plants reaching of woody in 101–200 and 201–500 m the 0–100 m ring and decreasing proportion of anthropo 2a), while the average rings (Fig. the distance from with increasing genic land use increased was no such 2b). There territory of the beaver centre (Fig. distribu the where sites, random the of case the in gradient the similar in remained types tion of the tested land cover which indicates that 0–100, 101–200 and 201–500 m rings, were sites and random areas beaver between the differences spatial distributions of the different by additionally increased interac land use types (see ANOVA and preferred avoided tion: beaver density on a large and beaver Habitat variability spatial scale the 0–500 Comparison of the habitat composition between territories and those in ran m buffers established in beaver these two cat between substantial overlap dom sites showed space, indicating that they were egories in DCA ordination 1 material Appendix 3a, Supplementary quite similar (Fig. to the gradient length according the same time, A3). At Table (Axis1 types. use land of composition the regarding diverse tively shrubs) or avoided anthropogenic lands (arable land, grass shrubs) anthropogenic or avoided to analyse the distribution of land, urban area) land, fallow buffers. The rmANOVA beaver classes within these broader - - - - -

distance* distance* distance* × × × ANOVA, Factor/design ANOVA, PERMANOVA, Factor/design PERMANOVA, Beaver vs random vs random Beaver Beaver vs random Beaver Beaver vs random Beaver vs random Beaver vs random) ID of sites nested in (beaver Residuals (unexplained) Total vs random Beaver vs random Beaver Detailed comparisons of all distinguished land cover types all distinguished land cover comparisons of Detailed Based on the SIMPER results, we grouped land use types grouped we results, on the SIMPER Based and trees woody plants (shrubs, forest, as either preferred Spatial distribution of resources within beaver within beaver Spatial distribution of resources territories family five times greater average cover of this land use type in all cover average times greater five than sites beaver in lower were area Urban rings. nested three mate rings (Supplementary in random site in all compared A2). Table 1 rial Appendix larity between beaver sites and randomly distributed sites was sites and randomly distributed beaver larity between for small water bodies (up to 3.1% in the 0–100 m revealed at least by characterized sites were ring), although the beaver and shrubs representing the greatest proportion and having the greatest and shrubs representing 20.7% reaching the highest contribution to the dissimilarity, contribution to the dissimi in the 0–100 m ring. The lowest random 28.1% in 0–100 m). Generally, beaver sites showed sites showed beaver random 28.1% in 0–100 m). Generally, all types of woody plant land cover for higher mean values vs 0.1% and 0.4% and shrubs vs 3.0%, forest 16.2% (trees shrubs 0.9% vs 0.3%; all in the 0–100 m range), with trees between respective rings (21.2%) with higher proportions in respective between sites, apart the ring clos the outer two rings in beaver from 26.2% vs territories (beaver of the beaver est to the centres observed dissimilarity in the 0–100 m ring. Fallow lands had observed in the 0–100 m ring. Fallow dissimilarity up to 28.1% in all ranges, reaching cover the highest average highest dissimilarity in the 0–100 m ring, and had the second distance from the colony centre (101–200 m, 55.7% and (101–200 m, centre the colony distance from the most differentiat all distances, 51.6%). For 201–500 m, land, which dominated the type was Arable ing land cover the of 23.6% to up reached and areas distributed randomly material Appendix 1 Table A2). The average dissimilarity A2). The average Table 1 material Appendix of 60% between the highest value the sites reached between gradually with increasing the 0–100 m rings and decreased tance from the nested colony centre point; Table 1a). Table point; the nested colony centre tance from were sites distributed and randomly sites beaver between analysis (Supplementary SIMPER the clearly described by distance from the territory vs random x dis distance from (beaver centre Anthropogenic (a) Variables Table 1. PERMANOVA comparisons of all distinguished land use types (a) and repeated measures ANOVA of woody plant cover (combina plant cover of woody ANOVA measures use types (a) and repeated all distinguished land comparisons of 1. PERMANOVA Table and urban area) grassland, land, land, arable of fallow areas (combination shrubs) and anthropogenic and shrubs, forest, and tion of trees in presented are ANOVA the from results hoc post The sites. unoccupied selected randomly and beavers by occupied sites the between (b) rings. and 201–500 m 2. *: 0–100, 101–200 Fig. Woody plants Woody (b) Variables All land use types Ecology in a human-dominated World 7.6% (Fig. 4a,Supplementary material Appendix 1 Table A5). beaver habitatvariability, andtheshared variation explained density thegeographicallocation explainedonly10.3%ofthe ver sites(efficiency largest fractionofvariability oflanduse compositioninbea nificant. Thegeographicallocationofacolonyexplained the and joinedvariation [pseudo-F position ofthecolony[pseudo-F ver density [pseudo-F on datafrom beaver sitesonly. Alltestedvariables (i.e.,bea ANOVA results see Table 1b. ** sons are presented orhorizontal asvertical brackets;* (CI). confidence intervals Significant posthoc Tukeytest compari Points represent mean values and the whiskers indicate the 95% (combination offallow land,arablegrassland,urbanarea). trees forests, (b)anthropogenic andshrubs, areas andshrubs); pied random sites (blue); (a) woody plant cover (combination of 200 and 201–500 m) surrounding beaver sites (red) and non-occu Figure 2. Land-usecharacteristicsinsequentialrings(0–100,101– 194 = p < 0.01and*** = =

46.7%), whileinthecaseoflocal beaver p = < ., df 9.9, 0.001.For adetaileddescriptionofthe = = , p 1, = ., p 5.3, ., df 5.2, = .0] geographical 0.001], =

0.001]) were sig = 6 p 16, = = p 0.001], < 0.05, - - - - - (Monte tions thatwere obtained,16were statisticallysignificant Among thespatialpredictors from the33PCOeigenfunc implies thatthedifferences between thehabitatsselected by and r beaver densityandwoodyplantcover (r the Dru bodies, clearly visible along the Vistula River but also within fluctuations attheintermediate-scalewithinmainwater r percentage ofwoodyplantcover (r ent showed nocorrelation withthelocalbeaver densityand the Dru ences inbeaver territoriesatthebroad scale,mostlybetween Table A7).Thefirsteigenvector (PCO.1)describesthediffer (Fig.most important 4d,Supplementary materialAppendix 1 bodies withinthespatialgradients,whichwere selectedas separation between the beaver sites from the different water material Appendix 1 Table A6).TheCCAplotpresents aclear PCO.1 (14.2%) and PCO.4 (7.2%) (Fig. 4b, Supplementary ofthespatialvariability werelargest portions explainedby material Appendix 1 Table A5).Amongthesepredictors, the Table in A2).Themostsubstantial difference wasobserved landuse types;Supplementary materialAppendixshrubs 1 for areas covered by woodyplants(Shrubs, Forest, Trees and from theavoidance ofarablelandsandtheclear preference cantly from thatofrandomareas. Thesedifferences resulted Habitat composition in beaver territories differed signifi Habitat selection Lucas 1969). through the spatialvariation ofthehabitat(Fretwell and not only by the densityof these territorial animals but also ideal despotic model of habitat selection (IDD) was affected previously reported inthisspecies.Our results implythatthe andecoregion scales,which hasnotbeen (family territory) the variability ofthesehabitatswasdependentonbothlocal arablelands.However,particularly stressing it isworth that andlowershrubs, cover by anthropogenic landusetypes, ofwoodyplants,such as ian habitatswithlargeproportions In the Vistula River delta,European beaver preferred ripar Discussion Fig. 5b). type (r 0.001; Fig. 5a)andnegatively with anthropogenic landuse plant cover (r vers waspositively correlated ofwoody withtheproportion centage ofoverlap ofthe0–500mbuffersoccupied by bea between thelocalbeaver densityandhabitatquality. Theper local densityofbeaver sitesandavailability offoodresources. bution, whileonamore localscale,theywere related tothe beavers dependedmostlyontheirbroad-scale spatialdistri s

= In addition,we foundahighlysignificantcorrelation 00, p –0.05, s

= s

- = 02, p –0.20, ż ż Carlo permutationtestp no Reserve, anditwassignificantlycorrelated with no Reserve andotherwaterbodies,thisgradi 07, p –0.74, = s

0.517 respectively; Fig. 4e).PCO.4illustrates = .9 p 0.69, < < 0.016, respectively; Fig. 4f). This result 0.016,respectively; Fig. 4f). 0.001; GAM R < 0.001,GAMR < s

= 0.042,Supplementary 2

00, p –0.08, = s 07, p 40.7%,

= 03, p –0.31, 2

= 81, p 28.1%, = .2 and 0.327 < < 0.001; 0.001 < ------

Ecology in a human-dominated World - - - 195 ski et al. ń no Reserve no Reserve ż , Rosell and and , Rosell , Fustec et al. –1 –1 , reaching relatively relatively , reaching –1 , Rosell et reason al. 1998). The no Reserve and the Vistula River River Vistula no Reserve and the ż –1 , Hartman 1994; 0.72 km –1 Beavers colonize the best habitats during the initial phase the best habitats during the initial phase colonize Beavers willow scrub that is favored by beavers (Nolet et al. beavers by scrub that is favored willow 1994, Fustec et al. 2009, Buli and Kostkan 2001, John vari habitat may be an important predictive Willow 2013). settlement in both optimal and subopti able for beaver before phases expansion the during habitats mal/marginal carryinga population reaches capacity (Fustec et al. 2001, 2010). John et al. (Fig. 3), and a large part of the variation was represented by by 3), and a large part was represented of the variation (Fig. occupied 1a). Beavers (Table in PERMANOVA the residuals optimal (Dru from of habitats, a wide variety to marginal River) and suboptimal (Nogat River) Vistula and the fact that which could reflect (other studied watercourses), been occupied, so coloniza habitats had already all preferred similarity between the Additionally, or stopped. tion slowed to the fact may be related occupied and non-occupied areas has territories food conditions in beaver that the current a pattern since settlement (Wright et al. 2002). Such altered number the represents that of a population is characteristic and density of animals close to the carrying capacity for the (John et al. 2010). area available increase numbers their and as development, population of (Fustec quality environments they switch to lower et al. 2001, Pinto et al. 2009, John et al. on the 2010). Based delta River Vistula conclude that the we of this study, results The carrying capacity. population was at, or near, beaver studied we the water bodies sites between densities of beaver km 0.05 to 0.86 colonies from differed in the Dru high values to other studies (0.12 km when compared 2003; 0.25 km 2001; 0.76 km Hovde for such a high density may be due to the high quality of by which was mostly overgrown the riparian environment, differed significantly (Hartmancontrast significantly 1996), in marked differed and random sites, that similarity in beaver The to our study. delta, was manifested by River Vistula document for the we areas random and sites beaver both within variation large the ------At the end of the last centuryAt in population the beaver Urban areas provided some of the lowest proportions in of the lowest some provided areas Urban Both the beaver and random areas were covered primar covered were areas and random Both the beaver Sweden was still increasing and animals settled only in opti was still increasing Sweden sites and non-occupied areas beaver mal habitats; therefore, However, this result was not related to building dams because was not related this result However, sites only. observed beaver such constructions in three were higher in beaver territories at all distances. This could reflect territories at all distances. This could higher in beaver from of the species, and it may result preference the real such as digging canals. the engineering activities of beaver, John et al. could be observed for preferences 2010). Some by type, which was represented the smallest land cover even type small water bodies. The proportion of this land cover were values and the average was higher near colony centres, both beaver and non-occupied habitats; however, this land this however, habitats; non-occupied and beaver both which also has animals in all ranges, by use type was avoided basin; River the Morava (e.g. from previously been reported of beavers, which can inhabit and reproduce in different different in reproduce inhabit and which can of beavers, landscapes cultivated semi-natural and including habitats, 1998). and Rosell (Nolet earlier work that showed how the importance of habitat vari how that showed earlier work (John et al. process ables may change during the colonization related to the flexibility result might also be 2010). This ingly contradictory results could be because the preferences ingly contradictory could be because the preferences results pro this land use type change with the towards of beavers with agree results These plants. woody portionpreferred of cover, which suggests that fallow lands could be treated as as lands could be treated fallow which suggests that cover, to land also corresponded neutral habitat elements. Fallow dissimilarity mea the second highest contribution to the seem Such occupied and unoccupied areas. between sure ily by fallow land, a remnant of high anthropogenic pres of high anthropogenic land, a remnant fallow ily by land fallow average similar also showed The two areas sure. ing, however, as many previous studies have reported similar reported studies have as many previous ing, however, observations (Hartman Fustec et al. 1996, and 2001, John 2010). 2009, John et al. 2009, Pinto et al. Kostkan the average Trees and shrubs cover, which was up to 5 times and shrubs cover, Trees the average territories (0–100 m ring) than of beaver higher in the centre result was not surpris of random sites. This in the centre Figure 3. Detrended correspondence analysis (DCA) ordination plot based on the composition of all distinguished land cover types within types of all distinguished land cover plot based on the composition (DCA) ordination analysis correspondence Detrended Figure 3. and (b) selected water bodies. points) and random sites (blue points) sites (red (a) the beaver the 0–500 m buffer for Ecology in a human-dominated World winter foodsupply, the percentage ofwoodyplantsdecreased watercourse. With increasing distancefrom alodge,den,or favoured and avoided land use types alongthe shore of a vers couldbedistinguishedby thespatialdistribution of Our studyalsoindicatedthatthehabitats preferred by bea Spatial habitatstructure withinafamily territory (PCO.1 andPCO.4,respectively) asafunctionoftheirgeographiclocationinthe Vistula River Fens. (d) SpatialValues distributionofbeaver sitesalongthe main watercourses inthe Vistula River Fens. ofkeyeigenfunctions (e)and(f) sites (points)inrelation tothemainspatialgradients(PCO.1 andPCO.4eigenfunctionsfrom PCNM)andthelocalbeaver sitesdensity. p spatial scales, derived from the PCNM analysis, whichwere model (Monte used inthevariation partitioning Carlo permutation test permutation testp that are explainedby bothgeographiclocation(usingthePCNMmethod)andlocaldensityofbeaver sites(all VP fractions,Monte Carlo (a) Venn presenting diagramofvariation partitioning uniqueandshared fractionsofthetotalvariation inbeaver siteslandusecomposition Figure withprincipalcoordinates 4.Results (VP) ofvariation ofneighbourmatricesanalysis(PCNM)andassociatedanalyses. partitioning 196 ≤ 0.042,Supplementary materialAppendix 1 Table A6).(c)CCAplotpresenting thevariability oflandcover compositionamongbeaver =

0.001, Supplementary materialAppendix 1 Table A5).(b)Bar ofsignificanteigenfunctionsillustratingdifferent chart - woody plants within a beaver territory (3rd-orderwoody plantswithinabeaver territory selection) tre, asindicatedby theSIMPER analysis. 0–100, 101–200,and201–500 mrangesfrom acolonycen beaver sitesandunoccupiedareas withinthesubsequent confirmed by thedecrease inthe totaldissimilarity between while anthropogenic area increased. Thispreference wasalso Such preferences for a clearly clumped distribution of -

Ecology in a human-dominated World ------197 Our study suggests that the spatial scale for habitat selec study suggests that Our habitat selection, which occurs at a local scale, ani During cipal coordinate (Legendre et al. the com cipal coordinate 2009). Moreover, scale was ecoregion territories at the broad position of beaver density and the proportion of woodyindependent of beaver habitat selection that 2nd-order plants. These findings showed on both local was spatially limited and dependent beavers by status of the population, development habitat quality and the the density of which was close to the carrying capacity. a broad ecoregion scale. ecoregion a broad important than the density of occupied terri tion is more that variation partitioningresults of the showed tories. The 16 significant by the importance of spatial scale, represented 14 times more over eigenfunctions in the PCNM, explained to the sites, compared beaver between of the habitat variation influential spatial gradients, The most density. local beaver two completely PCO.1 and PCO.4, provided by represented broad-scale of spatial scale. The interpretations different to the entire spatial component, which could be attributed coor the principal by was represented and level ecoregion density and of local beaver PCO.1, was independent dinate local PCO.4 described a more whereas woody plant cover, with correlated was gradient spatial the which in situation, both of these variables. as a consequence, the mals assess the quality of food resources; not considered spatial scales are at broader habitat differences 2009b). an animal during this selection stage (Mayor et al. by Vistula The habitats distributed along the water bodies of the a by which could not be evaluated delta varyRiver in quality, reason during dispersal. This factor likely is the single beaver that the com based on the PCNM showed why our results mostly with the largest territories differed position of beaver by them, which was reflected geographical distance between the first prin explained by the highest proportion of variation one of which may be the spatial distribution of beaver sites at of beaver one of which may be the spatial distribution ------(Fig. 5) also illustrate a large spread of points, which suggests 5) also illustrate a large spread (Fig. by other importantvariables, that the dataset was influenced 2010). We believe that this is a consequence of larger ter that this is a consequence of believe We 2010). densities to lower habitat, leading in poorer-quality ritories (Fustec et Wright al. 2001, et al. 2002). The GAM plots positively correlated with the abundance of woody plants and correlated positively lands, which confirms previ anthropogenic with negatively John et al. 2009, Pinto et al. 2001, ous findings (Fustec et al. the quality of selected habitats depends on population den a settlement decision is possible only in unoccu where sity, 1972). Based 1969, Fretwell and Lucas (Fretwell pied areas was density beaver local the that determined we GAM, the on selection The ideal despotic model of habitat selection describes how the long-term occupation and utilization of a certainthe long-term occupation and utilization territory 2002). (Wright et al. importance of a large spatial scale in habitat The be based not only on the local abundance of food resources of food resources be based not only on the local abundance of these resources but also on the clumped spatial distribution importantwithin a home range, which may be especially for and Pearson 1979, Schoener 1979, Jenkins 1980, Fryxell and Fryxell 1980, 1979, Schoener 1979, Jenkins and Pearson of process during the 1992). Therefore, 1991, Fryxell Doucet decision should colonization or natal dispersal, a settlement or winter food supply to reduce the cost of food transporta or winter food supply to reduce as central place foragers, and well-known are tion. Beavers with increas of food items decreases foraging time or size or pond (Orians a riverbank ing perpendicular distance from Our study shows that beaver prefers habitats where the most habitats where prefers that beaver study shows Our located near are resources and abundant available, valuable, vicinity of a family lodge a territory or in the direct centre may be beneficial from the optimal foraging theorymay be beneficial from view is localised in point because the majority of food resources 1966). and Pianka the vicinity of a family lodge (MacArthur Figure 5. General additive models (GAM) between the local density based on the percentage overlap of the beaver sites and (a) the percent of the beaver overlap the local density based on the percentage models (GAM) between additive 5. General Figure a colony centre. from lands for the 500 m buffer of woody plants and (b) anthropogenic age cover Ecology in a human-dominated World Cie Buczma, Wojciech Kosmalski, Mateusz Barcikowski andRobert Zmudczy Acknowledgements population densitiesare takenintoaccount. phase when both the spatial distributions of resources and study beaver habitatpreferences even intheclimax population capacity. Moreover, we demonstratedthatitwaspossibleto population from the Vistula River delta has reached carrying scale. potential future territory total amount and the spatial distribution of resources at the decisions hadtobebasedontheassessmentboth the anthropogenic landcover increased. Therefore, settlement plants declinedwithdistancefrom acolonycentre, while wasclumpedandnon-random; thecoverterritory ofwoody not atbroader scales(e.g.between largewatersheds). it applieswithinrestricted areas (e.g.asmallwatershed)but habitat selectionwaslimitedby scale,andinthecaseofbeavers beaver dispersal.Thissuggeststhattheidealdespoticmodelof population density, which may represent thescaleof depended on habitat quality and was correlated with local distant waterbodies. the beavers were notabletocompare theconditionsbetween settle probably occurred independentlyateachsitebecause ver populationdensity. Therefore, thebehaviouraldecisionto were independentoftheabundancewoodyplantsandbea large geographicaldistancesbetween theterritories,which in the Vistula River deltaecoregion mostlyresulted from the Differences inhabitatcompositionaround beaver colonies Conclusions delta area constraints(Kondracki withfew 2000). tions), whichallows forbeavers todispersethroughout the ofconnec and17000 km ditches (approximately 3500 km bodies are well connectedthrough by rivers anddrainage migration barriers. In the Vistula River delta, distant water differences inhabitatselection were notinfluenced byany fortheinterpretationimportant ofspatialvariability thatthe France: 8.8 Europe (Sweden: yr averaged 12–19.7 km than theaverage dispersaldistancesofcolonizingbeavers in Nogat Dru distances between theabove waterbodies(meandistances: water body. Thisdifference likely reflectsthefactthat nel mightberegarded asthebestpossible habitatinanother rivers. Asaresult, suboptimalhabitatinoneriver orchan bodies suchastheDru ver habitatselectionoccursindependentlywithinlargewater gradients onageographicalaxis(Fig. 4),we believe thatbea 198 The results of our studyalsoindicatedthatthebeaver The distributionofwoodyplantswithinabeaver family Within eachwatercourse, beaver habitat selection Based ontheCCAbiplotandpresentation ofspatial ś li ż ń no Reserve– Vistula ski fortheirassistanceduring fieldwork; Łukasz =

ń 21.55 km; Vistula –Nogat ska-Skarbek inplanning thestudy;Anna forhersupport ± 12.8 SD km yr – We wouldlike tothankKatarzyna ż no Reserveandthe Vistula orNogat =

–1 42.94 km;Dru , Fustec et al. 2003). It, Fustec et al. is also =

22.76 km)isgreater –1 , Hartman 1995; , Hartman ż no Reserve– Ś witaj, - - - - MW.M1/2/2011 andMW.M1/4/2013). Fundusz Ochrony Protection and Water Management inGda by the financially supported Regional Fund for Environmental pomorskiego z okre bobra europejskiego – damage causedby thebeaver (PLInwentaryzacja populacji delta fensinarea ofthePomeranian province andminimize ‘Inventory oftheEuropean beaver populationin Ochrony Environmental Protection inGda Pomorskiego wGda Gda Pomeranian Head ofMelioration andHydrotechnic Structures in Funding the manuscript. Douglas Kelt toimprove fortheirvaluable commentsandefforts would liketothanktheanonymousreviewers andtheeditor O We wouldalsoliketothankprof. LechStempniewicz, Agnieszka assistance indatamanagement;andthebestpilotJakub Leszczak. Manikowska- Małgorzata Kozak, Izabela Kulaszewicz, Izabela Fischer, Brygida Collen, P. and Gibson, R. 2001. The general ecology ofbeavers Clarke, K.R.andGorley, R.N.2006.User Manual/Tutorial. Clarke, K. R.1993.Non 2013.The Mammalian faunaof‘Jezioro Ciechanowski, M. et al. Cassini, M. H.2013.Distribution ecology:from individual habitat 2005. TerritoryCampbell, R.D. et al. and group sizes in Eurasian Buli Borcard, D.andLegendre, P. 2002.All-scalespatialanalysisof Beier, P. andBarrett, R.H.1987.Beaver habitatuseandimpactin 2013.Spatial spread ofEurasian beavers inriver Bartak, V. et al. Aszyk, M. and Kistowski, M. 2002. Monitoring bobra w gda Aszyk, K.1994.Bobry Anderson, M. et al. 2008.Permanova+ forPrimer: guidetosoftware References for theanalysisofaerialphotographs;ZdzisławBło ż arowska and Małgorzata Wydra Finally, for editing this article. review. –Rev. Fish. Biol. Fisher. 10:439–461. and riparianhabitats,the subsequent effectsonfish–a ( – Primer-E, Plymouth. changes incommunitystructure. –Aust. J.Ecol. 18:117–143. – Natural Monograph. Mantis Olsztyn. pp. 33–81. Dru use tospeciesbiogeographicalrange.–Springer. – Behav. Ecol. Sociobiol. 58:597–607. beavers ( Monograph. Mantis Olsztyn., pp. 33–81. Reserve. – In: Nitecki Cz. (ed.) Druzno Lake – Natural matrices. –Ecol. Model. 153:51–68. ecological databy meansofprincipalcoordinates ofneighbour 794–799. Truckee River basin,California.–J. Wildl. Manage. 51: Ecol. 82:587–597. networks: acomparisonofrangeexpansionrates.–J.Anim. – Pozna pomorskim. województwie Wyd. Uniwersytet Gda and statisticalmethods.–PRIMER-E,Plymouth. ń ń Castor ski, M. et al. 2013.The vegetation oftheLake Dru sk (PLZarz ż –Thisstudywascarriedoutatthe request ofthe no’ Nature Reserve. –In: Nitecki, C.(ed.),Druzno Lake spp.), asrelated totheirinfluence onstream ecosystems Ś ń rodowiska wGda Castor fiber . Ś lepowro ą d Melioracji iUrz Ś rodowiska i Gospodarki Wodnej w Gda ś leniem sposobu minimalizacji szkód)’, and ń ń Ż sku) andtheRegional Directorate for ): echoesofsettlementandreproduction? ska, Joanna Pardus and Wojciech Pardus uławy Wi ń ‐ skie. –DJ,Gda parametric multivariate analysesof ń sku) according totheproject ś ń ą lane w obszarze województwa lane wobszarzewojewództwa sk (PLRegionalna Dyrekcja dze ń Wodnych Województwa ń ń sk. sk (PL Wojewódzkisk Ż uławy, Vistula ń ski, Gda ż no Nature ń ski for ń sku; ń sk

Ecology in a human-dominated World

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