Dam Invaders: Impoundments Facilitate Biological Invasions into Freshwaters Author(s): Pieter T. J. Johnson, Julian D. Olden and M. Jake Vander Zanden Source: Frontiers in Ecology and the Environment, Vol. 6, No. 7 (Sep., 2008), pp. 357-363 Published by: Ecological Society of America Stable URL: http://www.jstor.org/stable/20440936 . Accessed: 08/10/2014 16:50 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 support@jstor.org. . Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Frontiers in Ecology and the Environment. http://www.jstor.org This content downloaded from 158.135.136.72 on Wed, 8 Oct 2014 16:50:55 PM All use subject to JSTOR Terms and Conditions RESEARCH COMMUNICATIONS Dam invaders: biological RESEARCH COMMUNICATIONS_ impoundments invasions into facilitate freshwaters PieterTJ Johnsonl*,JulianD Olden2, andM JakeVander Zanden3 of theglobal biodiversitycrisis,with more declining and extinct Freshwaterecosystemsare at the forefront in or marine environments. Hydrologic alterationsand biological invasions repre terrestrial species than sent twoof thegreatestthreatsto freshwater biota, yet the importanceof linkagesbetween thesedriversof environmentalchange remainsuncertain.Here,we quantitativelytestthehypothesisthat impoundments facilitatethe introductionand establishmentof aquatic invasivespecies in lake ecosystems.By combining and geographical distributionof fivenuisance data on boating activity,water body physicochemistry, invaders in theLaurentianGreat Lakes region,we show thatnon-indigenousspecies are 2.4 to 300 times more likelyto occur in impoundmentsthan in natural lakes,and that impoundmentsfrequentlysupport comparisonsof thecontemporaryand historical landscapes revealed that multiple invaders.Furthermore, impoundmentsenhance the invasion riskof natural lakesby increasingtheirproximityto invadedwater bodies, highlightingthe role of human-alteredecosystemsas "stepping-stone"habitats for the continued spreadof freshwaterinvaders. Front Ecol Environ 2008; 6(7): 357-363, doi:10.1890/070156 expandingglobalnetworkof dams and impound Shea andChesson 2002;Havel etal. 2005). In additionto habitats, ments has dramaticallychanged thedistribution of increasingthe abundance of standing-water largerandmore accessible are frequently freshwateracross the landscape.Broad-scale physical impoundments impactsof suchhydrologicalalterationsincluderiverine tohumans thanarenatural lakes;theyarebuilton river with hydrologicconnectionsto neighboring sedimentretention,enhanced evapora ine systems fragmentation, tion,and increasedgreenhouse-gas production(Nilssonet lakes,and are youngerin age and experiencegreaterdis turbancethando naturalsystems(Thorntonet al. 1990; al. 2005;Dudgeon etal. 2006; Poffetal. 2007). Less under arehypothesized interact Havel etal. 2005).All of theseinfluences stood,however,ishow dams or impoundments with otherlarge-scale agentsofglobalchange,suchas bio to enhance species colonization and establishment.If fornon also providegreateropportunities logicalinvasions.In theUS alone,more than80 000 large impoundments speciestospreadacrossthe landscape,identifi damsand 2.5million smallerimpoundments populatethe indigenous landscape(Graf1999; Smith et al. 2002), while close to cation of these alteredenvironmentsas invasionhubs and preventingfuture 1000 introducedfreshwater species contributeto the could aid efforts aimedat predicting invasions(MuirheadandMacIsaac 2005;Hulme 2006). ofnativebiodiversity endangerment (Rahel 2002;USGS on human behavior (ie boat By coupling information 2007). Together,dam constructionand biological inva and the characteristics, sions aremajor contributorsto the currentbiodiversity ing),waterbodyphysicochemical we of fivenuisance invaders, crisisin freshwater which exhibithigherrates geographicaldistributions ecosystems, and quantitativelyevaluated the hypothesisthat impound of extinctionand a greaterproportionof threatened ormarine environ ments facilitatebiological invasions into freshwaters. endangeredspeciesthan in terrestrial we addressedthefollowing questions: ments (Ricciardiand Rasmussen 1999; Dudgeon et al. Specifically, 2006). Despite the increasing ubiquityofboth impound ments and invasivespecies in freshwater (1) Are impoundmentsmore frequentlyinvaded ecosystems,a pressing, yetunansweredquestion remains:do impound comparedtonatural lakes? offactors relatedtocolonization ments enhance invasionriskbypromotingtheestablish (2)What combination ment and spreadofnon-indigenousspecies? potential(ie propagulepressure)and establishment Creation of impoundments mightbe expected to pro suitability (ie species'ecologicalniche) accountfor observed differencesin invader occurrences mote invasionsby increasingcolonizationopportunities waterbodies? betweennaturaland impounded fornon-indigenoustaxa and by enhancing theirsub sequent establishmentsuccess (Kolar and Lodge 2000; (3) Has thewidespread creation of impoundments ecosys facilitatedspecies invasionsin freshwater 'Ecologyand EvolutionaryBiology,UniversityofColorado, Boulder, tems? An CO *(pieter.johnson(colorado.edu);2SchoolofAquatic and Fishery Sciences, University of Washington, Seattle, WA; 'Center for Limnology,UniversityofWisconsin,Madison, WI Our studyexamined fivewidespreadnuisance invaders associated with substantial ecological or economic C) The Ecological Society of America This content downloaded from 158.135.136.72 on Wed, 8 Oct 2014 16:50:55 PM All use subject to JSTOR Terms and Conditions a rntersai PT) Johnsonetal. invasions and freshwater Impoundments N / 0 30 60 km Li.S / .LJLake Myriophyllum I I I Eurasianwatermilfoil 88"W 90>W 92tW _ Superior m. Zebra mussel ~~~~~~~~~~Dreissena polymorpha A 4 46"N spicatum x N - A Af ^_ * waterflea ~~~~~~~~~~~~~~~~Spiny longimanus X ww wBythotrephes 2&s A A~~ *A A t j Wisconsin A AAf AAA ~AA A A A 44?N 6 A A Rainbow smelt AOsmerusmordax MinnesotaA A* | lowi l \*1s A00-J 4wiQ Rusty crayfish Iowa A AA A Water 42*N rusticus ~~~~~~~~~Orconectes body type * Natural lake A Impoundment Illinois species forwhichat leastonenon-indigenous and impoundments (redtriangles) ofnaturallakes(bluecircles) Figure1. Distribution Eurasianwatermilfoil, zebra Michigan(n = 1080). Invasivespeciesincluded: inWisconsinandUpperPeninsula isestablished crayfish. andrusty smelt, waterfleas,rainbow mussels,spiny spicatum), hydropower,paper mills, and recreation,creating a impacts:Eurasianwatermilfoil(Myriophyllum waterbodies (Graf spiny water fleas diversemosaic ofnaturaland artificial polymorpha), mussels (Dreissena zebra mor 1999).Moreover,byvirtueof theircloseproximityto the rainbowsmelt(Osmerus longimanus), (Bythotrephes rusticus; Figure1). By highlyinvadedGreat Lakes, inland lakes in this region (Orconectes crayfish dax), and rusty of non-indige includinginvasivespeciesthatvarywidely in taxonomy, have been colonizedby a broad diversity introductionpathway, life history,and physiological nous species(Holeck etal. 2004). we providea robusttestof theaforemen requirements, tioned questions and reduce the likelihood that our * Methods results are idiosyncraticto a single species. The differ how naturallakesand impoundments LaurentianGreat Lakes regionoffersan ideal testing To determine and invader introduction affect likely to in inva attributes biological between groundto evaluate interactions for data physicochemical compiled in we establishment, rich Already impoundments. sions and hydrologic 1081 impoundments) and (4200 lakes bodies 5281 water has undergone lakes, this region natural glacially-formed acrossWisconsin and theupperpeninsulaof extensivedammingin the20th century,foragriculture, distributed _wwwvfonflersinecology-org of America. 0 TheEcological Society This content downloaded from 158.135.136.72 on Wed, 8 Oct 2014 16:50:55 PM All use subject to JSTOR Terms and Conditions PTJJohnsonetal. Impoundments and freshwater invasions Michigan.We focusedon environmentalcharacteristics ments ("historical"environment). We focusedthisanaly as these potentiallyimportantforaquatic invadercolonization sis on zebra mussels and Eurasian watermilfoil, and establishment, including surface area (km2), species are among themost likelyto be transported by upstreamwatershed area (km2),maximum depth (m), boating activity,and because theycontrast a recent mean summer water clarity(Secchi depth inm), specific invasion (eg zebramussels; firstrecordedinWisconsin conductance ([Lmhos cm-,), hydrologic connectivity in 1994) with a well-establishedinvasion (egwatermil (seepage versusdrainage),human accessibility(defined foil; firstrecorded inWisconsin in 1965). For each by thepresenceof a boat landingornavigablewaterway), uninvadednatural lake,we calculated thechange indis and straight-line distance to thenearestGreat Lake (as a tance to the nearest invadedwater body between the sourcepopulationofnon-indigenousspecies,particularly historical and contemporaryscenarios.We compared forspinywater fleas,zebramussels,and rainbowsmelt). observed changes in distance to the average distance Boatermovementamongwaterbodies isone of thedom traveledby90% of registered boaters inWisconsin using inanttransport vectorsforthespreadofmany freshwater trailers(48 km;Buchan and Padilla 1999), and consid non-indigenousspecies, so we followedLeung et al. ereduninvaded lakeswithin 48 km of an invadedwater (2006) in using thenumberof boat landingsperwater body highly vulnerable to a potential introduction bodyas a proxyforinvaderpropagulepressure.In support event. Importantly, because not all lakesand impound of thismetric,we founda strongpositive relationship ments inWisconsin were surveyed,calculated changes between thenumberof boat landingsand human visita indistancebetween invadedand uninvadedwater bod tionsfor937waterbodies,accordingtoboatermovement iesare consideredestimates.This analysiscapitalizeson data froma randomizedsurveyof >50 000 registered the observation that inter-lakedistance is negatively Wisconsin boaters(WebPanel 1). correlated to invasion risk and assumes that, in the on water body physico absence of impoundments, We combined information thenumberof invadednat and boatingbehaviorwith themost recentdis chemistry ural lakeswould not have increased(eg due to increased tributional data forEurasianwatermilfoil(n = 682 sam visitationratesbyboaters). pledwaterbodies), zebramussels (n = 353), rusty crayfish (n = 567), spinywater fleas(n = 341), and rainbowsmelt M Results (n = 83; Figure1). Only data thatmet strictcriteria were used; includeddata (1) pertainedonly toaquatic invaders Among the1080 sampledwaterbodies, the invasionlike reasonablyclassifiedas "nuisance"species; (2) contained lihoodof impoundments exceeded thatof natural lakes. presenceand absencedocumentation;(3) were collected For each of thefiveinvaders,impoundments were signifi validatedsamplingprotocols;and (4) cantlymore likelythannaturallakesto be invaded(two usingstandardized, were recordedby trainedbiologistsor fromvouchered tailedPearson chi-squaredanalysis,df = 1, P < 0.05; when theypertainedto invaderoccurrences. Figure2a). The associationbetween impoundments specimens, and Surveyedwater bodies (n = 1080) were broadlydistrib invaderoccurrencesvaried among the species:Eurasian utedacrossthestudyregion(Figure1) andwere similarto watermilfoil(logisticregression odds ratio = 2.4), rusty accessiblewater bodies in the comprehensivedatabase. crayfish (2.5), zebramussel (2.7), rainbowsmelt(3.0), and Sampling protocols,data sources,and an evaluation of spinywater flea (7.8). Among the 189 water bodies sur potentialsystematic biasesarepresentedinWebPanel 1. most commonnon-indigenousspecies veyedforthe three We used two-tailedchi-squaredtestsof independence (zebramussels,Eurasianwatermilfoil, and rustycrayfish), to compare the invasionlikelihoodof natural lakesand impoundments were also significantly more likelythan impoundments foreach invader,and assessed the effect naturallakesto support multiple invaders(Figure2b). sizeof significant using theodds ratio.The relationships Physicaland biological differences between lakesand odds ratioestimatesthe increaseinodds ofbeing inone contributedto theobserveddifferences in impoundments outcomecategory(eg invaded)when thevalue of a pre invasionprevalence. In comparisonwith natural lakes, dictorchanges (eg fromnatural lake to impoundment). impoundments were 1.4 times lower in averagewater We subsequentlyused logistic regressionto develop clarity, 2.1 timeshigherinconductivity, 4.3 timeshigher species-specific models thatpredictedtheprobabilityof in numberof boat landings,8.7 times largerin surface invaderoccurrenceas amultivariatefunction ofvariables area,and 44.6 timeslargerinwatershedarea (WebTable related to invader colonization and establishment, 1). Impoundments were also 68% more likelythannat includingimpoundment statusand thephysicochemical ural lakes to be accessible by humans and 54% more variablesdescribedabove (seeWebPanel 1). likelyto have hydrologicconnectionsvia streamnet Finally,we estimatedthepotential forimpoundments works (WebTable 1). Nevertheless,among thebest-sup to increasetheriskof invasionintonaturallakesbycom ported logistic regression models (based on Akaike's paring the straight-line distances between each unin information criterion[AIC] values and associateddiag vaded natural lakeand thatlake'snearestinvadedneigh nostics), impoundmentstatus remained a significant borunder twoscenarios:a landscapewith impoundments explanatory variableforpredictingtheoccurrenceof four (present-dayenvironment)and one without impound out of fiveinvaders(odds ratiorangingfrom2.1 to300), C The EcologicalSocietyofAmerica This content downloaded from 158.135.136.72 on Wed, 8 Oct 2014 16:50:55 PM All use subject to JSTOR Terms and Conditions Impoundments and freshwater invasions (a) 100 (b) Natural lakes -ge2( m Impoundments 50n T TI 0 60| inyIwatur las - Impoundments 80 V PTJJohnsonetal. v~~~~~~~ 0 ~~~~~~~~~~~~~40 I (~60 30 0 0 40 20 20 T 10 0 0 0 ,~~ ~~o Ole e~~~ C" 0 1 2 3 Number of invaders Figure 2. (a) Percentage of surveyed lakes and impoundmentsthatsupported spinywater fleas (n = 341 sampledwater bodies), zebra mussels (n = 353), rainbow smelt (n = 83), rustycrayfish(n = 567), and Eurasian watermilfoil (n = 682). Error bars represent 95% confidence intervals,for a proportion. According to two-tailed Pearson chi-squared tests of independence, ** = P < over-invaded relative tonatural lakes for each invader (C= P < 0.05, **= P < 0005, impoundmentswere significantly 0.0005). (b) Comparison of thenumber of invaders in lakesand impoundmentsfor the 189 water bodies sampled for the threemost common invaders (two-tailedPearson0~31 = 15.219, P = 0.002). ofotherenviron ered vulnerableto boatermovement froman invaded even afteraccountingforthe influence mental and anthropogenicfactorslikelyto affectintro waterbodyfrom45 lakesto67 lakes(-50% increase).For Eurasianwatermilfoil, impoundments themore established ductionand establishment (Table 1).Overall ratesofcor distanceby23.9% from rectclassification based on n-foldcrossvalidationwere reducednearestinvaded-neighbor 94% forspinywater fleas(Cohen's K = 0.6, P < 0.001), 17.6 km to 13.4 km (meanchange ? SE = 4.2 ? 1.2 km; 86% forrainbowsmelt (K = 0.655, P < 0.001), 86% for t95= 3.4; P = 0.001; Figure3b) and increasedthenumber zebramussels (K = 0.418, P < 0.05), 81% forEurasian ofvulnerablelakesfrom87 to95 (a 9.2% increase). watermilfoil(K = 0.603, P < 0.001), and 65% forrusty crayfish(K = 0.302, P < 0.05). Models for individual Discussion invadersincludedexplanatoryvariablesbroadlyconsis tentwith previousresearchon thesespeciesor aquatic A centralchallengeof invasionbiology is tounderstand how invasive species interactwith other large-scale invaders generally(Table 1;WebPanel 1). For zebramus orwatershed agentsofenvironmental change (Didham etal. 2007). Of sels alone, inclusionof eitherconductivity sta paramountimportanceto thistask is learninghow these area as predictorvariableseliminatedimpoundment driversare likelytochangeand interactin tusfromthebest-fit model (Table 1). Conductivity, par environmental Our find as itpredictstheconcentration ticularly of calcium ions, shapingfutureratesand patternsof invasions. constructionand theconver ingssuggestthatreservoir can limitzebramussel shellformation and, therefore pop riversto standingwatersmay ulti For water bodieswith dissolved sion of free-flowing ulation establishment. calciumconcentrations and conductivity measurements, mately facilitatethespreadof invasivespeciesacross the thesevariableswere strongly landscape.Although aquatic invasionshave previously positivelycorrelated(r = been linkedto impoundments (egMoyle andLight 1996; 0.80; n = 168; P < 0.001). and historicalland Kolar and Lodge 2000), the driversof thisassociation Comparisonsbetweenpresent-day and Most studiesof reservoirs scape scenarios (ie with and without impoundments,have remainedambiguous. have occurredin areaswith fewnatural respectively) revealedthat,forzebramussels,thepresence aquatic invaders of impoundments reducedmean distance to thenearest lakes,have focusedon lotic (runningwater) environ introduced game invadedwater body by 45%, from84.2 km to 46.4 km ments,or have examined intentionally (mean change ? SE = 37.7 ? 3.1 km; tI,4= 12.1, species, leaving open the possibilitythat any lentic P < 0.0001; Figure3a). Based on thedistancetraveledby (standing per water)habitat- ratherthanimpoundments 90% of registered Wisconsin boatersusing trailers,this se - would have producedan identicalincreasein inva increasedthenumberof uninvadednaturallakesconsid sion risk.Here, we overcome previous limitationsby ? The EcologicalSocietyofAmerica This content downloaded from 158.135.136.72 on Wed, 8 Oct 2014 16:50:55 PM All use subject to JSTOR Terms and Conditions PTJJohnsonetal. and freshwater invasions Impoundments Table 1. Best-fIt logistic regression models Model variables foreach freshwater Invasive species, as determined by AIC Eurasianwaterrnilfoil Zebra mussel fN= O.52 ,N-=0.58 n = 578 n = 306 Impoundmentstatus 0.73 i 0.29* (2.07) Number of boat landings 0.29 ? 0.09" (1.342) 0.20 ? 0.08* (1.218) -1.37 ? 0.59* Distance toGreat Lakes (0.255) Watershed area Maximum depth 4.36 ? 0.40* (78.1) Surfacearea Rustycayfish N.= 0.19 n 408 5.70 i 2.57 2.58 ? 0.96' 0.74 (298.9) (13.21) 1.54 ? 0.55w (0.012) (0.826) -5.23 ? 3.22 (0.005) 12.73 ? 6.04* 1.73? 0.65 (21.39) (336614) (5.617) 2.23 ? 0.87 -6.52 ? 3.92 (0.001) 11.93? 3.03w (151669) 5.50 ? 0.94** (244.92) -1.07 ? 0.28 -2.76 ? 0.927 (0.003) -1.16 ? 0.34* -6.45 ? I.0'* (0.002) (6.295) (1.637) 0.49 ? 0.22* (0.315) Constant 1.84 ? 0.5r 0.49 ? 0.22 (0.345) Hydrology -0.19 ? 0.06w (4.644) -4.39 ? 1.55* 00.26" (2.1) (2.307) (9.314) Conductance Rainbowsmelt rN= 0.16 n = 82 0.84 ? 0.28 3.06 ? 1.27 Secchi depth water flea Spiny 0N= 0.78 n = 120 (1.637) - 15.81? 4.38w (0.00001) 6.44 ? 6.94 (624) 10.21 ? 16.0 - 11.75? 3.0# (27121) (.0001) Notes: Listed in the firstrow are each invader;theNagelkerke r value for the best model, and the total sample size ofwater bodies. In the firstcolumn are the indepe variables included ineach analysis;all continuous variableswere logd-ransfofmed.Provided in each subsequent column are themaximum likelihoodparameter estimates ? I SE and the odds ratio (inparent ) foreach independentvariable in the finalmodel. A blank cell indicates thata given variablewas not selected in the finalmodel. =Wald chi-squared P < 0.05; Wald chi-squaredP < 0.005. quantitatively analyzingspatialpatternsof invasionsfora were weighted towardaccessiblewater bodies, notably diverseassemblageof species in a region underrepresenting taxonomically thethousands of lessaccessiblenatural richwith impoundments and naturallakes. lakesin thisregion, we expect thatthereportedinvasion Consistentwith the hypothesisthat impoundments bias in favorof impoundments isa considerableunderes facilitatefreshwater statuswas a invasions,impoundment timate;inaccessiblelakesare less likelyto be visitedby strongpredictorof non-indigenousspecies'occurrences. humansand, therefore, tobe invaded. For each of the fiveinvaders,impoundments were 2.5 to Despite inclusionofnumerousanthropogenic and envi 7.8 timesmore likelythannatural lakes to have estab ronmental variableshypothesizedtoaffecttheprobability lishedpopulations,and thesepatterns were robusttovari of invader introductionand establishment,impound ation in thedefinition of impoundment (seeWebPanel 1). ment statusremaineda significant predictorof occur Among thesubsetofwaterbodies surveyedforthe three rence forfourof the five invaders(Table 1). In several most common invaders(zebramussels,Eurasianwater cases, inclusion of additional explanatory variables milfoil,and rustycrayfish),impoundments were also sig enhanced the strengthof the impoundmenteffect,as nificantly more likelythannaturallakestosupport multi estimatedby theodds ratio(up to300 timesovernatural ple invaders. Considering thatcurrentsamplingefforts lakes).Models forindividualinvadersincludeda variety ? TheEcological Society America of This content downloaded from 158.135.136.72 on Wed, 8 Oct 2014 16:50:55 PM All use subject to JSTOR Terms and Conditions www.frontiersinecology.org Impoundments and freshwater invasions PTJJohnsonetal. (a) Zebra mussel Present 40 0 0 0 L Historical Historical 8 landscape(laes o~n landscape ~~~~~~~~~~Prsn-Paday - 20 0(A .0 1 (takesesenmpoundments 0 0 50 150 100 200 (b)Eurasian watermilfoil 50 Present Historical Historical landscape wakes only) ladcape Prsnt-day nearest invadelatebodyunderthe eakes + impoundments) 40 .0 0 30 o 20 0 theyoungage, increasedniche availabil ity,and high disturbanceregimecharac Natural teristicofmost impoundments. lakesin theGreat Lakes regionareat least 10 000 yearsold and formed duringglacial retreatin the Pleistocene,whereas the oldest impoundmentin our datasetwas 161 yearsold (median= 68 years),indi have had cating that thesecommunities lesstimetoassemble. substantially Newly assemblednative communities in im poundments may exhibitcorrespondingly lowerbiotic resistance,thereby making themparticularly vulnerable to species invasions(Levine and D'Antonio 1999; Kolar andLodge 2000;Shea andChesson 2002;Havel et al. 2005). Impoundments arealsomoredisturbedthannaturallakes, oftenexhibiting marked fluctuationsin water levels,temperature, fishstocking, and nutrientcontent (Thornton et al. 1990) - conditionsthatshould increase invasibility (egDavis etal. 2000). Broadly invasivespeciesmay alsohave distributed become adapted to impoundmentsin otherpartsof theirnativeor non-native theirsuccessin ranges,further facilitating North American impoundments. 0 120 40 60 80 Distance to nearest infested water bodly (kin) Figure 3. Comparison of the distance between uninvaded natural lakes and the nearestwater body invadedby (a) zebramussels and (b) Eurasian waternilfoil under two scenarios: present-day landscapewith lakesand impoundments(top stackedbars) and historical landscape with lakes and no impoundments(bottom stacked bars). Dashed lines indicate themean distance between uninvaded natural lakes and the nearest invadedwater body under thetwo scenarios; shaded regionreflectstheaverage inter-lakedistance traveledby boaters in thestudyregion. However,we cannotexcludethepossibil differin itythatlakesand impoundments invaderpropagulepressurebeyondwhat we showed in our analysis.Despite the ofwaterbodyareaand combinedstrength numberof boat landingsin predicting waterbodypopularity(seeWebPanel 1), additional differencesin boat landing quality,visitationrates,or typeof boater could generatedifferencesin invasion risk. An additionalchallengebeyondunder standingthefactors drivingthe impound its ment-invasion linkage is identifying lake consequencesfornatural ecosystems. to increase on We estimatedthepotentialforimpoundments ofpredictors broadlyconsistent with previousresearch these species.The numberof boat landingsand/orthe the invasionriskof natural lakesby comparingthe dis proximity of awaterbody to thenearestGreat Lake posi tancesbetweeneach uninvadednaturallakeand itsnearest tivelypredicted the occurrenceof spinywater fleas, invadedneighborunder two scenarios:a contemporary with impoundments and a historicallandscape Eurasianwatermilfoil, rainbowsmelt,and zebramussels, landscape highlightingthe importance of human boatingactivity devoid of impoundments. We foundthat impoundments and distanceto invadersourcepopulation(egMacdsaac et reducedthemean distancebetweenan uninvadedlakeand al. 2004; Leung etal. 2006). The importance neighborbyhalf,increas ofwatershed itsnearestzebramussel-invaded area forboth zebramussels and rustycrayfish highlyvulnera probably ingthenumberofnaturallakesconsidered reflectedincreasedcolonizationopportunitiesthrough ble to invasionby almost50%. For themore established increasedthenum impoundments upstreamlakeor riverconnections(eg Puth andAllen Eurasianwatermilfoil, 2005;Havel andMedley 2006). berofhighlyvulnerablelakesbyabout 10%.This finding We suggest that the strong association between suggeststhat the facilitativerole of impoundmentsin impoundments and non-indigenousspecies resultsfrom invasionmay attenuateas the landscapebecomes satu www.frontiersinecology.org a The Ecological This content downloaded from 158.135.136.72 on Wed, 8 Oct 2014 16:50:55 PM All use subject to JSTOR Terms and Conditions Society of America and freshwater invasions Impoundments PTJJohnsonetal. effects of habitat modification invasion and on native species rated with invadedwaterbodies.These resultsalso suggest speciesdecline. TrendsEcol Evol 22: 489-96. Ijlj increasetheriskof invasionintonat thatimpoundments Downing JA,PrairieYT, Cole JJ,etal 2006 The global abundance HB distanceand by ural lakesby both decreasinginter-lake size distribution and of lakes, ponds, and impoundments. LimnolOceanogr 51: 2388-97. increasingthe totalnumberof invadedwater bodies on et al 2006. Freshwater D, Arthington AH, Gessner MO, Dudgeon the landscape. chal threats, status and conservation importance, biodiversity: which ecosystems arevulnerableto invasions Predicting lenges. Biol Rev 81: 163-82. isan important goal of invasionbiology,and identificationGraf WL. 1999. Dam nation: a geographic census of American dams thatwill notonlybecome invadedbutwill ofenvironments and their large-scale hydrologie impacts. Water Resour Res 35: 1305-11. contribute to secondaryspreadof inva disproportionately and cost-effectiveHavel JE, Lee CE, and Vander Zanden MJ. 2005. Do reservoirs sivespeciesoffers thepromiseof targeted invasions into landscapes? BioScience facilitate 55: 518-25. management(Hulme2006).We arguethatimpoundmentsHavel KA. invasions across spatial 2006. Biological JE and Medley invaders may functionas invasion"hubs" forfreshwater scales: of clado intercontinental, regional and local dispersal ceran Zooplankton. theirsubse (MuirheadandMacIsaac 2005), facilitating Biol Invasions 8: 459-73. intonaturalwaterbodies. Holeck KT, Mills EL, Maclsaac HJ, et al 2004. Bridging troubled quent spreadand establishment waters: links between invasions, biological understanding are now a ubiquitousfeature of theglobal Impoundments in the transoceanic and other entry vectors shipping, hydrologic system(Smithetal. 2002;Downing etal. 2006), Laurentian Great Lakes. BioScience 54: 919-29. and vast regions where lakeswere formerly rareare now Hulme PE. 2006. Beyond control: wider implications for the man while otherregions home to hundredsof impoundments, agementof biological invasions.JAppl Ecol 43: 835-47. dominatedby natural lakesnow also include Kolar CS and Lodge DM. 2000. Freshwater nonindigenous historically species: interactions with other global changes. In: Mooney HA and numerous impoundments. climate-induced Furthermore, in a changing Invasive Hobbs world. (Eds). RJ species and increasing humandemand changestowateravailability DC: Island Press. Washington, ofnew reser Leung B, Bossenbroek JM, and Lodge DM. 2006. Boats, pathways, for waterare likelytoprompttheconstruction voirsto increase water suppliesinmany regions(Palmeret invasions: estimating dispersal potential and aquatic biological of contemporary with gravity models. Biol Invasions 8: 241-54. al. 2008). The widespreaddistribution impoundments and theongoingcreationofnew impound Levine JM and D'Antonio CM. 1999. Elton revisited: a review of evidence linking diversity and invasibility. Oikos 87: 15-26. ments could provideenhancedopportunities forinvasive Maclsaac PA. 2004. HJ, Borberly JVM, Muirhead JR, and Graniero speciesto establishinnaturallakesand spreadacrossthe invasions and forecasting of inland Backcasting biological landscape. lakes.Ecol Appl 14: 773-83. 1996. Biological invasions of freshwater: Moyle PB and LightT 0 Acknowledgements rules and assembly empirical theory. Biol Conserv 78: 149-61. Muirhead JRandMaclsaac HJ. 2005. Development of inland lakes The authorsgratefully acknowledgethe data contribu tionsof JMaxted, K Webster,P Soranno,L Herman,R Martin, T Mojo, D Balsiger,K Grande, T Asplund, D Olson, JFilbert,L Joppa,and S Knight.The following Wisconsin agencies generously shared information: DepartmentofNatural Resources (WDNR), Michigan Department of Environmental Quality, the North and TemperateLakes LTER andBiocomplexityprograms, theGreat Lakes Indian Fish andWildlife Commission. S Carpenter, JHavel, D Strayer,H MacIsaac, and E Stanleyprovidedfeedbackhelpfulin shapingthemanu script.This project was funded by grants from the WDNR, theWisconsin Coastal Management Program, and theNational Science Foundation (DEB-0411760, DEB-0217533, andDEB-0083545). in an invasion as hubs Nilsson C, Reidy Fragmentation systems. Science network. ] Appl CA, M, Dynesius and flow regulation 308: 80-90. Ecol 42: and Revenga of the world's change Front Ecol Environ options. of regional Homogenization river 2008. 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