Relationships among invasive common carp,native fishes and physicochemical characteristicsin upper Midwest (USA) lakesMichael J. Weber, Michael L. BrownDepartment of Wildlife and Fisheries Sciences, South Dakota State University, Brookings, SD, USA

Accepted for publication January 21, 2011

Abstract – Common carp Cyprinus carpio is a widespread invasive species that, in high abundance, can imposenumerous deleterious effects in aquatic ecosystems. Common carp increase turbidity and nutrient availability whilereducing invertebrate prey resources and aquatic macrophytes, transforming shallow lakes from the clear- to turbid-water state. However, potential effects of common carp on native fish communities have received limited attention.We evaluated the relationships among relative abundances of nine native fishes and common carp for 81 lakes ineastern South Dakota and their associated physicochemical characteristics. Inverse threshold relationships amongrelative abundances of native fishes and common carp were identified for black bullhead Ameiurus melas, blackcrappie Pomoxis nigromaculatus, bluegill Lepomis macrochirus, white bass Morone chrysops and northern pike Esoxlucius, while marginally significant relationships were detected for largemouth bass Micropterus salmoides andsmallmouth bass M. dolomieu. Lakes where common carp relative abundance exceeded 0.6 fish per net night had lowabundance of native fishes, whereas lower abundance of common carp resulted in variable abundance of native fishes.Lakes with abundance of common carp surpassing 0.6 fish per net night were also characterised by larger surfaceareas and watersheds and impaired water quality (higher dissolved solids and chlorophyll a concentrations and lowersecchi depth). Our results are consistent with the biotic-abiotic constraining hypothesis that proposes biotic factors canregulate fish populations regardless of abiotic conditions. Thus, common carp abundance may need to be reduced andsustained below ecological thresholds to improve water quality and increase abundance of native fishes.

Key words: invasive species; common carp; biotic-abiotic constraining hypothesis (BACH); alternative equilibria; water quality

Introduction

Invasive species pose one of the greatest threats to thebiotic integrity of aquatic ecosystems. Human-medi-ated distributions of non-native species often result inthe decline or extirpation of native organisms (Moyle1986). However, effects of invasive species on nativefauna are highly variable and extremely difficult topredict (Moyle 1986; Lodge 1993), because environ-mental conditions vary and the processes by whichspecies coexist are complex (Huisman & Weissing1999). Common carp Cyprinus carpio are distributedworldwide and considered one of the most wide-spread, detrimental invasive species (Lowe et al.

2004) because of their ability to attain extremedensities (up to 1000 kgÆha)1; Neess et al. 1957;Panek 1987; Koehn 2004) and alter freshwaterecosystems (Weber & Brown 2009). Since theirintroduction to the United States, common carp haveexpanded their range and currently occupy a majorityof freshwater lakes, reservoirs, rivers and streams(Zambrano et al. 2006a). While common carp arehighly regarded throughout much of Europe and Asia(e.g., Arlinghaus & Mehner 2003), substantial effortand cost are associated with management of thisinvasive species in areas of introduction (Koehn 2004;Schrage & Downing 2004; Weber & Brown 2009;Weber et al. in press). Common carp induce numerous

Correspondence: Michael J. Weber, Department of Wildlife and Fisheries Sciences, South Dakota State University, Brookings, SD 57007, USA; Tel.: (605) 688-6121,Fax: (605) 688-4515. E-mail: Michael.Weber@sdstate.edu

Ecology of Freshwater Fish 2011: 20: 270–278Printed in Malaysia Æ All rights reserved

� 2011 John Wiley & Sons A/S

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deleterious effects on shallow lakes, at both thecommunity and ecosystem level by increasing nutrientavailability, turbidity and phytoplankton abundance,reducing benthic macroinvertebrates and aquatic mac-rophytes, and modifying zooplankton assemblages(Lougheed et al. 1998; Parkos et al. 2003; Weber &Brown 2009). However, responses of higher trophiclevels to common carp perturbation are not well defined.

Ecosystem alterations induced by common carphave great potential to influence native fish popula-tions, but direct empirical evidence is limited. Byreducing or eliminating aquatic macrophytes anddisrupting substrates, common carp may indirectlyreduce abundance of other fishes through reductions inspawning and nursery habitats (Stuber et al. 1982;Durocher et al. 1984; Paukert et al. 2002). Addition-ally, common carp may adversely affect growth andsurvival of fishes that rely on invertebrate preyresources through reductions in these prey types.Centrarchids can experience reductions in growthand survival in the presence of common carp (Wolfeet al. 2009), and inverse relations between commoncarp and some fishes have been documented (Egert-son & Downing 2004; Jackson et al. 2010). Even ifprey resources are not reduced, increased turbidity,commonly associated with common carp popula-tions, may alter piscivore and planktivore foragingbehaviour and reduce success (Miner & Stein 1996;Reid et al. 1999), affecting fish condition, growthand survival. Conversely, poor foraging success ofpiscivores may result in increased survival of theirprey (Wolfe et al. 2009) and recruitment potential.Thus, common carp may interact with and haveeffects on native fishes through multiple complexmechanisms.

Common carp are hypothesised to reduce theabundance of native fishes, but current information isvague. Egertson & Downing (2004) and Jackson et al.(2010) indicated that common carp abundance wasnegatively related to abundance of bluegill Lepomismacrochirus, black crappie Pomoxis nigromaculatusand largemouth bass Micropterus salmoides, likelydriven by impaired water quality in these systems. Inthis study, we examined relationships between relativeabundances of nine native planktivorous and pisciv-orous fishes and invasive common carp across naturallakes and reservoirs. Given that invasive common carpimpose numerous deleterious effects on aquatic eco-systems that are often amplified with common carpabundance (Parkos et al. 2003; Chumchal et al. 2005;Weber & Brown 2009), we hypothesised that theabundance of native fishes is negatively related tocommon carp abundance and that systems withabundant common carp would have impaired waterquality. To test these hypotheses, we analysed a largestandardised lake data set to evaluate relationships

among relative abundances of invasive common carpand nine native species commonly found in upperMidwestern lakes: black bullheads Ameiurus melas,black crappies, bluegill, northern pike Esox lucius,white bass Morone chrysops, smallmouth bass M. dol-omieu, largemouth bass, walleye Sander vitreus andyellow perch Perca flavescens. Second, we evaluatedeight physicochemical variables [surface area, meandepth, watershed ⁄ lake surface area ratio, secchi depth,total Kjeldahl nitrogen (TKN), total phosphorus (TP),chlorophyll a and total dissolved solids (TDS)] toexplain differences in fish communities across lakemorphologies and limnological conditions in relationto common carp abundance.

Methods

Data collection

Relative abundances of adult common carp, blackbullheads, black crappies, bluegill, northern pike,white bass, smallmouth bass, largemouth bass, walleyeand yellow perch were obtained from standardised fishpopulation surveys conducted by South DakotaDepartment of Game, Fish and Parks (SDGFP) ineastern South Dakota (Fig. 1). Aforementioned sportfishes represent the most abundant species throughoutthe study region and provide an important recreationalvalue to anglers (St. Sauver et al. 2009). Waterbodieswere surveyed between 2003 and 2007 with acombination of trap nets, experimental gill nets andboat electrofishing; some fish communities weresurveyed multiple times between 2003 and 2007, butonly relative abundance data from the most recentsurvey were used to avoid pseudoreplication (Hurlbert1984). Standardisation of methods allowed for com-parisons of fish catch data among lakes, providing alarge and robust database. Trap nets were used in 81waterbodies to sample common carp, black bullhead,bluegill, black crappie, northern pike and smallmouthbass. Trap nets were constructed with 0.9 m high by1.5 m wide frames, 18.3-m long leads and 19-mm barmesh netting. Experimental gill nets were used in 71waterbodies to sample white bass, walleye and yellowperch. Experimental gill nets were 45.7 m long by1.8 m deep with one 7.6 m panel each of 13-, 19-, 25-,32-, 38- and 51-mm bar mesh monofilament netting.Nighttime boat electrofishing (pulsed direct current,200–250 V, 3–6 A) was conducted on 16 waterbodiesto sample largemouth bass. Effort invested for eachgear varied by lake and increased as a function of lakesurface area. All locations were randomly selectedwithin spatial strata, and nets were set for approxi-mately 24 h. Catch per unit effort (CPUE) wascalculated as the number of adult fish collected pertrap net or gill net night or per hour of electrofishing.

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Limnological and lake morphology data werecollected by the South Dakota Department of Envi-ronmental and Natural Resources from a subset of 34lakes in eastern South Dakota concurrent with fishsampling. Physicochemical variables measured weresurface area, mean depth, watershed ⁄ lake surface arearatio, secchi depth, TKN, TP, chlorophyll a and TDS.Sampling was conducted in accordance to the U.S.Environmental Protection Agency’s Clean Lakes Pro-gram standard operating procedures and analysis ofchemical parameters followed standard methods(physical solids, Methods 209A, 160.3, 209B, 160.1,209D and 160.2; water quality, Methods 365.2, 351.2,417F and 350.1; Stueven & Stewart 1996).

Analysis

Scatter plots of native fish and common carp CPUErelationships indicated that data were distinctlyclumped and related in a negative nonlinear fashion.Two-dimensional Kolmogorov–Smirnov (2DKS) testsare a useful tool for identifying nonrandom relation-ships in bivariate data (Garvey et al. 1998). Thus,

2DKS tests were used to identify significant relation-ships between native fishes and common carp. Whensignificant, the 2DKS test identified thresholds ofabundance between the native fish and common carpCPUE and categorised each lake as having either (i)high CPUE of common carp and low CPUE of nativefish (above the threshold) or (ii) low CPUE ofcommon carp and high CPUE of native fish (belowthe threshold; Jackson et al. 2010).

Fish communities and physicochemical characteris-tics were assessed using a Bray-Curtis similaritymatrix in Primer-E and four multivariate analyses(Clarke & Gorley 2001). To evaluate if characteristicsof fish communities were related to physicochemicalvariables, we used the RELATE function to detectsimilarities between data sets. Next, lake physico-chemical characteristics were analysed using ananalysis of similarity (ANOSIM) and nonmetricmultidimensional scaling (NMDS) to test for differ-ences in physicochemical parameters between systemsclassified as having high or low abundance of commoncarp, as determined by the aforementioned 2DKStests. NMDS grouped lakes with similar characteristics

Fig. 1. Locations of the 81 lakes and reservoirs sampled throughout eastern South Dakota, USA, denoted by a star. The star on the map ofNorth America indicates the location of South Dakota.

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near one another and those with dissimilar character-istics farther apart. ANOSIM calculates a global R-value ranging from 0 (no separation of groups) to 1(complete separation of groups) to describe similarityor differences between groups. Lastly, we used asimilarity percentage breakdown analysis (SIMPER)to determine which physicochemical parametersaccounted for at least 10% of the variation betweenlakes with high and low common carp CPUE.

Results

Lakes ranged in surface area from 15.6 to 6289 ha,had mean depths of 0.9–5.5 m and watershed ⁄ surfacearea ratios from 0 to 31.3 (Table 1). Relative abun-dances of all species populations were variable acrosslakes (Table 1). Mean trap-net CPUE of common carpwas 4.7 (±2.4 standard error, SE) and ranged from 0 to195.8. Mean trap-net CPUE of native fishes rangedfrom 0.3 to 123.2, mean gill-net CPUE of native fishesranged from 0.7 to 20.8, and mean largemouth basselectrofishing CPUE was 27.5.

Abundances of several native fishes were inverselyrelated to common carp abundance. Significant thresh-old relationships were identified for black bullhead,black crappie, bluegill, northern pike and white bass,whereas marginally significant threshold relationshipswere identified for largemouth bass and smallmouthbass (Figs 2 and 3). Threshold catch rates of sportfishes captured in trap nets ranged from 0.1 to 6.9,whereas the threshold for common carp ranged from

0.1 to 0.8 per net night (mean = 0.6). No relationshipswere identified between walleye or yellow perch andcommon carp relative abundance (Fig. 3).

Lake physicochemical characteristics varied acrosssystems in eastern South Dakota (Table 1) and wererelated to fish communities (RELATE: P = 0.006).Secchi depth ranged from 13 to 229 cm and TDSranged from 6.2 to 5202.5 mgÆl)1. Significant differ-ences in physicochemical parameters existed betweenlakes with high (>0.6 common carp per net night) andlow (<0.6 common carp per net night) common carpabundance (NMDS: stress = 0.04, Fig. 4; ANOSIM:Global R = 0.146, P = 0.02). Lakes with high com-mon carp CPUE had higher TDS (22% contribution),lower secchi depth (21% contribution), largerwatershed ⁄ surface area ratios (17% contribution) andsurface areas (16% contribution), and higher chloro-phyll a concentrations (15% contribution) compared tolakes with low common carp abundance (SIMPER:P < 0.05).

Discussion

Invasive species can influence population dynamics ofnative species or community interactions (Beisneret al. 2003; Simon & Townsend 2003), and theintroduction of some aquatic species has resulted inthe extirpation of some native fishes (Moyle 1986;Minckley & Deacon 1991; Witte et al. 1992).However, determining mechanisms and quantifyingthe effects of invasive fishes on native species is

Table 1. Number of lakes, mean, standard error, median, minimum and maximum values for lake physicochemical characteristics and catch per unit effort(CPUE) values for 10 fish species captured in trap nets, gill nets and electrofishing in eastern South Dakota. Lakes were surveyed once between 2003 and 2007.

Variable N Mean Standard error Median Minimum Maximum

PhysicochemicalSurface area (ha) 34 555.2 114.4 192.0 15.6 6289.0Mean depth (m) 34 2.6 0.1 2.4 0.9 5.5Watershed area:lake surface area 34 2.1 0.8 0.1 0.0 31.3Secchi depth (cm) 34 102.5 11.4 107.0 13.0 229.0Kjeldahl nitrogen (mgÆl)1) 34 1.9 0.2 1.7 0.5 8.6Total phosphorus (mgÆl)1) 34 0.5 0.3 0.1 0.0 10.4Chlorophyll a (ppm) 34 41.6 7.5 22.2 0.1 175.9Total dissolved solids (mgÆl)1) 34 1008.9 164.7 768.0 6.2 5202.5

Trap netCommon carp 81 4.7 2.4 0.3 0.0 195.8Black bullhead 81 123.2 32.5 14.8 0.0 1574.0Black crappie 81 12.6 4.0 1.2 0.0 243.6Bluegill 81 10.6 3.3 0.1 0.0 186.5Northern pike 81 1.2 0.3 0.4 0.0 15.4Smallmouth bass 81 0.3 0.1 0.0 0.0 8.7

Gill netWhite bass 70 0.7 0.4 0.0 0.0 25.5Walleye 70 10.1 1.6 5.7 0.0 68.0Yellow perch 70 20.8 3.0 12.0 0.0 117.7

ElectrofishingLargemouth bass 16 27.5 10.3 13.3 0.0 131.5

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D = 0.03P = 0.04(x,y) = (0.6, 6.9) (x,y) = (0.1, 0.1)

Fig. 2. Relationships among common carpcatch per unit effort (CPUE) and blackbullhead, black crappie, smallmouth bass,bluegill, and northern pike CPUE collectedwith trap nets. Note differences in y-axisscale for all species. Statistical results werederived from two-dimensional Kolmo-gorov-Smirnov tests.

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Fig. 3. Relationships among common carpcatch per unit effort (CPUE) in trap nets andwhite bass, walleye, yellow perch CPUEcollected with gill nets and largemouthbass CPUE collected with nighttime pulsedDC electrofishing. Note differences inx- and y-axis scale. Statistical results werederived from two-dimensional Kolmogorov-Smirnov tests.

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extremely difficult (Moyle & Light 1996). Abundanceof native species may be directly reduced (i.e.,predation) following the introduction of non-nativepiscivores into novel habitats (e.g., Moyle 1986;Moyle & Williams 1990; Eby et al. 2006). However,effects of omnivorous fish introductions, includingcommon carp, on native fish assemblages aredetected less frequently (Moyle 1986). Previous workhas documented negative relationships among com-mon carp and bluegill, black crappie and largemouthbass (Egertson & Downing 2004; Jackson et al. 2010).Our results build upon these and reveal that theabundance of several other native fishes is inverselyrelated to common carp abundance. However, ourresults are not surprising because dramatic decreasesin sport fish abundance have been identified followingthe establishment of common carp (Cahn 1929), andsuccessful common carp removal efforts have allowedsuccessful recruitment and re-establishment of fishpopulations (Ricker & Gottschalk 1940; Rose & Moen1953).

Dominant species often serve a major role indetermining community structure and ecosystemprocesses. Extremely high densities of common carpmay be found throughout the waterbodies we exam-ined (M. Weber, South Dakota State University,unpublished data) in addition to other parts ofMidwestern North America and south-central Austra-lia where densities up to 1000 kgÆha)1 have beenreported (Neess et al. 1957; Koehn 2004). Commoncarp abundance in these areas of introduction can be10-fold greater than common carp abundance inEurope (Crivelli 1983). Native fishes coexisting inecosystems with high common carp densities may beat the greatest risk and adversely affected by theirpopulations when compared to other regions where

native fishes may be more resilient to lower commoncarp densities. Inverse relationships between commoncarp and native fishes may be indicative of exploita-tion and interference competition where species thatcompete for limited resources replace one another(Seip 1997). Biomass of bluegill and black crappiemay decline linearly with increasing common carpbiomass (Egertson & Downing 2004). However, ourwork, in addition to others (Jackson et al. 2010),indicates that a threshold relationship exists betweencommon carp and native fishes, where the abundanceof common carp causes an abrupt change in abun-dance of native fishes.

Relationships identified between common carp andnative fishes support the biotic–abiotic constraininghypothesis (BACH) that suggests that biotic interac-tions can have an overriding negative influence onfishes despite abiotic conditions and may be particu-larly applicable for evaluating interactions betweennative and invasive or introduced species (Quist et al.2003, 2004; Quist & Hubert 2005). Instead of a linearrelationship, the BACH predicts a threshold to bioticstressors. High relative abundance of many nativefishes was only observed when common carp relativeabundance was low (<0.6 fish per net night) in thesystems we examined. When common carp relativeabundance was high (>0.6 fish per net night), abun-dance of native species within a waterbody wasconsistently low. If common carp did not have aneffect on abundance of native fishes, we would haveexpected to see variation in their abundance regardlessof common carp abundance, but this was rare acrossnumerous species and lakes. Thus, waterbodies withrelative abundances of common carp >0.6 fish per netnight are predicted to have relatively low abundancesof native fishes.

Multiple potential mechanisms exist where commoncarp may influence native fishes through predation andcompetitive interactions. First, predation is a majorforce structuring aquatic communities, and early lifestages of some native fishes may experience predationby common carp (Miller & Beckman 1996; Marsden1997), which may reduce recruitment. Second, nega-tive effects of common carp on native fishes may bebecause of competition for prey resources (Panek1987) as common carp can experience high dietaryoverlap with other fishes (Specziar et al. 1998; M.Weber, South Dakota State University, unpublisheddata). Adult common carp primarily consume benthicmacroinvertebrates and can reduce their densities(Specziar et al. 1998; Parkos et al. 2003; Weber &Brown 2009). Although adult common carp arenot zooplanktivorous, they may alter zooplanktoncommunities through indirect processes and reducedensities of large-bodied taxa such as Daphnia(Lougheed et al. 1998; Parkos et al. 2003; Weber &

COC CPUE > 0.6

COC CPUE < 0.6

NMDS stress = 0.04

Fig. 4. Nonmetric multidimensional scaling (NMDS) ordinationplot of physicochemical characteristics (surface area, mean depth,watershed ⁄ lake surface area ratio, secchi depth, Kjeldahl nitrogen,total phosphorus, chlorophyll a and total dissolved solids) collectedfrom lakes with low (<0.6 common carp per net night; open circle)and high (>0.6 common carp per net night; open triangle) commoncarp abundance.

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Brown 2009). All native fishes examined in this studyconsume zooplankton or benthic invertebrates duringsome life stages [black bullhead (Repsys et al. 1976),black crappie (Pope & Willis 1998), white bass,walleye (Beck et al. 1998), largemouth bass (Apple-gate & Kruckenberg 1978), bluegill (Mittelbach1984), smallmouth bass (Easton & Orth 1992),northern pike (Sammons et al. 1994), and yellowperch (Lott et al. 1996)], and competition for limitedprey resources may greatly reduce their abundance.

Effects of common carp on native fishes may alsobe because of habitat degradation. Common carp canincrease turbidity and nutrients in lakes through theirbenthic foraging behaviour, switching lakes from theclear- to turbid-water state (Lougheed et al. 1998;Parkos et al. 2003; Weber & Brown 2009). Lakes withabundant common carp populations were more turbid(higher TDS and lower secchi depth) and productive(higher chlorophyll a concentrations) compared tosystems with low common carp abundance. Althoughfish biomass is thought to increase with increasingproductivity (Bachmann et al. 1996), only benthivor-ous fish populations (i.e., common carp) may be ableto utilise increased production in eutrophic systemssuch as these (Egertson & Downing 2004). Commoncarp populations in eastern South Dakota have beenpreviously linked to suspended solid concentrations(Weber et al. 2010), whereas lakes in Iowa, USA,showed a shift in fish species composition from sportfish to common carp as turbidity and productivityincreased (Egertson & Downing 2004; Jackson et al.2010). Additionally, we found that high common carpabundance occurred in lakes with large surface areasand large watersheds. Egertson & Downing (2004)found higher common carp abundance in lakes withlarge surface areas, whereas Jackson et al. (2010)found high common carp abundance in lakes withsmall watersheds. Large watersheds may increasenutrient loading and sedimentation within lakes,interacting synergistically with common carp benthicforaging to impair water quality. Impaired physico-chemical habitats can alter fish communities andfavour those tolerant of poor water quality (Zambranoet al. 2006b; Jackson et al. 2010). Because commoncarp are generalists and tolerant of a wide range oflimnological conditions (Edwards & Twomey 1982;Weber et al. 2010), they are often associated withhighly turbid, productive systems, whereas the abun-dance of many sport fishes has been shown to declineunder similar conditions (Egertson & Downing 2004;Jackson et al. 2010).

Alternatively, habitat suitability and biotic resis-tance are important determinants of the success ofinvasive species (Moyle & Light 1996) and mayregulate common carp abundance. However, commoncarp, similar to other successful invaders, are gener-

alists, and are tolerant and thrive in a wide range ofhabitats (Edwards & Twomey 1982; Weber et al.2010). Thus, habitat is likely not a controlling factorlimiting their abundance below 0.6 fish per net night.The BACH suggests that interactions (i.e., nicheavailability, competition and predation) betweennative species and invaders may limit the abundanceof invasive species (Moyle & Light 1996), in this casebelow the BACH threshold. One potential bioticmechanism controlling common carp abundance ispredation. Predation may partially regulate commoncarp populations in their native range (Crivelli 1983),and we identified potential inverse relationshipsamong common carp and native predators (i.e.,largemouth bass, smallmouth bass, northern pike).Age-0 common carp have been found in diets ofwalleyes (Isaak et al. 1993), crappies (Marcy 1954)and northern pike (Sammons et al. 1994), and juvenilecommon carp size structure may shift because ofpredation (Nowlin et al. 2006). Thus, at appropriatedensities, predators may be able to regulate commoncarp recruitment, and hence abundance (Paukert et al.2003; Bajer & Sorensen 2010), and provide analternative explanation to our observed relationships.

We identified several relationships among relativeabundance of common carp, native fishes and physi-cochemical parameters. Lakes with abundant commoncarp (>0.6 fish per net night) had larger surface areasand watersheds and were more turbid and productivethan systems where common carp abundance was <0.6fish per net night. Additionally, the abundance of nativefishes was generally low in systems where commoncarp abundance was high. Our results build uponprevious research that suggests common carp canswitch aquatic ecosystems between alternative equilib-ria (Zambrano & Hinojosa 1999; Parkos et al. 2003;Weber & Brown 2009) and negatively affect nativefishes (Egertson & Downing 2004; Wolfe et al. 2009;Jackson et al. 2010). Thus, aquatic ecosystem restora-tions that incorporate common carp removals to reducetheir abundance below threshold levels (Weber et al. inpress) may be required to improve water quality andincrease the abundance of native fishes.

Acknowledgements

We would like to thank B. Blackwell, T. Kaufman, D. Lucchesi,T. St. Sauver and other SDGFP staff who provided fish surveydata. We thank P. Lorenzen and SDDENR for providingwater quality data, W. Schreck for providing maps, M. Fincelfor statistical consultation, and C. Guy and an anonymousreviewer for useful comments on a previous version of thismanuscript. Partial funding for this project was provided bythrough the Federal Aid in Sport FishRestorationAct Study 1513administered through South Dakota Department of Game, Fish,and Parks.

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