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Reviews in Fisheries Science & Aquaculture

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Catch-and-Release Influences on InlandRecreational Fisheries

Greg G. Sass & Stephanie L. Shaw

To cite this article: Greg G. Sass & Stephanie L. Shaw (2020) Catch-and-Release Influences onInland Recreational Fisheries, Reviews in Fisheries Science & Aquaculture, 28:2, 211-227, DOI:10.1080/23308249.2019.1701407

To link to this article: https://doi.org/10.1080/23308249.2019.1701407

Published online: 13 Dec 2019.

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Catch-and-Release Influences on Inland Recreational Fisheries

Greg G. Sass and Stephanie L. Shaw

Office of Applied Science, Wisconsin Department of Natural Resources, Escanaba Lake Research Station, Boulder Junction,Wisconsin, USA

ABSTRACTCatch-and-release (CR) has become a pervasive practice and “social norm” with anglers forsome inland recreational fisheries. This practice has been promoted for fish conservationand to meet angler and manager desires of greater fish abundances, angler catch rates, andtrophy growth potential. Catch-and-release in north-temperate inland recreational fisherieswas reviewed over time and documented the subsequent responses of fish populations tothe practice in catch rates, recruitment, abundance, size structure, growth, and trophypotential primarily focusing on black bass Micropterus spp., muskellunge Esox masquinongy,walleye Sander vitreus, and panfish (sunfishes Lepomis spp., crappies Pomoxis spp., yellowperch Perca flavescens). This review suggested that angler and manager desires may not bemet when fisheries are almost exclusively CR, CR may create situations where managers areunable to structure fish communities to meet such desires, and CR can cause imbalances infisheries managed for multiple species. Because CR may be one of the biggest challengesfacing inland recreational fisheries management in the 21st century and beyond, recommen-dations and future research considerations are provided and aimed to alleviate concernsidentified from this review to better balance fisheries, meet angler and manager desires,and to keep fisheries within a safe operating space.

KEYWORDSBass; inland recreationalfisheries; muskellunge;catch-and-release; walleye

Introduction

Conservative regulations and stocking in inland recre-ational fisheries have been used as conservation toolsto rehabilitate poorly recruiting, overexploited, or col-lapsed populations (Rypel et al. 2018), to keep fish-eries in a safe operating space (Carpenter et al. 2017),in biomanipulations to control invasive species (e.g.,rusty crayfish Orconectes rusticus; rainbow smeltOsmerus mordax) (Krueger and Hrabik 2005; Hein,Vander Zanden, and Magnuson 2007; Gaeta et al.2015), to increase angler catch rates, and to createput-and-take fisheries. Voluntary catch-and-releasepractices have also been promoted by managers, stake-holder groups, and anglers for certain species (e.g.,muskellunge Esox masquinongy, black bassesMicropterus spp.) with the goal of conservation.Likewise, anglers may be obligated to catch-and-release (CR) in certain fisheries managed for trophypotential because only a small subset of the popula-tion is legally harvestable. Often, CR practices are pro-moted under the assumption that catch rates,population size structure, and trophy potential will

improve as a result. Because fisheries management isaimed to maintain a balance between sustainability,catch rates, and trophy potential in harvested fisheries,fish community imbalances may occur when anglerschoose to CR certain fish species and/or species-spe-cific harvest regulations are ultra-conservative(Walters et al. 2000). Further, regulations requiringharvest are not typically used in North America; how-ever, this practice is more common in Europe(Arlinghaus 2007). Angler preferences for CR are gen-erally out of managerial control, therefore these idealsmay lead to imbalances in fish communities and notmeet angler expectations in the long-term; particularlyin fisheries managed for multiple species (Hansenet al. 2015; Miranda et al. 2017). Here, a review isprovided on the influences of CR practices in northtemperate inland lakes fisheries with primaryemphasis on largemouth bass M. salmoides, small-mouth bass M. dolomieu, muskellunge, walleye Sandervitreus, and panfish (sunfishes Lepomis spp., crappiesPomoxis spp., yellow perch Perca flavescens). A briefliterature review on CR research is provided, trends inCR behavior over time are identified, consequences of

CONTACT Greg G. Sass gregory.sass@wisconsin.gov Office of Applied Science, Wisconsin Department of Natural Resources, Escanaba LakeResearch Station, 3110 Trout Lake Station Drive, Boulder Junction, Wisconsin 54512, USA.� 2019 Taylor & Francis Group, LLC

REVIEWS IN FISHERIES SCIENCE & AQUACULTURE2020, VOL. 28, NO. 2, 211–227https://doi.org/10.1080/23308249.2019.1701407

CR behavior on catch rates, recruitment, relativeabundance, size structure, growth, and trophy poten-tial are discussed, and recommendations for betterbalancing fish communities in the future is provided.This review suggests that nearly exclusive CR behaviorand ultra-conservative harvest regulations may notachieve the perceived benefits of such practices tomeet angler, stakeholder, and manager expectations.

Catch-and-release in inland fisheries

Angler CR is a relatively recent socially-driven phe-nomenon for inland fisheries that has been promotedto conserve charismatic and previously harvested spe-cies, and to improve population size structure and tro-phy potential. For example, stakeholder groups suchas the Bass Anglers Sportsman Society (B.A.S.S.) andMuskies, Inc. have promoted CR as a mechanism toincrease trophy potential in the black basses and mus-kellunge. Prevalence of CR has also been commensur-ate with the increasing popularity andcommercialization of fishing tournaments over timebecause tournament sponsors often require and pro-mote CR. Nevertheless, many inland fisheries andanglers remain harvest-oriented (e.g., walleye, bluegillL. macrochirus, black and white crappie P. nigromacu-latus and P. annularis, yellow perch) (Gaeta et al.2013). Catch-and-release is assumed to be beneficialfor population conservation and increasing populationsize structure and catch rates in line with anglerexpectations. As a result, most studies of CR havefocused on measuring and reducing hooking mortalityto determine under what conditions and for what spe-cies CR will be most beneficial. Countless studies havebeen conducted on this topic across a range of speciesand habitats (e.g. Cooke and Suski 2005; Arlinghauset al. 2007; Cooke and Schramm 2007; Arlinghauset al. 2012; Kerns, Allen, and Harris 2012). In general,most studies examining hooking mortality of blackbasses and muskellunge have focused on hookinglocation on the fish, type of hook, water temperature,physiological stress associated with angling and tour-nament weigh-in procedures, and handling procedureeffects. Another body of literature has focused on CRinfluences on nesting black basses. Overall, estimatesof hooking mortality in the black basses have beenhighly variable (range 0-98%). Studies conducted inlaboratories or net pens have shown higher and morevariable hooking mortality (Schramm et al. 1987;Kwak and Henry 1995; Weathers and Newman 1997;Wilde 1998; Pollock and Pine 2007; Wilde and Pope2008), whereas mark-recapture studies (where released

fish are able to choose an unconfined natural habitatduring the refractory period) have shown lower andless variable hooking mortality (Cox 2000; Cline et al.2012; Sass et al. 2018). Tournament weigh-in proce-dures (holding in livewells for up to eight hours,exposure to air) have been shown to cause significantphysiological stress in the black basses (Edwards et al.2004a, Suski et al. 2005). Elevated water temperaturesand fight time have also been shown to increase blackbass hooking mortality rates (Edwards et al. 2004a,2004b, Keretz et al. 2018). Along with physiologicaland genetic effects, numerous studies have also sug-gested that CR of nesting black basses has negativenumerical population-level consequences (e.g. Phillipet al. 2009; Sutter et al. 2012; Parkos et al. 2013).Although individual nest success may be compro-mised by CR of nesting black basses (Kieffer et al.1995; Ostrand, Cooke, and Wahl 2004; Suski andPhilipp 2004; Trippel et al. 2017), no numerical popu-lation-level negative consequences of this practicehave been documented. For example, Jackson et al.(2015) found that CR for smallmouth bass had noeffect on recruitment, Trippel et al. (2017) observedno effect of bed-fishing on recruitment and overallreproductive success of Florida bass M. salmoides flor-idanus, and Zipkin et al. (2008) showed strong com-pensatory recruitment of smallmouth bass subjectedto experimental overharvest. Shaw and Allen (2016)found that successful Florida bass broods had to dropbelow about 5/ha before direct declines in recruitmentwere observed. In a northern Wisconsin lake, Sasset al. (2018) observed an increase in largemouth bassabundance when the population was subjected to CRfor five years suggesting increased recruitment.Therefore, hooking mortality and bed fishing do notappear to have direct negative consequences on theoverall abundance of black bass populations. For mus-kellunge, hooking mortality was found to be low inspecialized anglers (Landsman et al. 2011). Further,Shaw, Sass, and Eslinger (2019) showed that muskel-lunge hooking mortality was likely negligible usingdata from a long-term compulsory creel census on anorthern Wisconsin lake. Few studies have focused onthe potential negative implications of CR on popula-tion size structure and trophy potential.

Catch-and-release and angler behavior

Catch-and-release practices have increased in preva-lence over time for black bass and muskellungeanglers in Wisconsin. Wisconsin statewide largemouthbass release rates have exceeded 80% since the early

212 G. G. SASS AND S. L. SHAW

1990s and have averaged about 95% since 2000(Hansen et al. 2015) (Figure 1). Catch-and-release formuskellunge anglers in Vilas County, Wisconsin,increased in the mid-1980s and release rates haveexceeded about 90% since the mid-2000s (Gilbert and

Sass 2016) (Figure 1). On Escanaba Lake, Wisconsin,release rates have exceeded 75% since the mid-1980sdespite no size limit, bag limit, or closed season onmuskellunge. Escanaba Lake muskellunge release rateshave averaged >95% since 2010 (Eslinger et al. 2017)

Figure 1. Release rates of largemouth bass (Micropterus salmoides) in the Ceded Territory of Wisconsin during 1991-2011 (A)(Hansen et al. 2015), release rates of muskellunge (Esox masquinongy) in Vilas County, Wisconsin during 1978-2010 (B) (Gilbertand Sass 2016; hatched bars denote the percentage of muskellunge released), and release rates of muskellunge in Escanaba Lake,Wisconsin during 1987-2014 (C) (Eslinger et al. 2017). Reprinted with permission as G.G. Sass and/or S.L. Shaw was a coauthor onall cited articles.

REVIEWS IN FISHERIES SCIENCE & AQUACULTURE 213

(Figure 1). Further, Shaw, Sass, and Eslinger (2019)defined two distinct breakpoints (1995, 2011) in mus-kellunge harvest on Escanaba Lake during 1956-2016illustrating a transition in the fishery from harvest- tocatch-and-release-oriented. During the periods of1956-1994, 1995-2010, and 2011-2016, muskellungeharvest declined on Escanaba Lake and averagedabout 30, 7, and 1 fish per year, respectively (Shaw,Sass, and Eslinger 2019). The reduction in muskel-lunge harvest was unrelated to directed muskellungeangler effort, which was consistent during these timeperiods (Shaw, Sass, and Eslinger 2019). Using anangler diary survey, Gaeta et al. (2013) recordedrelease rates of 99% for muskellunge and 97% forblack bass in northern Wisconsin, with CR being themost common reason for release. Increases in CR ofmuskellunge in Canada have also been documented(Mosindy and Duffy 2007; Landsman et al. 2011).Similar patterns of increasing CR for black bass overtime have been observed across much of their rangein North America including the states of Alabama,California, Connecticut, Florida, Georgia, Iowa, Idaho,Michigan, Minnesota, Mississippi, Tennessee, Texas,and Wisconsin (Allen, Walters, and Myers 2008;Isermann, Maxwell, and McInerny 2013; Kerns, Allen,and Hightower 2016; Miranda et al. 2017; Hessenaueret al. 2018). Considerable evidence exists to suggestthat black bass and muskellunge anglers have transi-tioned from harvest-oriented to nearly exclusive CRregardless of regulations. This current “social fisherynorm” has rarely been evaluated to test whether thepractice of CR is achieving manager, stakeholder, andangler goals and/or creating imbalances in fish com-munities under an ecosystem-based fisheries manage-ment framework (but see Hansen et al. 2015; Mirandaet al. 2017; Hessenauer et al. 2018).

Catch-and-release and catch rates

One perceived goal of species-specific CR practices isan improvement in angler catch rates. The achievementof this goal appears to be species-specific and related tofish learning behavior. For the black basses, the increas-ing prevalence of CR has resulted in improvements infishery-independent and angler catch rates over time(Hansen et al. 2015; Rypel et al. 2016) (Figure 2).Increases in catch rates have also coincided with adecline in largemouth bass individual growth rates,population size structure, and trophy potential (Hansenet al. 2015; Rypel et al. 2016) (Figure 2). Fishery-inde-pendent and -dependent catch rate data for muskel-lunge during the transition to primarily CR is sparse

due to the inherent elusiveness of the species and alack of targeted fishery-independent surveys.Nevertheless, muskellunge compulsory creel censusangler data and fishery-independent surveyson Escanaba Lake have suggested that muskellungeangler catch rates (1 muskellunge/33 hours of directedangler effort) have not changed despite stable directedangler effort and a reduction in muskellunge abun-dance over time (Eslinger et al. 2017; Shaw, Sass, andEslinger 2019). Statewide fishery-independent surveysof muskellunge in Wisconsin have shown opposingtrends in fyke net and electrofishing catch per uniteffort (Rypel et al. 2016). Fyke net CPUE has increased,whereas electrofishing CPUE has decreased during therecent CR period (Rypel et al. 2016). This may suggestthat larger, mature individuals, which are more vulner-able to spring fyke netting, have increased in abun-dance, whereas smaller muskellunge, which are morevulnerable to fall electrofishing, may have declined inabundance. A similar pattern for smaller muskellungehas been observed in Escanaba Lake; however, adultabundance has declined over time (Eslinger et al.2017). Walleye anglers are harvest-oriented (Kempingerand Carline 1977; Gaeta et al. 2013; Hansen et al.2015). During 1946-2002, there was no size limit, baglimit, or closed season on walleye in Escanaba Lake(Kempinger and Carline 1977; Haglund, Isermann, andSass 2016). In 2003, a 711mm minimum length limitand a daily bag limit of one fish (i.e., essentially a CRregulation) was enacted for walleye on Escanaba Lake(Haglund, Isermann, and Sass 2016). Since 2003, nowalleye have been legally harvested from EscanabaLake by anglers, with only five walleye being legallyharvested by a Chippewa tribal member in December,2018. Despite an essentially CR regulation for walleye,adult density and angler catch rates did not differbetween the exploited and unexploited time periods(Haglund, Isermann, and Sass 2016). Plausible mecha-nisms for angler catch rates not being or negativelycorrelated with CR stems from learning behavior infishes. Learning behavior, lure avoidance, and the typeof bait (artificial vs. live) have all been shown to reduceangler catch rates across a number of species includingnorthern pike Esox lucius (Beukema 1970; Kuparinen,Klefoth, and Arlinghaus 2010), largemouth bass(Hackney and Linkous 1978; Hessenauer et al. 2016;Louison et al. 2019), rainbow trout Onchrhynchusmykiss (Askey et al. 2006), and walleye (Bailey et al.2018). Therefore, effects of voluntary or mandatory CRpractices on angler catch rates appear to be variable,species-specific, and related to fish learning behavior.

214 G. G. SASS AND S. L. SHAW

Figure 2. Trends in largemouth bass (Micropterus salmoides) electrofishing catch per unit effort (A), angler catch rates (B), andmean length of age-6 largemouth bass in Wisconsin lakes during 1991-2011 (Hansen et al. 2015). Reprinted with permission asG.G. Sass was a coauthor of Hansen et al. (2015).

REVIEWS IN FISHERIES SCIENCE & AQUACULTURE 215

Catch-and-release and recruitment

Catch-and-release angler behavior is presumed toincrease species-specific abundances over time.According to generalized Ricker and Beverton-Holtstock recruitment relationships, juvenile recruitmentwould be assumed to decline or stabilize withincreased adult stock size (Beverton and Holt 1957;Ricker 1975). For black basses, prevalence of CR haseither improved or had no effect on recruitment asstrong stock-recruitment relationships are lacking forthe black basses (Zipkin et al. 2008; Allen et al. 2011;Parkos et al. 2013; Sass et al. 2018). In Wisconsin,fisheries-dependent and -independent data haveshown that catch rates and relative abundances oflargemouth bass have increased over time with associ-ated declines in the mean length of age-6 fish(Hansen et al. 2015) (Figure 2). Sass et al. (2018) sug-gested that largemouth bass recruitment increasedafter a small, northern Wisconsin lake population wassubjected to CR angling for five years. Jackson et al.(2015) and Trippel et al. (2017) showed that CR hadno effect on smallmouth and Florida bass recruitment,respectively. Despite previous research suggesting thatvoluntary or mandatory CR of nesting black bass isdetrimental to individual nest success, no evidenceexists to support negative numerical population-levelconsequences. That said, climate change and associ-ated increases in growing degree days have beenshown to favor largemouth bass recruitment and rela-tive abundance (Hansen et al. 2017). Muskellungehave been shown to exhibit a Ricker stock-recruitmentrelationship (Eslinger, Dolan, and Newman 2010).Catch-and-release would be expected to increase adultmuskellunge abundances and therefore reduce juvenilesurvival unless the population was supplemented bystocking. Indeed, muskellunge recruitment hasdeclined to nearly undetectable levels on EscanabaLake over time (Eslinger et al. 2017) (Figure 3). Adultmuskellunge abundance has also declined significantlyover time suggesting that reproductive senescencemay occur in muskellunge and/or that there is adefined carrying capacity for relatively unexploitedmuskellunge populations in the absence of stocking(Eslinger et al. 2017; Shaw, Sass, and Eslinger 2019)(Figure 3). Given observations from the relativelyunexploited Escanaba Lake muskellunge population,CR may weaken resilience of the population toanthropogenic, abiotic, and biotic perturbations and/or the population may be seeking a new low adultdensity equilibrium like other unexploited fisheries inthe absence of stocking (Johnson 1976; Goedde andCoble 1981; Anderson et al. 2008). Like muskellunge,

elimination of legal harvest of walleye on EscanabaLake has resulted in low, stable recruitment (Haglund,Isermann, and Sass 2016). For walleye, boom and bustrecruitment events seem to be driven by harvest; how-ever, exploitation effects on recruitment appear to bepopulation-specific (Tsehaye, Roth, and Sass 2016;Sass and Shaw 2018). Cultivation/depensation effectson recruitment may prevent populations from recov-ery due to food web changes even if exploitation islimited or eliminated (Walters and Kitchell 2001).Ultimately, reproductive strategy (nest guarding, cen-trarchids; broadcast spawning with no parental care,muskellunge/walleye), exploitation, and cultivation/depensation influences on food web structure maydictate the relative influence of CR on recruitment.

Catch-and-release and relative abundance

The practice of species-specific CR is assumed toincrease the abundance of that species. Although thismay occur for some species, abundance responses toCR practices may also be species-specific and dictatedby aquatic ecosystem carrying capacity/productivityand the diversity of the fish community (Johnson1976; Goedde and Coble 1981; Anderson et al. 2008;Eslinger et al. 2017). For largemouth bass, CR hasgenerally led to increases in abundance regardless ofecosystem productivity, diversity of the fish commu-nity, and/or fishing for nesting males (Hansen et al.2015; Rypel et al. 2016; Sass et al. 2018) (Figure 2).During 1991-2011 (a period of high CR for large-mouth bass in Wisconsin), statewide electrofishingCPUE increased from about 15 to 30> 203mm bass/hr and from about 2 to 5> 356mm bass/hr (Hansenet al. 2015) (Figure 2). Over a 48-year period begin-ning in the 1960s, Rypel et al. (2016) showed thatlargemouth and smallmouth bass electrofishing CPUEin Wisconsin increased from about 10 to 32 and 3 to18 bass/hr, respectively. Sass et al. (2018) observed asignificant increase in largemouth bass abundance in anorthern Wisconsin lake subjected to five years ofexperimental CR fishing. Using mark-recapture tech-niques, largemouth bass density increased from about60 to 115 bass/ha (Sass et al. 2018). For largemouthand smallmouth bass, CR (regardless of bed fishing)appeared to have resulted in improved recruitmentleading to greater overall abundances (Rypel et al.2016; Sass et al. 2018). Although CR has resulted inincreased abundances of largemouth and smallmouthbass, abundance increases and overall carrying cap-acity appeared greater for largemouth than small-mouth bass. Despite the prevalence of CR for both

216 G. G. SASS AND S. L. SHAW

species, abundance increases observed in largemouthbass were about double that of smallmouth bass(Rypel et al. 2016). This may suggest that smallmouthbass are more specialized in their spawning habitatpreferences and diet, and may be more territorial dur-ing spawning than largemouth bass (Becker 1983;Rejwan et al. 1997). In small, northern Wisconsin lakebasins, largemouth bass nest densities ranged from52-129 nests/km of shoreline and nests were associ-ated with coarse woody habitat (CWH), boulders, finewoody habitat, human structures, and beaver struc-tures (Weis and Sass 2011). Lawson, Gaeta, andCarpenter (2011) also noted that largemouth bassnests were more associated with CWH in high

CWH:low lakeshore residentially developed northernWisconsin lakes, with nests found to be significantlydeeper in low CWH:high lakeshore residentially devel-oped lakes, and that docks did not serve as a surro-gate for CWH. To support this assertion ofsmallmouth bass being more spawning habitat special-ists than largemouth bass, Rejwan et al. (1997) foundthat smallmouth bass nests were patchily distributedwith high density nesting areas remaining stationaryacross years in Lake Opeongo, Ontario, Canada.Smallmouth bass nest densities ranged from 0-25nests/km of shoreline at the whole-lake scale andfrom 0-7 nests/100 m at the 100m scale (Rejwan et al.1997). These findings may suggest that largemouth

Figure 3. Trends in muskellunge (Esox masquinongy) age-0 electrofishing catch per unit effort (no./mile) (A) and adult density(no./acre) (B) on Escanaba Lake, Wisconsin during 1987-2014 (Eslinger et al. 2017). Reprinted with permission as G.G. Sass and S.L.Shaw were coauthors of Eslinger et al. (2017).

REVIEWS IN FISHERIES SCIENCE & AQUACULTURE 217

bass are naturally more abundant than smallmouthbass, thus explaining differences in recruitment andthe associated level of abundance increases observedwhen their fisheries are predominantly CR. Inaddition, smallmouth bass may be subjected tohigher hooking mortality rates than largemouth bass(Edwards et al. 2004a, 2004b) and climate change mayfavor largemouth bass over smallmouth bass (Hansenet al. 2017).

Catch-and-release of muskellunge over the past30 years does not appear to have caused an increase inWisconsin statewide or Escanaba Lake muskellungerelative abundances. Rypel et al. (2016) showedno change in muskellunge electrofishing CPUEin Wisconsin during 1944-2012. Muskellunge(�762mm) abundance declined significantly during1987-2014 in Escanaba Lake despite exploitation ratesbeing� about 5% during 2006-2014 (Eslinger et al.2017) (Figure 3). These results suggest that muskel-lunge abundance may be relatively unresponsive toCR. Muskellunge are a long-lived, apex piscivore andnaturally exist at low densities in the absence of stock-ing. In Wisconsin CR muskellunge fisheries that arenot stocked, average muskellunge density is about0.1 fish/ha (Eslinger et al. 2017). Therefore, carryingcapacities for muskellunge may be naturally low andCR may not result in increased total mortality rates inthe absence of exploitation, particularly when hookingmortality is low (Landsman et al. 2011; Shaw, Sass,and Eslinger 2019). In contrast, natural mortality ratesin older largemouth bass may increase in CR fisheriesallowing recruitment and overall abundances toincrease at the expense of reductions in populationsize structure (Sass et al. 2018). For muskellunge, CRmay not result in elevated abundances because fishingmortality is the strongest component of total mortality(Shaw, Sass, and Eslinger 2019). Therefore,unexploited muskellunge populations tend to showimproved size structure, a stable age distribution, highadult survival, and low recruitment (Eslinger et al.2017; Shaw, Sass, and Eslinger 2019).

Walleye anglers are harvest-oriented, thereforeabundance data on unexploited or predominantly CRfisheries is rare (Gaeta et al. 2013). Escanaba Lake iswell studied and transitioned from a no length limit,bag limit, and closed season walleye fishery toan unexploited walleye fishery beginning in 2003(711mm minimum length limit, daily bag limit of onefish, no closed season) (Haglund, Isermann, and Sass2016). The expectation of increased walleye densitywith the removal of exploitation did not occur outsideof an increase in female density; adult and male

density did not change between the exploited andunexploited time periods (Haglund, Isermann, andSass 2016). Despite observed reductions in adult wall-eye density and an associated release of density-dependent constraints on growth and maturation rateswith exploitation (Schueller et al. 2005; Schueller,Hansen, and Newman 2008; Schmalz et al. 2011;Tsehaye, Roth, and Sass 2016; Sass and Shaw 2018),the elimination of exploitation may not lead tomarked improvements in walleye density as carryingcapacities for this species may be constrained similarto muskellunge (Mrnak et al. 2018). Low natural mor-tality rates may act to stabilize species-specific fishabundances despite CR when hooking mortality is lowand fishing mortality is the primary component oftotal mortality (Payer, Pierce, and Pereira 1989;Hansen, Fayram, and Newman 2011; Landsman et al.2011; Tsehaye, Roth, and Sass 2016; Mrnak et al.2018; Sass et al. 2018; Shaw, Sass, and Eslinger 2019).Abiotic and biotic factors such as water temperatureand forage availability have been shown to be predic-tors of walleye and muskellunge recruitment(Madenjian et al. 1996; Hansen et al. 1998; Beard,Hansen, and Carpenter 2003; Eslinger, Dolan, andNewman 2010; Shaw, Sass, and VanDeHey 2018),whereas stock-recruitment relationships for Florida,largemouth, and smallmouth bass remain elusive(Zipkin et al. 2008; Allen et al. 2011; Parkos et al.2013). This may highlight an additional difference inspecies-specific abundance responses to CR. Parentalguarding species such as largemouth bass may bemore responsive to CR, whereas recruitment of broad-cast spawners such as muskellunge and walleye maybe driven by highly variable abiotic and biotic factors.Parental guarding smallmouth bass abundanceresponses to CR may lie in between those of large-mouth bass and walleye/muskellunge due to therequirement of specific, and sometimes limited,spawning habitat (Rejwan et al. 1997). Nevertheless,intentional overharvest of a smallmouth bass popula-tion resulted in strong compensatory recruitment sug-gesting that CR (including bed fishing) may have littleto no influence on population-level recruitment inthis species (Zipkin et al. 2008; Jackson et al. 2015)like Florida and largemouth bass (Allen et al. 2011;Shaw and Allen 2016).

Catch-and-release and size structure, growth, andtrophy potential

Anglers, stakeholders, popular articles, the scientificliterature, and managers have often promoted CR to

218 G. G. SASS AND S. L. SHAW

improve size structure, growth rates, and trophypotential in fish populations. Species-specific levels ofdensity-dependence and variability in aquatic ecosys-tem productivity can act in concert to decrease theprobability of reaching these goals (Sass and Kitchell2005; Hansen et al. 2015; Gilbert and Sass 2016;Pedersen et al. 2018; Noring et al. 2019). Increasedabundances of largemouth bass due to CR, increasedgrowing degree days, and lake warming due to climatechange in north temperate inland lakes (Rypel et al.2016; Hansen et al. 2017) have resulted in reductionsin size structure and individual growth rates, withdeclines in maximum growth potential. Proportionalsize distributions (PSD-Q and PSD-P) (Neumannet al. 2012) declined in a small northern Wisconsinlake where largemouth bass were experimentally sub-jected to five years of CR angling (Sass et al. 2018). Inheavily exploited largemouth bass populations withrelative low population sizes, CR mortality has alsobeen shown to negatively effect population size struc-ture (Hessenauer et al. 2018). During 1944-2012, themean length of largemouth bass in Wisconsinincreased significantly, whereas mean maximumlength was unchanged; this time period also corre-sponded with a statistically significant increase inlargemouth bass electrofishing CPUE suggestinghigher relative abundances over time (Rypel et al.2016). According to Rypel et al. (2016), mean andmean maximum length of largemouth bass were about250 and 450mm, respectively. In contrast, meanlength of age-6 largemouth bass declined from about381 to 330mm during 1992-2011 suggesting density-dependent effects on growth (Hansen et al. 2015)(Figure 2). Largemouth bass size structure and indi-vidual growth rates have declined as a result of CR,protection during the spawning season, and associatedincreases in abundance. As a result, maximum growthpotential has also been reduced.

Although relative abundances of smallmouth bassin Wisconsin lakes have increased significantly overtime, this species appears to be more resilient to dens-ity-dependent constraints on growth (Rypel et al.2016). Mean length and mean maximum length forsmallmouth bass in Wisconsin increased significantlyduring 1944-2012 (Rypel et al. 2016). Mean length forsmallmouth bass increased from about 200 to 250mmand mean maximum length has increased from about350 to 400mm (Rypel et al. 2016). As discussedabove, smallmouth bass may naturally be a lowerdensity species than largemouth bass. As such, dens-ity-dependent constraints on growth have not been asstrong and CR (and potentially protection during the

spawning season) has increased abundances and max-imum growth potential for smallmouth bass in themidwestern USA.

In general, CR in muskellunge fisheries has led toimprovements in size structure (Rypel et al. 2016;Eslinger et al. 2017); however, further improvementsin size structure appear to be capped by density-dependent constraints on growth, continued stocking,high minimum length limits, and low daily bag limitsto reduce maximum growth and trophy potential(e.g., muskellunge >1219mm and >1270mm)(Gilbert and Sass 2016; Shaw, Sass, and Eslinger2019). For example, muskellunge PSD-762mm, PSD-965mm, and PSD-1067mm increased significantlyon Escanaba Lake during 1987-2014; a period whenCR increased (Eslinger et al. 2017; Shaw, Sass, andVanDeHey 2018) (Figure 4). Similarly, mean lengthand mean maximum length of muskellunge inWisconsin have rebounded over time presumably dueto CR (Rypel et al. 2016). In Wisconsin, mean lengthand mean maximum length of muskellunge capturedby electrofishing has increased from about 600 to800mm and 1000 to 1200mm, respectively. Faustet al. (2015) found that mean asymptotic lengths offemale and male muskellunge in northern Wisconsinwere 1265mm and 1102mm, respectively, whereasminimum ultimate lengths were lower at 1143mm forfemales and 864mm for males. Faust et al. (2015)concluded that female maximum growth potential wascommensurate with trophy muskellunge managementin Wisconsin (1270mm minimum length limit anda daily bag limit of one fish) and adequate to meetangler desires. This review suggests that the frequencyof trophy muskellunge has been compromised overtime by CR, trophy regulations, density-dependentconstraints on growth, and continued stocking in nat-urally reproducing muskellunge populations (Gilbertand Sass 2016; Shaw, Sass, and Eslinger 2019). Usingdata from a long-term muskellunge fishing contest inVilas County, Wisconsin, Gilbert and Sass (2016)observed a 102mm and 4.2 kg decline in the meanlength and weight of the top ten muskellunge regis-tered in the contest during 1964-2010, which encom-passed the transition of muskellunge anglers fromharvest-oriented to CR (Eslinger et al. 2017; Shaw,Sass, and Eslinger 2019) (Figure 4). Mean length ofthe top ten muskellunge registered in the contestdeclined from about 1327mm in 1964 to about1233mm in 2010 (Gilbert and Sass 2016) (Figure 4).Similarly, mean weight of the top ten muskellungeregistered declined from about 18 kg to 14 kg between1964 and 2010, respectively (Figure 4). Therefore, CR,

REVIEWS IN FISHERIES SCIENCE & AQUACULTURE 219

highly conservative regulations, and continued stock-ing may be artificially overinflating muskellunge den-sities, reducing trophy growth potential, and notachieving angler expectations in these fisheries(Gilbert and Sass 2016; Shaw, Sass, and Eslinger2019). Given the current CR in muskellunge fisheries,adoption of liberal regulations (e.g., no closed season,no bag limit, and no minimum length limit) andreductions or elimination of stocking would likely bet-ter serve angler desires (Gilbert and Sass 2016;

Eslinger et al. 2017; Sass, Rypel, and Stafford 2017;Shaw, Sass, and Eslinger 2019).

Walleye anglers are predominantly harvest-oriented(Gaeta et al. 2013); however, certain anglers also desireand focus on trophy walleye opportunities. In instanceswhere there has been little to no exploitation on a walleyefishery (e.g. Escanaba Lake, 2003-present), size structure,growth, and trophy growth potential responses havebeen like those observed for muskellunge under nearlyexclusive CR (Haglund, Isermann, and Sass 2016).

Figure 4. Trends in muskellunge (Esox masquinongy) proportional size distributions on Escanaba Lake, Wisconsin during1987–2014 (A) (Eslinger et al. 2017) and the top ten mean lengths (B) and weights (C) of muskellunge registered in theVilas County, Wisconsin Musky Marathon fishing contest during 1964-2010 (Gilbert and Sass 2016). Reprinted with permissionas G.G. Sass and/or S.L. Shaw was a coauthor on all cited articles.

220 G. G. SASS AND S. L. SHAW

Walleye angler and stakeholder expectations wouldanticipate a quality fishery with high potential for pro-ducing trophies. Because walleye growth rates have beenshown to be weakly density-dependent and more relatedto ecosystem productivity (Sass and Kitchell 2005;Pedersen et al. 2018; Noring et al. 2019), reductions inwalleye exploitation through CR may improve size struc-ture, but would not be predicted to improve maximumgrowth potential. Mean length of males increased signifi-cantly after the elimination of legal harvest of walleye onEscanaba Lake; however, mean length of females andPSD-510mm did not differ between the exploited andunexploited time periods (Haglund, Isermann, and Sass2016). Interestingly, density-dependent responses ofwalleye in growth (increased), maturation schedule(reduced), and productivity (reduced) have been clearunder high exploitation scenarios; however, responses toa lack of exploitation appear to be capped based on thecarrying capacity of the system similar to muskellunge(Scheuller et al. 2005, 2008; Schmalz et al. 2011;Rypel et al. 2018; Sass and Shaw 2018). Collectively,fish responses to CR appear to be species-specific andmay not align with angler desires.

Recently, attempts to reduce panfish (i.e., bluegill,crappie, yellow perch) harvest through conservative dailybag limits and seasonal restrictions have been experi-mentally implemented on several Wisconsin lakes as atool to improve size structure and provide the sameamount of fillet yield to harvest-oriented anglers in thesefisheries (Rypel 2015; Lyons et al. 2017). This reviewquestions whether the promotion of CR for panfish andthe implementation of more conservative harvest regula-tions is warranted given the observed 20mm meanlength improvement in bluegill population size structure(Rypel 2015) in lieu of the unintended potential conse-quences of stunting and imbalances in fish communities.Evaluation of Wisconsins experimental panfish regula-tions are ongoing and will shed light on the question ofwhether size structure improvements were achieved or ifundesired consequences (stunting, imbalances in fishcommunities) were more prevalent. Again, promotion ofCR and the perceived benefits are likely species-specificand angler desires and single-species management goalsmay conflict with overall ecosystem-based managementgoals in fisheries managed for multiple species (Hansenet al. 2015).

Catch-and-release recommendations

Responses of fishes to CR are species-specific

Acknowledgement of species-specific differences toCR have been discussed previously; however, it has

been in the context of what anglers can do to minim-ize hooking mortality and sublethal effects wheninformation for a species is lacking (Cooke and Suski2005). This review has shown that realized hookingmortality is lower than previously thought for severalspecies when mark-recapture studies are used forevaluation (Landsman et al. 2011; Cline et al. 2012;Sass et al. 2018; Shaw, Sass, and Eslinger 2019).Catch-and-release can have variable species-specificconsequences with some responses meeting managerand angler expectations, whereas others may not meetthe defined goals of promoting this practice. Becauseof species-specific responses to CR, the promotionof this angling practice should not be treated asa “one size fits all” strategy to achieve angler, stake-holder, and manager desires in fisheries. At the sametime, this review acknowledges that this perspectiveis Wisconsin- and north temperate lake-centric andspecies in different environments and habitats mayrespond differently to CR.

CR may not achieve angler, stakeholder, andmanager desires

This review has provided clear examples where CRhas not resulted in the desired outcome for anglers,stakeholders, and managers outside of conservation.For example, CR of largemouth bass (including bedfishing) has resulted in greater abundances at theexpense of improved size structure, individual growth,and maximum growth potential (Cline et al. 2012;Hansen et al. 2015; Rypel et al. 2016; Sass et al. 2018).Outside of conservation, which has not been an issuefor this species, largemouth bass responses to CR havebeen in stark contrast to what anglers, stakeholders,and managers have desired. Similarly, anglers andstakeholders have desired trophy muskellungefisheries. The promotion of CR, stocking, and ultra-conservative regulations have been used as tools toincrease trophy potential of muskellunge populations.Although population size structure may be improved(Rypel et al. 2016; Eslinger et al. 2017), over 30 yearsof CR has not resulted in a greater frequency oftrophy muskellunge (Gilbert and Sass 2016) (Figure4). In contrast, CR for smallmouth bass (includingbed fishing) has resulted in increased abundances,size structure, and trophy growth potential in thisnaturally lower-density, long-lived, and slow-growingspecies, which is aligned with angler, stakeholder,and manager desires (Rypel et al. 2016). In harvestedspecies such as walleye, the promotion of CRto increase abundances, size structure, and trophy

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growth potential has had little effect to achieve thesegoals (Haglund, Isermann, and Sass 2016; Mrnaket al. 2018). Therefore, if conservation is not an issue,it would make little sense to promote CR and the useof conservative regulations, which would therebyreduce harvest opportunities for anglers for no reason.In the case of such species, system productivity andthe fish community appear to dictate the abundanceresponse, or lack thereof, to CR (Haglund, Isermann,and Sass 2016; Mrnak et al. 2018). Anglers, stakehold-ers, and managers should use the best species-specificscience available to predict the outcomes of promotingCR. When species-specific information is not avail-able, appropriate well-designed experiments and mod-eling should be conducted to test for responses(Carpenter 1998; Cooke and Suski 2005; Haglund,Isermann, and Sass 2016).

CR may create imbalances in multi-species fisheries

Catch-and-release is aimed to conserve fish popula-tions, and many studies have been conducted toreduce hooking mortality and sublethal effects of theangling process (Cooke and Suski 2005; Arlinghauset al. 2007; Cooke and Schramm 2007; Keretz et al.2018); however, the ecology of fishes must not beoverlooked. Nearly exclusive CR (e.g. black bass, mus-kellunge; Gaeta et al. 2013; Gilbert and Sass 2016) hasthe potential to create imbalances in fish communitiesmanaged for multiple species. Unnecessary stockingon top of natural recruitment can also overinflate nat-ural abundances of certain species compounding theseimbalances. Declines in walleye natural recruitmentand adult abundances in the Ceded Territory ofWisconsin provide an example of where such imbal-ances may have occurred (Hansen et al. 2015; Hansenet al. 2017; Rypel et al. 2018; Embke et al. 2019;Sullivan et al. 2019). Although increases in growingdegree days due to climate change have contributed tothe increasing abundances of largemouth bass innorthern Wisconsin (Hansen et al. 2017), nearlyexclusive CR has also greatly aided this species (Gaetaet al. 2013; Hansen et al. 2015; Rypel et al. 2016;Miranda et al. 2017; Hessenauer et al. 2018; Sullivanet al. 2019). Anecdotal, followed by empirical, obser-vations have confirmed reciprocal trends in walleye(negative) and largemouth bass (positive) abundancesin northern Wisconsin prompting collaborative effortsto restore balance between both recreationally import-ant species through walleye stocking, directed research(e.g., whole-lake centrarchid removal experiment), and

allowing harvest of largemouth bass during thespawning period (Schnell 2014; Hansen et al. 2015;Kelling et al. 2016; Hansen et al. 2017; Sullivan et al.2019). Nevertheless, restoring balance may be difficultto achieve if CR for bass remains pervasive and wall-eye and largemouth bass interact under cultivation-depensation dynamics (Walters and Kitchell 2001;Schnell 2014; Miranda et al. 2017; Hessenauer et al.2018; Sullivan et al. 2019). These changes observed incommunity composition related to changes in fishingpractices have been referred to as community over-fishing. This circumstance occurs when selective andheavy fishing is applied to only a few species in thecommunity, for example, walleye (Ontario Ministry ofNatural Resources 1983). The species being harvestedexperiences depensatory mortality from the speciesthat are favored by the new set of circumstances (e.g.,largemouth bass and CR) (Ontario Ministry ofNatural Resources 1983). Continued stocking on topof natural reproducing muskellunge populations andthe pervasive CR of this species may further hinderefforts to rebalance these fish communities (Gilbertand Sass 2016; Shaw, Sass, and Eslinger 2019).Ultimately, anglers, stakeholders, and managers shouldconsider unintended consequences of CR by usingecosystem-based fisheries management approachesin lieu of single species assessments in systems wheremultiple species are managed to avoid potentialimbalances (Walters et al. 2005; Hansen et al. 2015).

CR as a coupled social-ecological system

Catch-and-release practices inherently couple socialand ecological systems and this recognition has beenacknowledged previously (Arlinghaus et al. 2007;Lynch et al. 2016, 2017; Hessenauer et al. 2016; Wardet al. 2016). Because CR is aimed to conserve fisheries,and many studies have been conducted to furtherreduce hooking mortality and the sublethal effects ofangling, this practice has become a social norm inmany fisheries and generally assumed to be beneficialto the target species. This review suggests that this isa narrow and generalized view (i.e., fisheries utopia)of the practice that may be short-sighted by not takinga long-term and ecosystem-based approach (Sass,Rypel, and Stafford 2017). When this social norm ispervasive for a species, CR has the potential to createimbalances in fish communities and angler, stake-holder, and manager desires may not be met. In suchcircumstances, how does one then change the socialnorm to swing the pendulum back toward balanceand desirable multi-species fisheries? This review

222 G. G. SASS AND S. L. SHAW

suggests that this is currently one of the biggest chal-lenges facing inland fisheries management. In general,fisheries management is intended to regulate commer-cial, subsistence, and angler exploitation to conservefisheries. Managers use harvest of fish by anglergroups to structure fish populations, meet manage-ment objectives, and ensure sustainability of fisheries.Common tools used to regulate harvest include quo-tas, seasonal restrictions, length limits, and bag limits.Mandatory harvest regulations are not generally usedin North American recreational fisheries outside ofinvasive species because animal welfare is not consid-ered as strongly as it is in Europe (Arlinghaus 2007).Social norms that differ from management tools andobjectives create a situation out of managerial controland dominated by pervasive angler and stakeholderbeliefs that CR is always a good thing for fisheries. Assuch, changing the social norm of CR where the prac-tice is creating imbalanced fisheries and not satisfyingangler and stakeholder desires must derive from thesegroups themselves and rely on the best science avail-able. Scientists and managers can contribute to theseefforts by conducting deliberate CR experiments thatare objective and indifferent to the advocacy of CRand by providing reviews such as the one presentedhere. Ultimately, when CR is not achieving fisheriesmanagement goals, this must be effectively communi-cated to anglers and stakeholders to restore a healthylevel of harvest because pervasive CR can hinder regu-lation effectiveness (Miranda et al. 2017; Hessenaueret al. 2018). Further, management agencies generallylack the people power and funding to conduct whole-lake fish removals to rebalance fisheries themselvesand thus rely on angler harvest to achieve such goals.

CR can be used to maintain fish in a safeoperating space (SOS)

This review is not aimed to downplay the importanceof CR in fisheries management. Catch-and-release,whether through conservative harvest regulations orangler behavior, has resulted in many fisheries conser-vation success stories. The purpose here is to highlightsituations where CR has trumped efforts to structurefish populations and to meet angler desires throughmore commonly used output and input controls(Miranda et al. 2017; Hessenauer et al. 2018). Animportant instance where the promotion of CR can beparticularly helpful is in situations where mechanismscausing fishery declines are outside of managerial con-trol, such as climate change or angler effort in openaccess fisheries. Climate change cannot be managed in

the short-term and has negatively affected some northtemperate inland lake fishes (e.g. cisco Coregonusartedi, walleye) (Jacobson et al. 2012; Hansen et al.2017). In such situations, the promotion of CR mayprovide resilience in conserving effected species bykeeping populations within a “Safe Operating Space”(Carpenter et al. 2017). This form of CR would beindependent of angler desires and solely focused onthe true conservation of a species. Catch-and-releasemay also be important for sustainable management inhigh effort, open access fisheries even when CR mor-tality and harvest is low.

Future research

In some cases, pervasive CR by anglers has causedimbalances in fish communities and not met angler,stakeholder, and manager desires. When CR domi-nates in a fishery, commonly used input and outputcontrols may be rendered useless to structure fisheriesand meet angler desires. Studies of CR in recreationalfisheries have almost exclusively focused on hookingmortality rates and methods to reduce these rates.Because natural mortality rates can be plastic depend-ing on the level of fishing mortality, it is recom-mended that future studies focus on deliberateexperiments to test for the influence of CR practiceson the target species, fish community, and aquaticecosystem. Such studies should not be conducted inlaboratories, mesocosms, or with the use of net pens,but as deliberate ecosystem experiments (Carpenter1998; Cline et al. 2012; Sass et al. 2018). Whole-lakestudies (e.g., ecosystem experiments) that collect pre-and post-manipulation baseline data, have a referencesystem, and are monitored for enough time to detectpotential effects are the only type of study that is rele-vant to the scale of management (Carpenter 1998).

Future research could also use modeling as a toolto describe the contagious or viral social behavior thatCR has become in order to find a way to reverse thesesocial norms where needed. Such modeling effortsmay provide a better understanding of the behaviorand subsequently identify mechanisms that have beenimportant in establishing it as a social norm and iden-tifying where this behavior can be reversed whenneeded. Promotion of harvest may also borrow fromEuropean sentiments of fish welfare to mandate har-vest (Arlinghaus 2007; Arlinghaus et al. 2007); how-ever, this mechanism should be viewed cautiously asto not swing the pendulum too far toward potentialoverharvest situations and the elimination of CR as aconservation and management tool (Rypel et al.

REVIEWS IN FISHERIES SCIENCE & AQUACULTURE 223

2016). Fisheries management is aimed to conservefisheries through the control of exploitation in openaccess systems (i.e., to avoid the tragedy of the com-mons) and is not designed to dictate or control anglerbeliefs. Thus, CR, which is outside of managerial con-trol, may be one of the biggest challenges facing fish-eries management in the 21st century.

Acknowledgements

Special thanks to the University of Wisconsin-Madison,Center for Limnology, Trout Lake Station for providinglodging when this review was conceived and written.

Data availability statement

Data available on request from the authors – The data thatsupport the findings of this review are available from thecorresponding author; GGS, upon reasonable request.

Funding

This review was primarily funded by the United States Fishand Wildlife Service, Federal Aid in Sportfish Restorationand the Wisconsin Department of Natural Resources, F-95-P, project FHNH. Additional funding and support wereprovided by the U.S. National Science Foundation undergrant 1716066.

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