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Influence of Smallmouth Bass Predation onRecruitment of Age-0 Yellow Perch in South DakotaGlacial LakesDaniel J. Dembkowskia, D. W. Willisa, B. G. Blackwellb, S. R. Chippsc, T. D. Baculad & M. R.Wuellnera

a Department of Natural Resource Management, South Dakota State University, Box 2140B,Brookings, South Dakota 57006, USAb South Dakota Department of Game, Fish and Parks, 603 East 8th Avenue, Webster, SouthDakota 57274, USAc Department of Natural Resource Management, South Dakota State University, Box 2140B,Brookings, South Dakota 57006, USA; and U.S. Geological Survey, South Dakota CooperativeFish and Wildlife Research Unit, Brookings, South Dakota 57007, USAd Indiana Department of Natural Resources, 4320 West Toto Road, North Judson, Indiana46366, USAPublished online: 13 Jul 2015.

To cite this article: Daniel J. Dembkowski, D. W. Willis, B. G. Blackwell, S. R. Chipps, T. D. Bacula & M. R. Wuellner (2015)Influence of Smallmouth Bass Predation on Recruitment of Age-0 Yellow Perch in South Dakota Glacial Lakes, North AmericanJournal of Fisheries Management, 35:4, 736-747, DOI: 10.1080/02755947.2015.1044629

To link to this article: http://dx.doi.org/10.1080/02755947.2015.1044629

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ARTICLE

Influence of Smallmouth Bass Predation on Recruitmentof Age-0 Yellow Perch in South Dakota Glacial Lakes

Daniel J. Dembkowski*1 and D. W. WillisDepartment of Natural Resource Management, South Dakota State University, Box 2140B, Brookings,

South Dakota 57006, USA

B. G. BlackwellSouth Dakota Department of Game, Fish and Parks, 603 East 8th Avenue, Webster,

South Dakota 57274, USA

S. R. ChippsDepartment of Natural Resource Management, South Dakota State University, Box 2140B, Brookings,

South Dakota 57006, USA; and U.S. Geological Survey, South Dakota Cooperative Fish and Wildlife

Research Unit, Brookings, South Dakota 57007, USA

T. D. BaculaIndiana Department of Natural Resources, 4320 West Toto Road, North Judson, Indiana 46366, USA

M. R. WuellnerDepartment of Natural Resource Management, South Dakota State University, Box 2140B, Brookings,

South Dakota 57006, USA

AbstractWe estimated the influence of predation by Smallmouth Bass Micropterus dolomieu on recruitment of age-0

Yellow Perch Perca flavescens in two northeastern South Dakota glacial lakes. We estimated a likely range inconsumption of age-0 Yellow Perch using Smallmouth Bass diet information from two time periods when age-0Yellow Perch constituted high (2008) and low (2012 and 2013) proportions of Smallmouth Bass diets, and basspopulation size estimates as inputs in a bioenergetics model. The proportion of age-0 Yellow Perch consumed by theSmallmouth Bass populations was determined by comparing estimates of consumption with estimates of age-0perch production. During 2008, age-0 Yellow Perch constituted between 0% and 42% of Smallmouth Bass diets byweight, whereas during 2012 and 2013, age-0 perch constituted between 0% and 20% of bass diets by weight.Across both lakes and time periods, production of age-0 Yellow Perch ranged from 0.32 to 1.78 kg¢ha¡1¢week¡1.Estimates of Smallmouth Bass consumption measured during the same intervals ranged from 0.06 to0.33 kg¢ha¡1¢week¡1, equating to consumption of between 1% and 34% of the available Yellow Perch biomass.Given current conditions relative to Smallmouth Bass abundance and consumption dynamics and production ofage-0 Yellow Perch, it does not appear that Smallmouth Bass predation acts as a singular factor limitingrecruitment of age-0 Yellow Perch in our study lakes. However, future research and management initiatives shouldrecognize that the long-term impact of Smallmouth Bass predation is not static and will likely fluctuate dependingon environmental (e.g., temperature) and biotic (e.g., trends in macrophyte abundance, predator and preypopulation structure and abundance, and predatory fish assemblage dynamics) characteristics.

*Corresponding author email: dan.dembkowski@uwsp.edu1Present address: Fish Propagation Science Center, Wisconsin Cooperative Fishery Research Unit, College of Natural Resources,

University of Wisconsin–Stevens Point, 800 Reserve Street, Stevens Point, Wisconsin 54481, USA.Received December 19, 2014; accepted April 20, 2015

736

North American Journal of Fisheries Management 35:736–747, 2015

� American Fisheries Society 2015

ISSN: 0275-5947 print / 1548-8675 online

DOI: 10.1080/02755947.2015.1044629

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Black basses, especially Largemouth Bass Micropterus sal-

moides and Smallmouth Bass M. dolomieu, are some of the

most commonly introduced species worldwide (Jackson

2002). In the United States, Smallmouth Bass are native to 23

states but have been stocked into an additional 22 states (Rahel

2000). Despite a long history of Smallmouth Bass introduc-

tions, formal evaluation of the ecological consequences of

these introductions has only recently begun (Jackson 2002).

As predators with opportunistic feeding habits (Brown et al.

2009), Smallmouth Bass have the potential to create novel

interactions with other fishes in the form of resource competi-

tion and predation. Relative to their effects on prey fishes,

introductions of Smallmouth Bass outside of their native range

have been linked with changes in trophic interactions within

native fish assemblages (e.g., Whittier et al. 1997; Vander

Zanden et al. 1999), restructuring of aquatic communities

(e.g., Vander Zanden et al. 1999), and changes in prey fish

population dynamics (e.g., Johnson and Hale 1977; Tonn and

Magnuson 1983; Pflug and Pauly 1984).

The magnitude of predatory impacts of Smallmouth Bass

on prey fish populations encompasses a broad spectrum, rang-

ing from minimal impacts to complete extirpation (e.g.,

Jackson 2002). In a study of the influence of predation by

the predatory fish assemblage on abundance of out-migrat-

ing salmonid smolts in John Day Reservoir on the Columbia

River, Vigg et al. (1991) found that introduced Smallmouth

Bass had the lowest mean seasonal consumption of salmo-

nids in the reservoir and consumed only 0.04 salmonid

prey/predator per day. Similarly, Fayram and Sibley (2000)

concluded that Smallmouth Bass predation on juvenile

Sockeye Salmon Oncorhynchus nerka had little impact on

an observed temporal decrease in Sockeye Salmon abun-

dance in Lake Washington, Washington. Conversely, the

presence of Smallmouth Bass in a range of Minnesota lakes

corresponded to a substantial reduction in abundance of

Walleye Sander vitreus, possibly attributed to predation of

Walleyes by the bass (Johnson and Hale 1977). Several

studies report extensive predation on native cyprinids by

Smallmouth Bass in Ontario lakes, with predation high

enough to cause extirpation of cyprinids in some systems

(Jackson 2002; Jackson and Mandrak 2002).

In South Dakota, Smallmouth Bass are native to the Minne-

sota River drainage basin but were introduced outside of their

native range beginning in the mid-1980s to diversify angling

opportunities (Milewski and Willis 1990; Berry and Young

2004). Due to their short residence time, relatively little is

known about Smallmouth Bass interactions with native spe-

cies in most South Dakota waters. However, several studies

have examined potential predatory and competitive interac-

tions between Smallmouth Bass and Walleye in field and labo-

ratory settings as a response to angler concerns that bass were

preying on juvenile Walleyes and reducing abundance, condi-

tion, and growth rates of adult Walleyes via interspecific

resource competition (Wuellner et al. 2010, 2011a, 2011b;

Galster et al. 2012). While few studies have assessed the

impact of Smallmouth Bass predation on prey fish populations,

several have documented bass food habits in state waters (e.g.,

Lott 1996; Blackwell et al. 1999; Bacula 2009), which may

provide insight into potential predatory impacts. For example,

Bacula (2009) found that age-0 Yellow Perch Perca flavescens

comprised between 24% and 82% of Smallmouth Bass diets

by number and between 27% and 42% by weight across a

range of eastern South Dakota glacial lakes, prompting con-

cern that bass predation may negatively influence perch popu-

lations in systems where both species co-occur.

Yellow Perch are an important ecological and recreational

component of fish assemblages in South Dakota and through-

out much of their range (Hansen et al. 1998; Blackwell et al.

1999; Mayer et al. 2000; Gigliotti 2007; Isermann et al.

2007). Recruitment patterns of Yellow Perch are typically

characterized as erratic or inconsistent and exhibit a large

degree of interannual variation in year-class strength

(Anderson et al. 1998; Isermann et al. 2007; Isermann and

Willis 2008). Several studies have attempted to link interan-

nual variation in Yellow Perch year-class strength with abiotic

and climatological factors, but a large amount of variation

remains unexplained (e.g., Ward et al. 2004; Isermann and

Willis 2008; Jansen 2008). The findings of Bacula (2009)

prompted concern that Smallmouth Bass predation upon age-0

Yellow Perch may contribute to interannual variation in perch

year-class strength and ultimately act as a factor limiting

recruitment of Yellow Perch to sizes preferred by the recrea-

tional fishery. To address these concerns, we estimated the

potential impact of Smallmouth Bass predation on recruitment

of age-0 Yellow Perch (indexed as the abundance of age-0

perch) in two northeastern South Dakota glacial lakes. Specifi-

cally, we used Smallmouth Bass population size and diet infor-

mation as inputs in a bioenergetics model to estimate a likely

range in consumption of age-0 Yellow Perch. Consumption

estimates were compared with estimates of age-0 Yellow

Perch production to estimate the proportion of the age-0 perch

cohort consumed by the Smallmouth Bass populations.

METHODS

Study lakes.—This study was conducted on two glacial

lakes in northeastern South Dakota during 2012 and 2013.

Pickerel Lake (Day County) was sampled during May–Sep-

tember 2012, and Clear Lake (Marshall County) was sampled

during the same months in 2013. Pickerel Lake has a surface

area of 397 ha, mean depth of 4.8 m, maximum depth of

12.5 m, and shoreline development index of 2.2; Clear Lake

has a surface area of 474 ha, mean depth of 3.8 m, maximum

depth of 6.7 m, and shoreline development index of 1.5 (Stu-

even and Stewart 1996; Stukel 2003). Natural riparian vegeta-

tion surrounding both lakes is limited owing to anthropogenic

development, and littoral habitat consists largely of bare rock

and sand substrate interrupted by stands of submerged (sago

SMALLMOUTH BASS PREDATION AND YELLOW PERCH RECRUITMENT 737

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pondweed Stuckenia pectinata and coontail Ceratophyllum

demersum) and emergent (bulrushes Scirpus spp. and cattails

Typha spp.) macrophytes.

Fish assemblages in both lakes are relatively simple and con-

sist primarily of centrarchids, percids, ictalurids, cyprinids, eso-

cids, moronids, and catostomids. Pickerel Lake is managed as a

panfish (i.e., Black Crappie Pomoxis nigromaculatus, Bluegill

Lepomis macrochirus, and Yellow Perch), Smallmouth Bass,

and Walleye fishery. Management objectives in Clear Lake

also focus on Walleye and Smallmouth Bass but also include

Largemouth Bass. Smallmouth Bass regulations on both lakes

include a five-fish daily bag limit and a 355–457-mm protected

slot limit, and only one bass over 457 mm is allowed.

Smallmouth Bass sampling and diets.—Smallmouth Bass

were sampled from Pickerel and Clear lakes every 7–10 d dur-

ing May–September using boat electrofishing conducted at

night. Littoral transects with rocky substrate were randomly

selected at the onset of sampling and were fixed for the study

duration at each lake based on the recommendations of Bacula

et al. (2011). Specifically, six 10-min transects were sampled

using a boat electrofisher equipped with a Smith-Root 7.5 GPP

pulsator unit (Smith-Root, Vancouver, Washington). Pulsed

DC electricity was cycled at 60 Hz with voltage output

adjusted according to the specific conductance at each lake

and sampling event to maintain a constant output of 7–9 A.

All Smallmouth Bass were measured for TL (mm) and wet

weight (W; g). Scales were removed from a subsample of

Smallmouth Bass for age, growth, and mortality analyses. Sag-

ittal otoliths were removed from Smallmouth Bass subjected

to incidental mortality for verification of scale-based age esti-

mates. Diet items were collected from a target of 20 Small-

mouth Bass per length-group in August and September. Diet

sampling was only conducted during August and September

because previous studies indicated that Smallmouth Bass con-

sumption of age-0 Yellow Perch peaked during the late sum-

mer and early fall (Bacula 2009). Therefore, we assumed that

the potential impact of Smallmouth Bass predation on age-0

Yellow Perch would be greatest during this time period.

Length-groups consisted of substock (SS; <180 mm TL),

stock-quality (S-Q; 180–279 mm TL), quality-preferred (Q-P;

280–349 mm TL), and preferred or greater (PC; �350 mm

TL) (Gabelhouse 1984). To determine the diets of Smallmouth

Bass � 180 mm TL stomach contents were collected using

gastric lavage, a nonlethal sampling technique found to effec-

tively remove over 90% of diet items from black basses

(Hakala and Johnson 2004). When large diet items (e.g., cray-

fish) became lodged within esophagi, diet items were instead

removed using long-handled forceps. Stomach contents were

flushed onto a 500-mm-mesh sieve, transferred to a sample

container, and preserved in 70% ethyl alcohol. Smallmouth

Bass < 180 mm TL were sacrificed and preserved in 70%

ethyl alcohol for laboratory processing because small esoph-

ageal diameter inhibited efficiency of the gastric lavage

device. Stomach contents were identified to the lowest

practical taxonomic level, enumerated, and weighed. Prey

fishes were identified to species (Becker 1983) and macroin-

vertebrates were identified to order (Merritt and Cummins

1996) when possible. Unidentifiable prey fishes, miscellaneous

fish parts, and uncommon fish species were pooled as “other

fish.” Unidentifiable fishes with obvious centrarchid body

morphology were pooled as “unidentified centrarchids.” All

prey items were blotted to remove excess liquid and weighed

to the nearest 0.01 g. Diets stratified by length-group were

characterized using mean proportion of dominant diet items

by wet weight (Hanson et al. 1997; Chipps and Garvey 2007;

Hartman and Hayward 2007). Wet weights of dominant diet

items were not corrected for potential weight loss resulting

from storage and preservation in 70% ethyl alcohol.

Smallmouth Bass population size.—Smallmouth Bass popu-

lation size was estimated using a multiple census mark–recap-

ture design. During electrofishing sampling events, all

Smallmouth Bass were marked with a pectoral fin clip. Elec-

trofishing samples were periodically supplemented with

catches of Smallmouth Bass from mini-fyke nets and hook-

and-line angling to reduce gear selectivity biases. Smallmouth

Bass population size at each lake was estimated using the

Schumacher–Eschmeyer modification of the Schnabel popula-

tion estimator (Schnabel 1938; Schumacher and Eschmeyer

1943; Ricker 1975):

N DXt

iD2.ni £Mi/

Xt

iD2.mi C 1/

; (1)

where N is the estimated population size, t is the number of

sampling events, ni is the number of fish caught in the ith sam-

ple, mi is the number of fish with marks caught in the ith sam-

ple, and Mi is the number of marked fish at large for the ith

sample. Population size was estimated for whole Smallmouth

Bass populations and stratified by individual age-groups

(based on age-group-specific sample sizes and recapture rates).

Age-groups were assigned based on month- and population-

specific age–length keys developed from ages estimated from

scale samples (Ricker 1975; Isely and Grabowski 2007), and

sizes were estimated using a scalar modification of the

Schnabel population estimator (Schnabel 1938; Schumacher

and Eschmeyer 1943; Ricker 1975). Asymmetrical 95% Pois-

son CIs surrounding each population or age-group size esti-

mate were computed following equations in Ricker (1975).

Given the extended sampling period (i.e., May–September)

needed to reach marking and recapture rates sufficient for esti-

mating population and age-group sizes, assumptions relative

to closed population size estimators may have been violated.

However, the Schnabel population estimator is robust to mod-

erate violations of assumptions regarding recruitment and

mortality (Ricker 1975). Additionally, violations of these

assumptions typically result in inflated population and

738 DEMBKOWSKI ET AL.

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age-group size estimates (Ricker 1975), which would assist in

providing insight into the greatest potential impact of Small-

mouth Bass predation on recruitment of age-0 Yellow Perch

when used as inputs in bioenergetics models.

Yellow Perch sampling and production estimation.—Age-0

Yellow Perch were sampled at each lake at 6–10-d intervals

during August and September using a 27.4 £ 1.8-m beach

seine (Dembkowski et al. 2012). Dembkowski et al. (2011)

found that age-0 Yellow Perch in Pickerel and Clear lakes

were distributed around patches of submerged vegetation in

nearshore areas and maintain a mostly demersal existence;

thus, vegetated shoreline areas were selected as seine sample

sites. During each seine haul, density (number/m2) and W (g)

of age-0 Yellow Perch were recorded.

Production of age-0 Yellow Perch in Pickerel and Clear

lakes was estimated using the instantaneous growth rate

method (Ricker 1975; Hayes et al. 2007):

PD GB; (2)

where P is the estimated production for a given cohort within a

specified interval, G is the estimated instantaneous growth rate

for the cohort from time t to time t C 1, and B is the estimated

arithmetic mean cohort biomass from time t to time t C 1. Pro-

duction of age-0 Yellow Perch in the vegetated littoral zone

was expressed as kilograms per hectare and was estimated dur-

ing the period between each seine sampling event. Variance

surrounding each estimate of production was estimated using

equations available in Hayes et al. (2007).

Bioenergetics modeling.—A bioenergetics model (Hanson

et al. 1997) was used to estimate age-specific consumption of

age-0 Yellow Perch by the Smallmouth Bass populations in Pick-

erel and Clear lakes. Bioenergetics models are highly applicable

and function on the basis of an energymass-balance equation:

CDGCRCFCU ; (3)

where C is consumption, G is growth, R is respiration, F is eges-

tion, and U is excretion. Units of G are derived through popula-

tion measurements; R is a function of fish weight, temperature,

diet, and activity; and F and U are functions of temperature and

diet (Kitchell et al. 1977; Hanson et al. 1997; Hartman and Hay-

ward 2007; Fincel et al. 2014). Consumption modeling mini-

mally requires a set of predator-specific physiological

parameters, predator food habits, abundance, growth, predator

and prey energy densities, and water temperature. Species-spe-

cific physiological parameters were included implicitly in the bio-

energetics modeling software (Fish Bioenergetics 3.0: Hanson

et al. 1997). Smallmouth Bass food habits and age-group abun-

dance were derived from our population sampling and diet

collections during August and September. To provide an esti-

mate of the greatest potential impact of Smallmouth Bass

predation on recruitment of age-0 Yellow Perch, the upper

bounds (i.e., upper 95% CI) of bass age-group size estimates

were used as bioenergetics inputs. Additionally, previous

Smallmouth Bass food habits from Pickerel and Clear lakes

in 2008 were compiled from Bacula (2009); population and

diet sampling were conducted in a standardized fashion in

2008, 2012, and 2013. To incorporate this additional tempo-

ral dimension of Smallmouth Bass diet composition, we sim-

ply replaced our empirical food habits data with those

collected during 2008; all other bioenergetics inputs were

derived herein (Smallmouth Bass growth and abundance and

water temperature data were not collected during the 2008

study). In doing so, we assumed that 2012 and 2013 were

representative of typical age-0 Yellow Perch production in

our study lakes and that seasonal Smallmouth Bass growth

and abundance and thermal regimes were relatively similar

among years.

Because estimates of Smallmouth Bass abundance were strat-

ified by age and food habits were stratified by length-group, diets

were assigned to age-groups based on mean TL at age. Growth

of Smallmouth Bass was expressed as the difference between

final and initial weight (g) of each age-group between August 1

and September 30. Energy density values (J/g wet weight) of

individual prey items were gathered from the literature for mac-

roinvertebrates (Cummins and Wuycheck 1971; Hill 1997) and

fish (Kitchell et al. 1974; Rice et al. 1983; Bevelheimer et al.

1985; Bryan et al. 1996; Liao et al. 2004). For pooled diet items

(e.g., unidentified centrarchids), energy density was estimated as

the mean energy density of taxonomically similar diet items of

known identity. Daily mean littoral water temperatures were

recorded throughout the growing season by stationary data log-

gers affixed to dock pilings or cinderblock mounts at various

depths throughout the littoral zone in each lake.

Age-specific consumption of age-0Yellow Perch by the Small-

mouth Bass populations was expressed as the weight (g) of age-0

perch consumed by the Smallmouth Bass age-groups on any given

day. Total consumption of age-0 Yellow Perch was estimated as

the sum of daily consumption estimates for each Smallmouth

Bass age-group throughout the modeling period. Because previ-

ous studies indicated that predation of age-0 Yellow Perch by the

Smallmouth Bass populations peaked during August and Septem-

ber (Bacula 2009), consumption of age-0 perch was modeled

from August 1 to September 30, the period that directly corre-

sponded to our estimates of age-0 Yellow Perch production. The

percentage of total consumption of age-0 Yellow Perch by each

Smallmouth Bass age-group was estimated to further investigate

intrapopulation consumption dynamics and estimate which bass

age-groups contributedmost to perch consumption.

RESULTS

Smallmouth Bass Diets

We collected and analyzed stomach contents from 151 and

159 Smallmouth Bass at Pickerel and Clear lakes,

SMALLMOUTH BASS PREDATION AND YELLOW PERCH RECRUITMENT 739

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respectively. At Pickerel Lake 21 (12%) stomachs were

empty, whereas at Clear Lake, 15 (9%) were empty. For

Smallmouth Bass subjected to incidental mortality, scale- and

otolith-based age estimates were in general agreement. Thus,

scale-based age estimates were appropriate for developing

age–length keys and assigning Smallmouth Bass to different

age-groups. Based on mean TL at age of Smallmouth Bass,

diets observed for SS fish were used for the 0–2 age-group, S-

Q diets were used for age-group 3, Q-P diets were used for

age-group 4, and PC diets were used for age-groups 5 and 6Cat both Pickerel and Clear lakes. Prey composition of Small-

mouth Bass varied temporally through August and September

and among age-groups within each lake (Figure 1).

At Pickerel Lake, diet composition of Smallmouth Bass

less than quality length (i.e., age-groups 0–2 and 3) was domi-

nated by prey fishes (i.e., Bluegill, Black Crappie, unidentified

centrarchids, and Yellow Perch), whereas crayfish comprised

a large proportion of stomach contents for bass larger than the

minimum preferred length (i.e., age-groups 5 and 6C;

Figure 1). During August, age-0 Yellow Perch constituted

between 5% and 13% of Smallmouth Bass diets by weight;

perch contributed most to diets of Smallmouth Bass less

than minimum stock length (i.e., age-group 0–2). During

September, age-0 Yellow Perch constituted between 0% and

11% of Smallmouth Bass diets by weight and contributed

most to diets of bass of stock-quality length (i.e., age-group 3).

FIGURE 1. August and September prey composition (percent by wet weight) for Smallmouth Bass from Pickerel Lake during 2012 and Clear Lake during

2013. Consumption is stratified by length-group (Gabelhouse 1984: substock [SS], <180 mm TL; stock-quality [S-Q], 180–279 mm TL; quality-preferred

[Q-P], 280–349 mm TL; greater than preferred [PC], >350 mm TL). Numbers above each length-group represent the number of fish used for diet analyses.

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Similar trends were observed at Clear Lake, with a decreasing

reliance on prey fish and an increasing reliance on crayfish

through progressive Smallmouth Bass age-groups (Figure 1).

During August, age-0 Yellow Perch constituted between 0% and

20% of Smallmouth Bass diets by weight; perch contributed

most to the diets of stock-quality length bass (i.e., age-group 3).

During September, age-0 Yellow Perch also contributed most to

diets of stock-quality length Smallmouth Bass, but only consti-

tuted between 0% and 9% of bass diets by weight.

The contribution of age-0 Yellow Perch to Smallmouth

Bass diets in Pickerel and Clear lakes was substantially

greater in 2008 (Figure 2). In Pickerel Lake, age-0 Yellow

Perch constituted between 3% and 42% of bass diets by

weight during August and between 0% and 17% of diets

during September. In Clear Lake, age-0 Yellow Perch con-

stituted between 17% and 36% of Smallmouth Bass diets by

weight during August and between 0% and 12% of diets in

September. In contrast to diet composition patterns observed

in Pickerel and Clear lakes in 2012 and 2013, proportions of

age-0 Yellow Perch in 2008 diets were spread relatively

evenly across Smallmouth Bass age-groups and were not

limited to younger and smaller bass.

Smallmouth Bass Population Size Estimates

Throughout the sampling period, 1,206 and 893 Small-

mouth Bass were marked at Pickerel and Clear lakes, respec-

tively. Recapture rates of Smallmouth Bass were 13% at

FIGURE 2. August and September prey composition (percent by wet weight) for Smallmouth Bass from Pickerel and Clear lakes collected during 2008. Con-

sumption is stratified by length-group (Gabelhouse 1984: substock [SS], <180 mm TL; stock-quality [S-Q], 180–279 mm TL; quality-preferred [Q-P],

280–349 mm TL; greater than preferred [PC], >350 mm TL). Numbers above each length-group represent the number of fish used for diet analyses.

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Pickerel Lake and 11% at Clear Lake. Estimated Smallmouth

Bass population size was 5,321 (95% CI D 4,529–6,288) at

Pickerel Lake and 3,695 (95% CI D 3,197–4,346) at Clear

Lake (Table 1). The population size estimate at Pickerel Lake

yielded population density and standing crop estimates of 13.7

fish/ha and 3.3 kg/ha, respectively. At Clear Lake, population

density was 7.7 fish/ha and standing crop was 3.5 kg/ha.

Based on sample size and recapture rates among age-groups

at each lake, sizes were estimated for Smallmouth Bass age-

groups 0–2, 3, 4, 5, and 6C. At Pickerel Lake, the 0–2 age-

group was most abundant, followed by a trend of decreasing

abundance of age-groups 3 through 6C (Table 1). A similar

trend was observed at Clear Lake, but abundance of age-

groups 5 and 6C was increased relative to abundance of age-

group 3 (Table 1). Although 95% CIs varied substantially for

age-group size estimates at each lake, there were no instances

when the CIs encompassed zero.

Age-0 Yellow Perch Production

Seine sampling was conducted on nine occasions each at

Pickerel Lake in 2012 and Clear Lake in 2013. At Pickerel

Lake, density of age-0 Yellow Perch in seine hauls decreased

from 2.29 § 2.92 perch/m2 (mean § SD) on August 2 to

0.68 § 0.72 perch/m2 on October 4. Throughout the sampling

period, W of age-0 Yellow Perch increased from 0.91 §0.14 g to 2.94 § 0.67 g. At Clear Lake, density of age-0

Yellow Perch in seine hauls decreased from 4.06 § 3.89

perch/m2 on July 30 to 0.46 § 0.47 perch/m2 on October 2.

Weight of age-0 Yellow Perch increased from 0.68 § 0.10 g

to 2.46 § 0.52 g throughout the sampling period.

Production of age-0 Yellow Perch was estimated during eight

intervals at each lake ranging temporally from 6 to 10 d (hereaf-

ter, week) during August and September. At both lakes, weekly

production estimates were variable but showed a general

decrease from early August through late September (Figures 3,

4). Estimates of age-0 Yellow Perch production were comparable

between both lakes, with production estimates at Pickerel Lake

ranging from 0.48 kg¢ha¡1¢week¡1 (variance: § 0.07 kg¢ha¡1¢week¡1) to 1.38 kg¢ha¡1¢week¡1 (§0.29 kg¢ha¡1¢ week¡1)

(Figure 3) and production estimates at Clear Lake ranging from

0.32 kg¢ha¡1¢week¡1 (§0.04 kg¢ha¡1¢week¡1) to 1.78 kg¢ha¡1¢week¡1 (§0.29 kg¢ha¡1¢week¡1) (Figure 4).

Bioenergetics Modeling

Consumption of age-0 Yellow Perch by the Smallmouth

Bass populations was modeled during the same eight intervals

over which perch production was estimated. At Pickerel Lake

during 2012, weekly population-level consumption of age-0

Yellow Perch (estimated as the sum of daily age-specific con-

sumption) ranged from 0.01 to 0.10 kg¢ha¡1¢week¡1

(Figure 3), equating to consumption of between 1% and 7% of

available perch biomass. When 2012 diets were replaced in

the model with those from 2008 (the period when age-0 perch

constituted a substantially greater proportion of Smallmouth

Bass stomach contents), weekly consumption of age-0 Yellow

Perch ranged from 0.06 to 0.33 kg¢ha¡1¢week¡1 (Figure 3),

equating to the consumption of between 7% and 34% of avail-

able perch biomass. At Clear Lake during 2013, weekly popu-

lation-level consumption of age-0 Yellow Perch ranged from

0.02 to 0.04 kg¢ha¡1¢week¡1 (Figure 4), equating to the con-

sumption of between 2% and 6% of available perch biomass.

When 2013 diets were replaced in the model with those from

2008, population-level consumption ranged from 0.11 to

0.28 kg¢ha¡1¢week¡1 (Figure 4), equating to between 14%

and 34% of available age-0 Yellow Perch biomass.

Intrapopulation consumption dynamics of age-0 Yellow

Perch by Smallmouth Bass differed between the periods when

age-0 Yellow Perch constituted high (2008) and low (2012

and 2013) proportions of Smallmouth Bass diets. During 2012

and 2013 at Pickerel and Clear lakes, greater than 80% of the

total age-0 Yellow Perch consumption was from Smallmouth

Bass < 280 mm TL (Table 2). Conversely, consumption of

age-0 Yellow Perch during 2008 was spread relatively equally

throughout the Smallmouth Bass age-groups, and bass

> 280 mm TL were responsible for greater than 50% of the

total age-0 perch consumption (Table 2).

DISCUSSION

Given current conditions relative to Smallmouth Bass abun-

dance and consumption dynamics and production of age-0

Yellow Perch, it does not appear that Smallmouth Bass act as

a singular substantial factor limiting recruitment of age-0

TABLE 1. Smallmouth Bass population and age-group abundance estimates

for Pickerel Lake, South Dakota, during 2012 and Clear Lake, South Dakota,

during 2013. Population and age-group abundances were estimated using the

Shumacher–Eschmeyer modification of the Schnabel population estimator

(Schnabel 1938; Schumacher and Eschmeyer 1943; Ricker 1975). Numbers in

parentheses represent asymmetrical 95% Poisson CIs.

Age (years) Size estimate

Pickerel Lake

0–2 3,347 (2,636–4,559)

3 1,185 (954–1,565)

4 563 (397–901)

5 155 (93–348)

6C 64 (35–192)

Population 5,321 (4,529–6,288)

Clear Lake

0–2 1,361 (1,013–1,985)

3 935 (753–1,213)

4 319 (222–524)

5 441 (264–993)

6C 735 (490–1,323)

Population 3,695 (3,197–4,346)

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Yellow Perch in our study lakes. Few other studies have exam-

ined consumption dynamics of Smallmouth Bass populations

relative to prey production. Liao et al. (2004) found that

Smallmouth Bass consumption in Spirit Lake, Iowa, concen-

trated mainly on Yellow Perch and that summer and fall con-

sumption rates of perch ranged from approximately 0.08 to

0.38 kg/ha during 1995–1997, which are comparable with our

findings. However, Liao et al. (2004) did not estimate Yellow

Perch production or availability and thus were unable to con-

clude whether Smallmouth Bass predation represented a sub-

stantial influence on the perch population.

The extent of predatory interactions is likely dictated by the

degree of spatial and temporal overlap between Smallmouth

Bass and age-0 Yellow Perch (sensu Cushing 1975, 1990;

Chick and Van Den Avyle 1999; Romare et al. 2003;

Beauchamp et al. 2004). Dembkowski et al. (2011) found that

age-0 Yellow Perch in our study lakes were distributed in litto-

ral areas primarily around patches of submerged macrophytes

and maintained a mostly demersal existence. In contrast,

Smallmouth Bass generally occupy sites with cobble sub-

strates devoid of macrophytes (George and Hadley 1979; Ran-

kin 1986; Olson et al. 2003; Brown et al. 2009). However,

Smallmouth Bass do exhibit cover-seeking behavior at all life

stages and show no preference for cover type (e.g., Hubert and

Lackey 1980; Edwards et al. 1983), and juvenile bass

(i.e., <180 mm TL) were periodically sampled from vegetated

areas during age-0 Yellow Perch sampling efforts (D. J.

Dembkowski, unpublished data).

Temporally, consumption of age-0 Yellow Perch by Small-

mouth Bass in eastern South Dakota lakes increased in August

and September compared with May, June, and July (Bacula

2009). Temporal patterns in the occurrence of age-0 Yellow

Perch in Smallmouth Bass diets may have been a result of hab-

itat segregation between juvenile perch and bass in early

and mid-summer months or because perch in August and

September had grown to a size large enough to be used by

FIGURE 3. Weekly production of age-0 Yellow Perch and corresponding consumption of age-0 Yellow Perch by Smallmouth Bass at Pickerel Lake during

August and September, 2008 and 2012. Smallmouth Bass consumption was modeled during the same intervals in which age-0 Yellow Perch production was esti-

mated. Error bars represent variance as estimated by equations provided in Hayes et al. (2007).

SMALLMOUTH BASS PREDATION AND YELLOW PERCH RECRUITMENT 743

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bass as prey items and to be energetically profitable enough to

outweigh the energetic costs associated with pursuit and han-

dling (i.e., optimal forage: MacArthur and Pianka 1966).

Regardless, our results showed a general decrease in consump-

tion of age-0 Yellow Perch by Smallmouth Bass from August

to September, and we hypothesize that this trend continues

throughout later autumn months. Additionally, it is likely that

some age-0 Yellow Perch will outgrow gape limitations of a

proportion of the smaller Smallmouth Bass by the end of the

growing season, thus reducing their relative vulnerability to

predation (e.g., Hambright 1991; Schake et al. 2014). For

example, a cursory analysis evaluating trends in relative vul-

nerability of Yellow Perch to Smallmouth Bass predation in

our study lakes estimated that vulnerability to predation begins

to decrease once perch reach approximately 50 mm TL

(Dembkowski, unpublished data). Age-0 Yellow Perch

FIGURE 4. Weekly production of age-0 Yellow Perch and corresponding consumption of age-0 Yellow Perch by Smallmouth Bass at Clear Lake during

August and September, 2008 and 2013. Smallmouth Bass consumption was modeled during the same intervals in which age-0 Yellow Perch production was

estimated. Error bars represent variance as estimated by equations provided in Hayes et al. (2007).

TABLE 2. Age-specific consumption estimates (kg/ha) of age-0 Yellow Perch during August and September by Smallmouth Bass in Pickerel and Clear lakes,

South Dakota. Numbers in parentheses represent the percent of total consumption of age-0 Yellow Perch by each Smallmouth Bass age-group. Total consumption

was estimated as the sum of weekly age-specific consumption values across all sample dates.

Lake Year Age 0–2 Age 3 Age 4 Age 5 Age 6C Total

Pickerel 2008 0.34 (26) 0.32 (24) 0.04 (3) 0.37 (28) 0.26 (20) 1.33 (100)

Pickerel 2012 0.08 (38) 0.10 (43) 0.02 (9) 0.01 (5) 0.01 (5) 0.23 (100)

Clear 2008 0.07 (4) 0.27 (17) 0.20 (13) 0.41 (26) 0.63 (40) 1.57 (100)

Clear 2013 0.07 (29) 0.16 (67) 0.01 (4) 0 (0) 0 (0) 0.24 (100)

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collected during seining efforts in late September ranged from

approximately 65 to 90 mm TL. Thus, a proportion of these

Yellow Perch were theoretically invulnerable to predation by

smaller Smallmouth Bass. Given the relatively late appearance

of age-0 Yellow Perch in Smallmouth Bass diets in the grow-

ing season, the decreasing trend in consumption of age-0 perch

throughout the study duration observed herein, and the poten-

tial for age-0 perch to outgrow bass gape limitations by the

end of the growing season, it is likely that the duration of sub-

stantial predation on age-0 perch by bass is relatively short.

This short duration of predation, combined with the relatively

low degree of spatial overlap between Smallmouth Bass and

age-0 Yellow Perch, likely reduces the probability of bass pre-

dation imposing greater limitations upon the perch populations

in our study lakes.

The relative influence of Smallmouth Bass predation may

also vary depending on the availability of age-0 Yellow Perch.

Because we lacked age-0 Yellow Perch production data from

2008, we could not make direct insights relative to the func-

tional response of Smallmouth Bass predation to variation in

age-0 perch availability. However, Yellow Perch populations

in the northern Great Plains are often characterized by a large

degree of interannual variation in recruitment (e.g., Isermann

et al. 2007; Isermann and Willis 2008). Thus, availability of

age-0 Yellow Perch and subsequent consumption by Small-

mouth Bass may vary substantially from year to year. Given

the opportunistic feeding behavior of Smallmouth Bass

(Brown et al. 2009), it is likely that bass predation pressure is

positively related to prey availability. If it is a function of prey

availability, Smallmouth Bass predation may only strongly

influence Yellow Perch year-class strength periodically due to

erratic perch recruitment patterns (resulting in variable inter-

annual abundance of prey items; Sanderson et al. 1999;

Isermann and Willis 2008).

Differences in age-0 Yellow Perch availability may have

contributed to the observed differences in Smallmouth Bass

intrapopulation consumption dynamics when using diet

data from periods when perch comprised relatively high

(2008) and low (2012 and 2013) proportions of bass diets.

We hypothesize that differences in Smallmouth Bass intra-

population consumption dynamics between the two time

periods were mediated by trends in aquatic macrophyte

abundance. During 2008, submerged aquatic macrophyte

coverage in the study lakes was substantially less than that

in 2012 and 2013 (B. G. Blackwell, unpublished data).

Age-0 Yellow Perch vulnerability to predation may have

increased as a function of decreased protective habitat pro-

vided by submerged macrophytes, thereby explaining the

increased consumption of perch by Smallmouth Bass of all

sizes. Similarly, Gaeta et al. (2014) posited that consump-

tion of Yellow Perch by Largemouth Bass in a northern

Wisconsin lake may have increased due to a reduction in

protective habitat (coarse woody debris) that increased

predator–prey encounter rates.

It is important to note that we only estimated the potential

singular influence of Smallmouth Bass predation on recruit-

ment of age-0 Yellow Perch. Although we concluded that pre-

dation by Smallmouth Bass alone does not represent a

substantial factor limiting recruitment of age-0 Yellow Perch,

bass predation may have an additive or cumulative influence

when combined with consumption dynamics of the rest of the

predatory fish assemblages in these systems. For example,

although predation by Smallmouth Bass alone did not substan-

tially contribute to the annual loss of out-migrating salmonids

(Vigg et al. 1991), predation did substantially contribute to the

overall loss when consumption dynamics of the entire preda-

tory fish assemblage were considered collectively rather than

individually (Rieman et al. 1991). Other predators in our study

lakes included Walleye, Largemouth Bass, and Northern Pike

Esox lucius; all are opportunistic predators but are generally

more piscivorous than Smallmouth Bass (Becker 1983;

Blackwell et al. 1999). Blackwell et al. (1999) found that

Yellow Perch constituted between 30% and 50% of Walleye

and Northern Pike diets by weight in a study of seasonal diets

of top-level predators in a range of South Dakota glacial lakes

similar to those included in our study. If Walleye, Largemouth

Bass, and Northern Pike consumption of Yellow Perch in our

study lakes is of a similar magnitude, the influence of preda-

tion by the collective predatory fish assemblage (i.e., Small-

mouth Bass, Largemouth Bass, Walleye, and Northern Pike)

may act to limit recruitment of Yellow Perch. Alternatively, if

interspecific resource competition for age-0 Yellow Perch is

strong enough, the opportunistic feeding habits of Smallmouth

Bass may enable them to exploit other prey items (e.g., cray-

fish), thus lessening the collective impact of predation on age-

0 perch. Clearly, further research is warranted to estimate the

influence of the predation by the collective predatory fish

assemblages on recruitment of age-0 Yellow Perch in our

study lakes.

While results demonstrate that predation by Smallmouth

Bass does not represent a substantial factor limiting recruit-

ment of age-0 Yellow Perch under the observed conditions in

our study lakes, further research is needed to investigate poten-

tial changes in Smallmouth Bass consumption dynamics in

response to variable perch densities. Furthermore, and given

the presence of other predators in our study lakes, holistic

investigation of the impact of predation by the collective pred-

atory fish assemblage on prey fish population dynamics is

needed. If the impact of predatory interactions on recruitment

of age-0 Yellow Perch remains a concern in the future, manag-

ers may ultimately need to consider holistic predatory fish

assemblage management options to ensure the sustainability

of both predator (i.e., Smallmouth Bass, Largemouth Bass,

Walleye, and Northern Pike) and prey (i.e., Yellow Perch)

fisheries in South Dakota glacial lakes. Regardless, future

research and management initiatives should recognize that the

long-term impact of Smallmouth Bass predation is not static

and will likely fluctuate depending on environmental (e.g.,

SMALLMOUTH BASS PREDATION AND YELLOW PERCH RECRUITMENT 745

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temperature) and biotic (e.g., trends in macrophyte abundance,

predator and prey population structure and abundance, and

predatory fish assemblage dynamics) characteristics.

ACKNOWLEDGMENTS

Funding for this project was provided by Federal Aid in

Sport Fish Restoration funds (Project F-15-R, Study 1518)

administered by South Dakota Department of Game, Fish and

Parks (SDGFP) and South Dakota State University. We thank

SDGFP personnel (specifically T. Kaufman, S. Kennedy,

T. Moos, and R. Braun) and B. Graff, D. Benage, E. Gates,

J. Grote, M. Phayvanh, C. Schake, N. Scheibel, and B. Smith

for assistance in the field and laboratory. The South Dakota

Cooperative Fish and Wildlife Research Unit is jointly spon-

sored by the U.S. Geological Survey, SDGFP, South Dakota

State University, the Wildlife Management Institute, and the

U.S. Fish and Wildlife Service. Any use of trade names is for

descriptive purposes only and does not imply endorsement by

the U.S. Government.

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