habitat selection and abundance of young-of-year smallmouth bass in north temperate lakes

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Habitat Selection and Abundance ofYoung-of-Year Smallmouth Bass in NorthTemperate LakesPeter James Brown a & Michael Anthony Bozek aa U.S. Geological Survey, Wisconsin Cooperative FisheryResearch Unit , College of Natural Resources, University ofWisconsin–Stevens Point , Stevens Point, Wisconsin, 54481, USAPublished online: 09 Jan 2011.

To cite this article: Peter James Brown & Michael Anthony Bozek (2010) Habitat Selection andAbundance of Young-of-Year Smallmouth Bass in North Temperate Lakes, Transactions of theAmerican Fisheries Society, 139:4, 1247-1260, DOI: 10.1577/T09-049.1

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Habitat Selection and Abundance of Young-of-Year SmallmouthBass in North Temperate Lakes

PETER JAMES BROWN*1AND MICHAEL ANTHONY BOZEK

U.S. Geological Survey, Wisconsin Cooperative Fishery Research Unit, College of Natural Resources,University of Wisconsin–Stevens Point, Stevens Point, Wisconsin 54481, USA

Abstract.—Habitat use during early life history plays an important role in the ecology of smallmouth bass

Micropterus dolomieu in north temperate lakes. The highest levels of mortality occur during the first year of

life, and the habitat selected probably affects mortality. We used resource selection functions and abundance

data from two northern Wisconsin lakes to determine the habitats that influence the survival of smallmouth

bass. Coarse substrates were consistently important to both nesting locations and young-of-year smallmouth

bass. Young smallmouth bass used woody structure after swimming from their nests but disassociated

themselves from habitats with more complex woody structure by August. Nonwoody cobble areas offer

protection for young-of-year smallmouth bass without attracting predators, as woody habitats do. The decline

in the abundance of young-of-year smallmouth bass was best fit to an exponential decay function in woody

habitats, but in rock habitats it was linear. Habitat selection by young-of-year smallmouth bass shifts over

time, and the shift is linked to predation risk: woody habitats initially offer them an advantage with respect to

spawning but eventually provide their predators greater opportunities for ambush. This shift underscores the

importance of having a diversity of littoral habitats. This study provides the first quantifiable analyses

describing the habitat features selected by young-of-year smallmouth bass and links these descriptions to

population dynamics.

Habitat needs of young-of-the year smallmouth bass

Micropterus dolomieu have been well described when

fish are associated with nests, but descriptions are poor

after they leave the nest. Numerous studies have

provided descriptions of the habitat used by small-

mouth bass for nesting (Meehan 1911; Tester 1930;

Hubbs and Bailey 1938; Doan 1940; Webster 1945;

Pflieger 1966; Neves 1975; Schneider 1976; Lukas and

Orth 1995). This description of the spawning habitat

and chronology of spawning has been developed based

on general observations of the behavior of smallmouth

bass. Only recently has selection of nesting habitat

been quantified using comparisons between available

habitat and the habitat used for nesting. Bozek et al.

(2002) and Saunders et al. (2002) have provided a

more quantitative description of smallmouth bass

nesting habitat in north temperate lakes. However,

only general descriptions have been made of the habitat

used by young-of-year smallmouth bass after they

leave the nest (Okeyo and Hassler 1985; Bryan and

Scarnecchia 1992; Weaver et al. 1997; Brown et al.

2000). After yolk sack absorption, young-of-year

smallmouth bass remain proximal for several days,

while the male continues to defend the nest site for up

to 5 d (Hubbs and Bailey 1938; Ridgway 1988;

Wiegmann and Baylis 1995).

After swimming from the nest young-of-year small-

mouth bass are more difficult to track. Therefore,

indirect studies of littoral zone fish communities have

provided only general descriptions of the habitat used

by young-of-year smallmouth bass in lentic environ-

ments (Bryan and Scarnecchia 1992; Weaver et al.

1997; Brown et al. 2000). Young-of-year smallmouth

bass in lakes appear to use areas of shorelines that are

less complex (e.g., few macrophytes or woody

structure, Bryan and Scarnecchia 1992; Weaver et al.

1997). For example, young-of-year smallmouth bass

have been observed in higher densities associated with

sand and rock habitats compared with more structurally

complex woody habitats (Brown et al. 2000). The

habitat requirements of smallmouth bass during this

important life stage are poorly understood because no

studies have been specifically designed to describe the

habitat selected by young-of-year smallmouth bass in

lakes. Habitat selection, or the use of a specific habitat

type in greater proportion to its abundance, implies that

an animal is selecting an area for reasons other than

purely by chance (Johnson 1980; Manly et al. 1993;

Garshelis 2000).

Several approaches have been taken to model habitat

selection; however, logistic regression is ideal in this

case because it uses a binary response (presence or

absence) and continuous predictor variables (environ-

* Corresponding author: [email protected] Present address; Montana Cooperative Fishery Research

Unit, Department of Ecology, Montana State University,Bozeman, Montana 59717, USA.

Received March 9, 2009; accepted January 1, 2010Published online July 7, 2010

1247

Transactions of the American Fisheries Society 139:1247–1260, 2010American Fisheries Society 2010DOI: 10.1577/T09-049.1

[Article]

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mental characteristics, Press and Wilson 1978; Manly

et al. 1993). Logistic regression has been widely used

in studies of wildlife (Mace et al. 1996; Carroll et al.

1999) and fish (Parsons and Hubert 1988; Belaud et al.

1989) to determine the probability of resource

selection. Multiple logistic regression has been used

in studies of the early life history of smallmouth bass to

determine nest site selection (Bozek et al. 2002) and

the relative quality of a nest site (Saunders et al. 2002).

Habitat selection studies are valuable for quantifying

the probability of presence at a given site and for

identifying important characteristics of the habitat

selected, but they do not provide a measure of the

habitat’s quality. Measuring fish abundance trends in

different habitat types describes how well a habitat

suits the most basic aspect of life history, survival.

Combining studies of habitat selection with studies

of abundance provides insights into the relationship

between habitat and survival. This study calculates the

probability of resource selection and describes how

habitat affects population dynamics of young-of-year

smallmouth bass. The specific objectives were to

describe habitat selection by both nesting and young-

of-year smallmouth bass and to assess survival at four

time steps during early life history.

Methods

Study site.—This study was conducted on two

northern Wisconsin lakes, Big Crooked Lake, Vilas

County, and Yawkey Lake, Oneida County. Both lakes

are oligotrophic (i.e., very clear; Secchi disc depth, .5

m), have limited angling access, and do not contain

largemouth bass Micropterus salmoides (i.e., no

competition for spawning habitat). Big Crooked Lake

is a 276-ha lake with a shoreline length of 8.1 km, a

broad littoral zone, and a maximum depth of 11.6 m.

The riparian area is a second-growth, northern mixed-

hardwood forest, and the littoral zone is dominated by

sand with smaller areas of rock and trace amounts of

wood. Big Crooked Lake is encompassed by land

owned by a private club, Dairymen’s, Inc. Cottage

development is limited (a lodge and several cabins) and

encompasses approximately 25% of the shoreline.

Smallmouth bass fishing is restricted to catch-and-

release angling only. The fish community includes 20

species, with smallmouth bass, walleye Sander vitreus,

muskellunge Esox masquinongy, northern pike Esoxlucius, yellow perch Perca flavescens, rock bass

Ambloplites rupestris, mimic shiner Notropis volucel-lus, and white sucker Catostomus commersonii being

most abundant.

FIGURE 1.—Habitat used by smallmouth bass and available habitat with respect to the characteristics included in multiple

logistic regression models for nesting smallmouth bass in Big Crooked and Yawkey lakes. Substrate size-classes are as follows: 1

¼ fine organic matter, 2¼ silt (,0.2 mm in diameter), 3¼ sand (0.2–6.3 mm), 4¼gravel (6.4–76.0 mm), 5¼ cobble (76.1–149.9

mm), 6 ¼ rubble (150.0–303.9 mm), 7 ¼ small boulder (304.0–609.9 mm), 8 ¼ large boulder (.609.9 mm), and 9 ¼ bedrock

(consolidated parent material). Wood size-classes are as follows: small (,25.0 mm in diameter), medium (25.0–50.8 mm in

diameter, ,1.0 m long), and large (�50.8 mm in diameter, �1.0 m long).

1248 BROWN AND BOZEK

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Yawkey Lake is a 39-ha lake with a shoreline length

of 2.9 km, a steeper littoral zone than Big Crooked

Lake, and a maximum depth of 21.3 m. The riparian

area is a second-growth, northern mixed-hardwood

forest and the littoral zone habitat is dominated by sand

with small areas of rock and areas of wood. Yawkey

Lake is developed with 37 homes, of which 10 are used

by year-round residents, encompassing 75% of the

shoreline. There is no public access to the lake and

angling was monitored through a mail survey that

reported five smallmouth bass removed from the lake

during this study. There are eight fish species in the

lake of which smallmouth bass, walleye, and northern

pike dominate the littoral zone fish community.

To meet our objectives we described two aspects of

the young-of-year smallmouth bass population in these

lakes: habitat selection and abundance. We compared

the habitat used to that which was available four times

during summer to describe resource selection. We used

lake-wide measures of abundance at the egg, fry, and

fall young-of-year ontogenetic stages to determine the

population decline in the lakes. Also, we used counts of

fish in fixed areas of different habitat types to measure

the decline in population relative to habitat type.

Habitat selection.—Probability of resource selection

was calculated at four points in young-of-year small-

mouth bass development. The probability of selection

was calculated by paring information from four surveys

of habitat used (i.e., nesting, June, July, and August) to

information from a single survey of habitat availability

(Manly et al. 1993).

In May 2002, nests were located in the littoral zone

using three methods: (1) nests in water less than 1.0 m

deep were observed with polarized eye glasses from a

boat, (2) nests from 1.0 to 2.0 m deep were observed by

snorkeling or scuba diving, and (3) nests from 2.0 to

4.0 m deep were located by towing a scuba diver at

deeper depths (see Bozek et al. 2002; Saunders et al.

2002). No bass nests were located in water greater than

3.0 m deep during random surveys in these areas on

either lake, nor have smallmouth bass nests been

observed at these depths in similar surveys of proximal

lakes (Bozek et al. 2002; Saunders et al. 2002). Lakes

were searched for an entire day to avoid effects of light

angle, and lakes were visited in alternating days.

At each site where a nest was observed (i.e., used

site), a set of microhabitat characteristics was measured

within the nest rim or within a 1.0-m2 plot (for young-

of-year fish). For nests, depth, substrate percent

coverage, substrate embeddedness, percent coverage

of woody material, and distance to cover were

measured (Bozek et al. 2002; Saunders et al. 2002).

Depth was measured as the distance from the center of

the nest or 1-m2 plot to the water surface. Substrate was

classified using a modified Wentworth scale (Went-

worth 1922; Platts et al. 1983), as silt (,0.2 mm in

diameter), sand (0.2–6.3 mm), gravel (6.4–76.0 mm),

cobble (76.1–149.9 mm), rubble (150.0–303.9 mm),

small boulder (304.0–609.9 mm), large boulder

(.609.9 mm), and bedrock (consolidated parent

material). The percent coverage of each size-class

was visually estimated (Brown 2004). Substrates larger

than gravel were assigned an embeddedness code from

0 to 4 with a score of 0 indicating clean substrate with

no fine material (sand, silt) present in the top two layers

of the dominant larger substrates and a score of 4

indicating highly embedded substrates (Saunders 2001;

Brown 2004). Wood was divided into three size-

classes: small (,25.0 mm in diameter), medium (25.0–

50.8 mm in diameter, ,1.0 m long), and large (�50.8

mm in diameter, �1.0 m long, Brown 2004). The

percent coverage of each size-class was visually

estimated. In addition, the area within a 10-m radius

from the center of each nest was searched for protective

structure. Rocks larger than small boulder (�304 mm

diameter) and large pieces of woody structure (�50.8

mm in diameter, �1.0 m long) were considered to be

protective structures. The distance to structure items

was measured from the center of the nest to the closest

point on item.

After swim-up, the habitat used by young-of-year

smallmouth bass was quantified by a scuba diver along

random transects. Three post swim-up surveys were

conducted from 23 to 24 June, 21 to 23 July, and 20 to

23 August 2002. The amount of effort required to

survey the entire shoreline during these surveys was

time-prohibitive and, therefore, sampling was stratified

among six habitat types. In the littoral zone, four

habitat types were delineated: rock (sand–cobble

substrates with rock cover), wood (sand–silt substrates

with woody cover), sand (sand–silt substrates with no

cover), and macrophyte (sand substrates with macro-

phyte cover). In the pelagic zone, two habitat types

were delineated: deep bottom (bottom and transect 5.0–

10.0 m deep, silt substrate) and pelagic midwater

(bottom 10.0 m deep, transect 2.0 m deep). Each

habitat type was sampled proportional to its abundance

(i.e., length of time spent searching each habitat type

proportional to the area of shoreline covered by that

habitat type). Habitat use was evaluated by first

designating five random points in each habitat type

as start points. At each start point a scuba diver

searched along a compass bearing set 458 from

shoreline. To remain in the littoral zone the compass

bearing was changed by 908 when the diver reached 3-

m depth (�908 at the 3 m depth contour orþ908 at the

shoreline), making a zigzag through the littoral zone. In

pelagic areas, the diver searched from the randomly

SMALLMOUTH BASS IN NORTH TEMPERATE LAKES 1249

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determined point along a randomly determined com-

pass bearing for the appropriate time. The same

microhabitat characteristics measured at nest sites were

measured at each location where young of year

smallmouth bass were observed.

Available habitats for nesting and young-of-year

smallmouth bass were quantified along 100 randomly

placed transects around the shoreline. To select transect

locations the length of time to travel the entire lake

shoreline by boat was recorded in seconds and random

numbers were drawn that referred to the seconds

traveled from the start point. The lake was circumnav-

igated and transects were established when time

traveled equaled the random number. For each transect,

a 100-m tape measure was stretched perpendicular to

shore out to a depth of 3 m. Along this transect, the

same microhabitat measurements made at nest and

young-of-year bass sites were measured every other

meter.

Microhabitat measurements (independent variables)

were used to predict habitat use by smallmouth bass

(dependent variables) using logistic regression. Logis-

tic regression was used because the dependent variable

(site occupation) is binary (Cox and Snell 1989).

The �2 log likelihood statistic was used to test the

significance of each model. Lower values and higher

correct classification rates indicate improved fit

(Hosmer and Lemeshow 2000). Model building used

two steps. First, univariate models using each individ-

ual habitat characteristic were developed for nesting

and young-of-year smallmouth bass during June, July,

and August. Second, a multiple logistic regression

model was built from significant variables using

forward stepwise selection. At each step the Wald

chi-square statistic was used to determine significance

of including variables. Correct classification rates were

calculated to determine the predictive ability of each

model. If the number of used sites was less than 10% of

the number of available sites then correct classification

was calculated using a random subsample of the

available sites (available subsample size ¼ 10 3

number of used sites).

Abundance.—In both study lakes, population esti-

mates were made for the egg, pre-swim-up young-of-

year fish, and fall young-of-year ontogenetic stages.

The number of smallmouth bass eggs and pre-swim-up

young-of-year bass were visually enumerated by means

of scuba gear and a 36-cm 3 36-cm grid divided into 6-

cm 3 6-cm units. The same diver conducted all

surveys. At each nest, the number of eggs or fry in each

individual grid square was estimated and then all grid

square estimates were summed for an overall estimate

of total abundance. The grid was moved when eggs or

fry covered areas greater than 36 cm2. Estimates of fry

abundance were standardized for stage of development

and were estimated while fry were oriented on the top

of the substrate immediately before they dispersed

(Hubbs and Bailey 1938; Saunders et al. 2002). The

relationship between actual abundance and estimated

abundance (determined by manually removing all

individuals) was determined on six nests not included

in the study (number of eggs¼ 1.1012x� 40.923, r2¼0.99; number of fry ¼ 0.7499x þ 83.496, r2 ¼ 0.98).

Estimates of egg and fry abundance for each study nest

were corrected with these regression equations. The

corrected estimates of egg and fry abundance within

each study lake were summed to determine the lake-

wide population. Proportion of fish surviving at nest

sites was calculated as the number of fry divided by the

number of eggs.

In the fall, young-of-year smallmouth bass abun-

dance was estimated by first capturing young-of-year

bass with a bag seine and marking each with an upper

caudal fin clip. Young-of-year smallmouth bass were

then recaptured by electrofishing within 1 week of

marking. The population was estimated using the

adjusted Petersen method (Ricker 1975); the 95%confidence interval for the population estimate was

calculated by the Seber method (Seber 1982; Hayes et

al. 2007). Survival from swim-up fry to fall young of

year was calculated as the number of young of year at

swim-up divided by the number of young-of-year fish

in the fall.

Abundance of young-of-year smallmouth bass was

measured within specific habitat types throughout the

period of May through August. Abundance of young-

of-year smallmouth bass was evaluated among differ-

ent habitat types using weekly surveys over fixed areas

in both the littoral and pelagic zones. Three 1-m 3 10-

m plots were delineated with nylon string and anchored

at the corners in each littoral zone habitat type: rock,

wood, sand, and macrophyte. Littoral zone plots were

set in water 1.0–2.0 m deep. Pelagic plots were marked

with buoys. All observations were made by a single

diver; plots within a lake were observed at the same

time on a weekly basis. Species and general age-class

(young-of-year fish , 10 cm and adult fish � 10 cm)

were recorded for all fish observed. Counts of fish from

each of three replicate plots were averaged weekly to

develop a relative measure of abundance within each

habitat type. Trends in abundance were modeled using

three potential regression equations. Models using a

straight line, exponential decay, and exponential decay

with asymptote were fit to abundance estimates in each

habitat type. Within habitat types, Akaike’s informa-

tion criterion (AIC) values were compared to determine

the best model for predicting young-of-year small-

mouth bass abundance throughout summer. Again, the


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most parsimonious model was chosen when models

were within 2 AIC units (Burnham and Anderson

2002).

Results

Physical habitat in the littoral zones of Big Crooked

and Yawkey lakes differed with respect to morphology

and cover. The littoral zone of Big Crooked Lake was

largely broad and flat compared with the relatively

steep shorelines of Yawkey Lake (Figure 1). Big

Crooked Lake had little large woody structure (no

woody structure was recorded on transects), while 5%of available sites in Yawkey Lake had large woody

structure present (Figure 1). Big Crooked Lake had

abundant rock cover while there were few pieces of

substrate larger than small boulders in Yawkey Lake.

Both lakes had large areas of continuous sand substrate

and smaller areas of mixed sand–gravel and sand–

cobble substrates (Figure 1).

Habitat Selection

For nests and young-of-year smallmouth bass,

univariate logistic regression models associated these

sites with coarse substrates (e.g., gravel) and low

embeddedness. Other variables significantly related to

occupied nest or young-of-year bass sites included

depth and cover associated variables (e.g., distance to

coarse wood).

Nesting smallmouth bass in Big Crooked Lake

selected deeper sites having more sand, gravel, and

cobble substrates (Table 1; Figure 1). Correct classi-

fication of the best model was 94% with 84%predicting nest presence and 96% predicting nest

absence (Table 1). In Yawkey Lake, nesting small-

mouth bass selected sites with more gravel substrate

and less embedded gravel (Table 1; Figure 1). Correct

classification of the best model was 95% with 55%predicting nest presence and 98% predicting nest

absence (Table 1). Between lakes, nesting smallmouth

bass in Big Crooked Lake selected sites based on depth

and coarse substrates and nesting smallmouth bass in

Yawkey Lake selected sites based on proximity to large

wood and coarse substrates.

After leaving the nest in June young-of-year small-

mouth bass in Big Crooked Lake select deeper sites

with less embedded gravel and cobble (Table 2; Figure

2). Correct classification of the best model was 88%with 85% predicting young-of-year bass presence and

92% predicting absence (Table 2). After leaving the

nest young-of-year smallmouth bass in Yawkey Lake

selected sites that were closer to rock and farther from

woody cover with more sand substrate, less embedded

gravel, and more medium and large woody structure

(Table 2; Figure 2). Correct classification of the best

model was 84% with 81% predicting young-of-year

bass presence and 87% predicting absence (Table 2).

Between lakes, medium and large woody structure was

positively associated with young-of-year smallmouth

bass presence in Yawkey Lake; however, woody

structure did not enter the best model for Big Crooked

Lake. After leaving the nest young-of-year smallmouth

bass in Big Crooked Lake selected sites based solely on

substrate characteristics rather than a combination of

substrate and cover characteristics as in Yawkey Lake.

This would be expected because there are few wooded

areas in Big Crooked Lake.

By July, young-of-year smallmouth bass in Big

Crooked Lake selected sites having less embedded

gravel and more small and medium woody structure

(Table 3; Figure 3). The correct classification rate of

the best model was 94% with 41% predicting young-

of-year bass presence and 99% predicting absence

TABLE 1.—Best multiple logistic regression models of smallmouth bass nest presence in Big Crooked and Yawkey lakes.

Correct classification rates were calculated using a subsample of available sites.

Variable Coefficient (SE) Wald v2 P

Correct classification

Overall Presence Absence

Big Crooked Lake

Model 116.23 ,0.0001 94.1 83.9 95.8Depth 1.002 (0.207) 23.42 ,0.0001Sand substrate (%) 0.183 (0.064) 8.15 0.0043Gravel substrate (%) 0.255 (0.065) 15.55 ,0.0001Cobble substrate (%) 0.224 (0.065) 11.98 0.0005Intercept �26.800 (6.468) 17.17 ,0.0001

Yawkey Lake

Model 76.38 ,0.0001 94.6 55.4 98.4Gravel substrate (%) 0.023 (0.008) 8.54 0.0038Embeddedness of gravel �0.854 (0.240) 12.72 0.0004Distance to large

woody structure (m)�0.545 (0.111) 24.15 ,0.0001

Intercept 0.111 (0.887) 0.02 0.90


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TABLE 2.—Best multiple logistic regression models of young-of-year smallmouth bass presence in Big Crooked and Yawkey

lakes during June.

VariableCoefficient

(SE) Wald v2 P



Big Crooked Lake

Model 2030.91 ,0.0001 88.3 84.6 91.8Depth 0.690 (0.058) 142.97 ,0.0001Embeddedness of gravel �1.256 (0.046) 740.23 ,0.0001Embeddedness of cobble �0.513 (0.049) 112.49 ,0.0001Intercept 5.373 (0.171) 992.97 ,0.0001

Yawkey Lake

Model 579.65 ,0.0001 83.9 80.7 87.3Sand substrate (%) 0.019 (0.003) 57.73 ,0.0001Embeddedness of gravel �0.610 (0.060) 102.07 ,0.0001Medium woody structure (%) 0.326 (0.024) 180.02 ,0.0001Large woody structure (%) 0.378 (0.026) 219.41 ,0.0001Distance to large woody

structure (m) 0.222 (0.023) 95.45 ,0.0001Distance to rock .304 mm

in diameter (m) �0.194 (0.016) 148.50 ,0.0001Intercept �0.198 (0.315) 0.40 0.53


logistic regression models for smallmouth bass in Big Crooked and Yawkey lakes during June. Substrate and wood size-classes

are as described for Figure 1.


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(Table 3). In Yawkey Lake, young-of-year smallmouth

bass selected sites with more gravel substrate, less

embedded cobble, and more medium and large wood

(Table 3; Figure 3). Correct classification of the best

model was 88% with 27% predicting young-of-year

bass presence and 98% predicting absence (Table 3). In

both lakes, young-of-year smallmouth bass selected

sites based on a combination of coarse substrate

embeddedness and wood characteristics. Best models

for both lakes included two wood sizes with the

Yawkey Lake model including large wood rather than

small wood.

In August, young-of-year smallmouth bass in Big

Crooked Lake selected sites with less sand and gravel

substrates, more rubble substrates, and less embedded

gravel and cobble substrates (Table 4; Figure 4).

Correct classification of the best model was 95% with

66% predicting young-of-year bass presence and 98%


lakes during July.

VariableCoefficient

(SE) Wald v2 P



Big Crooked Lake

Model 322.08 ,0.0001 93.9a 41.2a 99.1a

Embeddedness of gravel �1.132 (0.066) 296.04 ,0.0001Small woody structure (%) 0.045 (0.006) 059.33 ,0.0001Medium woody structure (%) 1.616 (0.455) 12.61 0.0004Intercept 1.239 (0.230) 29.01 ,0.0001

Yawkey Lake

Model 183.77 ,0.0001 88.0 26.6 98.1Gravel substrate (%) 0.015 (0.003) 26.20 ,0.0001Embeddedness of cobble �0.381 (0.055) 48.18 ,0.0001Medium woody structure (%) 0.188 (0.025) 55.45 ,0.0001Large woody structure (%) 0.081 (0.025) 34.91 ,0.0001Intercept �0.727 (0.257) 7.98 0.0047

a Correct classification rates were calculated from a subsample of available sites.


logistic regression models for smallmouth bass in Big Crooked and Yawkey lakes during July. Substrate and wood size-classes

are as described for Figure 1.


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predicting absence (Table 4). In Yawkey Lake, young-

of-year smallmouth bass selected sites that were farther

from large woody structures with less sand and less

embedded gravel substrates (Table 4; Figure 4).

Correct classification of the best model was 93% with

55% predicting young-of-year bass presence, and 97%

predicting absence (Table 4). Between lakes, young-of-

year smallmouth bass selected sites based on the

presence of less-embedded substrates. In Yawkey

Lake, where wood was abundant, young-of-year bass

selected sites farther from coarse woody structure than

they did in Big Crooked Lake.

Abundance

Survival of young-of-year smallmouth bass at nest

sites was much higher than survival in the littoral zone


lakes during August.

VariableCoefficient

(SE) Wald v2 P



Big Crooked Lake

Model 376.51 ,0.0001 95.3a 66.0a 98.2a

Substrate (%) �0.013 (0.004) 9.26 0.0023Gravel substrate (%) �0.045 (0.007) 42.13 ,0.0001Rubble substrate (%) 0.053 (0.009) 34.04 ,0.0001Embeddedness of gravel �1.528 (0.134) 129.09 ,0.0001Embeddedness of cobble �0.351 (0.088) 15.94 ,0.0001Intercept 5.230 (0.605) 74.79 ,0.0001

Yawkey Lake

Model 238.74 ,0.0001 93.2 55.1 97.0Sand substrate (%) �0.022 (0.004) 39.16 ,0.0001Embeddedness of gravel �0.731 (0.077) 90.75 ,0.0001Distance to large

woody structure (m)0.296 (0.052) 31.91 ,0.0001

Intercept �1.541 (0.576) 7.15 0.0075

a Correct classification rates were calculated from a subsample of available sites.


logistic regression models for smallmouth bass in Big Crooked and Yawkey lakes during August. Substrate and wood size-

classes are as described for Figure 1.


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in both study lakes (Table 5). Survival was 44.6% in

Big Crooked Lake and 37.5% in Yawkey Lake from

the time eggs were laid to the time young-of-year bass

swam from the nest (Table 5). After leaving the nest

survival declined further with only 2.4% of the

smallmouth bass that swam away from Big Crooked

Lake nests and only 6.3% of the smallmouth bass that

swam away from Yawkey Lake nests surviving to fall

(Table 5; Figure 5).

A decline in abundance of young-of-year small-

mouth bass was evident along fixed plots in both lakes

(Figure 6). While abundance was highest in all habitat

types as young-of-year bass swam from their nests,

abundance declined through summer at different rates

in different habitat types. In both lakes, young-of-year

smallmouth bass used woody habitats and rock

habitats, but were not observed in open sand,

macrophyte, or pelagic areas throughout the summer.

Abundance of young-of-year smallmouth bass dropped

rapidly in woody habitats and steadily in rock habitats

(Figure 6). A straight line best described the decline in

young-of-year bass abundance in rock habitats while an

exponential decay function best fit the decline in

young-of-year bass in woody habitats (Table 6). Of

particular importance, the drop in abundance of young-

of-year smallmouth bass in woody habitats occurred at

the same time that adult smallmouth bass, yellow

perch, muskellunge, and walleyes were first observed

in woody habitats (Figure 6).

Discussion

The results of this study provide evidence that the

abundance of smallmouth bass is strongly influenced

during early life history and that habitat plays an

important role in this process. Resource selection

models show young-of-year smallmouth bass increas-

ingly select rock habitats and disassociate themselves

from woody habitats. Concurrent abundance measures

show an exponential decline in abundance in woody

habitats and slower linear decline in rock habitats.

Young-of-year smallmouth bass abundance unques-

tionably drops through the summer. But the behavior,

TABLE 5.—Numbers of young-of-year smallmouth bass in Big Crooked and Yawkey lakes at three life stages. Egg and fry

counts are absolute censuses. Fall estimates are 95% confidence intervals around the means. Survival values are not means, so

that confidence intervals could be calculated for them.

Lake Egg countPre swim-up

fry countFall young-of-year

estimateEgg

survivalYoung-of-year

survival

Big Crooked 329,621 147,124 3,456 6 2,865 44.6% 2.4%Yawkey 177,461 66,626 4,177 6 933 37.5% 6.3%

FIGURE 5.—Abundance of young-of-year smallmouth bass in Big Crooked (open circles) and Yawkey lakes (solid circles)

from nesting to August in 2002.


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and more specifically the resource selection, of young-

of-year bass has not been linked to abundance during

this time. Linking resource selection functions to

measures of abundance we demonstrated the key role

habitat plays in structuring year-class strength, and

hypothesized mechanisms explaining this decline.

Studies that incorporate bioenergetics and predation

risk would provide detail to our understanding of this

decline.

Nesting habitat features for smallmouth bass have

been studied in detail (Meehan 1911; Hubbs and Bailey

1938; Doan 1940; Pflieger 1966; Neves 1975; Bozek et

FIGURE 6.—Abundance of young-of-year smallmouth bass and presence of predators by habitat type in fixed plot surveys.

Young-of-year bass are represented by circles and predators by diamonds; open symbols represent rock habitats and filled

symbols woody habitats.


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al. 2002; Saunders et al. 2002). Previous descriptions

are of habitat preference; this study quantifies habitat

selection. While this study provides a model that can

predict the probability of nest or young-of-year small-

mouth bass presence, it is important to put the

mechanisms influencing survival into context. Survival

while young-of-year bass were on nests was 44.6% in

Big Crooked Lake and 37.5% in Yawkey Lake. The

amount of gravel at a site was a consistent predictor of

nest site presence in both Big Crooked Lake and

Yawkey Lake. Moreover, the embeddedness of gravel

and the amount of sand and cobble were variables that

predicted the presence of nest sites. Smallmouth bass

eggs are adhesive and are deposited on the surface of

substrates in the center of nests (Scott and Crossman

1973). This position, combined with continuous

fanning by the parental male, provides less opportunity

for fine sediments (e.g., silt) to occlude gas exchange

across the egg membrane (Eipper 1975). Additionally,

initial adhesion to substrates prevents eggs from

grouping together, limiting transfer of fungus growth,

which may occur as a result of egg mortality and can

spread through the entire egg mass (Webster 1945).

Selection of coarse nesting substrates with clean

interstices improves survival.

Structure adjacent to the nest is thought to help in

nest defense by providing fewer directions from which

a nest can be attacked. Previous studies have described

the presence of wood or boulder cover immediately

adjacent to nests (Beeman 1924; Hubbs and Bailey

1938; Saunders et al. 2002), and Hoff (1991) showed

that adjacent cover increased survival of smallmouth

bass on nests.

Water depth helped predict nest presence in Big

Cooked Lake, but not in Yawkey Lake. Variability in

nest depths between lakes may help explain the

contribution of depth into models. Nests in Yawkey

Lake were distributed more evenly between depths of

0–3 m while nests in Big Crooked Lake were more

often in slightly deeper water (1–3 m). Nests may not

have been built in the shallowest water (,1 m) in Big

Crooked Lake because the large fetch may create

extreme wave action that affects shallow nests.

Yawkey Lake, with a smaller fetch, is less conducive

to wave development and, therefore, less wave action

affects the nests.

Habitat plays an important role in survival on nests,

but parental protection probably plays the most

important role in survival. Parental protection lasts

from the time eggs are laid until several days after

young-of-year fish swim from the nest. Hubbs and

Bailey (1938) cited protection as the single most

important aspect of the smallmouth bass life cycle.

Survival on nests is lower if protection of the nest or

school stops even temporally (Ridgway and Shuter

1997).

Survival of young-of-year smallmouth bass from

nesting to fall was 2.4% in Big Crooked Lake and

6.3% in Yawkey Lake (Figure 5). Because of this low

survival rate, the period after young-of-year bass leave

the nest and before winter is probably the most

important factor in structuring populations of small-

mouth bass. Initially, young-of-year smallmouth bass

habitat selection was similar to nest-site selection and

may be characterized by coarse substrates and forms of

cover such as boulders or large woody structure.

However, 11 d after the last fry swam from their nests

the abundance of young-of-year smallmouth bass over

woody areas decreased dramatically in both lakes

(Figure 6). Parental protection was observed to last for

1 to 2 weeks after swim-up, which is similar to the

findings of Ridgway (1988). As parental protection

ended smallmouth bass abundance showed an expo-

nential decline in woody habitats and a linear decline in

rock habitats. While we did not quantify predation risk,

the difference in the shape of these declines suggests

TABLE 6.—Models of the abundance of young-of-year smallmouth bass (N) in rock and wood habitats through time (t). Model

coefficients (b) are not equivalent among models.

Model type Model r2Log

likelihoodAkaike information

criterion

Rock habitat

Linear Nt¼ b

0þ b

1t 0.84 �51.58 109.88

Exponential decay Nt¼ N

0� e�b

1t 0.87 �55.03 116.06

Exponential decaywith asymptote N

t¼ b

0þ N

0� e�b

1t 0.87 �54.98 117.97

Wood habitat

Linear Nt¼ b

0þ b

1t 0.77 �66.85 139.70

Exponential decay Nt¼ N

0� e�b

1t 0.84 �63.06 132.05

Exponential decaywith asymptote N

t¼ b

0þ N

0� e�b

1t 0.85 �62.52 133.03


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different predation risk in each habitat. We propose two

explanations for these differences; either predators are

more easily able to take young-of-year smallmouth

bass in woody habitats or young-of-year bass are more

easily able to avoid predators in rock habitats.

The first hypothesis (predator advantage) is support-

ed by several studies of predator–prey dynamics in

complex habitats. Newbrey et al. (2005) showed a

significant positive relationship between complexity of

large woody structure and abundance of large pisci-

vores. Further, Rodgers and Bergersen (1999) were

able to attract largemouth bass and northern pike to

artificial fishing structures. Savino and Stein (1989)

found that predator–prey encounter rates increased as

complexity increased. However, in the most complex

habitats (i.e., 1,000 stems/m2) prey were able to avoid

predators. Woody habitats in our study lakes were

characterized by low branching complexity and

probably provided a predatory advantage.

The second hypothesis (young-of-year fish avoid-

ance) is supported by direct study of young-of-year

smallmouth bass behavior. Previous studies have

shown that young-of-year smallmouth bass do not

necessarily prefer habitat that is structurally complex

(e.g., macrophytes) compared with habitats that

provide less cover (e.g., gravel and cobble). Young-

of-year smallmouth bass have been observed in sand

and rock habitats, completely segregated from adult

fish in Little Moose Lake, New York (Brown et al.

2000). Likewise, young-of-year smallmouth bass have

been observed in areas with lower macrophyte

complexity (Weaver et al. 1997) and areas where

macrophytes were removed from littoral zones leaving

only sand and gravel (Bryan and Scarnecchia 1992).

Because large predators are attracted to structurally

complex habitats and smallmouth bass select less

complex habitats, we suspect both of these factors

caused the differences observed in young-of-year

smallmouth bass abundance.

In this study, the abundance of young-of-year

smallmouth bass declined rapidly in complex habitats

and young-of-year smallmouth bass became less likely

to select complex habitats as they got older. These are

significant findings as large woody material in lakes is

positively correlated with fish abundance (Emery 1973;

Hubert and Lackey 1980; Newbrey et al. 2005) and are

often added to lakes to increase production (Hoff 1991;

Hunt and Annett 2002). Hoff (1991) increased the

survival of young-of-year smallmouth bass on nests by

adding woody structures that provided overhead cover

to parental males. Similarly, Hunt and Annett (2002)

found largemouth bass using supplemented woody

structure and natural woody structure equally for

spawning, and further, recommended that woody

structure be added to lakes to increase production.

Highly complex habitats offer protection to young-of-

year smallmouth bass in the form of escape refuge

(e.g., Savino and Stein 1989). However, highly

complex habitats become less complex rapidly and

these areas also concentrate predators (Newbrey et al.

2005). Therefore, lower young-of-year bass survival

would be expected in woody habitats where adult fish

congregate if the habitat is not complex. The

combination of habitat selection analyses and trends

in abundance show that areas of low habitat complexity

(i.e., sparse, slightly embedded gravel and cobble) are

important to young-of-year smallmouth bass.

The results of this research provide two important

findings with respect to north temperate lakes: (1)

young-of-year smallmouth bass survival on nests may

benefit from the proximity to large wood, but large

woody structure may not be beneficial to all life stages

of smallmouth bass; and (2) the abundance of habitat

that allows young-of-year bass to avoid predation

through summer is probably critical in shaping small-

mouth bass populations.

This study quantified resource selection probabilities

for young-of-year smallmouth bass and showed that

selection is linked to that survival. Moreover, these

changes may be related to differences in predation risk

within different habitats. It is clear that different habitat

types are important to smallmouth bass at different

times. Therefore, a diversity of habitats is important to

smallmouth bass populations and is probably important

to the ecology of lake systems. More importantly, the

functional differences in habitats and the role these

differences play in structuring populations underscores

the importance of maintaining diverse habitats in lake

littoral zones.

Acknowledgments

Melinda Brown, Case Brown, Pat Short, Rory

Saunders, Brian Achuff, Ben Torrinson, Matt Catalano,

Melissa Goerlitz, and Laura Rosenfield provided field

assistance. Mention of trade names is for information

purposes only and does not imply endorsement by the

U.S. Government.

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habitat selection and abundance of young-of-year smallmouth bass in north temperate lakes

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