North American Journal of Fisheries Management

ISSN: 0275-5947 (Print) 1548-8675 (Online) Journal homepage: http://www.tandfonline.com/loi/ujfm20

Effects of Variable Mortality and Recruitment

on Performance of Catch-Curve Residuals as
Indicators of Fish Year-Class Strength

Matthew J. Catalano , Andrew C. Dutterer , William E. Pine III & Micheal S.

Allen

To cite this article: Matthew J. Catalano , Andrew C. Dutterer , William E. Pine III & Micheal
S. Allen (2009) Effects of Variable Mortality and Recruitment on Performance of Catch-Curve
Residuals as Indicators of Fish Year-Class Strength, North American Journal of Fisheries
Management, 29:2, 295-305, DOI: 10.1577/M07-209.1

To link to this article: http://dx.doi.org/10.1577/M07-209.1

Published online: 08 Jan 2011.

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 North American Journal of Fisheries Management 29:295–305, 2009 [Article]
Ó Copyright by the American Fisheries Society 2009
DOI: 10.1577/M07-209.1

Effects of Variable Mortality and Recruitment on Performance of

Catch-Curve Residuals as Indicators of Fish Year-Class Strength
MATTHEW J. CATALANO,* ANDREW C. DUTTERER, WILLIAM E. PINE, III, AND MICHEAL S. ALLEN
Department of Fisheries and Aquatic Sciences, University of Florida,
7922 Northwest 71st Street, Gainesville, Florida 32653-3071, USA

Abstract.—We built a simulation model to assess the performance of catch-curve residuals as an index of
year-class strength for a short-lived and a long-lived fish life history type across a range of assumed values for
the variation in recruitment (CVR) and fishing mortality (CVF). The magnitude of CVR strongly influenced the
utility of catch-curve residuals in assessing year-class strength. The probability of finding a significant
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correlation between catch-curve residuals and true recruitment values exceeded 0.9 when CVR was greater
than 0.5 for the long-lived and greater than 1.0 for the short-lived life history types. This suggests that larger
recruitment values have a greater probability of being successfully ‘‘tracked’’ through the age structure.
Conversely, the magnitude of interannual variation in fishing mortality weakly influenced the performance of
catch-curve residuals. The inspection of individual catch-curve residuals relative to the known recruitment
values that produced them showed considerable scatter, indicating that the utility of this metric in assessing
individual year-class strength is small. Sensitivity analyses showed that the performance of catch-curve
residuals improved modestly with equal sampling vulnerability across ages and decreased slightly with
increased fishing mortality. Our results suggest that catch-curve residuals can serve as a rudimentary measure
of recruitment under ranges of recruitment and mortality variation similar to those frequently observed in field
studies.

Fish recruitment is commonly used as a metric with
which to assess the impacts of management actions or
changes in environmental conditions. Monitoring
recruitment can be costly because of the need for
repeated annual sampling to estimate recruitment time
series. A rapid assessment technique to associate
recruitment responses to these types of events would
aid fishery managers. One approach to evaluating the
correlation between abiotic or biotic factors and
recruitment patterns is the use of catch-curve residuals
to assess year-class strength (Maceina 1997). A catch
curve is a regression of the loge transformed number of
fish at each age on age, where the slope of the line
represents the instantaneous total mortality (Z; Ricker
1975). Maceina (1997) showed that residuals around a
catch curve can index the relative year-class strength of
cohorts, where positive residuals indicate above-
average recruitment levels and vice versa. If the
residual approach was reliable, fishery managers could
collect an age sample in a single year and make
inferences about relative year-class strength in the past,
and then relate these annual residuals to covariates such
as hydrology. Using catch curves of fish collected in a
single year is appealing to managers because of the
minimal data requirements and low cost of producing a
metric useful in evaluating recruitment trends. Increas-
* Corresponding author: catalm@ufl.edu
Received November 19, 2007; accepted July 22, 2008
Published online March 19, 2009
ingly this approach is being used to relate year-class
strength to environmental variables for several fish
species, including crappies Pomoxis spp. (Maceina and
Stimpert 1998; Isermann et al. 2002; Sammons et al.
2002; Maceina 2003), largemouth bass Micropterus
salmoides (Maceina 1997; Maceina and Bettoli 1998;
Bonvechio and Allen 2005), smallmouth bass M.
dolomieu (Slipke et al. 1998; Smith et al. 2005),
walleye Sander vitreus (Quist 2007), and white bass
Morone chrysops (DiCenzo and Duval 2002).
The utility of catch-curve residuals to assess year-
class strength is predicated on several assumptions that
may be difficult to satisfy in practice: (1) recruitment is
constant, (2) survival is constant among year-classes,
(3) survival is constant among ages, and (4) catches are
proportional to abundance (no gear selectivity). One
potential downfall to using catch-curve residuals is that
temporally variable natural and fishing mortality rates
could mask recruitment effects on residuals. Attribut-
ing unexplained variation in a catch curve solely to
recruitment variation may lead to bias in year-class
strength under variable survival, gear selectivity, or
both. Quist (2007) found that the relative abundance of
age-0 walleyes was strongly related to future catches at
ages 1 and 2 but poorly related to catches at ages 3 and
4. This author attributed the lack of association
between fish at young and older ages to fishing
mortality, where years with high recruitment attract
anglers leading to increased fishing mortality. Similar-
ly, changes in natural mortality can also affect the

295
 296
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FIGURE 1.—Sampling and harvest vulnerability with respect
to fish total length as a fraction of L‘ for the long-lived life
history type. This pattern also characterized the short-lived life
history type.
relationship between year-class strength and catch-
curve residuals. Missing year-classes preclude year-
class strength estimation using catch-curve residuals if
no fish of a given age are captured due to large natural
mortality events. These examples show that variable
mortality among years, cohorts, or both is possible,
which could make catch-curve residuals an unreliable
index of year-class strength.
The magnitude of recruitment variation within a
population may also affect the performance of catch-
curve residuals in describing recruitment history. The
influence of recruitment on age structure may be more
pronounced in populations exhibiting relatively high
degrees of recruitment variation when compared with
populations where recruitment variation is considered
low. Fish longevity could also influence the utility of
catch-curve residuals in indexing year-class strength
because long-lived fishes would have more observa-
tions (ages) in the catch curve than short-lived fishes.
The objective of this study was to evaluate the effects
of fish longevity, recruitment variation, and variation in
fishing mortality on reliability of catch-curve residuals
as a rapid measure of year-class strength using a
population simulation model.
Methods
We used a population simulation model to evaluate
the performance of catch-curve residuals as indicators
of year-class strength. The model represented an age-
structured fish population having density-dependent
CATALANO ET AL.
recruitment and lognormally distributed random re-
cruitment variation. We simulated a sampling and
aging process by which age–length samples were
collected from the simulated population and numbers at
age were estimated with an age–length key. The model
accounted for variation in the year signal in the
estimated numbers at age by incorporating annual
variation in fishing mortality, random errors in the
length–age sampling process, and aging errors. Catch
curves were constructed from the estimated age
frequencies, and residuals were obtained from a linear
regression of loge(catch) on age. Performance of catch-
curve residuals was evaluated by comparing the
residuals with the known recruitment values that
produced the estimated numbers at age. We assessed
the influence of the magnitude of variation in
recruitment and fishing mortality, and longevity (short
lived versus long lived) on catch-curve residual
performance using Monte Carlo simulation.
Model formulation.—Population numbers at age and
year (Na,t) were calculated from age 1 to age tmax using
(Ricker 1975), that is,
Na1;t ¼ Na;t1  eðMþFt1 va1 Þ :
where M is the instantaneous natural mortality rate,
Ft1 is the instantaneous fishing mortality rate, and
va1 represents age-specific harvest vulnerability.
Harvest vulnerability was knife edge and was attained
when fish exceeded a minimum length limit (MLL),
then declined with length thereafter (Figure 1).
Declining harvest vulnerability with age (or length)
has been documented for recreational line fisheries
(Miranda and Dorr 2000; Newby et al. 2000). The
catch-curve method assumes constant survival across
ages. Therefore, including declining fishing mortality
with age allowed us to evaluate the robustness of the
method to violations of the constant mortality assump-
tion.
Time-specific fishing mortality was calculated as
Ft ¼ F̄wt ;
where F̄ is the average instantaneous fishing mortality
rate and wt represents a random, lognormally distrib-
uted annual fishing effort deviate with a median of 1
and a variance of loge(CV2F þ 1), where CVF is the
coefficient of variation (SD/mean) in the instantaneous
fishing mortality rate (Limpert et al. 2001).
Numbers at age and year were driven by the annual
number of recruits to age 1. These were calculated as
the product of the average annual recruitment, R0, and a
random lognormally distributed deviate with a median
of 1 and a variance of loge(CV2R þ 1), where CVR is the
 ASSESSING PERFORMANCE OF CATCH-CURVE RESIDUALS
coefficient of variation in annual recruitment (Limpert
et al. 2001).
Sampling the simulated population.—The popula-
tion was projected forward for three times the
maximum age before simulated length–age samples
were collected. After this burn-in period, assessment
catches at age were simulated via the Poisson sampling
process, namely,
Ca ; Poiðka ¼ qNa sa Þ;
where q is an assumed catchability coefficient and sa is
the age schedule for dome-shaped vulnerability to the
sampling gear. In the analysis, q was scaled to result in
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a total catch of approximately 1,200 fish, of which
roughly 25–45% were aged, depending on longevity
and asymptotic length. Dome-shaped sampling vulner-
ability was modeled using the exponential logistic
function (Thompson 1994)
  
1 1  c c ebcðL50la Þ
sa ¼
1c c 1 þ ebðL50la Þ
where la is the age-specific total length (mm), L50 is
the length at which the vulnerability to capture is 0.5, b
is the slope of the function at L50, and c determines the
strength of the dome shape. Age-specific total length
was calculated using the von Bertalanffy (1938) model
h i
la ¼ L‘ 1  eKðat0 Þ ;
where L‘ is the asymptotic length parameter, K is the
metabolic parameter, and t0 is the time at zero length.
Note that we assumed sampling and harvest gears had
different vulnerability schedules. We chose dome-
shaped sampling gear vulnerability because typical
sampling gears such as electrofishing and gill nets
often capture intermediate-sized fish more effectively
than small and large individuals (Hansen et al. 1997;
Bayley and Austen 2002). However, the catch-curve
technique assumes that fish are sampled in proportion
to their abundance. Thus, we could assess the
robustness of the catch-curve residual method by
violating this assumption using dome-shaped sampling
vulnerability.
The population was sampled to mimic the field
practice of selecting a specified number of fish from a
specified size range (i.e., 10 fish per centimeter group)
for aging. Catch-at-age samples were converted to
length–age samples by choosing Ca lognormally
distributed random deviates for total length (mm) of
median la and SD (CVl 3 la) to approximate a
multiplicative error structure, where CVl is the
coefficient of variation in length at age. These catch-
at-age samples were subsampled for age analysis by
297
randomly selecting 10 fish per centimeter group. Aging
error was incorporated using methods described in
Richards et al. (1992) and Coggins and Quinn (1998).
We modeled increasing probability of incorrect age
assignment with fish age and errors were normally
distributed with no systematic bias in age estimation.
We set the SD for correct age assignment at 0.4 for
age-1 fish and 0.6 for age-tmax fish. Aged fish were
used to simulate an age–length key that was applied to
the simulated catch-at-length data to estimate the age
composition of the catch. Catch-curve residuals were
calculated from a linear regression analysis of the
natural log of catches at age versus age.
Correctly modeling how data analysts decide which
ages to include in the catch curve is critical to
realistically simulating the process. Typically, analysts
include ages beginning with the age having the largest
catch but may also use some judgment as to the
selective properties of the gear to guide their decision.
For example, if an age structure exhibited declining
catches beginning at age 2 but with an anomalous
maximum catch at age 7, presumably due to extreme
recruitment variation, the experienced analyst might
begin the catch curve at age 2 instead of age 7.
Therefore, our model assumed that analysts had some
knowledge of the selective properties of the gear but
would also begin including age samples at the age
with the largest catch. We specified a window of 1
year on either side of the age of maximum expected
catch (i.e., max[ka]) and began the catch curve at the
age of maximum catch within this 3-year window. In
doing so, we did not always begin the catch curve at
the age at maximum catch and were able to calculate
residuals for extreme recruitment events. Ages were
included up to the oldest age with at least five
individuals captured.
Model parameterization and analyses.—We evalu-
ated the performance of catch-curve residuals as
indicators of year-class strength across a range of
recruitment variation (CVR) and fishing mortality
variation (CVF) for two longevities (short lived and
long lived; see Table 1 for model parameter values).
Model parameters were set to generally represent North
American open-access recreational fisheries. Instanta-
neous natural mortality was calculated from tmax using
the empirical relationship of Hoenig (1983) and was
constant across years and ages. The coefficient of
variation in recruitment ranged from 0.4 to 1.2 (nine
levels) to reflect values observed for largemouth bass
(CV range ¼ 0.11–1.89; Allen and Pine 2000), crappies
(CV range ¼ 0.55–1.24; Allen and Pine 2000),
walleyes (CV ¼ 0.77; Hansen et al. 1998), and spotted
seatrout Cynoscion nebulosus (CV ¼ 0.35; Murphy
2006). Published time series data on the variation in
 298 CATALANO ET AL.

TABLE 1.—Input parameters used in the age-structured

models for simulations of the short-lived and long-lived life
history types. See text for additional details.
Parameter Units Short lived Long lived

tmax year 10 30
M year1 0.42 0.14
L‘ mm 500 1,000
K year1 0.28 0.092
t0 year 0 0
F̄ year1 0.2 0.2
CVF 0.1–0.6 0.1–0.6
CVR 0.4–1.2 0.4–1.2
R0 Recruits 1,000,000 1,000,000
CVl 0.1 0.1
MLL mm 300 600
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L50 mm 175 353

c 0.16 0.17
b 0.022 0.011

fishing mortality for open-access fisheries are rare. We

set the range of CVF from 0.1 to 0.6 (9 levels), which
exceeded the temporal variation in the exploitation
patterns for walleye (CV ¼ 0.43; Serns and Kempinger
1981) and spotted seatrout (CV ¼ 0.21; Murphy 2006)
fisheries by approximately 40–50%. The average
instantaneous fishing mortality rate (F̄) was 0.2/year
to approximate values for largemouth bass (median ¼
0.18; Allen et al. 2007), walleyes (mean ¼ 0.24;
Baccante and Colby 1996), spotted sea trout (mean ¼
0.23; Murphy 2006), crappies (mean ¼ 0.65; Allen et
al. 1998), and northern pike Esox lucius (mean ¼ 0.12;
Allen et al. 1998). The longevity of the short-lived life
history type was 10 years, whereas that of the long
lived was 30 years to represent the range observed for
recreationally exploited fish species. Sampling vulner-
ability parameters (L50, c, and b) were set such that
fish were (1) 50% vulnerable at 0.25  L‘, (2) fully
vulnerable at 0.5  L‘, and (3) 50% vulnerable at L‘,
which approximated the patterns from Bayley and
Austen (2002; Figure 2). Fish became vulnerable to
harvest at a minimum length limit (MLL) of 0.6  L‘,
and harvest vulnerability declined with length thereaf-
ter to 60% vulnerability at L‘ (Figure 2; Miranda and
Dorr 2000; Newby et al. 2000).
For each combination of CVR, CVF, and longevity,
we compared catch-curve residuals with the true
recruitment values that produced them. We were
interested in three levels of analysis: (1) the overall
pattern of residuals versus true recruitment values, (2)
how well individual recruitment values predicted
individual catch-curve residuals, and (3) how changes
in key underlying assumptions of the population model
affected performance of catch-curve residuals. For the
first analysis, we correlated catch-curve residuals with
the recruitment values that produced them using
Pearson’s correlations. We performed 5,000 Monte
FIGURE 2.—Representative 50-year time series of the
exploitation rate and annual recruitment under high, moderate,
and low levels of variation in fishing mortality (CVF) and
recruitment (CVR).
Carlo iterations of the process for each combination of
CVR and CVF, and for each we calculated the
probability of finding a statistically significant corre-
lation between catch-curve residuals and recruitment
values. The probability was calculated as the propor-
tion of Monte Carlo iterations resulting in a correlation
coefficient greater than or equal to 0.55 (i.e.,
approximate correlation at a ¼ 0.1). We kept the test
criteria the same for both longevities to allow for
comparisons between the two even though the sample
sizes differed due to differences in the number of ages
represented.
The second analysis evaluated how well we could
predict an individual catch-curve residual from a true
recruitment value. For this analysis, we chose two
levels for CVR (low ¼ 0.5; high ¼ 1.1) and CVF (low ¼
0.2; high ¼ 0.5), and we pooled all of the residual–
recruitment pairs across the 5,000 Monte Carlo
iterations for each combination of CVR, CVF, and
longevity. First, we evaluated the probability that we
would obtain a strong year-class (positive residual)
when the true recruitment value was the same (i.e.,
above-average recruitment) by calculating the propor-
 ASSESSING PERFORMANCE OF CATCH-CURVE RESIDUALS 299

tion of residual–recruitment pairs for which the sign

(6) agreed. We then used linear regression to model
the catch-curve residuals as a function of the true
recruitment values. We estimated the slope of this
relationship and calculated 95% prediction intervals for
the catch-curve residuals as a function of the
recruitment values.
Assessing model assumptions.—We evaluated the
effect of changes to assumptions of the base model on
the probability of detecting significant correlations
between catch-curve residuals and recruitment values.
This was accomplished by running the model with
systematic variations to the parameters and comparing
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the results with a base model. The base model was run
at intermediate values of CVF (0.35) and CVR (0.8) for
both longevities. We evaluated the effect of varying
input parameter values for_, M, aging error, and the
shape of the fishery and sampling vulnerability
schedules. We assessed the effect of higher fishing
mortality rates by increasing F̄ by 50% from 0.2 to 0.3/
year. The effect of changes in M was evaluated by
increasing M by 50%. The effect of aging error was
assessed by running the model without aging error. The
influence of changes in harvest vulnerability were
evaluated by assuming full vulnerability (knife-edge,
sigmoidal curve) to harvest upon reaching the mini-
mum length limit and constant vulnerability thereafter.
Similarly, changes in sampling vulnerability were
assessed by setting the vulnerability equal among
size-classes (ages). For each model assumption, we
calculated the proportion of 5,000 Monte Carlo
iterations that resulted in correlation coefficients
greater than or equal to 0.55 (i.e., approximate
correlation at a ¼ 0.1), and compared these values
with those of the base model. We calculated the percent
change in the probability of finding a significant
correlation as
ðPalt  Pbase Þ=Pbase ;
where Palt is the probability of finding a correlation
greater than 0.55 for an alternative model (i.e., models
with systematically varied assumptions) and Pbase is the
probability of finding a correlation greater than 0.55 for
the base model.
Results
The simulation model showed that the utility of
catch-curve residuals to index year-class strength was
positively related to the CV in annual recruitment
(CVR) but weakly affected by the CV in fishing
mortality (CVF). Less recruitment variation was
required to produce significant residual–recruitment
correlations for the long-lived than for the short-lived
life history type. For the short-lived life history type,
FIGURE 3.—Simulated probabilities (isopleths) of a signif-
icant correlation (a ¼ 0.10) between catch-curve residuals and
recruitment values as a function of the coefficient of variation
in recruitment and the coefficient of variation in fishing
mortality for the long-lived and short-lived life history types.
Each probability was calculated as a proportion of 5,000
Monte Carlo iterations that resulted in a correlation of more
than 0.55, which corresponded to a significance level of about
0.10.
the probability of finding a significant correlation
between catch-curve residuals and true recruitment
values increased with increasing CVR and exceeded 0.9
above a CVR of 1.0 (Figure 3, top panel). For the long-
lived life history type, the probability of finding
significant correlations between catch-curve residuals
and true recruitment values exceeded 0.9 above a CVR
of about 0.5 (Figure 3, bottom panel). Variation in
fishing mortality did not strongly influence the
probability of a significant correlation (Figure 3).
Unlike recruitment variation, the influence of variation
in fishing mortality was similar between life history
types.
Catch-curve residuals increased at less than a 1:1
ratio with increases in true recruitment values,
indicating a dampening of the magnitude of strong
and weak year-classes in the catch-at-age data (Table 2;
 300 CATALANO ET AL.

TABLE 2.—Results of 5,000 Monte Carlo simulations comparing catch-curve residuals with the recruitment values that
produced them. The symbols CVR and CVF stand for the coefficients of variation in annual recruitment and fishing mortality,
respectively. The probability of sign agreement is the probability of obtaining a positive catch-curve residual when the true
recruitment value is above average. The 95% prediction interval is an average prediction interval for expected catch-curve
residuals as a function of recruitment values from a linear regression, and the regression slope is the slope of the linear model.
Life history Cohort Probability of 95% Prediction Regression
type age CVR CVF sign agreement interval (6) slope

Short lived Age 5 Low Low 0.73 0.53 0.48

High 0.72 0.53 0.49
High Low 0.80 0.71 0.50
High 0.79 0.72 0.49
.Age 5 Low Low 0.71 0.64 0.51
High 0.70 0.65 0.51
High Low 0.76 0.82 0.48
High 0.75 0.82 0.47
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Long lived Age 9 Low Low 0.74 0.58 0.64

High 0.73 0.60 0.63
High Low 0.81 0.74 0.66
High 0.82 0.74 0.66
.Age 9 Low Low 0.75 0.70 0.71
High 0.74 0.72 0.70
High Low 0.81 0.88 0.67
High 0.81 0.89 0.67

Figures 4, 5). The slope of this relationship was steeper
for the long-lived than for the short-lived life history
type (Table 2). Despite this overall positive trend, there
was substantial variation in predictions of individual
catch-curve residuals as evident by the scatter around
the regression line. The probability of obtaining a
strong year-class when the true recruitment value was
positive (i.e., sign agreement) was around 0.75 for both
longevities and was slightly greater for old than for
young fish (age 5 for the short-lived life history type
and age 9 for the long-lived life history type; Table
2). The 95% prediction intervals for catch-curve
residuals as a function of true recruitment values
ranged from 0.53 to 0.82 for the short-lived life history
type, and from 0.58 to 0.89 for the long-lived life
history type, the highest values being observed for old
fish and at higher CVR values (Table 2). Thus, high
recruitment variation improved overall correlations
between catch-curve residuals and recruitment values,
but also worsened predictions regarding individual
catch-curve residuals.
Assessing Model Assumptions
Changes in fishing mortality and sampling vulner-
ability had the largest effect on the probability of
detecting a relationship between catch-curve residuals
and year-class strength (Table 3). When compared with
the base model, a 50% increase in the fishing mortality
rate resulted in 7% and 3% decreases in the probability
of detecting a significant correlation for the short-lived
and long-lived life history types, respectively. Assum-
ing equal sampling vulnerability across ages resulted in
a 6% and 1% increase in correlation probabilities for
the short-lived and long-lived life history type,
respectively. Changes in the natural mortality rate,
aging error, and harvest vulnerability had minimal
effect (1%) on correlations between catch-curve
residuals and recruitment values.
Discussion
Our results suggest that the magnitude of recruitment
variation and fish longevity strongly influence corre-
lations between catch-curve residuals and recruitment
values. Catch-curve residuals are a reasonable proxy
for year-class strength so long as recruitment variation
(CV in recruitment) exceeds about 50–80%. Allen and
Pine (2000) reported a recruitment CV of 82% (range,
55–124%) for crappie and 66% (range, 11–189%) for
largemouth bass, which are species primarily from the
southeastern United States. Given these CVs in
recruitment levels, catch-curve residuals would prob-
ably perform well for other species with similar levels
of recruitment variation. Recruitment variation less
than 50% probably results in a weak recruitment signal
that is obscured by other factors such as mortality
variation and sampling variability. We suggest that the
magnitude of recruitment variation be assessed using a
metric such as the recruitment coefficient of determi-
nation or the recruitment variability index to evaluate
the usefulness of catch-curve residuals for specific
populations before attempting the analysis (Isermann et
al. 2002).
Fishery managers are often interested in making
inferences regarding the strength of individual year-
classes. Our simulations show that although correla-
tions between catch-curve residuals and recruitment
 ASSESSING PERFORMANCE OF CATCH-CURVE RESIDUALS 301
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FIGURE 4.—Scatterplots of individual recruitment values versus catch-curve residuals for a short-lived life history type for two
levels of the coefficient of variation in recruitment (CVR; 0.5 and 1.1) and the coefficient of variation in fishing mortality (CVF;
0.2 and 0.5). This figure shows the deviations of individual catch-curve residuals from their respective true recruitment values.
Each point on the plot is the residual for a single age in a catch-curve. The 458 line represents a 1:1 ratio between residuals and
recruitment values. The dashed lines indicate regression slopes for young and old cohorts. These slopes were obtained from
5,000 Monte Carlo iterations of the model, but we only show the results from the first 30 iterations for clarity. The recruitment
values were loge transformed to linearize the relationship owing to the different scaling of the catch-curve residuals (normally
distributed; range ¼ ‘ to ‘) and recruitment values (lognormally distributed; range ¼ 0 to ‘).

values may be strong, there is much uncertainty in
making predictions regarding the strength of individual
year-classes. This uncertainty was relatively constant
across longevity and CVF values but depended on the
age of the year-class and the magnitude of recruitment
variation, as prediction intervals were wider for older
fish and higher CVR values. We propose that inferences
of individual year-classes should consider this uncer-
tainty and that assertions of individual year-class
strength should be made cautiously.
Several authors have attempted to validate catch-
curve residuals as indicators of year-class strength, and
the results thus far have been mixed. Catch-curve
residuals were significantly correlated with year-class
strength for crappie from trap nets and largemouth bass
collected via electrofishing in Alabama reservoirs
(Maceina 2004). Smith et al. (2005) reported signifi-
cant correlations between catch-curve residuals and
age-0 electrofishing catch per effort (CPE) for small-
mouth bass in three Virginia rivers. Bonvechio and
Allen (2005) verified catch-curve residuals using a sign
test to evaluate concordance of catch-curve residuals in
sequential years of electrofishing samples on eight
Florida water bodies. Allen et al. (2003) reported
general agreement between catch-curve residuals and
long-term electrofishing catch rates for age-0 large-
mouth bass at Lake Kissimmee, Florida. Conversely,
Quist (2007) found that age-0 CPE from gill nets was
poorly correlated with subsequent catch-curve residuals
for walleye in eight Kansas reservoirs. Isermann et al.
(2002) found poor associations between catch-curve
residuals and long-term recruitment estimates for
 302 CATALANO ET AL.
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FIGURE 5.—Scatterplots of individual recruitment values versus catch-curve residuals for a long-lived life history type for two
levels of the coefficient of variation in recruitment (CVR; 0.5 and 1.1) and the coefficient of variation in fishing mortality (CVF;
0.2 and 0.5). See Figure 4 for additional details.

crappie collected in trap nets, but the gear type was

thought to have poorly represented age-0 CPE due to
selectivity patterns. Field studies such as these are often
impeded by extreme variation in both independent
TABLE 3.—Effects of varying simulation model assumptions (errors-in-variables problem) and dependent variables.
on the probability of detecting a significant correlation Consequently, catch-curve residuals are not directly
between catch-curve residuals and recruitment values. The compared with true underlying recruitment values in
columns are the probabilities of detecting a correlation greater
than 0.55 for the base model and five alternative model field studies, but rather scientists correlate them with
formulations for the short- and long-lived life history types. age-0 or age-1 CPE, which is subject to substantial
The values in parentheses are the percent differences in sampling variability. In addition, comparing residuals
probability relative to the base model. Each assumption was to age-0 catches does not account for the possibility
tested individually and was based on 5,000 Monte Carlo that year-class strength may not be established during
iterations.
the first year of life.
Model Short-lived Long-lived The simulation approach that we used circumvented
Base model 0.88 0.98
the problem of validating catch-curve residuals against
Increase F̄ by 50% 0.81 (7%) 0.95 (3%) CPE indices, which are prone to sampling bias and
Increase M by 50% 0.87 (1%) 0.98 (0%) noise. Our approach provided perfect knowledge of the
Remove aging error 0.87 (1%) 0.98 (0%)
Knife-edge sigmoidal harvest true underlying recruitment time series and compared
vulnerability 0.87 (1%) 0.98 (0%) these to subsequent catch-curve residuals to evaluate
Constant sampling vulnerability performance of the approach. Another advantage of
(i.e., no size selectivity) 0.93 (þ6%) 0.99 (þ1%)
simulation is that we were able to purposefully violate
 ASSESSING PERFORMANCE OF CATCH-CURVE RESIDUALS
the assumptions of catch curves in a controlled manner
and evaluate the effects on catch-curve residual
performance. In this way, we could systematically test
the limits of the approach to determine conditions
under which it is most likely to succeed. This cannot be
done by comparing residuals and annual recruitment
indices from a few populations with unknown
underlying demographic parameters and sampling
variability. The disadvantage of a simulation approach
is that no model can represent the richness and
complexity of processes found in nature. In a purely
simulation framework, the model results are directly
dependent on the input parameters, which represent our
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imperfect knowledge of system structure. Such simu-
lations can lose context in the absence of data from real
fish populations. Nevertheless, we feel our simulations
provided insight into the conditions under which catch-
curve residuals should be used, which could be
compared with field investigations of this issue (e.g.,
Isermann et al. 2002; Allen et al. 2003; Maceina 2004;
Bonvechio and Allen 2005; Smith et al. 2005; Quist
2007).
Factors such as natural mortality variation, fishing
mortality, density-dependent survival, and growth
affect the performance of catch-curve residuals.
Maceina (2004) reported density-dependent growth
and mortality during the second year of life for crappie
in Alabama reservoirs. Our model assumed that all
density-dependent processes occurred before recruit-
ment to age 1, and thus was unable to account for
density dependence later in life. Our predictions
regarding catch-curve residual performance would be
overly optimistic for populations with strong density
dependence after the first year of life, particularly if
prerecruit age-classes were included in the catch curve.
Density-dependent fishing mortality may also be
important to the performance of catch-curve residuals.
Quist (2007) found poor performance of catch-curve
residuals because of high fishing mortality on large
year-classes due to shifts in angler effort. Such effort
shifts have been documented for many recreational
fisheries (Cox et al. 2002). Cox and Walters (2002)
showed that the spatial distribution of fishing effort
changes as a function of angler catch rate, and Cox et
al. (2003) developed a model that predicts how effort
limitations would be required to optimize recreational
fisheries due to angler movements to water bodies that
maximize their catch. Thus, catch-curve residual
performance may be hindered if fishing mortality is
positively correlated with recruitment, which would
reduce the magnitude of the residuals for strong year-
classes in the catch curve. Our model simulated
random variation in fishing mortality but did not
account for density dependent shifts in fishing effort
303
due to anglers targeting strong year-classes. However,
we did allow the fishing mortality rate to vary widely
among years, which reduced year-class abundances at
random periods through the simulations.
In evaluating the assumptions of our model, we
found that catch-curve residuals were robust to
violations of catch-curve assumptions. We found high
probabilities of significant residual–recruitment corre-
lations despite violating the equal capture probability
assumption by using a dome-shaped sampling vulner-
ability schedule. Another assumption of the catch curve
is constant mortality across ages. Our analysis detected
only a small increase in the performance of catch-curve
residuals when assuming a knife-edge sigmoidal
vulnerability function (i.e., fully vulnerable across all
ages after reaching the minimum length limit) instead
of declining fishing mortality with age. The catch-
curve technique also assumes constant mortality
through time. Our simulation violated this assumption
by allowing time-varying fishing mortality. However,
we found that the performance of catch-curve residuals
was only weakly reduced by increasing the CV in
fishing mortality. Our model did not allow time-
varying natural mortality, which could affect perfor-
mance and should be considered in systems with
episodic mass-mortality events. Finally, catch curves
assume constant recruitment such that residuals are the
result of normally distributed random recruitment
variation. We show that the magnitude of this variation
is important in determining the degree of correlation
between recruitment and catch-curve residuals. Catch-
curve residuals could be biased if there is a recent trend
in recruitment, but our model did not evaluate this
possible assumption violation. For example, several
recent years of high recruitment can overestimate the
total mortality rate, and residuals from such an analysis
may not depict recruitment patterns. This should be
considered in systems that are suspected of having
cyclic recruitment dynamics.
Management Implications
Catch-curve residuals represent a potentially rapid
way of assessing year-class strength that can be used to
relate recruitment to environmental variables. Our
results suggest that catch curves are likely to reveal
broad relationships between environmental variables
and recruitment at realistic levels of recruitment
variability as well as moderate variability in mortality
rates. Furthermore, the technique is robust to violations
of the underlying assumptions of the catch-curve
method. However, we propose several caveats for the
use of this technique: (1) assessing the magnitude of
specific year-classes using catch-curve residuals is
often not reliable, (2) the method is probably most
 304 CATALANO ET AL.
suitable as a rapid assessment method in systems for
which no historical data exist and the likelihood of
relatively static mortality is high, (3) estimates of the
variation in recruitment are needed for the species and
system of interest in order to fully assess the utility of
this technique, and (4) preliminary studies should
assess the degree of density-dependent mortality and
growth occurring after the age at recruitment.
Catch-curve residuals should be viewed as a
rudimentary but potentially useful measure of recruit-
ment under realistic ranges of recruitment and mortality
variation and could be helpful for fishery managers
faced with making management decisions in data-poor
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situations. Our analysis was requested by field fishery
managers with the Florida Fish and Wildlife Conser-
vation Commission working in the Apalachicola River
basin. Given current drought conditions in this river
basin, increases in water withdrawals to meet human
needs, and associated reductions in riverine flows, these
managers are involved in one of the most contentious
water allocation disputes in the eastern United States
(Barnett 2007). These managers sought to understand
the utility of catch-curve residuals to inform policy
decisions regarding the timing, duration, and magnitude
of flows in the lower portion of this basin by assessing
the performance of catch-curve residuals to track
relative year-class strength through time with varying
flow conditions. Thus, understanding the limits of the
approach should be of interest to fisheries managers,
researchers, and conservation groups.
Acknowledgments
We thank Rich Cailteux, Andy Strickland, and
Charlie Mesing for assistance with data development
and Lew Coggins, Carl Walters, and Steve Martell for
help formulating the model. We thank the Florida Fish
and Wildlife Conservation Commission for funding
this work.
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