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An individual-based modeling approach to

spawning-potential per-recruit models: an
application to blue crab (Callinectes sapidus) in
Chesapeake Bay
David B. Bunnell and Thomas J. Miller

Abstract: An individual-based modeling approach to estimate biological reference points for blue crabs (Callinectes
sapidus) in Chesapeake Bay offered several advantages over conventional models: (i) known individual variation in size
and growth rate could be incorporated, (ii) the underlying discontinuous growth pattern could be simulated, and
(iii) the complexity of the fishery, where vulnerability is based on size, shell status (e.g., soft, hard), maturity, and sex
could be accommodated. Across a range of natural mortality (M) scenarios (0.3751.2year1), we determined the exploitation fraction () and fishing mortality (F) that protected 20% of the spawning potential of an unfished population,
the current target. As M increased, 20% and F20% decreased. Assuming that M = 0.9year1, our models estimated
20% = 0.45, which is greater than field-based estimates of in 64% of the years since 1990. Hence, the commercial
fishery has likely contributed to the recent population decline in Chesapeake Bay. Comparisons of our results with conventional per-recruit approaches indicated that incorporating the complexity of the fishery was the most important advantage in our individual-based modeling approach.
Rsum : Une mthodologie de modlisation base sur lindividu pour estimer les points de rfrence biologique du
crabe bleu (Callinectes sapidus) dans la baie de Chesapeake prsente plusieurs avantages par rapport aux mthodes
conventionnelles : (i) on peut incorporer les variations individuelles connues de taille et de taux de croissance, (ii) on
peut simuler le patron sous-jacent de croissance discontinue et (iii) on peut tenir compte de la complexit de la pche
dans laquelle la vulnrabilit dpend de la taille, de ltat de la coquille (par exemple, molle, dure), de la maturit et
du sexe. Sur une gamme de scnarios de mortalit naturelle (M = 0,3751,2 an1), nous avons dtermin la fraction de
lexploitation () et la mortalit due la pche (F) qui protgent 20 % du potentiel de ponte de la population non
affecte par la pche, ce qui est lobjectif actuel. Lorsque M augmente, 20 % et F20 % diminuent. En assumant que
M = 0,9 an1, nos modles estiment 20 % 0,45, ce qui est plus que les estimations de terrain de dans 64 % des
annes depuis 1990. La pche commerciale a donc vraisemblablement contribu au dclin rcent de la population dans
la baie de Chesapeake. La comparaison de nos rsultats ceux des approches conventionnelles, qui font les calculs par
recrue, indique que lincorporation de la complexit de la pche est lavantage le plus significatif de notre mthodologie de modlisation base sur lindividu.
[Traduit par la Rdaction]

Bunnell and Miller

Introduction
Crustacean fisheries offer unique challenges for stock
assessment modelers (Smith and Addison 2003). First, crustaceans grow discontinuously through molting rather than
continuously as finfishes. Hence, models that use a von
Bertalanffy growth subroutine, which assumes continuous
growth, do not reflect the underlying biology of crustaceans.
Received 8 July 2004. Accepted 2 May 2005. Published on
the NRC Research Press Web site at http://cjfas.nrc.ca on
14 October 2005.
J18213
D.B. Bunnell1,2 and T.J. Miller. Chesapeake Biological
Laboratory, University of Maryland Center for Environmental
Science, P.O. Box 38, Solomons, MD 20688, USA.
1
2

Corresponding author (e-mail: dbunnell@usgs.gov).

Present address: USGS Great Lakes Science Center,
1451 Green Road, Ann Arbor, MI 48105-2807, USA.

Can. J. Fish. Aquat. Sci. 62: 25602572 (2005)

2572

Second, similar to many other organisms, the components of

the crustacean growth process (i.e., the intermolt period and
the growth per molt) can be highly variable among individuals. This variability, however, is ignored in many conventional fisheries models. Third, crustacean ages are difficult
to estimate because they do not have scales or otoliths that
are generally used to age finfishes (but alternatives may exist, see Ju et al. 2001). As a result, developing age-structured
models is problematic. Fourth, crustacean fisheries often
have regulations that are dependent on the sex, maturity, and
shell status (e.g., hard or soft) of the individual. Thus, an
ideal stock assessment model for crustaceans would be size
based and allow for individual variation in discontinuous
growth and for sex-, shell- and maturation-dependent harvest
regulations.
Individual-based models are perfectly suited for this task,
and this modeling approach has been used in previous stock
assessment models to accommodate complexities that are ill
suited for conventional models. Previous per-recruit assess-

doi: 10.1139/F05-153

2005 NRC Canada

Bunnell and Miller

ment models have used an individual-based approach to

incorporate spatial variation in fishing mortality (F) (Hart
2001), multiple spawning within a season (Lowerre-Barbieri
et al. 1998), size-selective mortality among individuals of
varying growth rate (Kristiansen and Svsand 1998), and the
molting growth process of crustaceans (Fogarty and Idoine
1988). In this paper, we build on the seminal work of
Fogarty and Idoines (1988) individual-based per-recruit
models for American lobster (Homarus americanus) by developing individual-based per-recruit models for blue crab
(Callinectes sapidus) in Chesapeake Bay.
Per-recruit models are generally used to evaluate how
changes in fishing mortality or size limits will influence the
yield or spawning potential (e.g., eggs produced, spawning
stock biomass) of a cohort of recruits. Beverton and Holt
(1957) developed a yield-per-recruit (YPR) model that used
the von Bertalanffy growth parameters to find the biological
reference point Fmax, or the level of fishing mortality that
maximizes yield. Because YPR models do not consider
whether resulting fishing mortality reference points are sustainable, spawning potential per recruit (SPPR) models (also
called egg-per-recruit models) were developed later (see
Goodyear 1993). SPPR models produce biological reference
points, Fx%, which represent the fishing mortality rate that
reduces the SPPR of the population to x% of the SPPR of a
virgin, unfished population (see Goodyear 1993). Resulting reference points have been recommended to be as
low as F10% for American lobsters (Anonymous 1993) and
as high as F50% for rockfishes (Sebastes spp.) in western
North America (Clark 2002).
In Chesapeake Bay, SPPR models have been used to set
biological reference points for the blue crab fishery, which
has averaged more than $45 million in annual market value
since 1981 (range = $25$72 million; US Department of
Commerce, NOAA Fisheries, http://www.st.nmfs.gov/st1/
commercial/index.html). Maryland, Virginia, and the Potomac River Fisheries Commission have management authority for their respective waters in the Bay. Although each
jurisdiction has its own unique and complex set of regulations, blue crabs in all waters of the Bay generally recruit
into and out of different fisheries based on their shell status
(hard shell, soft shell, or peeler), size, sex, and maturity. In
response to declining population sizes and harvests, the BiState Blue Crab Advisory Committee agreed on F20% as a
target reference point in 2001 (Chesapeake Bay Commission
2001). Their SPPR model estimated F20% = 0.7 (Chesapeake
Bay Commission 2001), which was slightly lower than a
previously published estimate of 0.8 (Rugolo et al. 1998).
Both of these estimates, however, derived from age-based
models that assumed continuous von Bertalanffy growth and
allowed recruitment to the fishery to be a function of only
crab age. Hence, these modeling approaches used a biologically inaccurate growth subroutine and did not incorporate
the complexity of the blue crab fishery.
Developing an accurate and biologically realistic SPPR
model for the blue crab fishery in the Chesapeake Bay is increasingly important given the blue crab population decline
in recent years. Although population abundance over the
past 50 years has varied considerably, abundances in the late
1990s and early 2000s have remained consistently low
(Lipcius and Stockhausen 2002; Sharov et al. 2003). Both

2561

spawning stock biomass and larval abundance are estimated

to be below the long-term average (Lipcius and Stockhausen
2002). In addition, the average size of males (Abbe 2002)
and mature females (Lipcius and Stockhausen 2002) has declined. Finally, rates of fishing mortality and exploitation
appear to have increased in recent years (Rugolo et al. 1998;
Sharov et al. 2003). Recent matrix-based modeling efforts
have indicated that current exploitation rates are not sustainable and should be reduced (Miller 2001, 2003).
In this paper, we develop an individual-based model that
incorporates individual variation in growth per molt (GPM)
and the intermolt period (IP) and incorporates the complexity of the fishery regulations. First, we provide a validation
of the growth subroutine of the model by comparing size
distributions of modeled crabs with size distributions of
crabs sampled during the year from the Chesapeake Bay.
Under a range of natural mortality (M) scenarios, we then
use our model to estimate F20% and 20%, where is the exploitation fraction. Finally, we perform a sensitivity analysis
to determine whether our individual-based reference point
estimation is influenced by (i) individual variation in size
and growth, (ii) the type of growth subroutine used (i.e.,
molting or von Bertalanffy growth), and (iii) the type of
fishery regulation that is modeled (i.e., one based on size,
sex, shell status, and maturity or one based only on putative
age). In short, we explored whether adding the complexity
and reality in our individual-based model provided different
results from more simplified models. Taken together, our results will provide specific reference points for the Chesapeake Bay blue crab fishery, but they also will inform other
crustacean stock assessment modelers as to whether incorporating complexities such as discontinuous growth and the appropriate fishing regulations is in fact necessary.

Methods
Blue crab life history
The blue crab ranges from South America to Nova Scotia
in the Atlantic Ocean and its estuarine tributaries. In Chesapeake Bay, blue crab larvae, termed zoea, are released by females from high-salinity waters during late spring and early
summer. Larvae are transported offshore but return to the
Chesapeake Bay as megalopae (i.e., the last larval stage) in
summer and autumn (Olmi 1995). Juvenile crabs, initially 2
3 mm carapace width (CW), settle in structured habitat in
the lower bay. Thereafter, blue crabs undergo a series of
molts in which their size is increased between 8% and 50%
(Tagatz 1968; Leffler 1972; Fitz and Wiegert 1991). Prior to
each molt, blue crabs are termed peelers as their hard shell
begins to soften and crack. After shedding the hard shell, the
soft-shelled crabs are highly vulnerable to predators while a
new hard shell hardens over the next 2448 h (Ryer et al.
1997). As juvenile blue crabs increase in size, they disperse
throughout the Bay. In the Chesapeake Bay, molting ceases
during winter with the onset of cold temperatures and blue
crabs burrow into the sediment. Crabs emerge from sediments in late spring and recommence growth. While maturing females are in their last soft-shell stage, males couple
with them and deposit sperm into female oviducts. After this
maturity molt, females are believed to cease molting (Hines
et al. 2003). Males, however, continue to molt after maturity.
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Throughout the Chesapeake Bay, mating occurs in late summer (Hines et al. 2003). These females can store the sperm
over the winter and fertilize and release their eggs the following summer. In the lower Bay, mating also is believed to
occur in late spring (Hines et al. 2003), and these females
likely fertilize and release their eggs during summer of the
same year. Between one and three broods are produced by
females each summer (Hines et al. 2003). Fecundity varies
linearly as a function of body size and typically ranges between 1 and 8 million eggs (Prager et al. 1990).
Individual-based per-recruit model
We developed a sex-specific, individual-based per-recruit
model that monitored the fate of individual blue crabs on a
daily time step. To accommodate the complexity of the fishery, we monitored the shell status (i.e., peeler, soft shell, or
hard shell) and maturity of individuals as they grew according to a molting interval that depended on temperature and
blue crab size. Each individual blue crab in the simulation
was characterized as a super-individual in that each represented some larger number of individuals (herein referred to
as their internal amount) in the population (sensu Scheffer et
al. 1995). The use of super-individuals allowed the total
number of individuals in the simulation to remain sufficiently large in the face of relatively high rates of fishing
and natural mortality.
Growth
In the model, blue crabs grew discontinuously by molting,
the magnitude and frequency of which were governed by
temperature (Tagatz 1968; Leffler 1972) and crab size
(Tagatz 1968; Fitz and Wiegert 1991). Empirical estimates
of the size increase associated with each molt (i.e., GPM)
have ranged from 1.08 to 1.50 times premolt CW (Tagatz
1968; Leffler 1972; Fitz and Wiegert 1991). Our model relied on the data from Tagatz (1968) because he provided the
widest size range of blue crabs. We modeled GPM to increase with size among females and to be constant across
sizes for males (Tagatz 1968). For females, the GPM was
drawn from a normal distribution with a premolt, CWdependent mean (1.218 + (7.09 104)CW) and a standard
deviation of 0.07. For males, the mean GPM also was drawn
from a normal distribution with a mean of 1.25 and an standard deviation of 0.06 (Tagatz 1968).
IP, or the time between molts, was represented by degreedays (degrees Celsius) and also was dependent on blue crab
CW. Each day in the model, blue crabs accumulated degreedays until some threshold had been reached, which, in turn,
resulted in molting. Degree-days were calculated by subtracting 8.9, the physiological minimum temperature for
blue crab growth (Smith 1997), from the mean daily water
temperature. Hence, molting does not occur when water
temperatures are less than 8.9 C (i.e., winter in the Chesapeake Bay) because degree-days will not accumulate. In
crustaceans, IP has been modeled to increase either linearly
(e.g., Smith 1997) or exponentially (e.g., Hoenig and
Restrepo 1989) as a function of size. A plot of IP versus
blue crab CW from several studies indicates that most have
focused on blue crabs less than 80 mm CW and that there is
high variation in IP among blue crabs of similar sizes
(Fig. 1). Once again, we based our growth model on the em-

Can. J. Fish. Aquat. Sci. Vol. 62, 2005

Fig. 1. Intermolt period as a function of blue crab (Callinectes
sapidus) size from several published studies. Because of the
wide size distribution, we relied on Tagatz (1968) to generate
size-dependent estimates of the two phases of the growth process
model (sensu Smith 1997): (broken line) represented the required or physiological minimum phase and represented the
variable phase (see eqs. 4 and 5). The sum of and (solid
line) represented the average intermolt period for a blue crab of
a given size.

pirical results of Tagatz (1968) because he provided data

with the broadest size distribution. For blue crab of a given
size, IP was drawn from a shifted exponential density function (Smith 1997):
(1)

f(IP) = (1/ ) e

IP

IP

where represents the required or physiological minimum

phase and represents an additional and variable phase. In
this distribution, the expected value, + , is relatively close
to the minimum value (i.e., ) but the distribution has a long
tail of larger values, which corresponds to the data from all
of the studies (Fig. 1). Assuming that the minimum IP reported by Tagatz (1968) represents the required phase (),
we modeled to increase exponentially with blue crab CW
as (Fig. 1):
(2)

= 69.70 (1.0149) CW

To ensure that the variable phase did not intersect with the
required phase, we also modeled to increase exponentially
with blue crab CW based on the mean IP from Tagatz
(1968) (Fig. 1):
(3)

= (166.39 (1.0115) CW)

In the model, IP was determined for each individual crab

at day 0 and at each molt. To do so, the model calculated a
cumulative exponential probability function for possible values of IP ranging from to the maximum IP value, which
was calculated as 2.11 times the expected IP (maximum
multiplier observed from Tagatz 1968). A random number
between 0 and 1 was generated, and the first IP for which
the value of the cumulative exponential probability function
exceeded the random number provided the IP for that individual. At each daily time step, degree-days were added to
degree-day exposure until the sum exceeded the IP. On that
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Bunnell and Miller

2563

day, individual blue crabs molted to their new size, their

degree-day exposure was reset to 0, a new IP was determined (as a function of their new size), and their shell status
was changed to soft.
We monitored the status of the blue crab shell (i.e., hard
shell, soft shell, or peeler) because the fishery regulations
change with shell status. Blue crabs were assumed to be
soft-shelled for the day of molting and the following day
only (i.e., individuals returned to hard-shell status 2 days after molting). Blue crab shells were classified as peelers for
approximately 1 week before molting. Because the model
does not calculate the day of molting in advance (i.e., future
temperature is unknown), we used a preliminary simulation
run to estimate the predicted proportion, , of the IP that is
obtained 1 week prior to molting:
degree-day exposure 1 week before molting
IP
In general, increased linearly with blue crab CW, but the
slope and intercept of the versus CW relationship differed
across months in which degree-day exposures were reset
(i.e., the month when molting occurred). During the warmer
months of MaySeptember, = 0.60 + (1.81 103)CW.
During October, because individual blue crabs carry a relatively high degree-day exposure with them during winter,
was the highest: = 0.8722 + (2.94 104)CW. Finally, during the cold months of NovemberApril, = 0.7355 +
(9.59 104)CW. On the day when degree-day exposure exceeded IP, the shell status of blue crabs was changed to
peeler until the crab molted to a soft shell.
(4)

Maturity
Maturity of individual blue crabs was assigned only to females in this model, as maturity was assumed not to influence growth of males. Female maturity was a function of
size and time of year. Using data from the Chesapeake Bay
Winter Dredge Survey from 1990 to 1998 (for details, see
Sharov et al. 2003), we calculated a female maturity ogive
based on 5-mm size bins (N = 23 610 female blue crabs). A
logistic function best described the maturity ogive, given as
(5)

Pr(maturity) =

0.9994
CW
1+

117.981

28.51

, r 2 = 0.99

Maturation was a function of time: females could mature between 1 April and 1 June or between 1 July and 1 October
corresponding to the two mating periods in the Bay (Hines
et al. 2003). During each molt within those periods, an
immature female reached maturity if a randomly generated
number between 0 and 1 was greater than her size-dependent
probability of maturing. Once a female matured, she remained a hard-shell crab and ceased molting for the remainder of her lifetime.
Mortality
Natural and fishing mortality rates were applied separately
in the model. Because there is considerable uncertainty surrounding estimates of natural mortality of blue crabs, we ran
our models with four different values of natural mortality. To
be consistent with previous blue crab population models, we

used an M = 0.375year1, which is based on the assumption

that 5% of blue crabs lived to a maximum age of 8 years
(Rugolo et al. 1998). To address recent concerns that M =
0.375year1 is an underestimate, we used life history invariant theory to provide alternative estimates of natural mortality.
Owing to evolutionary trade-offs among rates of mortality,
growth, and maturity, constant relationships (i.e., invariants)
among these life history parameters have emerged across
taxa (see Charnov 1993). Three caveats are worth noting
when applying life history invariants to blue crabs. First, uncertainty in blue crab aging also leads to uncertainty in estimates of age at maturity and growth rates. Second, invariants
are derived largely from fishes rather than from aquatic invertebrates owing to the larger sample size of fishes. Although the trade-offs that produce the invariants in fishes
also should produce invariants among aquatic invertebrates,
the values of invariants can differ (see Charnov 1993). Third,
estimates of total mortality (Z) derived from the Chesapeake
Bay Winter Dredge Survey (for details, see Sharov et al.
2003) constrain the upper value of natural mortality. These Z
estimates ranged from 0.34 to 1.47year1 (mean of 0.99) between 1990 and 2002 (Lynn Fegley, Maryland Department
of Natural Resources, 580 Taylor Avenue, Annapolis, MD
21401, USA, personal communication).
Given these caveats, we estimated natural mortality with
three different life history invariants. The product of natural
mortality and age at maturity is equal to 1.65 (Jensen 1996)
or 2.0 (Charnov and Berrigan 1990) in fishes. Female blue
crabs are believed to mature sometime between age 1 and
2 years (Hines et al. 2003); in our model, 50% of females
from the initial cohort matured at age 13 months. Considering the range of invariants and age at maturity, M
should be between 0.83 and 2.00year1 for female blue
crabs. A second invariant reveals M divided by k (the Brody
growth coefficient in the von Bertalanffy equation) to equal
1.50 (Jensen 1996) or 1.65 (Charnov 1993). Estimates of k
for blue crab have ranged from 0.49 to 1.09 (Rugolo et al.
1998; Ju et al. 2001), which, in turn, results in an M between
0.74 and 1.80year1. Finally, in a meta-analysis, Pauly
(1980) found natural mortality to be a function of mean annual temperature as well as Linf and k from the von
Bertalanffy equation. Using previous ranges of k, a range of
Linf (i.e., CWinf) from 18.1 cm (Ju et al. 2001) to 26.3 cm
(Rugolo et al. 1998) and an annual mean temperature of
16.5 C (grand mean of average annual temperature from
1991 to 2002 at the Virginia Institute of Marine Science
(VIMS) pier, Gloucester Point, Virginia) reveal M to range
from 1.02 to 1.57year1. Comparisons across the three estimates of natural mortality indicate a range of M between
0.74 and 1.8year1; given that Z has been estimated to be no
greater than 1.5year1, estimates of M greater than 1.3 appear highly unlikely. Nonetheless, the early age at maturity
and fast growth of blue crabs lead to considerably higher estimates of natural mortality using life history invariants than
those generated from the maximum age approach in previous blue crab models.
To span the range of potential natural mortality estimates,
we used four values of M (0.375, 0.6, 0.9, and 1.2year1) in
our models. In an individual-based model, mortality is modeled as a probability. The annual probability of surviving
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Can. J. Fish. Aquat. Sci. Vol. 62, 2005

sources of natural mortality was equal to eM. Thus, the daily

probability of natural mortality was equal to 1 eM/365. Individuals with soft shells were modeled to have twice the
probability of natural mortality as similar-sized peelers or
hard-shelled crabs owing to their greater vulnerability without the hard shell (Ryer et al. 1997). In the model, mortality
was represented by a reduction in the internal amount of
each super-individual. The internal amount (a) was reduced
by some number d, which represented the number of individuals within each super-individual that died. The value d
was drawn from a normal distribution when the product of a
and p was greater than 5, with
(6)

mean = ap
standard deviation = [ap(1 p)]0.5

where p is the daily probability of dying (Scheffer et al.

1995). Otherwise, d was drawn from a binomial distribution
(details in Scheffer et al. 1995). Given the internal amount a
of each super-individual blue crab and its shell-statusdependent daily probability of mortality p, our model estimated the number of individuals d within the superindividual that died using eq. 6.
We based the vulnerability to fishing mortality on Maryland fishing regulations, for which the season is 1 April
through 15 December. Male blue crabs and immature female
blue crabs with a hard shell were vulnerable when they
reached 130 mm CW; this size is a compromise between the
127 mm CW limit between 1 April and 14 July and the
133 mm CW size limit between 15 July and 15 December.
Mature females can be harvested regardless of their size. For
male and female crabs that were in peeler or soft shell
stages, the minimum harvestable size was set at 89 mm CW.
Note that because blue crabs can only have one shell status
(i.e., peeler, hard, or soft), each individual was vulnerable to
only one fishery on any given day.
In the model, the probability of daily fishing mortality
equaled 1 eF/259, where F represented a nominal annual
rate of fishing mortality and 259 is the number of days in the
fishing season. As with natural mortality calculations, the
model used eq. 6 to reduce the internal amount a for each
super-individual blue crab by some number d according to
the probability of daily fishing mortality. The nominal fishing mortality that was used to set the probability of daily
fishing mortality was not the realized fishing mortality on
the entire population because individuals in the model became vulnerable to the fishery at different times based on
their growth rate and shell status. As a result, we estimated
the realized fishing mortality by solving for F in Baranovs
catch equation (Quinn and Deriso 1999):
(7)

F=

C Z
N 0(1 e 2Z )

where C equaled the total number of crabs harvested over

the 2 years of the simulation, Z equaled the total mortality
rate, and N0 equaled the number of blue crabs alive on
1 April, year x + 1 (see Fig. 2), when the fishery begins. Total mortality Z was estimated as
(8)

Fig. 2. Timeline for the individual-based per-recruit model, which

began on 1 January of year x + 1 and ended on 31 December of
year x + 2, where x is the year of settlement.

Z =

ln(N 0) ln(N f )
2

where Nf equaled the number of blue crabs alive at the end

of the 2 years of the simulation (Quinn and Deriso 1999).
Reference points also can be presented in terms of , the
exploitation fraction. We calculated the exploitation fraction
as the number of crabs harvested divided by N0 (Quinn and
Deriso 1999). In our view, reference points for blue crabs in
Chesapeake Bay based on the exploitation fraction rather
than fishing mortality have two advantages. First, comparisons with empirical estimates are possible because both inputs to the exploitation fraction are measured reliably: the
winter dredge survey estimates N0 for each year and management agencies monitor the harvest data throughout the
year. Second, calculation of the exploitation fraction makes
no assumptions regarding natural mortality, over which there
is considerable uncertainty for blue crab. Because current
reference points rely upon fishing mortality, however, we
will report reference points in terms of both and fishing
mortality.
Validation of growth parameters
To validate the growth subroutine of the individual-based
model, we compared size distributions of blue crab generated from our model with size distributions of blue crab
sampled from the Chesapeake Bay during 19971999. Chesapeake Bay (herein field) distributions were derived from
two surveys: (i) the VIMS Juvenile Fish and Blue Crab
Trawl Survey, which occurs monthly in the Virginia waters
of the Chesapeake Bay (including the James, York, and
Rappahannock rivers), and (ii) the Chesapeake Bay Winter
Dredge Survey, which occurs between December and March
throughout the entire Chesapeake Bay. We used the dredge
survey for winter field distributions because it samples crabs
burrowed into the sediments more effectively than a trawl.
The trawl survey more effectively captures smaller crabs as a
result of its 6.4-mm liner compared with the 13-mm liner of
the dredge (for more survey details, see Montane et al.
(2003) for the trawl survey and Sharov et al. (2003) for the
dredge survey). In both surveys, all blue crabs were measured (nearest millimetre CW), sex was determined, and female blue crabs were assessed for maturity. Within each
year, we assumed that crabs smaller than 60 mm CW captured in September or later represented a new cohort (Sharov
et al. 2003) and those individuals were deleted from the field
distributions.
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For each year, we seeded the individual-based model with

the sex, maturity (for females), and CW of all blue crabs
captured in the winter dredge survey, which resulted in different numbers of crabs for each modeled year (1997: 5587,
1998: 3236, 1999: 1970). For each year, the internal amount
a of each individual crab equaled 400 million divided by the
total number of crabs modeled (i.e., we modeled a population of 400 million crabs, which is within the range of total
crab abundance in the 1990s; Sharov et al. 2003). Using the
mean daily temperature recorded at the VIMS pier, we simulated the growth and survival of those blue crabs during
1 year. We used an intermediate value of M, 0.75year1. We
used two different daily probabilities of fishing mortality to
produce exploitation fractions that bounded the minimum
(30%) and maximum (70%) exploitation fractions observed in
the Chesapeake Bay during the 1990s (Sharov et al. 2003). We
then compared size distributions predicted for May, July, September, and January from the model with size distributions
from the same months of blue crabs sampled in the field.
SPPR models and estimation of biological reference
points
For each SPPR model, we simulated a cohort of 2000 juvenile super-individual blue crabs through 2 years following
the year of initial settlement. The model began on 1 January
of year x + 1 and continued through 31 December of year
x + 2, where x is the year of settlement (see Fig. 2). By assigning each super-individual an internal amount of 150 000
at the start of each simulation, we modeled a cohort of 300
million individuals, which is within the range (95540 million) of new recruits estimated in the Chesapeake Bay during the first winter of life (Sharov et al. 2003). Sizes for
each super-individual in the cohort were drawn from a
lognormal distribution with a mean of 27.2 mm CW and a
standard deviation of 10.3, which is reflective of the size
distribution sampled in the winter dredge survey. We assumed an initial 1:1 sex ratio for each cohort. Water temperatures were equal across both years and equaled the mean
daily water temperature at the VIMS pier from 1991 to
2002. As with conventional per-recruit models (Beverton
and Holt 1957), we varied rates of natural mortality and
nominal fishing mortality to determine their effects on
spawning potential.
To estimate 20% and F20%, we estimated the spawning potential per recruit under various mortality regimes. Spawning
potential equaled the sum of the total numbers of eggs predicted to be spawned by females in the second year of the
simulation (see Fig. 2); no females were large enough to mature and spawn by 15 September of the first year of the
model. Mature females were randomly assigned a spawning
day between 15 May and 15 September. The spawning potential, or the number of eggs spawned, for each superindividual was the product of its size-based fecundity (millions of eggs = 2.248 + 0.337(CW); Prager et al. 1990) and
its internal amount a (i.e., the number of individuals that
were living) on the day of spawning. For each combination
of fishing and natural mortality, we summed the spawning
potential of each super-individual. We assumed that the virgin, unfished spawning potential occurred when or F =
0.0year1.

2565

Sensitivity to added model complexities

Our individual-based model was able to accommodate
several realistic complexities that most conventional lengthor age-based per-recruit models do not, including (i) individual variation in crab sizes and growth, (ii) discontinuous
growth, and (iii) vulnerability to harvest that is a function of
crab size, shell status, maturity, and sex. To determine the
impact of including these complexities on reference point estimation, we systematically removed one facet of complexity, replaced it with the more simplified facet, and compared
the resultant reference points. Specifically, using a factorial
design approach, we crossed three different growth subroutines ((i) discontinuous growth with individual variation in
initial sizes and growth (original model), (ii) discontinuous
growth without any individual variation, and (iii) continuous
growth without any individual variation) with two different
harvest subroutines ((i) function of size, sex, shell status,
and maturity (original model) and (ii) function of agedependent recruitment to the fishery). We ran these models
at one level of natural mortality (M = 0.375year1). For the
continuous growth subroutine, we calculated the average
daily size across a low and high level of fishing mortality
when M = 0.375year1 in the original model and used those
values to estimate the von Bertalanffy growth equation
(CW = 263.1(1 e0.6942(t0.007))). For the age-dependent harvest subroutine, vulnerability to the fishery was 0.75F for
the first year (i.e., age 1) and 0.95F for the second (i.e., age
2), which corresponds to previous age-based SPPR models
for blue crabs (Rugolo et al. 1998). For the models with the
von Bertalanffy growth subroutine, we could not use the maturity subroutine that was based on molting events at a given
size. Hence, maturity became a function of age: 10% of age1 females and 90% of age-2 females matured (Rugolo et al.
1998). As in the original model, spawning by mature females occurred in the second year of the simulation.

Results
Validation of growth parameters
Our intent with the validation was not to perfectly match
the field distributions by going through iterations of varying
rates of fishing pressure or natural mortality. Rather, we
wanted to demonstrate that the literature-derived growth parameters could effectively capture the general trajectory of
blue crab size frequencies, and hence growth rates, in the
Chesapeake Bay over the course of 1 year, within the range
of reasonable estimates for fishing and natural mortality. In
each year, the modeled size distributions were broadly similar to the field distributions, with the low fishing pressure
simulations performing slightly better than the high fishing
pressure ones (Fig. 3). As might be expected, models with
the low (0.33 in 1997, 0.46 in 1998, and 0.39 in 1999)
predicted more large crabs during each month of each year
than was observed in the field. Conversely, models with the
high (0.61 in 1997, 0.71 in 1998, and 0.65 in 1999) predicted considerably fewer blue crabs larger than 130 mm
CW than was observed in the field. Having validated this
growth subroutine, we subsequently used this individualbased population model as the basis of our per-recruit models.
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Can. J. Fish. Aquat. Sci. Vol. 62, 2005

Fig. 3. Validation of the growth parameters in the individual-based per-recruit model. Comparison of size frequency distributions of
blue crab (Callinectes sapidus) captured in the Chesapeake Bay (vertical shaded bars) with distributions of simulated blue crabs from
the model (lines) during May, July, September, and January of (a) 1997, (b) 1998, and (c) 1999. The broken line represents modeled
size distributions when exploitation fractions were low (i.e., 0.33 in 1997, 0.46 in 1998, and 0.39 in 1999), and the solid line represents modeled size distributions when exploitation fractions were high (i.e., 0.61 in 1997, 0.71 in 1998, and 0.65 in 1999).

SPPR model results

In our SPPR model, a cohort of newly recruited blue crabs
was simulated for 2 years beginning on 1 January. To describe the size distributions of the cohort through the
2 years, we present results from four representative model
runs of varying levels of natural (M) and fishing mortality
(exploitation fraction ): M = 0.375year1 and = 0.32
(Figs. 4a4c), M = 0.375year1 and = 0.65 (Figs. 4d4f),
M = 0.90year1 and = 0.30 (Figs. 4g4i), and M =
0.90year1 and = 0.48 (Figs. 4j4l). Initial size distributions were the same for all models and reflected size distributions of presumed newly recruited blue crabs in the
Chesapeake Bay (see Figs. 4a, 4d, 4g, and 4j and compare
with the first mode of fig. 4 in Sharov et al. 2003). Size distributions in the next two winters were somewhat influenced
by exploitation rate. By the second winter of life
(31 December, year x + 1) or the end of the first year of the
model run, the average surviving blue crab had recruited to
the hard-shell fishery, independent of the natural or fishing
mortality regime (Fig. 4), but within a level of natural mortality, the mean size of crabs was larger when exploitation
fractions were lower (e.g., compare Figs. 4b and 4h with
Figs. 4e and 4k). By the third winter of life (31 December,

year x + 2), the average crab had grown to more than

188 mm CW, but only a small percentage of the original cohort had survived (24% in Fig. 4c, 4% in Fig. 4f, 4% in
Fig. 4i, and 0.4% in Fig. 4l). Comparing survival between
the sexes, more males survived than females in all mortality
treatments, likely because all mature females were vulnerable to the fishery, independent of size.
Under all natural mortality regimes, the large majority of
blue crabs harvested were hard-shell ones. Within a level of
natural mortality, however, the percentage of soft-shell or
peeler crabs in the total harvest (by number) increased with
exploitation fraction. For example, when M = 0.9year1, the
soft-shell and peeler crab harvest increased from 13.5% of
the harvest when = 0.19 to 46.6% of the harvest when =
0.60. Because soft-shell and peeler blue crabs are vulnerable
to the fishery at smaller sizes than in the hard-shell fishery,
higher exploitation fractions in the model increased the number and overall percentage of soft-shell and peeler crabs that
were harvested.
For each rate of natural mortality, we estimated 20% (and
the associated F20%), which equaled the harvest rate that protected 20% of the spawning potential of an unfished population. Our 20% estimate decreased with increasing levels of
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2567

Fig. 4. Size frequency distributions of modeled blue crabs (Callinectes sapidus) under different mortality scenarios in the individualbased per-recruit model. Across each suite of panels (left to right), the size distributions during each of the three winters for each mortality scenario are provided: the leftmost panel is the distribution at the initiation of the model in the first winter (1 January, year x + 1),
the middle panel is the distribution in the second winter (1 January, year x + 2), and the rightmost panel is the distribution in the third
winter (31 December, year x + 2). (ac) = 0.32 and M = 0.375year1; (df) = 0.65 and M = 0.375year1; (gi) = 0.30 and M =
0.90year1; (jl) = 0.48 and M = 0.90year1. The mean on each panel represents the mean blue crab CW and N represents the number of blue crabs alive in the model on that day.

natural mortality: 20% = 0.67, 0.57, 0.45, and 0.36 when

M = 0.375, 0.6, 0.9, and 1.2year1, respectively. These exploitation fractions correspond to F20% ranging as high as
1.24year1 when M = 0.375year1 to as low as 0.90year1
when M = 1.2year1. We interpolated between these model
runs to create a contour plot that reveals the proportion of
the virgin, unfished spawning potential that is protected for a
given level of natural mortality and fishing pressure (Fig. 5).
For example, should M be determined to equal 1.0 and managers choose to protect 20% of the virgin spawning potential
(sensu Chesapeake Bay Commission 2001), then the plot reveals a corresponding allowable of 0.43 or less.
We also sought to determine whether adding replicate
model runs and changing the initial cohort size would influence our results. We ran four replicate simulations at the
lowest (M = 0.375year1 and = 0.18) and highest (M =
1.2year1 and = 0.50) mortality regime, expecting that
variability would be greater at the high-mortality regime
given the more frequent use of the uniform random number

generator subroutine (i.e., the source of the stochasticity). As

expected, a 1% difference between the smallest and largest
egg productions among the replicates existed within the lowmortality regime, whereas a 7% difference between the
smallest and largest egg productions among the replicates
existed within the highest mortality regime. Even this 7%
difference, however, had a small impact on the proportion of
maximum spawning potential that was calculated for each
model simulation and used to find 20%. For example, increasing and decreasing the spawning potential by 7% when
M = 0.9 and = 0.46 changed the proportion of maximum
spawning potential from 0.18 to either 0.17 or 0.19. As a result, we were convinced that running several replicate simulations for each level of fishing mortality would not change
the overall results. Next, we ran our model with an initial
abundance of only 150 million crabs rather than 300 million
crabs, given that the abundance of new recruits in the Chesapeake can be highly variable (Sharov et al. 2003). Perrecruit estimates versus fishing mortality revealed the bio 2005 NRC Canada

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Can. J. Fish. Aquat. Sci. Vol. 62, 2005

Table 1. Estimates of 20% (F20% in parentheses) when different subroutines were used in the individual based model and when M =
0.375year1.
Harvest subroutine
Growth subroutine

Complex fishery

Age-dependent fishery

Discontinuous, with individual variation in initial size and growth

Discontinuous, without individual variation
Continuous, without individual variation

0.67 (1.24)
0.67 (1.24)
0.67 (1.33)

0.73 (1.17)
0.73 (1.17)
0.73 (1.17)

Note: Discontinuous growth simulates the realistic molting growth pattern of blue crabs. Continuous growth uses the von Bertalanffy growth equation,
where the parameters were derived from the average daily size of crabs in the discontinuous growth subroutine, with individual variation. In the complex
fishery, vulnerability to the fishery is a function of crab size, sex, shell status, and maturity. In the age-dependent fishery, vulnerability to the fishery was
0.75F in the first year (i.e., age 1) and 0.95F in the second year (sensu Rugolo et al. 1998). The reference points provided elsewhere in the Results are
based on a discontinuous growth subroutine (with individual variation) and a complex fishery in the harvest subroutine.

Fig. 5. Contour plot of the spawning potential per recruit as a

function of (a) natural mortality and exploitation fraction and
(b) natural mortality and fishing mortality.

dependent, as has been used in previous SPPR models for

blue crabs, 20% increased from 0.67 to 0.73 (Table 1);
again 20% was independent of the growth subroutine. Reference points based on fishing mortality also were influenced
by the harvest subroutine, with higher values of F20% under
the complex harvest subroutine than under the agedependent harvest subroutine for a given growth subroutine
(Table 1). In addition, F20% was dependent on the growth
subroutine when the complex harvest subroutine was used.
When the simple harvest subroutine was used, F20% was
equal across all growth subroutines (Table 1).

Discussion

logical reference points to be the same, indicating that our

results are independent of the number of newly recruited
blue crabs for any given year.
Sensitivity to added model complexities
Exploitation-based reference points were sensitive to the
harvest subroutine but were insensitive to the growth subroutine (discontinuous with individual variation, discontinuous
growth without individual variation, continuous growth without individual variation). When we maintained the complex
fishery subroutine (i.e., harvest is a function of size, sex,
shell status, and maturity) from the original model, 20% =
0.67 independent of the growth subroutine (Table 1). However, when we changed the harvest subroutine to make it age

Using estimates of IP and GPM from the literature, we developed an individual-based model for blue crabs that simulated crustacean discontinuous growth and the complexity of
the Chesapeake Bay fishery and allowed for individual variation in blue crab sizes and growth. We used the model to
estimate the level of fishing that would protect 20% of the
spawning potential of an unfished Chesapeake Bay population, the target reference point (Chesapeake Bay Commission 2001). To validate the growth parameters, we first
seeded the model with size distributions of blue crabs in the
field, allowed blue crabs to grow in the model under field
temperatures, and compared size distributions of crabs in the
model with those in the field over the course of 1 year. We
expected some differences between the predicted and field
size distributions, given uncertainty in natural mortality and
the simplifications inherent in our model (i.e., lack of temporal variation in fishing mortality or spatial variation in
temperature, growth, and fishing mortality). However, given
these limitations, our predicted distributions compared favorably with the field distributions, even under extremely low
and high scenarios of harvest. Applying our model to a hypothetical cohort of newly recruited blue crabs, under a
range of natural mortality regimes (M = 0.3751.2year1),
we determined that protecting at least 20% of the spawning
potential of an unfished population required a management
regime that limited to range between 0.36 (when M =
1.2year1) to 0.67 (when M = 0.375year1). Finally, we
found that 20% was influenced by the harvest subroutine of
our model but was not influenced by the growth subroutine
or the presence of individual variation. Our estimate of F20%
was influenced by both the harvest subroutine and the
growth subroutine.
A few caveats to our model should be noted. First, the
spawning potential calculation in our model uses only fe 2005 NRC Canada

Bunnell and Miller

male size and abundance to predict egg production and thus

assumes that males are not limiting. Although a high percentage of females are mated in the Chesapeake Bay, recent
evidence suggests that the quantity of sperm delivered by
the male is declining, which may limit egg fertilization
(Hines et al. 2003). Second, we modeled natural mortality as
independent of blue crab size, despite evidence that among
crabs less than 70 mm CW, smaller crabs are more likely to
die from predation or cannibalism than larger crabs (Wilson
et al. 1987; Hines and Ruiz 1995). We completed simulations with a size-dependent natural mortality for blue crabs
less than 70 mm CW, but the high rate of mortality among
small crabs resulted in sustainable between 0.02 and 0.05,
as relatively few individuals from the cohort recruited to the
fishery. We removed this size dependency from our model
when we compared these exploitation fractions with those
measured in the winter dredge survey, which are between
0.40 and 0.70 (Sharov et al. 2003). Thus, even though sizedependent mortality among juvenile crabs likely occurs in
the field, including it in the model yielded unrealistic results.
Individual-based models were first used in fisheries ecology to explain the survival outcomes of fish cohorts. These
models revealed that consideration of the initial variation
among individual sizes and inclusion of time- or sizevarying relationships could lead to a different distribution of
survivor sizes than if a population of homogenous average
individuals with time- or size-independent relationships
were simulated (e.g., DeAngelis and Coutant 1982).
Individual-based models have enjoyed much broader application in recent years, including per-recruit models for fisheries stock assessment. Previous individual-based per-recruit
models have used the flexibility offered by this approach to
incorporate multiple spawning by weakfish (Cynoscion
regalis) within a season (Lowerre-Barbieri et al. 1998), allow for size-selective mortality among juvenile Atlantic cod
(Gadus morhua) that have individually varying growth rates
(Kristiansen and Svsand 1998), and apply spatially explicit
rates of fishing mortality for sea scallops (Placopecten
magellanicus) in the Atlantic Ocean (Hart 2001).
We chose an individual-based modeling approach for blue
crab for three reasons. First, we wanted to include variation
both in initial sizes within the cohort and in the GPM and IP
that has been observed in field and laboratory studies. The
sensitivity model runs revealed that including this individual
variation yielded the exact same reference point as an
individual-based model that included all blue crabs of the
same average size and the same average growth rate. Because harvest and IP were size dependent, we had expected
that including this individual variation would lead to different results than if we ignored it. Specifically, we had hypothesized that small individuals in the cohort might survive at a
higher rate because of a later recruitment to the fisheries,
and this asymmetrical survival would influence spawning
potential. Considering that spawning potential is a function
of blue crab size and the internal amount a (i.e., number still
alive) at the time of spawning, this hypothesis was incorrect
for two reasons. First, there was no relationship between initial size and size at maturity, even though large individuals
matured earlier than small ones. Larger individuals in the cohort accumulated more harvest mortalities, whereas smaller
individuals accumulated more natural mortalities. The latter

2569

is explained by smaller individuals having larger internal

amounts during the period when small individuals had not
yet recruited to the fishery. As a result of no differences in
total mortality and blue crab sizes at spawning among initially small and larger individuals, the differences in spawning potential were minimized.
Second, an individual-based approach permitted us to
simulate the true biology of the crustacean molt process.
However, the sensitivity analyses revealed 20% to be insensitive to either growth model (i.e., discontinuous growth versus continuous growth). Our estimate of F20% was slightly
influenced by the growth subroutines when the complex harvest subroutine was used: F20% was 6% higher in the continuous growth subroutine than in the discontinuous subroutine.
When the simple harvest subroutine was used, F20% was insensitive to the growth subroutines. Overall, these results
imply that blue crab per-recruit models that rely on a von
Bertalanffy growth subroutine should provide reference
point estimates that are as reliable as results from a perrecruit model that more accurately models blue crab growth
as discontinuous. This is an important finding because many
conventional stock assessment models rely on a von
Bertalanffy growth subroutine and these results provide
greater credibility for using this growth subroutine for crustaceans.
Third, we used an individual-based modeling approach
because it enabled us to simulate the complexity of the
Chesapeake Bay blue crab fishery, where regulations are a
function of crab size, sex, shell status, and maturity. Here,
the sensitivity results revealed that both 20% and F20% are
influenced by the harvest subroutine: within a growth subroutine, 20% was 9% greater with the age-based harvest subroutine than with complex harvest one, whereas F20% was
6%14% greater with the complex harvest subroutine than
with the age-based one. Before explaining this discrepancy,
it is important to note that the numbers and sizes of blue
crabs alive to spawn were roughly similar in the two harvest
subroutine models, which resulted in the equal spawning potentials between the two models. Thus, the reference point
differences are explained by differences in natural mortality
and harvest that occurred in the two models. In other words,
how can the two models have the same number of spawning
females when more females were harvested from the agebased harvest model than from the complex harvest model?
In the first year of the model, the number of crabs harvested
in the age-based harvest model was considerably higher.
With the complex harvest subroutine, most individuals did
not recruit to the hard-shell fishery until October of the first
year, meaning that individuals were only briefly vulnerable
to the fishery in earlier months (i.e., as peeler or soft-shelled
crabs). Conversely, in the age-dependent harvest model, all
age-1 crabs in the same year received 75% of the nominal
fishing mortality during the fishing season (sensu Rugolo et
al. 1998). Over the 2 years of the simulation, this first-year
difference resulted in many more crabs being vulnerable to
natural mortality in the complex harvest simulation. Hence,
despite the higher harvest in the age-based model, the higher
levels of natural mortality in the complex harvest model resulted in near equal numbers of spawning females between
the two models. The higher natural mortality in the complex
harvest model also resulted in a higher estimate of total mor 2005 NRC Canada

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tality than in the age-based one. Because total mortality influenced fishing mortality more than the catch (see eq. 7),
the higher total mortality explains the higher estimate of
F20% in the complex harvest subroutine. This examination
reveals how the timing of recruitment to the fishery can influence the reference point estimate. Because of the complexity of the Chesapeake Bay fishery, assessment models
that maintain this complexity should be favored over previous models that simplify the fishery into one that is based
only on age.
Our model produced an estimate of the target biological
reference point that is considerably higher than for previous
models. Currently, the Chesapeake Bay fishery is managed
with a target F20% = 0.7 (Chesapeake Bay Commission
2001), which is based on M = 0.375year1. This value is
slightly smaller than the 0.8 estimate from Rugolo et al.
(1998) and considerably smaller than our estimate of 1.24,
both assuming M = 0.375year1. To understand these differences, we used the NOAA Fisheries Toolbox Yield-PerRecruit model, an age-based model similar to the approach
of previous models. We should note that this model uses biomass of mature crabs to estimate spawning potential rather
than egg production as we used. Because egg production and
blue crab size are linearly related (Prager et al. 1990), however, these results should be much more similar than a situation where egg production is nonlinearly related to size.
First, we used the age-based model to attempt to duplicate
our results. We used the mean crab weight at age from our
model output, the same age-dependent harvest schedule used
in the sensitivity analysis, and the same proportion of F
(0.635) and M (0.75) before spawning reported in Rugolo et
al. (1998). For maturity schedule, we assigned 0% for age-1
crabs and 100% for age-2 crabs because eggs were produced
only in the second year of the simulation. These inputs yielded
an F20% = 1.19, well within the range of our individualbased modeling results (baseline F20% = 1.25, F20% from
sensitivity analyses = 1.171.33). This exercise provided a
validation of our individual-based modeling approach.
Second, we used Rugolo et al. (1998) and data that were
used to generate the Chesapeake Bay Commission (2001)
target reference point to recreate their respective model results and then attempt to understand why their results differed from ours. Compared with our inputs, those models
had different mean sizes at ages (generally larger than ours),
included more age classes (eight versus two in ours), and
had a different maturity schedule (several age classes with
mature females compared with only one age class in ours).
By changing only their maturity schedule to match ours, we
were able to generate F20% = 1.11 for both models, considerably higher than their original results and much closer to our
estimate.
This exercise revealed maturity schedule to have a clear
influence on spawning potential based reference points for
blue crabs in Chesapeake Bay. In the Rugolo et al. (1998)
model, they assumed the following schedule of maturity:
10% of age 1, 90% of age 2, 100% of age 3, 50% of age 4,
10% of age 5, and 0% of ages 68. Rugolo et al. (1998) acknowledged that a maturity schedule that more quickly reduced the percentage of mature females after the first full
year of spawning (i.e., 0% maturity of ages 48) was more

Can. J. Fish. Aquat. Sci. Vol. 62, 2005

biologically realistic than the one they chose. Nonetheless,

they chose their maturity schedule because it provided
smaller estimates of F20%, making it risk averse. Since those
analyses, however, aging blue crabs by the accumulation of
lipofuscin in their eyestalks has revealed that 90% of blue
crabs larger than 120 mm CW in Chesapeake Bay are age 1
and younger (Ju et al. 2003). The remaining 10% were age
2, although there was a statistically indistinguishable mode
of crabs within this percentage that may have been age 3
(Ju et al. 2003). Because of the demographic results of Ju et
al. (2003) and the conventional wisdom that female maturity
occurs sometime between age 1 and age 2 (Hines et al.
2003), we structured our model so that females mature at
age 1 and spawn at age 2, and at age-3 females are not included. Although our estimates of F20% and 20% are higher
than for previous models because of our truncated maturity
schedule, this schedule most accurately reflects current
knowledge of blue crab demography and maturity.
Finally, it is not surprising that our biological reference
point estimates were dependent on the assumed rate of natural mortality (sensu Beverton and Holt 1957). As natural
mortality increased, fewer blue crabs were available to be
harvested, which effectively led to an inverse relationship
between 20% and M as well as between F20% and M. Previous blue crab modeling efforts have assumed an M =
0.375year1, which was based on an assumed maximum
crab age of 8 years (Rugolo et al. 1998). Recently, Hewitt
and Hoenig (2005) suggested that even were the maximum
blue crab age to be 8 years, an estimate of M = 0.375year1
remains too low because of methodological concerns. As a
result of continued uncertainty in maximum age, we explored the use of life history invariants for finfishes as an alternative estimate for blue crab natural mortality. As
summarized in the Methods, the invariants provided a range
of M between 0.74 and 1.8year1, but we did not consider
estimates of M greater than 1.2year1 because field-based
estimates of Z have never exceeded 1.5year1 from 1990 to
2002. A revised Chesapeake Bay stock assessment is currently underway, and they have recommended that managers
use M = 0.9year1 for two primary reasons (T.J. Miller, personal communication). First, nearly all life-history invariant
estimates of M include 0.9year1. Second, the recent population decline observed in the Chesapeake Bay necessitated a
higher estimate of natural mortality than previous stock assessment models have used.
Assuming that a 20% spawning potential ratio is sufficient
to sustain the blue crab population and M = 0.9year1, our
model reveals 20% = 0.45 and F20% = 1.02. We advocate using the 20% reference point because annual estimates of exploitation factor can be calculated directly from the absolute
abundance data measured in the winter dredge survey and
from harvest monitoring data by the state management agencies. Hence, managers can measure the impact of the fishery
using exploitation factor directly and avoid using fishing
mortality, which would require some estimate of natural
mortality. From 1990 through 2003, has ranged from 0.34
to 0.71 (L. Fegley, Maryland Department of Natural Resources, 580 Taylor Avenue, Annapolis, MD 21401, USA,
personal communication), and exceeded 20% in 64% of
the years assuming M = 0.9year1. With lower estimates of
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Bunnell and Miller

natural mortality, our model predicts that exceeded 20% in

fewer years (29% of years when M = 0.6year1, 7% of years
when M = 0.375year1).
In conclusion, our individual-based modeling approach
provided estimates of F20% and 20% for a range of levels of
natural mortality. For a given level of natural mortality, we
found that our estimate of F20% was at least 55% larger than
previous estimates based on an age-based SPPR. This difference was explained by differences in the maturity schedule
of blue crabs. Limiting the percentage of females that reproduce in older age classes, which appears realistic given the
current knowledge of blue crab age structure (Ju et al. 2003),
leads to higher reference point estimates (sensu Rugolo et al.
1998). We found that some advantages of this individualbased approach, including incorporation of individual variation in blue crab size and growth and simulation of the real
molting growth process, did not provide reference point estimates that differed from a more simplified model where all
individuals were of average size and grew at the same average rate through a continuous growth process. Conversely,
the capacity to simulate the complexity of the Chesapeake
Bay blue crab fishery in the individual-based model did result in reference point estimates that were about 10% different than a more simplified model where recruitment to the
fishery was simply age dependent. Finally, a growing body
of evidence suggests that M is ~0.9year1 rather than
0.375year1 used in previous modeling efforts. With this assumption, the model predicts that 0.45 is necessary to
protect at least 20% of the unfished spawning potential.
With available field-based estimates of exploitation factor,
we found that the commercial fishery has exceeded 20% in
64% of years since 1990. Hence, these results join others
(Miller 2001, 2003) that have implicated the commercial
fishery as a likely contributor to the recent blue crab population decline in Chesapeake Bay.

Acknowledgments
We thank the numerous participators of the VIMS Juvenile Fish and Blue Crab Trawl Survey and the Chesapeake
Bay Winter Dredge Survey for collecting these valuable
data. We thank Kenny Rose for advice on modeling superindividuals. The comments of Dave Hewitt and three anonymous reviewers greatly improved this paper. This is contribution No. 3862 of the University of Maryland Center for
Environmental Science Chesapeake Biological Laboratory.
This work was supported by grants from Maryland Sea
Grant (R/F-93B) and by the NOAA Coastal Ocean Program
(NA17OP265).

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