Estuaries and Coasts
DOI 10.1007/s12237-016-0127-8
Population Structure of Adult Blue Crabs, Callinectes
sapidus, in Relation to Physical Characteristics in Barnegat Bay,
New Jersey
Paul R. Jivoff 1 & Jennifer M. Smith 1 & Valerie L. Sodi 2 & Stacy M. VanMorter 3 &
Kathryn M. Faugno 4 & Amy L. Werda 5 & Margaret J. Shaw 3
Received: 26 August 2015 / Revised: 9 June 2016 / Accepted: 22 June 2016
# Coastal and Estuarine Research Federation 2016
Abstract Blue crabs (Callinectes sapidus) are an important
species in coastal or lagoonal estuaries where adult population
characteristics may differ as compared to drowned-river estu-
aries. Barnegat Bay, in southern New Jersey, is composed of
two large embayments: one without and one with a salinity
gradient. We tested the influence of physical characteristics on
the abundance, sex ratio, and size of adult blue crabs and
examined variation in measures of reproductive potential
(e.g., sperm stores) in both sexes in Barnegat Bay from June
to September, 2008–2009. Population structure was distinct
between the embayments due to sex-specific responses to sa-
linity: male abundance was negatively correlated with salinity
whereas adult females were more abundant in high salinity
because of proximity to Barnegat Inlet. This produced high
sex ratios in low salinity areas and low sex ratios in high
salinity areas. Summer was a growing season for adult males
while in late summer-early fall, juvenile males recruited to the
Communicated by Nancy L. Jackson
* Paul R. Jivoff
pjivoff@rider.edu
1
Department of Biology, Rider University, 2083 Lawrenceville Rd,
Lawrenceville, NJ 08648, USA
2
Department of Biochemistry and Molecular Biology, Drexel
University, College of Medicine, Philadelphia, PA 19102, USA
3
Rutgers University Marine Field Station, 800 c/o 132 Great Bay
Blvd, Tuckerton, NJ 08087, USA
4
Greenwich Physical Therapy Center, 1171 East Putnam Ave,
Riverside, CT 06878, USA
5
943 Nassau St, North Brunswick, NJ 08902, USA
adult size class. The spawning season lasted from May to
August and ovigerous females were concentrated near the
inlets. Information on female sperm stores and ovarian devel-
opment identified two cohorts of adult females: females that
will spawn in the current summer and females that will not
spawn until the following summer. Thus, not all adult females
near the spawning grounds were members of the current
spawning stock. This suggests that annual estimates of spawning
stock size which overlook the proximity of females to spawning
are overestimating the current spawning stock in Barnegat Bay
and other estuaries.
Keywords Blue crab . Abundance . Sex ratio . Size .
Reproductive potential . Salinity
Introduction
Blue crabs support one of the most important commercial
fisheries in New Jersey (Kennish et al. 1984; Stehlik et al.
1998) and throughout the mid-Atlantic region (Jordan 1998).
Blue crab populations in drowned-river estuaries such as
Chesapeake Bay and Delaware Bay, the traditional locations
of important fisheries, have experienced considerable fluctua-
tions including temporary periods of significant decline (Abbe
and Stagg 1996; Cole 1998; Kahn et al. 1998). These reduc-
tions may stem from a number of factors including loss or
degradation of habitat for recruits and juveniles (Lipcius
et al. 2005) as a result of land use patterns (King et al.
2005), reduced water quality (Mistiaen et al. 2003), and signif-
icant natural and fishing mortality (Lipcius and Stockhausen
2002). In the mid-Atlantic region, during periods of reduced
blue crab catches in Delaware Bay, the relative importance of
New Jersey blue crab populations increased tenfold in terms
of both commercial landings and economic value (NOAA

fisheries unpublished data). Some of this increase comes from
New Jersey estuaries other than Delaware Bay. For example,
in the past decade, the percentage of New Jersey’s blue
crab catch from Barnegat Bay has quadrupled (New Jersey
Department of Environmental Protection fisheries unpub-
lished data). As the relative importance of blue crab popula-
tions in coastal or lagoonal estuaries like Barnegat Bay in-
creases, the extent of fishing effort and the potential for user
conflicts may also increase. Coastal or lagoonal estuaries can
differ from drowned-river estuaries in physical features (e.g.,
salinity regime, depth) that impact the population characteris-
tics of blue crabs. Therefore, it is critical to gather information
about the status and characteristics of the population of blue
crabs in estuaries like Barnegat Bay.
Blue crab population characteristics such as abundance,
size distribution, and sex ratio are influenced by both biotic
and abiotic features (Hines 2007). Temporal and/or spatial
variation in the blue crab population characteristics can be
influenced by factors including blue crab molt stage (Shirley
et al. 1990) and behavior (Ramach et al. 2009), vegetative
cover (Lipcius et al. 2005; Thomas et al. 1990), and intraspe-
cific interactions (Dittel et al. 1995). Physical aspects of the
environment such as temperature (Harding and Mann 2010);
dissolved oxygen, particularly hypoxic conditions (Bell and
Eggleston 2005); and local land-use patterns (King et al.
2005) also influence blue crab population characteristics.
Salinity is particularly influential in a variety of aspects of blue
crab life history leading to variation in abundance, sex ratio,
and size. Blue crabs can survive and grow successfully in a
wide range of salinities (Cadman and Weinstein 1988; Stickle
et al. 2007); however, especially in winter, salinity can interact
with temperature to influence mortality (Bauer and Miller
2010; Rome et al. 2005). The distribution of adults is moder-
ated by salinity resulting in spatial variation in the population
sex ratio, with adult females more prevalent at higher salinities
particularly during the spawning season (Archambault et al.
1990; Harding et al. 2010; Schaffner and Diaz 1988). In the
presence of an extensive salinity gradient, low salinity areas
represent important habitats for growth (molting) (Fisher
1999; Hines et al. 1987; Posey et al. 2005) and refuge from
predators (Posey et al. 2005) and therefore influence variation
in the spatial distribution of different-sized crabs. In the ab-
sence of extensive spatial variation in salinity, a characteristic
of lagoonal estuaries, blue crab population characteristics typ-
ically show relatively more temporal variation than spatial
variation (Buchanan and Stoner 1988; Murphy and Secor
2006). The lack of spatial variation, due to the absence of a
salinity gradient, may therefore influence a variety of aspects
of blue crab life history in lagoonal estuaries. Barnegat Bay, in
southern New Jersey, provides an interesting system to test the
influence of physical characteristics (especially salinity) be-
cause it is composed of two large continuous embayments
(see Fig. 1): one without and one with a salinity gradient.
Estuaries and Coasts
Variation in population characteristics of adults can have
important consequences on different aspects of blue crab life
history, particularly reproduction including competition for
access to mates (Jivoff 1997b; Jivoff and Hines 1998b;
Kendall and Wolcott 1999) and forms of reproductive invest-
ment (Jivoff 1997a; Kendall et al. 2002) which impact the
timing of reproduction and reproductive output (Hines et al.
2003). We know very little about the reproductive potential of
blue crab populations in New Jersey as compared to other
estuaries in the mid-Atlantic (Hines et al. 2003) or further
south along the east coast (Darnell et al. 2009; Gelpi et al.
2009; Ogburn and Habegger 2015) where offshore areas are
also important spawning grounds (Gelpi et al. 2009; Ogburn
and Habegger 2015). The reproductive potential of blue crab
populations may be an indicator of intense fishing pressure
(Carver et al. 2005; Hines et al. 2003; Jivoff 2003) and thus
represents important information for management efforts
(Miller et al. 2011; Prager et al. 1990).
Barnegat Bay is a lagoonal estuary composed of two con-
tinuous embayments that differ in physical characteristics,
particularly salinity, and thus provides a unique opportunity
to test the influence of physical characteristics on blue crab
population structure. The objective of this study was to test the
influence of physical characteristics on adult blue crab popu-
lation structure (e.g., abundance, sex ratio, size) by examining
temporal (e.g., annual and monthly) and spatial variation (e.g.,
within and between embayments) in blue crabs as well as
measures of reproductive potential (e.g., sperm stores of both
sexes, female brood production) in both sexes of crabs in
Barnegat Bay. Our working hypothesis is that physical char-
acteristics will influence aspects of population structure which
in turn will influence measures of reproductive potential.
Specifically, we hypothesize that lower salinities will produce
a greater abundance of males (and thus male-biased sex ratios)
which will lead to greater sperm stores in females as a result of
greater sexual competition among males.
Methods and Materials
Study Area
Barnegat Bay is a narrow (2–6 km wide) lagoonal estuary
extending approximately 67 km along the central New
Jersey coastline between Tuckerton and Pt. Pleasant, NJ (39°
31′ N, 74° 02′ W to 40° 06′ N, 74° 02′ W) composed of three
continuous embayments: Barnegat Bay, Manahawkin Bay,
and Little Egg Harbor (Fig. 1). For this study, BLittle Egg
Harbor^ will include both the Manahawkin Bay and Little
Egg Harbor embayments. Barnegat Bay is separated from
the Atlantic Ocean by a barrier island complex, including
Long Beach Island and Island Beach State Park, with two
inlets to the Atlantic Ocean: Barnegat Inlet, about 35 km south
Estuaries and Coasts
Fig. 1 Map of sampling locations in Barnegat Bay
of Pt. Pleasant, and Little Egg Inlet, at the southern boundary
(Fig. 1). Barnegat Bay is relatively shallow (1–6 m deep) with
most of the western side of the Bay (where sampling for this
study occurred) containing areas 1–3 m deep as compared to
the eastern side (∼1 m deep) (Kennish 2001). Shallow water
combined with tide- and wind-driven water movement result
in a vertically well-mixed system with weak salinity and ther-
mal stratification (Kennish 2001). Many small tributaries
drain into Barnegat Bay from the mainland; however, most
of the freshwater input comes from tributaries north of

Barnegat Inlet including Toms River and Cedar Creek (Kennish
2001). This creates a narrow (10–14 ppt) salinity gradient be-
tween Barnegat Inlet and Toms River within an entire system
that has an average summertime salinity of 20.5 ppt (Kennish
2001). Benthic sediments along the western side of the Bay are
composed of sand or a varying mixture of sand and mud
(Kennish 2001).
Field Sampling
This study tested the influence of physical characteristics on
adult blue crab population structure in Barnegat Bay by ex-
amining temporal and spatial variations in abundance, sex
ratio, size, and measures of reproductive potential in both
sexes. Focused sampling along the western shore (see Fig. 1)
was beneficial because it facilitated replication of sampling
sites (by sampling in multiple days per month) and minimized
variation in depth, bottom type, and benthic habitat among
sites along the entire north-south axis of the estuary, thus em-
phasizing physical characteristics. The drawback to this de-
sign is that spatial variation in depth, bottom type, benthic
habitats, and proximity to the ocean does exist (Kennish
2001) which can influence the population characteristics of
blue crabs in this system (Jivoff and Able 2001; Szedlmayer
and Able 1996). As such, after the completion of the present
study, we modified our sampling design so that it now incor-
porates spatial variation in both physical characteristics and in
benthic habitats to provide a more comprehensive understand-
ing of adult blue crabs in the bay.
In each year (2008–2009), we sampled adult blue crab
population structure (abundance, size composition, sex ratio)
using baited (menhaden) commercial-style traps (60 cm
high × 60 cm wide × 60 cm deep with 3.8-cm hexagonal mesh)
fished daily for four consecutive days, every other week from
June to August and 1 day every other week in September.
Barnegat Bay was divided into seven approximately equal-
sized sampling areas (each ∼8 km long) with four replicate sites
in each area (Fig. 1). Each sampling day, four traps were ran-
domly assigned to one of the four sites in each sampling area
(and placed at least 50 m apart from one another). Physical
characteristics near the first and last trap in each site including
depth, salinity, temperature, and dissolved oxygen were taken
with a hand-held YSI data logger (model 6800).
This study concentrated on adult crabs; therefore, traps
were used as a collecting gear because they are efficient at
capturing blue crabs in the adult size range (Guillory 1998)
and they are more efficient at capturing adults than other types
of gear (e.g., trawl) (Bellchambers and de Lestang 2005).
Traps have been criticized as a gear for estimating crab abun-
dance because variation among traps (e.g., soak time, presence
of bait and/or conspecifics) (Sturdivant and Clark 2011) and in
crab behavior (e.g., aggressiveness, feeding) may influence
free crabs from entering (Murray and Seed 2009; Sturdivant
Estuaries and Coasts
and Clark 2011) as well as captured crabs from escaping
the trap (Sturdivant and Clark 2011). In another Portunid
(Portunus pelagicus), there is also some evidence that
traps preferentially capture males as compared to females
(Bellchambers and de Lestang 2005); however, the sexes are
equally attracted to traps and put forth equal effort to enter
traps (Smith and Sumpton 1989). In this study, traps were used
systematically to minimize variation in trap soak time (24 h
for all traps), bait content (renewed daily), trap integrity (e.g.,
traps checked daily and damages repaired immediately), and
the extent of fouling on traps (traps were power-washed peri-
odically). Sturdivant and Clark (2011) found no evidence in
blue crabs that aggressive interactions with captured crabs
deterred free crabs from entering the traps. Murray and Seed
(2009) found that traps were especially ineffective when
colder temperatures reduced green crab feeding time. During
the current study, all of the physical variables were well within
tolerance limits of blue crabs suggesting the crabs were be-
having normally (e.g., feeding) throughout the study period.
Crabs were separated by trap in moist burlap bags, returned
to the Rutgers University Marine Field Station, and assessed
for carapace width between the tips of the lateral spines (to the
nearest mm), age, sex, sexual maturity (see below), molt stage
(see below), limb loss and regeneration, and ovigerous stage
(adult females only). Sexual maturity and molt stage were de-
termined using previously established methods (Jivoff 1997b).
Briefly, molt stage was determined by examining the
propodus on the fifth appendage for evidence of epidermal
retraction and color variation (Van Engel 1958) and, for re-
cently molted (postmolt) crabs, the relative hardness of the
newly formed carapace (Freeman et al. 1987). Males were
designated as adults according to the following criteria: the
secondary pleopods lay within the primary pleopods (intro-
mittent organs), the penes were inserted into the secondary
pleopods, and the abdomen easily pulled away from the ster-
num (Van Engel 1990). Crabs from these collections were also
used for measurements of reproductive potential.
Reproductive Potential Studies
Each sampling day, a subsample of crabs from each sampling
area in five size categories of each sex were frozen for subse-
quent dissection and measurement of reproductive potential:
weight (to the nearest 0.01 g) of the spermatophore and sem-
inal fluid sections of the vasa deferentia in males and weight
(to the nearest 0.01 g) of the seminal receptacles and ovary,
ovarian developmental stage, and brood stage in females using
previously established techniques (Hines et al. 2003; Jivoff
1997b). In females, seminal receptacle weight is a proxy for
the amount of sperm available for fertilization (Ogburn et al.
2014) and ovary weight provides an estimate for the size
(number of eggs) of the female’s next brood. Briefly, female
seminal receptacles and ovaries were removed and separated
Estuaries and Coasts
from one another using standard dissecting tools. Male vasa
deferentia were removed and the spermatophore and seminal
fluid sections were separated with standard dissecting tools
using the following criteria: diameter and color and texture
of the vasa deferentia section (indications of content) and by
applying pressure at the juncture of the spermatophore and
seminal fluid sections of the vasa deferentia to remove addi-
tional spermatophores.
Data Analysis
In each year, for each sampling area, site, month, and day,
crabs from the replicate traps were combined to produce
Bcomposite^ daily samples (preliminary analysis indicated
that trap did not explain a significant amount of variation in
any dependent variable). Physical variables were also treated
this way. Abundance of the crabs was converted to catch per
unit effort (CPUE) because the number of crabs caught was
not always based on an equal number of traps per site (e.g.,
due to missing or lost traps between sampling days) or days
per month. For each sampling area, site, month, and day,
CPUE was calculated as the number of crabs captured divided
by the number of traps used. Separate CPUE values were
calculated for males and females. For each sampling area, site,
month, and day, sex ratio was calculated as the number of
males divided by the number of females.
Temporal and spatial variation in abundance (CPUE of
males, adult, prepubertal and ovigerous females), size, and
sex ratio were analyzed using a general linear model (GLM)
with year (2008 and 2009), month (6–9), and embayment
(Little Egg Harbor and Barnegat Bay) as factors (including
each two-way and the three-way interaction) as well as site
(1–4) nested within the sampling area (1–7) and sampling area
nested within the embayment. In addition, the physical vari-
ables (temperature, salinity, dissolved oxygen, and depth)
were included in the model as covariates as well as the inter-
action between each physical variable and each factor. In our
case, these interaction terms not only serve to test the assump-
tion of no interaction between the covariates and independent
variables (Sokal and Rohlf 1981) but also to test for differ-
ences in the relative importance of physical variables on blue
crab population structure between the embayments (i.e., an
interaction between a physical variable and embayment)
(Engqvist 2005). If an interaction occurred, further analysis
(e.g., within particular values of the physical variable) ex-
plored the nature of the interaction. Temporal and spatial var-
iation in physical characteristics (temperature, salinity, dis-
solved oxygen, and depth) were analyzed using a GLM with
year (2008 and 2009), month (6–9), and embayment (Little
Egg Harbor and Barnegat Bay) as factors (including each two-
way and the three-way interaction) and site (1–4) nested with-
in the sampling area (1–7) and sampling area nested within the
embayment. Size distributions (sexes analyzed separately)
were compared among sampling areas and between embay-
ments using the Kolmogorov-Smirnov (KS) tests. The relative
distribution of size categories between the embayments was
compared using chi-square tests on abundances adjusted for
sample size. The weights of the reproductive tissues of each
sex were analyzed with the same GLM model as the physical
characteristics plus the following variables as covariates: car-
apace width (for both sexes), ovary weight for females, and
sex ratio for males. All GLM and KS analyses were
interpreted using P values adjusted for the number of compar-
isons (0.05/no. of comparisons). CPUE data were square root
transformed and other dependent variables were log (10)
transformed to produce homogeneous variances confirmed
by Levene’s test. The Tukey multiple comparisons test
(Tukey HSD) was used to identify significant differences
among multilevel independent variables with P values adjust-
ed for the number of comparisons (0.05/no. of comparisons).
The temporal and spatial distribution of ovigerous females,
and of females in different ovary stages, was analyzed with
chi-square. Statistical analyses were performed using SYSTAT
(SYSTAT 1992) and unless otherwise indicated, means ±1 SE
are shown.
Results
Physical Characteristics
Temperature varied by year (2008 > 2009) and month (Fig. 2a)
but not embayment (Fig. 2b; Table 1). The monthly differ-
ences in temperature varied by year (Table 1) based on how
rapidly peak temperatures were reached (Fig. 2a); however,
the greatest monthly temperature differences were, on aver-
age, less than 5 °C.
Salinity varied by year (2009 > 2008), month (Fig. 2c), and
embayment (Little Egg > Barnegat) (Fig. 2d; Table 1). Monthly
differences in salinity occurred between years (Table 1) but
apparently only due to a wetter spring in 2008 (Fig. 2c). In
contrast, monthly differences in salinity between the embay-
ments (Table 1) were the result of a consistent trend: average
salinities in Little Egg Harbor were greater than in Barnegat
Bay. There was also a different pattern of variation among the
sampling areas within the embayments (Table 1). In Little Egg
Harbor, salinities were similar among the areas although the
salinity in area 1 was slightly but significantly greater than that
in area 2 (P < 0.001) while in Barnegat Bay, a salinity gradient
was apparent; salinity significantly declined between each ar-
ea going from areas 4 to 7 (7 < 6 < 5 < 4, all comparisons
P < 0.001).
Dissolved oxygen varied by year (2009 > 2008) and
month (Fig. 2e) but not embayment (Fig. 2f; Table 1).
Annual differences in the monthly variation in dissolved
oxygen (Table 1) were solely driven by dissolved oxygen
 Estuaries and Coasts

Fig. 2 Monthly averages (±1 SE) of physical variables in each year: a temperature, c salinity, e dissolved oxygen, g depth relative to sea level. Average
(±1 SE) physical variables in each sampling area: b temperature, d salinity, f dissolved oxygen, h depth relative to sea level

values in July of each year (Fig. 2f); however, on average, Depth varied by year (2009 > 2008) and embayment (Little
dissolved oxygen was near or above saturation levels in Egg > Barnegat) (Fig. 2h), but not by month (Fig. 2g; Table 1).
each month. In each year, Little Egg Harbor typically had greater depths
Estuaries and Coasts

Table 1 Summary of statistical

results for the effect of each major Temperature (°C) Salinity (ppt) Dissolved Depth (m)
independent variable, including oxygen (mg/L)
interactions, on each physical
variable Independent variable F (276) P value F (276) P value F (274) P value F (280) P value

Year 42.1 0.0001 8.3 0.004 22.7 0.0001 17.1 0.0001

Month 66.4 0.0001 34.1 0.0001 16.2 0.0001 1.4 0.23
Embayment 2.4 0.12 559.7 0.0001 0.8 0.37 8.6 0.004
Area (embayment) 0.8 0.57 251.1 0.0001 0.3 0.90 28.5 0.0001
Year × month 77.6 0.0001 7.9 0.0001 76.7 0.0001 0.6 0.60
Year × embayment 0.1 0.72 25.6 0.0001 0.7 0.41 10.4 0.001
Year × month × 0.6 0.59 1.6 0.18 2.2 0.09 0.4 0.72
embayment

The F statistic (error degrees of freedom in parenthesis) and P value are shown

than Barnegat Bay (Table 1). The depths of the sampling areas
also varied within each embayment (Fig. 2f; Table 1); howev-
er, these differences were, on average, no greater than 40 cm.
Tide at the time of sampling is also a contributing factor to the
spatial variation in depth. The largest depth differences oc-
curred between areas that were sampled more often near high
tide as compared to areas sampled near low tide.
Adult Population Structure
Abundance
A total of 10,142 blue crabs were captured over the course of
this study: 7245 adult males, 245 juvenile males, 2117 adult
females, 532 prepubertal females, and 3 juvenile females.
Overall, male abundance was greater than that of females
(F1, 1233 = 533.5, P < 0.0001). Interestingly, the sexes exhib-
ited similar abundance patterns among the sampling areas
within Little Egg Harbor but divergent abundance patterns
across sampling areas within Barnegat Bay (F5, 1233 = 25.8,
P < 0.0001), suggesting that the physical differences between
the embayments may influence these sex-specific patterns. In
particular, salinity should help explain the differences between
the sexes since it is the only physical variable that exhibits
significant and systematic variation between the embayments.
Due to these significant differences between the sexes, the
abundance of males and females will be treated separately
below.
The abundance of males was influenced by embayment
(Barnegat > Little Egg, F1, 250 = 20.2, P < 0.0001), and there
was a different pattern of variation among the sampling areas
within the embayments (F5, 250 = 13.4, P < 0.0001). In Little
Egg Harbor, male abundance did not vary among the areas
whereas in Barnegat Bay, male abundance increased signifi-
cantly at each area between 4 and 7 (4 < 5 < 6 < 7, all com-
parisons P < 0.0001) (Fig. 3). Male abundance also exhibited
a different abundance pattern in each embayment based on
salinity (embayment × salinity interaction, F1, 250 = 31.9,
P < 0.0001): male abundance (square root transformed) was
not related to salinity in Little Egg Harbor (Y = −0.005X +
2.24, P = 0.88) whereas male abundance increased with de-
creasing salinity in Barnegat Bay (Y = −0.05X + 3.40,
P = 0.002). This interaction (produced by the lack of salinity
values less than 23 ppt in Little Egg Harbor) facilitated an
analysis only at those salinities shared most by both embay-
ments (25–27 ppt). This analysis included all of Little Egg
Harbor plus sampling areas 4 and 5 from Barnegat Bay and
showed that there was no significant variation in male abun-
dance. Thus, the distinguishing feature that drives the differ-
ences in male abundance between the embayments is the pres-
ence of lower salinity areas in Barnegat Bay.
The abundance of females varied with the percentage of
prepubertal females, such that female abundance (square root
transformed) decreased as the percentage of prepubertal fe-
males increased (Y = −0.013X + 1.96, P < 0.0001). Thus,
separate analyses were run for the abundance of adult females
and of prepubertal females. The abundance of adult females
exhibited a different pattern of variation among the sampling
areas within the embayments (F5, 250 = 3.8, P = 0.003). In
Little Egg Harbor, adult female abundance did not vary
among the areas whereas in Barnegat Bay, adult female abun-
dance declined significantly only between areas 4 and 5
(P = 0.004) (Fig. 3). The abundance of prepubertal females
varied only by month (F3, 270 = 29.2, P < 0.0001), with more
prepubertal females in August than in any other month (all
comparisons, P < 0.0001).
Sex Ratio
Sex ratio (no. of males/no. of females) varied by year
(2008 > 2009, F1, 250 = 7.7, P = 0.006) and exhibited a dif-
ferent pattern among sampling areas within each embayment
 Estuaries and Coasts

Fig. 3 Abundance expressed as

CPUE (±1 SE) of males, adult
females, pre-pubertal females,
and ovigerous females in each
sampling area

(F5, 250 = 6.8, P < 0.0001). In Little Egg Harbor, variation in
sex ratio among areas did not occur; however, in Barnegat
Bay, sex ratio variation was driven by two concurrent patterns:
sex ratio increased between areas 6 and 7 (P = 0.008) and area
4 had significantly smaller sex ratios than did all the other
areas (all comparisons, P < 0.0001) (Fig. 4). As with female
abundance, salinity alone did not explain a significant amount
of variation in sex ratio suggesting the low sex ratio values at
area 4 may be associated with the presence of a greater num-
ber of females due to area 4’s proximity to Barnegat Inlet. In
fact, an analysis focusing on sex ratios only at salinities of
greater than 25 ppt still shows lower sex ratios at area 4 as
compared to area 5 (P = 0.003), suggesting that some factor
other than salinity helps explain the low sex ratios at area 4.
Size
In each year, the average size of males exhibited a different
monthly pattern (year × month interaction, F3, 250 = 5.9,
P = 0.001). In 2008, the average male size in August was
greater than in June (P < 0.0001), whereas in 2009, no signif-
icant monthly variation in average male size was observed.
Adult females showed no significant temporal or spatial var-
iation in average size. Similarities in both male and female

Fig. 4 Average (±1 SE) sex ratio

(no. of males/no. of females) in
each sampling area
Estuaries and Coasts
size distributions among sampling areas, based on the KS
tests, distinguished three Bzones^ within Barnegat Bay: Little
Egg Harbor (areas 1–3, all comparisons Padjusted > 0.005),
Barnegat Bay-high salinity (areas 4–5, P > 0.05), and
Barnegat Bay-low salinity (areas 6–7, P > 0.05). In both sexes,
the size distribution in Barnegat Bay-high salinity was signifi-
cantly different from distributions in Barnegat Bay-low salinity
and Little Egg Harbor (KS tests, all comparisons, P < 0.0001).
Barnegat Bay-high salinity had fewer crabs in small size cate-
gories (90–110 mm) but more crabs in large size categories
(especially 140 mm and >150 mm) than Barnegat-low salinity
(frequencies adjusted for sample size, χ2, df = 6, P < 0.0001)
and Little Egg Harbor (frequencies adjusted for sample size, χ2,
df = 6, P < 0.0001) (Fig. 5a, b). The size distribution of crabs in
Barnegat Bay-low salinity was similar to Little Egg Harbor (KS
test, Padjusted > 0.05).
Reproductive Potential
Female Brood Production
The spawning season began in May (there were already fe-
males with late-stage broods captured in early June) and ended
in August (no ovigerous females were captured in September).
Of the adult females captured, the sampling areas containing
the greatest percentage of ovigerous females were those clos-
est to the inlets into Barnegat Bay: area 1, near Little Egg Inlet
(55.8 % ovigerous females) and area 4, near Barnegat Inlet
Fig. 5 Size distribution in three
zones based on salinity of
Barnegat Bay using frequencies
adjusted for sample size of: a
males; b females
(69.5 % ovigerous females) (χ2 = 661.5, df = 6, P < 0.0001)
(Fig. 6). The abundance of ovigerous females exhibited a dif-
ferent monthly pattern between the embayments (month ×
embayment interaction, F3, 228 = 14.0, P < 0.0001); in Little
Egg Harbor, no monthly variation occurred whereas in
Barnegat Bay, there were more ovigerous females in July than
in August (P < 0.0001). Overall, the relative percentage of
ovigerous females decreased in August (12.4 %) as compared
to June (41.0 %) and July (44.6 %) (χ2 = 107.0, df = 2,
P < 0.0001). The embayments also exhibited a different pat-
tern in the average abundance of ovigerous females among the
sampling areas; no variation occurred among the areas in
Little Egg Harbor, whereas area 4 contained more ovigerous
females did than all the other areas in Barnegat Bay (all com-
parisons, P < 0.0001). Incidentally, area 4 also contained more
ovigerous females than did all the areas in Little Egg Harbor
(all comparisons, P < 0.0001) (see Fig. 3).
Seminal Receptacle Contents
Much of the variation in the seminal receptacle weight of the
females was explained by the characteristics of the females
themselves including size and reproductive condition (ovary
developmental stage and ovary weight). Seminal receptacles
were heavier in females containing small (undeveloped) ova-
ries (Y = −0.109X + 1.89, P < 0.0001) and in larger females
(Y = 0.011X − 0.026, P = 0.012). Consistent with the effect of
ovary weight, female ovary stage also influenced the weight

Fig. 6 Percent of ovigerous and
non-ovigerous females in each
sampling area. Numbers inside
each bar are sample sizes of all
adult females
of seminal receptacles (F4, 668 = 30.8, P < 0.0001). Females in
early stages of ovary development (stages 0 and 1), indicating
they had mated during the current summer, had heavier sem-
inal receptacles than did females with more advanced ovary
stages (stages 3 and 4, all comparisons P < 0.0001) (Fig. 7),
suggesting they had mated in the previous summer and were
preparing to spawn. Only a small amount of variation in the
seminal receptacle weight of females was explained by spatial
factors. The embayments exhibited a different pattern in the
average weight of seminal receptacles among the sampling
areas (F5, 668 = 3.8, P = 0.002); in Little Egg Harbor, no
significant variation in seminal receptacle weight occurred
among the sampling areas, whereas in Barnegat Bay, females
in sampling area 4 had lighter seminal receptacles than fe-
males in areas 6 and 7 did (both comparisons P < 0.05), al-
though these differences were not statistically significant
using adjusted P values. The relative abundance of females
at different stages of ovary development helps explain the
variation in seminal receptacle weights among sampling areas
Fig. 7 Weight (±1 SE) of seminal
receptacles and ovaries of females
according to female ovarian
developmental stage
Estuaries and Coasts
in Barnegat Bay. As compared to other sampling areas, area 4
contained a small proportion of females with early ovary
stages (15.6 %) but many females with late-ovary stages
(55.7 %) (χ2 = 69.5, df = 124, P < 0.0001) indicating they
were preparing to spawn. This is consistent with the greater
abundance of ovigerous females observed at sampling area 4
as compared to all other areas (see Fig. 3).
Vasa Deferentia Contents
The weight of the spermatophore component of male vasa
deferentia varied significantly by month (F3, 242 = 24.2,
P < 0.0001) and male size (F1, 242 = 15.5, P < 0.0001) but
not by embayment (F1, 242 = 0.80, P = 0.373) or sampling area
within embayment (F5, 242 = 1.4, P = 0.226). Average sper-
matophore stores increased with male size (Y = 2.19X − 4.22,
P < 0.0001). Average spermatophore stores in June were less
than in all the other months (all comparisons, P < 0.001)
(Fig. 8).
Estuaries and Coasts
Fig. 8 Weight (±1 SE) of the
spermatophore and seminal fluid
portions of male vasa deferentia
in each month
The weight of the seminal fluid component of male vasa
deferentia varied significantly by month (F3, 242 = 6.7,
P < 0.0001) and male size (F1, 242 = 21.0, P < 0.0001) but
not by embayment (F1, 242 = 1.3, P = 0.257) or sampling area
within embayment (F5, 242 = 2.0, P = 0.075). Average seminal
fluid stores increased with male size (Y = 7.16X − 13.55,
P < 0.0001). Average seminal fluid stores in June were less
than in August (P < 0.0001) (Fig. 8).
Discussion
This study tested the influence of physical characteristics on
adult blue crab population structure in Barnegat Bay by ex-
amining temporal and spatial variations in abundance, sex
ratio, and size as well as measures of reproductive potential
in both sexes in the two main embayments of this lagoonal
estuary: Little Egg Harbor and Barnegat Bay (see Fig. 1).
Little Egg Harbor lacks spatial variation in physical character-
istics (particularly salinity), whereas Barnegat Bay contains a
salinity gradient spatially similar to a drowned-river estuary.
Abundance and sex ratio exhibited distinct spatial patterns in
the two embayments that were driven by sex-specific differ-
ences in response to salinity: in Barnegat Bay, males increased
whereas adult (and particularly ovigerous) females were in
consistently low abundance in areas other than near the inlet.
The male response was a gradual change along the whole
salinity gradient; however, the female response was a precip-
itous drop only between areas 4 and 5, suggesting that there
are factors in addition to salinity, proximity to Barnegat Inlet,
influencing females. In some coastal estuaries with spatial
variation in salinity, though not necessarily a spatial gradient,
blue crab population characteristics vary based on salinity
(Hammerschmidt 1982; Rogers et al. 1990; Steele and Perry
1990). Barnegat Bay is unique in that a portion of this lagoon-
al estuary contains a salinity gradient steep enough to impact
aspects of adult blue crab population characteristics consistent
with flooded-river estuaries (Archambault et al. 1990; Hines
et al. 1987).
In some lagoonal estuaries, temporal variation (especially
temperature) produces more predictable patterns in blue
crab population characteristics than spatial variation does
(Buchanan and Stoner 1988; Murphy and Secor 2006) due
to the lack of spatial variation in physical variables. We also
observed some temporal variation in abundance (males, pre-
pubertal, and ovigerous females) and size (males), which gen-
erally identify peaks in growth and spawning during the sum-
mer months. Both female and male reproductive potential
(sperm and seminal fluid stores) were positively related to
crab size. Female reproductive potential was primarily influ-
enced by the female’s temporal proximity to spawning. Male
reproductive potential varied temporally, suggesting males
need time to prepare sperm and seminal fluid supplies for
the summer. Thus, temporal variation in blue crab population
characteristics are tied to biological processes that are
temperature-related (e.g., growth and reproduction).
Salinity had a marked effect on adult blue crab abundance
because of sex-specific responses to salinity variation that
differed between the embayments. In Little Egg Harbor,
where little salinity variation exists, there was no spatial var-
iation in the abundance of either sex or in sex ratio. In
Barnegat Bay, where a salinity gradient similar to that of a
drowned-river estuary exists, males were more abundant
whereas adult females (especially ovigerous ones) rarely oc-
curred in low salinity areas, similar to what has been observed
in drowned-river estuaries (Archambault et al. 1990; Hines
et al. 1987). This pattern was produced by different responses
to salinity by the sexes. In Barnegat Bay, male abundance was
negatively correlated with salinity such that the number of
males steadily declined between the low salinity (area 7) and
high salinity (area 4) locations. An analysis focusing within a
salinity range shared by both embayments (25–27 ppt) result-
ed in the following: (1) low salinity areas (6 and 7) were
effectively eliminated, and (2) abundance differences between

the remaining areas in Barnegat Bay (areas 4 and 5) were
eliminated. Both results implicate salinity as the driver of the
spatial variation in male abundance patterns in Barnegat Bay.
As seen in drowned-river estuaries, males often move to lower
salinity areas for molting (Archambault et al. 1990; Hines
et al. 1987) and we noted that a greater percentage of molting
males (preparing to or recovering from molt) was captured
near lower salinity areas. In contrast, adult female abundance
did not correlate with salinity but there were more adult fe-
males in area 4 as compared to area 5 (where salinities were
lower), suggesting that area 4 is an important location for adult
females due to its proximity to Barnegat Inlet rather than high
salinity per se (but see sex ratio below). As seen in drowned-
river estuaries, adult females move to the mouth of the estuary
for spawning (Darnell et al. 2010; Schaffner and Diaz 1988;
Tankersley et al. 1998) and we observed that a large propor-
tion of ovigerous females were captured near Barnegat Inlet as
well as Little Egg Inlet. Thus, the influence of physical factors
on various aspects of blue crab life history events and the
different responses to those factors by the sexes help regulate
adult population structure of blue crabs.
The divergent responses by the sexes to salinity in the two
embayments also influenced variation in adult population sex
ratio (no. of males/no. of females). In Little Egg Harbor, there
was no spatial variation in sex ratio whereas in Barnegat Bay,
sex ratio was high in areas of low salinity (area 7) and lowest
in areas of high salinity (area 4). Unlike male abundance, sex
ratio and female abundance were not significantly correlated
with salinity, suggesting the lack of female response to salinity
per se diluted the effect of salinity on sex ratio. Indeed, an
analysis focusing only on high salinities (>25 ppt) still result-
ed in higher sex ratios at area 4 as compared to area 5, sug-
gesting that the peak in female abundance in high salinity may
be due to proximity to Barnegat Inlet. In Barnegat Bay, sex
ratios were consistently male-biased. The spatial variation in
sex ratio was considerable ranging from 2.3 (at Barnegat Inlet)
to 17.3 (in the lowest salinity area). In a variety of other estu-
aries, blue crab populations exhibit male-biased sex ra-
tios to varying degrees particularly in adults (Abbe 1983; Fitz
and Wiegert 1992; Hines et al. 1987; Hsueh et al. 1993;
McClintock et al. 1993) but sex ratios are closer to unity
especially among juveniles (Fitz and Wiegert 1992). Thus,
sex ratio variation is not expected due to reproduction but
can been attributed to a variety of factors including different
movement or habitat use patterns between the sexes that can
vary by season (Abbe 1983; Harding et al. 2010), age class
(Fitz and Wiegert 1992), potential fishery effects (Abbe and
Stagg 1996), or salinity (Hines et al. 1987). Indeed, Hines et al.
(1987) within the Rhode River, a subestuary of Chesapeake
Bay, observed sex ratios, of juveniles and adults combined,
ranging from 1.2 (near the river mouth) to 11.75 (near low
salinity). Sex ratio influences various factors in the blue crab
mating system. Male-biased sex ratios, heighten the degree of
Estuaries and Coasts
sexual competition among males (Jivoff 1997a, b), influence
the mating behavior of both sexes (Jivoff and Hines 1998a, b),
and can lead to larger sperm stores of recently mated females
(Ogburn et al. 2014). Thus, variation in sex ratio may lead to
spatial variation in the mating success and potentially the re-
productive output of both sexes. We have no evidence for the
influence of sex ratio on sperm stores in either sex, but more
information on the sperm stores of recently mated females
from across the sex-ratio gradient in Barnegat Bay may allow
us to test this idea in the future.
It is unlikely that aspects of the sampling design of this
study, namely, the sampling gear or sampling locations, pro-
duced the observed spatial variation in sex ratio. It has been
argued for another Portunid that traps as a collecting gear
produce male-biased catches (Bellchambers and de Lestang
2005). However, other studies using traps as a collecting gear
for blue crabs do not consistently show male-biased catches
(Abbe 1983; Harding and Mann 2010; Harding et al. 2010)
and one study in Barnegat Bay proper sampling via trawls
caught more adult males than females (Kennish et al. 1982).
Therefore, it is not clear that the sex ratios in this study are
solely due to the collecting gear or that gear alone can explain
the distinct spatial pattern in sex ratio observed here. The
sampling locations in our study were distributed along the
western shore of the estuary. Adult females may preferentially
use areas that were not sampled in this study, including along
the eastern side of Barnegat Bay and in off-shore areas, as seen
near some southern estuaries (Gelpi et al. 2009; Ogburn and
Habegger 2015). In other estuaries, adult females moving to
the spawning grounds often use the near-shore shallows
(Aguilar et al. 2005). The eastern side of Barnegat Bay is
not only dominated by relatively shallow water (Kennish
2001) but also contains the inlets where the spawning grounds
are likely to occur and therefore represents an area adult fe-
males may prefer and/or congregate. Despite the lack of east-
west spatial coverage in our study, the results are consistent
with Kennish et al. (1982) who had more extensive east-west
coverage (but less north-south coverage): adult males were
consistently more abundant than adult females, except near
Barnegat Inlet. Spatial variation in bottom-type and benthic
habitats (e.g., seagrass) also exists which can influence the
population characteristics of blue crabs in this system (Jivoff
and Able 2001; Szedlmayer and Able 1996), suggesting that
sampling efforts should incorporate biotic features that vary
spatially rather than different locations per se. Current sam-
pling now incorporates spatial variation in physical character-
istics (e.g., salinity) and in benthic habitats in an effort to
provide a more comprehensive examination of blue crab pop-
ulation characteristics in the bay in the near future.
The size of crabs exhibited only a small amount of varia-
tion and physical variables had little effect on the size of crabs.
Males exhibited more variation in size than females did. In
general, the average size of males increased from June to July
Estuaries and Coasts
(or August) then decreased in September. This temporal pat-
tern in size indicates that summer represents a growing season
for male blue crabs in Barnegat Bay, as seen in other estuaries
(Archambault et al. 1990; Hines et al. 1987; Stehlik et al. 1998).
The increase in average size from early to mid-summer
(125.5 ± 4.1 SD to 129.2 ± 3.0 SD) suggests growth of males
within the adult size class, whereas the decrease in average
size at the end of summer (to 124.7 ± 3.3 SD) suggests the
recruitment of juvenile males to adulthood. Spatially, an anal-
ysis of size frequency distributions in both sexes produced
three Bzones^ within Barnegat Bay based on salinity: Little
Egg Harbor, Barnegat Bay-high salinity, and Barnegat Bay-
low salinity. The Barnegat Bay-high salinity zone had fewer
crabs of both sexes in small size categories but more crabs in
large size categories than the Barnegat-low salinity and Little
Egg Harbor zones. Because the size distribution in the high
salinity zone of Barnegat Bay differs from another high salin-
ity area (Little Egg Harbor) and the low salinity area of
Barnegat Bay in a similar way, this suggests that it is not
salinity per se that is driving the differences in the size distri-
butions. The relative lack of small females but enhanced num-
ber of large females near Barnegat Inlet is consistent with a
low proportion of prepubertal females but high percentage of
adult (especially ovigerous) females near Barnegat Inlet.
For blue crabs, defining the temporal and spatial distribu-
tion of the spawning stock (Gelpi et al. 2009; Ogburn and
Habegger 2015) and identifying members of the spawning
stock, especially in the absence of ovigery (i.e., the presence
of a brood), are critical challenges for implementing current
management strategies that seek to maintain some level of
reproductive potential in the spawning stock (Miller et al.
2011). Based on the presence of ovigerous females, the
spawning season in Barnegat Bay began in May and ended
in August. This is consistent with previous observations in
New Jersey (Stehlik et al. 1998) and is a slightly shorter
spawning season than further south in Chesapeake Bay
(May–September) (McConaugha et al. 1983; Van Engel
1958) since ovarian and resultant larval development are
slowed by reduced temperatures (Millikin and Williams
1980). Consistent with other estuaries (Darnell et al. 2010;
Tankersley et al. 1998), ovigerous females were concentrated
near the inlets to Barnegat Bay; however, in New Jersey, it is
not known if females use offshore areas as spawning
grounds as observed elsewhere (Gelpi et al. 2009; Ogburn
and Habegger 2015). Non-ovigerous adult females were pres-
ent throughout the Bay but it can be difficult to determine their
membership in the spawning stock using external indicators.
Egg remnants remain on the female pleopods after spawning
but only for a relatively brief time (Van Engel 1958) and thus
only help identify females that have very recently spawned.
The extent of fouling on the carapace can be a proxy for the
interval since the previous molt (Hines et al. 1987; Van Engel
1958) and thus time of copulation, it but may not provide an
accurate estimate of an adult female’s subsequent reproductive
status. The measures of female reproductive potential used in
this study (e.g., seminal receptacle contents, ovarian condi-
tion) provided very useful information for identifying the
spawning stock of blue crabs in Barnegat Bay, even in the
absence of ovigery.
Observations on female seminal receptacle contents and
ovary condition identified two cohorts of non-ovigerous adult
females: members of the current-year spawning stock (i.e.,
light seminal receptacles containing primarily spermatophores
and advanced ovaries) who mated the previous summer and
members of the following-year spawning stock (i.e., heavy
seminal receptacles containing seminal fluid and spermato-
phores and undeveloped ovaries) who mated during the cur-
rent summer. These two cohorts dominated the population of
adult females at different times during the summer: current-
year spawners were prevalent in early summer (June and July)
whereas following-year spawners dominated in late summer
(August and September). This temporal pattern occurred at
each sampling area (except the low salinity area) including
near the inlets, although to a lesser degree; in August, 48 %
of the adult females near the inlets were following-year
spawners. Thus, not all adult females near the spawning
grounds throughout the spawning season were members of
the current-year spawning stock. This has implications for
how surveys of the spawning stock in Barnegat Bay should
occur and be interpreted. At a minimum, biological sampling
of female reproductive status is necessary to identify true
members of the spawning stock, and offshore sampling should
occur in order to assess the relative importance of these areas
as spawning grounds. In addition, even if previous surveys of
the spawning stock elsewhere enumerated adult females in the
spawning grounds throughout the spawning season (Jones
et al. 1990; Lipcius and Van Engel 1990), our results suggest
that they may have overestimated the actual size of the annual
spawning stock. If so, this may help explain some of the var-
iation observed in previous modeling estimates of spawning
stock size (Prager 1996) and in spawning stock-recruit rela-
tionships (Lipcius and Stockhausen 2002; Lipcius and Van
Engel 1990). Incorporating these types of complexities in blue
crab life history or in aspects of the blue crab fishery have
significantly improved modeling efforts to understand the im-
pacts of the fishery on blue crab populations (Bunnell and
Miller 2005; Miller 2003; Miller et al. 2011).
Measures of male reproductive potential, spermatophore,
and seminal fluid stores were influenced by male size and the
temporal proximity to the peak in mating activity. Large males
had greater stores of both the spermatophore and seminal fluid
portions of the vasa deferentia as in previous studies in other
estuaries (Carver et al. 2005; Jivoff 1997b; Kendall et al.
2001). On average, both components of the vasa deferentia
were relatively low in June as compared to the rest of the
summer months, particularly August. This pattern in male

spermatophore and seminal fluid stores suggests that males
need time to build these supplies in order to prepare for in-
creased opportunities to mate which, based on the relative
abundance of sexually receptive (prepubertal) females and of
mating pairs, peaked in August.
Summary
Physical characteristics, particularly salinity, influenced spa-
tial variation in the population structure of adult blue crabs.
Adult blue crab abundance and sex ratio were distinct between
the embayments, primarily in relation to salinity, with more
males and fewer females in lower salinity areas. This sex-
specific pattern apparently stems from males responding to
salinity while adult (and ovigerous) females respond to addi-
tional factors, including proximity to the inlets (especially
Barnegat Inlet). As a result of this divergent pattern, sex ratio
also varies spatially, with high sex ratios in low salinity areas
and low sex ratios in high salinity areas. Temporal variation in
the size of males suggests that early-to-mid summer represents
a growing season while late summer-to-fall is the time juve-
nile males (and prepubertal females) recruit to the adult size
class. The spawning season lasted from May to August (based
on the presence of ovigerous females), and ovigerous females
were concentrated near the inlets, especially Barnegat Inlet.
Information on female sperm stores and ovarian development
identified two cohorts of adult females in Barnegat Bay: fe-
males that will spawn in the current year and females that will
not spawn until the following year. Thus, not all adult females
near the spawning grounds are members of the current
spawning stock. Thus, we suggest that future surveys of the
spawning stock in Barnegat Bay include biological sampling
to assess the reproductive status of females and sampling in
offshore areas to assess their relative importance as spawning
habitat.
Acknowledgments We thank the Rutgers University Marine Field
Station for providing a considerable amount of logistical support as well
as student housing, lab space, and boat access. This work was funded by
the Barnegat Bay National Estuary Program, now known as the Barnegat
Bay Partnership.
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