See

discussions, stats, and author profiles for this publication at:

https://www.researchgate.net/publication/233632483

Spawning biology of the

blue crab, Callinectes
sapidus, in North Carolina

Article in Bulletin of Marine Science -Miami- · September 2006

CITATIONS READS

34 113

3 authors, including:

Gary H Dickinson Daniel Rittschof

The College of New J… Duke University
37 PUBLICATIONS 851 248 PUBLICATIONS 8,616
CITATIONS CITATIONS

SEE PROFILE SEE PROFILE

Some of the authors of this publication are also working on

these related projects:

barnacle glue View project

blue crab biology View project

All content following this page was uploaded by Daniel Rittschof on 24 November 2014.

The user has requested enhancement of the downloaded file.

BULLETIN OF MARINE SCIENCE, 79(2): 273–285, 2006

Spawning Biology of the Blue Crab,

Callinectes sapidus, in North Carolina

Gary H. Dickinson, Daniel Rittschof, and Catherine Latanich

ABSTRACT
The blue crab, Callinectes sapidus Rathbun, supports valuable fisheries in many
Atlantic and Gulf Coast states. We studied the spawning biology of female crabs
hand-captured in the Carrot Island Embayment, central North Carolina. Crabs
were retained sub-tidally in submerged, partially buried minnow traps and fed daily.
Of 124 experimental animals, 66% had two or more clutches of eggs with three in-
dividuals producing seven clutches over 18 wks. The longer the crabs were held, the
more clutches they produced. We infer that an average size crab (127 mm carapace
width) would produce eight clutches over a 25-wk spawning period. Since larger
crabs had larger clutches but produced them less frequently than smaller crabs, re-
productive output over 18 wks of the spawning season was statistically similar for
most size groups. Lipofuscin index values were higher for crabs that had recently
molted to maturity than for crabs that had been spawning for 18 wks. Sponge dam-
age of 2307 crabs, determined by the degree of deviation from the rounded form of
an intact sponge, indicated that animals caught by crab-pots had significantly more
sponge damage than hand-captured crabs. Sponge damage was most extensive dur-
ing mid-summer and the extent of damage differed among the smallest and largest
size groups during this time. Our finding that reproductive output is similar for
most size groups brings into question management plans that suggest the release of
the largest mature females.

The blue crab, Callinectes sapidus Rathbun, is common on the western Atlantic,
Gulf and Caribbean coasts from Massachusetts to Brazil (Van Engel, 1958). Blue crab
is the most valuable fishery for several Atlantic coast states including Maryland and
North Carolina (NMFS, 2003). Recent decreases in total catch (Chesapeake Bay Pro-
gram, 1997; Chesapeake Bay Commission, 2003), and abundance of spawning crabs
and larvae (Lipcius and Stockhausen, 2002) within the Chesapeake Bay has lead to
concerns over sustainability of the fishery. Among common strategies for manage-
ment of the fishery are measures to protect the spawning stock of mature females,
which have been employed in the Chesapeake Bay (Chesapeake Bay Program, 1997)
and recently in North Carolina (NCDMF, 2004).
To effectively manage the spawning stock using predictive models, an understand-
ing of blue crab spawning biology is necessary. Mating occurs with the female ter-
minal molt and females may store sperm sufficient for over a dozen clutches of eggs
(Hines et al., 2003). Quantity and viability of stored sperm are independent of female
body size (Wolcott et al., 2005). Females generate mature ovaries in warm months
in about six weeks. In central North Carolina, spawning occurs from April to No-
vember and to the greatest extent during summer months of June, July, and August
(Dudley and Judy, 1971). Females with clutches of eggs remain in, or move to, high
salinity waters (Van Engel, 1958; Mangum and Towle, 1977; Tankersley et al., 1998;
Turner et al., 2003) using ebb-tide transport (Tankersley et al., 1998; Hench et al.,
2004; Carr et al., 2004). In the Beaufort Inlet region of central North Carolina, crabs
move offshore through the inlet (Dudley and Judy, 1971; Carr et al., 2004). Work by
Hench et al. (2004) and Forward et al. (2005) suggest that blue crabs continue to

Bulletin of Marine Science 273

© 2006 Rosenstiel School of Marine and Atmospheric Science
of the University of Miami
274 BULLETIN OF MARINE SCIENCE, VOL. 79, NO. 2, 2006

move seaward using ebb-tide transport following larval release, contrary to previous
models suggesting a shift towards the estuary using flood-tide transport after larval
release.
Due to the mobile nature of mature female blue crabs, gaps exist in our basic knowl-
edge of blue crab spawning biology. The relationship between crab size and clutch
size has been well established for brachyuran crabs (Hines, 1982; Prager, 1990), but
other aspects of blue crab spawning biology, including an accurate estimate of how
many clutches a spawning female will produce in a given season, are not well under-
stood. Although local fishermen believe that blue crabs have only one clutch of eggs,
there is evidence that blue crabs produce multiple clutches of eggs (Van Engel, 1958;
Tagatz, 1968; Millikin and Williams, 1984; Prager et al., 1990; Hines et al., 2003).
The potential influence of the fishery on reproductive output of captured and
released ovigerous crabs is also not well understood. Ovigerous crabs captured in
crab-pots are often observed damaging their own sponge by picking at it with their
chelipeds (Rittschof, pers. obs.). This response is likely due to stressful environmen-
tal conditions, lack of food, and confinement with other crabs. Depending on the
number of clutches produced and the extent of sponge damage, a considerable por-
tion of a crab’s lifetime reproductive output may be lost due to trap stress even if the
crab is released.
Our primary objective for this study was to quantify several spawning parameters
for mature female blue crabs using crabs that were hand-captured in the vicinity of
the Carrot Island Embayment and retained in the field in submerged minnow traps.
Specifically, we addressed: (1) the total number of clutches produced during 18 wks
of the spawning season; (2) the relationship between crab size and clutch volume;
(3) differences in clutch production interval (number of weeks per clutch) among
size classes; (4) differences in the reproductive output among size classes and; (5)
since age is an important determinant of fecundity, we evaluated whether spawn-
ing parameters (number of clutches, clutch volume, clutch production interval, and
reproductive output) vary with lipofuscin index, which is a proxy of age in blue crabs
(Ju et al., 1999; 2001; 2002). Secondly, we analyzed the extent to which trap stress due
to potting may alter reproductive output using survey data on sponge damage (the
degree of deviation from the smooth rounded form of an intact sponge).

Materials and Methods

Two separate but related experiments were conducted in the vicinity of the Beaufort Inlet,
central North Carolina. These consisted of (1) field spawning trials with animals retained in
submerged, half-buried minnow traps (June–October 2004), and (2) a survey of sponge dam-
age of crabs in three locations (June 2000–November 2002). Data from both the field spawn-
ing trials and the sponge damage survey were analyzed using SigmaStat version 2.03.
Crab Collection For Field Spawning Trials.­—Our experimental design for the field
spawning trials entailed collecting 25 crabs in each of four months of May, June, July, and Au-
gust 2004 for a total of 100 crabs. This experimental design was chosen based on the number
of crabs we predicted we would be able to hand collect over a 2–3 d interval each month, and
to allow us to discern a seasonal trend in clutch production among crabs collected in different
months. We deviated from this design slightly due to an inability to collect 25 crabs in May,
and in order to replace crabs that died. During June, July, August, and September, at least 25
ovigerous crabs were collected by hand around the Carrot Island Embayment in the Rachael
Carson Estuarine Research Reserve, Beaufort, NC (latitude, 34°42′50″ N, longitude, 76°40′31″
 DICKINSON ET AL.: BLUE CRAB SPAWNING BIOLOGY 275

W). The July sample of 32 crabs included 18 crabs migrating at the surface at night from the
Morehead City turning basin (lat. 34°42′55″ N, long. 76°41′39″ W), approximately 1 km from
the Rachael Carson Estuarine Research Reserve.
Upon collection, each crab was labeled using a plastic poker chip with a unique identifica-
tion number attached across the back of the crab by looping and twisting coated 18-gauge
copper wire around the large lateral spines. Carapace width (CW) was measured as the dis-
tance between the tips of the large lateral spines. Crabs were examined for external eggs (a
sponge), and sponge dimensions (width-left/right, length-anterior/posterior, depth-dorsal/
ventral) were taken. Sponge color was noted and embryos from late-stage (brown or black)
sponges were sampled and examined using a dissecting microscope to determine viability.
Eggs with viable embryos are spherical, the exterior is dark yellow, and a dark brown/black
developing embryo can be seen in the interior. Non-viable eggs appear as a shriveled dark yel-
low mass or are spherical but contain no dark embryo in the interior (hollow). Eggs were taken
from each of four quadrants of the sponge (upper left, upper right, lower left, lower right), and
stored in alcohol at room temperature. The first fifteen eggs encountered in each quadrant
sample were scored for viability.
Experimental Procedure For Field Spawning Trials.—Crabs were confined in half
buried minnow traps below the low tide line (Ziegler, 2002) on the western side of Pivers Is-
land, Beaufort, NC. Minnow traps, 42 cm long and 23 cm in diameter, were buried vertically
in the sediment to a depth of approximately 20 cm to allow crabs to bury. Each trap consisted
of identical cylindrical halves hinged on one side and tapered to an inverted funnel measur-
ing 2.5 cm at either end and flattened on the bottom half of the trap that was buried. Traps
were buried in sediment that was shallow enough to be accessible but deep enough to remain
submerged at low tide (10–30 cm). A single female was confined to each trap and traps were
wired shut with 18-gauge coated copper wire. Each trap was assigned a number (1–100) and
tagged for reference. Tags and cages were periodically cleaned with a wire brush to remove
fouling. Crabs were fed daily with fresh and frozen baitfish (pinfish, croaker, spot, mullet, and
menhaden). In addition to the scheduled feedings, crabs caught and ate fish that entered the
traps.
The experiment ran from June through mid-October of 2004. Water temperature over the
course of the experiment ranged from 29.6 °C in early August to 21.0 °C in mid-October. Each
crab was sampled weekly for the presence of an egg sponge. Previous laboratory observations
have shown that egg development (from extrusion to release) takes at least seven days (Hines
et al., 2003; Rittschof, pers. obs.). Therefore, a weekly sampling interval ensured that all ex-
truded sponges were observed, although this interval was not frequent enough to discern
differences in the rate of development. Minnow traps were sampled for a total of 18 wks of the
spawning season. Examinations in the field took place at low tide. Crabs were handled with
a dip net and padded tongs. A cylindrical (61 cm height × 66 cm diameter) 1.0 cm galvanized
hardware cloth screen was placed around each trap to prevent crabs from escaping as they
were removed from the traps. Sponge dimensions and embryo viability samples were taken
as described above. The extent of external fouling on crabs was noted. Crabs that died during
the course of the experiment were replaced, so as to maintain one crab in each trap for the
duration of the experiment.
Lipofuscin Analysis of Animals Used in Field Spwaning Trails.—To determine
if any of the spawning parameters studied (number of clutches, clutch volume, clutch pro-
duction interval and reproductive output) vary with crab age, at the conclusion of the field
spawning trials (mid-October, 2004), 23 crabs that had been used in field spawning trials were
sacrificed for lipofuscin analysis, which is a proxy for age in blue crabs (Ju et al. 1999; 2001;
2002). For comparison, ten female and ten male crabs that had recently molted to maturity
(determined by examination of the abdomen) were collected by crab-pot in the North River
(lat. 34º45′36″ N, long. 76º36′09″ W) and subjected to lipofuscin analysis. Crabs were anes-
thetized on ice and carapace width and type/extent of fouling was recorded. The carapace was
removed and the presence of mature ovaries and gill parasites was noted.
276 BULLETIN OF MARINE SCIENCE, VOL. 79, NO. 2, 2006

Lipofuscin analysis was based on Ju et al. (1999) with minor modifications. Using forceps
and a sharp razor blade, external eyestalks were prepared for lipofuscin extraction by re-
moving the internal eyestalk and all retinal tissue. External eyestalks were cut lengthwise.
The inner neural tissue was retained and placed in a 50 ml conical polypropylene tube. The
chitonous outer coating was discarded. Total lipofuscin was extracted using HPLC grade di-
chloromethane (2.66 ml) and methanol (1.33 ml). Samples were sonicated using a microprobe
sonicator at 20 watts for 2 min and then centrifuged at 800× g for 10 min. The supernatant
was transferred to a clean glass test tube, dried under nitrogen to remove solvents, and the
resulting pellet re-dissolved in 4 ml methanol. Lipofuscin was quantified using a quinine
sulfate standard (dissolved in 0.1 N H2SO4) and analyzed on a Shimadzu RF-5301PC spectro-
flourophotometer with a 3 ml cuvette. Total protein analysis was conducted on the remaining
1 ml of sample. Samples were dried in a speed vac to remove methanol, and the pellet was
re-dissolved in 470 µl nanopure water. Total protein was analyzed using a Pierce Mirco BCA
kit with BSA standard, and quantified on a Molecular Devices SpectraMAX 190 spectro-
photometer. Lipofuscin index (LI) was calculated as the extractable lipofuscin concentration
calibrated vs quinine sulfate (µg ml–1) divided by the total protein concentration (mg ml–1).
Sponge Damage Survey.—Between June 2000 and November 2002, 2307 ovigerous crabs
were caught, marked, and released in three locations in central North Carolina: the Carrot
Island Embayment (lat. 34°42′50″ N, long. 76°40′31″ W), the North River (lat. 34°45′36″ N,
long. 76°36′09″ W), and the Newport River (lat. 34°45′36″ N, long. 76°42′09″ W). The survey
consisted of 341 crabs in the Carrot Island Embayment, 976 crabs in the North River, and
990 crabs in the Newport River. Carrot Island Embayment crabs were caught by hand. North
River and Newport River crabs were caught in crab-pots in the crab fishery. Carrot Island
Embayment crabs were collected June through November of each year. North River crabs
were collected in early summer (June and July) of each year, and Newport River crabs were
collected in mid-summer (July and August) of each year. All crabs were marked using a plas-
tic poker chip as described for the field spawning trails. CW was measured and each sponge
was examined visually and assigned a score for sponge damage where: 100%–71% of sponge
remaining was given a score of five; 70%–50% remaining, a four; 49%–41% remaining, a three;
40%–25% remaining, a two; and < 25% remaining, a one. Blue crab sponges follow a charac-
teristic smooth rounded form. The degree of deviation from this form was used to determine
extent of sponge damage.

Results

Spawning.—Blue crabs were found to have multiple clutches of eggs within a sin-
gle spawning season. Individuals that began spawning in June had as many as seven
clutches by October. Animals held in captivity for a longer period of time produced
more clutches than crabs held for short periods (One-way ANOVA: P < 0.001; Fig.
1). Of 124 experimental animals, 81 (66%) had at least two clutches, 50 (40%) at least
three, 33 (27%) at least four, 9 (7%) at least five, and 7 (6%) individuals had six or more
clutches. Of the 43 individuals producing only one clutch while in captivity, over half
(56%) spent two weeks or less in captivity.
Using the relationship between total number of clutches produced and total time
in captivity (Fig. 1) and assuming a 25-wk spawning period, we infer that average
size crabs (127 mm CW) that are mature at the beginning of the spawning period
produce eight clutches of eggs during a spawning season (6.5–9.3 clutches, 95% C.I.).
The assumption of a 25-wk spawning period (May 15–November 1) is based on: (1)
observations of ovigerous crabs in the Carrot Island Embayment by mid-May; (2) the
existence of crabs with sponges in mid-October; and (3) observation of mature ova-
ries in the majority of crabs (91%) sacrificed for lipofuscin analysis in mid-October,
 DICKINSON ET AL.: BLUE CRAB SPAWNING BIOLOGY 277

Figure 1. Total time in captivity vs number of clutches produced by blue crabs retained in the
field in submerged minnow traps. Mean value for all individuals held a given number of weeks is
shown with standard error bars. Points with no error bars represent data for a single crab. Line of
best fit (solid) and 95% confidence interval (dashed) is shown. The x-axis is extended to 25 wks
to allow estimation of clutch production over a full spawning season. Regression is significant (P
< 0.001).
indicating their potential to continue clutch production into November. We were not
able to discern a seasonal trend in clutch production.
Egg viability analyses were conducted on 98 late-stage sponges from 87 individu-
als. Viability was between 93%–100% for all sponges sampled. In late August, three
sponges were produced without embryo development (sponge did not change color
or was lost over a seven day interval). This occurred on the second clutch in captivity
for one animal and on the fourth clutch for two animals.
Fouling and parasitism will affect a crab’s ability to move and forage freely. Of ani-
mals used in spawning trials and sacrificed at the conclusion of the experiment, 80%
were fouled (any hard or soft fouling) and 96% of animals (22 of 23) had gill parasites
(parasitic gooseneck barnacle, Octolasmis muelleri Coker).
Clutch Volume.—Crabs producing three or more clutches showed a gradual de-
crease in clutch volume over successive clutches (Fig. 2). Between the first and the
seventh clutch, volume decreased by 56%. Of individuals having only two clutches
(n = 43), 77% had a decrease in clutch volume (average decrease from first to second
clutch, 25%), while 23% had an increase in clutch volume (average increase from first
to second clutch, 18%). Since we were only able to sample each sponge once, these
calculations do not discriminate between stages of sponge development.
Larger crabs had larger clutches of eggs (Fig. 3). Regression of clutch volume on
carapace width was significant for clutches 1, 2, 3, and 4 (One-way ANOVA: P <
0.001). As clutch number increased, the change in clutch volume per change in CW
decreased (Fig. 3). The difference in volume of clutches between large and small crabs
decreased by 9% from clutch one to clutch four, with large crabs producing relatively
smaller clutches on the fourth clutch than they did earlier.
Crab size vs Clutch Production Interval.—Total time an individual crab spent in
captivity divided by the total number of clutches that individual produced while in
captivity was taken as the clutch production interval. Clutch production interval was
longer for larger crabs than for smaller crabs (Fig. 4). Crabs were grouped into seven
size classes, each with a 15 mm range in CW: 72–86 mm (n = 6), 87–101 mm (n = 8),
278 BULLETIN OF MARINE SCIENCE, VOL. 79, NO. 2, 2006

Figure 2. Mean clutch volume (± SE) over successive clutches for blue crabs producing more than
two clutches in captivity. All clutch stages and all crab sizes are included. Crabs were retained in
the field in submerged minnow traps.
102–116 mm (n = 21), 117–131 mm (n = 36), 132–146 mm (n = 26), 147–161 mm (n =
13) and 162–176 mm (n = 10). The largest size group corresponds to crabs exceeding
the maximum size limit (171 mm CW, with 5% tolerance) proposed by the North
Carolina Blue Crab Fisheries Management Plan (NCDMF, 2004). Clutch interval dif-
fered significantly among size groups (Kruskal-Wallis one-way ANOVA on ranks: P
< 0.001). Clutch interval was significantly longer in the largest group (162–176 mm
CW) than in the two smallest groups (72–86 mm and 87–101 mm CW) and signifi-
cantly longer in the 132–146 mm CW group than in the smallest group (72–86 mm
CW; Dunn’s Method post-hoc test on ranks: P < 0.05).
Reproductive Output.—Reproductive output (calculated as an individual’s clutch
frequency (inverse of clutch interval) multiplied by that individual’s average clutch
volume) over 18 wks of the spawning season differed significantly among groups
(Kruskal-Wallis one-way ANOVA on ranks: P < 0.001; Fig. 5). The 147–161 mm CW
group differed significantly from the three smallest groups (72–86 mm, 87–101 mm
and 102–116 CW mm), whereas all other group comparisons indicated a statistically
similar reproductive output (Dunn’s Method post-hoc test on ranks: P < 0.05).
Lipofuscin Analysis.—None of the spawning parameters considered (numbers of
clutches, clutch volume, clutch production interval, and reproductive output) varied
significantly with lipofuscin index. Lipofuscin index values did not differ significant-
ly between the left and the right eyestalk, and did not vary with CW. Lipofuscin in-
dex differed significantly among groups (Kruskal-Wallis one-way ANOVA on ranks:
P < 0.001; Fig. 6), being significantly higher for recently molted females and males
than for spawning females (Dunn’s method post-hoc test on ranks: P < 0.05).
Sponge Damage Survey.—Of 2307 ovigerous crabs examined, extent of sponge
damage was significantly greater for crabs caught by crab-pots than for crabs caught
by hand (t-test: P < 0.001; Fig. 7). The majority of crabs caught by crab-pot (52% in
the Newport River, 51% in the North River) had 50%–70% of the sponge remaining,
whereas 97% of hand-captured crabs had 100% of the sponge remaining. Crabs were
grouped by size class as described for the field spawning trials, however, a 177–191
mm CW group was added to accommodate larger animals and the 72–86 mm CW
group was eliminated since no animals in the sponge damage survey were < 90 mm
 DICKINSON ET AL.: BLUE CRAB SPAWNING BIOLOGY 279

Figure 3. Carapace width vs clutch volume for the first four clutches produced after capture by
individual blue crabs retained in the field in submerged minnow traps. Best-fit lines for each of
the four clutches are shown on one plot for comparison. Regressions for all four clutches are
significant (P < 0.001).
CW. Sponge damage differed significantly among size groups for crabs caught in the
Newport River (crab-pot, mid-summer) (Kruskal-Wallis one-way ANOVA on ranks:
P < 0.001). The smallest group, 87–101 mm CW, was excluded from post-hoc analysis
due to very low sample size (2 crabs). The 102–116 mm CW group differed signifi-
cantly from the 132–146 mm, 147–161 mm and the 162–176 mm CW groups (Dunn’s
Method post-hoc test on ranks: P < 0.05). Sponge damage was statistically similar for
all size groups within a site for animals caught in the North River (early summer) in
the Carrot Island Embayment (June–November).

Discussion

Blue crabs produce multiple clutches of eggs within a single spawning season.
From our data, we deduce that on the central North Carolina coast an average sized
crab (127 mm CW) that is mature at the beginning of the spawning season produces
eight clutches within a 25-wk spawning season. Larger crabs produce larger clutches
but produce them less frequently than do smaller crabs. Therefore, reproductive out-
put (incorporating clutch volume and clutch frequency) is statistically equivalent for
most crabs. Crabs caught in crab-pots show significantly more sponge damage than
280 BULLETIN OF MARINE SCIENCE, VOL. 79, NO. 2, 2006

Figure 4. Mean clutch production interval (± SE) in weeks per clutch for blue crabs with carapace
width between 72–86 mm (n = 6), 87–101 mm (n = 8), 102–116 mm (n = 21), 117–131 mm (n = 36),
132–146 mm (n = 26), 147–161 (n = 13), and 162–176 (n = 10). Clutch production interval is the
total number of weeks an individual spent in captivity divided by the total number of clutches pro-
duced by that individual while in captivity. Crabs were retained in the field in submerged minnow
traps. Groups marked with different letters are significantly different as shown by Dunn’s Method
post-hoc analysis on ranks. The largest group, 162–176 mm carapace width, is crabs exceeding
the seasonal maximum size limit proposed for spawning stock protection in the North Carolina
Blue Crab Fisheries Management Plan (2004).
do hand-captured animals. The extent of sponge damage differs among size classes
in animals caught by crab-pots during mid-summer, with most larger animals hav-
ing more damage than the smallest animals.
Our primary objective was to determine if blue crabs collected within the Carrot
Island Embayment, central North Carolina, have multiple clutches within a single
spawning season. Most local fishermen believe that blue crabs produce one clutch
of eggs and then die whereas trawl surveys in the Chesapeake Bay (Van Engel, 1958;
Millikin and Williams, 1984; Prager et al., 1990) and Florida (Tagatz, 1968) suggest
that crabs may have two and at maximum three clutches in some years. Recently,
Hines et al. (2003), using crabs taken from the Sebastian Inlet, Florida, and kept in
large tanks, demonstrated that crabs may have up to seven clutches within a single
spawning season, and inferred that Florida crabs may have up to 18 clutches within
a lifetime. Carrot Island Embayment crabs kept for 18 wks in the field under optimal
conditions (fed daily, free from predators) also had up to seven clutches. Based on
the tight relationship of increasing clutch production with time in captivity, we infer
that over a full 25-wk spawning period, average sized crabs (127 mm CW) have eight
clutches if they are mature at the beginning of the spawning season. From the 95%
confidence interval of the regression, this estimate has a range of 6.5–9.3 clutches,
corresponding to large and small crabs, respectively. Since most of our crabs were
infected with gill parasites, we are hesitant to infer a lifetime clutch production for
Beaufort crabs similar to that suggested by Hines et al. (2003) for Florida crabs. Gill
parasites prevent crabs from burying, which may lead to incapacitation by growth of
fouling organisms (Rittschof, pers. obs.). All but one crab dissected at the conclusion
of the experiment had gill parasites (parasitic gooseneck barnacle, O. muelleri). Thus,
crabs were unlikely to live through another spawning season.
 DICKINSON ET AL.: BLUE CRAB SPAWNING BIOLOGY 281

Figure 5. Mean reproductive output (± SE) over 18 wks of the spawning season for blue crabs
with carapace width (CW) between 72–86 mm (n = 6), 87–101 mm (n = 8), 102–116 mm (n = 21),
117–131 mm (n = 36), 132–146 mm (n = 26), 147–161 (n = 13), and 162–176 (n = 10). Reproductive
output is the average clutch volume (cm 3) multiplied by clutch frequency (total number of clutches
divided by total time in captivity) produced by individual crabs after capture. Groups marked
with different letters are significantly different as shown by Dunn’s Method post-hoc analysis on
ranks. The largest group, 162–176 mm CW, is crabs exceeding the seasonal maximum size limit
proposed for spawning stock protection in the North Carolina Blue Crab Fisheries Management
Plan (2004).
Multiple clutch production should be considered within the context of blue crab
life history. Following mating, female blue crabs migrate to, or remain in, high salin-
ity waters to facilitate offshore larval development of salinity sensitive larvae (Mil-
likin and Williams, 1984). Benefits proposed for larval development offshore include
reduced predation (Morgan, 1990) and increased survivorship in high salinity waters
due to physiological limitations (Costlow and Bookhout, 1959; Mangum and Towle,
1977). Movement to higher salinity water is achieved through ebb-tide transport
(Tankersley et al., 1998; Carr et al., 2004; Hench et al., 2004). Production of multiple

Figure 6. Mean lipofuscin index (± SE) for recently molted blue crab males and females, and
spawning females. Lipofuscin index is the concentration of extractable lipofuscin in µg ml–1 (cali-
brated against quinine sulfate) divided by total protein concentration in mg ml–1. Groups marked
with different letters are significantly different as shown by Dunn’s Method post-hoc analysis on
ranks.
282 BULLETIN OF MARINE SCIENCE, VOL. 79, NO. 2, 2006

Figure 7. Mean sponge damage (± SE) for blue crabs in the Carrot Island Embayment, North
River, and Newport River. Crabs from Carrot Island embayment were collected by hand. Crabs
from the North River and Newport River were caught in pots. Sample sizes are displayed for
each size group. Newport River groups marked with different letters are significantly different
as shown by Dunn’s Method post-hoc analysis on ranks. The 87–101 mm carapace width group
was not included in post-hoc analysis due to very small sample size. North River and Embayment
showed no significant difference between groups within a site.
clutches is consistent with recent work by Hench et al. (2004) and Forward et al.
(2005) suggesting that females continue to move seaward using ebb-tide transport
after the release of a clutch. Continued movement offshore ensures that successive
clutches are released in conditions favorable to larval survival.
Reproductive output over the 18 wks of our study was similar for most size groups,
a finding consistent with recent work by Wolcott et al. (2005) showing that the quan-
tity and viability of stored sperm is independent of female CW. Reproductive poten-
tial, therefore, should be independent of size. Larger crabs produce larger clutches
of eggs (Hines, 1982; Prager et al., 1990), and based on interspecific comparisons of
brachyuran crabs, Hines (1982) suggests that body size is the principal determinant
of reproductive output. In the Hines (1982) study, however, the number of broods
produced was not related to body size, a pattern that emerged from our data. Further,
Hines (1982) notes a trade-off between the number of eggs per brood and the num-
ber of broods per year. Although physiological constraints will limit the size of each
clutch, given the same complement of sperm, smaller crabs will generate the same re-
productive output as larger crabs by producing clutches at a quicker rate. Therefore,
two independent lines of evidence support the notion that all spawning females have
similar reproductive potential: (1) all sizes of blue crabs store comparable amounts
of viable sperm (Wolcott et al., 2005), and (2) most size groups were shown to have
similar reproductive output over 18 wks of the spawning season (this study).
Various authors have suggested that lifetime reproductive potential of blue crabs
is sperm-limited (Kendall and Wolcott, 1999; Kendall et al., 2001; Hines et al., 2003).
Direct evidence for sperm limitation was shown in Florida crabs (kept in tanks)
where 25% of fifth clutches and 50% of sixth clutches were infertile (Hines et al.,
2003). Although in our study all sponges at a mature stage of embryo development
were at or near 100% viable, three sponges were observed at the end of the season
that did not develop, suggesting sperm limitation in some Beaufort crabs. Because
 DICKINSON ET AL.: BLUE CRAB SPAWNING BIOLOGY 283

all females store comparable amounts of sperm (Wolcott et al., 2005), and this level
of sperm storage is sufficient to produce more than 6–9 clutches, we suggest that
sperm limited crabs are those who have already reached their lifetime reproductive
potential and are in their second year of spawning.
Given that blue crabs produce multiple clutches of eggs over the course of their
lifetime, and that age is important in estimating fecundity, it would be useful to
determine how spawning parameters change as spawning crabs age. However, the
relationship between lipofuscin index (a proxy for age in blue crabs; Ju et al., 1999;
2001; 2002), and the spawning parameters we examined (numbers of clutches, clutch
volume, clutch production interval and reproductive output) varied widely and no
clear trends emerged. Lipofuscin values for recently molted (pre-spawning) female
and male crabs were significantly higher than that of spawning females, indicating
that lipofuscin may not be useful in determining age in spawning female blue crabs
at the resolution of interest (within 6 mo). Using lipofuscin to age spiny lobsters,
Maxwell et al. (2006) also showed a large degree of variation in lipofuscin derived age
estimates for spawning females.
We also considered the potential influence of the fishery on reproductive output
of crab-pot captured and released ovigerous crabs. The extent of sponge damage was
more severe in crabs captured by crab-pots than crabs captured by hand. On average,
crabs captured by crab-pots were found with 50%–70% of their sponge remaining,
whereas hand-captured crabs were rarely found with damaged sponges. Although
the relationship between the extent of sponge damage and crab size was not dra-
matic, crabs collected during mid-summer showed a size-specific difference in the
extent of sponge damage, with the smallest crabs showing the lowest levels of sponge
damage. The degree to which sponge damage reduces reproductive output is directly
dependent on the number of clutches produced. As larger crabs produce fewer, but
larger clutches than do smaller crabs, the potential decrease in reproductive output
due to sponge damage is most significant in larger crabs. For small crabs estimated
to produce nine clutches within a year, the decrease in reproductive output would be
minimal if the crabs were released to continue clutch production.
Results of this study showing that reproductive output is similar for most size
groups and that the loss of reproductive output due to sponge damage during pot-
ting is more severe in larger crabs, brings into question spawning stock protection
strategies that call for the release of the largest mature females (> 171 mm CW). This
management strategy is based on the relation of sponge size increasing with CW but
does not take into account differences among sizes in the overall number of clutches
produced. A more sensible management strategy that would protect the spawning
stock but impose minimal loss in profit to fisherman may be to release medium sized
mature females but retain animals in the largest size group, as these animals are
the most profitable. Further studies assessing reproductive output of blue crabs in
other locations, assessing reproductive potential later into the year and over mul-
tiple spawning seasons, and further investigations into the effect of trap stress on
reproductive output should be undertaken to fully describe the reproductive biol-
ogy of blue crabs. Based on our results we suggest a reworking of blue crab popula-
tion models utilizing a fecundity parameter based on multiple clutches rather than
a single clutch. In models such as Miller (2001), small changes in fecundity result in
substantial changes in the intrinsic rate of population increase. Therefore, a realistic
estimate of fecundity is essential to an accurate population model.
284 BULLETIN OF MARINE SCIENCE, VOL. 79, NO. 2, 2006

Acknowledgements

We would like to thank J. Miles for assistance with data processing and lipofuscin proce-
dure, M. Vandersea for use of the spectroflourometer, and T. Ziegler, C. Tepper and L. Nichols
for helpful comments on drafts. We thank two anonymous reviewers who provided valu-
able comments and suggestions on the manuscript. Funding was provided by NC Sea Grant
(NCSU Subaward 2004-1772-15), and NC Sea Grant 03-Bio-04.

Literature Cited

Carr, S. D., R. A. Tankersley, J. L. Hench, R. B. Forward, and R. A. Luettich. 2004. Movement

patterns and trajectories of ovigerous blue crabs Callinectes sapidus during the spawning
migration. Est. Coast Shelf. Sci. 60: 567–579.
Chesapeake Bay Commission. 2003. Blue crab 2003, status of the Chesapeake population and
its fisheries. Chesapeake Bay Commission, Blue Crab Technical Work Group, Annapolis.
20 p.
Chesapeake Bay Program. 1997. 1997 Chesapeake Bay blue crab fishery management plan.
Chesapeake Bay Program Office, US Environmental Protection Agency, Annapolis. 102 p.
Costlow, J. D. and C. G. Bookhout. 1959. The larval development of Callinectes sapidus Rath-
bun reared in the laboratory. Biol. Bull. 116: 373–396.
Dudley, D. L. and M. H. Judy. 1971. Occurrence of larval, juvenile, and mature crabs in the
vicinity of Beaufort Inlet, North Carolina. U.S. Dept Commer, NOAA Tech Rep NMFS
SSRF-637. 10 p.
Forward, R. B., J. H. Cohen, M. Z. Darnell, and A. Saal. 2005. The circatidal rhythm in vertical
swimming of female blue crabs, Callinectes sapidus, during their spawning migration: a
reconsideration. J. Shellfish Res. 24: 587–590.
Hench, J. L., R. B. Forward, S. D. Carr, D. Rittschof, and R. A. Luettich. 2004. Testing a selective
tidal-stream transport model: observations of female blue crab (Callinectes sapidus) verti-
cal migration during the spawning season. Limnol. Oceanogr. 49: 1857–1870.
Hines, A. H. 1982. Allometric constraints and variables of reproductive effort in brachyuran
crabs. Mar. Biol. 69: 309–320.
__________, P. R. Jivoff, P. J. Bushmann, J. van Montfrans, S. A. Reed, D. L. Wolcott, and T. G.
Wolcott. 2003. Evidence for sperm limitation in the blue crab, Callinectes sapidus. Bull.
Mar. Sci. 72: 287–310.
Ju, S. J., D. H. Secor, and H. R. Harvey. 1999. Use of extractable lipofuscin for age determination
of blue crab Callinectes sapidus. Mar. Ecol. Prog. Ser. 185: 171–179.
_______, _________, and ___________. 2001. Growth rate variability and lipofuscin accumula-
tion rates in the blue crab Callinectes sapidus. Mar. Ecol. Prog. Ser. 224: 197–205.
_______, _________, and ___________. 2003. Demographic assessment of the blue crab (Cal-
linectes sapidus) in Chesapeake Bay using extractable lipofuscins as age markers. Fish. Bull.
101: 312–320.
Kendall, M. S. and T. G. Wolcott. 1999. The influence of male mating history on male-male
competition and female choice in mating associations in the blue crab, Callinectes sapidus
(Rathbun). J. Exp. Mar. Biol. Ecol. 239: 23–32.
____________, D. L. Wolcott, T. G. Wolcott, and A. H. Hines. 2001. Reproductive potential of
individual male blue crabs, Callinectes sapidus, in a fished population: depletion and recov-
ery of sperm number and seminal fluid. Can. J. Fish. Aquat. Sci. 58: 1168–1177.
Lipcius, R. N. and W. T. Stockhausen. 2002. Concurrent decline of the spawning stock, recruit-
ment, larval abundance, and size of the blue crab Callinectes sapidus in Chesapeake Bay.
Mar. Ecol. Prog. Ser. 226: 45–61.
Mangum, C. and D. Towle. 1977. Physiological adaptation to unstable environments. Am. Sci.
65: 67–75.
 DICKINSON ET AL.: BLUE CRAB SPAWNING BIOLOGY 285

Maxwell, K. E., T. R. Matthews, M. R. J. Sheehy, R. D. Bertelsen, and C. D. Derby. 2006. Neu-

rolipofuscin is a measure of age in the Caribbean spiny lobster, Panulirus argus, in Florida.
Abstract. 35th Annual Benthic Ecology Meeting, Quebec City.
Miller, T. J. 2001. Matrix-based modeling of blue crab population dynamics with applications
to the Chesapeake Bay. Estuaries 24: 535–544.
Millikin, M. R. and A. B. Williams. 1984. Synopsis of biological data on the blue crab, Calli-
nectes sapidus Rathbun. FAO Fish. Synop. 138: 1–39.
Morgan, S. G. 1990. Impact of planktivorous fishes on dispersal, hatching, and morphology of
estuarine crab larvae. Ecology 71: 1639–1652.
NCDMF (North Carolina Division of Marine Fisheries). 2004. North Carolina fishery manage-
ment plan blue crab. Department of Environmental and Natural Resources, Morehead City.
230 p.
NMFS (National Marine Fisheries Service). Annual commercial landings electronic database.
Silver Spring, MD: National Marine Fisheries Service, Fisheries Statistics Division; ©2005,
18 November 2005. Available from: http://www.st.nmfs.gov/st1/commercial/landings/an-
nual_landings.html
Prager, M. H., J. R. Mcconaugha, C. M. Jones, and P. J. Geer. 1990. Fecundity of blue-crab, Calli-
nectes sapidus, in Chesapeake Bay - biological, statistical and management considerations.
Bull. Mar. Sci. 46: 170–179.
Tagatz, M. E. 1968. Biology of the blue crab, Callinectes sapidus Rathbun, in the St. Johns River,
Florida. Fish. Bull. 67: 17–33.
Tankersley, R. A., M. G. Wieber, M. A. Sigala, and K. A. Kachurak.1998. Migratory behavior
of ovigerous blue crabs Callinectes sapidus: evidence for selective tidal-stream transport.
Biol. Bull. 195: 168–173.
Turner, H. V., D. L. Wolcott, T. G. Wolcott, and A. H. Hines. 2003. Post-mating behavior, intra-
molt growth, and onset of migration to Chesapeake Bay spawning grounds by adult female
blue crabs, Callinectes sapidus Rathbun. J. Exp. Mar. Biol. Ecol. 295: 107–130.
Van Engel, W. A. 1958. The blue crab fishery in Chesapeake Bay: reproduction, early develop-
ment, growth, and migration. Commer. Fish. Rev. 20: 6–17.
Wolcott, D. L., C. Wynne Bost Hopkins, and T. G. Wolcott. 2005. Early events in seminal fluid
and sperm storage in the female blue crab Callinectes sapidus Rathbun: effects of male mat-
ing history, male size, and season. J. Exp. Mar. Biol. Ecol. 319: 43–55.
Ziegler, T. A. 2002. Larval release behavior of the blue crab, Callinectes sapidus: entrainment cues
and timing in different tidal regimes. M.S. Thesis, Florida Institute of Technology. 80 p.

Date Submitted: 11 May, 2005. Date Accepted: 1 May, 2006.

Addresses: (G.H.D.) University Program in Ecology, Nicholas School for the Environment and
Earth Sciences, 135 Duke Marine Lab Rd., Beaufort, North Carolina 28516. (D.R.) University
Program in Ecology, Department of Biology, Nicholas School for the Environment and Earth
Sciences, 135 Duke Marine Lab Rd., Beaufort, North Carolina 28516. (C.L.) Nicholas School
for the Environment and Earth Sciences, 135 Duke Marine Lab Rd., Beaufort, North Carolina
28516. Corresponding Author: (D.R.) E-mail: <ritt@duke.edu>.

View publication stats