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Reproductive migration and population dynamicsof the blue crab Callinectes danae in an estuary insoutheastern BrazilBruno Sampaio Sant'Anna a , Alexander Turra b & Fernando José Zara ca Universidade Estadual Paulista (UNESP) - Campus Experimental do Litoral Paulista , SãoVicente (SP) , Brazilb Universidade de São Paulo (USP) - Instituto Oceanográfico, Departamento deOceanografia Biológica , São Paulo (SP) , Brazilc Universidade Estadual Paulista - (UNESP), FCAV, Depto de Biologia Aplicada,Laboratório de Morfologia de Invertebrados and Aquaculture Center (CAUNESP) ,Jaboticabal (SP) , BrazilPublished online: 16 Feb 2012.

To cite this article: Bruno Sampaio Sant'Anna , Alexander Turra & Fernando José Zara (2012) Reproductive migration andpopulation dynamics of the blue crab Callinectes danae in an estuary in southeastern Brazil, Marine Biology Research, 8:4,354-362, DOI: 10.1080/17451000.2011.637563

To link to this article: http://dx.doi.org/10.1080/17451000.2011.637563

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ORIGINAL ARTICLE

Reproductive migration and population dynamics of the blue crabCallinectes danae in an estuary in southeastern Brazil

BRUNO SAMPAIO SANT’ANNA1*, ALEXANDER TURRA2 & FERNANDO JOSE ZARA3

1Universidade Estadual Paulista (UNESP) - Campus Experimental do Litoral Paulista, Sao Vicente (SP), Brazil;2Universidade de Sao Paulo (USP) - Instituto Oceanografico, Departamento de Oceanografia Biologica, Sao Paulo (SP),

Brazil; and 3Universidade Estadual Paulista - (UNESP), FCAV, Depto de Biologia Aplicada, Laboratorio de Morfologia de

Invertebrados and Aquaculture Center (CAUNESP), Jaboticabal (SP), Brazil

AbstractPortunid crabs are an important resource in estuaries, and require appropriate management to guarantee their long-termavailability. We investigated the population dynamics and reproduction of Callinectes danae in the Estuarine�Bay Complex ofSao Vicente, Sao Paulo, Brazil, to provide basic biological information for public policies for the management of this fishery.Monthly samples were obtained from March 2007 to February 2008 on eight transects, four in the estuary and four in thebay. A total of 2261 specimens (403 males, 1288 females, of which 570 were ovigerous) were collected. Males weresignificantly larger than females, and the size�frequency distribution was unimodal for males, females and ovigerousfemales. The sex ratio was nearly always skewed toward females (M:F - 1:4.6). C. danae showed seasonal-continuousreproduction, with high reproductive activity in the warmer season. C. danae breeds in the estuarine�bay complex, withmales and juvenile females concentrated in the estuary. After copulation, fertilized females migrate to the estuary entranceand the bay, where ovigerous females are commonly found spawning in high-salinity areas. Therefore, to manage thisimportant economic resource, both the estuary and the bay should be considered for protection, but special attentionshould be given to the estuary entrance during the summer months, when ovigerous females concentrate.

Key words: Callinectes danae, migration, Portunidae, population dynamics, reproduction

Introduction

Portunid crabs are an important fishery resource

worldwide (Costa & Negreiros-Fransozo 1998; Lee

& Hsu 2003; Carr et al. 2004; Stevens et al. 2008).

On western Atlantic coasts, the main portunid

species exploited is the blue crab Callinectes sapidus

Rathbun, 1896; its importance as a fishery resource

has motivated many studies (e.g. Van Engel 1958;

Millikin & Willians 1984; Hines et al. 1995; Turner

et al. 2003; Carr et al. 2004). These crabs are

intensively exploited, even out of season when soft-

shell crabs are fished (Secor et al. 2002), and the

management and protection of the stocks generally

involves the use of marine protected areas (Lipcius

et al. 2003).

In Brazilian waters, portunid crabs are exploited

principally by coastal artisanal fishing communities

(Severino-Rodrigues et al. 2001; Sforza et al. 2010)

or, as non-target species, by the shrimp fishery

(Loebmann & Vieira 2006; Keunecke et al. 2008).

Economically important species of portunid crabs on

the Brazilian coast include Callinectes danae Smith

1869, Callinects ornatus Ordway, 1863, and

Callinectes sapidus (Severino-Rodrigues et al. 2001;

Baptista-Metri et al. 2005; Loebmann & Vieira

2006). The relative abundance of these species varies

along the coast (Branco & Massunari 2000; Chacur

& Negreiros-Fransozo 2001; Severino-Rodrigues

et al. 2001, 2009; Baptista-Metri et al. 2005; Sforza

et al. 2010), with C. danae dominating in some areas.

However, some stocks, such as in the Santos area,

have undergone reductions (Severino-Rodrigues

et al. 2001), thus requiring studies to inform specific

public bodies to protect these populations.

*Corresponding author: Bruno Sant’Anna, Universidade Estadual Paulista � Campus Experimental do Litoral Paulista, CEP 11330-900 �Sao Vicente (SP), Brazil. E-mail: brunusant@hotmail.com

Published in collaboration with the University of Bergen and the Institute of Marine Research, Norway, and the Marine Biological Laboratory,

University of Copenhagen, Denmark

Marine Biology Research, 2012; 8: 354�362

(Accepted 2 October 2011; Published online 14 February 2012; Printed 29 February 2012)

ISSN 1745-1000 print/ISSN 1745-1019 online # 2012 Taylor & Francis

http://dx.doi.org/10.1080/17451000.2011.637563

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The spatial and temporal distribution, growth,

reproduction, and feeding ecology of C. danae have

been studied in Brazilian waters (Branco & Masu-

nari 1992, 2000; Branco & Verani 1997; Costa

& Negreiros-Fransozo 1998; Chacur & Negreiros-

Fransozo 2001). However, specific information on

where ovigerous females are concentrated and where

no-take policies should focus is not generally avail-

able. The relationship between population dynamics

and reproductive migration of C. danae suggested by

Pita et al. (1985b) and Branco & Masunari (2000)

needs further investigation, for application to other

parts of the coast.

Callinectes danae is the main artisanal and sub-

sistence fishery resource exploited in the Estuarine�Bay Complex of Sao Vicente, Sao Paulo, Brazil

(Severino-Rodrigues et al. 2001). This important

economic region harbours the Port of Santos, the

largest port in Latin America. Nevertheless, only

studies on the local portunid diversity, distribution,

and first maturation (Pita et al. 1985a, b) and on

artisanal fishing (Severino-Rodrigues et al. 2001) are

available. Very little information is available on the

ecology of this species for its management. There-

fore, the present study investigated the population

dynamics of C. danae in the Estuarine�Bay Complex

of Sao Vicente, focusing on the spatio-temporal

distribution with special emphasis on sex and size

variation along the estuarine�marine gradient.

Material and methods

The crabs were caught monthly in the Sao Vicente

Estuarine-Bay Complex from March 2007 through

to February 2008. A shrimp fishing boat equipped

with a semi-balloon otter-trawl net 4.0 m wide,

2.0 m high and 9.5 m long with a 15-mm mesh body

and 10-mm mesh cod liner was deployed for 20 min

in each trawl. Sampling was done on eight transects,

four (numbered 1 to 4) in the estuary and four

(numbered 5 to 8) in the bay (Figure 1). On each

transect, the bottom water was sampled with a

Nansen bottle to measure the temperature (8C)

and salinity.

The crabs were sorted from the catch, identified

according to Melo (1996), and sexed (mature and

immature males and females, and ovigerous fe-

males). The relative abundance of sex categories

was compared between samples from the estuary

(pooling areas 1�4) and the bay (pooling areas 5�8)

using total and partial chi-square tests. The monthly

and overall sex ratios (M:F) were tested with

the chi-square test against a 1:1 proportion. The

carapace width (CW) was recorded between the tips

of the lateral spines using a caliper (0.05 mm), and

for the size�frequency distribution analysis, size-

class intervals of 8 mm carapace width were used,

according to Sturges (1926). The size�frequency

distribution was evaluated for normality by the

Kolmogorov�Smirnov (KS) test, and the size was

compared between the sexes using the Kruskal�Wallis test.

All individuals were dissected, and the ovary and

testis development was classified macroscopically

according to Costa & Negreiros-Fransozo (1998).

Ovaries of adult females were classified in four

stages: rudimentary (RU) with distended seminal

receptacle (sperm plug), developing (DE), inter-

mediate (INT), and mature (MAT). Three stages

were used for testis development of males: rudi-

mentary (RU), developing (DE), and mature

(MAT). The moult stage of the crabs was classified

according to the degree of carapace calcification, in

five stages (adapted from Mantelatto & Fransozo

1998): 1, recent moult: when the new carapace was

recently formed, indicating a post-moult stage,

Figure 1. Estuarine�Bay Complex of Sao Vicente, Sao Paulo,

Brazil, showing the sampling locations in the estuary (transects 1,

2, 3 and 4) and in the bay (transects 5, 6, 7 and 8).

Reproductive migration of the blue crab Callinectes danae 355

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with a very soft carapace; 2, intermediate post-

moult: carapace begins to calcify; 3, intermoult:

carapace hard and dull, characteristic of the inter-

moult stage with no recent moulting; 4, pre-moult:

carapace hard and dull, being changed by the new

carapace fully formed, indicating a moult; and 5,

moulting: crab caught at the moment of the

carapace change.

The relationships between the monthly tempera-

ture records and both the abundance of individuals

and the occurrence of ovigerous females were ana-

lysed by the Spearman’s rank correlation coefficient.

A significance level of 5% was adopted for all

statistical analyses (Sokal & Rohlf 1995). The

reproductive period was determined from the per-

centage of individuals of the same sex with developed

gonads and the incidence of ovigerous females.

The data presentation and graphical analysis of

individual abundance refer to the austral seasons:

autumn (March�May), winter (June�August), spring

(September�November) and summer (December�February).

Results

A total of 2261 individuals of Callinectes danae (403

males, 1288 females, and 570 ovigerous females)

were collected in the Sao Vicente Estuarine�Bay

Complex. Crabs of different sexes used the area

differently (x2 �958.42; DF �4; P B0.001). Ma-

ture and immature males were significantly more

abundant in the estuary, as were juvenile females.

Mature and ovigerous females were significantly

more abundant in the bay (Table I). The sex ratio

was nearly always skewed toward females (M:F, 1:14

Table I. Number of mature males (MM), immature males (IM), mature females (MF), immature females (IF), and ovigerous females

(OF) of Callinectes danae captured in the bay and the estuary of the Estuarine�Bay Complex of Sao Vicente from March 2007 to February

2008.

Location MM IM MF IF OF Total

Bay 100 16 886 13 389 1404

Estuary 206 81 264 125 181 857

Total 306 97 1150 138 570 2261

Partial chi-square 225.00* 171.40* 218.32* 325.38* 18.31*

*P B0.001.

Table II. Monthly and overall sex ratios for the population of Callinectes danae in the Estuarine�Bay Complex of Sao Vicente from March

2007 to February 2008.

Months Males Females Total Sexratio (M:F) x2 P

March 2007 76 797 873 1:10.5 595.47 0.001

April 2007 80 114 194 1:1.4 5.96 0.018

May 2007 35 130 165 1:3.7 54.70 B0.001

June 2007 32 24 56 1:0.8 1.14 0.350ns

July 2007 23 42 65 1:1.8 5.55 0.026

August 2007 19 80 99 1:4.2 37.59 B0.001

September 2007 27 54 81 1:2.0 9.00 0.004

October 2007 20 48 68 1:2.4 11.53 0.001

November 2007 16 77 93 1:4.8 40.01 B0.001

December 2007 10 120 130 1:12.0 93.08 B0.001

January 2008 25 131 156 1:5.2 72.03 B0.001

February 2008 40 241 281 1:6.0 143.78 B0.001

Total 403 1858 2261 1:4.6 936.32 B0.001

ns, non-significant.

1000

900

800

700

600

500

400

300

200

100

0

100

200

20–I

28

28–I

36

36–I

44

44–I

52

52–I

60

60–I

68

68–I

76

76–I

84

84–I

92

92–I

100

100–

I 108

108–

I 116

Num

ber

of In

divi

dual

s

Size Classes (Carapace width)

OF F M

Figure 2. Size�frequency (mm) distribution of males (M),

females (F), and ovigerous females (OF) of Callinectes danae

caught in the Estuarine�Bay Complex of Sao Vicente from March

2007 to February 2008.

356 B. S. Sant’Anna et al.

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to 1:12.0), except in June 2007 (M:F, 1:0.8), with

highly variable proportions but with a clear year-

round pattern of high proportions of females in the

summer (Table II).

The overall size-frequency distribution of the C.

danae population showed a unimodal pattern for

males, females, and ovigerous females (Figure 2),

skewed from normality for males and females

(KS �0.15; P B0.01 and KS �0.10; P B0.01,

respectively) and with normal distributions for

ovigerous females (KS �0.04; P B0.20). Crab size

varied significantly in relation to sex (H �371.50;

DF �2; P B0.001; Table III): males were larger

than both females and ovigerous females, showing

sexual dimorphism in size. Ovigerous females were

larger on average than non-ovigerous females.

The monthly variation of abundance in the study

area was positively correlated with the temperature

(RS �0.767; DF �10; P �0.0168), with the crabs,

especially females, being more abundant in the

warmer months of the year (summer and autumn)

(Figure 3). Ovigerous females were recorded in

all seasons, but in higher abundance in the

warmer seasons. This indicated a heterogeneously

Table III. Number (N) and size [carapace width, mm; minimum

(Min), maximum (Max), mean (X), and standard deviation (SD)]

of males (M), non-ovigerous females (F), and ovigerous females

(OF) of Callinectes danae caught in the Estuarine�Bay Complex of

Sao Vicente from March 2007 to February 2008. Means followed

by the same superscript letter did not show significant differences

(P �0.05) in the Kruskal�Wallis test.

Sex N Min Max X9SD

M 403 20.00 114.01 73.9913.45a

F 1288 21.01 79.50 63.6997.40b

OF 570 52.06 79.06 66.0195.33c

Total 2261 20.00 114.01 66.0999.20

Size Classes (Carapace width)

Num

ber

of In

divi

dual

s

600

500

400

300

200

100

0

100

Autumn/2007 (n=1232)

600

500

400

300

200

100

0

100

20–2

8

28–3

6

36–4

4

44–5

2

52–6

0

60–6

8

68–7

6

76–8

4

84–9

2

92–1

00

100–

108

108–

116

Spring/2007 (n=242)

20–2

8

28–3

6

36–4

4

44–5

2

52–6

0

60–6

8

68–7

6

76–8

4

84–9

2

92–1

00

100–

108

108–

116

Summer/2008 (n=567)

Winter/2007 (n=220)

OF F M

Figure 3. Seasonal size�frequency (mm) distribution of Callinectes danae caught in the Estuarine�Bay Complex of Sao Vicente from March

2007 to February 2008. The austral seasons are autumn (March�May), winter (June�August), spring (September�November) and

summer (December�February).

0

20

40

60

80

100

March\07

April\07

May\07

June\07

July\07

August\07

Septem

ber\07

October\07

Novem

ber\07

Decem

ber\07

January\08

February\08

Per

cen

tag

e (%

)

Months

Figure 4. Seasonal percentage of ovigerous females of Callinectes

danae caught during the study period in the Estuarine�Bay

Complex of Sao Vicente from March 2007 to February 2008.

Reproductive migration of the blue crab Callinectes danae 357

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continuous reproductive pattern (Figure 4),

although with no significant correlation with water

temperature (RS �0.3958; DF �10; P �0.2027).

Mature males predominated throughout the year,

while mature females did not outnumber the other

stages in winter (Figure 5).

Discussion

The reproductive cycle of Callinectes danae is com-

pleted in this estuarine�bay complex, with the

females spawning in higher-salinity areas. This is

supported by the data for copulation occurring

inside the estuary, where moulting females (pre-

moult, moulting, and recent moult; Figure 6) and

males were relatively more abundant (Table I) as

recorded for Callinectes sapidus in the northern Gulf

of Mexico by Hines et al. (1987). In addition, as

shown in Figure 7, juvenile females and adult

primiparous females with rudimentary and develop-

ing ovaries with a distended seminal receptacle

(sperm plug) are concentrated inside the estuary,

and primiparous females with mature ovaries with

flaccid seminal receptacles and ovigerous females are

abundant in the estuary entrance (in the warmer

season), where the salinity is higher (Figure 8).

Generally in a population, the expected sex ratio is

1:1 (males to females) according to Fisher’s theory

(Fisher 1930). In many cases, Fisher’s model is

sufficient to explain the population sex ratio, but

there are many exceptions to equal investment that

are explained by other models in the sex-allocation

theory (Bull & Charnov 1988). In the Sao Vicente

area, the C. danae population was skewed toward

females. The same has been recorded for various

populations of different portunid crab species

around the world (Hines et al. 1987; Baptista-Metri

et al. 2005; Bonine et al. 2008; Pereira et al. 2009),

and there may be many reasons for this; for example,

habitat partitioning by sex, size and moult stages for

C. sapidus (Hines et al. 1987); differential adult

mortality rates between sexes in Liocarcinus depurator

(Linnaeus, 1758) observed by Abello (1989); and

differences in the spatial and temporal distribution

with migration of ovigerous females in C. danae

(Baptista-Metri et al. 2005; Pereira et al. 2009).

The biased sex ratio found in this study does not

appear to be influenced by the reproductive migra-

tion behaviour of primiparous ovigerous females to

spawn, or by the concentration of males and juvenile

females in the estuary and adult and ovigerous

females in the bay, because the biased sex ratio was

also observed for the total of the individuals sampled

in both the estuary and bay. It is possible that the

crabs are concentrated in other locations of the

estuary, i.e. shallow waters where no samples were

taken. However, fishery pressure, which was not

evaluated in this study, may be influencing the sex

Per

cen

tag

e (%

)

0

20

40

60

80

100

0

20

40

60

80

100

March\07

April\07

May\07

June\07

July\07

August\07

Septem

ber\07

October\07

Novem

ber\07

Decem

ber\07

January\08

February\08

Months

MATINT

DERU

B

A

Figure 5. Seasonal percentages of males (A) and primiparous

females (B) of Callinectes danae, with rudimentary (RU), devel-

oping (DE), intermediate development (INT), and mature

(MAT) gonads, caught in the Estuarine�Bay Complex of Sao

Vicente from March 2007 to February 2008.

0

5

10

15

20

25

30

T 1 T 2 T 3 T 4 T 5 T 6 T 7 T 8

Nu

mb

er o

f fem

ales

Figure 6. Total number of females in moulting stages (pre-moult,

moulting, recent moult, and intermediate post-moult) caught in

the Estuarine�Bay Complex of Sao Vicente from March 2007 to

February 2008. Transects 1�4 were located in the estuary, and

5�8 in the bay.

358 B. S. Sant’Anna et al.

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ratio, because males are preferentially targeted

because of their larger size. This hypothesis is based

on the long-term fishing study by Severino-Rodri-

gues et al. (2001) in this estuary, which found that

C. danae is the principal portunid crab exploited

with 93,684 individuals caught in the 8 years

analysed, with males comprising 94% of the crabs

marketed.

Males were larger than females, a common pattern

in portunid crabs (Avila & Branco 1996; Batista-

Metri et al. 2005; Fernandes et al. 2006). This

difference is associated with the puberty moult,

when the metabolic energy is directed to somatic

growth in males, and in females to ovary develop-

ment (Hartnoll 1985). The larger size of males helps

them to manipulate the females during copulation

(Santos et al. 1995). The overall size�frequency

distribution of this C. danae population showed a

unimodal distribution for males, females, and oviger-

ous females. According to Diaz & Conde (1989),

these data indicate a stable population with contin-

uous recruitment and constant mortality rates, a

common pattern in tropical decapods. This is con-

firmed by the occurrence of ovigerous females

throughout the year, although neither sex was

normally distributed. As suggested by Litulo et al.

(2005), this may be a result of the seasonal migration

suggested by our data for the Sao Vicente population.

Chacur & Negreiros-Fransozo (2001) obtained a

similar pattern to the present study, with crabs being

more abundant in the warmer seasons. Probably the

larger catches in the warmer seasons are associated

with the reproductive migration described above,

and in the other periods of low reproductive activity

the crabs are in shallow waters, as also suggested by

Chacur & Negreiros-Fransozo (2001). Habitat par-

titioning between adults and juveniles of C. danae,

with almost all juveniles recorded in low salinities,

was also recorded in other populations (Chacur &

Negreiros-Fransozo 2001). The present study found

the same spatial distribution pattern, with 87.66% of

the juvenile crabs inside the estuary.

At Sao Vicente, more than 80% of males showed

mature gonads throughout the year, whereas the

proportion of mature females varied from 30 to

52%, with ovigerous females being present in all

0

5

10

15

20

25

20

22

24

26

28

30

32

34

36

38

40

T1 T2 T3 T4 T5 T6 T7 T8

Mea

n n

um

ber

of

OF

Sal

init

y

Figure 8. Monthly variation of salinity mean and error (standard

deviation), dashed line and mean number of ovigerous females

(OF, solid line) on eight transects (T1�T8) in the Estuarine�Bay

Complex of Sao Vicente from March 2007 to February 2008.

0102030405060708090

100

% o

f Ind

ivid

uals

AutumnOF

RU

MAT

IM

0102030405060708090

100 Winter

0102030405060708090

100

T-1 T-2 T-3 T-4 T-5 T-6 T-7 T-8 T-1 T-2 T-3 T-4 T-5 T-6 T-7 T-8

% o

f Ind

ivid

uals

Spring

0102030405060708090

100 Summer

Figure 7. Seasonal percentage of immature females (IM), rudimentary (RU) and mature individuals (DE), and ovigerous females (OF,

bars) of Callinectes danae collected along the estuarine (T-1 to T-4) to marine (T-5 to T-8) gradient from March 2007 to February 2008.

Reproductive migration of the blue crab Callinectes danae 359

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months, mainly in the warmer months. In general,

crustaceans may develop two types of reproductive

patterns, continuous or discontinuous (Sastry

1983). Continuous reproduction occurs when

ovigerous individuals and/or females with mature

gonads are present throughout the year with similar

monthly frequencies, while discontinuous or seaso-

nal reproduction is defined when ovigerous females

and/or females with mature gonads are restricted to a

certain period (Pinheiro & Fransozo 2002). An

intermediate pattern may be observed when the

species reproduces throughout the year, with periods

of higher reproductive activity (Turra & Leite 2000;

Pinheiro & Fransozo 2002). This is the case for

C. danae in the Sao Vicente system, where it

reproduces year-round with high reproductive activ-

ity in the warmer season, suggesting a heteroge-

neously continuous reproduction, as in other parts of

the Brazilian coast (Costa & Negreiros-Fransozo

1998; Chacur & Negreiros-Fransozo 2001; Baptis-

ta-Metri et al. 2005). However, this needs to be

confirmed, because in the present study only one

year of reproductive biology was analysed.

Differences between the sexes in moving among

estuarine habitats during the life history of the

blue crab Callinectes sapidus are well known, with

ovigerous females migrating to high-salinity areas to

hatch their eggs (Hines et al. 1987). This behaviour

was first recorded for C. danae by Pita et al. (1985b).

In a more exhaustive analysis in a study of the

ecology of C. danae in southern Brazil, Branco &

Masunari (2000) suggested that mating occurs in-

side the lagoon system, where males are more

abundant, and that ovigerous females spawn in

locations with high salinities in the open sea. The

present study confirmed their interpretation. The

analysis of combined data for the abundance of

ovigerous females and stages of ovarian maturation

of the non-ovigerous females (primiparous) and the

moulting of females clearly showed this migration

pattern. In addition, the pattern of ovarian develop-

ment demonstrated that the migration behaviour is

closely related to the first brood production by

primiparous females.

Migratory behaviour may be synchronized with

abiotic factors such as precipitation, relative hu-

midity, and the lunar cycle (Adamczewska &

Morris 1998; Debelius 1999). Such synchronized

migration was demonstrated for Gecarcoidea natalis

(Pocock, 1888), which arrives at the sea in syn-

chrony with the highest tidal amplitudes (Debelius

1999). The association of the reproductive migra-

tion with the summer months is thus suggested for

the Sao Vicente population of C. danae, despite the

lack of a significant correlation with water tempera-

ture. This strategy, to migrate to release larvae in

more-saline waters provides a more favourable

environment for embryo and larval development,

and favors dispersal of the offspring (Baptista-

Mestri et al. 2005) in more stable physical and

chemical oceanographic conditions (Hines et al.

1987; Sforza et al. 2010). Another important

finding of this study is that the estuarine migration

is associated with primiparous females. A few

observations on C. sapidus indicate that the first

clutch is more fecund than subsequent broods,

which are smaller and contain non-viable eggs

(Hines et al. 2003; Jivoff et al. 2007). However,

Darnell et al. (2009) evaluated the reproductive

potential of this species, analyzing the viability and

size (volume) of broods of C. sapidus, and observed

that this crab may produce up to 7 broods over 1 or

2 seasons, and that in the first broods the viability

of eggs with embryos developing normally was very

similar. If this is also true for C. danae, the females

(primiparous or not) at the entrance of estuary may

constitute an important part of the population to

maintain the regular stocks of larvae.

Thus, for the management of this important

fishery resource, both areas (estuary and bay) should

be considered for protection, but special attention

should be given to no-take measures in the entrance

of the estuary during the peak reproductive

period, especially the summer months, due to the

higher concentration of ovigerous females in these

areas.

Acknowledgements

The authors thank FAPESP (Fundacao de Amparo

a Pesquisa do Estado de Sao Paulo, FJZ JP Proc.

2005/04707-5 and Biota Proc. 2010/50188-8) and

CNPq (PQ FJZ Proc. 308215/2010-9 and AT

301240/2006-0) for financial support, and Dr Alvaro

Reigada and the student Evandro Dias for help

during the field sampling and laboratory analysis.

We also thank Dr Janet W. Reid (JWR Associates) for

editing the English text. This research was con-

ducted according to Brazilian laws (proc. IBAMA/

MMA 02001.000946/2007-76, license 34/2007-

CGREP-IBAMA).

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