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Relative growth and determination of morphological sexual maturity of the

fiddler crab Uca thayeri Rathbun (Crustacea, Ocypodidae) in two mangrove
areas from Brazilian tropical coa...

Article in Instituto de Ecologia y Ciencias Ambientales · January 2012

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 Relative growth and determination of morphological sexual maturity
of the fiddler crab Uca thayeri Rathbun (Crustacea, Ocypodidae) in
two mangrove areas from Brazilian tropical coast
MARINA DE S. L. C. ARAÚJO¹*, PETRÔNIO A. COELHO¹,²†
& DANIELA DA S. CASTIGLIONI³.
¹Universidade Federal de Pernambuco (UFPE, Museu de Oceanografia, Departamento de Oceanografia, Laboratório
de Carcinologia. Av. Arquitetura, s/n, Cidade Universitária, CEP 50.740-550, Recife, PE, Brazil. *Corresponding
author: mslc.araujo@gmail.com
²Universidade Federal de Alagoas (UFAL, Curso de Engenharia de Pesca, Laboratório de Carcinologia. Av. Divaldo
Suruagy, s/n, CEP 57.200-000, Centro, Penedo, AL, Brazil.
³Universidade Federal de Santa Maria (UFSM, Centro Superior de Educação Norte do Rio Grande do Sul (CESNORS,
Departamento de Zootecnia e Ciências Biológicas, Laboratório de Zoologia e Ecologia. Av. Independência, 3751,
Vista Alegre, CEP 98.300-000, Palmeira das Missões, RS, Brazil.

Abstract. The aim of this paper was to study the relative growth and the morphological sexual maturity
of Uca thayeri from two mangrove areas of the State of Pernambuco, Northeast Brazil. The crabs were
manually collected monthly at the Ariquindá and Mamucabas Rivers, during spring low tide, from April
2008 to March 2009. Crabs were measured at the following dimensions: carapace width (CW); carapace
length (CL); cheliped propodus length (CPL); abdomen width (AW) and gonopod length (GL). The
growth was described by the equation y = a xb. The sexual maturity was estimated through a logistic
equation. A total of 2,711 individuals of U. thayeri were collected. Females presented larger CW than
males, an adaptation to increase the production of eggs. In the relationship CPL vs. CW, males showed
positive allometric and females, negative allometric growth. On the other hand, in the relationship AW vs.
CW, males showed negative allometric and females, positive allometric growth. This pattern is a reflex of
the growth of the cheliped in males and of the abdomen in females, due to their sexual functions. Males
reached maturity at larger sizes than females in both mangroves areas, probably due to their greater
investment in somatic growth, while females spend their energy in the reproductive process.

Keywords: allometric growth, biometric relationships, morphometry, sexual maturity

Resumo. Crescimento relativo e determinação da maturidade sexual morfológica do caranguejo

violinista Uca thayeri Rathbun (Crustacea, Ocypodidae) em duas áreas de manguezais da costa
tropical brasileira. O objetivo deste trabalho foi estudar o crescimento relativo e a maturidade sexual
morfológica de Uca thayeri em duas áreas de mangue do Estado de Pernambuco, Nordeste do Brasil. Os
caranguejos foram coletados manualmente nos Rios Ariquindá e Mamucabas, durante a maré baixa de
sizígia, mensalmente de abril/2008 a março/2009. Os caranguejos foram medidos nas seguintes
dimensões: largura da carapaça (LC); comprimento da carapaça (CC); comprimento do própodo do
quelípodo (CPQ); largura do abdome (LA) e comprimento do gonópodo (CG). O crescimento foi descrito
pela equação y = a xb. A maturidade sexual foi estimada através de uma equação logística. Um total de
2.711 indivíduos de U. thayeri foi coletado. As fêmeas apresentaram maior LC que os machos, uma
adaptação para aumentar a produção de ovos. Na relação CPQ vs. LC, os machos apresentaram
crescimento alométrico positivo e as fêmeas, crescimento alométrico negativo. Por outro lado, na relação
LA vs. LC, os machos apresentaram crescimento alométrico negativo e fêmeas, crescimento alométrico
positivo. Este padrão é um reflexo do crescimento do quelípodo nos machos e do abdômen nas fêmeas,
devido às suas funções sexuais. Os machos atingem a maturidade em tamanhos maiores do que as fêmeas
em ambas as áreas de manguezais, provavelmente devido ao seu maior investimento no crescimento
somático, enquanto as fêmeas gastam sua energia no processo reprodutivo.

Palavras chave: crescimento alométrico, maturidade sexual, morfometria, relações biométricas

Pan-American Journal of Aquatic Sciences (2012), 7(3):156-170

Sexual maturity of Uca thayeri 157

Introduction being one of them the independent (x), relative to

Among the macroinvertebrates found in the size of the animal, and the other dependent,
mangroves, the decapod crabs (mainly Brachyura) represented by a certain segment or tagma (y). The
are one of the most important taxa due to their high power function, or the equation of allometric
density and biomass (Litulo 2005 a,b). Among the growth, has been the most used in such studies
Infraorder Brachyura, the fiddler crabs of the genus (Hartnoll 1978, 1982, Araújo et al. 2012).
Uca Leach, 1814 (Ocypodidae) stand out for their The development of the crustaceans is
cosmopolitan character, being found in mangroves characterized by morphological changes related to
and estuarine beaches on temperate, subtropical and sex and puberty (Hartnoll 1982), which includes the
tropical zones (Crane 1975). differential growth of some tagmata. The chelipeds
This genus is relatively well studied, in males and the abdomen in females are some of the
especially on the Brazilian coast, with papers tagmata that are under this phenomenon, especially
dealing on many aspects of its biology, standing out in the transition of the last immature instar to the
those on their relative growth, for example: first mature, i.e., the puberty molt (Hartnoll 1974).
Masunari & Swiech-Ayoub (2003) for U. This transition can be detected by the inflection or
leptodactyla Rathbun, 1898, Negreiros-Fransozo et discontinuity in a series of linear or curvilinear
al. (2003) for U. thayeri Rathbun, 1900, Benetti & relationships of bivariate analysis (Haefner 1990)
Negreiros-Fransozo (2004) for U. burgersi Holthuis, and it is called morphological maturity. Thus,
1967, Castiglioni & Negreiros-Fransozo (2004) and studies on relative growth dealing with these two
Costa & Soares-Gomes (2008) for U. rapax (Smith, cited variables are the most used to detect the
1870), Masunari & Dissenha (2005) for U. mordax beginning of the morphological maturity (Negreiros-
(Smith, 1870), Masunari et al. (2005) and Hirose & Fransozo et al. 2003).
Negreiros-Fransozo (2007) for U. maracoani The aim of this study was to characterize the
(Latreille, 1802) and Pralon & Negreiros-Fransozo relative growth and to determine the morphological
(2008) for U. cumulanta Crane, 1943. sexual maturity of males and females of U. thayeri
Nearly 100 species of the genus Uca are from two mangrove areas located in Brazilian
known worldwide (Rosenberg 2001, Beinlich & von Northeast.
Hagen 2006, Shih et al. 2010) and for the Brazilian
coast, Melo (1996) registered ten species of this Materials and methods
genus, which nine of them have been recorded for Description of the sampling area
the littoral of the State of Pernambuco: Uca The State of Pernambuco is located in
leptodactyla, U. maracoani, U. mordax, U. rapax, Northeast Brazil (07º32’00” and 08º55’30” S;
U. burgersi, U. cumulanta, U. vocator (Herbst, 34º48’35” and 41º19’54” W). The coastal zone of
1804), U. victoriana von Hagen, 1987 and U. Pernambuco is 187 km long, being one of the most
thayeri (see Coelho 1994, Castiglioni et al. 2011a). important human population agglomerate of
The species U. thayeri occurs in muddy Northeast Brazil. The region has a climate within the
regions of the mangrove periphery, under shaded As' type of Köeppen system, i.e., hot and humid
areas, through the West coast of the Atlantic Ocean, with rainy autumn and winter. Both areas chosen for
in Florida, Gulf of Mexico, Antilles, Guatemala, the development of this work are located at the
Panama, Venezuela and Brazil (from the State of Southern littoral of Pernambuco: estuaries of
Maranhão to Santa Catarina) (Melo 1996). Ariquindá (8º 41’ 26” S, 35º 06’ 07” W) and
Negreiros-Fransozo et al. (2003) have accomplished Mamucabas (8º 46' 42” S, 35º 06’ 25” W) Rivers,
the only study on the relative growth and sexual Municipality of Tamandaré (Figure 1). The sampling
maturity of U. thayeri, in a population at Ubatuba, areas at each estuary were 9.69 km distant from each
State of São Paulo, Southeast Brazil. However, time, other. At Tamandaré, the mean air temperature
place and environmental factors may influence the varies from 25 to 30 ºC, the mean rainfall is around
growth rates of Brachyura (Baptista-Metri et al. 2,000 mm (Duarte 1993, Moura & Passavante
2005), and intraspecific variations in the size of 1995), the wind speed varies from 3.1 to 4.7 m.s-1
sexual maturity occur at different spatial scales (CPRH 2003), and the mean relative air humidity
(Díaz & Conde 1989, Hines 1982). Such facts during the sampling period was 74.93 %, according
highlight the importance of performing more studies to the database of the National System of
on growth and maturity, even if a given species has Environmental Data (SINDA).
been studied in other regions. The Ariquindá River is inserted at the
The studies on relative growth are based on Guadalupe Environmental Preservation Area.
the association between two biometric variables, Together with its affluent União, it becomes an

Pan-American Journal of Aquatic Sciences (2012), 7(3):156-170

158 M. ARAÚJO ET AL.

important component of Formoso River basin West of the Saltinho Biological Reserve. By
(CPRH 1999). It is considered one of the last non- entering in the Reserve, it suffers the first
polluted rivers of Pernambuco. The mangrove area environmental impact, being dammed to form the
chosen for the crabs' samplings is located near the reservoir that supplies Tamandaré. It meets the
confluence of Ariquindá and Formoso Rivers, at Ilhetas River and together they discharge into a spit
Carneiros Beach, and its vegetation is mainly (CPRH 2003). Besides the damming, it is considered
composed of Rhizophora mangle L., Laguncularia an impacted river due to the great deposition of solid
racemosa (L.) Gaertn. f. and, less frequently, waste and deforestation, as well as the housing
Avicennia schaueriana Stapf and Leechman ex occupation in the surrounding areas (Santos et al.
Moldenke. The substratum was visually 2001). The vegetation was mainly composed by L.
characterized as muddy sand. racemosa, R. mangle and few trees of A.
The Mamucabas River is located, almost schaueriana. The substratum was also visually
entirely, in the Municipality of Tamandaré. It rises at classified as muddy sand.

Figure 1. Map of the study area, at Ariquindá and Mamucabas Rivers, Pernambuco.

Field procedures Vitória, Vitória de Santo Antão Campus,

In each estuary, two sampling stations were Universidade Federal de Pernambuco, State of
chosen. The samplings of Uca thayeri were monthly Pernambuco (CAV-UFPE). Others were discarded
accomplished from April 2008 to March 2009, after study.
through manual collections during 30 minutes at Laboratory procedures
each estuary (15 minutes at each station), at low tide. The species U. thayeri was identified based
The crabs were captured above the substratum, as on Melo (1996). For the sexual differentiation,
well as extracted from the burrows with a gardening observations of the secondary sexual characters of
shovel. The sampling area had nearly 100 m². The the abdomen were made.
crabs were kept in plastic bags and preserved in ice, The following variables of each individual
and in the laboratory, they were preserved in alcohol were taken with a digital caliper (0.01 mm): CW –
70%. Some specimens were integrated into the carapace width, CL – carapace length, CPL – major
scientific collection of the Centro Acadêmico de cheliped propodus length of males and right cheliped

Pan-American Journal of Aquatic Sciences (2012), 7(3):156-170

Sexual maturity of Uca thayeri 159

of females, AW – abdomen width, GL – gonopod covariance (ANCOVA) (α = 0.05) (Zar 1996).

length, in males. The morphological sexual maturity was
Data analyses estimated to all allometric relationships, but the
The minimum, mean (± standard deviation) relationships CPL vs. CW for males and AW vs.
and maximum values of each variable were CW for females were chosen to represent the size at
estimated. The Student t test was applied to compare which 50% of males and females were already in the
the biometric values between sexes and estuaries (α adult phase (CW50%), taking into account the
= 0.05) (Zar 1996). importance of propodus and abdomen in the
For the determination of the age groups reproductive processes of males and females,
(juveniles and adults), the statistic methodology was respectively (Araújo et al. 2012). After the
based in the paper of Sampedro et al. (1999). For separation of groups, each age category was divided
that, the biometric data were log-transformed, and an in size classes (1.5 mm of CW) and the proportion of
analysis of K-means clustering, followed by a juveniles and adults in each class was calculated,
bivariate discriminant analysis, was performed. After with the proportion of adults being adjusted to the
that, the growth of Uca thayeri for each phase was logistic distribution [(y= a/(1+be-cx)], a commonly
described by the allometric equation y = a xb used normalized form of the logistic equation of
(Huxley 1950). The carapace width (CW) was Verhulst. After this, an interpolation was performed
considered the independent variable (x) and related to determine the CW50%.
to the other body dimensions (dependent variables,
y). The type of growth for each phase (juvenile Results
male, adult male, juvenile female and adult female) Ariquindá River
was established by the value of the constant 'b' of the A total of 1,411 individuals of Uca thayeri
potency function, which can be isometric (b = 1), were sampled at the estuary of Ariquindá River,
allometrically positive (b > 1) or allometrically being 627 males and 784 females.
negative (b < 1). The statistical significance of the The minimum, mean, standard deviation and
constant 'b' in relation to the unity was calculated maximum values of each evaluated variable are
with a t test (α = 0.05). To verify the possibility of shown in Table I. The t test showed that the sizes
grouping juveniles and adults under the same differed between sexes, with females showing larger
equation, the intercepts (a) and the regression CW (t = -4.18), CL (t = -4.10) and AW (t = -28.46),
coefficients (b) were tested by an analysis of and the males, larger CPL (t = 19.90) (p < 0.05).

Table I. Uca thayeri. Minimum, mean, standard deviation and maximum values of the measured morphological
variables for males and females at Ariquindá River, Pernambuco.

Sex CW CL CPL AW GL
Min 4.05 3.41 2.60 1.45 1.93
Mean 12.50 8.71 12.83 3.49 5.26
Males
Sd 3.06 2.22 6.12 1.44 1.16
Max 26.58 22.38 32.18 11.13 9.33
Min 4.65 1.31 1.90 1.36 (-)
Mean 13.16 9.20 5.00 7.31 (-)
Females
Sd 2.76 2.00 1.39 2.50 (-)
Max 23.01 19.47 15.64 14.14 (-)

Min, minimum; Sd, standard deviation; Max, maximum; CW, carapace width; CL, carapace length; CPL, cheliped
propodus length; AW, abdomen width; GL, gonopod length; (-), data not available.

The dispersion of points in the graphic of the males (b = 0.94, t = 3.14, p < 0.05) (Figure 2a). In
relation CL vs. CW was allometrically negative for females, the growth for this relationship was also
both juvenile (b = 0.91, t = 6.75, p < 0.05) and adult allometrically negative in both developmental

Pan-American Journal of Aquatic Sciences (2012), 7(3):156-170

160 M. ARAÚJO ET AL.

phases (juveniles: b = 0.94, t = 3.92, p < 0.05, adults (b = 1.19, t = -8.62, p < 0.05), with higher
adults: b = 0.97, t = 2.60, p < 0.05) (Figure 3a) growth rates in the juvenile phase (Figure 3c) (Table
(Table II). II).
In the relationship CPL vs. CW, males In the dispersion of points of the relationship
showed a positive allometric growth in both juvenile GL vs. CW for males, the growth was isometric in
and adult phases, with higher growth rate in adults the juvenile phase (b = 0.99, t = 1.53, p > 0.05) and
(b = 1.71, t = 17.43 and b = 1.82, t = 17.08, allometrically negative in the adult phase (b = 0.83, t
respectively) (p < 0.05) (Figure 2b), and the growth = 4.36, p < 0.05), i.e., the growth rates were higher
of females was isometric in both juvenile and adult in the juvenile phase (Figure 2d) (Table II).
phases (b = 0.97, t = 0.65 and b = 0.92, t = 1.93, The analysis of covariance showed
respectively) (p > 0.05) (Figure 3b) (Table II). significant differences in the intercepts and/or in the
In the relationship AW vs. CW, males regression coefficients between juveniles and adults
showed negative allometric growth in juveniles (b = of all analyzed relationships (p < 0.05) (Table II),
0.78, t = 6.73, p < 0.05), and adults (b = 0.82, t = which justifies the elaboration of graphics and
2.96, p < 0.05) (Figure 2c). However, the females equations separately for juveniles and adults from
showed an allometric positive growth, both for Ariquindá River.
juveniles (b = 1.39, t = -8.72, p < 0.05) and for

Table II. Uca thayeri. Linear equation, coefficient of determination (r²), results of the analysis of covariance (ANCOVA)
and t test, and allometry (Al) for males and females at Ariquindá River, Pernambuco.
Linear equation ANCOVA ANCOVA
Relation Phase n r² t test b Al CW50%
Log y = log a + b log x a b
JM 253 LogCL = -0.07+0.91LogCW 0.94 6.75* -
20.52* 0.00 12.6
CL vs. AM 290 LogCL = -0.10+0.94LogCW 0.91 3.14* -
CW JF 260 LogCL = -0.10+0.94LogCW 0.94 3.92* -
120.7* 120.81* 12.4
AF 464 LogCL = -0.12+0.97LogCW 0.93 2.60* -
109 LogCPL = -
JM 0.92 17.43* +
0.83+1.71LogCW
655.46* 630.04* 11.8
121 LogCPL = -
AM 0.88 17.08* +
CPL vs. 0.89+1.82LogCW
CW 119 LogCPL = -
JF 0.86 0.65 0
0.40+0.97LogCW
182.70* 168.28* 12.5
188 LogCPL = -
AF 0.76 1.93 0
0.34+0.92LogCW
120 LogAW = -
JM 0.83 6.73* -
0.38+0.78LogCW
12.12* 9.68* 19.0
150 LogAW = -
AM 0.68 2.96 -
AW vs. 0.31+0.82LogCW
CW 141 LogAW = -
JF 0.89 8.72* +
0.82+1.39LogCW
769.40* 815.81* 11.2
437 LogAW = -
AF 0.87 -8.62* +
0.45+1.19LogCW
GL vs. JM 101 LogGL = -0.37+0.99LogCW 0.90 1.53 0
138.84* 100.24* 12.6
CW AM 143 LogGL = -0.19+0.83LogCW 0.76 4.36* -

CW, carapace width; CL, carapace length; CPL, cheliped propodus length; AW, abdomen width, GL, gonopod length;
JM, juvenile male; AM, adult male; JF, juvenile female; AF, adult female; *, significant result (α = 0.05); +, positive
allometric growth; -, negative allometric growth; 0, isometric growth.

Mamucabas River maximum values of each evaluated variable are

A total of 1,300 individuals of Uca thayeri presented in Table III. The t test showed that the
were sampled at the estuary of Mamucabas River, sizes differed between sexes, with females
being 565 males and 735 females. presenting larger CW (t = -4,85), CL (t = -6,10) and
The minimum, mean, standard deviation and AW (t = -31,02), and the males, larger CPL (t =

Pan-American Journal of Aquatic Sciences (2012), 7(3):156-170

Sexual maturity of Uca thayeri 161

19,78) (p < 0.05). showed positive allometric growth in both juvenile

The dispersion of points in the graphic of the and adult phases, and changes in the degree of the
relationship CL vs. CW was allometrically negative cheliped allometry were more evident than at
for juvenile (b = 0.95, t = 6.04, p < 0.05), and adult Ariquindá River (juveniles: b = 1.43, t = 9.94,
males (b = 0.93, t = 8.00, p < 0.05) (Figure 4a), like adults: b = 1.88, t = 15.65, p < 0.05) (Figure 4b). On
at Ariquindá River. Both juvenile and adult females the other hand, females showed negative allometric
also showed negative allometric growth (b = 0.97, t growth in the juvenile phase (b = 0.89, t = 3.89, p <
= 6.09 and b = 0.99, t = 3.17, p < 0.05) (Figure 5a) 0.05) and isometric in the adult phase (b = 0.98, t =
(Table IV), like at Ariquindá River. 0.70, p < 0.05) (Figure 5b) (Table IV), a result a
In the relationship CPL vs. CW, males little different from the observed at Ariquindá River.

Figure 2. Uca thayeri. Dispersion of points and growth equations of the relationships between the carapace width (CW)
and the dependent variables [A - carapace length (CL), B – cheliped propodus length (CPL), C - abdomen width (AW),
D - gonopod length (GL)] for juveniles and adults of males at Ariquindá River, Pernambuco. Juveniles = gray points;
adults = black points.

In the relationship AW vs. CW, males 13.58, p < 0.05, adults: b = 1.27, t = 9.55, p < 0.05)
exhibited negative allometric growth, both in the (Figure 5c) (Table IV), like at Ariquindá River.
juvenile (b = 0.88, t = 2.54, p < 0.05) and adult In the relationship GL vs. CW, males
phase (b = 0.86, t = 4.43, p < 0.05) (Figure 4c). showed isometric growth in the juvenile phase (b =
However, juvenile and adult females showed 1.05, t = 1.43, p < 0.05) and negative allometric in
positive allometric growth, with a higher growth rate the adult phase (b = 0.80, t = 8.82, p < 0.05) (Figure
during the juvenile phase (juveniles: b = 1.51, t = - 4d) (Table IV), as also observed at Ariquindá River.

Pan-American Journal of Aquatic Sciences (2012), 7(3):156-170

162 M. ARAÚJO ET AL.

The analysis of covariance showed which justifies the elaboration of graphics and
significant differences in the intercepts and/or in the equations separately for juveniles and adults of U.
regression coefficients between juveniles and adults thayeri from Mamucabas River.
of all analyzed relationships (p < 0.05) (Table IV),

Figure 3. Uca thayeri. Dispersion of points and growth equations of the relationships between the carapace width (CW)
and the dependent variables [A - carapace length (CL), B – cheliped propodus length (CPL), C - abdomen width (AW)]
for juveniles and adults of females at Ariquindá River, Pernambuco. Juveniles = gray points; adults = black points.

Sexual Maturity these values were, respectively, 12.10 mm for males

(largest immature male CW = 13.75 mm, smallest
The size at morphological sexual maturity
mature male CW = 11.31 mm), and 11.90 mm for
was obtained for all biometric relationships for
females (largest immature female CW = 13.62 mm,
Ariquindá (Table II) and Mamucabas Rivers (Table
smallest mature female CW = 10.45 mm).
IV). However, the most adequate relationships to
determine the beginning of the reproductive activity Comparison of size between estuaries
are the CPL vs. CW in males and AW vs. CW in Comparing the size, it was possible to
females. At the Ariquindá River, the size at which observe that males showed larger CL and AW at
males of U. thayeri were considered Ariquindá River (p < 0.05), while the other
morphologically mature was 11.80 mm (largest morphological variables did not differ between
immature male CW = 12.98 mm, smallest mature estuaries (p > 0.05). The only variable to differ
male CW = 11.10 mm). Females reached the between estuaries for females was AW (p < 0.05),
morphological maturity at 11.20 mm (largest being larger at Ariquindá River. Due to these
immature female CW = 12.76 mm, smallest mature differences, the regressions were made for each
female CW = 9.56 mm). At the Mamucabas River, estuary separately.

Pan-American Journal of Aquatic Sciences (2012), 7(3):156-170

Sexual maturity of Uca thayeri 163

Discussion phases (Hartnoll 1978, 1982). Among the

The study of relative growth and the Brachyura, the body parts that exhibit these changes
determination of the size at sexual maturity are with more evidence are the cheliped, abdomen and
important attributes in crustaceans for a better pleopods of both sexes, during the transition from
comprehension of population and reproductive juvenile to adult phases (Hartnoll 1974, 1978). Thus,
biology (Castiglioni et al. 2011b). Crustacean patterns of growth can indicate at which size an
growth is discontinuous, and some body parts show organism reaches the size of sexual maturity
differential growth throughout the ontogenetic (González-Gurriáran & Freire 1994).

Table III. Uca thayeri. Minimum, mean, standard deviation and maximum values of the measured morphological
variables for males and females at Mamucabas River, Pernambuco.
Sex CW CL CPL AW GL
Min 4.30 3.24 2.42 1.40 1.58
Mean 12.26 8.35 12.16 3.19 5.15
Males
Sd 3.00 1.95 6.29 0.92 1.25
Max 22.38 14.62 35.20 10.15 8.50
Min 5.12 3.49 2.05 1.44 (-)
Mean 13.06 9.03 4.94 6.85 (-)
Females
Sd 2.88 2.00 1.11 2.69 (-)
Max 22.10 15.64 7.84 14.62 (-)

Min, minimum; Sd, standard deviation; Max, maximum; CW, carapace width; CL, carapace length; CPL, cheliped
propodus length; AW, abdomen width; GL, gonopod length; (-), data not available.

Table IV. Uca thayeri. Linear equation, coefficient of determination (r²), results of the analysis of covariance
(ANCOVA) and t test, and allometry (Al) for males and females at Mamucabas River, Pernambuco.
Linear equation ANCOVA ANCOVA
Relation Phase n r² t test b Al CW50%
Log y = log a + b log x a b
JM 235 LogCL = -0.12+0.95LogCW 0.94 6.04* -
10.73* 0.34 11.4
AM 322 LogCL = -0.08+0.93LogCW 0.94 8.00* -
CL vs CW
JF 267 LogCL = -0.13+0.97LogCW 0.96 6.09* -
31.43* 1.51 12.4
AF 462 LogCL = -0.15+0.99LogCW 0.94 3.17* -
LogCPL = -
JM 131 0.87 9.94* +
0.55+1.43LogCW
316.28* 391.67* 12.1
LogCPL = - -
AM 171 0.85 +
CPL vs 0.99+1.88LogCW 15.65*
CW LogCPL = -
JF 161 0.86 3.89* -
0.32+0.89LogCW
157.37* 94.88* 12.5
LogCPL = -
AF 265 0.82 0.70 0
0.41+0.98LogCW
LogAW = -
JM 139 0.74 2.54* -
0.48+0.88LogCW
298.09* 297.49* 11.4
LogAW = -
AM 215 0.80 4.43* -
AW vs 0.46+0.86LogCW
CW LogAW = - -
JF 211 0.88 +
0.95+1.51LogCW 13.58*
543.13* 42.24* 11.9
LogAW = -
AF 417 0.84 9.55* +
0.55+1.27LogCW
JM 121 LogGL = -0.43+1.05LogCW 0.85 1.43 0
GL vs CW 162.51* 189.49* 11.3
AM 217 LogGL = -0.16+0.80LogCW 0.86 8.82* -
CW, carapace width; CL, carapace length; CPL, cheliped propodus length; AW, abdomen width, GL, gonopod length;
JM, juvenile male; AM, adult male; JF, juvenile female; AF, adult female; *, significant result (α = 0.05); +, positive
allometric growth; -, negative allometric growth; 0, isometric growth.

Pan-American Journal of Aquatic Sciences (2012), 7(3):156-170

164 M. ARAÚJO ET AL.

Figure 4. Uca thayeri. Dispersion of points and growth equations of the relationships between the carapace width (CW)
and the dependent variables [A - carapace length (CL), B – cheliped propodus length (CPL), C - abdomen width (AW),
D - gonopod length (GL)] for juveniles and adults of males at Mamucabas River, Pernambuco. Juveniles = gray points;
adults = black points.

In the present study, in both estuaries, maturity at a smaller size, as it will be discussed
females were larger than males, an unusual result later, indicating that other factors besides growth,
among species of crab. Sexual dimorphism, with such as selective predation (Du Preez & Mclachlan
males larger than females, was observed in several 1984) may influence the size in which females attain
populations of Uca (Johnson 2003, Litulo 2005a,b, maturity. Besides, this dimorphism may be an
Benedetto & Masunari 2009). Bedê et al. (2008), adaptation to increase the production of eggs (Costa
studying eight species of the genus Uca, observed & Fransozo 2004, Castilho et al. 2008). Different
that males were larger than females in U. cumulanta, mortality rates, migration, higher tolerance of one
U. mordax and U. rapax, females were larger than sex to environmental adversities, differences in
males in U. thayeri and U. victoriana, and males and foraging efficiency, acquisition or assimilation of
females of U. leptodactyla, U. uruguayensis and U. food and behavioral patterns between sexes may be
vocator had similar sizes. Studies with U. thayeri cited as other factors that may influence these size
from São Paulo State coast also showed females differences (Crane 1975, Wolf et al. 1975, Johnson
larger than males (Costa & Negreiros-Fransozo 2003).
2003, Negreiros-Fransozo et al. 2003). However, In the present study, while females of U.
Bezerra & Matthews-Cascon (2007) did not observe thayeri showed larger AW, males showed larger
sexual dimorphism in size for Uca thayeri coming CPL. These secondary sexual characters are related
from another locality of northeastern Brazil. Despite with the roles played by each sex during
of being larger than males, females reached sexual reproduction (Tsuchida & Fujikura 2000, Castiglioni

Pan-American Journal of Aquatic Sciences (2012), 7(3):156-170

Sexual maturity of Uca thayeri 165

& Negreiros-Fransozo 2004). The crabs of the genus (Christy & Salmon 1984, Masunari & Swiech-
Uca are characterized by the presence of the Ayoub 2003, Costa & Soares-Gomes 2008). In
enormous chelae in one of the males' cheliped, being females, a large abdomen is important in the
the other of small size (Crane 1975). The giant perpetuation of the species, since the egg incubation
chelae increase the apparent size of the crab and play takes place inside it (Hartnoll 1982, Araújo et al.
an important role in agonistic encounters, defense, 2012).
food capture, territory demarcation and copula

Figure 5. Uca thayeri. Dispersion of points and growth equations of the relationships between the carapace width (CW)
and the dependent variables [A - carapace length (CL), B – cheliped propodus length (CPL), C - abdomen width (AW)]
for juveniles and adults of females at Mamucabas River, Pernambuco. Juveniles = gray points; adults = black points.

The CW has been generally considered the Castiglioni & Negreiros-Fransozo (2004) for U.
independent variable in morphometric studies of rapax, except for adult females, which showed
Brachyura because it exhibits few morphological isometric growth for this relationship in both studies.
changes throughout the crab's life (Castiglioni & However, this relationship is not the most
Negreiros-Fransozo 2004, Araújo et al. 2012, appropriate to express biological changes with
Castiglioni & Coelho 2011). In the relationship CL reproductive meaning that occur in the life of crabs
vs. CW, males and females of both estuaries showed (Santos et al. 1995). Other body parts, as the
negative allometric growth, indicating a higher abdomen, cheliped and gonopod shows changes in
growth of the CW in relation to the CL. This result the degree of allometry with puberty (Castiglioni &
is very similar to that obtained by Negreiros- Negreiros-Fransozo 2004, Castiglioni & Coelho
Fransozo et al. (2003) for the same species and by 2011) and are the most adequate for studies of

Pan-American Journal of Aquatic Sciences (2012), 7(3):156-170

166 M. ARAÚJO ET AL.

relative growth and determination of morphological Fransozo et al. (2003), the differences in the size and
maturity. shape of the chelae between sexes are noticeable
In both estuaries, in the relation CPL vs. even when the crabs are still small. This high
CW, males showed positive allometric and females, allometry of males is very important because the
negative allometric growth. On the other hand, in the courtship behavior on this genus is mainly visual,
relationship AW vs. CW, males showed negative through the waving with the giant chela during the
allometric and females positive allometric growth. reproductive period (Crane 1975). The genus Uca
This observed pattern is a reflex of the growth of the presents, among the brachyuran, males with the
cheliped in males and of the abdomen in females, highest values of b for the growth of the cheliped
due to their sexual functions, as discussed before. In (Masunari & Dissenha 2005). In other species of the
some studies with Uca, the growth of the abdomen superfamily Ocypodoidea, as in Ucides cordatus, the
was only analyzed for females and the growth of the values of b do not exceed 1.50 (Pinheiro & Hattori
cheliped, only for males (Masunari & Swiech- 2006, Castiglioni et al. 2011b), while in Uca, b
Ayoub 2003, Masunari & Dissenha 2005, Masunari usually exceeds this value (Benetti & Negreiros-
et al. 2005, Costa & Soares-Gomes 2008). However, Fransozo 2004, Castiglioni & Negreiros-Fransozo
according to Negreiros-Fransozo et al. (2003), the 2004, Masunari & Dissenha 2005), which stands out
abdomen and the cheliped are the dimensions that again the importance of the giant chela on the
better distinguish the growth of males and females. ecology of these animals.
This was observed in the present study, which In the U. thayeri females studied herein, the
reveals the importance of analyzing these structures relation AW vs. CW follows a pattern distinct from
for both sexes, as in the studies made by Castiglioni what is found in other brachyuran crabs, showing
& Negreiros-Fransozo (2004), Hirose & Negreiros- positive allometry in both juvenile and adult phases.
Fransozo (2007) and Pralon & Negreiros-Fransozo However, this fact corroborates previous studies on
(2008). U. thayeri (Negreiros-Fransozo et al. 2003), U.
The growth pattern determined by Hartnoll burgersi (Benetti & Negreiros-Fransozo 2004), U.
(1974) shows a considerable positive allometry in mordax (Masunari & Dissenha 2005) and U. rapax
the growth of the largest cheliped in males, both in (Castiglioni & Negreiros-Fransozo 2004). Thus, this
the juvenile and adult phases, as observed in several seems to be a pattern for the genus Uca. Castiglioni
species of Brachyura, as Ucides cordatus (Linnaeus, & Negreiros-Fransozo (2004) described that juvenile
1763) (Ucididae) (Castiglioni et al. 2011b) and in females of U. rapax present a higher rate of
species of the genus Uca, as in U. thayeri abdomen growth than adult ones, which was also
(Negreiros-Fransozo et al. 2003), U. burgersi found by Masunari & Swiech-Ayoub (2003) for U.
(Benetti & Negreiros-Fransozo 2004), U. mordax leptodactyla and by Negreiros-Fransozo et al. (2003)
(Masunari & Dissenha 2005) and in the present for U. thayeri, as well as in the present study.
study. This high level of the cheliped allometry is Probably the juvenile females invest in the abdomen
responsible for its larger size in males of Uca, growth and, by reaching the sexual maturity, they
resulting in crabs that can exhibit chelae larger than will be ready to incubate the eggs. As the abdomen
their carapaces (Crane 1975, Masunari et al. 2005). is not an independent appendage as chelipeds, and it
Even though at some phase of their development, must work associated to the sternum, any
females may show a positive allometric growth of disproportional growth after the puberty molt would
their chelipeds, as in U. maracoani (Hirose & difficult the movement of the pereopods (Masunari
Negreiros-Fransozo 2007), the values of b are & Dissenha 2005).
always smaller than the observed in males In the relation AW vs. CW, males exhibited
(Negreiros-Fransozo et al. 2003, Hirose & negative allometric growth in both phases and
Negreiros-Fransozo 2007). estuaries, as observed by Negreiros-Fransozo et al.
In many male Brachyura, the growth of the (2003) for the same species. The abdomen of male
chelipeds after the maturity becomes greater than in Brachyura stores two pairs of gonopods, responsible
the juvenile phase, as a secondary sexual character for the transference of spermatophores during copula
(Hartnoll 1982), as observed in U. thayeri (Hartnoll 1982). Due to this simple function,
(Negreiros-Fransozo et al. 2003), U. burgersi especially when compared to the functionality of the
(Benetti & Negreiros-Fransozo 2004), U. maracoani female's abdomen, the growth of male's abdomen is
(Hirose & Negreiros-Fransozo 2007), U. cumulanta allometrically negative or isometric in all
(Pralon & Negreiros-Fransozo 2008) and in the ontogenetic phases (Benetti & Negreiros-Fransozo
present study, a fact evidenced by the increase of b 2004, Castiglioni & Negreiros-Fransozo 2004,
in the adult phase. However, as cited by Negreiros- Pralon & Negreiros-Fransozo 2008).

Pan-American Journal of Aquatic Sciences (2012), 7(3):156-170

Sexual maturity of Uca thayeri 167

In the relation GL vs. CW, juvenile males probably due to their greater investment in somatic
presented higher growth rates than adult males, as growth, while females spend their energy in the
observed by Negreiros-Fransozo et al. (2003) for the reproductive process, saving energy for the
same species, Castiglioni & Negreiros-Fransozo production of eggs (Hartnoll 2006, Castiglioni &
(2004) for U. rapax and Pralon & Negreiros- Negreiros-Fransozo 2006, Araújo et al. 2012).
Fransozo (2008) for U. cumulanta. Juvenile males According to Crane (1975), small males avoid
may invest in the growth of the gonopods and, by combating with large and dominant males. Thus, it
reaching sexual maturity, they will be ready to is more advantageous to invest more energy in
fertilize females. The lower growth rates of growth than in reproduction.
gonopods in adults may be explained by the fact that Despite of impact signs observed in
they have to be accommodated inside the abdomen, Mamucabas River mangrove, males and females of
which in turn exhibited negative allometric growth, this area reached maturity at sizes larger than those
which restricts the length of the gonopods. observed at Ariquindá River. Among the
The regression coefficients and/or the environmental factors that regulate the size at sexual
intercepts of all relations differed between juveniles maturity, stands out the availability of food (Hines
and adults in both sexes of U. thayeri, indicating 1989). It is probable that the population of U. thayeri
contrasting growth patterns, also observed by of Mamucabas River is taking advantage from the
Negreiros-Fransozo et al. (2003). This points out the greater availability of organic matter from pollution,
need to analyze the relative growth separately for as observed for U. burgersi by Benetti & Negreiros-
juveniles and adults, which was observed in other Fransozo (2004).
studies with species from the genus Uca (Negreiros- Variations in the size at sexual maturity for a
Fransozo et al. 2003 for U. thayeri; Masunari & same species may occur in different spatial scales
Swiech-Ayoub 2003 for U. leptodactyla; Masunari (Díaz & Conde 1989, Hines 1982, Araújo et al.
& Dissenha 2005 for U. mordax; Masunari et al. 2012), even at near localities, as the Ariquindá and
2005 for U. maracoani). These differences also Mamucabas Rivers. Such fact also justifies the
demonstrate the changes in body parts throughout differences between the sizes at morphological
the ontogeny (Hartnoll 1974, 1978, Negreiros- sexual maturity in the present study and those
Fransozo et al. 2003). observed by Negreiros-Fransozo et al. (2003)
Through statistical analyses, it was possible (CW50%♂ = 13.80 mm; CW50%♀ = between 10.70 and
to observe that males presented larger CL and AW at 16.80 mm).
Ariquindá River, and the females, larger AW at The present study is the first contribution on
Ariquindá River. Regarding the dispersion of points, the relative growth and morphological sexual
the relationships presented similar growth patterns maturity of U. thayeri in tropical areas, where
between estuaries, except for the relation CPL vs. despite of the abundance of this species, no study
CW for females. Baptista-Metri et al. (2005) and had been carried out on such aspects. As it could be
Araújo et al. (2012) listed the locality, time and observed, this species presented growth patterns
abiotic variables as factors that may determine similar to those found for other species of Uca and
variations in the growth rate of organisms, which also with a population of U. thayeri from subtropical
may explain the differences observed between the areas, São Paulo State (Negreiros-Fransozo et al.
estuaries here studied. 2003). However, the size at morphological sexual
It is well known that in Brachyura the most maturity differed from the observed by these
appropriate dimensions to determine the onset of authors, showing latitudinal intraspecific differences
morphological sexual maturity are the cheliped and in these values. Meanwhile, more studies are needed
abdomen for males and females, respectively to better comprehend the ecology of this species in
(Masunari & Swiech-Ayoub 2003, Araújo et al. the tropical region.
2012). Thus, the CPL and AW were the variables
chosen in the present study to better illustrate the Acknowledgements
maturity in U. thayeri. In both studied estuaries, The authors are thankful to: Fundação de
males reached maturity at larger CW than females. Amparo à Ciência e Tecnologia do Estado de
This phenomenon was already observed in other Pernambuco (FACEPE), for financial support (APQ:
species from the genus Uca: U. burgersi by Benetti 0108-2.04/07); to CNPq and CAPES for the
& Negreiros-Fransozo (2004), U. rapax by fellowships; to the anonymous reviewers, for the
Castiglioni & Negreiros-Fransozo (2004, 2006) and special attention on our manuscript; to Adriano
U. cumulanta by Pralon & Negreiros-Fransozo Martins for his help during fieldwork; to Dr. Tereza
(2008). Males mature with larger sizes than females Cristina dos Santos Calado (UFAL), Dr. Jesser

Pan-American Journal of Aquatic Sciences (2012), 7(3):156-170

168 M. ARAÚJO ET AL.

Fidelis de Souza Filho (UFPE) and Dr. Sigrid Paulo, Brazil. Revista Brasileira de Zoologia,
Neumann Leitão (UFPE) for the important 21(1): 137-144.
suggestions on this doctorate qualification; and Castiglioni, D. S. & Negreiros-Fransozo, M. L.
especially to Dr. Petrônio Alves Coelho in 2006. Physiologic sexual maturity of the
memoriam, for being a great supervisor and friend. fiddler crab Uca rapax (Smith, 1870)
All sampling in this study was conducted in (Crustacea, Ocypodidae) from two mangroves
compliance with current applicable state and federal in Ubatuba, Brazil. Brazilian Archives of
laws (ICMBio 14340-1). Biology and Technology, 49(2): 239-248.
Castiglioni, D. S. & Coelho, P. A. 2011.
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Received June 2012

Accepted October 2012
Published online November 2012

Pan-American Journal of Aquatic Sciences (2012), 7(3):156-170

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