Bioecology of the crab Ucides cordatus (Crustacea,

Decapoda) in mangroves influenced by the Amazon

River, Brazil
Kárlia Dalla Santa AMARAL1*, Inácia Maria VIEIRA2, Frederico Moreira OSÓRIO3,
Joana D’arc Mauricio ROCHA4, Jô de Farias LIMA1
1
Empresa Brasileira de Pesquisa Agropecuária – Embrapa Amapá, Rodovia Juscelino Kubitschek, km 5, n° 2600, CEP: 68903-419, Phone: +55 (96) 4009-9501. Macapá,
Amapá, Brazil. kdsa5@yahoo.com.br, jo.lima@embrapa.br
2
Instituto de Pesquisas Científicas e Tecnológica do Estado do Amapá – IEPA, Centro de Pesquisas Zoobotânicas e Geológicas. Rodovia Juscelino Kubitschek, km 10, Fazendinha,
CEP: 68903-419, Phone: +55 (96) 3212-5358. Macapá, Amapá, Brazil. inaciavieira2011@gmail.com
3
Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais Renováveis – IBAMA/RN. Avenida Alm. Alexandrino Alencar, 1399, CEP: 59.015-350, Phone: +55 (84) 3342-
0410. Natal, Rio Grande do Norte, Brazil. fredosorio@gmail.com
4
Universidade do Estado do Amapá – UEAP. Avenida Presidente Vargas, nº 650, CEP: 68.906-970. Macapá, Amapá, Brazil. j0anar0cha@hotmail.com
* Corresponding Author: kdsa5@yahoo.com.br

ABSTRACT
The mangrove crab (Ucides cordatus) is a valuable fishery resource, overfished along the Brazilian coast. This study aimed to
obtain bioecological data on this crab along the coast of the State of Amapá. Six bimonthly samplings were conducted between
December 2008 and January 2010. Transects were used to estimate the density (burrows m-2) and population abundance
(individuals m-2). All the animals were subjected to biometrics, with females being classified according to their stage of gonadal
maturation. The mean density (1.09 burrows m-2) and abundance (0.31 individuals m-2) were influenced by the climate with the
highest values ​​in summer (1.17 burrows m-2 and 0.34 individuals m-2). The male to female ratio was 1.38:1 showing significant
difference in the proportion of sexes. The individuals showed sexual dimorphism, with linear measurements significantly higher
in males. The sampled animals also had larger carapace length and width (CL and CW) compared to crabs studied in other
Brazilian states. There was a positive relationship between CW and CL and individual weight (IW) and CW for males (R2
= 0.83 and 0.90) and females (R2 = 0.79 and 0.84). The growth was negative allometric (CL increases to a lesser extent than
CW) for both sexes. The highest frequency of ovigerous females (78%) and in maturation stage IV (38%) occurred in the
CW size class between 59.8 and 67.5 mm. The peak of mature females occurred in May and August, showing a reproductive
period different from those in other Brazilian states.
KEYWORDS: State of Amapá, reproductive dynamics, population structure, fishery resource

Bioecologia do caranguejo Ucides cordatus (Crustacea, Decapoda) em

manguezais influenciados pelo Rio Amazonas, Brasil
RESUMO
O caranguejo-uçá (Ucides cordatus) é um valioso recurso pesqueiro, sobreexplorado ao longo do litoral brasileiro. O objetivo
deste trabalho foi obter dados bioecológicos deste caranguejo em toda a costa do Estado do Amapá. Foram realizadas seis
coletas de frequência bimensal, compreendidas entre os meses de dezembro de 2008 a janeiro de 2010. Por meio de transectos
foram estimadas a densidade (tocas m-2) e a abundância (indivíduos m-2) populacional. Todos os animais foram submetidos
à biometria, sendo somente as fêmeas, classificadas quanto ao estágio de maturação gonadal. A densidade (1,09 tocas m-2)
e a abundância (0,31 indivíduos m-2) foram influenciadas pelo clima, sendo os maiores valores observados durante o verão
(1,17 tocas m-2 e 0,34 indivíduos m-2). A razão sexual foi de 1,38M:1F, mostrando diferença significativa entre a proporção
de sexos. Os indivíduos apresentaram dimorfometria sexual, com medidas lineares significativamente superiores nos machos.
Os animais coletados também apresentaram maior comprimento e largura de cefalotórax (CC e LC), quando comparados aos
caranguejos estudados em outros estados brasileiros. Houve relação tanto entre LC e CC, como entre peso individual (PI) e
LC para machos (R2 = 0,83 e 0,90) e fêmeas (R2 = 0,79 e 0,84). O crescimento foi alométrico negativo (menor crescimento
de CC em relação à LC) para ambos os sexos. A maior frequência de fêmeas ovígeras (78%) e no estágio IV de maturação
(38%) pertenciam à classe de tamanho de LC 59,8 – 67,5 mm. O maior pico de fêmeas maduras ocorreu nos meses de maio
e agosto, mostrando um período reprodutivo diferente de outros estados do Brasil.
PALAVRAS CHAVES: Estado do Amapá, dinâmica reprodutiva, estrutura populacional, recurso pesqueiro.

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 Bioecology of the crab Ucides cordatus (Crustacea, Decapoda)
in mangroves influenced by the Amazon River, Brazil

INTRODUCTION three crustacean species of greatest economic importance

to the region. Therefore, this work aimed at studying some
The mangrove crab Ucides cordatus is distributed from
bioecological parameters of the species along the coast of
Florida (USA) to Santa Catarina, Southern Brazil (Coelho
Amapá in order to contribute to provision of standards for
and Ramos 1972; Melo 1996). Its growth rate is relatively
sustainable exploitation of the species, since this state includes
slow, making only one molting per year and reaching a life
traditional fishing communities, as well as many protected
span of over 10 years (Diele 2000; Pinheiro et al. 2005).
areas.
Reproduction is seasonal and the reproductive period is
inversely proportional to latitude, being longer in Northern
Brazil (Diele 2000; Pinheiro and Fiscarelli 2001). MATERIALS AND METHODS
Ucides cordatus is the second largest crustacean found in Study area
the Brazilian mangroves, and it is the most exploited species The Atlantic Coastal Zone of the State of Amapá is located
for human consumption (Castro et al. 2008). Some factors in the equatorial region and characterized by low temperature
such as the continued increase in fishing effort, habitat variation (mean = 27 °C), high relative humidity (above 80%),
degradation and the onset of diseases, contributed to this high rainfall (mean = 2,500 mm year-1), a short dry season
species being included in the “National List of Species of (August to November) and a long wet season (December to
Aquatic Invertebrates and Fishes Overexploited or Threatened July) (Vásquez and Rabelo 1999). The coastal ecosystems
with Overexploitation” (IBAMA, 2004). are strongly influenced by river discharges and macrotides,
Although mangrove crab is widely studied along the which can reach more than 10 m. Among these ecosystems,
Brazilian coast, there are few bioecology studies conducted mangroves of the region are considered the largest ones in
on the Amazonian coastal zone (Diele and Smith 2006; Diele the Brazilian coast and distributed in an almost continuous
and Smith 2007; Nordhaus et al. 2009), especially in the State range, from the mouth of the Oiapoque River to the Amazon
of Amapá (Fernandes and Carvalho 2007). Studies of this estuary (Costa-Neto et al. 2006).
nature are extremely important to assist government agencies The crab U. cordatus (Linnaeus, 1763) (Crustacea,
in developing protective measures such as: establishment of Decapoda, Ucididae) occurs in three municipalities in the
the closed season, which is the temporary interruption of the State of Amapá - Amapá, Calçoene and Oiapoque - and its
capture during the species reproduction period and setting capture for trading is performed in these three municipalities
minimum sizes and capture quotas. The regulation of crab (Figure 1). Two sampling sites were randomly selected in each
collection is responsibility of the Brazilian Environmental municipality, totaling six sampling areas. Mangroves in these
Agencies. However, due to the lack of studies, the legally areas are composed of two species: Rhizophora mangle L. (red
established closed season often does not coincide with the mangrove) and Avicennia germinans L. (black mangrove), the
real time of reproduction. latter being predominant.
The State of Amapá has a total area of ​​142,814,585 km²
and a coastline of 598 km. Along the coast, mangrove areas Bioecological characterization
represent approximately 1.94% (2784.97 km2) of State To investigate the seasonality influence on the population
territory. Mangroves are located in the lower parts of the dynamics of U. cordatus, six bimonthly diurnal samplings
coastal plain, extending continuously along the coast (IEPA were performed at each site in: December 2008; March, May,
2002). August and November 2009; and January 2010. Monthly
Hydrodynamics of mangrove waters bordering the State samples were not performed due to the difficult access to the
of Amapá, as well as other abiotic factors, suffer profound sampling areas that require a long period to perform each
influence of the Amazon River, which takes a northwest expedition. Samples were collected with permission of IBAMA
direction depending on the influence of the Guiana Current and ICMBio (Nº. 17545-1/2009).
(IEPA 2002). Since this river has the largest water flow on the Three transects were performed in each sampling area
planet, it causes marked changes to the mangrove ecosystem, with 3-m minimum spacing between them to determine
which are manifested in the externalization of structural and the number of existing burrows, the number of burrows with
functional patterns completely different from other coastal inhabitants and, the number of open and closed burrows.
regions in the country. Three squares (25 m2 each) were sampled in each transect.
In the resident fauna of these mangroves, there is the crab The collected data were used to estimate the abundance
U. cordatus (Linnaeus, 1763) (IEPA 2002), which along with and density existing in the studied areas. The number of
the shrimps Macrobrachium amazonicum (Heller, 1862) and individuals collected (individuals m-2) was used to determine
Macrobrachium carcinus (Linnaeus, 1758) are regarded the the abundance, while data on the number of inhabited

214 VOL. 44(2) 2014: 213 - 222  AMARAL et al.

 Bioecology of the crab Ucides cordatus (Crustacea, Decapoda)
in mangroves influenced by the Amazon River, Brazil

Figure 1 - Municipalities in the State of Amapá (Amapá, Calçoene and Oiapoque) with occurrence of the crab Ucides cordatus, showing the mangroves,
protected areas (Cabo Orange National Park and Maracá-Jipioca Ecological Station) and collection areas.

burrows, empty burrows and capped burrows (total burrows the gonadal maturation of females. The maturation stages
m-2) were used in the density calculation. were not identified in males, since the analyses were carried
The animal capture was performed manually by hands. out in the field and did not allow more accurate observation
After collection, the specimens were washed and packed of the gonads.
in polyethylene bags for transport to appropriate location
The sexual characterization of individuals was made
to perform the biometrics. Subsequently, the animals were
macroscopically because the animals showed evident sexual
stored ice coolers to induce them to a lethargic stage, thus
facilitating the acquisition of biometric data (sex, weight and dimorphism in the abdomen. The statistical significance of
carapace’s linear measurements), as well as determining the sex ratio on stocks composition was calculated by using the
gonadal maturation stage of females. The evaluation criteria Chi-square test (x²) for each month and the total period of
established by Mota Alves (1975) were used for determining data collection.

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 Bioecology of the crab Ucides cordatus (Crustacea, Decapoda)
in mangroves influenced by the Amazon River, Brazil

The individual weight (IW) was determined on analytical March (c² = 3.16), November (c² = 0.7) and December (c² =
balance (0.1 g accuracy). Linear measurements (mm) such as 1.32). Regarding the total individuals collected (c² = 21.8),
carapace length and width (CL and CW) were determined there was a significant difference for sex ratios in January (c²
with steel caliper (0.1 mm accuracy), according to the = 15.86), May (c² = 5.54) and August (c² = 13.78).
methodology of Pinheiro and Fiscarelli (2001). During Mean values obtained from the linear measurements and
weighing, the individuals without any of the claws were weights showed that the collected animals had larger carapace
discarded. Crabs without some of the pereiopods had the width (64.35 mm) and length (49 mm) and heavier weight
weight of the pereiopod symmetrical to that one lost added (125.3 mm). According to the statistical analysis (t-test), all
to their total weight. The t test was used to compare biomass variables were significantly different for males and females, as
data, CL and CW between males and females at 0.05% shown by the P values ​​(a = 0.05) (Table 1). Thus, males had
probability. The ratios between linear measures of carapace heavier weight, larger carapace length (CL), width (CW) and
width and length (CW x CL) for males and females were set higher height values (CH) than females, which clearly shows
to the linear equations y = a + bx. Ratios between carapace the dimorphometrics between sexes (Table 1).
width and individual weight (CW x IW) were set to the
exponential equations y = Axb. Regression analysis by power Ratios between linear measurements of males and females
function was used to check for the growth types through the showed a relationship between carapace length and width,
b value (Hartnoll 1978). Regarding CW x CL, the growth which was stronger for male individuals (R² = 0.8319) (Figure
types may be: Isometric (b = 1), positive allometric (b > 1) or 3). Regarding the growth type (CW x CL), the b values of
negative allometric (b < 1). Concerning to CW x IW, growth males (b=0.99) and females (b=0.83) indicated negative
can be isometric (b = 3), positive allometric (b > 3) or negative allometric growth tending to the isometry.
allometric (b < 3). Ratios between carapace width and individual weight (CW
x IW) was stronger for males (R² = 0.9055) (Figure 3). The
RESULTS CW x IW equations for males (b=2.7) and females (b=2.4)
showed b constant value less than 3 and indicated negative
The density (total burrows m-2) was higher than abundance
allometric growth.
(individuals m-2) in January, March, August and November
(Figure 2).The mean density (1.09 burrows m-2) and mean Most females collected (37.5%) showed CW between 59.8
abundance (0.31 individuals m-2) were low in the study period. and 67.5 mm. As for males, the highest frequency (28.9%) for
The highest values of individuals (0.49 individuals m-2) and CW class was 67.5 - 75.1 mm (Figure 4). The low number of
burrows (1.45 burrows m-2) were found in August, November crabs in smaller size classes (juveniles and small adults found
and December, when there is low rainfall (Figure 2). The mean in the mangroves) is related to the difficulty of finding their
density and mean abundance in the dry season (1.17 burrows burrows and due to the method for animal capture used by
m-2 and 0.34 individuals m-2) were higher than those found in the collectors.
the wet season (1.01 burrows m-2 and 0.28 individuals m-2). Regarding the reproduction, the highest frequency of
A total of 859 crabs of the species Ucides cordatus, 498 ovigerous females (78%) and those on stage IV of maturation
males and 361 females, were collected for bioecological studies (38%) were found in CW class 59.8 to 67.5 mm. Most females
in six areas along the coast of Amapá from December 2008 were sexually mature in the late May and early August due to
to January 2010 (bimonthly samplings). According to the the the higher percentage of females with dark brown and wine
Chi-square test for c² (0.05) = 3.84, there was no significant colored ovaries (Figure 5). In addition, 100% of the ovigerous
difference between the frequency of males and females in females were found in August, suggesting that the reproductive
peak occurs in the last quarter of wet season and in the first
quarter of dry season.

DISCUSSION
The populations of Ucides cordatus showed lower density
values than those determined for some States from the North
and Northeast regions of Brazil. This fact may be related to
the environmental characteristics of the study areas, such as
vegetation structure and composition (Pedersen et al. 2003),
freshwater influence (Wolcott 1988), sediments (Frusher et
Figure 2 - Density and abundance distributions for Ucides cordatus during al. 1994) and topography (Blankensteyn et al. 1997). Mean
the sampling months densities of 6.0 (Almeida and Melo 1996) and 1.65 burrows

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 Bioecology of the crab Ucides cordatus (Crustacea, Decapoda)
in mangroves influenced by the Amazon River, Brazil

Table 1 - Carapace linear measurements and individual weights of males and females of Ucides cordatus from the coast of Amapá.

Carapace measurements (mm)

Weight (g)
Values Length Width Height
Male Female Male Female Male Female Male Female
Mean (+SD) 51.9 (± 8.0) 46.1 (± 5.5) 69.1 (±11.2) 59.6 (± 7.7) 27.5 (± 11.3) 23.5 (± 3.6) 155.4 (± 64.8) 95,2 (± 28.8)
Minimum 19.4 28.1 19.0 15.2 13.8 12.3 11 21
Maximum 72.2 62.8 96.5 78.6 55.0 48.9 372 195
N 498 368 498 360 497 361 342 247
P (³0,05) 9.16 x 10 -34
3.0 x 10 -44
3.9 x 10 -13
1.43 x 10 -43

SD = Standard Deviation.

m-2 (Diele 2000) were recorded in Northern Brazil (Pará State). Once the State law of Amapá prohibits collecting females
In the Northeast, densities values were 5.58 in Maranhão during all months of the year, it was expected that the
(Castro et al. 2008); 1.26 in Rio Grande do Norte (Ivo et al. frequency of male individuals was lower than that of females.
2000) and 1.70 burrows m-2 in Paraíba (Alves and Nishida However, the total number of males collected exceeded the
2004). Mean densities were higher than the abundance number of females. The same pattern occurred in December,
values in most months, suggesting that the same individual May and August. In the States where the capture of females
may be occupying more than one burrow for shelter. In is allowed, except for the closed season (reproduction period),
addition, marks left by predators on the edge of the burrows the greater capture of males may be related to their larger
indicate that some of these burrows may be empty due to size. Glaser and Diele (2004) found that market demand for
predation, anthropogenic actions, or may have been caused females in the estuary Caeté, Pará State is very low and only
by other animals such as raccoons (Procyon sp.). Reproductive male individuals are landed in the local ports. A study in the
migration can also influence the abundance; however, this estuary of Ceará River, located in the municipality of Caucaia,
phenomenon was not observed in this study. Ceará State with a great tradition of crab consumption and
whose stocks are almost exhausted, showed the predominance
The abundance and density were associated in some of females over males (Alcântara-Filho 1978). The same
studies with reduced forest fragment size and/or increased situation occurred in Pernambuco with the predominance of
pressure on capture species of fish, mollusks and crustaceans females in the mangroves of Rio Formoso Estuary (Botelho et
in other parts of the world (Oribhabor and Udo 2011). al. 1999). Araújo and Calado (2008) indicated a numerically
According to Lara (2003), mangroves in the Amazon coast balanced population in Alagoas with no significant difference
are well preserved due to the lower anthropogenic pressure between the number of males and females. In this work, the
on this region. However, with increasing habitat destruction same situation was found in November, December and March,
and increased overexploitation of fishery stocks, those species despite the statistical analyses showing a significant difference
with low population densities are more likely to be impacted. between the total frequency of males and females.
In addition, mean density and mean abundance were Another difference observed between males and females
higher in the dry season months than in those of wet season. was regarding the linear measurements studied (height, width
Glaser and Diele (2004) reported that the number of and length of the carapace - CH, CW and CL) and the weight
animals captured by gatherers in Caeté, Pará State during the of individuals. In São Paulo (Pinheiro and Fiscarelli 2009),
rainy season was lower. One explanation for the significant Santa Catarina (Wunderlich et al. 2008), Piauí (Ivo et al. 1999)
reduction in the production of U. cordatus in wet season is and Rio Grande do Norte (Vasconcelos et al. 1999) males were
mainly due to the individuals’ behavior of crawling into the larger and heavier than females. On the other hand, Araujo
deepest parts of the galleries, making them difficult to collect and Calado (2008) found no significant difference between
(Oliveira et al. 2007) as observed in this study. The climate these parameters in Alagoas.
influence on other population parameters, such as density, Overall, males and females collected in the State of
may also be corroborated by other studies. In the State of Amapá had larger CW and CL compared to crabs studied
Maranhão (Castro 1986) changes were observed in the mean in the States of Ceará (Alcântara-Filho 1978), Maranhão
density of crabs between the rainy and dry seasons. However, (Castro 1986), Pernambuco (Botelho et al. 1999), Piauí (Ivo
in another work performed on the island of São Luis and the et al. 1999), Rio Grande do Norte (Vasconselos et al. 1999),
eastern coast of the State of Maranhão, Castro et al. (2008) Paraná (Dalabona et al. 2005), Pará (Diele et al. 2005) and
observed equivalent densities for both periods. Alagoas (Araújo and Calado 2008). The fact of the Amapá

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 Bioecology of the crab Ucides cordatus (Crustacea, Decapoda)
in mangroves influenced by the Amazon River, Brazil

Figure 3 - Relationship between carapace width and length for females (A) and males (B); and the relationship between carapace width and individual weight
for females (C) and males (D) of the crab Ucides cordatus collected in the State of Amapá.

State having population stocks consisting of large individuals size class 59.8 - 67.5 mm, but mature females also occurred in
may be related in part to the anthropogenic factors. The the smallest size class (14 - 21.6 mm). Moreover, most females
tradition of consuming the species in the North of Brazil is were sexually mature in the late May and early August. In the
not as intense as in the Northeast, which accounts for the Northeast, Alcântara-Filho (1978) and Costa (1979) described
highest percentage of captures in the country. In the latter, the reproductive period occurring from December to May,
the closed season coincides with the period of most intense although shorter periods - from January to May (Mota-Alves
tourism, being demanded a lot of animals to supply bars and 1975) and January to March (Castro 1986) - have also been
restaurants. Amapá is still a sparsely populated state and where described in the literature. Branco (1993) reported the mating
tourism is underdeveloped, which may also be contributing to of species in January to the South of Brazil only based on
the maintenance of large individuals. Moreover, the isolation observations on the reproductive behavior of individuals. In
of areas where U. cordatus occurs make capture more difficult Santa Catarina, Wunderlich et al. (2008) observed females
in some way. with mature gonads during five months of the year (November
In the ratios CW x CL and CW x IW of males and to March), with two major annual incidences in November
females collected in this study, the samples distribution (95.2%) and February (18.2%).
showed a straight line trend, therefore showing a relationship According to Sastry (1983), crustaceans can reproduce
between the parameters investigated. Considering both CW during all months of the year (continuous pattern) or be
x CL and CW x IW (males and females), the growth type restricted to a few months, where environmental conditions
was characterized as negative allometric. Regarding the CW are more favorable (discontinuous or seasonal pattern).
x CL, the animals showed negative allometric (males) and Sastry (1983) and Pinheiro and Fransozo (2002) report that
isometric (females) growth types in Ceará (Leite et al. 2006) U. cordatus has seasonal reproduction, once the record of
and Paraná (Dalabona et al. 2005). This indicates that the females with mature gonads occurs during only five months
carapace length increases to a lesser extent in males and with of the year (November to March), including the occurrence of
the same proportion in females. In São Paulo (Pinheiro and ovigerous females (December and January). Such information
Hattori 2006), Rio de Janeiro (Passos and Di-Beneditto 2005) corroborates Diele (2000) in the State of Pará, who shows
and Alagoas (Araújo and Calado 2008), negative allometry was the reproduction of this species restricted to the rainy season,
detected for both sexes. Concerning to the ratio IW x CW, following similar pattern to that of other semi-terrestrial and
studies by Pinheiro and Fiscarelli (2009) indicated isometric terrestrial crabs already studied (Henmi and Kaneto 1989;
growth for males and negative allometry for females. Adamczewska and Morris 2001). Therefore, the reproductive
Regarding the reproduction, the highest frequency of period of crabs collected in the State of Amapá is different
ovigerous and mature (stage IV) females was found in the CW from those in the aforementioned states.

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 Bioecology of the crab Ucides cordatus (Crustacea, Decapoda)
in mangroves influenced by the Amazon River, Brazil

Figure 4 - Size classes of carapace width for males and females of Ucides Figure 5 - Relative frequency for the female different gonadal maturation stages
cordatus. (Ovigerous; I-V) of Ucides cordatus during the sampling months.

ACKNOWLEDGMENTS Branco, J. O. 1993. Aspectos Bioecológicos do Caranguejo Ucides

cordatus cordatus (Linnaeus,1763) (Crustacea, Decapoda)
Funding was provided by Empresa Brasileira de do Manguezal do Itacurubi, Santa Catarina, BR. Arquivos de
Pesquisa Agropecuária (EMBRAPA). The authors thank the Biologia e Tecnologia, 36: 133–148.
environmental analysts and technicians from the “Instituto Botelho, E. R. O.; Dias, A. F.; Ivo, C. T. 1999. Estudo sobre a
Brasileiro do Meio Ambiente e dos Recursos Naturais biologia do caranguejo-uçá, Ucides cordatus (Linnaeus, 1763),
Renováveis - IBAMA” and from the “Instituto Chico capturado nos estuários dos Rios Formoso (Rio Formoso) e
Mendes de Conservação da Biodiversidade - ICMBio” for Ilhetas (Tamandaré), no Estado de Pernambuco. Boletim Técnico
Científico do CEPENE, 7: 117-145.
their logistical support offered to this work. Also, we thank
Frederico Fonseca Galvão de Oliveira for the map confection Castro, A. C. L. 1986. Aspectos Bio-Ecológicos do Caranguejo-uçá,
Ucides cordatus cordatus (Linnaeus, 1763), no estuário do Rio
and the crab collectors for their assistance with the fieldwork.
dos Cachorros e Estreito do Coqueiro, São Luís – MA. Boletim
do Laboratório de Hidrobiologia, 7: 7-26.
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Recebido em 28/02/2013
Aceito em 01/08/2013

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