Relative growth of the mangrove crab Ucides cordatus (Linnaeus, 1763) (Crustacea, Brachyura, Ocypodidae) at Iguape, São Paulo, Brazil

BIOLOGICAL AND APPLIED SCIENCES

Relative growth of the mangrove crab Ucides cordatus (Linnaeus, 1763) (Crustacea, Brachyura, Ocypodidae) at Iguape, São Paulo, Brazil

Marcelo Antonio Amaro PinheiroI, ^*; Gustavo Yomar Hattori^I,II

^IGrupo de Pesquisa em Biologia de Crustáceos - CRUSTA; UNESP São Vicente; CSV; [e-mail]; Pç. Infante D. Henrique, s/n; 11330-900; São Vicente - SP - Brasil

^IIPrograma de Pós-Graduação em Zootecnia; Área de Concentração Produção Animal; UNESP Jaboticabal; FCAV; [e-mail]; R. Prof. Paulo Donato Castellane, s/n; 14884-900; Jaboticabal - SP - Brasil

ABSTRACT

A total of 2,130 individuals of Ucides cordatus (1,255 males and 875 females) were captured in a mangrove forest at Iguape, São Paulo, Brazil. For each crab, the following body structures were measured: carapace (width = CW; length = CL; depth = CD), 5^th abdominal somite (AW), major chelar propodus (length = PL; width = PW; depth = PD), and 1^st and 2^nd gonopod pairs (length = GL₁ and GL₂). The Student "t" and Snedecor "F" tests were used to verify any changes in growth allometric rates during ontogeny. The relationships CLxCW, PLxCW (for both sexes), GL₁xCW and GL₂xCW (males) and AWxCW (females), showed a better fit by two equations for the juvenile and adult phases (p<0.01). The inflexion point size between regression lines, indicated by each morphometric relationship, allowed to propose four morphotypes for U. cordatus. Males were classified in juvenile (CW < 32 mm), pre-puberty (32 < CW < 51 mm), sub-adult (51 < CW < 59 mm) and adult (CW > 59 mm). Females showed a similar size interval: (juvenile CW < 39 mm, pre-puberty 39 < CW < 53 mm, sub-adult 53 < CW < 58 mm, adult CW > 58 mm).

Key words: Relative growth, Brachyura, Ocypodidae, Ucides

RESUMO

Um total de 2,130 indivíduos de U. cordatus (1,255 machos e 875 fêmeas) foi coletado em Iguape (SP), Brasil. Cada exemplar foi submetido à biometria das seguintes estruturas: cefalotórax (largura = CW; comprimento = CL; altura = CD), quinto somito abdominal (AW), própodo quelar (comprimento = PL; espessura = PW; altura = PD), e 1^º e 2^º par de gonopódios (comprimento = GL₁ e GL₂). O teste "t" Student e "F" de Snedecor foram utilizados para identificar diferenças no grau de alometria e alterações ontogenéticas na taxa de crescimento, respectivamente. As relações CLxCW, PLxCW (ambos os sexos), GL₁xCW e GL₂xCW (machos) e AWxCW (fêmeas), apresentaram ajuste por duas equações representando a fase jovem e adulta (p<0.01). A determinação dos tamanhos indicados pelas relações morfométricas permitiu a divisão de cada sexo em quatro morfotipos. Os machos foram classificados como jovens (CW<32mm), pré-púberes (32<CW<51mm), sub-adultos (51<CW<59mm) e adultos (CW>59mm), com tamanho similar ao dos morfotipos das fêmeas (jovens CW<39mm; pré-púberes 39<CW<53mm; sub-adultas 53<CW<58mm; adultas CW>58mm).

INTRODUCTION

Ucides cordatus (Linnaeus, 1763) is a semiterrestrial crab that lives only in mangrove areas and occurs throughout the western Atlantic Ocean, from Florida, USA to Santa Catarina State, Brazil (Melo, 1996). Because of its large size and tasty meat, this crab has been exploited in many parts of Brazil (Rodrigues et al., 2000). The relative growth of several species of ocypodids has been studied (Crane, 1941; Barnes, 1968; Haley, 1969, 1973; Miller, 1973; von Hagen, 1987), including fiddler crabs of the genus Uca (Frith and Brunmeister, 1983; Negreiros-Fransozo et al., 2003). The biometric studies of U. cordatus have emphasized dimorphism of the chelae (Santos and Garcia-Mendes, 1982), or allometric growth of the carapace (Branco, 1993). Recently, Dalabona et al. (2005) described the relative growth of this species from a mangrove area on southern Brazilian coast.

Ontogeny in brachyurans is marked by morphological changes related to sex and puberty, which were first observed by Huxley (1924, 1950). Biometric changes in the chelae, abdomen and pleopods are evident in both sexes and in the developmental stages of crustaceans, principally during the transition from the immature to the mature stage, when the puberty molt occurs (Hartnoll, 1974, 1982). Analyses of these changes allow mathematical equations to be determined that could be used to convert biometric variables (Pinheiro and Fransozo, 1993), estimate the size at puberty (Pinheiro and Fransozo, 1998), and also to identify brachyuran species (Huber, 1985).

The present study describes the relative growth of U. cordatus, analyzing the biometric relationships between the carapace (length and width), the major chelar propodus (length, width and depth) and the abdomen for each sex. For males, the lengths of the first and second pair of gonopods were also analyzed. These data were used to identify possible changes in the allometric growth rates between development phases (juvenile and adult), and to propose a description of the morphotypes, based on the size estimated at the puberty molt.

MATERIALS AND METHODS

Individuals of U. cordatus were captured monthly from September 1998 to September 2000, in mangrove forests near Icapara Bar (24°41'S), Iguape, state of São Paulo, Brazil, by digging or trapping. The specimens were kept frozen separately until the biometric analysis. Each crab was sexed, and then measured with a vernier caliper to the nearest 0.05 mm. The following dimensions were selected for morphometric analysis: carapace (CW = width; CL = length; CD = depth), abdomen (AW = width of 5^th somite), major chelar propodus (PL = length; PW = width; PD = depth) and male gonopods (GL₁ and GL₂ = lengths of the 1^st and 2^nd pair of gonopods, respectively) (Fig. 1). All the biometric relationships were submitted to regression analyses by a power function (y = ax^b) (Huxley, 1950) with respect to the variable CW. The biometric relationships were verified by a coefficient of determination (R²), while Snedecor's F test (a = 0.01) (Sokal and Rolf, 1995) was used to verify the presence of one or two regression lines for the empirical points. All the biometric relationships were submitted to MATURE 1 (Somerton, 1980) or MATURE 2 software (Somerton and MacIntosh, 1983), to identify the inflection point at puberty size.

The allometric growth rate of each developmental phase was established by the "b" value, and considered as isometry (b = 1), positive allometric (b > 1) or negative allometric (b < 1). Student "t" test was used to verify the significance of this value related to the unit (a = 0.01).

The morphometric relationships, which showed allometric growth rate changes according to the MATURE software, had their biometric equation constants submitted to a "t" (z) test (Santos, 1978), to determine if juveniles of each sex could be clustered by a single equation. The same procedure was used for adults.

RESULTS

Table 1 lists all the variables obtained by biometric analyses of 2,130 specimens (1,255 males and 785 females). All the equations calculated for biometric relationships showed a close fit, with the coefficient of determination higher than 0.9 in 67.6% of all cases (Table 2). The CLxCW relationship evidenced negative allometric growth for both males (t = 11.4; p<0.01) and females (t = 2.9; p<0.01). Even then, the constant "b" showed a strong tendency toward isometry (b_males = 0.95; b_females = 0.96). The empirical points of each sex were better fitted by two equations, with changes in allometric rates during ontogeny (F_males = 6.9; F_females = 6.6; p<0.01). The comparison of the cut point size between juvenile and adult was similar in the size estimated (CW_males = 59.1 mm, Fig. 2-A; CW_females = 58.2 mm; Fig. 2-B). The juveniles of each sex tended toward isometry, but this tendency was statistically significant only for females (b_females = 0.99; t = 0.2; p>0.01). The adults showed negative allometry (b_males = 0.89; b_females = 0.84; p<0.01), which could be represented by a single equation (p>0.05) (Table 3).

The CDxCW relationship did not reveal any change in growth rate during ontogeny. This relationship indicated isometry for females (t = 0.16; p>0.01) and negative allometry for males (t = 15.2; p<0.01).

The points of the PLxCW relationship showed a better fit for two equations in both sexes (F_males = 4.9; F_females = 11.2; p<0.01), the inflexion point was 51.3 mm for males (Fig. 3-A) and 52.6 mm for females (Fig. 3-B). The sexes differed in the degree of allometry: males showed positive allometric growth (b_adults = 1.47), whereas growth in females was isometric (b_adults = 1.01).

The PWxCW relationship for both sexes was represented by two equations, (F_males = 23.99; F_females = 17.38; p<0.01), with the cut points at 63.8 mm (males) and 51.4 mm (females). The degree of allometry did not differ between the sexes; it was positive allometric for juveniles (b_males = 1.34; b_females = 1.20) and isometric in adults (b_males = 0.94; b_females = 0.91). The adult phase could be clustered and represented by one equation (p>0.05) (Table 3).

For both sexes, the PDxCW relationship was best expressed by two equations (F_males = 16.2; F_females = 19.2; p<0.01), with differences in the degree of allometry between the developmental phases. In juveniles, growth was positive allometric, more strongly in males (b_males = 1.25; b_females = 1.18), but showing a different growth pattern in the adult phase. Isometry was verified for males (b = 0.99) and negative allometry for females (b = 0.89). In no phase of development, the biometric variable of this relationship could be clustered (p<0.05).

The AWxCW relationship for males showed negative allometric growth (b = 0.88), best fitted by a single regression line (F = 0.53; p>0.01). Contrariwise, females showed positive allometry and could be represented by two lines (F=229.8; p<0.01), with the inflexion point at 39.1mm (Fig. 4).

The GL₁xCW was best fitted by two regression lines (F = 40.9; p<0.01) and was characterized by positive allometric growth in the juvenile phase (b = 1.49) and negative allometry in adults (b = 0.95), with the cut point at 31.9 mm. The same pattern occurred for the GL₂xCW relationship (F = 59.6; p<0.01), with the inflexion point 58.9% higher than estimated by the GL₁xCW (50.7 mm). The inflexion point sizes obtained for all the morphometric relationships allowed us to propose four morphotypes for each sex. Males were classified in juvenile (CW < 32 mm), pre-puberty (32 < CW < 51 mm), sub-adult (51 < CW < 59 mm) and adult (CW > 59 mm), with a similar division for females (juvenile CW < 39 mm, pre-puberty 39 < CW < 53 mm, sub-adult 53 < CW < 58 mm, and adult CW > 58 mm).

DISCUSSION

In the studies on relative growth, Hartnoll (1974, 1978, 1982) observed that morphometric variables related to the carapace were characterized by isometry, which was represented by some authors as an interval of 0.9 < b < 1.1 (Kuris et al., 1987; Pinheiro and Fransozo, 1993). However, the use of statistical tests invalidated the hypothesis of isometric growth for this biometric relationship in certain species (Finney and Abele, 1981; Davidson and Marsden, 1987). This growth pattern was demonstrated by Barnes (1968) in biometric analyses of the ocypodids Macrophthalmus spp., and it was also observed in the present study.

In many of the brachyurans that have been studied, the carapace relationships did not indicate a change in growth pattern during ontogeny. This body structure has not been used to estimate the size at morphological maturity (Somerton, 1980; Somerton and MacIntosh, 1983; Pinheiro and Fransozo, 1998). However, Aguilar and Spina (1988) observed for females of the callapid Mursia gaudichaudi (H. Milne Edwards, 1837), a synchrony between the size at onset of sexual maturity and the inflexion point size obtained by CLxCW. A similar synchrony was not seen in U. cordatus, since the size at gonadal maturity was lower than indicated by the CLxCW relationship (CW_males = 51 mm and CW_females = 43 mm, according to Hattori and Pinheiro, submitted). The greater increment of CW size from 59 mm might be associated with the increase in the size of the gill chamber in the adult phase. A similar fact was observed by Gifford (1962) for Cardisoma guanhumi (Latreille, 1825), permitting a characterization of three morphotypes for this species. The CLxCW relationship could be used as an important character to propose morphotypes, since the juveniles' growth was different between the sexes and the adult phase could be represented by a single equation (Table 3). Branco (1993) recorded isometric growth in U. cordatus using a linear function (y = a + bx), similar observed by other authors (Botelho et al., 1999; Ivo et al., 1999; Vasconcelos et al., 1999). However, these authors did not subject this biometric relationship data to statistical analyses that could recognize differences in allometric growth between the developmental phases. Dalabona et al. (2005) studied this biometric relationship for this species, but were not observed a difference in growth rates during the ontogeny. This pattern was registered by these authors, probably due to the reduced number of individuals used in their biometric regression analyses.

There are few published studies of the morphometric relationships between gonopod size and the carapace, particularly in regard to the 2^nd gonopod pair. The two gonopod pairs of U. cordatus showed a growth type similar to "A" described by Somerton (1980), with a distinct cut point between juvenile and adult phases. The same pattern was observed by Haley (1969) for Ocypode quadrata (Fabricius, 1787) and the xanthids Eriphia smithi MacLeay 1838 and E. gonagra (Fabricius, 1781), studied by Vaninni and Gherardi (1988) and Góes and Fransozo (1997), respectively.

Hartnoll (1965) and Flores and Negreiros-Fransozo (1999) also observed a low growth rate of the 1^st gonopod pair after puberty in grapsid crabs. This slow growth could be a reproductive advantage, because males could copulate with females of a wide range of sizes, improving their reproductive output (Hartnoll, 1974). Male brachyurans use this abdominal appendix as a copulatory organ, this structure protects the 2^nd gonopod pair, which is smaller and poorly calcified, and it is incapable by itself of executing the mating (Pinheiro and Fransozo, 1999). Males showed the 1^st gonopod pair size at the same proportion of adults with 31.9 mm CW, but they required other characters such as a larger chela and mature gonads for mating success (Góes et al., 2000). Haley (1969) observed a coincidence between the allometric changes in the 1^st gonopod pair and the size at gonadal maturity for males of Ocypode quadrata. However, in U. cordatus the 2^nd gonopod pair showed a better association with gonadal maturation. The inflection point size obtained (50.7 mm) was near that estimated by male gonadal maturity, according to Hattori and Pinheiro (submitted).

The PLxCW relationship for both sexes was similar to the "A" growth type described by Somerton (1980), with a 7.3% increase in the male's chelar propodus length from 51.3 mm CW. The same growth pattern was observed for the other males' chelar variable; however, the chelae length was more evident, similar to data for the portunid Arenaeus cribrarius, analyzed by Pinheiro and Fransozo (1993) and U. cordatus by Dalabona et al. (2005). According to Dalabona et al. (2005), the major and minor males chelae show a different pattern in allometric growth rates. The major chela had the same allometry rates throughout the ontogeny, which was observed in the present study. However, the minor chela of U. cordatus males studied by those authors, showed a positive allometry in juveniles and isometry in adults, this difference in allometric growth rates became the heterochely in males more evident.

The high rate of chelae growth in brachyuran males makes the reproductive behavioral display efficient, because they use it to manipulate the females during mating (Hartnoll, 1969; Hazlett, 1975; Pinheiro and Fransozo, 1999). Jivoff (1997a, b) studied the reproductive behavior of Callinectes sapidus, and showed that larger-sized males with large chelae had an advantage in partner selection. This assumes great importance for semiterrestrial and terrestrial crabs, where visual and tactile stimuli are most important for couple formation (Hartnoll, 1969; Pinheiro and Fransozo, 1999; Góes et al., 2000). According to Góes et al. (2000), during the mating period, the male of U. cordatus uses the major chela to strike another male's carapace when they fight over a female. Another hypothesis is of sex recognition by the males' chelae size, because females showed 17% reduction of chelae growth rates at 52.6 mm CW, whereas the inverse occurred in males of similar size.

In brachyurans, the relative growth of the abdomen has been used only to estimate the females' size at puberty, because certain somites showed striking modification in growth and morphology during ontogeny (Huxley and Richards, 1931), whereas similar changes were not observed in males (Hartnoll, 1974). Development of this female body structure serves to bring them to an efficient size and shape to carry and protect the incubating eggs (Simons, 1981). In males, the abdomen is only used as a support structure for the pleopods, with a copulatory function (Pinheiro and Fransozo, 1993).

All the biometric relationships for U. cordatus characterized by two regression lines allowed us to distinguish four morphotypes in this species: juvenile, pre-puberty, sub-adults and adults. In Table 4 describes each one in detail. The relative growth was helpful to establishe the morphological maturity of this species. This information could be useful in future studies, for instance projects related to management of this mangrove crab.

ACKNOWLEDGEMENTS

We thank FAPESP to providing financial support for the Uçá Project (#1998/6055-0), and CAPES for fellowship to the second author. Thanks are also due to Biologist Ana G. Fiscarelli for the illustrations and her assistance in fieldwork, and to Biologist Maristela D. Baveloni and other members of CRUSTA (Research Group in Crustacean Biology) for their help in fieldwork and laboratory analyses.

Received: February 14, 2005;

Revised: August 29, 2005;

Accepted: March 30, 2006.

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*

Author for correspondence

Publication Dates

• Publication in this collection
  23 Feb 2007
• Date of issue
  Sept 2006

History

• Accepted
  30 Mar 2006
• Reviewed
  29 Aug 2005
• Received
  14 Feb 2005